Feron Kinsey, 1937

Figs 1–513

Etymology. Kinsey (1937) did not indicate the etymology of this genus name in the original description. The name Feron could come from either the Greek φέρω (phérō) or Latin fero, both of which are verbs meaning “to carry” in which case its gender would be neuter; it could alternatively come from Φηρῶν which is a mythological name of a centaur, in which case it would be masculine. The suffixes in the epithets of the original species described under the name Feron, correspond to a neuter word, thus it must refer to a verb and not to a noun. Here, all species names’ terminals have been changed to neuter when applicable.

Type species: Feron verutum Kinsey, 1937 (according to the original description).

Kinsey (1937) described this genus with six new Mexican species, Feron tibiale, F. tostum, F. uterinum, F. validum, F. verutum and F. vitreum, which induce crystalline galls on leaves of white oaks in the subgenus Quercus (section Quercus, subsection Leucomexicana; Denk et al. 2017, Manos & Hipp 2021). Kinsey (1937: 70) commented that Feron should include numerous species that had previously been assigned to Andricus and other Western Palearctic genera; in addition, two other genera, Dros and Druon, were described in the same paper and Kinsey mentioned that the limits of all three genera would be determined in a monographic revision of the entire group. However, this revision was never published. Moreover, Kinsey constantly compared his new Feron species with Andricus tecturnarum but never transferred that species to Feron; our phylogenetic study shows that A. tecturnarum falls within the genus Feron. According to Weld (1951; 1952a), morphologically Feron closely resembles Andricus, and Weld (1951: 632) synonymized both genera. However, the original description of Feron does not mention several important characters that differentiate it from Andricus so for this reason, Feron is re-described below.

Diagnosis. Feron has the lower face with two smooth or almost smooth areas between the clypeus and eye, the malar sulcus absent; gena broadened or not behind the eye in frontal view, toruli usually located in the upper half of the head, the antenna always with 11–12 flagellomeres, the notaulus is usually complete, the mesopleuron is smooth or with a few vertical carinae on the most posterodorsal part of the speculum or with weak longitudinal carinae anteriorly; the transscutal articulation is present; lateral propodeal carinae bent outwards, sometimes strongly; the prominent part of the ventral spine of the hypopygium relatively long, usually around 4.0–6.0× as long as broad. According to these characters, Feron is similar to some sexual Andricus species. Andricus do not exhibit smooth areas on the lower face, toruli are always located at mid-height of the head, and the prominent part of the ventral spine of the hypopygium is usually shorter (around 2.0–3.5× as long as broad). Exceptionally, Andricus with longer ventral spines of the hypopygium have lateral propodeal carinae subparallel or bent slightly outwards and the mesoscutum is completely smooth or partially delicately alutaceous; even in some sexual female Andricus the antenna has 13 flagellomeres. Biologically, an important character to differentiate Feron from Andricus is the galls: one group of Feron species induces pubescent, crystalline or conical galls with rigid extensions on leaves, while the galls of Andricus never form gregarious pubescent masses on leaves; also, other lineages of Feron species induce disc-shaped, often star-like, spangle galls on leaves, a morphology that never occurs in Andricus. A few sexual-generation Andricus species, for example, the Western Palearctic A. quercusramuli (Linnaeus, 1761), induce Feron -like pubescent galls but only on catkins; in addition, those sexual females differ from Feron in the absence of smooth areas on the lower face, the mesopleuron is uniformly and very delicately transversally striate and the prominent part of the ventral spine of the hypopygium is usually short. The galls of some Feron species are similar to those induced by Druon species, but in Druon galls are covered in softer, more wool-like pubescent compared to the more crystalline hairs on Feron galls. The adults of Druon differ from Feron by the mesopleuron striate, and the lower face is entirely sculptured without smooth areas. Finally, some species of Feron can resemble Dros, but in Dros the frons is completely smooth (always sculptured in Feron) and the galls are significantly different.

Re-description. Females. Body yellowish to light brown, chestnut brown or dark brown, legs always lighter; antenna light brown to dark brown, with scape, pedicel and flagellomeres from F1 until F4 lighter than subsequent flagellomeres.

Head trapezoid, rounded, slightly ovate or semiquadrangular, with sparse white setae, denser on lower face, 1.1–1.2× as broad as high, slightly broader than mesosoma in small specimens in frontal view; 1.6–2.3× as broad as long in dorsal view. Gena alutaceous-reticulate, not broadened or very slightly broadened behind eye in frontal view, narrower than the transverse diameter of the eye in lateral view. Malar space alutaceous-reticulate, with striae radiating from clypeus sometimes reaching the eye, malar sulcus absent; eye 1.7–6.1× as high as the length of malar space. Inner margins of eyes parallel, converging, or slightly divergent. POL 1.3–3.0× as long as OOL (longer in males), OOL 1.0–3.3× as long as the diameter of lateral ocellus and slightly shorter to 2.0× as long as LOL, all ocelli ovate, of the same size. Transfacial distance longer or slightly shorter than the height of eye (shorter in some sexual forms); toruli usually located in the upper half of head and then frons shorter than lower face, the diameter of antennal torulus 1.0-2.0× as long as the distance between them, distance between torulus and eye around 1.0–1.5× as long as diameter of torulus; lower face smooth or slightly alutaceous between clypeus and eye, glabrous or with sparse white setae; slightly elevated median area alutaceous to smooth, with a few setae. Clypeus trapezoid, rectangular or quadrangular, smooth, or delicately coriaceous or striate, glabrous or with a few long setae scattered all over; ventrally rounded, emarginate, with or without median incision; anterior tentorial pit small, rounded, indistinct, epistomal sulcus distinct, clypeo-pleurostomal line well impressed. Frons alutaceous-reticulate or imbricate; interocellar area and vertex alutaceous-reticulate, without striae or setae; area under central ocellus impressed, alutaceous; occiput, postocciput and postgena alutaceous-reticulate, with few setae; 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 longer than head+mesosoma, with 11–12 flagellomeres (13 in males); pedicel longer than broad; F1 1.5–2.6× as long as pedicel and from equal with to 1.4× as long as F2; F2 equal to or longer than F3; all subsequent flagellomeres shorter, sometimes thicker; placodeal sensilla from F2 to F 5 in females (in all flagellomeres in males); F 1 in males with a variable shape, sometimes 1.6× as long as F2.

Mesosoma as long as high, with a few white setae along propleura. Pronotum smooth, glabrous or more or less pubescent laterally, foveolate or not along propleuron, laterally smooth or with delicate parallel striae in anteroposterior part; propleuron smooth, glabrous or setose. Mesoscutum as long as broad or slightly longer than broad (greatest width measured across mesoscutum level with base of tegulae), uniformly delicately coriaceous or alutaceous to rugose-reticulate, sometimes with smooth areas and punctures (smooth in some sexual forms). Notaulus complete or not, deep or shallow, posteriorly converging and usually broader than anteriorly, bottom smooth, glabrous; in most posterior section the distance between notauli shorter than distance between notaulus and side of mesoscutum. Anterior parallel line indistinct or distinct, impressed, smooth, glabrous, reaching at most 1/2 length of mesoscutum; parapsidal line indistinct or marked with broad smooth, glabrous stripe; median mesoscutal line absent or in the form of a short triangle; parascutal carina broad, reaching notaulus. Mesoscutellum ovate or trapezoid, longer than broad, broader at posterior end; disc of mesoscutellum uniformly rugoso-coriaceous, reticulate or smooth in central part, rugose in most posterior section and along sides, with some longitudinal rugae, overhanging metanotum. Mesoscutellar foveae present, divided by a rugose elevated central area or fused in the form of an anterior transverse impression, with smooth, glabrous bottom. Mesopleuron entirely smooth or with a few vertical carinae in most posterodorsal section, glabrous or with setae only along ventral edge; mesopleural triangle delicately coriaceous, glabrous or with white setae; dorsal and lateral axillar areas smooth, with or without dense setae; axillula with delicate parallel longitudinal striae; subaxillular bar smooth, glabrous, triangular, posteriorly as high as height of metanotal trough; metapleural sulcus reaching mesopleuron at half of its height, in some species slightly higher or lower, lower part of sulcus delimiting broad triangular coriaceous area; upper part of sulcus either distinct, separating smooth, glabrous area or indistinct. Metascutellum coriaceous or smooth, glabrous, as high as height of smooth, glabrous ventral impressed area; metanotal trough smooth, glabrous or with short setae; central propodeal area lyre-shaped, smooth, without or with only a few strong short longitudinal rugae; lateral propodeal carinae strong, broad and high, bent outwards in posterior 1/3; lateral propodeal area smooth, rarely alutaceous, glabrous or with long white setae, each seta with piliferous point at the base. Nucha with parallel longitudinal sulci dorsally and laterally or with a net of short irregular rugae. Tarsal claws with basal lobe.

Fore wing longer than body, except in brachypterous forms, hyaline, with dense cilia on margin, veins brown, radial cell open, 3.5–5.2× as long as broad; Rs and R1 not or nearly reaching wing margin; areolet triangular, enclosed by distinct veins. Rs+M distinct along entire length or along 1/3 to 2/3 of distance between areolet and basalis, its projection reaching basalis at half of its height.

Metasoma longer than head+mesosoma; 2nd metasomal tergum extending to 1/3–5/6 length of metasoma in dorsal view, with patch of dense white setae anterolaterally, with or without micropunctures; all subsequent terga and hypopygium with or without micropunctures; prominent part of ventral spine of hypopygium variable, 2.0–8.0× as long as broad in ventral view, with or without a few short white setae ventrally.

Body length 0.9–3.1 mm.

Gall. The asexual generation’s galls are always found on leaves but have a wide range of morphologies that can mainly be divided into two subgroups: One with pubescent masses, crystalline (resembling mineral crystalline structures, usually erected tubular structures with short rigid and brittle filaments radiating from it) or conical galls with rigid extensions on leaves; and a second with disc-shaped, often star-like, spangle galls (disc-shaped with a central round protrusion). The sexual generation’s gall

Biology. Adults galling on oaks from Section Quercus. The galls of the asexual generation mature during autumn, and adults overwinter in the galls and emerge between late January and May. The sexual generation occurs in late spring. The galls maduration and emergence of adults spans between April and May.

Distribution. Southern Canada, USA, and Mexico.

Descriptions of the sexual females and males are given in the species section below.

Thirty-four species are currently recognized within Feron:

F. albicomus (Weld, 1952), comb. nov., F. amphorus (Weld, 1926), comb. nov., F. apiarium (Weld, 1944), comb. nov., F. atrimentum (Kinsey, 1922), comb. nov., F. bakkeri (Lyon, 1984), comb. nov., F. caepula (Weld, 1926), comb. nov., F. californicum (Beutenmueller, 1911), comb. nov., F. clarkei (Bassett, 1890), comb. nov., F. comatum (Weld, 1952), comb. nov., F. crystallinum (Bassett, 1900), comb. nov., F. cylindratum (Kinsey, 1937), comb. nov., F. discale (Weld, 1926), comb. nov., F. discularis (Weld, 1926), comb. nov., F. dumosae (Weld, 1957), comb. nov., F. gigas (Kinsey, 1922), comb. nov., F. izabellae Melika, Nicholls & Stone, sp. nov., F. kingi (Bassett, 1900), comb. nov., F. parmula (Bassett, 1900), comb. nov., F. pattersonae (Fullaway, 1911), comb. nov. (= Andricus pedicellatus (Kinsey, 1922), syn. nov.), F. roberti Melika, Nicholls & Stone, sp. nov., F. rucklei Melika, Nicholls & Stone, sp. nov., F. scutellum (Weld, 1930), comb. nov., F. serranoae Pujade-Villar & Cuesta-Porta, sp. nov., F. splendens (Weld, 1919), comb. nov., F. stellare (Weld, 1926), comb. nov., F. stellulum (Burnett, 1974), comb. nov., F. sulfureum (Weld, 1926), comb. nov., F. syndicorum Pujade-Villar & Cuesta-Porta, sp. nov., F. tecturnarum (Kinsey, 1920), comb. nov., F. tetyanae Melika, sp. nov., F. tibiale Kinsey, 1937, comb. rev. (= F. tostum Kinsey, 1937, syn. nov., Feron uterinum Kinsey, 1937, syn. nov.), F. tubifaciens (Weld, 1926), comb. nov., F. verutum Kinsey, 1937, comb. rev. and F. vitreum Kinsey, 1937, comb. rev. (= F. validum Kinsey, 1937, syn. nov.).

Key to Feron species

1. Males (Figs 62–67, 110–115, 146–147, 245–249, 288–292).................................................... 2

- Females (Figs 1–11, 51–61, 68, 116–120).................................................................. 9

2. Ocelli strongly elevated above frons in frontal view (Fig. 62), OOL shorter than diameter of lateral ocellus and inner margins of eyes subparallel (Figs 62–63); mesoscutum uniformly reticulate between notauli in anterior half and laterad to notauli, smooth between notauli in posterior half....................................................... atrimentum comb. nov.

- Ocelli moderately or not elevated above frons; if not then inner margins of eyes strongly converging laterally and/or OOL longer (Figs 110–111, 146, 245, 247, 288–289); mesoscutum smooth or partially alutaceous only anteriorly.............. 3

3. Notaulus narrow, distinct only posteriorly, fragmented or absent anteriorly where mesoscutum delicately coriaceous.............................................................................................. gigas comb. nov.

- Notaulus distinct, complete, reaching pronotum............................................................. 4

4. Inner margins of eyes slightly diverging ventrally (Fig. 146); mesoscutellar foveae absent, present in the form of a transverse smooth anterior impression, continuing into smooth mesoscutellar disc (Fig. 146); body yellowish to amber (Figs 146–147)................................................................................. crystallinum comb. nov.

- Inner margins of eyes parallel or slightly converging ventrally (Figs 110, 245, 288); mesoscutellar foveae present, delimited posteriorly and delimited or not medially; mesoscutellar disc sometimes smooth; body brown to black (Figs 110–115, 245–248, 288–292)............................................................................................ 5

5. Mesoscutellum with irregular rugae at least in lateral and posterior part; sometimes dorsocentral part smooth, shining, without piliferous points...................................................................................... 6

- mesoscutellum uniformly alutaceous, with numerous setae on piliferous points; body brown or chestnut (Fig. 115); body black (Fig. 116)........................................................................................... 7

6. F1 1.6× as long as F2; transfacial distance as long as or slightly shorter than height of eye............ dumosae comb. nov.

- F1 1.3× as long as F2; transfacial distance longer than height of eye............................... clarkei comb. nov.

7. F1 equal in length to scape+pedicel, 1.45× as long as F2, slightly broadened and curved, flagellomeres lighter than scape and pedicel.............................................................................. comatum comb. nov.

- F1 longer than scape+pedicel, 1.3× as long as F2, straight, not broadened and curved, flagellomeres, scape and pedicel uniformly coloured............................................................................................ 8

8. Head ovate in frontal view; internal margin of eyes parallel; space between central elevated area of lower space to lateral margin of eye sculptured; OOL shorter than diameter of lateral ocellus..................................... kingi comb nov.

- Head triangular in frontal view (Fig. 288); inner margin of eyes converging ventrally; space between central elevated area of lower face to lateral margin of eye smooth (Fig. 288); OOL as long as diameter of lateral ocellus (Fig. 289)................................................................................................ pattersonae comb. nov.

9. Fore wing rudimentary, as long as head+mesosoma but not reaching the mid-length of metasoma, with indistinct veins (Fig. 94)........................................................................ californicum comb. nov. (asex.)

- Fore wing longer than length of body with distinct veins (Figs 184–185)........................................ 10

10. Mesoscutum smooth or partially finely alutaceous anteriorly; always glabrous (Figs 20–21, 32–33, 104–105, 118, 120, 143– 144, 156–157, 169–170, 196–197, 251–252, 295–296)....................................................... 11

- Mesoscutum alutaceous to coriaceous, rugose-reticulate, reticulate, sometimes with small smooth areas and piliferous points; glabrous or pubescent (Figs 8–9, 44–45, 70, 131, 210–211, 305, 436)........................................... 21

11. Mesoscutellum uniformly reticulate (Fig. 157)...................................... cylindratum comb. nov. (asex)

- Mesoscutellum with different sculpture (Figs 21, 105, 118, 120, 170, 277)....................................... 12

12. Frons bulging in frontal view (Fig. 162); inner margins of eyes strongly converging ventrally (Fig. 162); mesoscutellum in central part smooth, glabrous, posteriorly and laterally rugose (Figs 170–171); body yellowish to reddish brown (Figs 162– 172)............................................................................ discale comb. nov. (asex)

- Frons flat, not or only slightly bulging in frontal view (Fig. 121); inner margins of eyes parallel or only slightly converging ventrally; mesoscutellum rugose and/or with piliferous points (Figs 116, 118, 120); darker specimens (Figs 13–23, 25–35, 97–106, 116–121, 241–243, 250–253, 254, 293–299)........................................................ 13

13. mesoscutellum uniformly alutaceous with numerous setae on piliferous points (Figs 33, 120, 252, 296); body black (Figs 25–35, 116–121, 241–243, 250–253, 254, 293–299)......................................................... 14

- Mesoscutellum with irregular rugae at least in lateral and posterior parts, sometimes central part smooth, shining, without piliferous points (Figs 21, 105, 144, 197); body light brown to chestnut brown, rarely darker (Figs 13–23, 97–106, 136–145, 189–198)........................................................................................... 18

14. Notaulus narrow, distinct only posteriorly, discontinuous or absent and finely coriaceous in anterior part; mesopleuron smooth, with transverse striae in central part...................................................... gigas comb. nov. (sex)

- Notaulus distinct, complete, reaching pronotum (Figs 32, 116, 118, 251, 295); mesopleuron entirely smooth, without transverse striae in central part (Figs 31, 117–119, 250, 294)........................................................... 15

15. Toruli at half the height of head (Fig. 121); scape, pedicel, F1–F2 yellowish (Fig. 116); legs including hind coxa yellowish (Fig. 117); prominent part of ventral spine of hypopygium without setae ventrally.................. comatum comb. nov. (sex)

- Toruli in the upper half of head (Figs 25, 241, 284; scape, pedicel, F1–F2 dark brown, sometimes light brown but never yellowish (Figs 29, 244, 287); legs reddish brown, at least hind coxa darker; prominent part of ventral spine of hypopygium with setae ventrally................................................................................... 16

16. Eyes parallel; eye 1.7× as high as length of malar space (Fig. 25); antennomeres with long and erect setae (Fig. 29); OOL only slightly longer than diameter of lateral ocellus (Fig. 26); notaulus complete, deep (Fig. 32); prominent part of ventral spine of hypopygium with long dense white setae ventrally (Fig. 35).............................. apiarium comb. nov. (asex)

- Eyes converging ventrally; eye at least 4.0× as high as length of malar space (Figs 241, 284); antennomeres with short setae (Figs 244, 287); OOL 1.5× as long as diameter of lateral ocellus (Figs 242, 286); notaulus complete, sometimes weakly impressed anteriorly (Figs 251, 295); prominent part of ventral spine of hypopygium with few short setae ventrally (Figs 255, 299)............................................................................................... 17

17. Antenna with 11 flagellomeres; F1 subequal to F2 and at most 2.0× as long as pedicel (Fig. 244); head distinctly narrower than mesosoma and genae broadened in frontal view (Fig. 241); mesoscutellar foveae separated by narrow elevated coriaceous central carina; areolet inconspicuous (Figs 251–252)........................................ kingi comb. nov. (sex)

- Antenna with 12 flagellomeres; F1 1.2× as long as F2 and more than 2.0× as long as pedicel (Fig. 287); head only slightly narrower than mesosoma and genae not broadened (Fig. 284); mesoscutellar foveae fused in the form of a narrow, semi-lunar depression (Figs 296–297); areolet triangular (Fig. 298)................................ pattersonae comb. nov. (sex)

18. Transfacial distance as long as or slightly shorter than height of eye; eye more than 3.6× as high as length of malar space (Figs 136, 189); pronotum laterally with short carinae only along posterior margin (Figs 142, 195)........................ 19

- Transfacial distance longer than height of eye; eye less than 3.0× as high as length of malar space (Figs 13, 97); pronotum with carinae going across entire lateral surface (Figs 19, 103)..................................................... 20

19. Eyes converging ventrally (Fig. 189); F1 longer than F2 (Fig. 193); mesopleuron smooth, shining, with delicate transverse subparallel striae in central part (Fig. 195); prominent part of ventral spine of hypopygium slightly longer than broad, without setae (Fig. 200); brown coloured (Figs 189–198, 200).................................... dumosae comb. nov. (sex)

- Eyes parallel or only slightly converging posteriorly (Fig. 136); F1= F2; mesopleuron completely smooth, without striae (Fig. 139); ventral spine of hypopygium 3.7× as long as broad, with short white setae ventrally (Fig. 145; dark-coloured body(Figs 136–138, 142–145)............................................................. crystallinum comb. nov. (sex)

20. Gena not broadened behind eye in frontal view (Fig. 13); mesoscutellar foveae in the form of an anterior transverse impression, posteriorly not defined, continuing into disc of mesoscutellum (Fig. 21); mesopleuron completely smooth; metapleural sulcus reaching mesopleuron at half of its height (Fig. 19); 2nd metasomal tergum with setae anteriorly (Fig. 23); body light brown (Figs 13–23)................................................................... amphorus comb. nov. (asex)

- Gena slightly broadened behind eye in frontal view (Fig. 97); mesoscutellar foveae well-delimited by a carina posteriorly, separated medially (Fig. 105); mesopleuron with delicate indistinct transverse subparallel striae in anterodorsal part at mid height; metapleural sulcus reaching mesopleuron above half of its height (Fig. 103); 2nd metasomal tergum without setae (Fig. 108); body dark brown to chestnut brown (Figs 97–106, 108)............................... clarkei comb. nov. (sex)

21. Pronotum laterally smooth to coriaceous, without carinae (Figs 7, 43, 130, 222, 275, 378)........................... 22

- Pronotum with longitudinal carinae laterally (Figs 57, 68, 78, 209, 304, 360, 401, 470)............................. 27

22. Frons bulging in frontal view (Figs 1, 216); ocelli not elevated above head; transfacial distance 1.4× longer than height of eye (Figs 1, 216); eyes slightly converging ventrally, metapleural sulcus reaching mesopleuron on upper 2/3 of its height (Figs 7, 222)............................................................................................... 23

- Frons not bulging in frontal view, ocelli sometimes elevated above head; transfacial distance subequal or 1.2× longer than height of eye (Figs 37, 124, 269, 372); metapleural sulcus reaching mesopleuron at half of its height (Figs 43, 130, 275, 378). .................................................................................................. 24

23. Median mesoscutal line in the form of a short smooth, shining triangle (Fig. 8); mesoscutellar foveae divided by a thin carina (Fig. 9); ventral spine of hypopygium 11.0× longer than broad in ventral view (Fig. 11); head and mesosoma uniformly reddish brown (Figs 1–10).............................................................. albicomus comb. nov. (asex)

- Median mesoscutal line absent (Figs 223–224); mesoscutellar foveae divided by an elevated coriaceous triangle (Fig. 224); ventral spine of hypopygium 8.0× as long as broad in ventral view (Fig. 226); head and mesosoma amber (Figs 216–225).................................................................................... izabellae sp. nov. (asex)

24. Pronotum smooth, with sparse setae and piliferous points along dorsal edge (Fig. 43); mesoscutum longer than broad, scarcely pubescent, without black stripes, alutaceous-reticulate between notauli in anterior 2/3 length and lateral to notauli, smooth between notauli in posterior 1/3; anterior parallel line indistinct, parapsidal line absent (Figs 44–45)................................................................................................. atrimentum comb. nov. (asex)

- Pronotum coriaceous, if smooth then with dense setae and piliferous points (Figs 130, 275, 378); mesoscutum only slightly longer than broad (subequal), pubescent (Figs 131, 276, 379); mesoscutum dark brown in between notauli in anterior 1/3 length of mesoscutum and along parapsidal line, with other surface sculpture; anterior parallel line and parapsidal line present (Figs 131, 276, 379)....................................................................................... 25

25. Head ovate in frontal view (Fig. 269); pronotum smooth with dense setae and piliferous points (Fig. 275); mesoscutellar foveae fused (Fig. 277); 2nd metasomal tergum extending to 1/3 the length of metasoma in dorsal view (Fig. 280)........................................................................................... pattersonae comb. nov. (asex)

- Head trapezoid in frontal view (Figs 124, 372); pronotum coriaceous, if smooth then with dense setae and without piliferous points (Figs 130, 378); mesoscutellar foveae divided by median elevated narrow triangular area or carina (Figs 132, 379); 2nd metasomal tergum smooth, extending at least to 5/6 length of metasoma in dorsal view (Figs 134, 382)................ 26

26. Pronotum coriaceous in posterolateral area, with piliferous points (Fig. 130); mesoscutum rugose-reticulate between notauli and laterad to notaulus in anterior half, alutaceous glabrous between notauli in posterior half (Fig. 131); eye 2.7× as high as length of malar space (Fig. 124); distal flagellomeres broader than basal ones, F1 shorter than scape+pedicel (Fig. 128); prominent part of ventral spine of hypopygium 6.5x as long as broad in ventral view (Fig. 134)................................................................................................... crystallinum comb. nov. (asex)

- Pronotum smooth, without piliferous points (Fig. 378); mesoscutum uniformly delicately coriaceous (Fig. 379); eye 3.7× as high as length of mala space (Fig. 372); all flagellomeres uniformly broad, F1 longer than scape+pedicel (Fig. 376); prominent part of ventral spine of hypopygium more than 5.5× as long a broad in ventral view (Fig. 382)...................................................................................................... sulfureum comb. nov. (asex)

27. Mesoscutum with distinct and deep piliferous points (Figs 235–236, 471)........................................ 28

- Mesoscutum without piliferous points or shallow almost inconspicuous points (Figs 58, 70, 79, 210, 305, 361, 390, 402–403, 420, 436)........................................................................................... 30

28. Frons bulging in frontal view, ocelli not elevated above head (Fig. 186); toruli at mid-height of head (Fig. 186); median mesoscutal line in the form of a short triangle; entire body black (Figs 184–186)............. discularis comb. nov. (asex)

- Frons not bulging in frontal view, ocelli not elevated above head; toruli in upper half of head (Figs 228, 464); median mesoscutal line absent (Figs 235, 471); head and mesosoma never black (Figs 228–238, 464–472, 474)......................... 29

29. Gena broadened behind eye in frontal view (Fig. 228); clypeus smooth; eye 2.9× as high as length of malar space (Fig. 228); metapleural sulcus reaching mesopleuron at half of its height (Fig. 234); fore wing longer than body; metasomal terga 6 and 7 with micropunctures; prominent part of ventral spine of hypopygium around 8.0× as long as broad in ventral view (Fig. 238); light specimens, rusty brown and rarely uniformly dark brown (Figs 228–238)................... kingi comb. nov. (asex)

- Head not broadened behind eye in frontal view; clypeus with short interrupted delicate transverse striae; eye 1.9× as high as length of malar space (Fig. 464); metapleural sulcus reaching mesopleuron at upper 1/3 of its height (Fig. 470); fore wing only slightly longer than length of body (Fig. 473); all metasomal terga smooth, without micropunctures; prominent part of ventral spine of hypopygium around 5.0× as long as broad (Fig. 474); dark specimens, head dark brown to black, mesosoma reddish brown with some darker tints (Figs 464–472, 474).................................... tubifaciens comb. nov. (asex)

30. Eyes strongly converging ventrally; transfacial distance shorter than height of eye; antennal toruli located at half or slightly above height of eyes; eye around 6.0× as high as length of malar space (Fig. 51); body black (Figs 51–61)............................................................................................. atrimentum comb. nov. (sex)

- Eyes parallel or very slightly converging ventrally; transfacial distance equal to or longer than height of eye; toruli located above mid-height of eyes; eye less than 3.0× as high as length of malar space (Figs 69, 72, 300, 329, 342, 476, 489, 502); body yellowish, reddish brown, brown to chestnut brown (Figs 68–70, 72–82, 203–212, 256–265, 324–327, 329–338, 354–363) 31

31. Head quadrangular or ovate in frontal view, with ocelli always elevated above frons (Figs 342, 476, 489, 502); lateral ocelli large, OOL subequal or at most 1.7× as long as diameter of ocellus (Figs 344, 477, 490, 503)........................ 32

- Head rounded or trapezoid to triangular in frontal view, ocelli usually not elevated above frons (Figs 69, 72, 203, 256, 300, 354, 384, 395, 429); lateral ocelli smaller, OOL at least 2.5× as long as diameter of ocellus, if shorter then head and mesosoma are not yellowish or light brown (Figs 73, 204, 257, 302, 356, 386, 397, 430)........................................ 34

32. F1 distinctly longer than scape+pedicel (Fig. 493); transfacial distance equal to or slightly shorter than height of eye (Fig. 489)............................................................................ vitreum comb. rev. (asex)

- F1 subequal to or slightly longer than scape+pedicel (Figs 346, 480, 506); transfacial distance longer than height of eye (Figs 342, 476, 502)....................................................................................... 33.. 33. Mesoscutum with some delicate transverse striae in anterior part between notauli; anterior parallel lines indistinct (Figs 349–350); mesoscutellar foveae transverse, ovate, separated by a broad carina, posteriorly delimited by a carina (Fig. 350)............................................................................... splendens comb. nov. (asex)

- Mesoscutum without striae, anterior parallel line in the form of a bare, smooth stripe, extending to 1/3 length of mesoscutum (Figs 483–484); foveae quadrangular, separated by a thin carina, posteriorly not delimited by a carina (Fig. 484).......................................................................................... verutum comb. rev. (asex)

34. Mesopleuron with transverse reticulate-carinate band at mid-height (Figs 209, 262, 370, 389, 435).................... 35

- Mesopleuron entirely smooth or slightly sculptured on anterior margin (Figs 68, 78, 304, 317, 327, 335, 360, 401, 419, 435). .................................................................................................. 39

35. Gena broadened behind eye in frontal view (Figs 384, 429); 2nd metasomal tergum without micropunctures posteriorly and subsequent terga with or without micropunctures (Figs 393, 439).............................................. 36

- Gena not broadened behind eye in frontal view (Figs 203, 256, 369); 2nd metasomal tergum with a band of micropunctures posteriorly and all subsequent terga with micropunctures (Figs 214, 267, 368).................................... 37

36. Mesoscutum coarsely reticulated; notaulus complete, notaular furrow smooth and deep along its length or alutaceous on anterior 1/4 (Fig. 436); mesoscutellar disk flat in lateral view, following curvature of mesoscutum (Fig. 437); subaxillular bar reaching half-length of mesoscutellum in lateral view (Fig. 435)................................ tibiale comb. rev. (asex), part

- Mesoscutum finely alutaceous; notaulus complete, but indistinct and shallow at least on anterior 3/4 of its length (Fig. 390); mesoscutellar disk strongly curved in lateral view; mesoscutum and mesoscutellum forming two independent lobes in lateral view (Fig. 391); subaxillular bar reaching 1/3 of height of mesoscutellum (Fig. 389)........... syndicorum sp. nov. (asex)

37. Malar space without striae (Fig. 256); notaulus complete, indistinct in anterior 1/4 of mesoscutum (Fig. 263); mesoscutellar foveae absent or slightly impressed (Fig. 264); veins of fore wing pale (Fig. 266); prominent part of ventral spine of hypopygium 7.0× as long as broad in ventral view (Fig. 267)........................................ parmula comb. nov. (asex)

- Malar space with a few delicate striae radiating from clypeus (Figs 203, 369); notaulus complete, distinct through all its length (Figs 210, 367); mesoscutellar foveae defined (Figs 211, 367); veins of fore wing light brown to brown (Figs 213, 367); prominent part of ventral spine around 4.5× as long as broad in ventral view (Figs 214, 368)......................... 38

38. Antenna with 11 flagellomeres (Figs 368–370); eye 2.7× as high as length of malar space (Fig. 369); veins of fore wing brown; areolet present (Fig. 367); central propodeal area with rugae.............................. stellulum comb. nov. (asex)

- Antenna with 12 flagellomeres (Fig. 207); eye 2.2× as high as length of malar space (Fig. 203); veins of fore wing lighter; areolet absent (Fig. 213); central propodeal area without rugae (Fig. 212)....................... gigas comb. nov. (asex)

39. Head trapezoid (Fig. 300); gena not broadened behind eye; transfacial distance equal to height of eye (Fig. 300); antenna with 11 flagellomeres; F1=F2; mesoscutellar disc reticulate (Fig. 303); metapleural sulcus reaching mesopleuron in lower 1/3 of its height (Fig. 304); 2nd metasomal tergum extending to 3/4 of metasoma length in dorsal view (Fig. 309)................................................................................................... roberti sp. nov. (asex)

- Head rounded or ovate (Figs 69, 72, 311, 324, 329, 354, 395, 409, 429); gena at least slightly broadened behind eyes; transfacial distance longer than height of eye (Figs 69, 72, 311, 324, 329, 354, 395, 409, 429); antenna with 12 flagellomeres, sometimes suture between F11 and F12 incomplete; F1 longer than F2; distal flagellomeres broader than basal ones (Figs 76, 315, 326, 333, 358, 399, 413, 418, 433); mesoscutellar disc smooth or with a different sculpture; metapleural sulcus reaching mesopleuron at least at 1/2 of its height (Figs 70, 80, 319, 325, 337, 362, 403, 420, 424, 437); 2nd metasomal tergum extending to 1/2–2/3 length of metasoma in dorsal view (Figs 68, 82, 321, 327, 340, 365, 406, 426–427, 439)............................ 40

40. Head transversely ovate in frontal view (Figs 329, 395, 409, 429); slightly elevated median area of lower face alutaceous to delicately coriaceous, matte (Figs 329, 395, 409, 429); lateral sides of pronotum longitudinally striated at least on posterior half, rest of pronotum alutaceous, matte (Figs 335, 401, 419, 435); mesoscutum coarsely reticulated, each individual cell formed by the reticula bulging (Figs 336, 402, 420, 436); mesopleuron finely striated to alutaceous on anterior margin (Figs 335, 401, 419, 435)............................................................................................... 41

- Head rounded (Figs 69, 72, 311, 324, 354); slightly elevated median area of lower face smooth, shining (Figs 69, 72, 311, 324, 354); lateral sides of pronotum with fine longitudinal striations on posterior margin, never reaching half the length of pronotum, rest of the pronotum shining, smooth with piliferous points on dorsal margin (Figs 68, 78, 317, 327, 360); mesoscutum finely reticulated, the cells formed by the reticula flat (Figs 70, 79, 318, 325, 361), mesopleuron completely smooth (Figs 68, 78, 317, 327, 360)........................................................................................... 44

41. Mesoscutellum 1.3× longer than broad and margined posteriorly by a strong circumscutellar carina (Fig. 337)............................................................................................. serranoae sp. nov. (asex)

- Mesoscutellum slightly longer than broad and circumscutellar carina absent (Figs 403, 420, 424, 437)................. 42

42. Frons bulging in frontal view (Fig. 429); POL subequal to OOL (Fig. 430); gena broadened behind eye in frontal view (Fig. 429); mesoscutum uniformly reticulate-coriaceous (Fig. 436); central mesoscutellar disk coriaceous (Fig. 437); radial cell around 4.8× as long as broad; third and subsequent metasomal terga with sparse micropunctures (Fig. 439); body yellowish to light brown (Figs 429–439)...................................................... tibiale comb. rev. (asex), part

- Frons not bulging in frontal view (Figs 395, 409, 414), POL at least 1.4× longer than OOL (Figs 397, 410, 415); gena not or slightly broadened behind eye in frontal view (Figs 395, 409, 414); mesoscutum alutaceous to reticulate (Figs 402, 420, 424); center of mesoscutellar disk smooth or finely rugose (Figs 403, 420, 424); radial cell around 3.7× as long as broad (Figs 405, 422); all metasomal terga without micropunctures (Figs 406, 426–427); body chestnut brown or rusty brown (Figs 395–406, 409–427, 429–439)................................................................................... 43

43. Mesoscutum sometimes with dark brown areas around anterior parallel lines and parapsidal lines (Figs 420, 424); frons coarsely coriaceous (Figs 409–410); mesoscutellar foveae divided by a fine carina (Fig. 420); chestnut brown, head always darker than mesosoma (Figs 409–427)............................................................ tetyanae sp. nov. (asex)

- Frons finely alutaceous (Fig. 395); mesoscutellar foveae divided by a triangular elevated coriaceous central carina (Fig. 403); rusty brown to light brown, head and mesosoma of the same colour and without dark areas (Figs 395–404, 406)...................................................................................... tecturnarum comb. nov. (asex)

44. Frons bulging in frontal view; eyes strongly converging ventrally; eye 3.0× as high as length of malar space (Fig. 72); central part of mesoscutellum smooth (Fig. 80); prominent part of ventral spine of hypopygium around 3.8× as long as broad in ventral view (Fig. 82); yellowish to light brown (Figs 72–82).................................... caepula comb. nov. (asex)

- Head rounded; eyes parallel or very slightly converging ventrally; height of eye less than 2.6× as high as length of malar space (Figs 69, 311, 324, 354); central part of mesoscutellum coriaceous to rugose (Figs 70, 319, 325, 362); prominent part of ventral spine of hypopygium at least 5.5× as long as broad in ventral view (Figs 68, 321, 327, 365); brown, sometimes with darker marks (Figs 68–70, 311–315, 317–321, 324–327, 354–363, 365)............................................... 45

45. Mesoscutellar foveae absent, only slightly impressed on anterior part, with the bottom finely rugose like the rest of the disk, without central carina (Fig. 70); metasoma longer than high in lateral view (Fig. 68)............ bakkeri, comb. nov. (asex)

- Mesoscutellar foveae conspicuous and bottom smooth, fused or divided by a central carina or a triangular elevated area (Figs 319, 325, 362); metasoma as high as long or higher in lateral view (Figs 321, 327, 365)............................. 46

46. Mesoscutellum disk faintly reticulate, with an elevated central area; mesoscutellar foveae not delimited posteriorly (Fig. 325)............................................................................. scutellum comb. nov. (asex)

- Mesoscutellar disk coarsely rugose, without elevated area; mesoscutellar fovea completely delimited by rugose sculpture or by a carina (Figs 319, 362)............................................................................... 47

47. Notaulus incomplete (Fig. 361); mesoscutellar foveae delimited all around by strong black carina; mesoscutellum rugose-coriaceous (Fig. 362); all metasomal terga without micropunctures (Fig. 365); first flagellomeres lighter than subsequent (Fig. 358)............................................................................. stellare comb nov. (asex)

- Notaulus complete (Fig. 318); mesoscutellar foveae separated by triangular elevated coriaceous central carina; mesoscutellum rugose (Fig. 319); third and subsequent terga with sparse micropunctures (Fig. 321); all flagellomeres dark (Fig. 315)........................................................................................ rucklei sp. nov. (asex)