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Loxaulus Mayr, 1881 Currently 14 species of Loxaulus are known from America north of Mexico, which are associated with different Quercus sections (Dailey 1977, Burks 1979, Dailey & Sprenger 1983, Melika & Abrahamson 2000): five species gall section Quercus, one species galls section Virentes, one galls section Lobatae, three gall section Protobalanus and for the other four species the hosts are unknown. Four new species of Loxaulus have recently been described from Panama and Mexico (Medianero et al. 2011, Pujade-Villar et al. 2014b), three from section Quercus oaks and one from a section Lobatae oak (although the latter record must be reviewed). As is the case with Dryocosmus, this variation in host section implies this genus may be polyphyletic. Herein we describe one new species, L. virginianae Melika & Buss, which is associated with Q. virginiana (in the oak section Virentes)., Published as part of Melika, George, Nicholls, James A., Abrahamson, Warren G., Buss, Eileen A. & Stone, Graham N., 2021, New species of Nearctic oak gall wasps (Hymenoptera: Cynipidae, Cynipini), pp. 1-131 in Zootaxa 5084 (1) on page 86, DOI: 10.11646/zootaxa.5084.1.1, http://zenodo.org/record/5800716, {"references":["Mayr, G. (1881) Die Genera der gallenbewohnenden Cynipiden. Jahresberichte der Communal-Oberrealschule im I. Bezirke, Wien, 20, 1 - 38.","Dailey, D. C. (1977) Elevation of Loxaulus brunneus variety atrior (Kinsey) to full species status (Hymenoptera: Cynipidae). The Pan-Pacific Entomologist, 53 (2), 145 - 146.","Burks, B. D. (1979) Superfamily Cynipoidea. In: Krombein, K. V., Hurd, P. D. Jr., Smith, D. R. & Burks, B. D. (Eds.), Catalog of Hymenoptera in America North of Mexico. Vol. 1. Symphyta and Apocrita. Smithsonian Institution Press, Washington, D. C., pp. 1045 - 1107.","Dailey, D. C. & Sprenger, C. M. (1983) Gall-inducing cynipid wasps from Quercus dunnii Kellogg (Hymenoptera). The Pan- Pacific Entomologist, 59 (1 - 4), 42 - 49.","Melika, G. & Abrahamson, W. G. (2000) Review of the cynipid gall wasps of the genus Loxaulus Mayr (Hymenoptera: Cynipidae), with descriptions of new species. Proceedings of the Entomological Society of Washington, 102 (1), 198 - 211.","Medianero, E., Nieves-Aldrey, J. L. & Melika, G. (2011) Two new Neotropical species of oak gall wasps of the genus Loxaulus Mayr (Hymenoptera: Cynipidae: Cynipini) from Panama. Zootaxa, 2811, 37 - 46."]}

Loxaulus virginianae Melika & Buss, sp. nov. Figs. 317–328 urn:lsid:zoobank.org:act: A7C8E359-5013-4D15-B686-02589B18F2CB Type material: HOLOTYPE female “ USA, Alabama, Grand Bay, Oak Hill Tree Farm, coll. 18-26 May 2013, leg. Albert van Hoogmoed; ex galls in twigs of Quercus virginiana ”; PARATYPES: 4 females with the same labels as the holotype. The holotype is deposited at the USNM, 4 females at the PHDNRL. Etymology. Named after the host plant, Q. virginiana. Diagnosis. A key to all 14 Loxaulus species known from America north of Mexico was published by Melika & Abrahamson (2000). There are two species of Loxaulus, L. pattersoni (Kinsey, 1922) and L. virginianae, which are associated with Q. virginiana. Loxaulus pattersoni, known from Texas, induces galls on roots of Q. virginiana. In L. pattersonae the forewing margin has dense cilia, antenna with 13 flagellomeres, while in L. virginianae the forewing margin is without cilia, antenna with 12 flagellomeres. Loxaulus virginianae somewhat resembles L. laeta Pujade-Villar, 2014 described from Mexico (Pujade-Villar et al. 2014b), but the latter has antennae with only 10 flagellomeres, the forewing with cilia on the margin and the body is dark brown to black, while L. virginianae has antennae with 12 flagellomers, the forewing without cilia on margins, and the body is uniformly reddish brown. Description. Asexual female (Figs. 317–327). Head, mesosoma, metasoma, antennae, mouthparts, legs all uniformly reddish brown. Head delicately reticulate, with a few white setae, 1.2× as broad as high and slightly broader than mesosoma in frontal view, 2.0× as broad as long in dorsal view. Gena reticulate, broadened behind eye in frontal view, broader than transverse diameter of eye in lateral view. Malar space reticulate, with malar sulcus, eye 3.2× as high as length of malar space. Inner margins of eyes parallel. POL 2.0× as long as OOL, OOL 1.8× as long as diameter of lateral ocellus, 1.2× as long as LOL, all ocelli ovate, of same size. Transfacial distance as long as height of eye, diameter of antennal torulus 1.9× as long as distance between them, distance between torulus and eye 1.2× as long as diameter of torulus; lower face and slightly elevated median area alutaceous, with a few short white setae, without striae. Clypeus trapezoid, slightly broader than high, smooth, glabrous; ventrally rounded, not emarginate, without median incision and with a few long setae; anterior tentorial pit large, deep, rounded, epistomal sulcus and clypeo-pleurostomal line distinct. Frons and interocellar area reticulate, without setae. Vertex and occiput reticulate, postocciput smooth, glabrous, postgena reticulate, with a few setae; posterior tentorial pit large, ovate, area below impressed, occipital foramen 1.6× as high as height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which are not united, running parallel to half the length of postgenal bridge, diverge in anterior half; anterior half of postgenal bridge as broad as occipital foramen. Antenna slightly longer than head+mesosoma, with 12 flagellomeres (in some paratypes suture between F11 and F12 indistinct), pedicel 1.8× as long as broad, F1 1.6× as long as pedicel and equal to F2; F2=F3=F4; F4 to F11 equal in length, slightly shorter than F3; F12 slightly shorter than F11; placodeal sensilla on F3–F12. Mesosoma longer than high, with rare short white setae. Pronotum uniformly reticulate, invaginated anterior margin of pronotum foveolate, dorsally with a few sparse setae. Propleuron delicately coriaceous, glabrous. Mesoscutum uniformly reticulate, longer than broad (greatest width measured across mesoscutum at level of base of tegulae). Notaulus incomplete, deep, with smooth, glabrous bottom in posterior 1/3 of mesoscutum, hardly traceable in anterior 2/3, converging posteriorly; anterior parallel line indistinct, extends to 1/4 of mesoscutum length; parapsidal line visible in posterior 2/3 of mesoscutum, marked with smooth, glabrous areas; median mesoscutal line absent or in the form of a short triangle posteriorly; circumscutellar carina narrow, reaching slightly above tegula. Mesoscutellum rounded slightly longer than broad, with net of irregular rugae, posteriorly rounded. Mesoscutellar foveae not separated medially, in the form of a transverse anterior impression, with smooth glabrous bottom and parallel longitudinal rugae. Mesopleuron and speculum glabrous, uniformly delicately reticulate, without setae; mesopleural triangle glabrous, with some transverse delicate rugae; dorsal and lateral axillar areas reticulate; subaxillular bar smooth, glabrous, narrow, with subparallel margins, at posterior end slightly shorter than height of metanotal trough; metapleural sulcus delimiting broad rugose, glabrous area, reaching mesopleuron in upper 1/3, upper part of sulcus indistinct. Metascutellum coriaceous, 2.0× as high as height of smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous, without setae, with distinct longitudinal parallel rugae; central propodeal area smooth, glabrous, without setae, broader anteriorly, narrowing posteriorly, with strong complete central longitudinal carinae and numerous transverse rugae dividing central propodeal area into rectangular units; lateral propodeal area glabrous, without setae, with net of irregular rugae. Nucha with net of short irregular delicate sulci dorsally and laterally. Tarsal claws simple, without basal lobe. Forewing longer than body, hyaline, with dark brown veins, margin without cilia, with dark brown stripe going across radial cell and extend to Cu1b, radial cell open, 3.5× as long as broad, R1and Rs nearly reaching wing margin, areolet triangular, well-delimited, Rs+M distinct along full length, reaching basalis at mid height. Metasoma longer than head+mesosoma, slightly longer than high in lateral view; second metasomal tergite extending to 1/4 length of metasoma in dorsal view, with a few white setae anterolaterally, without micropunctures, subsequent tergites glabrous, with few rare indistinct micropunctures. Hypopygium with delicate dense micropunctures, prominent part of ventral spine of hypopygium narrowing slightly towards apex, more than 2.0× as long as broad in ventral view, with few long setae extend beyond apex of spine. Body length 2.7–3.1 mm (n = 5). Gall. (Fig. 328). Galls are tiny cryptic cells hidden under the bark of twigs, with little or no visible swelling of the twig. The galls are 2 mm long and consist only of the larval chamber. The galls are commonly induced in a longitudinal cluster and are most easily detected by the emergence holes in the bark. Biology. Only an asexual generation is known, which induces stem galls in twigs of Q. virginiana. Galls mature in May, adults emerge soon afterwards. Distribution. USA, Alabama, Grand Bay.

Neuroterus aliceae Melika, Nicholls & Stone, sp. nov. Figs. 344–367 urn:lsid:zoobank.org:act: 23F3C805-6588-4327-8BDA-6FA255705A33 Alternate asexual and sexual generations are described. Etymology. Named after Alice I. Nicholls, daughter of Dr. J. A. Nicholls (Institute of Evolutionary Biology, University of Edinburgh, UK and Australian National Insect Collection, CSIRO, Canberra). Asexual generation Figs. 344–351, 367 Type material: HOLOTYPE female “ USA, Arizona, Molino Basin campground, Santa Catalina Mtns., leg. J. Nicholls, 2007.10.28. Code AZ632, spAZl7; ex Quercus oblongifolia ”. PARATYPES: 26 females with the same labels as the holotype, 3 with the same collecting code, others with the following codes: AZ629— 1 female, AZ637— 1 female, AZ638— 3 females, AZ645— 3 females, AZ651— 1 female, AZ652— 5 females, AZ654— 1 female, AZ656— 1 female, AZ657— 2 females, AZ658— 1 female, AZ660— 1 female, AZ662— 1 female, AZ664— 2 females. The holotype and 3 female paratypes are deposited at the USNM, 23 female paratypes at the PHDNRL. Diagnosis. Neuroterus alicae belongs to Kinsey’s subgenus Diplobius. The asexual generation of Neuroterus aliceae induces dehiscent leaf galls on Q. oblongifoliae. The only previously known Neuroterus species from the US southwest to induce detachable leaf galls, N. argentatus Weld, 1944 asexual generation, induces small, flat, rounded spangle galls on the underside of the leaves of Q. gambelii, Q. oblongifolia and Q. turbinella, which are obviously different from the aggregated brown fluffy leaf galls induced by N. aliceae. Description. Asexual female (Figs. 344–351). Head, mesosoma and metasoma dark brown to black; mandibles, maxillary and labial palpi light brown; antennae brown, gradually darker towards apex; legs dark brown. Head alutaceous, with sparse white setae, denser on lower face and postgena, 1.2× as broad as high and broader than mesosoma in front view, 2.2× as broad as long in dorsal view. Gena alutaceous, slightly broadened behind eye in frontal view, as broad as transverse diameter of eye in lateral view. Malar space alutaceous, with malar sulcus; eye 3.7× as high as length of malar space. Inner margins of eyes parallel. POL 2.7× as long as OOL; OOL 1.4× as long as diameter of lateral ocellus and slightly shorter than LOL; all ocelli ovate, of same size. Transfacial distance length equal to height of eye; diameter of antennal torulus 1.3× as long as distance between them, distance between torulus and eye 1.3× as long as diameter of torulus; lower face and slightly elevated median area uniformly alutaceous, without striae, with some setae. Clypeus trapezoid, alutaceous, broader than high; ventrally rounded, emarginate, without median incision and with a few long setae; anterior tentorial pit large, rounded, epistomal sulcus and clypeo-pleurostomal line distinct, broad. Frons and slightly elevated interocellar area alutaceous, with short white setae. Vertex, occiput, postocciput, postgena alutaceous, with a few setae; occipital foramen slightly shorter than height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which are not united; postgenal bridge anteriorly as broad as occipital foramen. Antenna slightly longer than head+mesosoma, with 10 flagellomeres, pedicel 1.4× as long as broad, F1 1.5× as long as pedicel and 1.4× as long as F2, F2 slightly longer than F3, F3 shorter than F4, subsequent flagellomeres shorter, nearly equal in length, F12 slightly longer than F11, placodeal sensilla on F2–F12. Mesosoma slightly longer than high, with a few white setae. Pronotum uniformly alutaceous; propleuron smooth, glabrous, with few setae; anterior margin invaginated, smooth, glabrous, not foveolate. Mesoscutum uniformly alutaceous, with a few setae anteriorly; slightly longer than broad (greatest width measured across mesoscutum at level of base of tegulae). Notaulus, anterior parallel line, parapsidal line and median mesoscutal line absent; circumscutellar carina narrow, extending slightly above tegula. Mesoscutum emarginate posterolaterally, slightly elevated above dorsal axillar area, V-shaped posteriocentrally. Transscutal articulation absent. Mesoscutellum rounded, slightly broader than high, smooth, glabrous, with a few white short setae on disk; anteriorly with short and narrow strongly impressed smooth, glabrous area, not overhanging metanotum. Mesopleuron and speculum alutaceous, glabrous; mesopleural triangle smooth, glabrous, with some delicate rugae; dorsal and lateral axillar areas smooth, glabrous; subaxillular bar smooth, glabrous, triangular, at posterior end higher than height of metanotal trough; metapleural sulcus delimiting smooth, glabrous area, impressed, distinct, reaching mesopleuron in lower half, upper part of sulcus absent, instead some parallel rugae radiating from mesopleuron and reaching subaxillular bar. Metascutellum smooth, glabrous, as high as height of smooth, glabrous ventral impressed area, metanotal trough smooth, glabrous; central propodeal area ovate, broadest at mid height, glabrous, with delicate numerous interrupted rugae, which fill in the entire central propodeal area; no distinct lateral propodeal carinae; lateral propodeal area smooth, glabrous, without setae. Nucha smooth, glabrous, without rugae. Tarsal claws simple, without basal lobe. Forewing longer than body, hyaline, margin with long dense cilia, veins dark brown, radial cell open, 4.1× as long as broad, R1 and Rs nearly reaching wing margin, areolet triangular, well-delimited, Rs+M distinct along full length, reaching basalis at lower 1/3. Metasoma as long as head+mesosoma, taller than long in lateral view; second metasomal tergite occupying 1/2 the length of metasoma in dorsal view, without setae anterolaterally, without micropunctures, subsequent tergites 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 setae ventrally which do not extend beyond apex of spine. Body length 1.8–2.1 mm (n = 15). Gall. (Fig. 367). Brown fluffy leaf galls, consisting of an aggregated group of larval chambers ≥ 5 mm across, on midrib on the underside of leaf. Covered in a dense, somewhat woolly, coat of reddish-brown hairs. Similar in structure to Andricus pattoni (Bassett, 1881) from Florida. Sexual generation Figs. 352–366 Material examined. Seven females and 7 males “ USA, Arizona, Molino Basin campground, Santa Catalina Mtns., leg. J. Nicholls, 2008.04.09. Code AZ1946, spAZc3; ex Quercus oblongifolia ”; 3 males “ USA, Arizona, Molino Basin campground, Santa Catalina Mtns., leg. J. Nicholls, 2008.04.09. Code AZ1872, ex Quercus oblongifolia ”, and 3 males “ USA, Arizona, Molino Basin campground, Santa Catalina Mtns., leg. J. Nicholls, 2008.04.09. Code AZ1887, ex Quercus oblongifolia ”. Diagnosis. The sexual generation of N. aliceae induces catkin galls on Q. oblongifolia. The only known Neuroterus species from the US southwest which induces flower galls (also on white oaks) is N. floricomus Weld, 1957, sexual generation. The galls of N. floricomus are dense wooly enlargements of the catkin axis (Weld 1957b) and are clearly different from those of N. aliceae (Fig. 366). In N. floricomus the head of females is coriaceous, genae broadened behind eyes, the body of females is black, while in N. aliceae, head is alutaceous, genae are not broadened behind eyes, and entire body is brown. Description. Sexual female (Figs. 352–354, 357, 359–364). Head black, mesosoma and metasoma brown; mandibles, maxillary and labial palpi yellow; antennae brown; legs light brown to yellow. Head alutaceous, with sparse white setae, denser 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, with delicate parallel striae radiating from clypeus and reaching eye, malar sulcus absent or inconspicuous between striae; eye 4.6× as high as length of malar space. Inner margins of eyes parallel. POL 1.9× as long as OOL, OOL 1.5× as long as diameter of lateral ocellus and equal to LOL; all ocelli ovate, of same size. Transfacial distance equal to height of eye; diameter of antennal torulus 1.3× as long as distance between them, distance between torulus and eye nearly equal to diameter of torulus; lower face and slightly elevated median area uniformly alutaceous, without striae. Clypeus trapezoid, alutaceous, broader than high; ventrally rounded, emarginate, without median incision and with a few long setae ventrally; anterior tentorial pit large, rounded, epistomal sulcus and clypeo-pleurostomal line distinct, broad. Frons and slightly elevated interocellar area alutaceous, with a few short white setae. Vertex, occiput alutaceous, with rare setae; postocciput, postgena smooth, glabrous; occipital foramen as high as height of postgenal bridge; hypostomal carina emarginate, continuing into indistinct united postgenal sulci, postgenal bridge narrow. Antenna longer than head+mesosoma, with 12 flagellomeres (in some paratypes suture between F12 and F11 indistinct), pedicel subglobular; F1 1.7× as long as pedicel and 1.25× as long as F2, F2 slightly longer than F3, F3=F4, F5–F10 equal in length, F12 slightly longer than F11; placodeal sensilla on F2–F12. Mesosoma slightly longer than high, with a few white setae. Propleuron smooth, glabrous. Pronotum alutaceous, with a few setae and some delicate striae laterally; anterior margin invaginated, smooth, glabrous, foveolate. Mesoscutum alutaceous, with a few setae; longer than broad (greatest width measured across mesoscutum at level of base of tegulae). Notaulus impressed in posterior 2/3 of mesoscutum length, only a trace in anterior 1/3, anterior parallel line invisible, parapsidal line marked with impressed smooth, glabrous area; median mesoscutal line in the form of a short triangle; circumscutellar carina broad, extending above tegula. Mesoscutum emarginate posterolaterally, slightly elevated above dorsal axillar area. Transscutal articulation absent. Mesoscutellum slightly longer than broad, ovate, uniformly alutaceous, with some white short setae, posteriorly rounded, clearly projecting over metanotum. Mesoscutellar foveae in the form of a transverse, narrow impressed anterior area, with smooth, glabrous bottom. Mesopleuron and speculum alutaceous, glabrous, invaginated in lower half; mesopleural triangle smooth, glabrous, with a few white setae; dorsal and lateral axillar areas smooth, glabrous; subaxillular bar smooth, glabrous, with parallel margins, shorter than height of metanotal trough; metapleural sulcus distinct, reaching mesopleuron at half its height. Metascutellum smooth, glabrous, as tall as height of smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous; propodeum posterodorsally smooth, glabrous, without carinae, lateral propodeal carina present only at the most posterior end of propodeum. Nucha short, smooth, glabrous. Tarsal claws simple, without basal lobe. Forewing longer than body, hyaline, margin with long dense cilia, with dark brown veins, radial cell open, 4.3× as long as broad, R1 and Rs reaching wing margin; areolet triangular, well-delimited, Rs+M distinct along full length, reaching basalis in lower 1/3 of its height. Metasoma as long as head+mesosoma, shorter than high in lateral view; second metasomal tergite extending to 1/3 of metasoma length in dorsal view, with a few white setae anterolaterally, without micropunctures, subsequent tergites smooth, glabrous, without micropunctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium 4.4× as long as broad in ventral view, without setae. Body length 1.7–2.1 mm (n = 6). Male (Figs. 355–356, 358, 365). Similar to female but eye 1.2× as high as length of transfacial distance, eye 5.3× as high as length of malar space, POL 3.4× as long as OOL, LOL 1.75× as long as OOL; antenna with 12 flagellomeres, F1 slightly curved, placodeal sensilla on F1–F13. Body length 1.7–1.9 mm (n = 6). Gall. (Fig. 366). A whitish swelling surrounding individual flowers, with sparse very short pubescence. A swollen flower is 1–2 mm in diameter; multiple flowers per catkin are galled. Biology. Alternate asexual and sexual generations are known, both of which induce galls on Q. oblongifolia. Sexual catkin galls mature in April, adults emerge soon afterwards. The asexual fluffy leaf galls mature in October-November; adults overwinter in the galls and emerge the following spring. Alternate generations (and males/females within the sexual generation) were matched using DNA data. Five individuals were sequenced for cytb, with between 0 and 0.23% divergence apparent (GenBank OK346301 – OK346305), while four individuals showed at most 0.20% divergence for ITS2 (OK 350661 – OK350664). Distribution. USA, Arizona, Santa Catalina Mountains.

Neuroterus rosieae Melika, Nicholls & Stone, sp. nov. Figs. 406–421 urn:lsid:zoobank.org:act: 621BA670-0C0C-4835-96D6-D4D6438879A0 Type material: HOLOTYPE female “ USA, Arizona, 5km N of Payson. Leg. J. Nicholls. 2008.04.12. Code AZ1865, spAZb22; ex Quercus turbinella ”. PARATYPES: (6 females and 5 males): 3 females and 3 males with the same labels as the holotype; 1 female “ USA, Arizona, Lower Oak Creek Canyon, leg. J. Nicholls, 2008.04.11. Code AZ1933, ex Quercus arizonica ”; 2 females and 2 males “ USA, Arizona, Stoneman Lake Road at I17, 2008.04.10. Code AZ1964, leg. J. Nicholls, ex Quercus turbinella ”. The holotype and one male are deposited at the USNM, 6 females and 4 males at the PHDNRL. Etymology. Named after Rosie Stone, daughter of Prof. G.N. Stone (Institute of Evolutionary Biology, University of Edinburgh, UK). Diagnosis. Neuroterus rosieae belongs to Kinsey’s subgenus Diplobius. It possesses an unusual character, namely that the entire metasoma is covered with parallel longitudinal striae, with distinct punctures. No other Neuroterus species is known to have such peculiar metasomal sculpturing. Only N. stonei has some delicate longitudinal striae but only on the fifth tergum. Description. Sexual female (Figs. 406–408, 411, 413–418). Head, mesosoma and metasoma black; mandibles, maxillary and labial palpi, antennae brown; eyes silvery, legs dark brown. Head alutaceous, with sparse white setae, 1.25× 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 in frontal view, narrower than transverse diameter of eye in lateral view. Malar space with delicate parallel striae radiating from clypeus and reaching eye margin, malar sulcus absent. Eye 3.0× as high as length of malar space. Inner margins of eyes diverge ventrally. POL 2.3× as long as OOL; OOL as long as diameter of lateral ocellus and equal to LOL; all ocelli ovate, of same size. Transfacial distance shorter than height of eye; diameter of antennal torulus 1.2× as long as distance between them, distance between torulus and eye nearly equal to diameter of torulus; lower face and slightly elevated median area uniformly alutaceous, without striae. Clypeus large, trapezoid, alutaceous, broader than high; ventrally rounded, emarginate, without median incision and with a few long setae; anterior tentorial pits large, rounded, epistomal sulcus and clypeo-pleurostomal line distinct, broad. Frons and elevated interocellar area alutaceous, with some short white setae. Vertex, occiput, postocciput, postgena uniformly alutaceous, with some setae; occipital foramen longer than height of postgenal bridge; hypostomal carina emarginate, continuing into distinct postgenal sulci which are distinctly bent outwards; postgenal bridge broad. Antenna longer than head+mesosoma with 12 flagellomeres, pedicel 1.9× as long as broad; F1 1.6× as long as pedicel, F1=F2, F2 slightly longer than F3, F3=F4, subsequent flagellomeres slightly shorter and nearly equal in length, placodeal sensilla on F2–F12. Mesosoma slightly longer than high, with a few white setae. Propleuron alutaceous. Pronotum alutaceous, with a few setae and some delicate striae laterally; anterior margin invaginated, smooth, glabrous, foveolate. Mesoscutum alutaceous, with a few setae; longer than broad (greatest width measured across mesoscutum at level of base of tegulae). Notaulus and median mesoscutal line absent; anterior parallel lines, parapsidal lines hardly visible, marked with smooth, glabrous areas; circumscutellar carina broad, reaching above tegulae. Mesoscutum emarginate posterolaterally, slightly elevated above dorsal axillar area. Transscutal articulation absent. Mesoscutellum longer than broad, ovate, posteriorly rounded; uniformly alutaceous, with some white short setae, clearly overhanging metanotum. Mesoscutellar foveae in the form of a transverse, narrow semilunar impression, with smooth, glabrous bottom. Mesopleuron and speculum alutaceous; mesopleural triangle smooth, glabrous, with a few white setae; dorsal and lateral axillar areas alutaceous; subaxillular bar smooth, glabrous, with parallel margins, shorter than height of metanotal trough; metapleural sulcus indistinct, reaching mesopleuron at half of its height. Metascutellum smooth, glabrous, as high as height of smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous; propodeum posterodorsally smooth, glabrous, lateral propodeal carinae distinct, bent slightly outwards at mid height; central propodeal area smooth, glabrous, with some short striae. Nucha short, smooth, glabrous, with delicate subparallel longitudinal striae. Tarsal claws simple, without basal lobe. Forewing longer than body, hyaline, veins light brown, margin with long dense cilia, radial cell open, 3.2× as long as broad; R1 and Rs reaching wing margin; areolet triangular, well-delimited, Rs+M indistinct along entire length, reaching basalis at half height. Metasoma as long as head+mesosoma, as long as high in lateral view; second metasomal tergite extending to 1/4 of the metasoma length in dorsal view, smooth glabrous, with white setae anterolaterally; subsequent tergites with delicate parallel longitudinal striae, and punctures. Entire metasoma covered with distinct punctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium 2.0× as long as broad in ventral view, with setae along ventral margin. Body length 2.4–2.8 mm (n = 5). Male (Figs. 409–410, 412, 419). Similar to female but antennae dark brown to black, metasoma brown with long petiole; eyes and ocelli large, eye broader than transfacial distance in frontal view; height of eye 1.75× as long as than transfacial distance, malar space 0.1× the length of the height of eye; lateral ocellus nearly reaching eye; POL 21.0× as long as OOL, diameter of ocellus nearly 2.0× as long as LOL; interocellar area strongly elevated in frontal view; antenna with 13 flagellomeres, longer than length of body, F1 curved and broadened in distal half, placodeal sensilla on F1–F13. Body length 2.1–2.6 mm (n = 5). Gall. (Figs. 420–421). A small monolocular bud gall, pear-shaped with a spherical body that is drawn out to an apical point, greenish to purple, 3 mm across, 5 mm tall. The gall bears tiny bracts on its surface and is covered in dense, short, pale pubescence. Biology. Only a sexual generation is known, which induces bud galls on Q. arizonica and Q. turbinella. Galls mature in April, adults emerge soon afterwards. Within-species genetic diversity was estimated using cytb data from six individuals (males and females, collected from multiple locations; including some female paratypes); divergence among individuals ranged from 0 to 1.62% (GenBank OK346318 – OK346323). Distribution. USA, Arizona, along Mogollon Rim in area south of Flagstaff., Published as part of Melika, George, Nicholls, James A., Abrahamson, Warren G., Buss, Eileen A. & Stone, Graham N., 2021, New species of Nearctic oak gall wasps (Hymenoptera: Cynipidae, Cynipini), pp. 1-131 in Zootaxa 5084 (1) on pages 111-114, DOI: 10.11646/zootaxa.5084.1.1, http://zenodo.org/record/5800716

Andricus coconinoensis Melika, Nicholls & Stone, sp. nov. Figs. 77–88 urn:lsid:zoobank.org:act: 937DAF1F-8D20-49EA-AD69-840DE6304DE1 Type material: HOLOTYPE female “ USA, Arizona, 5 km N of Payson, leg. J. Nicholls, 2008.04.12. Code AZ1865, spAZb8; ex Q. turbinella ”. PARATYPES: three females and 2 males; 3 females with the same labels as the holotype, 2 males “ USA, Arizona, Molino Basin campground, Santa Catalina Mnts., leg. J. Nicholls, 2008.04.09. Code AZ1886, spAZb8; ex Q. oblongifolia ”. The holotype female is deposited at the USNM, 3 females and 2 males at the PHDNRL. Etymology. The species is named after the Coconino County in Arizona where it was first found. Diagnosis. Andricus coconinoensis induces a small bud gall on Q. turbinella and Q. oblongifolia (Fig. 88). Only two other described Andricus species are known to induce bud galls in the southwest of the USA: Andricus cellularius Gillette, 1892 and A. multicostatus Weld, 1952. In A. coconinoensis the tarsal claws are simple, without a basal lobe, while in the other two species the tarsal claws have a basal lobe. Also resembles A. chiricahuensis. In A. cococinoensis the mesosoma is uniformly light brown, the mesoscutum smooth, glabrous, notaulus complete, the median mesoscutal line in the form of a short triangle, the mesoscutellum rectangular, lateral propodeal carinae bent slightly outwards in their posterior half, the metasoma is longer than high in lateral view, while in A.chiricahuensis the mesosoma is dark brown, the mesoscutum uniformly alutaceous, notaulus incomplete, extending to 2/3 of mesoscutum length, the median mesoscutal line absent, the mesoscutellum trapezoid, lateral propodeal carinae bent outwards at their mid height; the metasoma is higher than long in lateral view. Description. Sexual female (Figs. 77–87). Head dark brown, except light brown lower face, mesosoma uniformly light brown; maxillary and labial palpi yellow; scapus and last three flagellomeres dark brown, rest of antennae yellow; legs light brown to yellowish; metasoma dark brown. Head alutaceous, with contrasting sparse white setae, nearly as broad as long and slightly broader than mesosoma in frontal view, 1.9× as broad as long in dorsal view. Gena alutaceous, not broadened behind eye, 1.5× as short as transverse diameter of eye. Malar space alutaceous, glabrous, with a few distinct parallel striae radiating from clypeus and reaching eye; eye 2.4× as high as length of malar space. Inner margins of eyes converging ventrally. POL 1.7× as broad as OOL, OOL 1.8× as long as diameter of lateral ocellus and 1.2× as long as LOL; all ocelli ovate, of same size. Transfacial distance 1.2× as long as height of eye; diameter of antennal torulus 1.7× as long as distance between them, distance between torulus and eye 1.4× as long as diameter of torulus; lower face alutaceous, glabrous, without striae and setae, slightly elevated median area smooth, glabrous, with few setae. Clypeus smooth, glabrous, rectangular, 1.6× as broad as high; ventrally rounded, not emarginate, without median incision; anterior tentorial pit large, rounded, epistomal sulcus distinct, deep, clypeo-pleurostomal line well impressed. Frons alutaceous, glabrous with few setae and without striae; interocellar area elevated, alutaceous, with some setae. Vertex, postgena alutaceous, with rare white setae; occiput and postocciput smooth, glabrous, without setae; posterior tentorial pit small, rounded, area below impressed; occipital foramen shorter than height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which diverge strongly; anteriorly postgenal bridge as broad as occipital foramen. Antenna longer than head+mesosoma, with 12 flagellomeres; pedicel slightly longer than broad; flagellomeres slightly broadened till apex; F1 1.7× as long as pedicel and 1.1× as long as F2; F2 1.2× as long as F3; F3=F4, F5–F11 nearly equal in length, F12 1.6× as long as F11, placodeal sensilla on F3–F12. Mesosoma 1.2× as long as high. Pronotum smooth, glabrous, with row of dense setae along anterolateral margin; propleuron uniformly smooth, glabrous. Mesoscutum smooth, glabrous, slightly longer than broad (greatest width measured across mesoscutum at level of base of tegulae), with some scattered, white setae; setae denser anteriorly alongside notaulus. Notaulus complete, broader posteriorly and converging; bottom of notaulus smooth, glabrous; anterior parallel line invisible; parapsidal line indistinct; median mesoscutal line in the form of a short triangle; circumscutellar carina reaching notaulus. Mesoscutellum rectangular, slightly longer than broad, with subparallel sides, smooth, glabrous, with some irregular rugae posteriorly and posterolaterally, posteriorly rounded, overhanging metanotum. Mesoscutellar foveae separated by elevated central carina, rounded, with smooth, glabrous bottom. Mesopleuron and speculum uniformly smooth, glabrous, with a few setae anteroventrally; mesopleural triangle smooth, glabrous, with some delicate rugae, with sparse white setae; dorsal and lateral axillar areas smooth, with a few white setae; subaxillular bar smooth, glabrous, posterior part as high as height of metanotal trough; metapleural sulcus reaching mesopleuron in upper 1/3, upper part of sulcus indistinct, narrow. Metascutellum smooth, glabrous, as high as height of smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous, without setae; central propodeal area narrow, smooth, glabrous, without rugae; lateral propodeal carinae bent slightly outwards in posterior half; lateral propodeal area smooth, glabrous, with long white setae. Nucha with delicate sulci dorsally and laterally. Tarsal claws simple, without basal lobe. Forewing longer than body, hyaline, margin with long, dense cilia, veins dark brown, radial cell open, 4.5× as long as broad; R1and Rs reaching wing margin; areolet triangular, closed, distinct. Rs+M distinct at 3/4 of distance to basalis and its projection reaching basalis at mid height. Metasoma as long as head+mesosoma, longer than high in lateral view; second metasomal tergite extending to half the length of metasoma in dorsal view, with white setae anterolaterally; all tergites smooth, glabrous, without micropunctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium 5.4× as long as broad in ventral view, with some setae ventrally which not extend beyond apex of spine. Body length 1.3–1.5 mm (n = 4). Male. Similar to female, but eyes bigger, ratio of malar space length to height of eye greater than in female; ocelli slightly bigger; antenna longer than body, with 13 flagellomeres, F1 slightly curved, apically slightly swollen. Body length 1.2–1.3 mm (n = 2). Gall (Fig. 88). A small unilocular bud gall, roughly spherical, about 3 mm in diameter, with a smooth hard surface. Pale brown colour, sometimes with a slight darker brown mottled pattern. Similar to or possibly the same as the undescribed gall in Figure 20 of Weld (1960). Biology. Only a sexual generation is known,which induces galls on Q.arizonica, Q.turbinella and Q.oblongifolia. Adults emerged in April. Despite coming from different locations, the pairing of male and female collections was confirmed using DNA data, with cytb sequences differing by 1.15% (GenBank OK346262 – OK346263) and ITS2 sequences differing only by an insertion of a single dinucleotide repeat unit (OK350628 – OK350629). Distribution. USA, Arizona: Santa Catalina Mountains, Chiricahua Mountains, Mogollon Rim., Published as part of Melika, George, Nicholls, James A., Abrahamson, Warren G., Buss, Eileen A. & Stone, Graham N., 2021, New species of Nearctic oak gall wasps (Hymenoptera: Cynipidae, Cynipini), pp. 1-131 in Zootaxa 5084 (1) on pages 25-28, DOI: 10.11646/zootaxa.5084.1.1, http://zenodo.org/record/5800716, {"references":["Gillette, C. P. (1892) Colorado Cynipidae. Entomological News, 3 (1892), 246 - 248.","Weld, L. H. (1960) Cynipid galls of the Southwest. Privately printed, Ann Arbor, Michigan, 50 pp."]}

Zapatella abrahamsoni Melika, sp. nov. Figs. 435–445 urn:lsid:zoobank.org:act: 718E0718-850D-4CDA-8A4A-C5C57A289402 Type material: HOLOTYPE female “ USA, FL., Highlands Co., Lake Placid, Archbold Biological Station, coll. G. Melika on 6-8 February 1995, dissected from galls 13–17 March 1995; ex Quercus inopina ”. PARATYPES: 11 female paratypes with the same labels as the holotype. The holotype female and 3 females are deposited at the USNM, 8 females at the PHDNRL. Etymology. Named in recognition of the significant contribution of Prof. Warren G. Abrahamson (Bucknell University, Lewisburg, Pennsylvania, USA) to studies of oak gall wasps. Diagnosis. Most closely resembles Z. cryptica. In Z. abrahamsoni the head and mesosoma are light reddish brown, the median mesoscutal line extending to 1/3 of the mesoscutum length; lateral propodeal area entirely covered with dense felt-like white setae, while in Z. cryptica the head and mesosoma are darker, the median mesoscutal line extending at least to 2/3 of the mesoscutum length or even longer; the lateral propodeal area without dense felt-like white setae. Description. Asexual female (Figs. 435–444). Body, antennae and legs uniformly reddish brown, only tips of mandibles, postocciput, propleura and tarsal claws always darker. Head uniformly delicately reticulate, with a few white, sparse, short and inconspicuous setae, denser on lower face and postgena; 1.2× as broad as high and slightly broader than mesosoma in frontal view, 1.8× as broad as long in dorsal view. Gena broadened behind eye in frontal view, broader than transverse diameter of eye in lateral view. Malar space with striae radiating from clypeus and nearly reaching eye, without sulcus. Eye 1.5× as high as length of malar space; lower face delicately coriaceous, without elevated area medially. Clypeus slightly impressed, setose, alutaceous; ventrally rounded, slightly emarginate, medially not incised; anterior tentorial pits small, indistinct, epistomal sulcus and clypeo-pleurostomal line distinct. POL 1.2× as long as OOL, OOL 2.5× as long as length of lateral ocellus and 1.75× as long as LOL, interocellar area reticulate, not elevated. Frons, vertex, occiput, postgenae reticulate; postocciput alutaceous. Antenna slightly longer than head+mesosoma, with 12 flagellomeres, pedicel slightly longer than broad, F1 slightly shorter than F2 and equal to F3; F6–F11 shorter and broader than preceding flagellomeres; F12 longer than F11 (in some specimens the suture between F12 and F11 indistinct); placodeal sensilla on F5–F12, hardly discernable or invisible on F1–F4. Mesosoma slightly longer than high, mesoscutum dorsally concave in lateral view. Pronotum along anterolateral side with dense white setae, laterally delicately reticulate, without carinae posterolaterally. Mesoscutum slightly broader than long in dorsal view, with sparse scattered setae, with uniform regular reticulate surface. Notaulus deep and uniformly broad, reaching pronotum, with smooth glabrous bottom; median mesoscutal line extending to 1/3 length of mesoscutum, parapsidal line distinct, impressed, extending to half the length of mesoscutum; anterior parallel line invisible. Mesopleuron uniformly reticulate. Mesoscutellum trapezoid, uniformly reticulate; scutellar foveae transversely ovate, with smooth and glabrous bottom, clearly separated by a narrow elevated reticulate area. Metascutellum smooth, higher than height of smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous, with dense white setae. Metapleural sulcus reaching mesopleuron at upper 1/3 of its height. Propodeum laterally with dense white setae that hide the surface sculpture, with smooth, glabrous central propodeal area, delimited by distinct subparallel lateral carinae, which bend slightly outwards in posterior 1/3. Nucha with irregular rugae. Forewing nearly as long as body, pubescent, margin without cilia; radial cell open, around 2.8× as long as broad; veins very light, hardly traceable; areolet indistinct, usually invisible; vein Rs+M points slightly below midway along basalis; R1 and Rs never reach wing margin, very inconspicuous, often invisible or absent. Tarsal claws simple, without basal lobe, but with broad base. Metasoma shorter than head+mesosoma, slightly longer than high and as long as high in lateral view; all metasomal tergites smooth, glabrous; base of second metasomal tergite with felt-like dense ring of white setae, interrupted dorsally, and with a few scattered setae on lateral surface of tergite. Narrow posterior band on second metasomal tergite and all subsequent tergites with very delicate, dense micropunctures. Prominent part of ventral spine of hypopygium needle-like, tapering to apex, 9.0–10.0× as long as broad, with two parallel rows of short, white, scattered setae not reaching beyond apex of spine. Body length 2.5–2.7 mm (n = 10). Gall. (Fig. 445). The galls are rounded, irregular woody expansions of lateral buds, 5.0– 7.5 mm in diameter, 4–6 mm high. The tissue of the gall is hard, typically with 2–4 larval cells (very rarely only one) embedded in the surrounding woody tissues. The walls of the larval chambers are distinct, slightly paler than the surrounding woody tissue. The galls persist on the trees for years. Biology. Only an asexual generation is known, which induces galls on Q. inopina, an oak species endemic to the Lake Wales Ridge. The galls become visible in late September and develop through November.The larvae (sometimes pupae) overwinter in the galls. Adult wasps were dissected from galls from the end of January through March; emerge in late April-May. One individual from the type series was sequenced for cytb (GenBank OK346326). Distribution. USA, Florida, Highlands Co., Lake Placid, Archbold Biological Station. This species is likely endemic to south-central Florida and particularly to the Lake Wales Ridge., Published as part of Melika, George, Nicholls, James A., Abrahamson, Warren G., Buss, Eileen A. & Stone, Graham N., 2021, New species of Nearctic oak gall wasps (Hymenoptera: Cynipidae, Cynipini), pp. 1-131 in Zootaxa 5084 (1) on pages 118-122, DOI: 10.11646/zootaxa.5084.1.1, http://zenodo.org/record/5800716

Andricus mellificus Nicholls, Stone & Melika, sp. nov. Figs. 159–171 urn:lsid:zoobank.org:act: FA848B4B-F890-4807-A424-7D9C7D50A190 Type material: HOLOTYPE female “ USA, Arizona, Chiricahua Mnts nr Portal. Leg. J. Nicholls. 2008.04.06. Code AZ1973, spAZb7; ex Quercus arizonica ”. PARATYPES: 2 females with the same labels as the holotype. The holotype is deposited at the USNM, 2 female at the PHDNRL. Etymology. Named after the behavior of this species in inducing galls that secrete nectar. Diagnosis. The gall is very distinctive; no other species from Arizona are known to induce such a specifically shaped and coloured gall (Fig. 171). Description. Female (Figs. 159–170). Head, mesosoma black, metasoma dark brown, antennae, mouthparts, tegulae, legs brown. Head delicately coriaceous, with sparse white setae, denser on lower face; rounded, only slightly broader than high and slightly broader than mesosoma in frontal view, 1.9× as broad as long in dorsal view. Gena coriaceous, slightly broadened behind eye in frontal view, narrower than transverse diameter of eye in lateral view. Malar space delicately coriaceous, glabrous, with a few delicate striae radiating from clypeus and not reaching eye; eye 2.9× as high as length of malar space. Eyes slightly converging ventrally. POL 2.1× as long as OOL; OOL 1.3× as long as diameter of lateral ocellus and equal to LOL; lateral ocelli slightly larger than central ocellus. Transfacial distance as long as height of eye; diameter of antennal torulus 1.8× as long as distance between them, distance between torulus and eye 1.3× as long as diameter of torulus; lower face coriaceous, with white long setae, without striae; slightly elevated median area coriaceous. Clypeus trapezoid, slightly broader than high, smooth, glabrous, with a few long setae; ventrally rounded, not emarginate and without median incision; anterior tentorial pit small, rounded, epistomal sulcus and clypeo-pleurostomal line distinct. Frons uniformly and delicately coriaceous, without striae; interocellar area delicately coriaceous and slightly elevated. Vertex delicately coriaceous; occiput with delicate transverse parallel rugae; postocciput smooth, glabrous; postgena coriaceous, 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 towards occipital foramen, postgenal bridge anteriorly broader. Antenna longer than head+mesosoma, with 12 flagellomeres (in some paratypes suture between F12 and F11 indistinct), pedicel slightly longer than broad, all flagellomeres slightly broadened towards apex, F1 1.3× as long as pedicel and equal in length to F2, F2 slightly longer than F3, F3=F4, F5–F7 equal in length, F7–F12 shorter than preceding flagellomeres and equal in length, placodeal sensilla on F4–F12. Mesosoma longer than high, with sparse short white setae. Pronotum smooth, glabrous dorsally; coriaceous, with net of delicate striae laterally; propleuron with delicate transverse striae, glabrous. Mesoscutum reticulate in posterior 2/3, with more transversely orientated striae in anterior 1/3, with sparse white setae, denser along lateral and anterior sides; slightly longer than broad (greatest width measured across mesoscutum at level of base of tegulae). Notaulus complete, reaching pronotum; deeper and broader, with smooth glabrous bottom in posterior 2/3, less impressed but still distinct in anterior 1/3 of mesoscutum; posteriorly converging. Anterior parallel line visible in anterior 1/3 of mesoscutum, delimited by smooth surface; parapsidal line indistinct, hardly traceable; median mesoscutal line short; circumscutellar carina narrow, smooth, reaching notaulus. Mesoscutellum rounded, slightly longer than broad, posteriorly rounded, uniformly dull rugose, overhanging metanotum; mesoscutellar foveae separated by narrow elevated coriaceous central carina, ovate, broader than high, with smooth, glabrous bottom. Mesopleuron with delicate transverse striae at mid height, running across full length of mesopleuron; speculum smooth, glabrous; ventral part uniformly smooth, glabrous, without striae. Mesopleural triangle coriaceous; dorsal and lateral axillar areas alutaceous, glabrous, with some white short setae; subaxillular bar smooth, glabrous, most posterior part slightly narrower than height of metanotal trough; metapleural sulcus reaching mesopleuron in upper 1/3 of height, upper part of sulcus distinct. Metascutellum coriaceous, at least 3.0× as high as height of smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous, with dense white setae; central propodeal area quadrangular, smooth, glabrous; lateral propodeal carinae parallel; lateral propodeal area smooth, glabrous, with long white setae. Nucha rugose, with strong irregular sulci. Tarsal claws with basal lobe. Forewing longer than body, hyaline, margin without cilia, veins dark brown, radial cell open, 3.1× as long as broad, areolet triangular, distinct, Rs+M inconspicuous, its projection reaching basalis at mid height. Metasoma as long as head+mesosoma, longer than high in lateral view; second metasomal tergite extending to 3/4 of metasoma length in dorsal view, with white setae anterolaterally; second and third tergites without micropunctures, fourth tergite with micropunctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium 3.2× as long as broad in ventral view, narrowing towards apex, with some short setae ventrally which do not extend beyond apex of spine. Body length 2.1–2.3 mm (n = 3). Gall (Fig. 171). A small unilocular bud gall, reaching 4–5 mm in diameter, green at first then becoming redpurple, in an axillary bud. Gall ovoid with a flattened top and a central nipple; nectar-secreting. Also illustrated in Figure 24 of Weld (1960). Biology. Only females are known, which induce small bud galls on Q. arizonica; Weld (1960) also lists Q. rugosa as a host. Galls mature in April, and adults emerge soon afterwards. Although only females are known, the spring phenology of this gall strongly implies that this is a sexual generation and males remain to be discovered. This species was included under the species code spAZb 7 in the Nicholls et al. (2017) study of nectar secretion in oak gall wasps. This gall was collected at several sites in Arizona, with DNA data used to confirm the morphological assessment that emerged individuals were conspecific. Two individuals were sequenced for cytb, differing by 1.62% (GenBank KX683628, OK346276). Distribution. USA, Arizona: Chiricahua Mountains, Sonoita., Published as part of Melika, George, Nicholls, James A., Abrahamson, Warren G., Buss, Eileen A. & Stone, Graham N., 2021, New species of Nearctic oak gall wasps (Hymenoptera: Cynipidae, Cynipini), pp. 1-131 in Zootaxa 5084 (1) on pages 46-49, DOI: 10.11646/zootaxa.5084.1.1, http://zenodo.org/record/5800716, {"references":["Weld, L. H. (1960) Cynipid galls of the Southwest. Privately printed, Ann Arbor, Michigan, 50 pp.","Nicholls, J. A., Melika, G. & Stone, G. N. (2017) Sweet tetra-trophic interactions: multiple evolution of nectar secretion, a defensive extended phenotype in cynipid gallwasps. American Naturalist, 189, 67 - 77. https: // doi. org / 10.5061 / dryad. bj 82 r."]}

Andricus nichollsi Melika & Stone, sp. nov. Figs. 194–204 urn:lsid:zoobank.org:act: 46627F86-0717-496A-B75A-FAE0FF6C1F3F Type material: HOLOTYPE female “ USA, Arizona, Molino Basin campground. Santa Catalina Mnts. Leg. J. Nicholls. 2008.04.09. Code AZ1916, spAZl21; ex Quercus oblongifolia ”. PARATYPES: 3 females with the same labels as the holotype. The holotype is deposited at the USNM, 3 females at the PHDNRL. Etymology. In recognition of the continuing contribution of Dr. James A. Nicholls (Institute of Evolutionary Biology, University of Edinburgh, UK, and Australian National Insect Collection, Canberra, Australia) to studies on oak gall wasps. Diagnosis. The gall somewhat resembles the sexual gall of Andricus kingi Bassett, 1900 but it is larger and greenish. The entire habitus of the female is more like Dryocosmus and is quite different from that of A. kingi. Description. Sexual female (Figs. 194–203). Head and mesosoma light brown, metasoma slightly darker. maxillary and labial palpi, antennae pale yellow, legs uniformly whitish yellow. Head alutaceous, with sparse white setae, denser on lower face, 1.2× as broad as high and as broad as mesosoma in frontal view, 2.2× as broad as long in dorsal view. Gena alutaceous, not broadened behind eye in frontal view, at least 2.5× as narrow as transverse diameter of eye in lateral view. Malar space alutaceous, glabrous, with delicate striae, radiating from clypeus and nearly reaching eye. Eyes large, occupying nearly whole length of head in frontal view, 6.6× as high as height of malar space, inner margins of eyes nearly parallel. POL 2.1× as long as OOL; OOL nearly equal in length to diameter of lateral ocellus and equal to LOL; all ocelli ovate, of same size. Transfacial distance narrower than height of eye; diameter of antennal torulus 1.9× as long as distance between them, distance between torulus and eye equal to diameter of torulus; lower face smooth, glabrous, with white setae, without striae; slightly elevated median area alutaceous, glabrous, without setae. Clypeus rectangular, slightly broader than high, smooth, glabrous, with a few long setae; ventrally rounded, not emarginate and without median incision; anterior tentorial pit large, rounded, epistomal sulcus and clypeo-pleurostomal line broad, well impressed. Frons alutaceous, without striae, with a few short setae and impressed area below central ocellus; interocellar area coriaceous, with some irregular short striae. Vertex, occiput, postgena alutaceous, with sparse white setae; postocciput smooth, glabrous; posterior tentorial pit large, ovate, area below impressed; occipital foramen higher than height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which bend outwards, postgenal bridge anteriorly broadened, as broad as width of occipital foramen. Antenna longer than head+mesosoma, with 12 flagellomeres, pedicel 1.5× as long as broad, F1 2.3× as long as pedicel and 1.4× as long as F2; F2 1.25× as long as F3; F3=F4=F5, subsequent flagellomeres shorter, nearly equal in length, F12 slightly longer than F11; placodeal sensilla on F2– F12. Mesosoma slightly longer than high, with sparse short white setae. Pronotum smooth, glabrous, with distinct parallel rugae dorsolaterally; propleuron smooth, glabrous. Mesoscutum smooth, glabrous, with few white setae along notaulus; slightly longer than broad (greatest width measured across mesoscutum at level of base of tegulae). Notaulus complete, deep, broad, with smooth, glabrous bottom, slightly converging posteriorly; anterior parallel, parapsidal and median mesoscutal lines absent; circumscutellar carina narrow, smooth, reaching notaulus. Mesoscutellum rectangular, with parallel sides, slightly longer than broad, centrally smooth, glabrous, with strong rugae laterally and posteriorly; posteriorly slightly rounded, overhanging metanotum. Mesoscutellar foveae in the form of a semilunar impression, with smooth, glabrous bottom, not separated. Mesopleuron and speculum uniformly smooth, glabrous, with a few setae ventrally; mesopleural triangle smooth, glabrous, with strong irregular striae in anterior half; dorsal and lateral axillar areas smooth, glabrous, with a few white short setae; subaxillular bar smooth, glabrous, with subparallel sides, posteriorly as high as height of metanotal trough; metapleural sulcus reaching mesopleuron in lower half, upper part of sulcus indistinct. Metascutellum smooth, glabrous, as high as height of smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous, without setae; central propodeal area smooth, glabrous, broad, with some incomplete longitudinal parallel rugae; lateral propodeal carinae strongly curved outwards at mid height; lateral propodeal area smooth, glabrous, with long dense white setae. Nucha smooth, glabrous dorsally, with numerous sulci laterally. Tarsal claws simple, without basal lobe. Forewing longer than body, hyaline, margin with long dense cilia, veins pale yellow, radial cell open, 5.0× as long as broad, R1 and Rs reaching wing margin, areolet triangular, distinct, Rs+M inconspicuous, traceable along 1/2 of its length, its projection reaching basalis at lower half of its height. Metasoma as long as head+mesosoma, nearly as long as high in lateral view; second metasomal tergite extending to half the length of metasoma in dorsal view, with few white setae anterolaterally; all tergites without micropunctures, smooth, glabrous. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium 4.0× as long as broad in ventral view, with a few short setae ventrally which do not extend beyond apex of spine. Body length 1.8–2.1 mm (n = 4). Male. Unknown. Gall. (Fig. 204). A small pointed unilocular leaf gall, 4–5 mm tall, 2 mm across, on edge of leaf, which causes the leaf to curve inwards at the point of gall attachment. Gall pale green with scattered brown flecks and covered with a dense coating of short hairs. Some of the hairs and the gall tip become red and then brown as the gall matures. Biology. Only females are known. Their morphology strongly suggests that they belong to the sexual generation rather than the asexual generation. This is also suggested by the gall’s rapid development in spring. In addition, very similar galls with comparable morphology (small, unilocular, roughly conical, developing on the edge of leaves) are known from the sexual generations of some Nearctic Andricus species (see Doutt 1960, Dailey & Sprenger 1973a,b). Galls develop on Q. oblongifolia, mature in April; adults emerge soon afterwards. Since this species induces a rather generic sexual generation gall morphology, two individuals were sequenced to confirm the morphological assessment of conspecificity. Cytb sequences were 1.15% divergent (GenBank OK346279 – OK346280), and ITS2sequences were 0.41% divergent (OK 350644 – OK350645), matching expectations they were conspecific. Distribution. USA, Arizona, Santa Catalina Mountains., Published as part of Melika, George, Nicholls, James A., Abrahamson, Warren G., Buss, Eileen A. & Stone, Graham N., 2021, New species of Nearctic oak gall wasps (Hymenoptera: Cynipidae, Cynipini), pp. 1-131 in Zootaxa 5084 (1) on pages 56-59, DOI: 10.11646/zootaxa.5084.1.1, http://zenodo.org/record/5800716, {"references":["Bassett, H. F. (1900) New species of North American Cynipidae. Transactions of the American Entomological Society, 26, 310 - 336.","Doutt, R. L. (1960) Heterogony in Andricus crystallinus Bassett (Hymenoptera: Cynipidae). Pan-Pacific Entomologist, 36, 167 - 170.","Dailey, D. C. & Sprenger, C. M. (1973 a) Synonyny of Andricus gigas and the bisexual generation of Andricus crenatus (Hymenoptera: Cynipidae). The Pan-Pacific Entomologist, 49, 188 - 191."]}