taxonID	type	description	language	source
03E987BFFFB6CE0F4E9D544CA805A943.taxon	description	Figs 1 – 11, 13 Synonyms: Cynips quercus hirta Bassett (1864: 688), females and galls. Cynips (Teras) hirta combination by Osten Sacken (1865: 333); Biorhiza hirta combination by Ashmead (1885: 296); Biorrhiza hirta combination and incorrect spelling by Dalla Torre (1893: 60); Philonix hirta combination by Smith (1900: 548); Phylonyx hirta combination and incorrect spelling by Smith (1910: 598); Trichoteras quercus-hirtum combination by Dalla Torre & Kieffer (1910: 404); Biorhiza (Xystoteras) hirta combination by Britton (1920: 319); Acraspis hirta combination by Weld (1922 a: 10); Cynips (Acraspis) hirta var. hirta combination by Kinsey (1930: 426); Cynips (Acraspis) hirta combination by Kinsey (1936: 285); Acraspis quercushirta corrected spelling by Burks (1979: 1079).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFB6CE0F4E9D544CA805A943.taxon	materials_examined	Material examined: Six sexual females labelled as “ Canada, Alberta, Edmonton, Royal Alberta Museum, 53.54218 ° N, - 113.54289 ° W, GPS ± 10 m, S. Digweed, Collected 22. v. 2005 ex Quercus macrocarpa, Reared 25. v. 2005 ex gall of Acraspis macrocarpae ♂ ♀ ”, determined originally as Acraspis macrocarpae sexual generation females by S. Digweed 2005. Six sexual females have been deposited at the PHDNRL.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFB6CE0F4E9D544CA805A943.taxon	diagnosis	Diagnosis. Closely resembles A. villosa. In A. quercushirta the radial cell of the forewing 4.2 × as long as broad; the eye 4.5 × as high as length of malar space, inner margins of eyes slightly converge ventrally; POL 2.2 × as long as OOL, OOL 1.2 × as long as diameter of lateral ocellus. In A. villosa the radial cell of the forewing 5.0 × as long as broad, the eye 2.5 × as high as length of malar space, inner margins of eyes are parallel; POL 1.8 × as long as OOL, OOL 1.75 × as long as diameter of lateral ocellus.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFB6CE0F4E9D544CA805A943.taxon	description	Description. Sexual female (Figs 1 – 11). Head, mesosoma, metasoma uniformly dark brown; antenna slightly lighter than body; mandibles, palpi maxillaris and labialis, legs light brown to yellowish. Head alutaceous, with white setae, denser on lower face and postgena, with distinct rows of setae along inner margin of eye; rounded, 1.1 × as broad as high and as broad as mesosoma in frontal view, 1.8 × as broad as long from dorsal view. Gena alutaceous, not broadened behind eye in frontal view, narrower than transverse diameter of eye in lateral view. Malar space alutaceous, glabrous, with very delicate striae that radiate from clypeus and do not reach eye; eye 4.5 × as high as length of malar space. Inner margins of eyes slightly converge ventrally. POL 2.2 × as long as OOL, OOL 1.2 × as long as diameter of lateral ocellus and slightly shorter than LOL, all ocelli small, ovate, of same size. Transfacial distance as great as height of eye; diameter of antennal torulus 1.3 × as long as distance between them, distance between torulus and eye as long as diameter of torulus; lower face alutaceous, matte, slightly elevated median area alutaceous. Clypeus rounded, nearly as broad as high, alutaceous, matte, with a few long setae; ventrally rounded, slightly emarginate, without median incision; anterior tentorial pit small, rounded, epistomal sulcus distinct, clypeo-pleurostomal line well impressed. Frons and interocellar area uniformly alutaceous, without striae and setae. Vertex, occiput and postocciput alutaceous; postgena smooth, glabrous, with dense white setae; posterior tentorial pit small, rounded, area below impressed; occipital foramen as high as height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which are not united and diverge toward occipital foramen, postgenal bridge anteriorly as broad as occipital foramen. Antenna longer than head + mesosoma, with 12 flagellomeres, pedicel slightly longer than broad, flagellomeres slightly broader towards apex; F 1 2.4 × as long as pedicel and 1.4 × as long as F 2; F 2 slightly longer than F 3; F 3 1.2 × as long as F 4, F 5 slightly longer than F 6, all subsequent flagellomeres equal in length; placodeal sensilla on F 2 – F 12, absent on F 1. Mesosoma 1.1 × as long as high, with sparse white setae, except for dense setae on lateral propodeal area, mesopleural triangle and propleuron. Pronotum smooth, matte, with sparse setae, without striae laterally; propleuron alutaceous, matte. Mesoscutum smooth, glabrous, with sparse white setae along notauli, slightly longer than broad (greatest width measured across mesoscutum level with base of tegulae). Notaulus incomplete, extending to 2 / 3 of mesoscutum length, shallow, broad, bottom smooth, glabrous, posteriorly broader and slightly converging; anterior parallel line and median mesoscutal line absent; parapsidal line indistinct, in the form of an impressed line; circumscutellar carina narrow, distinct, reaching notaulus. Mesoscutellum ovate, slightly longer than broad, with semi-parallel sides, coriaceous, overhanging metanotum, posteriorly rounded; disk alutaceous with delicate rugae laterally and posteriorly. Mesoscutellar foveae absent, anterior part of mesoscutellum is not impressed, in the same plane as the mesoscutellar disk, alutaceous to smooth. Mesopleuron smooth, matte, setae only on posteroventral quarter of mesopleuron; mesopleural triangle delicately coriaceous, matte, with some delicate longitudinal parallel striae and a few setae; dorsal and lateral axillar areas smooth, glabrous, with dense setae; subaxillular bar smooth, glabrous, triangulate, gradually higher toward posterior end, at posterior end as high as height of metanotal trough; metapleural sulcus reaching mesopleuron slightly below half of its height, upper part of sulcus indistinct. Metascutellum coriaceous, as high as height of coriaceous ventral impressed area; metanotal trough smooth, matte, without setae; central propodeal area lyre-shaped, rugose, matte, with net of irregular rugae continuing onto nucha; lateral propodeal carinae bent slightly outwards in posterior 1 / 3, complete or incomplete, do not reaching anterior end of propodeum; lateral propodeal area coriaceous, matte, with long white setae. Nucha with net of irregular rugae dorsally and laterally. Tarsal claws with basal lobe. Forewing longer than body, hyaline, with long dense cilia on margin, veins brown, radial cell open, 4.2 × as long as broad; R 1 and Rs reaching wing margin; areolet small, triangular, closed, distinct. Rs + M narrow, reaching basalis at mid height. Metasoma as long as head + mesosoma, longer than high in lateral view; 2 nd metasomal tergum extending to 1 / 3 length of metasoma in dorsal view, without setae or micropunctures; subsequent terga smooth, glabrous, without micropunctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium longer than broad in ventral view, with a few short setae ventrally which extend beyond apex of spine. Body length 1.8 – 1.9 mm (n = 4). Male. The male as in Acraspis villosa; it is impossible to distinguish males of the two species using morphological characters (see male description for A. villosa). Gall (Fig. 13). Sexual galls develop in the innermost scales in apical and lateral buds, a brittle, thin-walled, seed-like, sub-ovoid cell approximately 2 mm long, tan or orange-brown when mature, with faint longitudinal striations and sparse, pale hairs. Galls form in the basal portion of the scale and often leave the scale apex intact, which extends as a flat flap beyond the gall.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFB6CE0F4E9D544CA805A943.taxon	biology_ecology	Biology. See Bassett (1864), Weld (1922 a, 1926), and Kinsey (1930, 1936) for the biology of the asexual generation. Asexual generation galls (Fig. 12) have been recorded on leaves of Q. chapmanii, Q. gambelii, Q. macrocarpa, Q. michauxii, and Q. montana. Adult females of the asexual generation emerged in Edmonton from late September to early November, with the peak occurring during the last half of October. Bassett (1864) noted that adult females of the asexual generation bore exit holes in their galls some time prior to emerging. This behaviour was observed in Edmonton as well, with females often remaining for days or weeks just inside emergence holes. Completion of emergence was often triggered by cooler temperatures. Leaves with galls have mostly fallen to the ground when the brachypterous females emerge in Edmonton, so they walk to a bur oak tree bole, climb it, and oviposit into bud scales, remaining active for up to 2 – 3 weeks, often at temperatures below freezing. Galls of the sexual generation usually occur singly, but up to six galls were found around a single bud. Galls and inhabitants survived freezing in late April 2005 in Edmonton that destroyed new shoots on many bur oaks. Emergence typically occurs through holes chewed near gall apices. Empty galls can persist around bases of new shoots for weeks or months, but are easily detached as bud scales slough off during shoot maturation. Adults of the sexual generation emerged in Edmonton from late May to early June. Galls of the asexual generation appeared on leaves starting in late June or early July.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFB6CE0F4E9D544CA805A943.taxon	distribution	Distribution. USA: Connecticut south to Virginia, Illinois, Michigan, Texas, Utah (Burks 1979), Colorado, New Mexico, Oklahoma (Kinsey 1930). Canada: Alberta, Saskatchewan, Manitoba, Ontario, Quebec, New Brunswick. Probably everywhere east of the Rocky Mountains in the USA and Canada where white oaks occur (Kinsey 1930) or have been introduced as urban trees (e. g., bur oak in Alberta). Molecular taxonomy. Alternating generations were matched using DNA data, with four individuals (two asexual females, two sexual females) from Canadian Q. macrocarpa rearings sequenced for both cytb and ITS 2. Cytb sequences were 0 – 0.23 % divergent (GenBank accessions OM 321608 – OM 321610, OM 321612); ITS 2 sequences were identical (OM 331800 – OM 331802, OM 331805). The conspecific status of A. macrocarpae and A. quercushirta was confirmed by comparing sequences from the four Canadian individuals (originally determined as A. macrocarpae) with those from two south-eastern USA individuals (determined as A. quercushirta; cytb: KX 683594, OM 321611; ITS 2: OM 331803 – OM 331804). Cytb haplotype divergence between the two regions averaged 2.08 % (range 2.54 – 1.62 %), with greater variation (3.23 %) actually observed between the two USA haplotypes. ITS 2 sequences were 0.38 % different between Canadian and USA specimens, with four additional single base indels in polyA or polyT repeat units.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFBACE0B4E9D56F4A8F4AE0F.taxon	description	Figs 14 – 26, 28 Synonyms: Acraspis villosus Gillette (1888: 218), females and galls. Acraspis villosa corrected spelling by Dalla Torre (1893: 64); Philonix villosus combination by Felt (1906: 713); Philonix villosa combination by Beutenmueller (1909: 249); Cynips (Acraspis) villosa var. villosa combination by Kinsey (1930: 355). Kinsey (1930, 1936) described the asexual generation of six additional species within his “ villosa ” complex: Cynips (Acraspis) apache Kinsey, 1930 from Q. grisea and Q. arizonica; Cynips (Acraspis) calvescens Kinsey, 1930 from Q. gambelii; Cynips (Acraspis) consocians Kinsey, 1930 from Q. macrocarpa; Cynips (Acraspis) erutor Kinsey, 1936 from Q. repanda; Cynips (Acraspis) expletor Kinsey, 1936 from Q. oblongifolia; and Cynips (Acraspis) expositor Kinsey, 1930 from Q. grisea. Weld (1951) and Burks (1979) treated four of these species as varieties of A. villosa; hence Acraspis villosa var. apache (Kinsey), syn. nov., Acraspis villosa var. calvescens (Kinsey), syn. nov., Acraspis villosa var. consocians (Kinsey), syn. nov., and Acraspis villosa var. expositor (Kinsey), syn. nov. are all synonyms of A. villosa. Two species known only from Mexico are valid and treated as Cynips erutor Kinsey and C. expletor Kinsey (Pujade-Villar & Ferrer-Suay 2015).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFBACE0B4E9D56F4A8F4AE0F.taxon	materials_examined	Material examined: 2 sexual females labelled as “ Canada, Alberta, Edmonton, Northern Forestry Centre, 53.49165 ° N, - 113.54291 ° W, ± 10 m, Collected 4. vi. 2007 ex in situ mesh bag on branch of Quercus macrocarpa for rearing Acraspis villosa ♂ ♀ ”, determined originally as Acraspis villosa sexual generation females by S. Digweed, 2007. Two males labelled as “ Canada, Alberta Edmonton, NoFC, Reared 1. VI. 2007, ex Quercus macrocarpa, bag for rearing. Acraspis villosa, sexual. S. Digweed ”. Specimens have been deposited at the PHDNRL and RAM.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFBACE0B4E9D56F4A8F4AE0F.taxon	diagnosis	Diagnosis. Closely resembles Acraspis gemula (Bassett, 1881) and A. quercushirta. In A. villosa notauli are incomplete, entire body dark brown to black. In A. gemula notauli are complete, entire body reddish brown. See also Diagnosis to A. quercushirta.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFBACE0B4E9D56F4A8F4AE0F.taxon	description	Description. Sexual female (Figs 14 – 17, 22 – 26). Head, mesosoma, metasoma uniformly dark brown to black; scapus, pedicel, F 1 light brown, subsequent flagellomeres darker, slightly lighter than body; mouthparts and legs light brown to yellowish. Head alutaceous, with white setae, denser on lower face and postgena, rounded, 1.2 × as broad as high and as broad as mesosoma in frontal view, 1.6 x as broad as long from dorsal view. Gena alutaceous, not broadened behind eye in frontal view, narrower than transverse diameter of eye in lateral view. Malar space alutaceous, glabrous, without striae; eye 2.5 × as high as length of malar space. Inner margins of eyes parallel. POL 1.8 × as long as OOL, OOL 1.75 × as long as diameter of lateral ocellus and 1.2 × as long as LOL, all ocelli small, ovate, of same size. Transfacial distance 1.2 × as long as height of eye; diameter of antennal torulus 1.9 × as long as distance between them, distance between torulus and eye 1.3 × as long as diameter of torulus; lower face alutaceous, matte, with dense white setae, slightly elevated median area alutaceous. Clypeus rectangular, nearly 2.0 × as broad as high, smooth, matte, with long setae; ventrally rounded, slightly emarginate, without median incision; anterior tentorial pit small, rounded, epistomal sulcus distinct, clypeo-pleurostomal line well impressed. Frons and interocellar area uniformly alutaceous, without striae or setae. Vertex, occiput and postocciput alutaceous; postgena smooth, glabrous, with dense white setae; posterior tentorial pit small, rounded, area below impressed; occipital foramen greater than 2.0 × as high as height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which diverge toward occipital foramen, postgenal bridge anteriorly as broad as occipital foramen. Antenna longer than head + mesosoma, nearly as long as body, with 12 flagellomeres, pedicel slightly longer than broad, flagellomeres slightly broader towards apex; F 1 2.5 × as long as pedicel and 1.3 × as long as F 2; F 2 slightly longer than F 3; F 3 1.1 × as long as F 4, F 5 slightly longer than F 6, all subsequent flagellomeres equal in length; placodeal sensilla on F 2 – F 12, absent on F 1. Mesosoma 1.2 × as long as high, with sparse white setae, except for dense setae on lateral propodeal area. Pronotum smooth, matte, with sparse setae, without striae laterally; propleuron alutaceous, matte. Mesoscutum smooth, glabrous, with sparse white setae, slightly longer than broad (greatest width measured across mesoscutum level with base of tegulae). Notaulus incomplete, extending to 2 / 3 of mesoscutum length, shallow, broad, bottom smooth, glabrous, posteriorly broader and slightly converging; anterior parallel line, parapsidal line, median mesoscutal line absent; circumscutellar carina narrow, distinct, reaching notaulus. Mesoscutellum ovate, slightly longer than broad, with semi-parallel sides, coriaceous, overhanging metanotum, posteriorly rounded. Mesoscutellar foveae absent, anterior part of mesoscutellum is not impressed, in the same plane as mesoscutellar disk, with the same sculpture as the entire mesoscutellum. Mesopleuron smooth, matte, setae only on posteroventral quarter of mesopleuron; mesopleural triangle delicately coriaceous, matte, with some delicate longitudinal parallel striae and a few setae; dorsal and lateral axillar areas smooth, glabrous, with a few setae; subaxillular bar smooth, glabrous, triangulate, gradually higher toward posterior end, at posterior end as high as height of metanotal trough; metapleural sulcus reaching mesopleuron slightly below half of its height, upper part of sulcus indistinct. Metascutellum coriaceous, as high as height of coriaceous ventral impressed area; metanotal trough smooth, matte, without setae; central propodeal area lyre-shaped, rugose, matte, with few rugae in posterior half which continue onto nucha; lateral propodeal carinae bent slightly outwards in posterior 1 / 3, complete or incomplete, do not reach anterior end of propodeum; lateral propodeal area coriaceous, matte, with long white setae. Nucha with net of irregular rugae dorsally and laterally. Tarsal claws with small basal lobe. Forewing longer than body, hyaline, with distinct dense cilia on margin, veins brown, radial cell open, 5.0 × as long as broad; R 1 and Rs reaching wing margin; areolet large, triangular, closed, distinct. Rs + M distinct, reaching basalis slightly below its mid height. Metasoma slightly longer than head + mesosoma, longer than high in lateral view; 2 nd metasomal tergum extending to half length of metasoma in dorsal view, with sparse scattered setae laterally; all terga alutaceous, matte, without micropunctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium longer than broad in ventral view, with some short setae ventrally which extend beyond apex of spine. Body length 1.8 mm (n = 1). Male (Figs 18 – 21). Similar to female, head, mesosoma and metasoma black to dark brown, mandibles and palpi maxillaris and labialis light brown, antenna brown; legs uniformly light brown to yellow. Eye 3.3 × as high as length of malar space; ocelli much bigger than in female. Antenna as long as body, forewing longer than body; metasoma slightly shorter than mesosoma. Antenna with 13 flagellomeres, scapus + pedicel shorter than F 1; F 1 slightly curved, excavated, apically broadened, 1.2 × as long as F 2; F 13 longer than F 12; placodeal sensilla on all flagellomeres. Metasoma with long narrow petiole, 2 nd metasomal tergum extending to half length of metasoma, with a few white setae laterally; all terga smooth, glabrous, without micropunctures. Gall (Fig. 28). The sexual galls are not apparently distinguishable from galls of the sexual generation of Acraspis quercushirta, described above.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFBACE0B4E9D56F4A8F4AE0F.taxon	biology_ecology	Biology. See Weld (1926) and Kinsey (1930, 1936) for the biology of the asexual generation. Asexual galls (Fig. 27) from species in Kinsey’s (1936) “ villosa ” complex have been recorded on leaves of Q. alba, Q. arizonica, Q. gambelii, Q. grisea, Q. macrocarpa and Q. oblongifolia. Adult females of the asexual generation emerged (from galls collected in Manitoba but reared in Edmonton) during the second half of October and early November. Adults of the sexual generation emerged in Edmonton from late May to early June. Galls of the asexual generation appeared on leaves in Edmonton in July.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFBACE0B4E9D56F4A8F4AE0F.taxon	distribution	Distribution. USA: New York, Michigan, Iowa, Kansas, Texas, Utah, Arizona (Burks 1979); Canada: Manitoba. Molecular taxonomy. Alternate sexual and asexual generations were proposed by Digweed (2010) and are herein matched using DNA data, with four individuals (two asexual females, two sexual females) sequenced for both cytb and ITS 2. Cytb haplotype sequences were identical among the four individuals (GenBank accessions MW 388885, OM 321613 – OM 321615), as were ITS 2 sequences (OM 331806 – OM 331809).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFBECE174E9D560CAD4FAC4F.taxon	description	Figs 29 – 41, 43 Synonyms: Amphibolips spinosa Ashmead (1887: 141), females and galls. Amphibolips spinosus corrected spelling by Dalla Torre & Kieffer (1910: 591).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFBECE174E9D560CAD4FAC4F.taxon	materials_examined	Material examined: 5 females labelled as “ USA, FL., Walton Co., Gratin Beach Island, Coast Scrub, coll. 3 May 1995; emerged 16 May 1995. G. Melika; ex Quercus myrtifolia ”. One male labelled as “ USA, FL., Highlands Co., Josephine Creek, leg. J. Nicholls, 2008.04.16. Code FL 1075; ex Quercus incana ”. 2 females have been deposited at the USNM, Washington, DC; 3 females and 1 male at the PHDNRL.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFBECE174E9D560CAD4FAC4F.taxon	diagnosis	Diagnosis. The sexual galls resemble those of A. quercuscitriformis (Ashmead, 1881), but differ from them in having much thicker and harder walls, the surface of the gall is rough, and they are appreciably narrower, and turn dirty brown when mature; in contrast the galls of A. quercuscitriformis are much more fragile at maturity, with thinner walls, the surface of the gall is smooth and glabrous, are much broader in diameter, remain lemon-yellow after adult emergence. Below the description of the sexual female and male are given according to the current morpho-description requirements.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFBECE174E9D560CAD4FAC4F.taxon	description	Description. Sexual female (Figs 29 – 33, 36 – 41). Head, mesosoma, metasoma, antennae, legs uniformly reddish brown. Head dull rugose, with sparse short white setae, denser on lower face, gena and postgena, quadrangular, 1.3 × as broad as high, narrower than mesosoma in frontal view; 1.9 x as broad as long in dorsal view. Gena dull rugose, not broadened behind eye in frontal view, narrower than transverse diameter of eye in lateral view. Malar space rugose, without striae. Eye 2.0 × as high as length of malar space. POL 2.0 × as long as OOL, OOL longer than LOL, lateral ocellus 2.2 × as long as LOL; ocelli ovate, of same size. Transfacial distance slightly longer than height of eye; diameter of antennal torulus 1.7 x as long as distance between toruli, and 1.4 x as long as distance between torulus and eye; lower face dull rugose, without striae and with narrow elevated rugose median area. Clypeus coriaceous, rectangular, broader than high; ventrally rounded, emarginate, without median incision; anterior tentorial pits small, indistinct, epistomal sulcus impressed, clypeo-pleurostomal line indistinct. Frons, vertex and occiput uniformly dull rugose. Interocellar area dull rugose, with large strongly elevated ocelli. Postocciput and postgena smooth, glabrous, impressed around occipital foramen; posterior tentorial pits large, deep, area around them strongly impressed; occipital foramen at least 4.0 × as high as height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which are not united. Antenna slightly longer than head + mesosoma, with 10 flagellomeres; scape 2.0 × as long as pedicel, pedicel subglobose, F 1 2.8 × as long as pedicel and 1.4 × as long as F 2; F 2 slightly longer than F 3, F 3 = F 4, subsequent flagellomeres shorter, F 10 2.0 × as long as F 9, placodeal sensilla on all flagellomeres. Mesosoma slightly longer than high in lateral view. Pronotum rugose dorsally, with numerous transversely orientated strong irregular rugae laterally which are absent in smooth, glabrous anterolateral part. Mesoscutum uniformly dull rugose, slightly longer than broad (greatest width measured across mesoscutum level with tegulae base). Notaulus indistinct in dull rugose sculpture; anterior parallel line extending to 1 / 3 length of mesoscutum, slightly impressed, marked by more delicate sculpture; parapsidal line distinct, originating away from posterior margin and extending to nearly half the length of mesoscutum, marked by elevated lines; median mesoscutal line absent; circumscutellar carina broad, extending to level of tegula. Mesoscutellum uniformly dull rugose, trapezoid, slightly longer than broad, without posteromedian depression, with median incision, overhanging metanotum. Mesoscutellar foveae large, deep, slightly longer than broad, with a few striae on smooth, glabrous bottom, with distinct elevated narrow median carina; lateral sides of foveae with strong carinae, separating them from dorsoaxillar areas. Mesopleuron, including speculum, uniformly dull rugose. Mesopleural triangle smooth, glabrous, with dense white setae; dorsal axillar area delicately rugose with dense setae; lateral axillar area coriaceous, with a few short, white setae; subaxillular bar smooth, glabrous, with parallel sides, its height less than height of metanotal trough, posterior end extending to half height of mesoscutellum; metapleural sulcus hidden in dull rugose sculpture. Metascutellum uniformly coriaceous, metanotal trough coriaceous, with dense white setae; ventral impressed area smooth, slightly shorter than height of metascutellum; central propodeal area lyre-shaped, smooth, glabrous, with strong median longitudinal carina; lateral propodeal carinae bent slightly outwards medially; lateral propodeal area with irregular strong rugae and dense white setae. Nucha short, with irregular sulci. All legs with dense short white setae; tarsal claws with acute basal lobe. Forewing longer than body, infuscate, margin with short dense cilia and dark brown rounded spot in radial cell; R 1 and Rs weakly pigmented, light, nearly reaching wing margin; radial cell narrow, open, slightly longer than broad; areolet small, triangular, closed, distinct; Rs + M reaching basalis (M) slightly above half its height. Metasoma longer than head + mesosoma, longer than high in lateral view; 2 nd metasomal tergum extending to 2 / 3 of metasoma length, smooth, glabrous, with short dense setae anterolaterally, with band of micropunctures posteriorly; all subsequent terga and hypopygium uniformly and entirely micropunctate. Ventral spine of hypopygium robust, long, needle-like, prominent part 6.5 × as long as broad, with two rows of white setae ventrally, extending beyond apex of spine. Body length 4.5 – 4.8 mm (n = 5). Male (Figs 34 – 35). Similar to female but head more rounded in frontal view, eye 4.2 × as high as length of malar space; POL 2.5 × as long as OOL, OOL longer than LOL; interocellar area strongly elevated in frontal view; antenna with 13 flagellomeres, pedicel as long as broad, F 1 the longest flagellomere, slightly curved and broadened in distal half and 1.4 × as long as F 2, F 2 = F 3 = F 4, subsequent flagellomeres shorter and nearly equal in length, placodeal sensilla on all flagellomeres. Body length 2.1 mm (n = 1). Gall (Fig. 43). Spindle-like bud galls, green and soft while young and growing, 20 – 25 mm in length, 5 – 7 mm in diameter. The surface is rough, not shiny, with pale spots, interior filled with dense radiating filaments that support the central larval cell. After the gall matures, it turns dirty brown, the walls are very hard and approximately 1 mm thick.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFBECE174E9D560CAD4FAC4F.taxon	biology_ecology	Biology. Alternate asexual and sexual generations are known. Asexual galls (Fig. 42) are ovate, about 10 mm long, in buds on Q. laurifolia and Q. myrtifolia. The sexual bud galls begin to develop in early May, mature in May-June, adults emerge in May through June. We recorded sexual galls on Q. myrtifolia, Q. laurifolia, Q. inopina and Q. incana.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFBECE174E9D560CAD4FAC4F.taxon	distribution	Distribution. USA: Florida, Highlands, Manatee, Martin, Walton, Liberty, Brevard Counties. We found this species also in North Carolina (Patsy Pond Area and Croatan National Forest, Carperet Co.). Molecular taxonomy. Alternating generations were matched using DNA data, with seven individuals (five asexual females, one sexual female, one sexual male) sequenced for cytb and six individuals (four asexual females, one sexual female, one sexual male) sequenced for ITS 2. Cytb sequences were on average 0.99 % divergent (range 0 – 1.85 %; GenBank accessions OM 321616 – OM 321622); ITS 2 sequences were all identical (OM 331810 – OM 331815).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFA2CE134E9D53F4AC91AC28.taxon	description	Figs 44 – 57 Synonyms: Callirhytis balanaspis Weld, 1922 b: 22, female, gall.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFA2CE134E9D53F4AC91AC28.taxon	materials_examined	Material examined: 5 females and 5 males “ USA, Florida, Archbold BioStation, leg. J. Nicholls, 2008.04.15. Code FL 1073, spFLc 1 a; ex Quercus inopina ”. Specimens have been deposited at the USNM and PHDNRL.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFA2CE134E9D53F4AC91AC28.taxon	diagnosis	Diagnosis. The sexual generation of A. balanaspis most closely resembles three species from Florida that also gall catkins, Callirhytis myrtifoliae (Beutenmueller, 1917) on Q. myrtifolia, Callirhytis quercusturnerii (Ashmead, 1881) on Q. nigra and Andricus cooki Melika, Nicholls & Stone, 2021 on Q. myrtifolia and Q. laurifolia. In C. myrtifoliae the clusters of catkin galls are smooth and shiny, without pubescence, while in A. balanaspis the catkin galls are covered with sparse white setae. In A. cooki, clusters of galls are dark green to purple with dense short pubescence, contain 10 - 20 individual galls tightly grouped together that cause shortening of the catkin axis; in A. balanaspis the galls are pale green with longer pubescence, and are scattered along a normal-length catkin. In A. cooki the head, antennae, mouthparts, mesosoma and legs reddish brown, metasoma brown, with darker brown areas, OOL 1.7 × as long as LOL, antenna with 12 flagellomeres, the mesopleuron and speculum uniformly reticulate, lateral propodeal carinae bent slightly outwards in posterior 1 / 3 of propodeum, the prominent part of the ventral spine of hypopygium 8.3 × as long as broad in ventral view. In the sexual generation of A. balanaspis the head, mesosoma, metasoma, antenna, mouthparts and legs are uniformly light brown to reddish, OOL 1.2 × as long as LOL, female antenna with 11 flagellomeres, the mesopleuron with delicate transverse subparallel striae and only the speculum is uniformly reticulate, lateral propodeal carinae bent slightly outwards at the mid-height of propodeum, the prominent part of the ventral spine of hypopygium short, slightly longer than broad in ventral view. In C. quercusturnerii the head of male entirely reddish brown; in females the mesoscutellar disk uniformly covered with strong rugae; the mesopleuron uniformly reticulate, the mesosoma and metasoma reddish brown. In the sexual generation of A. balanaspis the head of male is black; in females the center of the mesoscutellar disk is coriaceous, with strong irregular rugae laterally and posteriorly, the mesopleuron with delicate transverse subparallel striae, the mesosoma and metasoma light brown to reddish.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFA2CE134E9D53F4AC91AC28.taxon	description	Description. Sexual female (Figs 44 – 47, 51 – 55). Head, mesosoma, metasoma, antenna, mouthparts, legs uniformly light brown to reddish. Head alutaceous, with sparse white setae on lower face, 1.2 × as broad as high and slightly broader than mesosoma in frontal view, 2.0 × as broad as long in dorsal view. Gena alutaceous, not broadened behind eye in frontal view, narrower than transverse diameter of eye in lateral view. Malar space alutaceous, glabrous, with delicate striae radiating from clypeus and reaching eye, malar sulcus absent; eye 1.7 × as high as length of malar space. Eyes slightly converging ventrally. POL 1.6 × as long as OOL, OOL 2.3 × as long as diameter of lateral ocellus and 1.2 × as long as LOL, all ocelli ovate, of same size. Transfacial distance 1.7 × as long as height of eye; diameter of antennal torulus as great as distance between them, distance between torulus and eye 1.3 × as long as diameter of torulus; lower face alutaceous, glabrous, with white setae, without striae radiating from clypeus; slightly elevated median area alutaceous, glabrous, without setae. Clypeus rectangular, nearly 2.0 x as broad as high, smooth, glabrous, with long setae; ventrally rounded, not emarginate and without median incision; anterior tentorial pit large, rounded, indistinct, epistomal sulcus distinct, clypeo-pleurostomal line inconspicuous. Frons uniformly reticulate, without striae or setae, interocellar area reticulate. Vertex and occiput reticulate, with sparse white setae; postocciput smooth, glabrous; postgena mainly reticulate, smooth and glabrous along occipital foramen, postgenal bridge and hypostomata; posterior tentorial pit large, ovate, area below impressed; occipital foramen higher than height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which diverge strongly toward occipital foramen, postgenal bridge anteriorly as broad as occipital foramen. Antenna longer than head + mesosoma, with 11 flagellomeres, pedicel slightly longer than broad, flagellomeres slightly broader towards apex; F 1 2.0 × as long as pedicel and slightly longer than F 2; F 2 1.2 × as long as F 3; F 3 = F 4, F 5 = F 6 = F 7, subsequent flagellomeres nearly equal in length; placodeal sensilla indistinct on all flagellomeres. Mesosoma slightly longer than high, with sparse white setae, denser on lateral propodeal area. Pronotum reticulate, with sparse dense setae, with some delicate striae anteroventrally; propleuron smooth, glabrous. Mesoscutum uniformly reticulate, with sparse white setae, slightly longer than broad (greatest width measured across mesoscutum level with base of tegulae). Notaulus complete, deep, broad, bottom smooth, posteriorly broader and slightly converging; anterior parallel line indistinct, marked with smooth line; parapsidal line distinct, marked with smooth line; median mesoscutal line absent; circumscutellar carina broad, reaching notaulus. Mesoscutellum trapezoid, slightly longer than broad, glabrous, with net of strong irregular rugae, posteriorly rounded, overhanging metanotum. Mesoscutellar foveae separated by narrow elevated coriaceous central carina, ovate, with smooth, glabrous bottom. Mesopleuron with delicate transverse subparallel striae, speculum uniformly reticulate; mesopleural triangle smooth, glabrous, with some delicate striae and sparse white setae; dorsal and lateral axillar areas delicately coriaceous, with a few white setae; subaxillular bar smooth, glabrous, with parallel sides, as high as height of metanotal trough, slightly higher at posterior end; metapleural sulcus reaching mesopleuron slightly above half of its height, delimiting coriaceous area, upper part of sulcus distinct. Metascutellum smooth, glabrous, slightly shorter than height of smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous, without setae; central propodeal area smooth, glabrous, with a few irregular rugae; lateral propodeal carinae bent slightly outwards at midheight; lateral propodeal area smooth, glabrous, with dense white setae. Nucha with numerous sulci dorsally and laterally. Tarsal claws simple, without basal lobes. Forewing longer than body, hyaline, without cilia on margin, veins light brown, hardly visible, radial cell open, 2.2 × as long as broad; R 1 and Rs reaching wing margin; areolet invisible. Rs + M narrow, inconspicuous, its projection reaching basalis at its mid height. Metasoma as long as head + mesosoma, longer than high in lateral view; 2 nd metasomal tergum extending to 2 / 3 length of metasoma in dorsal view, without setae and micropunctures; all terga smooth, glabrous, without micropunctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium short, slightly longer than broad in ventral view, with a few short setae ventrally which do not extend beyond apex of spine. Body length 1.8 – 2.0 mm (n = 5). Male (Figs 48 – 50). Similar to female, but head dark brown to black, with lighter lower face and malar space; mesosoma and metasoma dark brown to black; eye 2.5 × as high as length of malar space; POL 1.7 × as long as OOL, OOL equal LOL and 1.6 × as long as diameter of lateral ocellus, ocelli ovate, larger than in female; interocellar area elevated in frontal view; antenna with 13 flagellomeres, F 1 slightly curved, broadened in apical 2 / 3, F 1 1.3 × as long as F 2, F 13 equal F 12; placodeal sensilla on F 2 – F 13, absent on F 1. Body length 1.7 – 1.9 mm (n = 5). Gall (Figs 56 – 57). Sexual galls are aggregated on catkins, each rounded, to 3 mm in diameter, pale green with white pubescence. In sparse clusters, rarely singly, along axis of catkin; sometimes causing the catkin to curl but never stunting its development.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFA2CE134E9D53F4AC91AC28.taxon	biology_ecology	Biology. Alternate asexual and sexual generations are known, inducing galls on red oaks in (Quercus section Lobatae). The asexual generation galls (Figs 58 – 60) have been found on acorns of Q. marilandica and Q. myrtifolia, mature in October, adults emerge in spring of the following year. To date, sexual catkin galls have only been recognised on Q. inopina and Q. myrtifolia, mature in April, adults emerge soon afterwards.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFA2CE134E9D53F4AC91AC28.taxon	distribution	Distribution. USA: Washington DC, Missouri, Arkansas, Texas, Virginia, Florida. Molecular taxonomy. Alternating generations were matched using DNA data, with eight individuals (two asexual females, three sexual females, three sexual males) sequenced for cytb and two individuals (one asexual female, one sexual female) sequenced for ITS 2. Cytb sequences were on average 0.28 % divergent (range 0 – 0.69 %; GenBank accessions KX 683615, OM 321627 – OM 321633); ITS 2 sequences were identical except for a single base indel within a polyT repeat region (OM 331821 – OM 331822). The authors’ unpublished sequence data show that A. balanaspis is part of a clade of species galling red oaks whose sexual generations, where known, are on catkins and asexual generations, where known, are acorn galls. This clade includes species such as Callirhytis carmelensis Weld, 1922, A. cooki, C. myrtifoliae, A. fitpatricki Melika & Abrahamson, 2021 and C. balanacea Weld, 1944.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFA2CE134E9D53F4AC91AC28.taxon	discussion	Comments. Detailed descriptions of the asexual generation females, galls and biology is given in Weld (1922 b).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFA6CE1F4E9D53DCA90DAE98.taxon	description	Figs 61 – 78, 80 – 81 Synonyms: Cynips quercusnubila Bassett (1881 b: 56), females and galls. Dryophanta nubila combination and corrected spelling by Mayr (1881); Dryophanta nubila combination by Ashmead (1885: 296); Diplolepis quercus-nubila combination by Dalla Torre and Kieffer (1910); Cynips nubila combination by Cresson (1923); Diplolepis nubila combination by Weld (1926); Cynips (Acraspis) nubila var. nubila combination by Kinsey (1930); Antron nubila combination by Weld (1951); Antron quercusnubila corrected spelling by Burks (1979). Kinsey (1930, 1936, 1938) included 10 additional species in his “ nubila ” complex, found on a range of Mexican oaks: Cynips (Acraspis) chica Kinsey 1936; Cynips (Acraspis) incompta (Kinsey, 1920); Cynips (Acraspis) nigricula Kinsey, 1936; Cynips (Acraspis) radialis Kinsey, 1936; Cynips (Acraspis) rufula Kinsey, 1936; Cynips (Acraspis) russa Kinsey, 1930; Cynips (Acraspis) subtincta Kinsey, 1936; Cynips (Acraspis) tincta Kinsey, 1936, Cynips (nubila) lanaris Kinsey, 1938 and Cynips (nubila) molucrum Kinsey, 1938.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFA6CE1F4E9D53DCA90DAE98.taxon	materials_examined	Material examined: 1 female and 2 males “ USA, Arizona, Molino Basin campground, Santa Catalina Mtns. leg. J. Nicholls, 2008.04.09. Code AZ 1875, spAZb 9; ex Quercus oblongifolia ”. Specimens have been deposited at the USNM.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFA6CE1F4E9D53DCA90DAE98.taxon	diagnosis	Diagnosis. Two species of Antron Kinsey, 1930, A. douglasii (Ashmead, 1896) and A. quercusechinus (Osten Sacken, 1870), are known from a sexual generation in California; both induce succulent green bud galls on Q. douglasii, Q. dumosa, and Q. lobata. One other species, Antron pileus Weld, 1952, is known from Arizona from its sexual generation; this species induces bud galls on Q. turbinella (Weld 1952 b). Galls of A. pileus are mushroomlike and without hairs while the bud galls of A. quercusnubila are oblong with long hairs. In A. pileus the female head is coriaceous, antenna with 13 flagellomeres, the disk of mesoscutellum finely rugose, and the male antenna has 14 flagellomeres while in A. quercusnubila, the sexual female head is alutaceous, antenna with 12 flagellomeres, the disk of mesoscutellum uniformly smooth, glabrous, with net of strong irregular rugae and the male antenna has 13 flagellomeres.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFA6CE1F4E9D53DCA90DAE98.taxon	description	Description. Sexual female (Figs 61 – 65, 70 – 72, 76 – 77). Head, mesosoma, metasoma, maxillary and labial palpi, legs uniformly yellow; antenna darker than head. Head alutaceous, with sparse white setae, 1.3 × as broad as high and slightly broader than mesosoma in frontal view, 2.2 × as broad as long in dorsal view. Gena alutaceous, not broadened behind eye, invisible in frontal view, narrower than transverse diameter of eye in lateral view. Malar space smooth, glabrous, without striae; eye 4.1 × as high as length of malar space. Inner margins of eyes parallel. POL 3.9 × as long as OOL, OOL slightly shorter than diameter of lateral ocellus, 2.0 × shorter than LOL, all ocelli ovate, of same size. Transfacial distance slightly shorter than height of eye, diameter of antennal torulus 2.0 × as long as distance between them, distance between torulus and eye slightly shorter than diameter of torulus; lower face smooth, glabrous, with a few short white setae, without striae; slightly elevated median area smooth, glabrous, without striae, with some short setae; small area under torulus, between toruli and area between torulus and eye delicately alutaceous. Clypeus trapezoid, slightly broader than high, smooth, glabrous; ventrally rounded, not emarginate, without median incision, with a few long setae; anterior tentorial pit large, deep, rounded, epistomal sulcus and clypeo-pleurostomal line distinct. Frons and slightly elevated interocellar area alutaceous, with short white setae. Vertex alutaceous; occiput, postocciput, postgena smooth, glabrous, with a few setae; posterior tentorial pit large, elongated, area below impressed; occipital foramen as high as height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which do not unite, diverging until postgenal bridge which is as broad as occipital foramen. Antenna longer than head + mesosoma, with 12 flagellomeres, scape, pedicel and F 1 lighter than rest of flagellomeres, pedicel subglobular; F 1 2.7 × as long as pedicel and 1.3 x as long as F 2, F 2 = F 3 = F 4, F 4 slightly longer than F 5; F 7 to F 11 equal in length; placodeal sensilla on F 5 – F 12, absent on F 1 – F 4. Mesosoma distinctly longer than high, without setae. Pronotum smooth, glabrous, without striae along posterior margin, with a few setae; invaginated anterior margin of pronotum smooth, glabrous. Propleuron smooth, glabrous, without setae. Mesoscutum smooth, longer than broad (greatest width measured across mesoscutum level with base of tegulae). Notaulus complete, deep, with smooth, glabrous bottom, converging posteriorly; anterior parallel line invisible; parapsidal line and median mesoscutal line absent; circumscutellar carina narrow, reaching notaulus. Mesoscutellum slightly longer than broad, with nearly parallel sides, uniformly smooth, glabrous, with net of strong irregular rugae, overhanging metanotum. Mesoscutellar foveae in the form of a transverse anterior impression, not separated medially, with smooth, glabrous bottom. Mesopleuron smooth, glabrous, without setae; mesopleural triangle smooth, glabrous, with white setae, without striae; dorsal and lateral axillar areas smooth, glabrous, with white setae; subaxillular bar smooth, glabrous, narrow, at posterior end higher than height of metanotal trough; metapleural sulcus delimiting very narrow area, reaching mesopleuron at its mid height, upper part of sulcus distinct. Metascutellum smooth, glabrous, slightly higher than smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous, without setae; central propodeal area smooth, glabrous, without setae, with a few delicate short striae in anterocentral part; lateral propodeal carinae nearly parallel, bent outwards at most posterior end; lateral propodeal area smooth, glabrous, with long setae. Nucha without sulci. Tarsal claws with distinct basal lobe. Forewing longer than body, hyaline, with dark brown veins, margin without cilia; radial cell open, 2.0 × as long as broad, R 1 and Rs nearly reaching wing margin, areolet triangular, well-delimited, Rs + M distinct along entire length, reaching basalis in lower 1 / 3. Metasoma slightly longer than head + mesosoma, slightly longer than high in lateral view; 2 nd metasomal tergum extending to 2 / 3 of metasoma length in dorsal view, without setae anterolaterally, without micropunctures; subsequent terga smooth, glabrous, without micropunctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium as broad as long in ventral view, with long setae that extend far beyond apex of spine and form a tuft. Body length 2.6 – 2.9 mm (n = 4). Male (Figs 66 – 69, 73 – 75, 78). Similar to female but head, mesosoma, metasoma dark brown to black, legs yellow. Head slightly broader than high in frontal view; malar space shorter; eyes larger, ocelli larger, ovate, POL 4.6 × as long as OOL, OOL 2.0 × shorter than length of ocellus and shorter than LOL; interocellar area elevated. Antenna with 13 flagellomeres, F 1 straight; placodeal sensilla on all flagellomeres. Gall (Figs 80 – 81). The sexual generation develops in a small oblong bud gall, 5 mm long, 3 mm in diameter, pale green with white or pink pubescence, hairs several mm long. The gall develops on lenticel buds on stems or on the bark of larger branches.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFA6CE1F4E9D53DCA90DAE98.taxon	biology_ecology	Biology. Asexual galls (Fig. 79) develop on leaves, sexual galls in lenticel buds; both generations on Q. arizonica, Q. rugosa and Q. oblongifolia.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFA6CE1F4E9D53DCA90DAE98.taxon	distribution	Distribution. USA: Arizona (Burks 1979). Molecular taxonomy. Alternating generations were matched using DNA data, with four individuals (two asexual females, two sexual males) sequenced for both cytb and ITS 2. Two cytb haplotypes were found across the four individuals, differing by 0.92 % (GenBank accessions OM 321623 – OM 321626); two ITS 2 alleles were also observed, differing by 0.19 % (with a single base indel within a polyT repeat region; OM 331816 – OM 331819).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFAACE254E9D552DAD5EADE0.taxon	description	Figs 82 – 93 Synonyms: Neuroterus flavipes Gillette 1889: 281. Male, female and gall. Description reprinted by Gillette (1890: 21). Beutenmüller (1910: 136) suggested that this species should probably be placed in Callirhytis or Andricus rather than Neuroterus. Kinsey (1923: 136) agreed, and left this species unplaced in his revision of Neuroterus. Callirhytis flavipes (Gillette): combination by Weld (1926: 91). Melika & Abrahamson (2002: 168) considered this species should belong to Andricus, however, this transfer was made only informally, without following the ICZN requirements.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFAACE254E9D552DAD5EADE0.taxon	materials_examined	Material examined: Fifteen females “ Canada, Alberta, University of Alberta, Edmonton, leg. S. Digweed, 27 / 04 / 2007, Q. macrocarpa ”.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFAACE254E9D552DAD5EADE0.taxon	diagnosis	Diagnosis. This species possesses the diagnostic characters of Bassettia Ashmead, 1887: absence of a malar sulcus; the transversely rugose mesoscutum, dorsoventrally compressed mesosoma not arched in lateral view like in most Cynipini; subparallel lateral propodeal carinae; and a ring of dense white setae basally on 2 nd metasomal tergum (Melika & Abrahamson 2007; Medianero & Nieves-Aldrey 2010). Bassettia flavipes also forms inconspicuous twig galls in the asexual generation and integral leaf galls in the sexual generation, as in other Bassettia species (Melika & Abrahamson 2007; Medianero & Nieves-Aldrey 2010). Bassettia flavipes most closely resembles B. tenuana Weld, 1921. In B. flavipes the body is dark brown, the eye 2.0 × as high as the length of the malar space, the notaulus incomplete, traceable at most along half the length of the mesoscutum, the forewing veins are light to transparent, the central propodeal area with numerous irregular short rugae, the metasoma as long as head + mesosoma and nearly as long as high in lateral view, 2 nd metasomal tergum extending to 1 / 3 of metasoma length in dorsal view. In B. tenuana the body is black, the eye equal in height to the length of malar space, the notaulus complete, the forewing veins dark brown, the central propodeal area reticulate, the metasoma longer than head + mesosoma and distinctly longer than high in lateral view, 2 nd metasomal tergum extending to 2 / 3 of metasoma length in dorsal view. Bassettia flavipes is also similar to B. gemmae Ashmead, 1896; both species have been recorded from Q. alba (Weld 1926). In B. flavipes the head and body dark brown, the mesoscutum uniformly alutaceous, the prominent part of the ventral spine of hypopygium more than 2.0 × as long as broad in ventral view. In B. gemmae the head and body black, the mesoscutum transversely dull rugose, the prominent part of the ventral spine of hypopygium over 4.0 × as long as broad in ventral view. In addition, the length-to-width ratio of the radial cell of B. gemmae is greater than that of B. flavipes. It must be also mentioned that the asexual and sexual females within Bassettia are very similar and only a few characters differentiate them.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFAACE254E9D552DAD5EADE0.taxon	description	Description. Asexual female (Figs 82 – 92) Head, mesosoma, metasoma black; mouthparts, lower genal and clypeal margins, antennae, tegulae, coxal tips, bases and tips of femora and tibiae, tarsi apices, and ventral spine of hypopygium, all dark to light brown. Wing venation pale yellow or colourless. Head alutaceous without setae; rounded, 1.3 × as broad as high and as broad as mesosoma in frontal view, 2.2 × as broad as long in dorsal view. Gena alutaceous, broadened behind eye in frontal view, as broad as transverse diameter of eye in lateral view. Malar space alutaceous, with very delicate, indistinct striae radiating from clypeus and reaching eye, eye 2.0 × as high as height of malar space. Eyes slightly converging ventrally. POL 1.9 × as long as OOL; OOL 2.0 × as long as diameter of lateral ocellus and as long as LOL; lateral ocelli rounded, median ocellus ovate. Transfacial distance 1.3 × as long as height of eye; diameter of antennal torulus 2.0 × as long as distance between them, distance between torulus and eye as great as diameter of torulus. Antennal toruli level with the lower half of eye. Lower face alutaceous, with sparse short white setae, as high as frons, with short delicate striae radiating from clypeus and reaching eye, striae only alongside median part of lower face; sculpture of lower face rougher than frons; median area alutaceous, not elevated; lower face with two impressed smooth glabrous sulci starting from lateral edges of clypeus and extending to 1 / 3 of lower face height. Clypeus rectangular, 1.6 × as high as broad, uniformly alutaceous, ventrally rounded, emarginate, without median incision, reaches beyond ventral edge of lower face; anterior tentorial pit small, rounded, inconspicuous; smooth, glabrous epistomal sulcus and smooth, glabrous clypeo-pleurostomal line distinct, strongly impressed. Frons and interocellar area uniformly alutaceous, without setae. Vertex, occiput, postocciput, postgena uniformly alutaceous, without parallel striae, with white setae; posterior tentorial pit large, ovate, area below impressed; occipital foramen slightly higher than height of postgenal bridge; hypostomal carina emarginate, continuing into united postgenal sulci which diverge toward occipital foramen, postgenal bridge anteriorly broader than high. Antenna longer than head + mesosoma; with 12 flagellomeres (suture between F 12 and F 11 indistinct), scape and pedicel 2.0 × as broad as F 1, scape and pedicel shorter than F 1, F 1 1.4 × as long as pedicel and 1.4 × as long as F 2, F 2 longer than F 3, F 3 = F 4, F 5 = F 6 and shorter than previous one, F 7 to F 11 nearly equal in length, F 11 longer than F 10; placodeal sensilla on F 3 – F 12. Mesosoma slightly longer than high. Pronotum alutaceous, with white setae along margin, laterally broader than width of mesopleuron, anterolateral edge impressed, smooth, without sculpture. Propleuron alutaceous, with sparse setae. Mesoscutum uniformly alutaceous; slightly longer than broad (greatest width measured across mesoscutum level with base of tegulae). Notaulus incomplete, very weakly impressed, traceable at most to half the length of mesoscutum; anterior parallel line and median mesoscutal sulcus absent; parapsidal line in the form of a smooth stripe, extending slightly above tegula; circumscutellar carina broad, smooth along tegula. Mesoscutellum ovate, slightly longer than broad; posteriorly rounded and emarginate; uniformly rugoso-reticulate, overhanging metanotum. Mesoscutellum with narrow short transverse semilunar impression anteriorly, with smooth, glabrous bottom, with central broad triangular elevated area. Mesopleuron with delicate transverse striae starting posteroventrally and extending across mesopleuron and widening towards anterodorsal end, anteroventral quarter of mesopleuron smooth, shiny; speculum shiny, with very delicate, hardly traceable striae; mesopleural triangle rugose, with white setae; dorsal and lateral axillar areas alutaceous, without setae; subaxillular bar smooth, glabrous, at posterior end as high as height of metanotal trough; metapleural sulcus reaching mesopleuron at half its height, delimiting smooth, shiny triangular area along mesopleuron; upper part of mesopleural sulcus indistinct. Metascutellum rugose, as high as height of smooth, glabrous ventral impressed area; metanotal trough alutaceous, glabrous, without setae; propodeum uniformly alutaceous, lateral propodeal carinae strong, slightly bent outwards in posterior 1 / 3 of propodeum height, delimiting smooth, shiny central propodeal area with numerous irregular short rugae; lateral propodeal area smooth, shiny with dense white setae. Nucha very short, with a few logitudinal sulci laterally and dorsally. Tarsal claws simple, without basal lobe. Forewing longer than body, hyaline, with long dense cilia on margin, veins indistinct, pale, radial cell open, 4.2 × as long as broad; R 1 and Rs nearly reaching wing margin; areolet and Rs + M indistinct. Metasoma as long as head + mesosoma, nearly as long as high in lateral view; 2 nd metasomal tergum extending to 1 / 3 of metasoma length in dorsal view, with dense white setae anterolaterally; all terga smooth, glabrous, without micropunctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium 2.1 × as long as broad in ventral view, with a few short setae ventrally. Body length 1.4 – 2.1 mm long (n = 15). Gall. Asexual galls (Fig. 93) are inconspicuous stem galls comprised of individual larval cells within corky bark on 2 – 3 year old twigs of Q. macrocarpa. Galls are not evident as swellings on twig surfaces. Galls are not induced on twigs of the previous year’s growth, nor are they present in smooth twigs.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFAACE254E9D552DAD5EADE0.taxon	biology_ecology	Biology. Asexual females of B. flavipes emerged in mid- to late April in Edmonton and oviposited into swelling buds, preferring those having the bud scales slightly open with green showing between them. Females actively palpated prospective buds, and briefly probed them by inserting their ovipositors between bud scales. Females then quickly plunged their ovipositors fully into the bud, oviposited for 10 - 20 seconds, retracted their ovipositors, and departed. Females mostly seemed to avoid buds into which they or other females had previously oviposited. Sexual generation galls are multilocular, fleshy swellings along the leaf midrib or major lateral veins (Fig. 94) and are visible starting in mid- to late May in Edmonton. Adults emerged starting in late June or early July, and males tended to begin emerging earlier than females. Each emerging adult appeared to cut its own exit hole from the gall. Adult females of the sexual generation of B. flavipes oviposited into twig bark. We did not assess whether unmated females oviposited or produced viable eggs. Sexual generation galls have been recorded from Q. macrocarpa and Q. alba (Burks 1979, Weld 1926), and the authors additionally observed them on Q. × bebbiana (Q. alba x Q. macrocarpa) and Q. × schuettei (Q. bicolor x Q. macrocarpa) at the Jardin Botanique de Montreal in August 2007.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFAACE254E9D552DAD5EADE0.taxon	distribution	Distribution. USA: Illinois, Indiana, Iowa, Minnesota, North Dakota, Virginia; Canada: Alberta, Ontario (Weld 1926, Burks 1979, Ives & Wong 1988, Bergdahl 2015). Within Canada, the authors have observed galls of the sexual generation on introduced bur oaks in Alberta, and on bur oaks within their native range in Manitoba and Quebec. Molecular taxonomy. This species was first described from adults of the sexual generation and galls on Q. macrocarpa (Gillette 1889); Ives & Wong (1988) tentatively identified adults and galls of the asexual generation of B. flavipes but did not provide any confirmatory evidence. Herein, alternating generations are matched for the first time using DNA data, with four individuals (two asexual females, two sexual females) sequenced for both cytb and ITS 2. Cytb sequences were on average 0.54 % divergent (range 0 – 0.92 %; GenBank accessions MN 184628 – MN 184631). Six ITS 2 sequences were observed across the four individuals (two individuals being heterozygous; MN 184632 – MN 184637); those sequences were 0 – 0.60 % divergent with three indels observed across all sequences. The B. flavipes cytb haplotypes were on average 8.24 % divergent from the only other published Bassettia cytb sequence, obtained from B. pallida (GenBank accession MG 821072; Nicholls et al. 2018 b). In addition, ITS 2 sequences among the two species were about 6.64 % different, with 21 indels (GenBank accession OM 331820 for B. pallida). These divergences are consistent with those expected for congeneric species, confirming our generic-level re-classification of this species.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFAACE254E9D552DAD5EADE0.taxon	discussion	Comments. Suspected galls of the asexual generation of B. flavipes have been identified as a source of damage on ornamental bur oaks because, at high population levels, they may attract birds which damage twigs while extracting larvae from galls (Ives & Wong 1988; Bergdahl 2015). Photos of this damage suggest that galls containing larvae may occur on a greater variety of stem sizes and maturities than we observed, perhaps as a function of higher population densities. It is also possible that old galls bearing exit holes, which persist on larger and older stems, house other insects that are sought by birds. Rearings of asexual galls of B. flavipes from Edmonton, Alberta produced multiple other gall occupants. The most common (88 %) alternative occupant was a species of Ceroptres Hartig, 1840; this genus is an inquiline in cynipid galls, although it is not clear if it is lethal to the original gall inducer (Ronquist et al. 2015). It appeared to specialize on B. flavipes in Edmonton, as it was not reared from other bur-oak-galling cynipids there (S. Digweed, unpublished data). We also reared the chalcid parasitoids Ormyrus labotus Walker, 1843, and Sycophila marylandica (Girault, 1916), from asexual galls. All parasitoids emerged from mid-May to mid-June. We also reared these three species from sexual galls of B. flavipes, as well as adults of the parasitoids Baryscapus racemariae (Ashmead, 1886), Sycophila dubia (Walsh, 1870), and Eurytoma studiosa Say, 1836. These alternative occupants of sexual B. flavipes galls emerged between early July and mid-August. Sycophila foliatae (Ashmead, 1881), S. quercilanae (Fitch, 1859), and S. xanthochroa (Ashmead, 1894), have been reared elsewhere from sexual galls of B. flavipes (Noyes 2016).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF90CE214E9D5314AF1DAD0B.taxon	description	Figs 95 – 111, 113 Synonyms: Bassettia archboldi Melika & Abrahamson, 2007, syn. nov. This species was initially described based only on males, without knowledge of the females or the galls they induce; a male was obtained in a Malaise trap in November in Florida, at the Archbold Biological Station (Melika & Abrahamson 2007).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF90CE214E9D5314AF1DAD0B.taxon	materials_examined	Material examined: Two females and one male “ Skinners nursery, Crescent City., FL, gall coll. 04 / 03 / 24. Tree 20 C, Bassettia gall. ”. Four males “ USA, FL, Highlands Co., Lake Placid, Archbold Bio Sta, 1995.10.11. MT. leg. G. Melika ” (in the collection of the Archbold Biological Station, Highlands Co., Florida). Two males have been deposited at the USNM, two female and three males at the PHDNRL.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF90CE214E9D5314AF1DAD0B.taxon	diagnosis	Diagnosis. Both females and males most closely resemble B. ligni Kinsey, 1922. In B. pallida the mesoscutum is longer than broad in dorsal view, the mesoscutellar foveae in the form of a narrow transverse semi-lunar impression, without a median carina, while in B. ligni the mesoscutum nearly as long as broad, mesoscutellar foveae broad, separated by a distinct median carina. For species identification the key to Bassettia can also be used (Melika & Abrahamson 2007).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF90CE214E9D5314AF1DAD0B.taxon	description	Description. Sexual female (Figs 95 – 98, 103 – 104, 109 – 111). Head and mesosoma black, antenna brown, metasoma dark brown; legs light brown with dark brown coxae, trochanter and tibia. Head alutaceous without setae; rounded, 1.2 × as broad as high and broader than mesosoma in frontal view, 1.9 × as broad as long in dorsal view. Gena alutaceous, slightly broadened behind eye in frontal view, narrower than transverse diameter of eye in lateral view. Malar space alutaceous, without sulcus or striae; eye 4.8 × as high as height of malar space. Eyes slightly converging ventrally. POL 5.4 × as long as OOL; OOL nearly 2.0 × as long as diameter of lateral ocellus and 2.0 × as long as LOL; all ocelli ovate, of same size. Transfacial distance 1.3 × as long as height of eye; diameter of antennal torulus 2.0 × as long as distance between them, distance between torulus and eye 1.2 × as long as diameter of torulus. Lower face alutaceous, 2.0 x as long as frons in dorsal view, without setae or striae; sculpture of lower face rougher than frons; median area alutaceous, not elevated; lower face with two impressed smooth glabrous sulci starting from lateral edges of clypeus and extending to 1 / 3 of lower face height. Clypeus rectangular, 1.4 × as high as broad, uniformly alutaceous, ventrally emarginate, without median incision, reaches beyond ventral edge of lower face; anterior tentorial pit small, rounded, inconspicuous; smooth, glabrous epistomal sulcus and smooth, glabrous clypeo-pleurostomal line distinct, strongly impressed. Frons and interocellar area uniformly alutaceous, without setae. Vertex, occiput, postocciput, postgena uniformly alutaceous, without parallel striae, with white setae; posterior tentorial pit large, ovate, area below impressed; occipital foramen 1.6 × as long as height of postgenal bridge; hypostomal carina emarginate, continuing into united postgenal sulci which strongly diverge toward occipital foramen, postgenal bridge anteriorly broader than high. Antenna longer than head + mesosoma; with 11 flagellomeres; scape, pedicel, F 1 and F 2 much lighter than F 3 – F 11; pedicel slightly longer than broad, F 1 1.7 × as long as pedicel and 1.4 × as long as F 2, F 2 to F 6 nearly equal in length, subsequent flagellomeres slightly shorter but equal in length; placodeal sensilla on F 2 – F 11. Mesosoma 1.5 × as long as high. Pronotum alutaceous, with white setae along margin, laterally broader than width of mesopleuron. Propleuron alutaceous. Mesoscutum alutaceous, with sparse white setae lateral of parapsidal line; slightly longer than broad (greatest width measured across mesoscutum level with base of tegulae). Notaulus, anterior parallel line, median mesoscutal sulcus absent; parapsidal line in the form of a smooth stripe; circumscutellar carina broad, smooth along tegula. Mesoscutellum ovate, slightly longer than broad; posteriorly rounded; anterior 2 / 3 with delicate net of rugae; posteriorly alutaceous, without setae, not overhanging metanotum. Mesoscutellum with narrow short transverse semilunar impression anteriorly, with smooth, glabrous bottom. Mesopleuron alutaceous, with transverse invagination in ventral part; mesopleural triangle alutaceous, without striae or setae; dorsal and lateral axillar areas alutaceous, without setae; subaxillular bar smooth, glabrous, at posterior end shorter than height of metanotal trough; metapleural sulcus reaching mesopleuron below half its height, delimiting broad alutaceous area along mesopleuron; upper part of mesopleural sulcus absent, area above sulcus uniformly alutaceous. Metascutellum smooth, slightly higher than height of smooth, glabrous ventral impressed area; metanotal trough alutaceous, glabrous, without setae; propodeum uniformly alutaceous, without propodeal carinae, without rugae or setae. Nucha very short, smooth. Tarsal claws simple, without basal lobe. Forewing longer than body, hyaline, with dense long cilia on margin, veins brown, radial cell open, 4.8 × as long as broad; R 1 and Rs nearly reaching wing margin; areolet small, triangular, distinct, Rs + M distinct along 2 / 3 of its length, projection reaching basalis slightly below its mid height. Metasoma as long as head + mesosoma, nearly as long as high in lateral view; 2 nd metasomal tergum extending to 1 / 3 of metasoma length in dorsal view, with very few setae scattered anterolaterally; all terga smooth, glabrous, without micropunctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium short, as long as broad in ventral view, without setae ventrally. Body length 2.7 – 2.8 mm (n = 2). Male (Figs 99 – 102, 105 – 108). Similar to female, body uniformly brown, lighter than female, legs uniformly light brown. Head 1.8 × as broad as long in dorsal view, ocelli larger than in female, POL 2.4 × as long as OOL, OOL slightly shorter than diameter of ocellus and equal to LOL. Antenna slightly longer than body, light brown, with last four flagellomeres darker, with 14 flagellomeres. F 1 equal to scape + pedicel, 1.5 × as long as F 2, F 2 slightly longer than F 3, F 3 slightly longer than F 4, F 4 = F 5 = F 6, subsequent flagellomeres nearly equal in length, F 14 slightly shorter than F 13, placodeal sensilla on F 4 – F 14, absent on F 1 – F 3. Metasoma nearly 2.0 × as long as head + mesosoma, 2 nd metasomal tergum extending to half the length of metasoma in dorsal view; all terga smooth, glabrous, without punctures. Gall. Asexual galls (Fig. 112) are in twigs, larval cells develop as cryptic elliptical chambers 3.2 mm long by 1.2 mm in diameter under the bark of small twigs. Sexual galls remain to be confirmed but may well be found within hypertrophied buds that occur in clusters on stems, with bud scales open, although with no obvious cavity observed within the bud (Fig. 113).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF90CE214E9D5314AF1DAD0B.taxon	biology_ecology	Biology. Bassettia pallida, previously known only from asexual females, is now also known from its sexual generation. The asexual galls develop under the bark of twigs on Q. virginiana and Q. geminata. Galls mature in late September-October; adults emerge in November. Both sexual females and males were reared from galls within the same twig with enlarged and distorted buds on it, collected at the Archbold Biological Station (Florida) from Q. geminata in October. See Weinersmith et al. (2020) for the community ecology and natural enemy assemblage associated with galls.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF90CE214E9D5314AF1DAD0B.taxon	distribution	Distribution. USA: Florida, Georgia, Missouri (Burks 1979).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF97CE2C4E9D50C0A849AB6F.taxon	description	Figs 114 – 118, 120 Synonyms: Cynips tumifica Osten Sacken (1865: 356), female, male, gall. Andricus (Callirhytis) tumifica combination by Mayr (1881); Andricus (Callirhytis) tumificus corrected spelling by Dalla Torre (1893); Callirhytis tumifica combination by Dalla Torre & Kieffer (1910); Melikaiella tumifica combination by Pujade-Villar et al. (2014).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF97CE2C4E9D50C0A849AB6F.taxon	materials_examined	Material examined: 15 asexual females labelled as “ Germany, Baden-Württemberg, Schwetzinger Hardt, 10 km SW of Heidelberg, ex acorns on Q. rubra, coll. R. Beiderbeck, 2014.10.01, emerge 2017.03.22 ”. Three asexual females have been deposited at the USNM, 12 females at the PHDNRL.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF97CE2C4E9D50C0A849AB6F.taxon	diagnosis	Diagnosis. Three Melikaiella species from eastern North America are known only from their asexual generations: M. fructuosa (Weld, 1922), M. fructicola (Ashmead, 1896), and M. corrugis (Bassett, 1881), all of which produce galls in acorns of red oaks (Pujade-Villar et al. 2014). In M. fructuosa the female antenna has 10 flagellomeres and is shorter than in M. tumifica, reaching at most the level of the tegulae. Melikaiella tumifica adults most closely resemble those of M. fructicola and M. corrugis. However, in M. fructicola and M. corrugis the mesoscutum is uniformly dull rugose in between and on either side of the notauli, mesoscutellar foveae without subparallel distinct longitudinal rugae while in asexual females of M. tumifica the mesoscutum is dull rugose between the notauli and delicately reticulate on either side of the notauli, mesoscutellar foveae distinctly divided by a central elevated carina and the bottom of foveae has strong longitudinal rugae.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF97CE2C4E9D50C0A849AB6F.taxon	description	Description. Asexual female (Figs 114 – 118). Head and antenna rusty brown, except dark brown to black frons and interocellar area, mouthparts light brown; mesoscutellum, metascutellum, propodeum dark brown to black, legs rusty brown, metasoma dark brown. Head reticulate, with sparse white setae, transverse, 1.3 × as broad as high and broader than mesosoma in frontal view, 2.5 × as broad as long in dorsal view. Gena reticulate, broadened behind eye in frontal view, as broad as transverse diameter of eye in lateral view. Malar space with malar sulcus and multiple delicate parallel striae on both sides of sulcus, radiating from clypeus and reaching eye; eye 2.2 × as high as height of malar space. Inner margins of eyes diverging slightly ventrally. POL 1.4 × as long as OOL; OOL 2.7 × as long as diameter of lateral ocellus and 1.8 × as long as LOL; all ocelli rounded, of same size. Antennal toruli in the ventral half of head height. Transfacial distance 1.1 × as long as height of eye; diameter of antennal torulus 1.2 × as long as distance between them, distance between torulus and eye 1.7 × as long as diameter of torulus. Lower face reticulate, with white setae, without striae; median area delicately coriaceous, slightly elevated. Clypeus quadrangular, slightly broader than high, smooth, glabrous, with dense long white setae, ventrally slightly emarginate, without median incision; anterior tentorial pit large, rounded, epistomal sulcus and clypeo-pleurostomal line broad, well impressed. Frons reticulate, area under central ocellus and interocellar area coriaceous. Vertex uniformly reticulate, occiput with delicate transverse parallel rugae; postocciput reticulate, glabrous; postgena reticulate, with white setae; posterior tentorial pit large, ovate, area below impressed; occipital foramen higher than height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which diverge strongly toward occipital foramen, postgenal bridge anteriorly broader than high. Antenna shorter than head + mesosoma, with 12 flagellomeres, pedicel slightly longer than broad, F 1 2.2 × as long as pedicel and 1.4 x as long as F 2, F 2 slightly longer than F 3, subsequent flagellomeres gradually shorter until F 10, F 11 2.2 × as long as F 10 (in some specimens a suture visible so antenna has 13 flagellomeres), placodeal sensilla on F 5 – F 12. Mesosoma slightly longer than high. Pronotum reticulate with uniformly distributed parallel rugae; propleuron smooth, glabrous. Mesoscutum uniformly rugose between notauli, reticulate on outer side of notaulus, without setae; slightly longer than broad (greatest width measured across mesoscutum level with base of tegulae). Notaulus complete, deep, broad; bottom smooth, glabrous, with transverse rugae; posteriorly slightly broader and slightly converging; anterior parallel line impressed in anterior 1 / 3 of mesoscutum length; median mesoscutal line in the form of a short impression; parapsidal line distinct, broad, delimited by smooth glabrous area, extending to 2 / 3 of mesoscutum length; circumscutellar carina broad, smooth, foveolate. Mesoscutellum elongated, slightly longer than broad, trapezoid, posteriorly rounded, uniformly dull rugose, overhanging metanotum. Mesoscutellar foveae separated by narrow elevated rugose central area, transversely ovate, 2.0 × as broad as high; bottom smooth, glabrous, with strong longitudinal parallel rugae. Mesopleuron uniformly reticulate; mesopleural triangle rugose, with irregular striae; dorsal and lateral axillar areas reticulate, with a few white short setae; subaxillular bar smooth, glabrous, at posterior end as high as height of metanotal trough; metapleural sulcus reaching mesopleuron in upper 1 / 3 of its height, upper part of sulcus distinct. Metascutellum rugose, 3.0 x as high as height of smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous, with strong longitudinal parallel rugae; central propodeal area smooth, glabrous, with strong irregular rugae; lateral propodeal carinae strong, nearly parallel; lateral propodeal area smooth, glabrous, with strong irregular rugae. Nucha with strong sulci dorsally and laterally. Coxae reticulate, rest of legs smooth, glabrous; tarsal claws simple, without basal lobe. Forewing longer than body, hyaline, margin without cilia, veins dark brown, radial cell open, 2.6 × as long as broad; R 1 and Rs reaching wing margin; areolet absent, Rs + M distinct along entire length, reaching basalis slightly above its mid height. Metasoma longer than head + mesosoma, longer than high in lateral view; 2 nd metasomal tergum extending to 1 / 3 length of metasoma in dorsal view, without setae anterolaterally, microreticulate in posterior 1 / 3; all subsequent terga microreticulate, glabrous. Prominent part of ventral spine of hypopygium slightly longer than broad in ventral view, narrowing towards apex, with short setae ventrally which do not extend beyond apex of spine. Body length 3.1 – 3.3 mm (n = 5). Gall. Sexual generation gall (Fig. 119) is a large, multilocular thickening of the leaf midrib / petiole. Asexual galls (Fig. 120) are in acorns (Beiderbeck & Nicholls 2014). The acorns are stunted, smaller than a non-infected acorn, and contain up to 6 – 7 cryptic larval chambers; the cotyledons are brown or used up by the larvae (Beiderbeck & Nicholls 2014).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF97CE2C4E9D50C0A849AB6F.taxon	biology_ecology	Biology. See Beiderbeck (2012), Beiderbeck & Nicholls (2014), and Pujade-Villar et al. (2014) for the biology of this species. The sexual generation develops on Q. rubra and Q. velutina; the asexual only known so far from Q. rubra. Sexual generation leaf galls develop in early spring; adults emerge in May-June. Asexual acorn galls were found in October - November, maturing in November. Adults overwinter in galls and emerge in the following spring.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF97CE2C4E9D50C0A849AB6F.taxon	distribution	Distribution. USA: New York, Virginia, North Carolina, Illinois, Iowa (Burks 1979). Also an introduced species in Germany (Heidelberg) on introduced Q. rubra (Beiderbeck 2012, Beiderbeck & Nicholls 2014). Molecular taxonomy. Alternate generations (using larvae of the asexual generation) were matched with sequence data by Beiderbeck & Nicholls (2014); here we supplement those data with additional sequences from sexual and asexual adults. In total, nine individuals (four asexual females, two sexual females, three sexual males) were sequenced for cytb and seven individuals (four asexual females, two sexual females, one sexual male) were sequenced for ITS 2; these individuals were sourced from an introduced population in Germany as well as a nativerange population in North Carolina, USA. Cytb sequences were on average 0.18 % divergent (range 0 – 0.46 %; GenBank accessions MG 821060, MW 326694, OM 321634 – OM 321640); ITS 2 sequences were 0 – 0.20 % divergent (OM 331823 – OM 331829).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF97CE2C4E9D50C0A849AB6F.taxon	discussion	Comments. The genus Melikaiella Pujade-Villar, 2014 (Pujade-Villar et al. 2014) is allied to “ true ” Callirhytis from the Western Palaearctic, or as previously called, Callirhytis ‘ sensu stricto’ (Pujade-Villar et al. 2012 a). It thus makes good sense that their asexual generations develop in cryptic galls inside acorns, as is also the case for Callirhytis sensu stricto. Melikaiella was described with three new species from Mexico: M. amphibolensis Pujade-Villar, 2014, M. bicolor Pujade-Villar, 2014 and M. reticulata Pujade-Villar, 2014; and one from the USA, M. ostensackeni Pujade-Villar, 2014. The species Callirhytis sonorae Weld, 1944 from Mexico, C. corrugis, C. flora Weld, 1922, C. fructicola, C. fructuosa, C. lupana Weld, 1944 and C. tumifica were also transferred to Melikaiella, and Callirhytis petrosa Weld, 1922 was synonymized to Melikaiella corrugis (Pujade-Villar et al. 2014). The species status of Cynips papula Bassett, 1881 from the USA, an earlier synonym of Callirhytis quercusmodesta (Osten Sacken, 1861), was restored and this species was also transferred to Melikaiella, under the new name combination M. papula (Pujade-Villar et al. 2014). Subsequently, Medianero & Nieves-Aldrey (2014) described the species Callirhytis cameroni from Panama, and although they mentioned that it most likely belonged to the genus Melikaiella, it was not formally transferred. Thus, the number of species in this genus is currently 12.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF99CE2B4E9D5498AC96AFC7.taxon	description	Figs 121 – 150 Synonyms: Cynips floccosa Bassett (1881 b: 111), asexual female, gall. Neuroterus floccosus combination by Mayr (1881: 37). Neuroterus verrucarum var. floccosus combination by Kinsey (1923: 70); combination of Mayr (1881) restored by Weld (1951: 621).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF99CE2B4E9D5498AC96AFC7.taxon	materials_examined	Material examined: Nine asexual females labelled as “ Canada, Manitoba, Assiniboine Forest, 49.85862 ° N, - 97.25015 ° W, GPS ± 10 m, S. Digweed, Collected 2005. viii. 30 ex Quercus macrocarpa, Reared 15 - 25. v. 2006 ex gall of Neuroterus quercusverrucarum agamic gen. ”, determined originally as Neuroterus quercusverrucarum asexual generation females by S. Digweed. Seven sexual females and 4 males labelled as “ Canada, Alberta, Edmonton, Northern Forest Centre, 53.49165 ° N, - 113.54291 ° W, GPS ± 10 m, Reared 4 - 13. vii. 2006 ex galls of Neuroterus quercusverrucarum ♂ ♀ ex Quercus macrocarpa, S. Digweed ”, determined originally as Neuroterus quercusverrucarum sexual generation females by S. Digweed. Specimens have been deposited at the USNM, PHDNRL and RAM.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF99CE2B4E9D5498AC96AFC7.taxon	diagnosis	Diagnosis. Asexual females of N. floccosus most closely resemble those of N. quercusverrucarum. In N. floccosus, asexual female the gena 0.5 × as wide as the transverse diameter of eye in lateral view, diameter of antennal torulus 1.9 × as long as distance between them, F 1 as long as pedicel; the mesoscutellum trapezoid, the posterior half of which gradually narrows towards its posterior end; the metascutellum 3.0 × as high as height of ventral impressed area; the prominent part of the ventral spine of hypopygium 2.0 × as long as broad in ventral view, with short sparse setae ventrally. In N. quercusverrucarum the gena 1 / 3 the width of transverse diameter of eye in lateral view, diameter of antennal torulus slightly greater than distance between them, antenna F 1 1.3 × as long as pedicel; the mesoscutellum posteriorly rounded; the metascutellum as high as height of ventral impressed area; the prominent part of the ventral spine of hypopygium as long as broad in ventral view, without setae ventrally. Below the description of the asexual female is given according to current morpho-description requirements (the original description by Bassett (1881 b) is very superficial).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF99CE2B4E9D5498AC96AFC7.taxon	description	Description. Asexual female (Figs 121 – 131). Head, mesosoma, metasoma brown; antenna light brown; legs yellowish to white, with darker femora and tibiae. Head alutaceous, with a few setae on lower face, 1.2 × as broad as high and as broad as mesosoma in frontal view, 1.9 × as broad as long in dorsal view. Gena alutaceous, not broadened behind eye in frontal view, 0.5 × as wide as the transverse diameter of eye in lateral view. Malar space alutaceous, with distinct malar sulcus, eye 3.6 × as high as height of malar space. Inner margins of eyes converge slightly ventrally. POL 2.0 × as long as OOL; OOL 1.4 × as long as diameter of lateral ocellus and equal to LOL; all ocelli ovate, of same size. Transfacial distance slightly longer than height of eye; diameter of antennal torulus 1.9 × as long as distance between them, distance between torulus and eye 1.5 × as long as diameter of torulus. Lower face and median slightly elevated area alutaceous, with white setae, without striae. Clypeus quadrangular, as broad as high, alutaceous, with long setae along ventral edge; ventrally not emarginate, without median incision; anterior tentorial pit small, rounded, indistinct; epistomal sulcus and clypeo-pleurostomal line impressed, distinct. Frons and interocellar area alutaceous, without setae. Vertex, occiput, postocciput, postgena alutaceous; posterior tentorial pit large, ovate, area below impressed; occipital foramen 1.5 × shorter than height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which are widely spaced, diverge only slightly toward occipital foramen, postgenal bridge only slightly broader than high anteriorly. Antenna longer than head + mesosoma, with 11 flagellomeres, flagellomeres half as wide as scape and pedicel, gradually broadening until F 11; pedicel as long as scape, 1.5 × as long as broad; F 1 as long as pedicel, 1.2 × as long as F 2, F 2 1.2 × as long as F 3, F 3 = F 4, F 5 = F 6, F 7 = F 8 = F 9 = F 10, F 11 slightly longer than F 10; placodeal sensilla on F 2 – F 11. Mesosoma slightly longer than high. Pronotum alutaceous, without setae in front view; alutaceous laterally; propleuron alutaceous. Mesoscutum uniformly alutaceous, without setae; slightly longer than broad (greatest width measured across mesoscutum level with base of tegulae). Notaulus, anterior parallel line, median mesoscutal line and parapsidal line absent; circumscutellar carina present, narrow. Transscutal articulation absent, mesoscutum fused with mesoscutellum; as a result of fusion of the mesoscutum and mesoscutellum, the boundary between these structures is not straight but slightly curved towards the mesoscutum; the mesoscutum is emarginate posterolaterally and slightly elevated above the dorsoaxillar area. Mesoscutellum trapezoid, longer than broad, posterior half gradually narrowing towards posterior end, uniformly smooth, glabrous, without setae, overhanging metanotum. Mesopleuron higher than broad in lateral view, smooth, glabrous; mesopleural triangle smooth, glabrous, with interrupted delicate striae, without setae; dorsal and lateral axillar areas smooth, glabrous, without setae; subaxillular bar smooth, glabrous, at posterior end as high as height of metanotal trough; metapleural sulcus reaching mesopleuron at half its height, delimiting a narrow smooth area along mesopleuron; upper part of sulcus absent. Metascutellum smooth, 3.0 x as high as height of narrow, smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous, without setae; propodeum smooth, glabrous; propodeum uniformly alutaceous, without carinae, striae or setae. Nucha short, smooth, without sulci. Tarsal claws simple, without basal lobe. Forewing longer than body, hyaline, margin with dense long cilia, veins brown, distinct, radial cell open, 5.0 × as long as broad; R 1 and Rs reaching wing margin, Rs reaching wing margin; areolet large, triangular, distinct, Rs + M brown, visible along 2 / 3 of its length, projection reaching basalis in the lower 1 / 3 of its height. Metasoma slightly longer than head + mesosoma, slightly longer than high in lateral view; all terga weakly sclerotized, thus limits of terga hardly visible. 2 nd metasomal tergum extending to 1 / 3 of length of metasoma in dorsal view, without setae; all terga smooth, glabrous, without micropunctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium short, 2.0 × as long as broad in ventral view, with short sparse setae ventrally. Body length 1.1 – 1.3 mm (n = 4). Sexual female (Figs 132 – 136, 142 – 147). Head, mesosoma, metasoma brown; mandibles, palpi, antenna, legs lighter than body. Head alutaceous, with sparse white setae, denser on lower face; slightly higher than broad and narrower than mesosoma in frontal view, slightly broader than long in dorsal view. Gena alutaceous, not broadened behind eye in frontal view, narrower than transverse diameter of eye in lateral view. Malar space alutaceous, with deep malar sulcus; eye 3.7 × as high as height of malar space. Inner margins of eyes parallel. POL 2.5 × as long as OOL; OOL slightly longer than diameter of lateral ocellus and slightly shorter than LOL; all ocelli elongate, ovate, of same size. Transfacial distance shorter than height of eye; diameter of antennal torulus longer than distance between them, distance between torulus and eye longer than diameter of torulus. Toruli located in upper 1 / 3 of face (lower face + frons) height. Lower face alutaceous, without striae, median area not elevated. Clypeus trapezoid, longer than broad, alutaceous, with a few setae scattered all around; ventrally rounded, not emarginate, without median incision; anterior tentorial pit large, rounded, distinct; epistomal sulcus and clypeo-pleurostomal line impressed, distinct. Frons and interocellar area alutaceous, without setae; area under frontal ocellus impressed, alutaceous. Vertex, occiput, postocciput, postgena alutaceous; posterior tentorial pit large, ovate, area below impressed; occipital foramen less than 0.5 × as long as height of smooth, matte postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which diverge toward occipital foramen. Antenna as long as head + mesosoma, with 11 flagellomeres, scape shorter than pedicel, scape + pedicel longer than F 1, pedicel 2.0 × as long as broad, F 1 1.2 × as long as F 2, F 2 – F 4 nearly equal in length; F 5 shorter than F 4; F 5 – F 10 equal in length; F 11 slightly longer than F 10; placodeal sensilla on F 3 – F 11, absent on F 1 – F 2. Mesosoma slightly higher than long in lateral view, without setae. Pronotum alutaceous, without striae laterally; propleuron alutaceous. Mesoscutum smooth, glabrous, as long as broad (greatest width measured across mesoscutum level with base of tegulae). Notaulus, anterior parallel line, median mesoscutal line and parapsidal line absent; circumscutellar carina narrow but present. Transscutal articulation absent, mesoscutum fused with mesoscutellum; as a result of fusion of the mesoscutum and mesoscutellum, the boundary between these structures is not straight but slightly curved towards the mesoscutum; the mesoscutum is emarginate posterolaterally and slightly elevated above the dorsoaxillar area. Mesoscutellum rounded, as long as broad (broadest part at mid height); posteriorly rounded, uniformly smooth, glabrous, with very few short scattered white setae, slightly overhanging metanotum. Mesoscutellar foveae absent, mesoscutellum with semilunar impression anteriorly. Mesopleuron and mesopleural triangle uniformly alutaceous, without setae; dorsal and lateral axillar areas alutaceous, without setae; subaxillular bar smooth, glabrous, triangular, highest part at anterior end and gradually narrowing towards posterior end; most anterior end higher than height of metanotal trough; metapleural sulcus reaching mesopleuron slightly below half its height, delimiting broad smooth area along mesopleuron; upper part of sulcus distinct, delimiting a narrow area along mesopleuron. Metascutellum smooth, 3.0 x as high as height of smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous, without setae; propodeum smooth, glabrous; central propodeal area alutaceous, with delicate longitudinal interrupted rugae; lateral propodeal carinae absent; lateral propodeal area smooth, glabrous, without setae. Nucha short, smooth, with strong longitudinal sulci dorsally and laterally. Tarsal claws simple, without basal lobe. Forewing longer than body, hyaline, margin with dense long cilia, veins light brown, radial cell open, 3.5 × as long as broad; R 1 and Rs reaching wing margin; areolet small, triangular, distinct, Rs + M indistinct, projection reaching basalis in the lower 1 / 3 of its height. Metasoma as long as head + mesosoma, 2.0 × as high as long; all terga weakly sclerotized, thus limits of terga hardly visible. 2 nd metasomal tergum extending to half the length of metasoma in dorsal view, without setae; all terga smooth, glabrous, without micropunctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium short, as long as broad in ventral view, with a few long setae ventrally. Body length 1.0 – 1.2 mm (n = 6). Male (Figs 137 – 141, 148). Similar to female but body light brown, mandibles, palpi, antenna, legs yellow. Antenna nearly as long as body, ocelli at least 2.0 × bigger than in female; forewing longer than body, 2 nd metasomal tergum in the form of a narrow long petiole; metasoma shorter than mesosoma. Antenna with 12 flagellomeres, scape shorter than pedicel, scape + pedicel nearly as long as F 1, pedicel 1.8 × as long as broad, F 1 = F 2, F 2 = F 3 = F 4; F 5 shorter than F 4; F 5 = F 6, F 7 – F 10 equal in length; F 12 slightly longer than F 11; placodeal sensilla on F 1 – F 12. Body length 0.8 – 1.0 mm (n = 4). Gall. Sexual generation minute (approximately 1 mm long), ovoid, seed-like integral leaf galls, smooth and only covered with normal leaf pubescence (Fig. 150). Galls are visible and project slightly from both upper and lower leaf surfaces, and often but not always occur alongside major leaf veins. Galls appear reddish-brown from above and paler tan from below. Exit holes of adult insects generally occur on the underside of the leaf.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF99CE2B4E9D5498AC96AFC7.taxon	biology_ecology	Biology. See Bassett (1881 b), Beutenmueller (1910), and Weld (1926) for the biology of the asexual generation. Asexual galls (Fig. 149) have been recorded on leaves of Q. bicolor and Q. macrocarpa. Galls become apparent in August as small depressions filled with loose, pale wool on the underside of new, terminal, long shoot leaves. Multiple galls often co-occur on a single leaf, which causes the leaf to curl, and galls tend to be found on stumps or damaged trees bearing vigorous, long late-season shoots. Adult females of the asexual generation emerge the spring following gall induction, from mid-April to late May. Galls of the sexual generation first appeared in late May on Q. macrocarpa in Edmonton, and adults emerged in late July and early August.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF99CE2B4E9D5498AC96AFC7.taxon	distribution	Distribution. USA: from Virginia west to Illinois (Burks 1979). Canada: Ontario (Burks 1979), Manitoba. This species was only reared in sleeves in Edmonton, Alberta, and does not naturally occur on bur oak there. Molecular taxonomy. Alternating generations were matched using DNA data, with five individuals (three asexual females, two sexual females) sequenced for both cytb and ITS 2. Two cytb haplotypes were found across the five individuals, differing by 0.23 % (GenBank accessions OM 321641 – OM 321645); ITS 2 sequences were identical (OM 331830 – OM 331834). The distinctiveness of N. floccosus from its close relative N. quercusverrucarum was confirmed with both cytb and ITS 2 data (GenBank accessions OM 321650 – OM 321651 and OM 331839 – OM 331840 for N. quercusverrucarum). On average, cytb haplotypes differed between the two species by 5.20 % (range 4.85 – 5.54 %), while ITS 2 sequences differed by 2.38 % (with two indels). These values are consistent with those expected between distinct congeneric species.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF9ECE334E9D5170AF4AAE77.taxon	description	Figs 151 – 182 Synonyms: Neuroterus nigrum Gillette (1888: 218), females and galls. Revised spelling by Dalla Torre and Kieffer (1902: 51).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF9ECE334E9D5170AF4AAE77.taxon	description	The fifth variety described by Kinsey is Neuroterus niger var. alimas Kinsey, 1923 (sexual generation, Texas, on Q. virginiana) that was still considered as form of N. niger by Burks (1979). According to Kinsey (1923) this variety induces a sexual generation spring gall, while the other four varieties induce autumn asexual generation galls. Moreover, N. niger var. alimas is the only variety associated with section Virentes oaks while the other four are on section Quercus oaks, and as mentioned in the Introduction, no Nearctic cynipid species are shared across the oak sections (Abrahamson et al. 1998 a, 2003; Stone et al. 2009). Hence biological information suggests that N. niger var. alimas is a species distinct from N. niger (sensu Gillette 1888) and the other four of Kinsey’s varieties. Kinsey (1923) described N. niger var. alimas based on one female (holotype – lost) and 13 males and gave no detailed morphological description for this variety. Thus, there is no morphological evidence for synonymisation of N. niger var. alimas to N. niger while at the same time it cannot be treated as a valid species so Melika et al. (2021) moved it to nomen dubium. Types examined. Asexual generation of Neuroterus niger Gillette: one female labelled as “ Ac. Cat. 851 ”, “ Ag. Coll. Michigan 3 - 25 - 88 ”, red label “ Type No 1898 USNM ”, handwritten “ N. nigrum ”. Neuroterus niger var. alimas Kinsey, 1923: holotype female labelled as “ Acc. 24856 ”, pink ” N. alimas, female, Holo- Paratype ”, ” Q. virginiana ”, ” Austin, Tex. 4.16.21., Patterson ” – insect absent, only gall. Neuroterus niger var. arizonicae Kinsey, 1923: holotype female labelled as ” Ac. 24856 ”, pink ” N. arizonica, Holo- Paratype’, ” Q. arizonica, Kinsey coll. ”, ” Bisbee, Ariz. Gall 1.15.20. ” – insect absent, only gall. Neuroterus niger var. griseae Kinsey, 1923: holotype female labelled as ” Ac. 24856 ”, pink ” N. griseae, Holo-Paratype’, ” Q. grisea, Kinsey coll. ”, ” Fort Davis, Tex., Gall. 12.16.19. ” – insect absent, only gall. Neuroterus niger var. nigripes Kinsey, 1923: holotype female labelled as ” Ac. 24856. ”, pink ” Neuroterus nigripes, Holo- Paratype’, ” Q. stellata, Kinsey coll. ”, ” Austin, Tex. 12.4.19. ” Neuroterus niger var. pattersoni f. pattersoni Kinsey, 1923: holotype female labelled as ” Ac. 24856. ”, pink ” Neuroterus pattersoni, Holo- Paratype’, ” Q. stellata, Patterson coll. ”, ” Austin, Tex. 5.7.21., gall 310. 21. ” Sexual generation of Neuroterus niger: 3 females on one pin “ Io ”, “ Ac. Cat. 996 ”, red label “ Type ”, handwritten label “ Neuroterus vernus Gill. ”; 5 females on one pin: handwritten label “ Iowa, Gillette ”, white label “ Type ”, “ colCF Baker ”; also one label with galls and red label “ Type ”. Neuroterus papillosus Beutenmueller, 1910: two females on two pins, both labelled as “ Bronx Park, New York City, W. Beutenmüller ”, red label “ Type ”, “ Beut. Coll. rec’d 1935 ”, handwritten label “ Neuroterus papillosus Beutnm ”. Types were examined by GM during his multiple visits to the AMNH, NYC and USNM, Washington, DC.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF9ECE334E9D5170AF4AAE77.taxon	materials_examined	Material examined. Five asexual females labelled as “ CANADA: Alberta, Edmonton, Grand Trunk Pool, SC 30, 2002.04.12. S. Digweed, Neuroterus niger Gillette females of the asexual generation, det. S. Digweed 2005 ”. Sexual generation: 5 females labelled as “ CANADA, Alberta, Edmonton. Reared 18. iv. 2005, ex Q. macrocarpa. N. niger (?) gall, Grand Trunk, coll. 22. ix. 04. S. Digweed ”; 5 males labelled as “ CANADA, Alberta, Edmonton. Reared 3 - 13. x. 2005, ex Q. macrocarpa. N. niger (?) gall, Grand Trunk, coll. 22. ix. 04. S. Digweed ”. Specimens have been deposited at the USNM, PHDNRL and RAM. Asexual females have been deposited at the PHDNRL and RAM.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF9ECE334E9D5170AF4AAE77.taxon	diagnosis	Diagnosis. Belongs to Kinsey’s (1923) subgenus Diplobius. The oak gallwasp (Cynipini) fauna of the Nearctic north of Mexico is divided into three geographic zones, with distinct species: the eastern United States (Weld 1959), the US Southwest (Weld 1960) and the Pacific Slope (Weld 1957). In the eastern US four species, including N. niger, induce parenchyma thickening integral leaf galls; however, the three other Neuroterus species are all only known from their sexual generations. Neuroterus fugiens Weld, 1926, induces galls on Q. macrocarpa in spring in Illinois, USA; N. papillosus Beutenmueller, 1910, induces galls on Q. bicolor in USA: NY, NJ, IN, IL; N. perminimus Bassett, 1900, induces galls on Q. alba in USA: NY, MD, VA, OH, IL and Canada: ON (Burks 1979). Galls of N. fugiens are quite different, being covered in rose-red hairs on their lower surface and not emerging from the upper leaf surface (Weld 1926). However, the galls of N. papillosus are the same as in the sexual generation of N. niger designated herein. Kinsey (1923) treated N. papillosus as Neuroterus niger var. papillosus and N. perminimus as N. niger var. perminimus and our assessment of morphology supports this synonymy. Below the descriptions of the asexual and sexual females and males are given according to the current morpho-description requirements (the original descriptions by Gillette (1888, 1889) are very superficial).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF9ECE334E9D5170AF4AAE77.taxon	description	Description. Asexual female (Figs 151 – 161). Body, antenna, legs uniformly dark brown. Head alutaceous, with sparse white setae, denser on lower face; strongly transverse, 1.3 × as broad as high and broader than mesosoma in frontal view, 1.9 × as broad as long in dorsal view. Gena alutaceous, broadened behind eye in frontal view, just over half as wide as transverse diameter of eye in lateral view. Malar space alutaceous, with malar sulcus, eye 3.8 × as high as height of malar space. Inner margins of eyes parallel. POL 1.8 × as long as OOL; OOL 2.1 × as long as diameter of lateral ocellus and as long as LOL; all ocelli slightly ovate, of same size. Transfacial distance 1.2 × as long as height of eye; diameter of antennal torulus slightly longer than distance between them, distance between torulus and eye 1.3 × as long as diameter of torulus. Lower face and median elevated area alutaceous, with white setae, without striae. Clypeus rectangular, 1.8 × as broad as high, alutaceous, with long setae along ventral edge; ventrally not emarginate, without median incision; anterior tentorial pit small, rounded, indistinct; epistomal sulcus and clypeo-pleurostomal line impressed, distinct. Frons and interocellar area alutaceous, without setae. Vertex, occiput, postocciput, postgena alutaceous; posterior tentorial pit large, ovate, area below impressed; occipital foramen shorter than height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which are widely spaced, only slightly diverge toward occipital foramen, postgenal bridge anteriorly only slightly broader than high. Antenna longer than head + mesosoma, with 10 flagellomeres, pedicel 1.8 × as long as broad, F 1 1.2 × as long as pedicel and 1.4 x as long as F 2, F 2 = F 3, F 4 slightly longer than F 3, F 5 shorter than F 4 and shorter than F 6, F 7 to F 9 gradually shorter until F 9; F 10 2.0 × as long as F 9 (F 10 probably consists of two flagellomeres, with invisible suture between two flagellomeres); placodeal sensilla on F 5 – F 10. Mesosoma as long as high. Pronotum smooth, polished, glabrous, without setae in frontal view; alutaceous laterally; propleuron smooth, glabrous. Mesoscutum smooth, glabrous, with setae only along sides; as long as broad (greatest width measured across mesoscutum level with base of tegulae). Notaulus, anterior parallel line, median mesoscutal line and parapsidal line absent; circumscutellar carina narrow present. Transscutal articulation absent, mesoscutum fused with mesoscutellum; as a result of fusion of the mesoscutum and mesoscutellum, the boundary between these structures is not straight but slightly curved towards the mesoscutum; the mesoscutum is emarginate posterolaterally and slightly elevated above the dorsoaxillar area. Mesoscutellum nearly as broad as long, posteriorly rounded, uniformly smooth, glabrous, without setae, overhanging metanotum; mesoscutellar foveae absent. Mesopleuron with speculum and mesopleural triangle uniformly alutaceous, without setae; transverse impressed line present in ventral 1 / 4 of height of mesopleuron; dorsal and lateral axillar areas alutaceous, without setae; subaxillular bar smooth, glabrous, at posterior end as high as height of metanotal trough; metapleural sulcus reaching mesopleuron slightly below half its height, delimiting narrow smooth area along mesopleuron. Metascutellum smooth, as high as height of smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous, without setae; propodeum smooth, glabrous; central propodeal area smooth, glabrous, with some longitudinal interrupted rugae; lateral propodeal carinae absent; lateral propodeal area smooth, glabrous, without setae. Nucha short, smooth, without sulci. Tarsal claws simple, without basal lobe. Forewing longer than body, hyaline, margin with dense long cilia, veins dark brown, radial cell open, 5.0 × as long as broad; R 1 nearly reaching wing margin, Rs reaching wing margin; areolet large, triangular, indistinct, Rs + M light brown, visible along 1 / 3 of its length, projection reaching basalis in the lower 1 / 3 of its height. Metasoma slightly longer than head + mesosoma, nearly as long as high in lateral view; all terga weakly sclerotized, thus limits of terga hardly visible. 2 nd metasomal tergum extending to half the length of metasoma in dorsal view, without setae; all terga smooth, glabrous, without micropunctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium short, greater than 2.0 × as long as broad in ventral view, without setae ventrally. Body length 1.4 – 1.5 mm (n = 3). Sexual female (= N. vernus) (Figs 162 – 166, 172 – 177). Body, antenna, legs uniformly reddish brown. Head alutaceous, with sparse white setae on lower face; transverse, 1.3 × as broad as high and broader than mesosoma in frontal view, 2.0 × as broad as long in dorsal view. Gena alutaceous, slightly broadened behind eye in frontal view, narrower than transverse diameter of eye in lateral view. Malar space alutaceous, with malar sulcus in the form of a deep impressed stripe; eye 4.5 × as high as height of malar space. Inner margins of eyes parallel. POL 2.5 × as long as OOL; OOL 1.4 × as long as diameter of lateral ocellus and as slightly shorter than LOL; all ocelli ovate, of same size. Transfacial distance 1.2 × as long as height of eye; diameter of antennal torulus slightly longer than distance between them, distance between torulus and eye 1.5 × as long as diameter of torulus. Lower face and median elevated area alutaceous, without setae and striae. Clypeus rounded, as long as broad, alutaceous, with a few setae scattered all over; ventrally rounded, not emarginate, without median incision; anterior tentorial pit large, rounded, distinct; epistomal sulcus and clypeo-pleurostomal line impressed, distinct. Frons and interocellar area alutaceous, without setae; area under frontal ocellus impressed, alutaceous. Vertex, occiput, postocciput, postgena alutaceous; posterior tentorial pit large, ovate, area below impressed; occipital foramen shorter than height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which are widely spaced and only diverge slightly toward occipital foramen, postgenal bridge anteriorly only slightly broader than high. Antenna longer than head + mesosoma, with 11 flagellomeres, pedicel 1.6 × as long as broad, F 1 1.5 × as long as pedicel and 1.6 × as long as F 2, F 2 = F 3, F 4 slightly longer than F 3 and equal F 5 and F 6, all subsequent flagellomeres slightly shorter than F 6 and all equal in length; placodeal sensilla on F 2 – F 11. Mesosoma as long as high. Pronotum alutaceous, with a few setae in frontal view; alutaceous laterally; propleuron alutaceous. Mesoscutum smooth, glabrous, with setae only along sides; as long as broad (greatest width measured across mesoscutum level with base of tegulae). Notaulus, anterior parallel line, median mesoscutal line and parapsidal line absent; circumscutellar carina narrow present. Transscutal articulation absent, mesoscutum fused with mesoscutellum; as a result of fusion of the mesoscutum and mesoscutellum, the boundary between these structures is not straight but slightly curved towards the mesoscutum; the mesoscutum is emarginate posterolaterally and slightly elevated above the dorsoaxillar area. Mesoscutellum nearly as broad as long, posteriorly rounded, uniformly smooth, glabrous, without setae, overhanging metanotum; mesoscutellar foveae absent. Mesopleuron and mesopleural triangle uniformly alutaceous, without setae; speculum smooth, glabrous; transverse impressed line present in ventral 1 / 4 of height of mesopleuron, extending across entire width of mesopleuron; dorsal and lateral axillar areas alutaceous, without setae; subaxillular bar smooth, glabrous, at posterior end as high as height of metanotal trough; metapleural sulcus reaching mesopleuron slightly below half its height, delimiting narrow smooth area along mesopleuron. Metascutellum smooth, 3.0 × as high as height of smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous, without setae; propodeum smooth, glabrous; central propodeal area coriaceous, with longitudinal interrupted rugae; lateral propodeal carinae absent; lateral propodeal area smooth, glabrous, without setae. Nucha short, smooth, without sulci. Tarsal claws simple, without basal lobe. Forewing longer than body, hyaline, margin with dense long cilia, veins light brown, radial cell open, 5.0 × as long as broad; R 1 and Rs reaching wing margin; areolet small, triangular, indistinct, Rs + M light brown, visible along 2 / 3 of its length, projection reaching basalis slightly below its mid height. Metasoma as long as head + mesosoma, nearly as long as high in lateral view; all terga weakly sclerotized, thus limits of terga hardly visible. 2 nd metasomal tergum extending to half the length of metasoma in dorsal view, without setae; all terga smooth, glabrous, without micropunctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium short, as long as broad in ventral view, without setae ventrally. Body length 1.4 – 1.6 mm (n = 4). Male (Figs 167 – 171, 178 – 180). Similar to female, body brown, antenna and legs slightly lighter; ocelli bigger than in female. Body length 0.9 – 1.4 mm (n = 5). Originally Gillette (1889) described only females, no males were observed or reared. Gall. The asexual generation gall is described by Gillette (1888) and are small blisters integral to the leaf blade (Fig. 181). The sexual generation galls (Fig. 182), first recorded as N. vernus, are cells hidden within new shoots, on the young leaf petioles and midribs and on the catkins of Q. macrocarpa; these plant organs are sometimes distorted as a result (Digweed 2010).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF9ECE334E9D5170AF4AAE77.taxon	biology_ecology	Biology. See Gillette (1890), Beutenmueller (1910), Kinsey (1923), and Weld (1926) for the biology of one or both generations. The minute, integral leaf galls of the asexual generation have been recorded on Q. macrocarpa. Galls became evident in Edmonton in August and matured in September. This species represents a rare case where males are present in the asexual generation. Males of the asexual generation comprised a tiny fraction of the cynipids reared from asexual generation galls, with only 62 males reared from gall collections made in Edmonton during 2004 - 2006, as opposed to 20979 females. In addition, most males were reared in the autumn (October-December) of the year of gall formation, whereas almost all females emerged in April-May the following spring after gall formation. Thus, most males were temporally isolated from ovipositing females and therefore essentially functionless. This is similar to Patterson’s (1928) discovery of functionless males of the asexual generations of Neuroterus contortus (Weld, 1921) and N. quercusrileyi (Bassett, 1881). In his study, males and females co-occurred, but they appeared unwilling to mate under laboratory conditions. Galls of the sexual generation are on Q. alba, Q. bicolor and Q. macrocarpa (Gillette 1890, Burks 1979, Digweed 2010). We observed that clusters of galls (e. g., on one catkin) often produced adults of only one gender. Galls were visible in Edmonton in May, with adults emerging in late May or early June. Sexual generation females were observed ovipositing mostly on the undersides of leaves and often on leaves that were on long shoots and / or that were not yet fully mature.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF9ECE334E9D5170AF4AAE77.taxon	distribution	Distribution. In the USA the asexual generation has been recorded from Massachusetts, Michigan, Wisconsin, Iowa (Burks 1979), Illinois, Indiana, New York, New Jersey, Pennsylvania, Ohio (Beutenmueller 1910); Arizona, Texas (Kinsey 1923), and Alberta and Manitoba in Canada. The sexual generation has been recorded from Illinois (Weld 1926), Iowa (Burks 1979), and TX (Kinsey 1923) in the USA and Alberta in Canada. Molecular taxonomy. Alternating generations of this species, which have been previously described under different names (Digweed 2010), were matched using DNA data from four individuals (two asexual females, two sexual females) sequenced for both cytb and ITS 2. Cytb sequences were identical among the four individuals (GenBank accessions OM 321646 – OM 321649), as were ITS 2 sequences (OM 331835 – OM 331838).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF9ECE334E9D5170AF4AAE77.taxon	discussion	Comments. In the Southwest of the USA (AZ, UT) one species, N. howertoni Bassett, 1890 asexual generation, is known to induce leaf parenchyma thickening galls on Q. arizonica and Q. oblongifolia which are like those of N. niger (Weld 1960), while in California there is another species, N. engelmannii Kinsey, 1922, which induces similar galls on Q. engelmannii (Kinsey 1922 a, 1923; Weld 1957).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF86CE384E9D51E1A859ABB3.taxon	description	Figs 183 – 202, 204 Synonyms: Philonix fulvicollis Fitch (1859: 783), asexual females; Cynips (Teras) fulvicollis combination by Osten Sacken (1865: 379); Biorhiza fulvicollis combination by Ashmead (1885: 296); Biorrhiza fulvicollis combination and incorrect spelling by Dalla Torre (1893: 61); Philonyx fulvicollis combination and incorrect spelling by Smith (1910: 598); Cynips (Philonix) fulvicollis var. fulvicollis form fulvicollis combination by Kinsey (1930: 262); Cynips (Philonix) fulvicollis combination by Kinsey (1936: 207).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF86CE384E9D51E1A859ABB3.taxon	materials_examined	Material examined: One sexual female and one male labelled as “ CANADA: Alberta, Edmonton, 3761 - 20 Street, 53.47371 ° N, 113.37202 ° W, GPS ± 10 m, reared 1. vi. 2008 ex bud galls on Quercus macrocarpa, S. Digweed, det. Philonix fulvicollis Fitch, sexual generation female, S. Digweed 2008 ”. The female and male specimens have been deposited at the PHDNRL.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF86CE384E9D51E1A859ABB3.taxon	diagnosis	Diagnosis. Sexual Philonix most closely resemble sexual Acraspis. In Acraspis the postgenal bridge shorter than the height of occipital foramen, the mesoscutum entirely smooth, notaulus incomplete, the mesoscutellum rounded, uniformly rugose; the metasoma only slightly longer than high. In Philonix the postgenal bridge as high as the occipital foramen, most of the mesoscutum smooth, except delicately alutaceous in the anterior part between notauli, notaulus complete, the mesoscutellum elongated with a smooth, glabrous surface in anterior half; the metasoma much higher in lateral view.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF86CE384E9D51E1A859ABB3.taxon	description	Description. Sexual female (Figs 183 – 187, 192 – 196, 200 – 201). Body uniformly reddish brown, antenna slightly lighter, palpi yellow, legs uniformly yellow. Head alutaceous, with sparse white setae, denser on lower face, with distinct rows of setae along inner margin of eye; slightly broader than high and as broad as mesosoma in frontal view, 1.9 × as broad as long in dorsal view. Gena alutaceous, not broadened behind eye in frontal view, 0.5 × width of transverse diameter of eye in lateral view. Malar space alutaceous, with delicate striae radiating from clypeus and reaching eye margin; eye 3.8 × as high as length of malar space. Inner margins of eyes parallel. POL 1.5 × as long as OOL, OOL 1.9 × as long as diameter of lateral ocellus and 1.7 × as long as LOL, all ocelli ovate, of same size. Transfacial distance as long as height of eye; diameter of antennal torulus shorter than distance between them, distance between torulus and eye as long as diameter of torulus; lower face alutaceous, slightly elevated median area alutaceous. Clypeus rectangular, nearly 2.0 × as broad as high, alutaceous, with a few long setae along ventral edge; ventrally rounded, slightly emarginate, without median incision; anterior tentorial pit rounded, distinct, epistomal sulcus distinct, clypeo-pleurostomal line well impressed. Frons and interocellar area uniformly alutaceous, without striae or setae. Vertex, occiput, postocciput and postgena alutaceous, with sparse white setae; posterior tentorial pit large, ovate, area below impressed; occipital foramen slightly higher than height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which are not united and diverge toward occipital foramen, postgenal bridge anteriorly as broad as occipital foramen. Antenna longer than head + mesosoma, with 12 flagellomeres, scape only slightly longer than pedicel, pedicel slightly longer than broad, flagellomeres slightly broader towards apex; F 1 2.0 × as long as pedicel and 1.2 × as long as F 2; F 2 slightly longer than F 3; F 3 slightly longer than F 4, F 5 = F 6, all subsequent flagellomeres equal in length; F 12 slightly longer than F 11; placodeal sensilla on F 3 – F 12, absent on F 1 – F 2. Mesosoma slightly longer than high. Pronotum smooth, glabrous, with sparse setae in dorsolateral part, with delicate short parallel striae along posterior edge; propleuron smooth, glabrous, with sparse white setae. Mesoscutum smooth, glabrous, delicately alutaceous at the anterior end between notauli, with a few white setae along notauli, slightly longer than broad (greatest width measured across mesoscutum level with base of tegulae). Notaulus complete, reaching pronotum, well impressed along entire length, with smooth bottom, posteriorly slightly converging; anterior parallel line inconspicuous, marked with slightly impressed narrow area, extending to 1 / 6 of mesoscutum length; median mesoscutal line absent; parapsidal line indistinct, circumscutellar carina narrow but distinct, reaching notaulus. Mesoscutellum quadrangular, longer than broad, with parallel sides, uniformly dull coriaceous, with strong irregular rugae along sides and posteriorly; posteriorly rounded, overhanging metanotum. Anterior part of mesoscutellum impressed, smooth, glabrous, in the form of a semilunar impression, with rugose triangular central area dividing anterior part into two halves. Mesopleuron smooth, glabrous, with a few white setae on the most posteroventral part; mesopleural triangle smooth, glabrous, with a few delicate longitudinally orientated striae and a few setae; dorsal and lateral axillar areas smooth, glabrous, with sparse setae; subaxillular bar smooth, glabrous, triangular, gradually higher toward posterior end, at posterior end as high as height of metanotal trough; metapleural sulcus distinct, reaching mesopleuron at half of its height. Metascutellum smooth, 0.5 × as long as height of smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous, without setae; central propodeal area lyre-shaped, smooth, glabrous, with a few short rugae, in posterior half the rugae continue onto nucha; lateral propodeal carinae distinct, complete, bent slightly outwards in posterior 1 / 3; lateral propodeal area smooth, glabrous, with sparse long white setae. Nucha with parallel longitudinal rugae dorsally and laterally. Tarsal claws simple, without basal lobe. dorsal view. Forewing longer than body, hyaline, with short cilia on margin, veins brown, radial cell open, 3.8 x as long as broad; R 1 and Rs nearly reaching wing margin; areolet triangular, closed and distinct. Rs + M narrow, indistinct, reaching basalis slightly below mid height. Metasoma as long as head + mesosoma, as long as high in lateral view; 2 nd metasomal tergum extending to 1 / 3 length of metasoma in dorsal view, with a few setae laterally, without micropunctures; subsequent terga smooth, glabrous, without micropunctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium short, longer than broad in ventral view, with some long setae ventrally which extend far beyond apex of spine. Body length 1.9 mm (n = 1). Male (Figs 188 – 191, 197 – 199, 202). Similar to female but head, antenna, mesosoma, metasoma uniformly brown; mandibles light brown, mouthparts and legs uniformly yellow. Head transverse in frontal view; ocelli bigger than in female; malar space with striae, radiating from clypeus and reaching eye margin; ocelli elevated. Antenna as long as body, with 13 flagellomeres; scape + pedicel as long as F 1, F 1 slightly curved and broadened apically, longer than F 2; F 13 shorter than F 12; placodeal sensilla on all flagellomeres. Metasoma shorter than mesosoma; 2 nd metasomal tergum in the form of a narrow long petiole; metasomal tergum 3 longer than all subsequent terga together; all terga smooth, without micropunctures. Body length 1.8 mm (n = 1). Gall. The sexual generation galls (Fig. 204) occur s ingly in terminal buds of twigs of Q. macrocarpa. The gall is a relatively robust, seed-like, sub-ovoid cell approximately 2 mm long, dark brown when mature with longitudinal striations. Occupies the entire bud when mature, causing stunting or suppression of shoots and leaves produced from the bud.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF86CE384E9D51E1A859ABB3.taxon	biology_ecology	Biology. See Fitch (1859), Weld (1922 a, 1926), and Kinsey (1930, 1936) for the biology of the asexual generation. Asexual generation galls (Fig. 203) in Kinsey’s (1936) “ fulvicollis ” complex have been recorded on leaves of Q. alba, Q. bicolor, Q. chapmanii, Q. gambelii, Q. lyrata, Q. macrocarpa, Q. michauxii, Q. muehlenbergii, and Q. stellata, (Kinsey 1930, 1936; Burks 1979). Asexual generation females only begin to emerge from galls in the autumn of the year following the year of gall formation (Kinsey 1930), and we recorded females emerging up to four years after the year of gall formation. Adult females of the asexual generation (from galls collected in Manitoba but reared in Edmonton) emerged during the second half of October and early November. Sexual generation galls matured in May in Edmonton, and adults emerged 1 – 3 June 2008. Galls of the asexual generation appeared on leaves in Edmonton in July.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF86CE384E9D51E1A859ABB3.taxon	distribution	Distribution. USA: New York, Michigan, Tennessee, Illinois, Kansas (Burks 1979); Canada: Manitoba, Ontario, Quebec, New Brunswick. Molecular taxonomy. Alternate sexual and asexual generations were proposed by Digweed (2010) and are herein confirmed on the basis of similarity in cytb and ITS 2 sequence data generated for four individuals (three asexual females, one sexual female). Cytb sequences were on average 1.04 % divergent (range 0.69 – 1.39 %; GenBank accessions OM 321652 – OM 321655). Two ITS 2 alleles were observed across the four individuals that differed by 0.20 % (OM 331841 – OM 331844).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF86CE384E9D51E1A859ABB3.taxon	discussion	Comments. Kinsey (1930) transferred Dryophanta pallipes Bassett, 1900 to Philonix and proposed it as the sexual generation of P. fulvicollis, although no justification for this synonymy was provided. Weld (1959) wrote that P. pallipes might be well a synonym of Acraspis gemula. Melika & Abrahamson (2002) compared the three specimens of P. pallipes (from Beutenmueller collection, USNM, Washington, DC) with the paratypes of the sexual generation of A. gemula and found no differences, thus Philonix pallipes was synonymized with A. gemula. Digweed (2010) treated Philonix gigas (Weld), Philonix insulensis (Kinsey), and Philonix nigra (Gillette) as synonyms of P. fulvicollis, although without formal synonymization. Molecular analysis, however, shows that P. fulvicollis and P. nigra are distinct species. Two cytb haplotypes obtained from P. nigra (GenBank MW 388894, OM 321656) were on average 3.46 % divergent (range 3.00 – 3.93 %) from cytb haplotypes of P. fulvicollis.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF8CCE464E9D563AA909AA47.taxon	description	Figs 206 – 223, 225 Synonyms: Cecidomyia poculum Osten Sacken (1862: 201), galls. Xystoteras poculum combination and description of adult females by Weld (1922 a: 7); Phylloteras poculum combination by Lyon (1993: 133).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF8CCE464E9D563AA909AA47.taxon	materials_examined	Material examined: Five males and five females of the sexual generation, labelled as “ CANADA: Alberta, Edmonton, Brennan Crescent, 53.51746 ° N, - 113.682957 ° W, Google Maps, bud / shoot / ament galls collected 21. v. 2018 ex Quercus macrocarpa, S. Digweed, reared 24 - 30. v. 2018 ”. Specimens have been deposited at the USNM, PHDNRL, RAM.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF8CCE464E9D563AA909AA47.taxon	diagnosis	Diagnosis. This is the first sexual generation to be described for Phylloteras. Lyon (1993) gave a key to adults and galls of the asexual generation.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF8CCE464E9D563AA909AA47.taxon	description	Description. Asexual female (Fig. 205). Apterous, ant-like. Head, antenna, mesosoma, legs entirely black; metasoma dark brown. Sexual female (Figs 206 – 210, 216 – 221). Head and mesosoma black, metasoma dark brown to black; mandibles dark brown, palpi yellow; scape, pedicel, F 1 and F 2 yellow, all subsequent flagellomeres uniformly dark brown; legs yellow with brown coxae. Head alutaceous, with sparse white setae on lower face; nearly rounded, only 1.2 × as broad as high and broader than mesosoma in frontal view, 2.0 × as broad as long in dorsal view. Gena alutaceous, not broadened behind eye in frontal view, 1 / 3 width of transverse diameter of eye in lateral view. Malar space alutaceous, without malar sulcus or striae; eye 3.9 × as high as height of malar space. Inner margins of eyes slightly converging ventrally. POL 2.0 × as long as OOL; OOL 1.3 × as long as diameter of lateral ocellus and slightly shorter than LOL; all ocelli ovate, of same size. Transfacial distance as long as height of eye; diameter of antennal torulus slightly longer than distance between them, distance between torulus and eye as long as diameter of torulus. Lower face nearly as high as height of frons, alutaceous, without striae, with white scattered setae. Clypeus rounded, nearly as long as broad, alutaceous, with a few setae scattered all over; ventrally rounded, emarginate, without median incision; anterior tentorial pit small, indistinct; epistomal sulcus impressed, distinct; clypeo-pleurostomal line semi-lunar, impressed, distinct. Frons and interocellar area alutaceous, without setae. Vertex, occiput, postocciput, postgena alutaceous; posterior tentorial pit large, ovate, area below impressed; occipital foramen shorter than height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which are widely separated and diverge toward occipital foramen, postgenal bridge only slightly broader than high anteriorly. Antenna longer than head + mesosoma, with 12 flagellomeres, scape slightly longer than pedicel; pedicel longer than broad, F 1 1.4 × as long as pedicel and as long as F 2, F 2 1.4 × as long as F 3, F 3 = F 4, F 5 to F 11 equal in length; F 12 longer than F 11; placodeal sensilla on F 3 – F 12, absent on F 1 – F 2. Mesosoma slightly longer than high. Pronotum dorsally alutaceous, with white setae along anterior rim in frontal view; smooth, matte laterally, with impressed foveolate sculpture along anterior edge; propleuron alutaceous, matte. Mesoscutum smooth, glabrous, with a few scattered setae only along sides; slightly longer than broad (greatest width measured across mesoscutum level with base of tegulae). Notaulus, anterior parallel line, median mesoscutal line and parapsidal line absent; circumscutellar carina present but narrow. Transscutal articulation absent, mesoscutum fused with mesoscutellum; as a result of fusion of the mesoscutum and mesoscutellum, the boundary between these structures is not straight but slightly curved towards the mesoscutum; the mesoscutum is emarginate posterolaterally and slightly elevated above the dorsoaxillar area. Mesoscutellum longer than broad, ovate, posteriorly rounded, uniformly dull rugose, with more delicate coriaceous sculpture toward anterior end; strongly overhanging metanotum. Mesoscutellar foveae in the form of a narrow semilunar impression with smooth glabrous bottom, without central longitudinal carina or elevation. Mesopleuron, including speculum smooth, matte, with short delicate striae, together the striae form a line extending across mesopleuron from posteroventral to anterodorsal edges; mesopleural triangle smooth, matte; dorsal and lateral axillar areas smooth, matte, with a few setae; subaxillular bar smooth, glabrous, at posterior end as high as height of metanotal trough; metapleural sulcus reaching mesopleuron in upper 1 / 3, delimiting broad smooth and wrinkled area along mesopleuron; upper part of sulcus hardly traceable. Metascutellum narrow, smooth, as high as height of smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous, without setae; propodeum smooth, glabrous; central propodeal area broad, smooth, matte, with a few irregular interrupted rugae; lateral propodeal carinae strong, elevated, bent outwards in posterior 1 / 3 of propodeum height; lateral propodeal area smooth, glabrous, without setae, with irregular short rugae, with dense long white setae in most lateroventral parts. Nucha short, with irregular rugae dorsally and laterally. Tarsal claws with acute basal lobe. Forewing longer than body, hyaline, margin with long cilia, veins dark brown, radial cell open, 5.25 × as long as broad; R 1 and Rs reaching wing margin and extending along margin; areolet small, triangular, distinct; Rs + M dark brown, visible along 2 / 3 of its length, projection reaching basalis at its mid height. Metasoma longer than head + mesosoma, slightly higher than long in lateral view; 2 nd metasomal tergum extending to nearly 1 / 3 length of metasoma in lateral view, with a few setae anterolaterally; all terga smooth, matte, without micropunctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium needle-like, 3.5 × as long as broad in ventral view, with a few setae ventrally. Body length 1.8 – 1.9 mm (n = 3). Male (Figs 211 – 215, 222 – 223). Similar to female, also in coloration, but head with larger eyes, interocellar area and ocelli strongly elevated; ocelli bigger than in female; antenna slightly longer than body, with 13 flagellomeres; scape short, as long as pedicel, pedicel slightly longer than broad; scape + pedicel slightly longer than F 1, F 1 slightly curved, not broadened apically, longer than F 2; F 2 longer than F 3; F 4 and F 5 nearly equal in length, both shorter than F 3; F 5 twisted, curved, apically broader; F 7 – F 12 with distinct long setae apically; F 13 longer than F 12; placodeal sensilla on all flagellomeres. Mesosoma and nucha slightly longer than in female; disc of mesoscutellum smooth, glabrous; metasoma as long as mesosoma; 2 nd metasomal tergum extending to half the length of metasoma in dorsal view; all terga smooth, matte, without micropunctures. Body length 1.7 mm (n = 3). Gall. Sexual galls (Fig. 225) are exposed, thin-walled, seed-like, ovoid, brown chambers in buds, catkins, and proximal regions of immature leaf midribs and laminae. Sometimes with a sparse pubescence of fibrous, tan hairs; sometimes these hairs are more abundant as a distal tuft. Adults emerge through holes chewed in the distal half of galls. Aging galls can persist on buds and leaves into autumn. Galls of the asexual generation (Fig. 224) on Q. alba are as described by Osten Sacken (1862) and Weld (1922 a).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF8CCE464E9D563AA909AA47.taxon	biology_ecology	Biology. See Weld (1922 a) for the biology of the asexual generation. Galls of the asexual generation became evident on bur oak leaves in Edmonton in July, and were fully mature in September, when many separate easily from the leaf lamina, falling to the ground. Galls are susceptible to drying, and rearing experience suggests that desiccated galls produce no emergents. Females of the asexual generation overwinter and pass the entire subsequent summer in their galls, emerging in late autumn and winter – generally from late October to December with a peak in mid-November, but when a large number of galls was reared over winter 2016 - 2017 in Edmonton, a small number of females emerged in all months from January through April 2017. Weld (1922 a) reared adults in Evanston, Illinois, in March. The brachypterous females were observed in Edmonton to crawl up through any accumulated snowpack, climb bur oak stems, and oviposit in buds. Emergence and adult female activity appeared delayed in autumn during periods of warmer weather, and was observed during a period of sustained temperatures ranging from - 10 ° C to - 17 ° C during early January 2017 in Edmonton. Galls of the sexual generation became apparent in early May in Edmonton, with adults emerging from mid-May through mid-June. This species has not been previously recorded from Q. macrocarpa; earlier records are from Q. alba and Q. montana. Another congeneric species, Phylloteras volutellae (Ashmead, 1897), is also found on Q. macrocarpa.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFF8CCE464E9D563AA909AA47.taxon	distribution	Distribution. USA: Virginia, New York, Vermont, New Hampshire, Illinois, Missouri (Burks 1979). Canada: Manitoba, Alberta. Molecular taxonomy. Alternating generations were matched using DNA data, with six individuals (four asexual females, two sexual females) sequenced for both cytb and ITS 2. Cytb sequences were on average 0.85 % divergent (range 0 – 1.62 %; GenBank accessions MZ 935719, OK 041500, OM 321657 – OM 321660). Two ITS 2 alleles were observed that differed only by a 5 bp indel (OM 331845 – OM 331850); four Canadian samples (sexual and asexual females) shared the same ITS 2 allele while the other allele was present in two North Carolina asexual females.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFF3CE404E9D55F1A9E3AFEB.taxon	description	Figs 226 – 235, 237 Synonyms: Dryophanta carolina Ashmead (1887: 145), asexual females, galls. Sphaeroteras carolina (Burks 1979). Atrusca carolina (Melika & Abrahamson 2002).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFF3CE404E9D55F1A9E3AFEB.taxon	materials_examined	Material examined: 2 females as “ USA, Florida, Manatee Co., Lake Manatee SRA, leg. J. Nicholls, 2008.04.16. FL 1097, spFLb 4; ex Quercus chapmanii ”. One female has been deposited at the USNM, one female at the PHDNRL.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFF3CE404E9D55F1A9E3AFEB.taxon	diagnosis	Diagnosis. No other sexual generation Sphaeroteras species are known from Florida on Q. chapmanii, especially in spring. The sexual gall resembles those of A. kingi and A. dumosae. However, those two species are known only from California, and gall other species of oaks which are distributed only on the Pacific slope.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFF3CE404E9D55F1A9E3AFEB.taxon	description	Description. Sexual female (Figs 226 – 235). Head, mesosoma light brown, metasoma darker; antenna dark brown; mouthparts yellow, legs yellow, except dark brown tibiae and tarsi. Head smooth, shiny, polished, without surface sculpture, with sparse white setae on frons and lower face; strongly transverse, 1.4 × as broad as high and broader than mesosoma in frontal view, 2.4 × as broad as long in dorsal view. Gena smooth, not broadened behind eye in frontal view, narrower than transverse diameter of eye in lateral view. Malar space smooth, without striae, eye 3.5 × as high as height of malar space. Inner margins of eyes parallel. POL 1.7 × as long as OOL; OOL 1.5 × as long as diameter of lateral ocellus and 1.4 × as long as LOL; all ocelli slightly ovate, of same size. Transfacial distance slightly longer than height of eye; diameter of antennal torulus 1.4 × as long as distance between them, distance between torulus and eye 1.1 × as long as diameter of torulus. Lower face smooth, polished, with white setae, without striae; median area smooth, not elevated. Clypeus nearly quadrangular, smooth, polished, with a few setae, ventrally not emarginate, without median incision; anterior tentorial pit small, rounded, inconspicuous; epistomal sulcus and clypeo-pleurostomal line narrow, slightly impressed. Frons smooth, with a few white setae; area under central ocellus smooth, glabrous; interocellar area smooth, with some setae. Vertex smooth; occiput smooth, without parallel striae; postocciput smooth, glabrous; postgena smooth, without setae; posterior tentorial pit large, ovate, area below impressed; occipital foramen about as high as height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which diverge strongly toward occipital foramen, postgenal bridge anteriorly broader than high. Antenna longer than head + mesosoma, with 12 flagellomeres, pedicel slightly longer than broad, F 1 2.8 × as long as pedicel and 1.1 x as long as F 2, F 2 slightly longer than F 3, subsequent flagellomeres gradually shorter until F 11, F 12 slightly longer than F 11; placodeal sensilla on F 4 – F 11. Mesosoma slightly longer than high, with sparse short white setae. Pronotum smooth, with white setae; propleuron smooth, glabrous. Mesoscutum smooth, with sparse white setae; slightly longer than broad (greatest width measured across mesoscutum level with base of tegulae). Notaulus complete, reaches pronotum, deep, broad, with smooth, glabrous bottom, posteriorly broader and strongly converging; anterior parallel line invisible; median mesoscutal line and parapsidal line absent; circumscutellar carina broad, smooth along tegula. Mesoscutellum broader than long, posteriorly rounded, uniformly rugose, overhanging metanotum, with strong, uniformly distributed white setae. Mesoscutellar foveae separated by narrow elevated smooth central carina, transverse, 1.8 × as broad as high, with smooth, glabrous bottom. Mesopleuron and speculum smooth, with white setae; mesopleural triangle delicately coriaceous, without striae, with dense white setae; dorsal and lateral axillar areas smooth, with white setae; subaxillular bar smooth, glabrous, at posterior end as high as height of metanotal trough; metapleural sulcus reaching mesopleuron at half of its height, delimiting broad area along mesopleuron. Metascutellum coriaceous, as high as height of smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous, with dense setae; central propodeal area smooth, glabrous, without striae; lateral propodeal carinae bent outwards at mid-height of propodeum; lateral propodeal area smooth, glabrous, with dense white setae. Nucha short, with delicate sulci dorsally and laterally. Tarsal claws simple, without basal lobe. Forewing longer than body, hyaline, margin with short cilia, veins dark brown, with dark stripes along all veins, radial cell open, 1.8 × as long as broad; R 1 and Rs nearly reaching wing margin; areolet small, triangular, distinct, Rs + M projection reaching basalis slightly below its mid height. Metasoma as long as head + mesosoma, nearly as long as high in lateral view; 2 nd metasomal tergum extending to more than half the length of metasoma in dorsal view, with white setae scattered all over the tergum; all terga smooth, glabrous, without micropunctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium short, slightly longer than broad in ventral view, with long setae ventrally which extend beyond apex of spine. Body length 3.8 – 3.9 mm (n = 2). Males unknown. Gall. Sexual galls (Fig. 237) on small twigs. Initially red / yellow, becoming darker as they mature, with pale pubescence, pear-shaped, tip bent to one side, 6 – 7 mm long, 3 mm diameter at widest point.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFF3CE404E9D55F1A9E3AFEB.taxon	biology_ecology	Biology. Asexual detachable rounded galls (Fig. 236) on leaves on Q. alba, Q. chapmanii and Q. stellata (Burks 1979). Sexual galls mature in April; adults emerge soon afterwards; to date found only on Q. chapmanii.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFF3CE404E9D55F1A9E3AFEB.taxon	distribution	Distribution. USA: North Carolina, Florida, Missouri, Arkansas, Texas (Burks 1979). Molecular taxonomy. Alternating generations were matched using DNA data, with four individuals (three asexual females, one sexual female) sequenced for both cytb and ITS 2. Cytb sequences were on average 1.12 % divergent (range 0 – 2.08 %; GenBank accessions OM 321661 – OM 321664); ITS 2 sequences identical (OM 331851 – OM 331854).	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
03E987BFFFF3CE404E9D55F1A9E3AFEB.taxon	discussion	Comments. The genus Sphaeroteras was described by Ashmead (1897). Later, Mayr (1902), Beutenmueller (1909), Dalla Torre & Kieffer (1910) treated Sphaeroteras as a synonym of Biorhiza Westwood, 1840. Weld (1951) re-established the validity of this genus and transferred in S. carolina from Diplolepis Geoffroy, 1762. Melika & Abrahamson (2002) again synonymized Sphaeroteras with Biorhiza. This species was erroneously transferred to Atrusca Kinsey, 1930, by Melika & Abrahamson (2002). Pujade-Villar et al. (2018) re-established Sphaeroteras; however, they did not mention S. carolina in the list of taxa returned to that genus. Phylogenetic reconstruction puts this species in the same clade as some species of Antron (Nicholls unpubl. data), but this clade does not contain all current Sphaeroteras species nor all current Antron species, so delimitation of both genera requires further work.	en	Nicholls, James A., Melika, George, Digweed, Scott C., Stone, Graham N. (2022): Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini). Zootaxa 5145 (1): 1-79, DOI: 10.11646/zootaxa.5145.1.1, URL: http://zoobank.org/1f909f98-7d98-4930-93d8-dd55008d9c76
