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BELONOCNEMA TREATAE (MAYR, 1881) (FIG. 2) Dryorhizoxenus floridanus Ashmead, 1881, female, male, sexual generation, gall. Belenocnema [sic!] treatae Mayr, 1881, female, sexual generation. Material examined: Asexual generation ��� 12F ��� USA: FL: Jacksonville Type 2813, USNM, Dryorhizoxenus floridanus ���; 6F ��� USA:FL: E. Florida, USNM���; 1F ��� USA:FL: LaBelle, IV-20���21, USNM���; 1F ��� USA: FL: Tampa 14-4, USNM���; 1F ��� USA: FL: Manatee Co., 111��� 26, R. F. Tinker, USNM���; 5F ��� USA: AL, Dauphin Island, 30.2504, -88.1325, 29/X/2016, Ott Lab, Q. virginiana ���; 2F ��� USA: FL, Archbold Biol. Stn., 27.1846, -81.3521, 28/X/2016, Ott Lab, Q. virginiana ���; 5F ��� USA: FL, Kissimmee River, 27.3780, -81.0968, 18/X/2016, Ott Lab, Q. virginiana ���; 5F ��� USA: FL, Perry, 30.1161, -83.5895, 14/X/2015, Ott Lab, Q. virginiana ���; 5F ��� USA: GA, Jekyll Island, 31.0174, -81.4297, 28/X/2016, Ott Lab, Q. virginiana ���; 3F ��� USA: MS, Gautier, 30.3803, -88.6104, 28/X/2016, Ott Lab, Q. virginiana ���; 5F ��� USA: NC, Fort Macon, 34.6951, -76.6862, 30/X/2016, Ott Lab, Q. virginiana ���; 5F ��� USA: SC, Charleston, 32.7688, -79.9734, 30/X/2016, Ott Lab, Q. virginiana ���. Sexual generation ��� 1F Lectotype (photo only) ���Brief mai 78 N. Amer., Collect G. Mayr, Bel. Treatae det. G. Mayr, LECTOTYPE Belonocnema treatae Mayr desig. G. Melika 998��� NHMW. 5F 6M ��� USA: AL, Gulf Shores, 30.2558, -87.7205, 31/X/2016, Ott Lab, Q. virginiana ���; 5F 5M ��� USA: FL, Kissimmee River, 27.3780, -81.0968, III/2016, Ott Lab, Q.virginiana ���; 10M ��� USA: FL, Okeechobee, 27.2434, -80.8276, III/2017, Ott Lab, Q. virginiana ���; 5F 5M ��� USA: FL, Perry, 30.1161, -83.5895, III/2016, Ott Lab, Q. virginiana ���. Diagnosis: Belonocnema treatae can be distinguished from B. fossoria by the spur on the anterior side of fore tibia shorter than basitarsus and tibial spurs in both generations. It can also be separated from B. kinseyi in the sexual generation by the weakly delimited scutellar foveae separated broadly by a ridge and the yellowish brown colour along with a distinctive areolet in the asexual generation. Description Asexual female (Fig. 2E): Body length 2.8���3.2 mm (N = 35). Yellowish brown; tip of mandibles, scutellum, propodeum, mesopleural triangle, metapleuron, scutellum, hind tibia dark brown (Fig. 2E). Head finely coriaceous with sparse white setae; slightly rounded in dorsal view; 2�� as broad as long in dorsal view; 1.3�� as broad as long in frontal view; slightly broader than mesosoma. Gena alutaceous; not broadened behind eye in dorsal view; 1.1�� broader than the cross diameter of eye. Malar space alutaceous, without striae radiating from clypeus; eye 2.2�� higher than length of malar space. Inner margins of eyes parallel. OOL 1.2�� longer than POL; OOL 2.4�� longer than LOL; ocelli ovate, all equal in size. Transfacial distance 1.7�� longer than height of eye and 1.8�� longer than height of lower face; diameter of antennal torulus 1.3�� longer than distance between them, distance between torulus and eye margin 3.3�� longer than diameter of torulus. Lower face finely coriaceous, with white setae, without striae radiating from clypeus, median area not elevated. Clypeus trapezoid, flat, broader than high, with deep anterior tentorial pits, distinct epistomal sulcus and clypeo-pleurostomal line. Frons finely coriaceous, glabrous; vertex, interocellar area, occiput is finely coriaceous. Postgena coriaceous, glabrous. Antenna 13 segmented, longer than head + mesosoma; F1 shorter than the length of scape + pedicel, 1.6�� longer than F2 (Fig. 2E). Mesosoma longer than high in lateral view. Propleuron evenly setose. Mesoscutum smooth, glabrous between notauli, alutaceous lateral to notaulus; longer than broad; notauli complete, deeply impressed for full length; median mesoscutal line distinct; anterior parallel lines and parapsidal lines absent; mesoscutellum only slightly longer than broad, slightly narrower posteriorly; shorter than mesoscutum, uniformly rugose, overhanging metanotum; scutellar foveae present. Mesopleural triangle covered with dense white setae, mesopleuron smooth, glabrous, with a few white setae along ventral and anterior margins. Lateral propodeal carinae distinctly raised and bent outwards, central propodeal area rugose; lateral propodeal area alutaceous, with dense white setae; nucha short, coriaceous. Tibia setose on anterior edge. Fore tibia prolonged on the anterior side into a curved spine, much shorter than tibial spur and basitarsus; tarsal claws simple with a slight ridge but never a full tooth. Middle and hind tibia with two spurs. Fore wing hyaline, longer than body, margin with dense cilia. Radial cell 2.5�� as long as wide; 2r infumated, Rs curved upwards and thickened at apex; radial cell open; areolet large and distinct; Rs + M reaching to M; cu-a absent; cu1 broken (Fig. 2E). Metasoma shorter than head + mesosoma, 1.1�� longer than high in lateral view, smooth and glabrous. Second metasomal tergite setose medially; all subsequent tergites without setae, smooth, glossy. Ventral spine of the hypopygium short, prominent part 1.2�� as long as broad in ventral view, with white setae extending beyond the apex of spine (Fig. 2E). Sexual female (Fig. 2A���C, F): Body length 3.5���4.0 mm (N = 15). Yellowish brown; flagellomeres, tip of mandibles, metascutellum, propodeum, wing veins, middle and hind tibia, and tarsi dark brown (Fig. 2A). Head finely coriaceous with sparse white setae; slightly rounded in dorsal view; 1.8�� as broad as long in dorsal view; 1.2�� as broad as long in frontal view; slightly broader than mesosoma. Gena alutaceous, not broadened behind eye in dorsal view; 1.3�� broader than cross diameter of eye. Malar space alutaceous, without striae radiating from clypeus; eye 3�� higher than length of malar space. Inner margins of eyes parallel. OOL 1.3�� longer than POL; OOL 3.5�� longer than LOL; ocelli ovate, all equal in size. Transfacial distance 1.4�� longer than height of eye and 1.4�� longer than height of lower face (Fig. 2A); diameter of antennal torulus 2�� longer than distance between them, distance between torulus and eye margin equal to diameter of torulus. Lower face finely coriaceous, with white setae, without striae radiating from clypeus, median area not elevated. Clypeus trapezoid, flat, broader than high, with deep anterior tentorial pits, distinct epistomal sulcus and clypeo-pleurostomal line. Frons finely coriaceous, glabrous; vertex, interocellar area, occiput is finely coriaceous. Postgena coriaceous, glabrous. Antenna 14 segmented, longer than head + mesosoma; F1 shorter than the length of scape + pedicel, 1.4�� longer than F2 (Fig. 2C). Mesosoma longer than high in lateral view. Propleuron alutaceous, with few setae. Mesoscutum smooth, glabrous between notauli, alutaceous lateral to notaulus; longer than broad; notauli complete, deeply impressed for full length (Fig. 2B); median mesoscutal line distinct; anterior parallel lines and parapsidal lines absent. Mesoscutellum only slightly longer than broad, slightly narrower posteriorly; shorter than mesoscutum, uniformly rugose, overhanging metanotum (Fig. 2B); scutellar foveae shallow, weakly delimited posteriorly, separated widely by a ridge (Fig. 2F). Mesopleuron smooth, glabrous, with a few white setae along ventral and anterior margins. Lateral propodeal carinae distinct, curved; central propodeal area rugose; lateral propodeal area alutaceous, with dense white setae; nucha short, coriaceous. Tibia setose on anterior edge. Fore tibia prolonged on the anterior side into a curved spine, much shorter than tibial spur and basitarsus (Fig. 2C); tarsal claws simple with a slight ridge but never a full tooth. Middle and hind tibia with two spurs. Fore wing hyaline, longer than body, margin with dense cilia. Radial cell 2.3�� as long as wide; 2r infumated, Rs curved upwards and thickened at apex; radial cell open; areolet small and indistinct; Rs + M reaching to M; cu-a absent; cu1 broken (Fig. 2A). Metasoma shorter than head + mesosoma, 1.2�� longer than high in lateral view, smooth and glabrous; second metasomal tergite setose medially; all subsequent tergites without setae, smooth, glossy. Ventral spine of the hypopygium short, prominent part 1.3�� as long as broad in ventral view, with white setae extending beyond the apex of spine (Fig. 2A). Male: 3.7 mm (N = 26). Colour and sculptures like the sexual female, antenna 15 segmented; F1 is curved, excavated and incised medially. Metasoma smaller than head + mesosoma. Gall: Smooth, pea-like galls (4.92���5.50 mm) on the ventral side of leaves for the asexual generation, irregular shaped, multilocular galls often in clusters on the small rootlets for the sexual generation (Egan et al., 2013). The leaf galls produced by B. treatae on Q. virginiana are indistinguishable from the leaf gall produced by B. kinseyi on both Q. fusiformis (Fig. 1E) and Q. virginiana. Host plant: Quercus virginiana and rarely on Q. geminata. Distribution: Alabama, Florida, Georgia, Mississippi, North Carolina, South Carolina. Biology: The asexual generation has long, straight wings and is capable of flight (Table 2). Sexualgeneration adults emerge from mid-March to end of April, corresponding with the timing of leaf flush of their main host Q. virginiana (Hood et al., 2019). Remarks: Few specimens of this species have been collected from Q. geminata, which is unsurprising given the much later leaf flushing time of Q. geminata (Hood et al., 2019). While the lectotype of B. treatae designated from Gustav Mayr���s collection at NHMW by Melika & Bechtold (2001) was not examined physically, the high-quality image of the habitus clearly shows the diagnostic characters of B. treatae (small fore tibial spine shorter than tibial spur, broadly separated scutellar foveae)., Published as part of Zhang, Y. Miles, Egan, Scott P., Driscoe, Amanda L. & Ott, James R., 2021, One hundred and sixty years of taxonomic confusion resolved: Belonocnema (Hymenoptera: Cynipidae: Cynipini) gall wasps associated with live oaks in the USA, pp. 1234-1255 in Zoological Journal of the Linnean Society 193 (4) on pages 1250-1252, DOI: 10.1093/zoolinnean/zlab001, http://zenodo.org/record/5761660, {"references":["Mayr GL. 1881. Die Genera der gallenbewohnenden Cynipiden. Jahresbericht der Rossauer Communal-Oberrealschule: Wien 20: 1 - 38.","Ashmead WH. 1881. On the cynipidous galls of Florida. Transactions of the American Entomological Society and Proceedings of the Entomological Section of the Academy of Natural Sciences 9: 9 - 28.","Egan SP, Hood GR, DeVela G, Ott JR. 2013. Parallel patterns of morphological and behavioral variation among host-associated populations of two gall wasp species. PLoS One 8: e 54690.","Hood GR, Zhang L, Hu EG, Ott JR, Egan SP. 2019. Cascading reproductive isolation: plant phenology drives temporal isolation among populations of a host-specific herbivore. Evolution 73: 554 - 568.","Melika G, Bechtold M. 2001. Taxonomic notes and type designations of gall inducing cynipid wasps described by G. Mayr (Insecta: Hymenoptera: Cynipidae). Annalen des Naturhistorischen Museums in Wien. Serie B fur Botanik und Zoologie 103: 327 - 339."]}

Belonocnema quercusvirens, Burks, 1979. Material examined: Asexual generation ��� Syntype 1F ��� USA: FL, Clearwater, Reared Dec-13���19, Quercus geminata, L.H.Weld Collector, Hopk. U.S. 15634f, Type 24099, USNMENT 00802094���; Paratype 9F same locality as Syntype USNMENT 00893032, 00893095, 00893115, 00893123, 00893218; 4F ��� USA:SC, Charleston Co. S. Car ���43, Q. virginiana, 1160a, USNM���; 2F ��� USA: FL, Archbold Biol. Stn., 27.1846, -81.3521, 19/X/2016, Ott Lab, Q. geminata ���; 5F ��� USA: FL, Dickinson State Pk., 27.0261, -80.1090, 18/X/2016, Ott Lab, Q. geminata ���; 5F ��� USA: FL, Oceanside Village, 29.9542, -85.4277, 30/X/2016, Ott Lab, Q. geminata ���. Sexual Generation ��� 7F3M ��� USA: FL, Archbold Biol. Stn., 27.1846, -81.3521, IV/2018, Egan Lab, Q. geminata ���; 5F5M ��� USA: FL, Dickinson State Pk., 27.0261, -80.1090, III/2017, Egan Lab, Q. geminata ���; 5F5M ��� USA: FL, Lake Lizzie, 28.2277, -81.1800, III/2018, Egan Lab, Q. geminata ���; 4M ��� USA: FL, Volusia Co. Daytona Beach, IV-6-1998, Urban beachside UV light trap, C Yorke, S. Fullerton, UCFC 0017693, 0017694, 0017695, 0017696 ���; 1F2M ��� USA: FL, Brevard Co. Malabar, Malabar Rd. 30 Mar ��� 25 May 2000, P.J. Russell, Z. Prusak, S.M. Fullerton, UCFC 0079161, 0100100, 0101364 ���; 2F3M ��� USA: FL, Brevard Co. Titusville, SR 405, 21 Feb���15 May 2001, Z. Prusak, P.J. Russell, S.M. Fullerton, UCFC 0078923, 0079318, 0093022, 0093669, 0103143 ���; 2F2M ��� USA: FL, Orange Co. Rk. Spr. Rn. St. Res. IV-13���1995, S.M. Fullerton, UCFC 0201995, 0202004, 0202411, 0202415 ���; 5F5M ��� USA: FL, Orange Co. UCF 28��36���37���N 81��12���01���W LLP Flatwds, M. Carey, S.L. Kelly, S.M. Fullerton, III-28���2008, UCFC 0463902, 0463926, 0463936, 0463954, 0463957, 0464351, 0464355, 0464384, 0464390, 0464524. 8F4M ��� USA: FL, Orange / Osceola Co. Walt Disney World, 24 Mar���28 Apr 1998, Z. Prusak, S. Fullerton, UCFC 0017132, 0017146, 0017173, 0017230, 0017232, 0017601, 0017602, 0017603, 0017604, 0017608, 0017787, 0017793 ���; 2M ��� USA: FL, Sarasota Co. MCC- Venice Campus, III-31- 1997, K.J. Maharay, S.M. Fullerton, UCFC 0018361, 0018363 ���; 2F1M ��� USA: FL, Seminole Co. Econ. Wild. Area IV-8-2000, T. Smith, UCFC 0054591, 0054607, 0060190 ���; 2F1M ��� USA: FL, Seminole Co. Lower Wekiva River St. Preserve, 28-IV-2001, P.J. Russell, S.M. Fullerton, UCFC 0108938, 0109646, 0109778 ���; 3F ��� USA: FL, Seminole Co. Oveido, rural yard, Malaise trap, IV-7-1994, S.M. Fullerton, UCFC 0202173, 0202175, 0202190 ���; 4F6M ��� USA: FL, Seminole Co. Oveido, rural yard, UV light, 28��39���25���N 81��10���44���W, S.M. Fullerton, III-28���IV-12-2009, UCFC 0446980, 0446991, 0446992, 0446993, 0446994, 0446996, 0448649, 0448658, 0448659, 0448660 ���. Diagnosis: Belonocnema fossoria can be distinguished from the other two known species by the spur on the anterior side of fore tibia longer than basitarsus and tibial spurs (Fig. 3A). The asexual generation has small, non-functional wings, lack of areolet in the front wing, and the middle tibia with an additional spur. Description Asexual female (Figs 1D, 3A���C): Body length 2.5��� 3.5 mm (N = 12). Reddish brown; tip of mandibles, wing veins dark brown (Fig. 3A). Head finely coriaceous with sparse white setae; slightly rounded in dorsal view; 2.3�� as broad as long in dorsal view; 1.2�� as broad as long in frontal view; slightly broader than mesosoma. Gena alutaceous, not broadened behind eye in dorsal view; 1.2�� as broad as cross diameter of eye. Malar space alutaceous, without striae radiating from clypeus; eye 2.1�� higher than length of malar space. Inner margins of eyes parallel. OOL 1.6�� longer than POL; OOL 2.3�� longer than LOL; ocelli ovate, all equal in size. Transfacial distance 1.8�� longer than height of eye and 1.5�� longer than height of lower face (Fig. 3C); diameter of antennal torulus 2.3�� longer than distance between them, distance between torulus and eye margin 1.4�� longer than diameter of torulus. Lower face finely coriaceous, with white setae, without striae radiating from clypeus, median area not elevated. Clypeus trapezoid, flat, broader than high, with deep anterior tentorial pits, distinct epistomal sulcus and clypeopleurostomal line. Frons finely coriaceous, glabrous; vertex, interocellar area, occiput is finely coriaceous. Postgena coriaceous, glabrous. Antenna 13 segmented, longer than head + mesosoma; F1 shorter than the length of scape + pedicel, 1.7�� longer than F2 (Fig. 3C). Mesosoma longer than high in lateral view. Propleuron alutaceous, with few setae. Mesoscutum smooth, glabrous between notauli, alutaceous lateral to notaulus; longer than broad (width measured across base of tegulae); notauli complete, deeply impressed for full length; median mesoscutal line distinct; anterior parallel lines and parapsidal lines absent (Fig. 3B); Mesoscutellum only slightly longer than broad, slightly broader posteriorly; shorter than mesoscutum, uniformly rugose, overhanging metanotum; scutellar foveae present but shallow and indistinct. Mesopleural triangle large, sparsely setose; Mesopleuron smooth, glabrous, with a few white setae along ventral and anterior margins. Lateral propodeal carinae distinct, central propodeal area glabrous, with rugae; lateral propodeal area alutaceous, with dense white setae; nucha short, coriaceous. Legs short and stout; tibia setose on anterior edge; fore tibia prolonged on the anterior side into a curved spine (Fig. 3A), longer than basitarsus and the tibial spur; tarsal claws simple with a slight ridge but never a full tooth. Middle tibia also with curved spine but smaller than basitarsus and tibial spurs; middle and hind tibia with two spurs. Tarsi covered in setae. Fore wing hyaline, shorter than body and often curved upwards, margin with dense cilia; Radial cell 2�� as long as wide; 2r and Rs infumated, Rs curved upwards and thickened at apex; radial cell open; areolet absent; Rs + medial vein (M) reaching to M; cubitus-anal crossvein (cu-a) absent; first cubitus vein (cu1) broken (Fig. 3B). Metasoma slightly longer than head + mesosoma, as long as high in lateral view, smooth and glabrous; second metasomal tergite occupies setose medially; all subsequent tergites without setae, smooth, glossy; ventral spine of the hypopygium short, prominent part as long as broad in ventral view, with white setae extending beyond the apex of spine (Fig. 3A). N = number tested; * see Supporting Information, Table S1 for corresponding site abbreviations. Sexual female (Figs 1B, 3E): Body length 3.5���4.5 mm (N = 16). Reddish brown; pedicel, flagellomeres, vertex, tip of mandibles, tarsal claws, wing veins dark brown (Fig. 3E). Head finely coriaceous with sparse white setae; slightly rounded in dorsal view; 1.8�� as broad as long in dorsal view; 1.3�� as broad as long in frontal view; slightly broader than mesosoma. Gena alutaceous, not broadened behind eye in dorsal view; 0.7�� as broad as cross diameter of eye. Malar space alutaceous, without striae radiating from clypeus; eye 2.1�� higher than length of malar space. Inner margins of eyes parallel. OOL 1.6�� longer than POL; OOL 1.4�� longer than LOL; ocelli ovate, all equal in size. Transfacial distance 1.6�� longer than height of eye and 1.3�� longer than height of lower face; diameter of antennal torulus 2�� longer than distance between them, distance between torulus and eye margin 1.3�� longer than diameter of torulus. Lower face finely coriaceous, with white setae, without striae radiating from clypeus, median area not elevated. Clypeus trapezoid, flat, broader than high, with deep anterior tentorial pits, distinct epistomal sulcus and clypeo-pleurostomal line. Frons finely coriaceous, glabrous; vertex, interocellar area, occiput is finely coriaceous. Postgena coriaceous, glabrous. Antenna 14 segmented, longer than head + mesosoma; F1 shorter than the length of scape + pedicel, 1.3�� longer than F2 (Fig. 3E). Mesosoma longer than high in lateral view. Propleuron alutaceous, with few setae. Mesoscutum smooth, glabrous between notauli, alutaceous lateral to notaulus; longer than broad (width measured across base of tegulae); notauli complete, deeply impressed for full length; median mesoscutal line distinct; anterior parallel lines and parapsidal lines absent; mesoscutellum only slightly longer than broad, slightly broader posteriorly; shorter than mesoscutum, uniformly rugose, overhanging metanotum; scutellar foveae present. Mesopleuron smooth, glabrous, with a few white setae along ventral and anterior margins; mesopleural triangle setose. Lateral propodeal carinae bent outwards, central propodeal area glabrous, with rugae; lateral propodeal area alutaceous, with dense white setae; nucha short, coriaceous. Legs short and stout; tibia setose on anterior edge; fore tibia prolonged on the anterior side into a curved spine, longer than basitarsus and the tibial spur; tarsal claws simple with a slight ridge but never a full tooth. Middle tibia also with curved spine but smaller than basitarsus and tibial spurs; middle and hind tibia with two spurs. Tarsi covered in setae (Fig. 3E). Fore wing hyaline, shorter than body and often curved upwards, margin with dense cilia; radial cell 2�� as long as wide; 2r and Rs infumated, Rs curved upwards and thickened at apex; radial cell open; areolet present; Rs + M reaching to M; cu-a absent; cu1 broken (Fig. 3E). Metasoma slightly longer than head + mesosoma, as long as high in lateral view, smooth and glabrous; second metasomal tergite occupies setose medially; all subsequent tergites without setae, smooth, glossy; ventral spine of the hypopygium short, prominent part 1.5�� as long as broad in ventral view, with white setae extending beyond the apex of spine (Fig. 3E). Male: Body length 3.2���4.1 mm (N = 15). Colour and sculptures like the sexual female, Antenna 15 segmented; F1 is curved, excavated, and incised medially. Metasoma smaller than head + mesosoma. Gall: Smooth, single-chambered pea galls (5.88��� 6.45mm in diameter) on the ventral side of leaves for the asexual generation (Fig. 1C); irregular shaped, multilocular galls often in clusters on the small rootlets for the sexual generation (Fig. 1A; Egan et al., 2013). Host plant: Quercus geminata. Distribution: Georgia, Florida (USA). Biology: The small, often bent wings of the asexual generation (Figs 1D, 3A) appear to be non-functional (Table 2), and the large tibial spur and short, stout legs are likely fossorial adaptations that allows B. fossoria to reach the rootlets of Q. geminata in sandy soil where these species co-occur. The sexual generation emerges from early March to mid-April, corresponding with the timing of leaf flush of their host Q. geminata (Hood et al., 2019). Remarks: Osten Sacken���s (1861) original description of C. quercusvirens was based on the asexual galls alone collected on ���live oak��� in Georgia, the wasp itself was described by Weld (1921) as B. fossoria. As both B. treatae and B. fossoria can be found in the region and both induce similar galls, it is difficult to know which species Osten Sacken collected and named. However, names described before 1930, and which were applied only to the product of an animal, are valid names for the organism itself under ICZN Article 1, section 3. Therefore, we here propose the name B. quercusvirens as species inquirenda., Published as part of Zhang, Y. Miles, Egan, Scott P., Driscoe, Amanda L. & Ott, James R., 2021, One hundred and sixty years of taxonomic confusion resolved: Belonocnema (Hymenoptera: Cynipidae: Cynipini) gall wasps associated with live oaks in the USA, pp. 1234-1255 in Zoological Journal of the Linnean Society 193 (4) on pages 1244-1248, DOI: 10.1093/zoolinnean/zlab001, http://zenodo.org/record/5761660, {"references":["Burks B. 1979. Superfamily Cynipoidea. In: Krombein KV, Hurd P, Smith DR, Burks B, eds. Catalog of Hymenoptera in America north of Mexico. Washington, DC: Smithsonian Institution Press, 1045 - 1107.","Driscoe AL, Nice CC, Busbee RW, Hood GR, Egan SP, Ott JR. 2019. Host plant associations and geography interact to shape diversification in a specialist insect herbivore. Molecular Ecology 28: 4197 - 4211.","Egan SP, Hood GR, DeVela G, Ott JR. 2013. Parallel patterns of morphological and behavioral variation among host-associated populations of two gall wasp species. PLoS One 8: e 54690.","Hood GR, Zhang L, Hu EG, Ott JR, Egan SP. 2019. Cascading reproductive isolation: plant phenology drives temporal isolation among populations of a host-specific herbivore. Evolution 73: 554 - 568.","Osten Sacken CR. 1861. On the Cynipidae of the North American oaks, and their galls. Proceedings of the Entomological Society of Philadelphia 1: 47 - 72.","Weld LH. 1921. American gallflies of the family Cynipidae producing subterranean galls on oak. Proceedings of the United States National Museum 59: 187 - 246."]}

Zhang, Y. Miles, Egan, Scott P., Driscoe, Amanda L., Ott, James R. (2021): One hundred and sixty years of taxonomic confusion resolved: Belonocnema (Hymenoptera: Cynipidae: Cynipini) gall wasps associated with live oaks in the USA. Zoological Journal of the Linnean Society 193 (4): 1234-1255, DOI: 10.1093/zoolinnean/zlab001, URL: https://doi.org/10.1093/zoolinnean/zlab001

BELONOCNEMA FOSSORIA, WELD, 1921, REV. STAT. (FIGS 1A���D, 3), Published as part of Zhang, Y. Miles, Egan, Scott P., Driscoe, Amanda L. & Ott, James R., 2021, One hundred and sixty years of taxonomic confusion resolved: Belonocnema (Hymenoptera: Cynipidae: Cynipini) gall wasps associated with live oaks in the USA, pp. 1234-1255 in Zoological Journal of the Linnean Society 193 (4) on page 1244, DOI: 10.1093/zoolinnean/zlab001, http://zenodo.org/record/5761660, {"references":["Weld LH. 1921. American gallflies of the family Cynipidae producing subterranean galls on oak. Proceedings of the United States National Museum 59: 187 - 246."]}

BELONOCNEMA KINSEYI, WELD, 1921, REV. STAT. (FIGS 1E, F, 4) Belonocnema treatae, Lund et al., 1998, female, male, asexual, sexual generation, gall. Material examined: Asexual ��� Syntype 1F ��� USA: TX, Boerne, Nov. -15���1917, Cotype 22832, USNMENT 00802145���; 46F ���same locality as Syntype, Nov.-15��� Dec.1-1917, USNM���; 5F ��� USA: LA: Golden Meadow. 29.3939, -90.2729, 22/X/2016, Ott Lab, Q. virginiana ���; 5F ��� USA: LA, Oak Grove Hwy, 29.7668, -92.9750, 21/X/2016, Ott Lab, Q. virginiana ���; 2F ��� USA: MS, Gautier, 30.3803, -88.6104, 28/X/2016, Ott Lab, Q. virginiana ���; 5F ��� USA: MS, Picayune, 30.5271, -89.6813, 30/X/2016, Ott Lab, Q. virginiana ���; 5F ��� USA: OK, Quartz Mountain, 34.8901, -99.3011, 17/X/2018, Ott Lab, Q. fusiformis ���; 5F ��� USA: TX, Encino, 26.8942, -98.1352, 13/IX/2015, Ott Lab, Q. fusiformis ���; 5F ��� USA: TX, High Island, 29.5612, -94.3918, 17/X/2016, Ott Lab, Q. virginiana ���; 5F ��� USA: TX, Live Oak Park, 27.8544, -97.2105, 1/XI/2015, Ott Lab, Q. fusiformis ���; 5F ��� USA: TX, Luling, 29.6739, -97.6350, 5/XI/2016, Ott Lab, Q. fusiformis ���; 5F ��� USA: TX, Pleasanton, 28.9523, -98.4509, 23/X/2016, Ott Lab, Q. fusiformis ���; 5F ��� USA: TX, Rocksprings, 29.8751, -100.1086, 13/XI/2016, Ott Lab, Q. fusiformis ���. Sexual generation ��� 4F 6M ��� USA: TX, Rice University, 28.7174, -95.4023, III/2018, Egan Lab, Q. virginiana ���; 2F UTIC 200066, 200067 ��� USA, TX, Travis Co: Austin nr Austin Mem. Park Cemetery, 30.3281, -97.7543, 210 m 14.III.2016, A.L.Wild, UV Light, 20���2300 h���; 6F 6M ��� USA: LA, Golden Meadow, 29.3939, -90.2729. III/2018, Egan Lab, Q. virginiana ���; 5F 5M ��� USA: MS, Picayune, 30.5271, -89.6813. III/2016, Ott Lab, Q. virginiana ���; 5F 5M ��� USA: OK, Quartz Mountain, 34.8901, -99.3011. III/2016, Ott Lab, Q. fusiformis ���; 4F 5M ��� USA: TX, Encino, 26.8942, -98.1352, III/2015, Ott Lab, Q. fusiformis ���; 5F 4M ��� USA: TX, Live Oak Park, 27.8544, -97.2105, III/2015, Ott Lab, Q. fusiformis ���; 4F 4M ��� USA: TX, San Marcos, 29.9373. -98.0099, 11/ XI/2016, Ott Lab, Q. fusiformis ���. Diagnosis: Belonocnema kinseyi can be distinguished from B. fossoria by the spur on the anterior side of fore tibia shorter than basitarsus and tibial spurs in both generations. It can also be separated from B. treatae in the sexual generation by the deeply delimited scutellar foveae separated narrowly by a carina, and the reddish brown colour along with an indistinctive areolet in the asexual generation. Description Asexual female (Figs 1F, 4A���C): Body length 2.6��� 3.3 mm (N = 22). Reddish brown; tip of mandibles, mesosoma (except for mesoscutum), wing veins, anterior third of first gastral tergite, anterior edge of fore, meso, and metacoxae, and distal edge of hind femora black (Fig. 4A). Head finely coriaceous with sparse white setae; slightly rounded in dorsal view; 2.1�� as broad as long in dorsal view; 1.4�� as broad as long in frontal view; slightly broader than mesosoma. Gena alutaceous, not broadened behind eye in dorsal view; equally broad as cross diameter of eye. Malar space alutaceous, without striae radiating from clypeus; eye 2.3�� higher than length of malar space. Inner margins of eyes parallel. OOL 1.1�� longer than POL; OOL 2.2�� longer than LOL; ocelli ovate, all equal in size. Transfacial distance 1.4�� longer than height of eye and 1.3�� longer than height of lower face (Fig. 4C); diameter of antennal torulus 2�� longer than distance between them, distance between torulus and eye margin 2�� longer than diameter of torulus. Lower face finely coriaceous, with white setae, without striae radiating from clypeus, median area not elevated. Clypeus trapezoid, flat, broader than high, with deep anterior tentorial pits, distinct epistomal sulcus and clypeo-pleurostomal line. Frons finely coriaceous, glabrous; vertex, interocellar area, occiput is finely coriaceous. Postgena coriaceous, glabrous. Antenna 13 segmented, longer than head + mesosoma; F1 shorter than the length of scape + pedicel, 1.6�� longer than F2 (Fig. 4A). Mesosoma longer than high in lateral view. Propleuron alutaceous, with few setae. Mesoscutum smooth, glabrous between notauli, alutaceous lateral to notaulus; longer than broad; notauli complete, deeply impressed for full length; median mesoscutal line distinct; anterior parallel lines and parapsidal lines absent (Fig. 4B); mesoscutellum only slightly longer than broad, slightly narrower posteriorly; shorter than mesoscutum, uniformly rugose, overhanging metanotum; scutellar foveae present. Mesopleural triangle covered with dense white setae, mesopleuron smooth, glabrous, with a few white setae along ventral and anterior margins. Lateral propodeal carinae distinct, bent outwards; central propodeal area rugose; lateral propodeal area alutaceous, with dense white setae; nucha short, coriaceous. Tibia setose on anterior edge; Fore tibia prolonged on the anterior side into a curved spine, much shorter than tibial spur and basitarsus; tarsal claws simple with a slight ridge but never a full tooth. Middle and hind tibia with two spurs (Fig. 4A). Fore wing hyaline, longer than body, margin with dense cilia; radial cell 2�� as long as wide; 2r infumated, Rs curved upwards and thickened at apex; radial cell open; areolet small and indistinct; Rs + M reaching to M; cu-a absent; cu1 broken (Fig. 4A). Metasoma shorter than head + mesosoma, 1.2�� longer than high in lateral view, smooth and glabrous; second metasomal tergite setose medially; all subsequent tergites without setae, smooth, glossy; ventral spine of the hypopygium short, prominent part 1.3�� as long as broad in ventral view, with white setae extending beyond the apex of spine (Fig. 4A). Sexual female (Fig. 4E, F): Body length 3.5���4.0 mm (N = 15). Yellowish brown; scape, flagellomeres, tip of mandibles, propodeum, wing veins, distal edge of hind femora, hind tibia, and tarsi dark brown (Fig. 4E). Head finely coriaceous with sparse white setae; slightly rounded in dorsal view; 2.4�� as broad as long in dorsal view; 1.2�� as broad as long in frontal view; slightly broader than mesosoma. Gena alutaceous, not broadened behind eye in dorsal view; equally broad as cross diameter of eye. Malar space alutaceous, without striae radiating from clypeus; eye 2.3�� higher than length of malar space. Inner margins of eyes parallel. OOL 1.1�� longer than POL; OOL 2.2�� longer than LOL; ocelli ovate, all equal in size. Transfacial distance 1.8�� longer than height of eye and 1.7�� longer than height of lower face; diameter of antennal torulus 2.3�� longer than distance between them, distance between torulus and eye margin 1.6�� longer than diameter of torulus. Lower face finely coriaceous, with white setae, without striae radiating from clypeus, median area not elevated. Clypeus trapezoid, flat, broader than high, with deep anterior tentorial pits, distinct epistomal sulcus and clypeo-pleurostomal line. Frons finely coriaceous, glabrous; vertex, interocellar area, occiput is finely coriaceous. Postgena coriaceous, glabrous. Antenna 14 segmented, longer than head + mesosoma; F1 shorter than the length of scape + pedicel, 1.6�� longer than F2 (Fig. 4E). Mesosoma longer than high in lateral view. Propleuron alutaceous, with few setae. Mesoscutum smooth, glabrous between notauli, alutaceous lateral to notaulus; longer than broad; notauli complete, deeply impressed for full length; median mesoscutal line distinct; anterior parallel lines and parapsidal lines absent; mesoscutellum only slightly longer than broad, slightly narrower posteriorly; shorter than mesoscutum, uniformly rugose, overhanging metanotum; scutellar foveae deeply excavated, fully delimited on all sides, separated narrowly by carina (Fig. 4H). Mesopleuron smooth, glabrous, with a few white setae along ventral and anterior margins. Lateral propodeal carinae distinct, straight; central propodeal area punctate; lateral propodeal area alutaceous, with dense white setae; nucha short, coriaceous (Fig. 4F). Tibia setose on anterior edge; fore tibia prolonged on the anterior side into a curved spine, much shorter than tibial spur and basitarsus; tarsal claws simple with a slight ridge but never a full tooth. Middle and hind tibia with two spurs (Fig. 4E). Fore wing hyaline, longer than body, margin with dense cilia; radial cell 2.3�� as long as wide; 2r infumated, Rs curved upwards and thickened at apex; radial cell open; areolet small and indistinct; Rs + M reaching to M; cu-a absent; cu1 broken (Fig. 4E). Metasoma shorter than head + mesosoma, 1.2�� longer than high in lateral view, smooth and glabrous; second metasomal tergite setose medially; all subsequent tergites without setae, smooth, glossy. Ventral spine of the hypopygium short, prominent part 1.5�� as long as broad in ventral view, with white setae extending beyond the apex of spine (Fig. 4E). Male: Body length 2.6���3.9 mm (N = 16). Colour darker than sexual female, scape, pronotum, mesoscutum and mesopleuron, metasoma dark brown, sculptures like the sexual female. Antenna 15 segmented; F1 is curved, excavated, and incised medially. Metasoma smaller than head + mesosoma; see Lund et al. (1998: fig. 7) for comparison of lateral habitus of sexual generation male and female. Gall: Smooth, unilocular pea-like galls on the ventral side of leaves for the asexual generation (Fig. 1E; Lund et al. 1998: figs 1, 2), irregular shaped, multilocular clusters of galls on the small rootlets for the sexual generation (Lund et al. 1998: figs 3, 4). Host plant: Quercus fusiformis (Oklahoma, Texas, Mexico) and Q. virginiana (east Texas to Florida, north to North Carolina). Distribution: Louisiana, Mississippi, Oklahoma, Texas. Biology: The asexual generation has long, straight wings and is capable of flight (Table 2). Given the distribution of its host Q. fusiformis (Cavender-Bares et al., 2015), this species likely also occurs in northern Mexico. Remarks: Belonocnema kinseyi was previously synonymized by Lund et al. (1998) based on experimental rearing that linked the asexual and sexual generations developing on Q. fusiformis in Texas. Given the molecular and morphological evidence showing that populations west of the Gulf of Mexico represent a distinct species, the synonymy by Lund et al. (1998) is, therefore, rejected and B. kinseyi 17.15 is restored as a valid species. The genome of this species, B. kinseyi, has been sequenced and annotated (NCBI SRA: PRJNA 623416) originally under the name B. treatae., Published as part of Zhang, Y. Miles, Egan, Scott P., Driscoe, Amanda L. & Ott, James R., 2021, One hundred and sixty years of taxonomic confusion resolved: Belonocnema (Hymenoptera: Cynipidae: Cynipini) gall wasps associated with live oaks in the USA, pp. 1234-1255 in Zoological Journal of the Linnean Society 193 (4) on pages 1248-1250, DOI: 10.1093/zoolinnean/zlab001, http://zenodo.org/record/5761660, {"references":["Weld LH. 1921. American gallflies of the family Cynipidae producing subterranean galls on oak. Proceedings of the United States National Museum 59: 187 - 246.","Lund JN, Ott JR, Lyon RJ. 1998. Heterogony in Belenocnema treatae Mayr (Hymenoptera: Cynipidae). Proceedings of the Entomological Society of Washington 100: 755 - 763.","Cavender-Bares J, Gonzalez-Rodriguez A, Eaton DA, Hipp AA, Beulke A, Manos PS. 2015. Phylogeny and biogeography of the American live oaks (Quercus subsection Virentes): a genomic and population genetics approach. Molecular Ecology 24: 3668 - 3687."]}

Phenological differences between host plants can promote temporal isolation among host-associated populations of insects with life cycles tightly coupled to plant phenology. Divergence in the timing of spring budbreak between two sympatric sister oak species has been shown to promote temporal isolation between host plants and their host-associated populations of a cynipid gall wasp. Here, we examined the generality of this mechanism by testing the hypothesis of cascading temporal isolation for five additional gall-formers and three natural enemy species associated with these same oak species. The timing of adult emergence from galls differed significantly between host-associated populations for all nine species and parallels the direction of the phenological differences between host plants. Differences in emergence timing can reduce gene flow between host-associated populations by diminishing mating opportunities and/or reducing the fitness of immigrants due to differences in the availability of ephemeral resources. Our study suggests that cascading temporal isolation could be a powerful ‘biodiversity generator’ across multiple trophic levels in tightly coupled plant–insect systems.