One hundred and sixty years of taxonomic confusion resolved: Belonocnema (Hymenoptera: Cynipidae: Cynipini) gall wasps associated with live oaks in the USA Author Zhang, Y. Miles Author Egan, Scott P. Author Driscoe, Amanda L. Author Ott, James R. text Zoological Journal of the Linnean Society 2021 2021-02-27 193 4 1234 1255 https://doi.org/10.1093/zoolinnean/zlab001 journal article 3109 10.1093/zoolinnean/zlab001 9182a649-923b-4a69-bac0-d41873ded32e 0024-4082 5761660 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 .