Feron pattersonae ( Fullaway, 1911 ) 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 : 94-101

Feron pattersonae ( Fullaway, 1911) , comb. nov.

Figs 269–299

Andricus pattersonae Fullaway, 1911: 352 , female (asex), gall; Burks, 1978: 1087.

Liodora pattersonae (Fullaway) : Evans 1972: 1813.

Sexual generation:

Andricus pedicellatus Kinsey, 1922: 284–285 , male, female, gall; Melika & Abrahamson, 2002: 162, syn. nov.

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

Types examined: Andricus pattersonae : SYNTYPES: No. 5822 is deposited at the California Academy of Sciences Collection , was not examined by the authors. A cotype asexual female “ L.S.Jr.U. Lot 508 Sub52”, handwriting label “Stanford Univ., Calif.”, handwriting label “ Andricus pattersonae Full. ” “Beut. Coll. rec’d 1935”, red label “ USNM cotype N52974”, a Weld handwriting label “ Andricus pattersonae Full. ” was examined by GM. Andricus pedicellatus : HOLOTYPE: Sexual female “Three Rivers Cal., Gall 3.23.20.”, “ Q. Douglasii, Kinsey coll.”, red label “ Neuroterus pedicellatus , Holo- COTYPE”, deposited in, NYC, examined by GM. Four sexual female paratypes, two males and many galls in the general collection of AMNH were also examined by GM .

Additional material. Andricus pattersonae , 12 asexual females: 3 females as “ USA, California, 30km SW of Williams, ex Quercus douglasii, CA 16, coll. 2007.11.02. leg. J. A. Nicholls ” and 9 females as “ USA, California, Dye Creek Preserve, ex Quercus douglasii, CA1177 , coll. 2001.09.27 leg. J. A. Nicholls ”. Dros pedicellatum : 3 sexual females labelled as “ USA, California, Quail Ridge Reserve , ex Quercus douglasii, CA1038 , coll. 2008.04.04. leg. J.A. Nicholls ” .

Diagnosis. Asexual females belong to the group of Feron species in which the pronotum is without carinae, with dense setae and piliferous points; the mesoscutum is dark brown between notauli in anterior 1/3 of mesoscutum length; as in F. crystallinum (asex) and F. sulfureum (asex). Feron pattersonae differs from these species in the ovate head in frontal view; the pronotum is smooth, with dense setae and piliferous points; mesoscutellar foveae are fused; second metasomal tergum extending to 1/3 length of the metasoma in dorsal view. Sexual females belong to the group of Feron species with 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 are never strongly converging ventrally; the mesoscutum is smooth or partially alutaceous anteriorly and glabrous, the notaulus complete; the mesopleuron is entirely smooth without transverse striae in its central part; the mesoscutellum is 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. kingi (sex). Differs from F. apiarium in the eyes converging ventrally, the eye is at least 4.0x 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 closely resembles F. kingi but differs in the number of flagellomeres (12 in F. pattersonae , 11 in F. kingi ) and in other characters mentioned at couplet 17 in the key. Males are characterised by a black body, ocelli are only moderately or not elevated above the frons, the notaulus reaching the pronotum and the mesoscutellum is uniformly alutaceous with numerous setae on piliferous points; as in F. comatum and F. kingi . In F. pattersonae F1 longer than scape+pedicel, straight, not broadened and curved; flagellomeres, scape and pedicel are uniformly coloured (F1 is equal in length to scape+pedicel, slightly broadened and curved, flagellomeres lighter than scape and pedicel in F. comatum ); the head is triangular in frontal view, inner margins of eyes converging ventrally, space between central elevated area of lower face to lateral margin of eye is smooth, OOL as long as diameter of the lateral ocellus (in F. kingi the head is ovate, inner margins of eyes are parallel, central elevated area of the lower face is sculptured, OOL shorter than the diameter of the lateral ocellus).

Re-description. Asexual female ( Figs 269–280 ). Entire body, antenna, legs are uniformly reddish brown, with dark brown to black metascutellum and central propodeal area. In some specimens internotaular area in anterior half, posteroventral part of mesopleuron black; last flagellomeres darker than rest of antenna.

Head with a few long setae, denser on lower face, alutaceous below toruli, transverse, 1.2× as broad as high and slightly broader than mesosoma in frontal view; 1.8× as broad as long from dorsal view. Gena alutaceous, broadened behind eye in frontal view, narrower than transverse diameter of eye in lateral view. Malar space alutaceous, glabrous, with numerous delicate parallel striae radiating from clypeus and reaching eye; malar sulcus absent; eye 2.3× as high as length of malar space. Eyes converging slightly ventrally. POL 2.1× as long as OOL, OOL 1.9× as long as diameter of lateral ocellus and slightly shorter than LOL, all ocelli slightly ovate, of same size. Transfacial distance 1.25× as long as height of eye; diameter of antennal torulus 1.7× as long as distance between them, distance between torulus and eye as long as diameter of torulus; lower face smooth, with dense white setae, without striae; slightly elevated median area alutaceous with a few setae. Clypeus impressed, trapezoid, 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 reticulate, without striae and setae, interocellar area alutaceous. Vertex, occiput reticulate, without setae; postgena alutaceous with dense setae; postocciput alutaceous with delicate, longitudinal parallel striae and dense white setae; posterior tentorial pit large, elongated, area below impressed; occipital foramen higher than height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which strongly diverge toward occipital foramen, postgenal bridge anteriorly broader than occipital foramen. Antenna longer than head+mesosoma, with 13 flagellomeres, pedicel 1.6x as long as broad; F1 2.6× as long as pedicel and 1.3× as long as F2; F2=F3, F4=F5, F5 1.3x as long as F6, all subsequent flagellomeres nearly equal in length; F13 slightly longer than F12; placodeal sensilla on F5–F11, absent on F1–F4.

Mesosoma slightly longer than high, with dense white setae, except bare central propodeal area. Pronotum smooth without striae laterally, with dense setae and piliferous points; propleuron alutaceous, glabrous. Mesoscutum with dense white setae, slightly longer than broad (greatest width measured across mesoscutum level with base of tegulae), alutaceous alongside notaulus, in between notauli rugose-reticulate in anterior 1/3 and smooth shining in posterior 2/3. Notaulus deep, with smooth bottom, incomplete, indistinct in anterior 1/5 of mesoscutum length; anterior parallel lines in the form of a narrow elevated smooth, glabrous stripes, extending to 1/3 of mesoscutum length; parapsidal line distinct, marked with broad slightly impressed line; median mesoscutal line absent; parascutal carina broad, reaching notaulus. Mesoscutellum ovate, longer than broad; overhanging metanotum, with dense long setae; disk of mesoscutellum delicately coriaceous, laterally dull rugose. Mesoscutellar foveae in the form of a semi-lunar depression, with smooth, glabrous bottom, without median carina separating them. Mesopleuron entirely smooth, with dense setae anteroventrally and ventrally; mesopleural triangle smooth, with dense setae; dorsal and lateral axillar areas smooth, with dense white setae; subaxillular bar smooth, glabrous, triangular, gradually higher towards the most posterior end, posteriorly as high as height of metanotal trough; metapleural sulcus reaching mesopleuron at half of its height, upper part of sulcus indistinct, lower part of sulcus separating smooth, glabrous area, with a few setae in lower part. Metascutellum rugose, 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 in posterior 1/3; lateral propodeal carinae strong, bent outwards in posterior 1/3; lateral propodeal area smooth, with long dense white setae. Nucha with strong sulci dorsally and laterally. Tarsal claws with basal lobe.

Fore wing longer than body, hyaline, with distinct dense cilia on margin, veins brown, distinct, radial cell open, 4.8× as long as broad; R1 and Rs nearly reaching wing margin; areolet small, triangular, closed, indistinct. Rs+M narrow, distinct, extending to 2/3 of length from areolet to basalis; 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 1/3 length of metasoma in dorsal view, with numerous white setae anteroventrally and without micropunctures; subsequent terga smooth, glabrous, with micropunctures. Hypopygium with micropunctures, prominent part of ventral spine of hypopygium 4.7× as long as broad in ventral view, with two rows of short setae ventrally which do not extend beyond apex of spine.

Body length 2.0– 2.2 mm (n = 5).

Sexual female ( Figs 284–287 , 293–299 ). Head, antenna, mesosoma, metasoma dark brown to black, legs yellowish with darker mid and hind coxae. Fore wings hyaline, veins brown.

Head alutaceous, with a few setae, denser on lower face, rounded, very slightly higher than broad and broader than mesosoma in frontal view; 1.9× as broad as long from dorsal view. Gena alutaceous, not broadened behind eye in frontal view, narrower than transverse diameter of eye in lateral view. Malar space alutaceous, glabrous, without striae; eye 5.0× as high as length of malar space. Eyes strongly converging ventrally. POL 2.4× as long as OOL, OOL 1.7× as long as diameter of lateral ocellus and slightly shorter than LOL, all ocelli slightly ovate, of same size. Transfacial distance shorter than height of eye; diameter of antennal torulus 1.7× as long as distance between them, distance between torulus and eye slightly shorter than diameter of torulus; lower face and slightly elevated median area alutaceous, with dense white setae, without striae. Clypeus rounded, smooth, with a few setae; ventrally rounded, emarginate, without median incision; anterior tentorial pit large, rounded, distinct, epistomal sulcus distinct, clypeo-pleurostomal line well impressed. Frons uniformly delicately coriaceous to reticulate, without striae and setae, interocellar area alutaceous. Vertex, occiput and postocciput alutaceous; postgena smooth, with setae and piliferous points; posterior tentorial pit large, elongated, area below impressed; occipital foramen as high as 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 12 flagellomeres, all flagellomeres with dense long setae, not broadened distally; pedicel slightly longer than broad; F1 3.0× as long as pedicel and 1.2× as long as F2; F2 1.3x as long as F3; F3=F4, F5=F6, subsequent flagellomeres nearly equal in length; F12 1.2× as long as F11; placodeal sensilla on F3–F12.

Mesosoma as long as high, with sparse white setae along notauli, laterally on pronotum, mesoscutum and mesoscutellum; denser setae on dorsal and lateral axillar areas and lateral propodeal area. Pronotum alutaceous to smooth, with sparse setae, with very delicate parallel striae in lateroposterior part; propleuron alutaceous, glabrous. Mesoscutum alutaceous, glabrous in anterior half, smooth and glabrous in posterior half, with sparse white setae, slightly longer than broad (greatest width measured across mesoscutum level with base of tegulae). Notaulus narrow, complete, only slightly impressed anteriorly, bottom smooth, slightly converging posteriorly; anterior parallel line invisible; parapsidal line distinct, marked with broad slightly impressed line; median mesoscutal line absent or sometimes in the form of a short triangle; parascutal carina broad, reaching notaulus. Mesoscutellum slightly longer than broad; disk of mesoscutellum smooth, glabrous, sparsely pubescent laterally, with rugae laterally and posteriorly, overhanging metanotum, with sparse long setae. Mesoscutellar foveae in the form of a narrow semi-lunar depression, with smooth, glabrous bottom. Mesopleuron entirely smooth, glabrous, with a few setae in anteroventral part; mesopleural triangle smooth, with some irregular striae and a few setae; dorsal and lateral axillar areas smooth, with dense white setae; subaxillular bar smooth, glabrous, triangulate, posteriorly as high as height of metanotal trough; metapleural sulcus reaching mesopleuron in the lower half of its height, upper part of sulcus indistinct, lower part of sulcus delimiting smooth, glabrous area, with setae and which is as broad as high. Metascutellum coriaceous, as high as height of smooth, glabrous ventral impressed area; metanotal trough smooth, with dense white setae; central propodeal area lyre-shaped, smooth, glabrous, with irregular interrupted rugae basally; lateral propodeal carinae strong, bent outwards in posterior 1/3; lateral propodeal area smooth, with long dense white setae. Nucha with few sulci dorsally and laterally. Tarsal claws with basal lobe.

Fore wing longer than body, hyaline, with distinct dense cilia on margin, veins brown, radial cell open, 5.2× as long as broad; R1 and Rs reaching wing margin; areolet triangular, closed, distinct. Rs+M narrow, distinct, reaching basalis at its mid height.

Metasoma as long as head+mesosoma, as long as high in lateral view; 2nd metasomal tergum extending to more than half-length of metasoma in dorsal view, with 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 2.8–2.9 mm (n = 2).

Male ( Figs 288–292 ). Similar to female, legs slightly darker; head triangular in frontal view; eye 4.5× as high as length of malar space; head posteriorly uniformly and entirely alutaceous; POL 2.7× as long as OOL, OOL as long as diameter of lateral ocellus and slightly shorter than LOL, ocelli ovate, larger than in female; interocellar area elevated in frontal view; antenna longer than body, pedicel as long as broad, with 13 flagellomeres, F1 straight, not swollen and curved, longer than scape+pedicel, F1 1.3× as long as F2, F2=F3=F4, all subsequent flagellomeres nearly equal in length, F13 slightly shorter than F12; placodeal sensilla on all flagellomeres. Body length 2.7 mm (n = 1).

Gall. Asexual gall ( Figs 281–282 ) Relatively large, thin, flat spangle gall on underside of leaves, detachable, monolocular. about 9 mm in diameter, with irregular way margin. Green when young, becoming yellow or tan when mature. Sexual gall ( Fig. 283 ) is a small, elongate capsule, 5–7 mm long, 1.0– 1.5 mm wide, attached by a slender pedicel to the edge of the leaf, unilocular. The gall is sharp-tipped, broadest near the tip, continuing into a slender, thread-like stem, 5–12 mm long, evidently a continuation of a leaf vein. The gall is thin-walled, entirely hollow.

Biology. Andricus pattersonae and Liodora dumosae were erroneously paired by Evans (1972; restated by Dailey & Menke 1980). Evans (1972) appears to have mis-identified the asexual gall he found as A. pattersonae ; his Figure 15 shows a gall much more similar to the new species F. rucklei described herein. Molecular data also refute Evans’ pairing, instead establishing the correct matching of A. pattersonae with A. pedicellatus , with the associated synonymisation herein. Four individuals (two asexual females [= A. pattersonae ], two sexual females [= A. pedicellatus ]) were sequenced for cytb and ITS2. Cytb sequences were on average 1.14% divergent (range 0.23– 1.90%; GenBank accessions OQ446204–OQ446206) and ITS2 sequences were on average 0.25% divergent (range 0–0.42%, including identical alleles in one asexual and one sexual individual; GenBank accessions OQ448243– OQ448246).

Asexual galls have been recorded from leaves on section Quercus , subsection Dumosae oaks: Q. douglasii , Q. dumosa , Q. garryana , and Q. lobata ( Burks 1979) , the gall matures by late autumn, adults emerge in late winter. Sexual galls occur on leaves of Q. douglasii ( Burks 1979) and mature in April; adults emerge soon after.

Distribution. USA: California ( Burks 1979).

Figures

FIGURES 269–273. Feron pattersonae, comb. nov., asexual female. 269–272, head: 269, frontal view, 270, dorsal view, 271, lateral view, 272, posterior view. 273, antenna.

FIGURES 274–278. Feron pattersonae, comb. nov., asexual female. 274, mesosoma and propleura, frontal view. 275, mesosoma, lateral view. 276, mesoscutum, dorsal view. 277, mesoscutellum, dorsal view. 278, metascutellum and propodeum, posterodorsal view.

FIGURES 279–283. Feron pattersonae, comb. nov.. 279–280, asexual female: 279, fore wing, part, 280, metasoma, lateral view. 281–283, galls: 281, mature asexual gall. 282, growing young asexual gall, 283, sexual galls.

FIGURES 284–292. Feron pattersonae, comb. nov., sexual generation. 284–287, sexual female: 284–286, head: 284, frontal view, 285, posterior view, 286, dorsal view; 287, antenna. 288–292, male: 288–290, head: 288, frontal view, 289, dorsal view, 290, posterior view; 291, antenna, 292, scape, pedicel, F1, F2.

FIGURES 293–297. Feron pattersonae, comb. nov., sexual female. 293, head, lateral view. 294, mesosoma, lateral view. 295, mesoscutum, dorsal view. 296, mesoscutellum, dorsal view. 297, metascutellum and propodeum, posterodorsal view.

FIGURES 298–299. Feron pattersonae, comb. nov., sexual female. 298, fore wing, part, 299, metasoma, lateral view.

FIGURES 13–18. Feron amphorus, comb. nov., asexual female. 13–16, head: 13, frontal view, 14, dorsal view, 15, lateral view, 16, posterior view. 17, antenna. 18, mesosoma and propleura, frontal view.