Feron kingi ( Bassett, 1900 ) Cuesta-Porta & Melika & Nicholls & Stone & Pujade-Villar, 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 : 82-90

Feron kingi ( Bassett, 1900) , comb. nov.

Figs 228–255

Andricus kingi Bassett, 1900: 316 , female (asex), gall.

Andricus pistillaris Trotter, 1910: 117 , female (asex), gall; synonymized in Weld, 1951: 634. Type examined. Andricus kingi Bassett : Type No. 10495, asexual female, deposited at the Entomology Type Collection at the Academy of Natural Sciences of Philadelphia, not examined by the authors.

Additional material. Asexual females: 10 females “ USA, CA, Stockton, Q. lobata , leg. K. Schick, 2004.02.29 ”; 6 females “ USA, CA, Dye Creek Preserve, Q. lobata , leg. J. de Martini, 2001.09.22.”; 28 females “ USA, CA, Clearlake , CA29 , galltype 160, Q. lobata , leg. J.A. Nicholls, 2007.11.03.”; 46 females “ USA, CA , Delta College , Stockton, CA905 , CA906 , galltype 160, Q. lobata , leg. J.A. Nicholls, 2007.11.06.”; 9 females “ USA, CA , Cosumnes River Preserve, CA927 , 906 , galltype 160, Q. lobata , leg. J.A. Nicholls, 2007.11.07.” Sexual generation: 6 males “ USA, CA , Dye Creek Preserve, CA1050 , galltype 237, Q. douglasii , leg. J.A. Nicholls, 2008.03.30 ”; 2 females and 1 male “ USA, CA, Quail Ridge Reserve, CA1069 , galltype 237, Q. berberidifolia , leg. J.A. Nicholls, 2008.04.04 ”; 1 female and 3 males “ USA, CA, Delta college, Stockton, CA1047 , galltype 237, Q. lobata , leg. J.A. Nicholls, 2008.03.28 ”. 3 females and 10 males: 6 males “ USA, CA, Dye Creek Preserve, Q. douglasii , coll. J.A. Nicholls, 2008.03.30; CA1050 ”, 2 females and 1 male “ USA, CA, Quail Ridge Reserve, Q. berberidifolia , coll. J.A. Nicholls, 2008.04.04; CA1069 ”; 1 female and 1 male “ USA, CA, Delta College, Stockton, Q. lobata , 2008.03.28; CA1047 ”; 2 males “ USA, CA, Cosumnes River Preserve, Q. lobata , coll. J.A. Nicholls, 2008.03.28; CA1060 ”. Specimens have been deposited at the USNM and PHDNRL .

Diagnosis. Asexual females belong to the Feron species group in which the pronotum laterally has longitudinal carinae and the mesoscutum is alutaceous to coriaceous, rugose-reticulate, reticulate, with piliferous points; as in F. discularis (asex) and F. tubifaciens (asex). Differs from F. discularis in the body colour, which is never black; the frons not bulging in frontal view, ocelli are not elevated above the head; toruli located in the upper half of the head; the median mesoscutal line is absent. The most similar species is F. tubifaciens but differs in colour, the gena is broadened behind the eye in frontal view, the clypeus is smooth, the fore wing longer than the body; for other characters see couplet 29 in the key. Sexual females belong to the group of Feron species with a black body, scape to F2 are dark brown (sometimes light brown but never yellowish), legs are reddish brown with at least the hind coxa darker, the frons is flat not or only slightly bulging in frontal view; toruli are located in the upper half of the head, inner margins of eyes never strongly converging ventrally, the mesoscutum is smooth or partially alutaceous anteriorly and glabrous, the notaulus is complete, the mesopleuron entirely smooth without transverse striae in the central part, the mesoscutellum uniformly alutaceous with numerous setae on piliferous points and the prominent part of the ventral spine of the hypopygium with setae; as in F. apiarium (asex) and F. pattersonae (sex). Differs from F. apiarium as the eyes converge ventrally, the eye at least 4.0× as high as length of the malar space, antennomeres with short setae, OOL 1.5× as long as diameter of the lateral ocellus, the notaulus sometimes weakly impressed anteriorly and the prominent part of the ventral spine of the hypopygium with few short setae ventrally. Most similar morphologically is F. pattersonae ; see diagnostic characters at couplet 17 in the key. Males are characterised by a black body, ocelli are moderately or not elevated above the frons, the notaulus reaches the pronotum and the mesoscutellum is uniformly alutaceous with numerous setae on piliferous points; these characters are as in F. comatum and F. pattersonae , but in F. kingi F1 longer than scape+pedicel, straight, not broadened and curved; all flagellomeres, scape and pedicel are uniformly coloured (F1 equal in length to scape+pedicel, slightly broadened and curved; all flagellomeres lighter than the scape and pedicel in F. comatum ) and the head is ovate in frontal view, inner margins of eyes are parallel, space between central elevated area of the lower face to the lateral margin of the eye is sculptured, OOL shorter than the diameter of the lateral ocellus (triangular head, inner margins of eyes converging vertrally, central elevated area smooth and OOL as long as diameter of the lateral ocellus in F. pattersonae ).

Re-description. Asexual female ( Figs 228–238 ). Body, antenna, legs, uniformly rusty brown, some specimens uniformly dark brown.

Head with white setae, denser on lower face, occiput, postocciput and postgena, slightly broader than high and as broad as mesosoma in frontal view; 1.9× as broad as long in dorsal view. Gena alutaceous, broadened behind eye in frontal view, narrower than transverse diameter of eye in lateral view. Malar space with striae radiating from clypeus and reaching eye; eye 2.9× as high as length of malar space; malar sulcus absent. Inner margins of eyes parallel. POL 2.2× as long as OOL, OOL 1.6× as long as diameter of lateral ocellus and slightly shorter than LOL, all ocelli ovate, lateral ocelli slightly larger than central ocellus. Transfacial distance slightly longer than height of eye; toruli located in the upper half of head and frons definitely shorter than lower face, diameter of antennal torulus 1.9× as long as distance between them, distance between torulus and eye nearly as long as diameter of torulus; lower face smooth with white setae; slightly elevated median area and area between toruli smooth. Clypeus ovate, slightly broader than high, smooth, with a few long setae scattered all over; ventrally rounded, emarginate, without median incision; anterior tentorial pit small, rounded, distinct, epistomal sulcus distinct, clypeo-pleurostomal line well impressed. Frons, interocellar area, vertex, occiput uniformly reticulate, without striae and setae; area under central ocellus impressed, smooth, glabrous; postocciput and postgena reticulate, with a few setae; posterior tentorial pit large, elongated, area below impressed; occipital foramen as high as height of postgenal bridge; hypostomal carina emarginate, continuing into distinct postgenal sulci which strongly diverge toward occipital foramen, postgenal bridge anteriorly slightly broader than occipital foramen. Antenna longer than head+mesosoma, with 12 flagellomeres; all flagellomeres with dense setae; pedicel 1.7× as long as broad; F1 2.0× as long as pedicel and 1.4× as long as F2; F2=F3; F4 to F6 equal in length, F7 to F11 equal in length, F12 slightly shorter than F11; placodeal sensilla on F4–F12.

Mesosoma slightly longer than high, with a few white setae. Propleura alutaceous with setae. Pronotum smooth, glabrous dorsally; laterally coriaceous, with dense setae; propleuron smooth, with dense white setae. Mesoscutum longer than broad (greatest width measured across mesoscutum level with base of tegulae), uniformly coriaceous; entirely and uniformly with scattered long white setae. Notaulus complete, shallow, posteriorly converging and broader than anteriorly, bottom smooth, glabrous; at posterior end the distance between notauli shorter than distance between notaulus and side of mesoscutum; anterior parallel line and parapsidal line indistinct or invisible; median mesoscutal line absent; parascutal carina broad, reaching notaulus. Mesoscutellum slightly longer than broad, with subparallel sides, broadest part in posterior 1/3; mesoscutellum rugose-coriaceous, overhanging metanotum, with dense long setae. Mesoscutellar foveae ovate, broader than high, with smooth, glabrous bottom, delimited all around by strong black carina. Mesopleuron entirely smooth, with setae only along ventral edge and some sparse setae along mid-height; 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 as high as height of metanotal trough; metapleural sulcus reaching mesopleuron at half of its height, upper part of sulcus also distinct, sulcus separating smooth, glabrous area, with dense setae in lower part. Metascutellum rugose, higher than height of smooth, glabrous ventral impressed area; metanotal trough smooth, with dense setae; central propodeal area lyre-shaped, smooth, glabrous, with a few strong short irregular rugae; lateral propodeal carinae strong, broad and high, bent outwards in posterior 1/3; lateral propodeal area smooth, with long white setae. Nucha with net of numerous irregular rugae. Tarsal claws with acute basal lobe.

Fore wing longer than body, hyaline, with short cilia on margin, veins brown, radial cell open, 4.4× as long as broad; Rs reaches wing margin, R1 nearly reaching wing margin; areolet absent. Rs+M indistinct, visible on 1/3 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 to half-length of metasoma in dorsal view, with patch of dense white setae anterolaterally, with inconspicuous micropunctures extending to half-length of tergum; all subsequent terga until tergum 5 and hypopygium without micropunctures; tergum 6 and 7 with micropunctures, prominent part of ventral spine of hypopygium 6.8× as long as broad in ventral view, with short white setae ventrally.

Body length 1.5–2.1 mm (n = 10).

The description of the sexual generation is given below for the first time.

Sexual female ( Figs 241–244 , 250–255 ). Head, mesosoma, metasoma dark brown, antenna slightly lighter; mouthparts, legs light brown with darker coxae.

Head alutaceous, with a few setae, denser on lower face, rounded, 1.2× as broad as high and slightly broader than mesosoma in frontal view; 2.2× as broad as long in dorsal view. Gena alutaceous, slightly broadened behind eye in frontal view, narrower than tranverse diameter of eye in lateral view. Malar space alutaceous, shining, glabrous, without sulcus and striae radiating from clypeus and reaching eye; eye 4.4× as high as length of malar space. Eyes slightly converging ventrally. POL 2.4× as long as OOL, OOL 1.5× as long as diameter of lateral ocellus and 0.8× as long as LOL, all ocelli slightly ovate, of same size. Transfacial distance as long as height of eye or slightly shorter; diameter of antennal torulus 1.4× as long as distance between them, distance between torulus and eye slightly shorter than diameter of torulus; lower face smooth, with dense white setae, without striae; slightly elevated median area delicately coriaceous, with few setae. Clypeus rectangular, nearly 2.0× as broad as high, smooth, with long setae; ventrally rounded, not emarginate and without median incision; anterior tentorial pit large, rounded, distinct, epistomal sulcus distinct, clypeo-pleurostomal line well impressed. Frons uniformly alutaceous, without striae and setae; interocellar area alutaceous. Vertex, occiput and postocciput alutaceous; postgena alutaceous, without setae, with longitudinal delicate parallel striae along gular sulci; posterior tentorial pit large, elongated, area below impressed; occipital foramen slightly shorter than height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which diverge strongly toward occipital foramen, postgenal bridge anteriorly broader than occipital foramen. Antenna longer than head+mesosoma, with 11 flagellomeres, pedicel longer than broad; F1 1.7x as long as pedicel and 1.2× as long as F2; F2 1.1× as long as F3; F3 slightly longer than F4, F5 1.1× as long as F6, subsequent flagellomeres nearly equal in length, F11 1.7× as long as F10; placodeal sensilla on F5–F11, absent on F1–F4.

Mesosoma 1.2× as long as high, with sparse white setae, except dense setae on lateral propodeal area. Pronotum smooth, with a few sparse setae, with some delicate striae laterally; propleuron alutaceous, glabrous. Mesoscutum smooth, with sparse white setae laterally and along notaulus, slightly longer than broad (greratest width measured across mesoscutum level with base of tegulae). Notaulus complete, deep, bottom smooth, posteriorly broader and strongly converging; at posterior end the distance between notauli shorter than distance between notaulus and side of mesoscutum; anterior parallel line invisible; parapsidal line distinct, marked with broad slightly impressed line; median mesoscutal line absent; parascutal carina broad, reaching notaulus. Mesoscutellum ovate, slightly longer than broad; disk of mesoscutellum smooth, glabrous, dull rugose laterally and posteriorly, overhanging metanotum, with sparse long setae. Mesoscutellar foveae in the form of a narrow semi-lunar depression, not separated medially, with smooth, glabrous bottom. Mesopleuron entirely smooth, with setae in posteroventral part; mesopleural triangle smooth, glabrous, with some irregular striae and a few setae; dorsal and lateral axillar areas smooth, glabrous, with dense white setae; subaxillular bar smooth, glabrous, with parallel sides, as high as height of metanotal trough; metapleural sulcus reaching mesopleuron at half of its height, upper part of sulcus distinct, sulcus separating smooth, glabrous area, with a few setae in lower part. Metascutellum delicately coriaceous, 3.0× as high as height of smooth, glabrous ventral impressed area; metanotal trough smooth, with some setae; central propodeal area lyre-shaped, smooth, glabrous, with net of irregular interrupted rugae; lateral propodeal carinae distinct, bent outwards in posterior 1/3; lateral propodeal area smooth, with long dense white setae. Nucha with numerous sulci dorsally and laterally. Tarsal claws with very tiny basal lobe.

Fore wing longer than body, hyaline, with distinct dense cilia on margin, veins light brown, radial cell open, 4.6× as long as broad; R1 and Rs reaching wing margin; areolet small, triangular, closed, indistinct. Rs+M narrow, inconspicuous, its projection reaching basalis slightly below its mid height.

Metasoma as long as head+mesosoma, slightly longer than high in lateral view; 2nd metasomal tergum extending to half-length of metasoma in dorsal view, with a few short white setae and without micropunctures laterally; subsequent terga smooth, glabrous, without micropunctures. Hypopygium without micropunctures, prominent part of ventral spine of hypopygium 2.0× as long as broad in ventral view, with some short setae ventrally which do not extend beyond apex of spine.

Body length 2.0– 2.2 mm (n = 5).

Male ( Figs 245–249 ). Similar to female, body slightly darker; eye 4.8× as high as length of malar space; POL 3.6× as long as OOL, OOL 0.6× as long as LOL, diameter of lateral ocellus 1.4× as long as OOL, ocelli ovate, larger than in female; interocellar area elevated in frontal view; antenna longer than body, with 13 flagellomeres, F1 straight, not broadened and curved, F1 1.3× as long as F2, F2=F3, all subsequent flagellomeres nearly equal in length, F13=F12; placodeal sensilla on all flagellomeres.

Body length 1.7–1.9 mm (n = 5).

Galls. The sexual gall ( Fig. 239 ) is small, 2–3 mm, conical gall on the leaf margin and/or catkin, slightly pubescent. Fresh galls are pinkish to pale brown, turn dark purple-black when mature, the leaf blade twists slightly at the gall. The gall is difficult to distinguish from similar sexual galls of F. comatum , F. crystallinum and F. dumosae . The asexual gall ( Fig. 240 ) is red, monolocular, detachable, cone-shaped on both sides of the leaves, 5 mm high and 3–5 mm broad at the base. The cone rises from a flared, cup-shaped base, which is narrowly attached to the leaf. The apex of the gall is pointed, with a blunt tip. The gall is covered by fine pubescence; the larval chamber is large, occupying the most of the gall base. Occurs singly or in groups, up to dozens ( Russo 2006, 2021).

Biology. Liodora dumosae was erroneously named as the sexual generation of A. kingi ( Rosenthal 1968; Rosenthal & Koehler 1971, Burks 1979), but subsequent detailed examination of the type material showed that L. dumosae is a distinct species and Rosenthal’s material represented the previously unknown and undescribed sexual generation of A. kingi ( Dailey & Menke 1980; our examinations); herein we formally describe sexual A. kingi for the first time. Alternate sexual and asexual generations are also confirmed herein on the basis of similarity in cytb sequence data. Four individuals (two asexual females, one sexual female, one sexual male) had cytb sequences that were on average 0.23% divergent (range 0–0.47%; GenBank accessions OK041496, OQ446201–OQ446203).

Asexual galls have been collected from Q. douglasii , Q. dumosa , Q. garryana , and Q. lobata (section Quercus , subsection Dumosae) in late September and October; adults emerge early the following spring ( Burks 1979). Mature sexual galls have been collected in late March and the beginning of April from Q. berberidifolia , Q. douglasii , and Q. lobata (section Quercus , subsection Dumosae); adults emerge soon after under laboratory conditions.

Distribution. USA: California.

Figures

FIGURES 228–233. Feron kingi, comb. nov., asexual female. 228–231, head: 228, frontal view, 229, dorsal view, 230, lateral view, 231, posterior view. 232, antenna. 233, mesosoma and propleura, frontal view.

FIGURES 234–237. Feron kingi, comb. nov., asexual female. 234, mesosoma, lateral view. 235, mesoscutum, dorsal view. 236, mesoscutellum, dorsal view. 237, metascutellum and propodeum, posterodorsal view.

FIGURES 238–240. Feron kingi, comb. nov., 238, asexual female, metasoma, lateral view. 239–240, gall: 239, sexual galls, 240, asexual galls.

FIGURES 241–249. Feron kingi, comb. nov., sexual generation. 241–244, sexual female: 241–243 head: 241, frontal view, 242, dorsal view, 243, lateral view. 244, antenna. 245–249 male: 245–248, head: 245, frontal view, 246, posterior view, 247, dorsal view, 248, lateral view; 249, antenna.

FIGURES 250–255. Feron kingi, comb. nov., sexual female. 250, mesosoma, lateral view. 251, mesoscutum, dorsal view. 252, mesoscutellum, dorsal view. 253, metascutellum and propodeum, posterodorsal view. 254, fore wing, part. 255, metasoma, lateral view.