Feron atrimentum ( Kinsey, 1922 ), 2023

Cuesta-Porta, Victor, Melika, George, Nicholls, James A., Stone, Graham N. & Pujade-Villar, Juli, 2023, Re-establishment of the Nearctic oak cynipid gall wasp genus Feron Kinsey, 1937 (Hymenoptera: Cynipidae: Cynipini), including the description of six new species, Zootaxa 5366 (1), pp. 1-174 : 26-34

Feron atrimentum ( Kinsey, 1922)

Figs 37–67

Andricus atrimentus Kinsey, 1922: 279 , female, male, gall.

Dros atrimentum (Kinsey) : Weld, 1951: 629.

Types examined. HOLOTYPE: sexual female “Three Rivers Cal., gall 3.23.20.’, ‘ Q. douglasii, Kinsey coll.”, red ‘ And. atrimentus Holo- COTYPE” deposited in AMNH, NYC, examined by GM . PARATYPES (2 sexual females) in the general collection in AMNH also examined by GM .

Additional material. Asexual females: 27 females label as “ USA: CA, Dye Creek Preserve, CA1170 , galltype 137; Q. douglasii ; leg. J.A. Nicholls, 2001.09.27 ” ; 2 females “ USA: CA, Del Puerto Canyon , Q. douglasii ; leg. R. Challis, 2004.10.21 ” ; 4 females “ USA: CA, Clearlake , CA941 , galltype 137; Q. douglasii ; leg. J.A. Nicholls, 2007.11.03 ”. Sexual generation : 3 females “ USA: CA, Quail Ridge Reserve , CA1044 , galltype 231, Q. douglasii ; leg. J.A. Nicholls, 2008.04.04 ” ; 55 females and 37 males “ USA: CA, Dye Creek Preserve, CA1048 , 1049 , galltype 231; Q. douglasii ; leg. J.A. Nicholls, 2008.03.30 ” .

Diagnosis. Asexual females belong to the group of Feron species with the pronotum laterally smooth to coriaceous, without carinae; the mesoscutum is alutaceous to coriaceous, rugose-reticulate, reticulate, sometimes with smooth areas and piliferous points, glabrous or pubescent as occurs in F. albicomus (asex.), F. crystallinum (asex), F. izabellae (asex), F. pattersonae (asex) and F. sulfureum (asex). Feron atrimentum differs from F. albicomus and F. izabellae in having the ocelli elevated above the head and the pronotum laterally with setae on the entire lateral surface; differs from F. crystallinum , F. pattersonae and F. sulfureum in the pronotum which is smooth (coriaceous in the other three species); the mesoscutum is longer than broad, scarcely pubescent, without black stripes and the anterior parallel line is indistinct. For other characters consult couplet 24 in the key. Sexual females belong to the group of Feron species in which the mesoscutum is alutaceous to coriaceous, rugose-reticulate, reticulate, without piliferous points and the pronotum laterally with longitudinal carinae as in F. splendens (asex), F. vitreum (asex) and F. verutum (asex); nevertheless, it differs from these asexual forms by having a black body, eyes are strongly converging ventrally, the transfacial distance shorter than the height of eye, antennal toruli located slightly above half the eyes’ height and the eye around 6.0× as high as length of the malar space. Males differs from all known Feron males in having strongly elevated ocelli above the frons in frontal view, OOL shorter than the diameter of lateral ocellus and inner margins of eyes subparallel, the mesoscutum is uniformly reticulate between notauli in anterior half and laterad to the notaulus but smooth between notauli in the posterior half.

Re-description. Asexual female ( Figs 37–48 ). Body uniformly brown, with darker propodeum.

Head with sparse setae on lower face, gena and posteriorly, 1.2× as broad as high and slightly narrower than mesosoma in frontal view; 1.9× as broad as long in dorsal view. Gena alutaceous-reticulate, not broadened behind eye in frontal view, narrower than transverse diameter of eye in lateral view. Malar space alutaceous, shining, with short striae radiating from clypeus reaching to 1/3 of malar space length; eye 3.0× as high as length of malar space; malar sulcus absent. Inner margins of eyes parallel. POL 2.6× as long as OOL, OOL 1.7× as long as diameter of lateral ocellus and slightly shorter than LOL, all ocelli ovate, of same size. Antennal toruli located above mid-height of eyes. Transfacial distance 1.2× as long as height of eye; diameter of antennal torulus slightly longer than distance between them, distance between torulus and eye as long as diameter of torulus; lower face smooth with white setae; slightly elevated median area smooth, without setae. Clypeus impressed, rectangular, 1.75× as broad as high, delicately coriaceous, with a few setae scattered all over; ventrally rounded, emarginate, without median incision; anterior tentorial pit large, rounded, deep, epistomal sulcus broad and deep, clypeo-pleurostomal line well impressed. Frons, interocellar area, vertex uniformly reticulate, without striae and setae; area under central ocellus impressed, smooth, glabrous; occiput alutaceous, with dense setae; postocciput glabrous, with numerous delicate longitudinal interrupted parallel striae; postgena smooth, with a few setae; posterior tentorial pit large, elongated, area below impressed; occipital carina black, well-visible alongside occipital foramen; occipital foramen as high as height of postgenal bridge; hypostomal carina emarginate, continuing into strong postgenal sulci which diverge until half height of postgenal bridge, further sulci parallel and running alongside foramen until 1/3 of its height. Antenna slightly longer than head+mesosoma, with 12 flagellomeres; flagellomeres gradually broadened towards apex; F1–F3 with sparse white setae, F4–F12 with dense setae; pedicel 1.7× as long as broad; F1 2.2× as long as pedicel and 1.4× as long as F2; F2=F3=F4; F5=F6 and shorter than previous; all subsequent flagellomeres shorter and equal in length; F12 as long as F11, suture between F11 and F12 distinct; placodeal sensilla on F4–F12.

Mesosoma longer than high, with white setae, setae denser on pronotum laterally, mesopleural triangle and mesoscutellum. Pronotum smooth without striae, with sparse setae, piliferous points along dorsal edge; propleuron smooth, with sparse white setae. Mesoscutum longer than broad (greatest width measured across mesoscutum level with base of tegulae), alutaceous-reticulate between notauli in anterior 2/3 of length and laterad to notauli, smooth, glabrous in between notauli in posterior half. Notaulus complete, deep, posteriorly strongly converging and broader than anteriorly, with smooth, glabrous bottom; at posterior end the distance between notauli shorter than distance between notaulus and side of mesoscutum; anterior parallel line indistinct; parapsidal line absent, not marked; median mesoscutal line absent; parascutal carina broad, reaching notaulus. Mesoscutellum trapezoid, longer than broad, broadest part in posterior 1/3; center part of mesoscutellum disc rugoso-coriaceous, with stronger rugae posteriorly and laterally, overhanging metanotum, with long setae. Mesoscutellar foveae ovate, broader than high, with smooth, glabrous bottom, divided by a elevated narrow carina. Mesopleuron entirely smooth, with setae only along ventral edge; mesopleural triangle smooth, with dense white setae and piliferous points; dorsal and lateral axillar areas smooth, with setae; axillula with delicate parallel longitudinal striae; subaxillular bar smooth, glabrous, with subparallel sides, posteriorly as high as height of metanotal trough; metapleural sulcus reaching mesopleuron at half of its height, lower part delimiting smooth area with a few setae, upper part of sulcus absent. Metascutellum coriaceous, as high as height of smooth, glabrous ventral impressed area; metanotal trough smooth, with few setae; central propodeal area lyre-shaped, smooth, glabrous; lateral propodeal carinae strong, bent outwards in posterior 1/3; lateral propodeal area smooth, with long white setae. Nucha with longitudinal sulci dorsally and laterally. Tarsal claws with acute basal lobe.

Fore wing longer than body, hyaline, with short dense cilia on margin, veins brown, radial cell open, 4.2× as long as broad; Rs and R1 nearly reaching wing margin; areolet absent. Rs+M indistinct, visible on 2/3 of distance between areolet and basalis, its projection reaching basalis at half of its height.

Metasoma longer than head+mesosoma, higher than long in lateral view; 2nd metasomal tergum extending nearly to half-length of metasoma in dorsal view, with patch of dense white setae anterolaterally, without micropunctures; all subsequent terga and hypopygium with fine micropunctures; prominent part of ventral spine of hypopygium 6.7× as long as broad in ventral view, with a few short white setae ventrally.

Body length 1.7–1.9 mm (n = 2).

Sexual female ( Figs 51–61 ). Head, mesosoma, metasoma black; antenna dark brown, except yellowish scape, pedicel and F1; metasoma chestnut, hypopygium light brown; legs yellow with dark brown to black coxae.

Head with sparse setae, denser on lower face and postgena, 1.2× as broad as high and slightly broader than mesosoma in frontal view; 2.1× 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, without striae; eye 6.1× as high as length of malar space; malar sulcus absent. Inner margins of eyes strongly converging ventrally. POL 2.4× as long as OOL, OOL 1.2× as long as diameter of lateral ocellus and equal in length to LOL, all ocelli ovate, of same size. Transfacial distance shorter than height of eye; toruli located at half of eyes height, frons shorter than lower face, diameter of antennal torulus 1.3× as long as distance between them, distance between torulus and eye as long as diameter of torulus; lower face smooth with white setae, alutaceous in most ventral part above malar space; slightly elevated median area and area between toruli alutaceous. Clypeus trapezoid, as broad as high, delicately coriaceous, with a few long setae along ventral edge; ventrally emarginate, with median incision; anterior tentorial pit large, ovate, epistomal sulcus distinct, deep, clypeo-pleurostomal line well impressed, deep. Frons, interocellar area, vertex uniformly reticulate, without striae and setae; area under central ocellus impressed, alutaceous; occiput and postgena alutaceous; postocciput with numerous delicate interrupted parallel striae dorsally and laterally of occipital foramen; posterior tentorial pit large, elongated, area below impressed; occipital foramen as high as height of postgenal bridge; hypostomal carina emarginate, continuing into strong postgenal sulci which diverge strongly toward occipital foramen, postgenal bridge anteriorly slightly broader than occipital foramen. Antenna slightly longer than head+mesosoma, with 12 flagellomeres; F1–F3 with rare setae, subsequent flagellomeres with denser setae; scape, pedicel, F1 yellowish brown, all subsequent flagellomeres dark brown; pedicel only slightly longer than broad; F1 2.5× as long as pedicel and slightly longer than F2; F2 1.2× as long as F3, F3=F4, all subsequent flagellomeres nearly equal in length, F12 longer than F11; placodeal sensilla on F3–F12.

Mesosoma longer than high, with a few white setae, denser on mesoscutellum and lateral propodeal area. Pronotum delicately coriaceous, with sparse setae and delicate parallel striae laterally; propleuron alutaceous. Mesoscutum slightly longer than broad (greatest width measured across mesoscutum level with base of tegulae), uniformly reticulate between notauli in anterior half and laterad to notauli, smooth and glabrous in between notauli in posterior half. Notaulus complete, shallow, posteriorly converging and broader than anteriorly, bottom smooth, glabrous; in most posterior part distance between notauli shorter than distance between notaulus and side of mesoscutum; anterior parallel line distinct, smooth, glabrous, reaching to 1/3 length of mesoscutum; parapsidal line marked with broad smooth, glabrous stripe; median mesoscutal line absent; parascutal carina broad, reaching notaulus. Mesoscutellum trapezoid, longer than broad, broadest part in posterior 1/3, uniformly rugoso-coriaceous, overhanging metanotum, with sparse long setae. Mesoscutellar foveae quadrangular, as broad as high, with smooth, glabrous bottom, divided by a rugose elevated triangular area. Mesopleuron entirely smooth, with setae only along ventral edge; mesopleural triangle smooth, with dense white setae; dorsal and lateral axillar areas smooth, glabrous, without setae; axillula with delicate parallel longitudinal striae; subaxillular bar smooth, glabrous, with parallel sides, posteriorly shorter than height of smooth, glabrous metanotal trough; metapleural sulcus reaching mesopleuron at half of its height, upper part of sulcus indistinct, lower part of sulcus delimiting smooth, glabrous area, with few setae. Metascutellum coriaceous, slightly higher than height of smooth, glabrous ventral impressed area; metanotal trough smooth, with few setae and few rugae; central propodeal area smooth, glabrous, with a few strong short longitudinal rugae posteriorly; lateral propodeal carinae strong, broad and high, bent strongly outwards at mid-height; lateral propodeal area smooth, with long white setae, each seta with piliferous point at the base. Nucha with numerous irregular rugae. Tarsal claws with small basal lobe.

Fore wing longer than body, hyaline, with short cilia on margin, veins brown, radial cell open, 4.3× as long as broad; Rs and R1 reaching wing margin; areolet distinct, triangular. Rs+M distinct, reaching basalis slightly below half of its height.

Metasoma slightly longer than head+mesosoma, longer than high in lateral view; 2nd metasomal tergum extending to 1/3 length of metasoma in dorsal view, with patch of white setae anterolaterally, without micropunctures; all subsequent terga and hypopygium without micropunctures; prominent part of ventral spine of hypopygium 4.5× as long as broad in ventral view, with short white setae ventrally.

Body length 2.7–3.1 mm (n = 10).

Male ( Figs 62–67 ). Like female but antenna longer than body, with 13 flagellomeres, F1 with longitudinal yellow stripe on the inner side, placodeal sensilla on all flagellomeres, F13 slightly shorter than F12. Eye and ocelli larger than in female; POL 3.0× as long as OOL, OOL slightly longer than diameter of ocellus and slightly shorter than LOL; transfacial distance shorter than height of eye, height of eye 6.0× as long as length of malar space.

Body length 2.2–2.7 mm (n = 10).

Gall. The sexual gall ( Fig. 49 ) is monolocular, spherical, 3–5 mm in diameter, smooth, green when growing, becoming paler later, mature galls when dried are yellow-brown; larval cell is blue-black and sometimes visible though outer layers of the gall. Walls of the gall are thin, the larval chamber is also thin-walled, 1.5 mm in diameter and connected with the outside wall by a tangle of dense fibers. The gall is in the leaf tissue, projecting equally on both sides. Asexual galls ( Fig. 50 ) are monolocular, conical, flat-based, detachable, on lower leaf margin. Sides nearly straight, flaring slightly at base. Red and yellow striped when fresh, turning to brown and yellow with age. Basal larval cell oval from lateral view, 1.6 mm in diameter, 0.8 mm high, pubescent. Upper surface of larval cell coriaceous with sparse pubescence; brown, darker towards center. Gall height 3.6–4.6 mm, width 3.0– 4.6 mm.

Biology. Matching of generations was first presented in Dailey & Sprenger (1973b). Alternate sexual and asexual generations are also confirmed herein using DNA data, with four individuals (three asexual females, one sexual female) sequenced for cytb. Cytb sequences were on average 0.35% divergent between individuals (range 0–0.70%; GenBank accessions OK041496, OQ446191–OQ446193).

Feron atrimentum is associated with Q. douglasii , Q. dumosa , Q. john-tuckeri and Q. lobata (section Quercus , subsection Dumosae). The sexual generation females emerge throughout April and usually oviposit in the lower leaf surface near the margin. Asexual galls develop over the following three months. Pupation occurs during late October to November; asexual females emerge in late winter and oviposit in leaf buds. Sexual generation galls usually reach full size by mid-March ( Dailey & Sprenger 1973b).

Distribution. USA: California.

Figures

FIGURES 37–42. Feron atrimentum, comb. nov., asexual female. 37–40, head: 37, frontal view, 38, dorsal view, 39, lateral view, 40, posterior view. 41, antenna. 42, mesosoma and propleura, frontal view.

FIGURES 43–46. Feron atrimentum, comb. nov., asexual female. 43, mesosoma, lateral view. 44, mesoscutum, dorsal view. 45, mesoscutellum, dorsal view. 46, metascutellum and propodeum, posterodorsal view.

FIGURES 47–50. Feron atrimentum, comb. nov.. 47–48, asexual female: 47, fore wing, part, 48, metasoma, lateral view. 49, sexual gall, 50, asexual gall.

FIGURES 51–56. Feron atrimentum, comb. nov., sexual female. 51–54, head: 51, frontal view, 52, dorsal view, 53, lateral view, 54, posterior view. 55, antenna. 56, mesosoma and propleura, frontal view.

FIGURES 57–60. Feron atrimentum, comb. nov., sexual female. 57, mesosoma, lateral view. 58, mesoscutum, dorsal view. 59, mesoscutellum, dorsal view. 60, metascutellum and propodeum, posterodorsal view.

FIGURES 61–67. Feron atrimentum, comb. nov., sexual generation. 61, sexual female, metasoma, lateral view. 62–67, male: 62–65, head: 62, frontal view, 63, dorsal view, 64, lateral view, 65, posterior view. 66, antenna. 67, metasoma, lateral view.