Atrusca Kinsey, 1930

Atrusca Kinsey, 1930 View in CoL

Figs 1–627 View FIGURES 1–6 View FIGURES 7–10 View FIGURES 11–14 View FIGURES 15–20 View FIGURES 21–24 View FIGURES 25–28 View FIGURES 29–32 View FIGURES 33–38 View FIGURES 39–42 View FIGURES 43–46 View FIGURES 47–52 View FIGURES 53–56 View FIGURES 57–60 View FIGURES 61–66 View FIGURES 67–70 View FIGURES 71–74 View FIGURES 75–80 View FIGURES 81–84 View FIGURES 85–88 View FIGURES 89–94 View FIGURES 95–98 View FIGURES 99–102 View FIGURES 103–108 View FIGURES 109–112 View FIGURES 113–116 View FIGURE 117 View FIGURES 118–122 View FIGURES 123–128 View FIGURES 129–132 View FIGURES 133–136 View FIGURES 137–142 View FIGURES 143–146 View FIGURES 147–150 View FIGURES 151–154 View FIGURES 155–160 View FIGURES 161–164 View FIGURES 165–168 View FIGURES 169–172 View FIGURES 173–178 View FIGURES 179–182 View FIGURES 183–185 View FIGURES 186–191 View FIGURES 192–195 View FIGURES 196–199 View FIGURES 200–204 View FIGURES 205–206 View FIGURES 207–212 View FIGURES 213–216 View FIGURES 217–220 View FIGURES 221–226 View FIGURES 227–230 View FIGURES 231–234 View FIGURES 235–240 View FIGURES 241–244 View FIGURES 245–248 View FIGURES 249–253 View FIGURES 254–257 View FIGURES 258–261 View FIGURES 262–267 View FIGURES 268–271 View FIGURES 272–275 View FIGURES 276–281 View FIGURES 282–285 View FIGURES 286–289 View FIGURES 290–295 View FIGURES 296–299 View FIGURES 300–301 View FIGURES 302–303 View FIGURES 304–309 View FIGURES 310–313 View FIGURES 314–317 View FIGURES 318–323 View FIGURES 324–327 View FIGURES 328–330 View FIGURES 331–336 View FIGURES 337–340 View FIGURES 341–343 View FIGURES 344–345 View FIGURES 346–350 View FIGURE 351 View FIGURES 352–357 View FIGURES 358–361 View FIGURES 362–365 View FIGURES 366–367 View FIGURES 368–373 View FIGURES 374–377 View FIGURES 378–381 View FIGURES 382–387 View FIGURES 388–391 View FIGURES 392–395 View FIGURES 396–401 View FIGURES 402–405 View FIGURES 406–409 View FIGURES 410–415 View FIGURES 416–419 View FIGURES 420–423 View FIGURES 424–429 View FIGURES 430–433 View FIGURES 434–437 View FIGURES 438–442 View FIGURES 443–446 View FIGURES 447–452 View FIGURES 453–456 View FIGURES 457–460 View FIGURES 461–466 View FIGURES 467–470 View FIGURES 471–474 View FIGURES 475–480 View FIGURES 481–484 View FIGURES 485–488 View FIGURES 489–494 View FIGURES 495–498 View FIGURES 499–502 View FIGURES 503–507 View FIGURES 508–511 View FIGURES 512–515 View FIGURES 516–521 View FIGURES 522–525 View FIGURES 526–529 View FIGURES 530–535 View FIGURES 536–539 View FIGURES 540–543 View FIGURES 544–549 View FIGURES 550–553 View FIGURES 554–557 View FIGURES 558–563 View FIGURES 564–567 View FIGURES 568–571 View FIGURES 572–577 View FIGURES 578–581 View FIGURES 582–585 View FIGURES 586–591 View FIGURES 592–595 View FIGURES 596–599 View FIGURES 600–605 View FIGURES 606–609 View FIGURES 610–613 View FIGURES 614–619 View FIGURES 620–623 View FIGURES 624–627

Cynips View in CoL subgenus Atrusca Kinsey, 1930: 276–278 , asexual female, gall.

Cynips View in CoL subgenus Atrusca Kinsey, 1936: 79 .

Atrusca Kinsey, 1930 View in CoL in Weld, 1951: 637.

Type species: Cynips dugesi variety simulatrix Kinsey 1930: 280 , female, gall. Original designation by Kinsey 1930: 277. Current valid name: Atrusca simulatrix ( Kinsey, 1930) .

All species are known from the asexual generation only and associated with white oaks only. Kinsey mentioned that all species induce nearly the same shaped, coloured galls and it is impossible to tell apart species based on galls only. Burks (1979) listed 15 species in the USA.

Diagnosis. Atrusca belongs to a group of Cynipini genera with a straight, complete transscutal articulation; the mesoscutum is never emarginate posterolaterally, not fused with the mesoscutellum. Many species are fully winged, and others are brachypterous. The malar sulcus is always absent, antennae with 12 flagellomeres; the mesoscutum alutaceous to alutaceous-reticulate or delicately coriaceous, with or without piliferous points; in the fore wing the second abscissa of Rs usually strongly curved, the radial cell short, opened, 1.4–2.6× as long as broad, with dark spots and/or dark stripes along veins; the hind tibia without a lobe apically; the ventral spine of the hypopygium with variable length, usually with subapical setae forming a tuft and reaching beyond the apex of the spine, tarsal claws with a basal lobe.

Three Nearctic genera, Antron, Atrusca , Cynips (= Besbicus ), have a transcutal articulation, toothed tarsal claws, and fore wings with fuscate stripes and/or spots. Atrusca differs from all other mentioned genera by the shape or the radial cell in the fore wing; in Atrusca the second abscissa of Rs is always curved usually strongly curved or angulated distally, then the radial cell short (around 1.6–2.4× as long as broad, rarely slightly longer), while in Antron, Cynips (= Besbicus ) the second abscissa of Rs is always straight or only slightly curved, with radial cell longer (around 2.5–3.6× as long as broad, rarely shorter). Also, in Atrusca the mesoscutellum is always elongated, with parallel or subparallel sides, posteriorly rounded or elongated, with shallow invagination posteromedially or gradually narrowed till central part, and the prominent part of the ventral spine of the hypopygium is long or extra-long, needle-like shaped, while in Antron the mesoscutellum as long as broad or slightly longer than broad, posteriorly always rounded and in Cynips (= Besbicus ) the ventral spine of the hypopygium has strong lateral lobes in asexual forms or it is triangular in ventral and lateral views, broadest at the base and uniformly tapering to a blunt point at the apex in the sexual generations. Atrusca also differs from Antron and Cynips (= Besbicus ) in the morphology of galls; no Antron or Cynips (= Besbicus ) species have galls with a fragile wall and with the larval chamber located centrally attached by radiating filaments to the outer wall.

Re-description. Asexual female ( Figs 1–627 View FIGURES 1–6 View FIGURES 7–10 View FIGURES 11–14 View FIGURES 15–20 View FIGURES 21–24 View FIGURES 25–28 View FIGURES 29–32 View FIGURES 33–38 View FIGURES 39–42 View FIGURES 43–46 View FIGURES 47–52 View FIGURES 53–56 View FIGURES 57–60 View FIGURES 61–66 View FIGURES 67–70 View FIGURES 71–74 View FIGURES 75–80 View FIGURES 81–84 View FIGURES 85–88 View FIGURES 89–94 View FIGURES 95–98 View FIGURES 99–102 View FIGURES 103–108 View FIGURES 109–112 View FIGURES 113–116 View FIGURE 117 View FIGURES 118–122 View FIGURES 123–128 View FIGURES 129–132 View FIGURES 133–136 View FIGURES 137–142 View FIGURES 143–146 View FIGURES 147–150 View FIGURES 151–154 View FIGURES 155–160 View FIGURES 161–164 View FIGURES 165–168 View FIGURES 169–172 View FIGURES 173–178 View FIGURES 179–182 View FIGURES 183–185 View FIGURES 186–191 View FIGURES 192–195 View FIGURES 196–199 View FIGURES 200–204 View FIGURES 205–206 View FIGURES 207–212 View FIGURES 213–216 View FIGURES 217–220 View FIGURES 221–226 View FIGURES 227–230 View FIGURES 231–234 View FIGURES 235–240 View FIGURES 241–244 View FIGURES 245–248 View FIGURES 249–253 View FIGURES 254–257 View FIGURES 258–261 View FIGURES 262–267 View FIGURES 268–271 View FIGURES 272–275 View FIGURES 276–281 View FIGURES 282–285 View FIGURES 286–289 View FIGURES 290–295 View FIGURES 296–299 View FIGURES 300–301 View FIGURES 302–303 View FIGURES 304–309 View FIGURES 310–313 View FIGURES 314–317 View FIGURES 318–323 View FIGURES 324–327 View FIGURES 328–330 View FIGURES 331–336 View FIGURES 337–340 View FIGURES 341–343 View FIGURES 344–345 View FIGURES 346–350 View FIGURE 351 View FIGURES 352–357 View FIGURES 358–361 View FIGURES 362–365 View FIGURES 366–367 View FIGURES 368–373 View FIGURES 374–377 View FIGURES 378–381 View FIGURES 382–387 View FIGURES 388–391 View FIGURES 392–395 View FIGURES 396–401 View FIGURES 402–405 View FIGURES 406–409 View FIGURES 410–415 View FIGURES 416–419 View FIGURES 420–423 View FIGURES 424–429 View FIGURES 430–433 View FIGURES 434–437 View FIGURES 438–442 View FIGURES 443–446 View FIGURES 447–452 View FIGURES 453–456 View FIGURES 457–460 View FIGURES 461–466 View FIGURES 467–470 View FIGURES 471–474 View FIGURES 475–480 View FIGURES 481–484 View FIGURES 485–488 View FIGURES 489–494 View FIGURES 495–498 View FIGURES 499–502 View FIGURES 503–507 View FIGURES 508–511 View FIGURES 512–515 View FIGURES 516–521 View FIGURES 522–525 View FIGURES 526–529 View FIGURES 530–535 View FIGURES 536–539 View FIGURES 540–543 View FIGURES 544–549 View FIGURES 550–553 View FIGURES 554–557 View FIGURES 558–563 View FIGURES 564–567 View FIGURES 568–571 View FIGURES 572–577 View FIGURES 578–581 View FIGURES 582–585 View FIGURES 586–591 View FIGURES 592–595 View FIGURES 596–599 View FIGURES 600–605 View FIGURES 606–609 View FIGURES 610–613 View FIGURES 614–619 View FIGURES 620–623 View FIGURES 624–627 ). Body uniformly brown, reddish brown, sometimes with black areas, in some species very extensive; black stripes usually along anterior parallel and parapsidal lines and/or between notauli in anterior 2/3 of mesoscutum length.

Head transversely quadrangular to ovate in frontal view, sometimes rounded or quadrangular, with sparse white setae, 1.1–1.4× as broad as high or as broad as high, narrower than mesosoma in frontal view. Gena alutaceous to coriaceous, slightly or not broadened behind eye in frontal view, narrower than transverse diameter of eye in lateral view. Malar space delicately coriaceous to alutaceous, with or without delicate striae, radiating from clypeus, malar sulcus absent. Inner margins of eyes parallel or slightly diverging ventrally. Lower face alutaceous to coriaceous, sometimes rugose in central part below toruli, with setae; slightly elevated median area usually rugose, with or without setae. Interocellar area elevated or not above head in frontal view; lateral ocelli prominent or not. Clypeus impressed, uniformly smooth to coriaceous, rectangular, trapezoid or quadrangular, with few long setae; ventrally rounded, emarginate, usually without median incision, sometimes with weak median incision; anterior tentorial pit large, rounded, epistomal sulcus and clypeo-pleurostomal line broad, well impressed. Frons and interocellar area coriaceous, with few short setae. Vertex, occiput alutaceous to coriaceous, sometimes with some rugae; postocciput, postgena alutaceous to smooth, shining with sparse white setae; posterior tentorial pit large, ovate, area below impressed; occipital foramen higher or shorter than height of postgenal bridge; hypostomal carina emarginate, continuing into postgenal sulci which run towards occipital foramen, dorsally bent outwards, postgenal bridge smooth, shining, anteriorly broader than posteriorly and slightly broader than width of occipital foramen. Antenna longer than head+mesosoma, with 12 flagellomeres.

Mesosoma with sparse short white setae. Pronotum laterally alutaceous in dorsal half, smooth, shining on rest surface, without irregular rugae, with sparse setae; propleuron alutaceous, shining, with dense white setae. Mesoscutum uniformly alutaceous-reticulate, with sparse white setae and numerous dense distinct piliferous points; slightly longer than broad (greatest width measured across mesoscutum at level of base of tegulae). Notaulus complete, deep, broad, with alutaceous bottom, posteriorly broader and slightly converging; anterior parallel and parapsidal lines indistinct, hardly traceable, indicated by smooth stripes; median mesoscutal line absent; parascutal carina narrow, smooth, reaching notaulus. Mesoscutellum delicately coriaceous to coriaceous-rugose, ovate, slightly longer than broad, sometimes with strong rugae laterally and posteriorly; posteriorly rounded, in some species with shallow invagination posterocentrally or posteriorly gradually narrowed into a centro-median tip; overhanging metanotum; mesoscutellar foveae quadrangular or transverse, broader than high; bottom smooth, shining, with or without irregular rugae, separated by narrow elevated coriaceous or rugose central area. Circumscutellar carina complete or not. Mesopleuron smooth to delicately alutaceous, speculum uniformly alutaceous to smooth, with few setae; mesopleural triangle smooth to delicate rugose, with dense setae; dorsal and lateral axillar areas smooth or alutaceous to coriaceous, shining, with dense white setae; subaxillular bar smooth, shining, posteriorly elongated towards mesoscutellum, most posterior part higher than height of metanotal trough; metapleural sulcus reaching mesopleuron at its midheight or in upper 1/3 of mesopleuron height; upper part of sulcus present or not. Metascutellum smooth or sculptured, as high or higher than the smooth ventral impressed area; metanotal trough smooth, shining, with sparse setae; central propodeal area smooth, shining, with or without delicate irregular rugae; lateral propodeal carinae bent outwards at mid height or in posterior 1/3 of propodeum height; lateral propodeal area smooth, shining, with or without piliferous points and long dense white setae. Nucha with irregular rugae, sometimes smooth and shining dorsocentrally. Tarsal claws toothed, with basal lobe.

Fore wing longer than body or brachypterous, hyaline, margin with long dense cilia, multiple slightly darker spots all over the wing, veins light brown, radial cell open, always short, 1.6–2.6× as long as broad; second abscissa of Rs always curved, usually strongly curved distally; R1 and Rs nearly reaching wing margin; areolet triangular, sometimes indistinct, Rs+M distinct or inconspicuous, traceable along entire length, reaching basalis in lower half of its height, rarely at mid-height.

Metasoma usually longer than head+mesosoma, sometimes as long as head+mesosoma, rarely shorter, higher than long in lateral view; second metasomal tergum extending over 1/2 to 4/5 of the metasoma length in dorsal view, with white setae anterolaterally, rarely setae absent, without or with band of micropunctures posteriorly; subsequent terga without micropunctures, rarely with micropunctures. Hypopygium without micropunctures except in a few species, prominent part of ventral spine of hypopygium needle-like shaped, 2.0–11.5× as long as broad in ventral view, usually with setae extending beyond apex of spine.

Body length 1.5–5.5 mm ( Kinsey 1936).

All species of Atrusca are known from the asexual generations only and all species are associated with white oaks only, Quercus (Quercus) section Quercus s.s. Phylogenetically Atrusca forms a very distinct clade, most closely related to Cynips (= Besbicus ), Philonix and Xanthoteras .

Below a key to species is given.

Key to Atrusca species

1. Fore wing with veins often faint, not impressed on all length, discontinuous or absent ( Figs 378 View FIGURES 378–381 , 392 View FIGURES 392–395 , 406 View FIGURES 406–409 ), always shorter than body, brachypterous............................................................................... pictor

1’. Fore wing with complete venation, brachypterous or normal length (e.g.: Figs 25 View FIGURES 25–28 , 85 View FIGURES 85–88 , 113 View FIGURES 113–116 )........................... 2

2. Rs of radial cell gradually curved, almost straight ( Figs 183 View FIGURES 183–185 , 457 View FIGURES 457–460 , 526 View FIGURES 526–529 , 540 View FIGURES 540–543 , 554 View FIGURES 554–557 )................................... 3

2’. Rs of radial cell strongly curved distally or the distal part of Rs with a different angle (e.g.: Figs 11 View FIGURES 11–14 , 25 View FIGURES 25–28 , 43 View FIGURES 43–46 , 57 View FIGURES 57–60 , 71 View FIGURES 71–74 ), never almost straight............................................................................................. 8

3. Body brown; head transversally ovate ( Figs 173 View FIGURES 173–178 , 447 View FIGURES 447–452 , 558 View FIGURES 558–563 ); interocellar area strongly elevated above head in frontal view and lateral ocelli prominent ( Figs 173 View FIGURES 173–178 , 44 View FIGURES 43–46 , 558 View FIGURES 558–563 ; ventral spine of hypopygium short, 2.5–3.0× as long as broad ( Figs 182 View FIGURES 179–182 , 458 View FIGURES 457–460 , 569 View FIGURES 568–571 ) ................................................................................................... 4

3’. Body reddish; head rounded ( Figs 516 View FIGURES 516–521 , 554 View FIGURES 554–557 ) or more or less quadrangular ( Fig. 530 View FIGURES 530–535 ); interocellar area not elevated above head in frontal view and lateral ocelli not or only slightly prominent ( Figs 516 View FIGURES 516–521 , 530 View FIGURES 530–535 , 544 View FIGURES 544–549 ); ventral spine of hypopygium long, 9.2–11.3× as long as broad ( Figs 527 View FIGURES 526–529 , 541 View FIGURES 540–543 , 555 View FIGURES 554–557 )............................................................. 6

4. Pronotum laterally alutaceous ( Fig. 179 View FIGURES 179–182 ); median mesoscutal line absent ( Fig. 181 View FIGURES 179–182 ); mesopleuron and speculum uniformly alutaceous to smooth, without piliferous points ( Fig. 179 View FIGURES 179–182 ).............................................. clivorum

4’. Pronotum laterally coriaceous-reticulate ( Fig. 453 View FIGURES 453–456 ) or with irregular rugae ( Fig. 564 View FIGURES 564–567 ); median mesoscutal line present ( Figs 454 View FIGURES 453–456 , 565 View FIGURES 564–567 ); mesopleuron and speculum with distinct sculpture and piliferous points ( Figs 453 View FIGURES 453–456 , 564 View FIGURES 564–567 )........................ 5

5. Pronotum rugose, with irregular rugae laterally, pubescent ( Fig. 564 View FIGURES 564–567 ); median mesoscutal line long, impressed, extending to 1/4 of mesoscutum length ( Fig. 565 View FIGURES 564–567 ); mesopleuron setose, centro-ventrally with transverse parallel delicate striae, dorsal part alutaceous-reticulate ( Fig. 564 View FIGURES 564–567 ); speculum alutaceous-reticulate ( Fig. 569 View FIGURES 568–571 ).................................. strians

5’. Pronotum uniformly alutaceous-reticulate, without setae ( Fig. 453 View FIGURES 453–456 ); median mesoscutal line superficial ( Fig. 454 View FIGURES 453–456 ); mesopleuron and speculum uniformly alutaceous-reticulate, without setae ( Fig. 453 View FIGURES 453–456 )............................. quercuscentricola

6. Lateral ocelli not prominent in frontal view ( Fig. 544 View FIGURES 544–549 ); pronotum with weak carinae only in upper half of posterior part ( Fig. 550 View FIGURES 550–553 ); mesoscutum predominantly weakly alutaceous, smooth between notauli in posterior 2/3 ( Fig. 552 View FIGURES 550–553 ); mesoscutellum with semilunar invagination posterocentrally ( Fig. 552 View FIGURES 550–553 )...................................................... spinifera

6’. Lateral ocelli slightly prominent in frontal view ( Figs 516 View FIGURES 516–521 , 530 View FIGURES 530–535 ); pronotum with rugae in posterior part ( Figs 522 View FIGURES 522–525 , 536 View FIGURES 536–539 ); mesoscutum uniformly coriaceous ( Figs 523 View FIGURES 522–525 , 537 View FIGURES 536–539 ); mesoscutellum rounded posteriorly ( Figs 524 View FIGURES 522–525 , 538 View FIGURES 536–539 )................. 7

7. Mesoscutum without darker stripes ( Fig. 523 View FIGURES 522–525 ); median mesoscutal line present in the form of a short, impressed triangle ( Fig. 523 View FIGURES 522–525 ); mesoscutellar foveae smooth, separated by a central area delimited by two carinae ( Figs 523–524 View FIGURES 522–525 )............ spinalis

7’. Mesoscutum with darker brown stripes along anterior parallel and parapsidal lines ( Fig. 537 View FIGURES 536–539 ); medial mesoscutal line absent ( Figs 537–538 View FIGURES 536–539 ); mesoscutellar foveae with irregular short rugae, separated by narrow elevated central carina ( Figs 537–538 View FIGURES 536–539 )............................................................................................ spinescens

8. Mesoscutellum coriaceous, sometimes with some lateral rugae (e.g.: Figs 40–41 View FIGURES 39–42 , 55 View FIGURES 53–56 , 69 View FIGURES 67–70 , 82–82 View FIGURES 81–84 , 144–145 View FIGURES 143–146 )............... 9

8’. Mesoscutellum entirely rugose (e.g: Figs 9 View FIGURES 7–10 , 23 View FIGURES 21–24 , 96 View FIGURES 95–98 , 111 View FIGURES 109–112 , 119 View FIGURES 118–122 ).................................................. 17

9. Body reddish to reddish brown, mesoscutum without darker stripes or with stripes only slightly darker than the color of the mesoscutum ( Figs 82 View FIGURES 81–84 , 214 View FIGURES 213–216 , 256 View FIGURES 254–257 ); fore wings ( Figs 85 View FIGURES 85–88 , 217 View FIGURES 217–220 , 258 View FIGURES 258–261 ) short, 0.76–1.15× as body length..................... 10

9’. Body brown to black (e.g.: Figs 33–42 View FIGURES 33–38 View FIGURES 39–42 , 44–45 View FIGURES 43–46 , 47–56 View FIGURES 47–52 View FIGURES 53–56 , 58 View FIGURES 57–60 , 61–70 View FIGURES 61–66 View FIGURES 67–70 ), rarely with lemon areas ( Figs 137–146 View FIGURES 137–142 View FIGURES 143–146 , 148–149 View FIGURES 147–150 ); mesoscutum with darker stripes ( Figs 40 View FIGURES 39–42 , 54 View FIGURES 53–56 , 68 View FIGURES 67–70 , 144 View FIGURES 143–146 , 169 View FIGURES 169–172 , 203 View FIGURES 200–204 , 269 View FIGURES 268–271 ); fore wing usually of normal size (1.21–1.40× as long as body length, ( Figs 43 View FIGURES 43–46 , 200 View FIGURES 200–204 , 272 View FIGURES 272–275 ), shorter only in A. cava (1.15x, Fig. 170 View FIGURES 169–172 ).................................................. 12

10. Pronotum delicately coriaceous in dorsolateral part, rest of pronotum laterally smooth, shining ( Fig. 213 View FIGURES 213–216 ); mesoscutellum posteriorly gradually narrowed into a centromedian tip, finished in a point ( Fig. 215 View FIGURES 213–216 ); brachypterous ( Fig. 217 View FIGURES 217–220 ), fore wing 0.76–0.95× than body length....................................................................... cubitalis

10’. Pronotum coriaceous with or without delicate rugae laterally ( Figs 81 View FIGURES 81–84 , 255 View FIGURES 254–257 ); mesoscutellum rounded distally ( Figs 83 View FIGURES 81–84 , 256 View FIGURES 254–257 ); brachypterous ( Figs 85 View FIGURES 85–88 , 258 View FIGURES 258–261 ), fore wing equal or longer than body but shorter than normal size....................... 11

11. Mesoscutum with conspicuous piliferous points ( Figs 82–83 View FIGURES 81–84 ); central propodeal area with transverse parallel delicate striae in anterior 1/3 and some irregular rugae posterolaterally ( Fig. 84 View FIGURES 81–84 ); fore wing ( Fig. 85 View FIGURES 85–88 ) shorter or subequal to body length (0.81–097× as long as body)................................................................... brevipennata

11’. Mesoscutum with very scarce or without piliferous points ( Fig. 256 View FIGURES 254–257 ); central propodeal area smooth ( Fig. 257 View FIGURES 254–257 ); fore wing ( Fig. 258 View FIGURES 258–261 ) longer than body (1.10–1.15× as long as body) but shorter than normal size............................... dugesi

12. Head, antenna, metasoma yellowish, mesosoma yellowish brown with darker brown areas ( Figs 137–146 View FIGURES 137–142 View FIGURES 143–146 , 148–149 View FIGURES 147–150 ); median mesoscutal line present, short, triangular ( Fig. 144 View FIGURES 143–146 ).................................................... capronae

12’. Head, mesosoma, metasoma reddish brown with dark to black areas ( Figs 33–42 View FIGURES 33–38 View FIGURES 39–42 , 44–45 View FIGURES 43–46 , 47–56 View FIGURES 47–52 View FIGURES 53–56 , 58 View FIGURES 57–60 , 61–70 View FIGURES 61–66 View FIGURES 67–70 , 72–73 View FIGURES 71–74 ); median mesoscutal line absent ( Figs 40–41 View FIGURES 39–42 , 55 View FIGURES 53–56 , 699, 169, 203, 269)................................................... 13

13. Mesoscutum with dense or sparse piliferous points, uniformly distributed all over mesoscutum ( Fig. 40 View FIGURES 39–42 ); radial cell with or without spot ( Figs 43 View FIGURES 43–46 , 170 View FIGURES 169–172 )............................................................................. 14

13’. Mesoscutum without piliferous points or with only few piliferous points along anterior parallel lines ( Figs 54–55 View FIGURES 53–56 , 68 View FIGURES 67–70 , 203 View FIGURES 200–204 , 269 View FIGURES 268–271 ); radial cell always with a spot ( Figs 57 View FIGURES 57–60 , 71 View FIGURES 71–74 , 201 View FIGURES 200–204 , 272 View FIGURES 272–275 )......................................................... 15

14. Head transversely quadrangular in frontal view with dense setae ( Fig. 171 View FIGURES 169–172 ); pronotum laterally with parallel rugae posteroventrally, extending to half width of pronotum ( Fig. 170 View FIGURES 169–172 ); mesoscutum with sparse piliferous points ( Fig. 169 View FIGURES 169–172 ); fore wing somewhat reduced; radial cell hyaline and Rs curved distally ( Fig. 170 View FIGURES 169–172 )................................... cava

14’. Head transversely ovate in frontal view with sparse setae ( Fig. 33 View FIGURES 33–38 ); pronotum without irregular rugae ( Fig. 39 View FIGURES 39–42 ); mesoscutum with numerous dense distinct piliferous points ( Fig. 40 View FIGURES 39–42 ); fore wing of normal size; radial cell with a spot and Rs strongly curved distally, forming a straight angle with margin of fore wing ( Fig. 43 View FIGURES 43–46 ).......................................... aspera

15. Head strongly transverse in frontal view ( Fig. 205 View FIGURES 205–206 ), narrower than mesosoma in front view; radial cell very short, around 1.3× as long as broad ( Fig. 201 View FIGURES 200–204 )....................................................................... congesta

15’. Head ovate in frontal view, as broad or slightly broader than mesosoma in front view ( Figs 47 View FIGURES 47–52 , 61 View FIGURES 61–66 , 262 View FIGURES 262–267 ); radial cell 1.8–2.1× as long as broad ( Figs 57 View FIGURES 57–60 , 71 View FIGURES 71–74 , 272 View FIGURES 272–275 )......................................................................... 16

16. Pronotum laterally smooth to alutaceous ( Figs 53 View FIGURES 53–56 , 67 View FIGURES 67–70 ); mesopleuron smooth, shining, speculum smooth to weakly alutaceous ( Figs 53 View FIGURES 53–56 , 67 View FIGURES 67–70 ); mesoscutellar foveae in a form of transverse impression ( Figs 55 View FIGURES 53–56 , 69 View FIGURES 67–70 ); lateral propodeal carinae weak, complete, sometimes inconspicuous subparallel or bent outwards at mid height of propodeum ( Figs 56 View FIGURES 53–56 , 70 View FIGURES 67–70 ); central propodeal with delicate subparallel longitudinal rugae along lateral propodeal carina in posterior part ( Figs 56 View FIGURES 53–56 , 70 View FIGURES 67–70 ); Rs uniformly curved ( Figs 57 View FIGURES 57–60 , 71 View FIGURES 71–74 )....................................................................... bella ( = vanescens , syn. nov.)

16’. Pronotum delicately coriaceous laterally ( Fig. 268 View FIGURES 268–271 ); mesopleuron and speculum uniformly delicately coriaceous ( Fig. 268 View FIGURES 268–271 ); mesoscutellar foveae separated by a medial elevated area ( Fig. 270 View FIGURES 268–271 ); lateral propodeal carinae complete, lyre–shaped, bent outwards in the middle ( Fig. 271 View FIGURES 268–271 ); central propodeal area smooth, without rugae ( Fig. 271 View FIGURES 268–271 ); Rs strongly curved distally ( Fig. 272 View FIGURES 272–275 )......................................................................................... dumosae

17. Ventral spine of hypopygium very long (10.0–10.5× as long as broad) with a few short setae ventrally which do not extend beyond apex of spine ( Figs 12–13 View FIGURES 11–14 , 232–233 View FIGURES 231–234 ); mesoscutellum with an invagination posterocentrally ( Figs 8 View FIGURES 7–10 , 228–229 View FIGURES 227–230 )..... 18

17’. Ventral spine of hypopygium variable in length, setose, with setae extending beyond apex of spine (e.g.: Figs 26 View FIGURES 25–28 , 100 View FIGURES 99–102 , 114 View FIGURES 113–116 , 118 View FIGURES 118–122 , 134 View FIGURES 133–136 ); mesoscutellum with or without an invagination posterocentrally........................................... 19

18. Head quadrangular in frontal view ( Fig. 1 View FIGURES 1–6 ); interocellar area not elevated in frontal view but ocelli prominent ( Fig. 1 View FIGURES 1–6 ); notauli with the same width along its entire length ( Fig. 8 View FIGURES 7–10 ); mesoscutum alutaceous-reticualte ( Fig. 9 View FIGURES 7–10 )................... aequalis

18’. Head ovate in frontal view ( Fig. 221 View FIGURES 221–226 ); interocellar area elevated above head ( Fig. 221 View FIGURES 221–226 ); notauli wider posteriorly ( Fig. 228 View FIGURES 227–230 ); mesoscutum uniformly rugose ( Fig. 228 View FIGURES 227–230 )............................................................ cucurbita

19. Mesopleuron and speculum completely smooth, always without piliferous points ( Figs 161 View FIGURES 161–164 , 192 View FIGURES 192–195 , 296 View FIGURES 296–299 , 310 View FIGURES 310–313 , 324 View FIGURES 324–327 , 508 View FIGURES 508–511 , 592 View FIGURES 592–595 ). 20

19’. Mesopleuron and/or speculum partially or completely sculptured, with or without piliferous points (e.g.: Figs 21 View FIGURES 21–24 , 95 View FIGURES 95–98 , 109 View FIGURES 109–112 , 118 View FIGURES 118–122 , 129 View FIGURES 129–132 )............................................................................................... 26

20. Color of body brown to black, mesoscutum with black stripes ( Figs 297 View FIGURES 296–299 , 311 View FIGURES 310–313 , 509 View FIGURES 508–511 )................................. 21

20’. Color of body reddish to reddish brown, mesoscutum without darker stripes or with stripes very little darker than the color of the mesoscutum, never black ( Figs 162 View FIGURES 161–164 , 193 View FIGURES 192–195 , 325 View FIGURES 324–327 , 593 View FIGURES 592–595 ).......................................................23

21. Pronotum delicately coriaceous ( Fig. 310 View FIGURES 310–313 ); mesoscutum alutaceous, between notauli in posterior 1/3 smooth ( Figs 311–312 View FIGURES 310–313 ); mesoscutellum posteriorly gradually narrowed till central part ( Fig. 312 View FIGURES 310–313 ); lateral propodeal carinae bent outwards in posterior 1/3 ( Fig. 313 View FIGURES 310–313 ); prominent part of ventral spine of hypopygium short, 5.5× as long as broad ( Fig. 315 View FIGURES 314–317 )............... longa

21’. Pronotum rugose laterally ( Figs 296 View FIGURES 296–299 , 508 View FIGURES 508–511 ); mesoscutum alutaceous to delicately coriaceous without smooth areas ( Figs 297 View FIGURES 296–299 , 509 View FIGURES 508–511 ); mesoscutellum posteriorly rounded, with distinct shallow invagination posterocentrally ( Figs 297–298 View FIGURES 296–299 , 510 View FIGURES 508–511 ); lateral propodeal carinae subparallel or slightly bent ( Figs 299 View FIGURES 296–299 , 511 View FIGURES 508–511 ); prominent part of ventral spine of hypopygium longer, 7.2–10.8× as long as broad ( Figs 300–301 View FIGURES 300–301 , 513–514 View FIGURES 512–515 )................................................................. 22

22. Inner margins of eyes parallel ( Fig. 290 View FIGURES 290–295 ); F9–F10 at least 2.0× as long as broad ( Fig. 294 View FIGURES 290–295 ); lateral propodeal area smooth ( Fig. 299 View FIGURES 296–299 ); prominent part of ventral spine of hypopygium very long, 10.8× as long as broad in ventral view ( Figs 300–301 View FIGURES 300–301 ).................................................................................................... finitima

22’. Inner margins of eyes slightly diverging ventrally ( Fig. 503 View FIGURES 503–507 ); F9–F10 subquadrate or slightly longer than broad ( Fig. 507 View FIGURES 503–507 ); lateral propodeal area coriaceous ( Fig. 511 View FIGURES 508–511 ); ventral spine of hypopygium shorter, 7.2× as long as broad ( Figs 513–514 View FIGURES 512–515 )................................................................................................... spiculi

23. Ventral spine of hypopygium long, around 10.0× as long as wide ( Figs 597–598 View FIGURES 596–599 ); last flagellomere segments long, 2.0× as long as wide ( Fig. 590 View FIGURES 586–591 )................................................................................. tigrina

23’. Ventral spine of hypopygium shorter 2.8–4.0× as long as wide ( Figs 166–167 View FIGURES 165–168 , 197–198 View FIGURES 196–199 , 329–330 View FIGURES 328–330 ); last flagellomere segments shorter ( Figs 190 View FIGURES 186–191 , 322 View FIGURES 318–323 )................................................................................ 24

24. In frontal view lateral ocelli small and shortly prominent ( Fig. 186 View FIGURES 186–191 ); inner margins of eyes strongly divergent ventrally ( Fig. 186 View FIGURES 186–191 ); OOL equal to diameter of lateral ocellus ( Fig. 187 View FIGURES 186–191 ); Rs arcuate-angulate ( Fig. 196 View FIGURES 196–199 )........................ conexa

24’. In frontal view lateral ocelli big and strongly prominent ( Figs 155 View FIGURES 155–160 , 318 View FIGURES 318–323 ); inner margins of eyes parallel or shortly diverging ventrally ( Figs 155 View FIGURES 155–160 , 318 View FIGURES 318–323 ); OOL longer than 2.0× the diameter of lateral ocellus ( Figs 156 View FIGURES 155–160 , 319 View FIGURES 318–323 ); Rs curved ( Figs 165 View FIGURES 165–168 , 328 View FIGURES 328–330 )... .................................................................................................. 25

25. Head rusty brown ( Fig. 155 View FIGURES 155–160 ); mesoscutum with light brown stripes ( Fig. 162 View FIGURES 161–164 ); mesoscutellum posteriorly rounded ( Fig. 163 View FIGURES 161–164 ); central propodeal area smooth without rugae and striae ( Fig. 164 View FIGURES 161–164 ); central propodeal area smooth ( Fig. 164 View FIGURES 161–164 ); fore wings hyaline, with few slightly darker spots and stripes ( Fig. 165 View FIGURES 165–168 )...................................................... catena

25’. Head dark brown with central area of lower face black ( Fig. 318 View FIGURES 318–323 ); mesoscutum with dark brown stripes ( Fig. 325 View FIGURES 324–327 ); mesoscutellum distinctly invaginated posterocentrally ( Figs 325–326 View FIGURES 324–327 ); central propodeal area with strong irregular short rugae ( Fig. 327 View FIGURES 324–327 ); fore wings infuscated, with numerous darker spots and infuscations ( Fig. 328 View FIGURES 328–330 ).................................... lucaris

26. Body reddish to reddish brown, mesoscutum without darker stripes or with stripes slightly darker than the color of mesoscutum ( Figs 338 View FIGURES 337–340 , 347 View FIGURES 346–350 , 439 View FIGURES 438–442 , 359 View FIGURES 358–361 , 496 View FIGURES 495–498 , 607 View FIGURES 606–609 )....................................................................... 27

26’. Body brown to black with black stripes (e.g.: Fig. 22 View FIGURES 21–24 , 96 View FIGURES 95–98 , 110 View FIGURES 109–112 , 119 View FIGURES 118–122 , 130 View FIGURES 129–132 )......................................... 32

27. Mesopleuron anteriorly and posteriorly rugose, with transversely oriented parallel rugae, in central part smooth ( Fig. 337 View FIGURES 337–340 )................................................................................................ luminata

27’. Mesopleuron without linear elements ( Figs 346 View FIGURES 346–350 , 358 View FIGURES 358–361 , 440 View FIGURES 438–442 , 495 View FIGURES 495–498 , 606 View FIGURES 606–609 )............................................ 28

28. Mesopleuron and speculum uniformly alutaceous ( Fig. 346 View FIGURES 346–350 ); Rs strongly angulated distally ( Fig. 345 View FIGURES 344–345 )............. occidua

28’. Mesopleuron with smooth areas, speculum sculptured or not ( Figs 358 View FIGURES 358–361 , 440 View FIGURES 438–442 , 495 View FIGURES 495–498 , 606 View FIGURES 606–609 ); Rs arcuate, sometimes slightly angulated distally ( Figs 362 View FIGURES 362–365 , 442 View FIGURES 438–442 , 499 View FIGURES 499–502 , 610 View FIGURES 610–613 )........................................................................ 29

29. OOL 1.2× as long as diameter of lateral ocellus ( Fig. 490 View FIGURES 489–494 )................................................. spadix

29’. OOL longer, 2.1–2.6× as long as diameter of lateral ocellus ( Figs 353 View FIGURES 352–357 , 444 View FIGURES 443–446 , 601 View FIGURES 600–605 )................................... 30

30. Mesoscutum uniformly alutaceous-reticulate, with sparse white setae and numerous distinct piliferous points ( Fig. 439 View FIGURES 438–442 ).................................................................................................. pupoides

30’. Mesoscutum smooth and shining at least between notauli in posterior half with sparse indistinct piliferous points in the posterior half ( Figs 359 View FIGURES 358–361 , 607 View FIGURES 606–609 )................................................................................... 31

31. Pronotum laterally delicately alutaceous, without linear elements or with very weak and short rugae between coriaceous sculpture ( Fig. 358 View FIGURES 358–361 ); mesopleuron without piliferous points ( Fig. 358 View FIGURES 358–361 ); radial cell relatively long, around 2.1× as long as broad ( Fig. 362 View FIGURES 362–365 )........................................................................................ oriens

31’. Pronotum with rugae or carinae ( Fig. 606 View FIGURES 606–609 ); mesopleuron with piliferous points ( Fig. 606 View FIGURES 606–609 ); radial cell shorter, around 1.6× as long as broad ( Fig. 610 View FIGURES 610–613 ).............................................................................. vasta

32. Ventral spine of hypopygium long,7.7–11.3× as long as wide ( Figs 26 View FIGURES 25–28 , 100 View FIGURES 99–102 , 114 View FIGURES 113–116 , 118 View FIGURES 118–122 , 134 View FIGURES 133–136 , 472 View FIGURES 471–474 )....................... 33

32’. Ventral spine of hypopygium shorter 2.5–4.5× as long as wide ( Figs 246 View FIGURES 245–248 , 287 View FIGURES 286–289 , 435 View FIGURES 434–437 , 486 View FIGURES 485–488 , 583 View FIGURES 582–585 )........................ 38

33. Mesoscutum without piliferous points or very indistinct at least in the posterior area between notauli ( Figs 96–97 View FIGURES 95–98 , 468 View FIGURES 467–470 ).... 34

33’. Mesoscutum with conspicuous piliferous points ( Figs 22 View FIGURES 21–24 , 110 View FIGURES 109–112 , 119 View FIGURES 118–122 , 130 View FIGURES 129–132 )......................................... 35

34. Mesoscutum uniformly alutaceous ( Fig. 96 View FIGURES 95–98 ); mesopleuron and speculum uniformly alutaceous ( Fig 95 View FIGURES 95–98 )........... bulbacea

34’. Mesoscutum rugose in anterior half, smooth to alutaceous in posterior half ( Fig. 468 View FIGURES 467–470 ); Mesopleuron along anterior edge alutaceous, rest smooth, speculum smooth ( Fig. 467 View FIGURES 467–470 )..................................................... sierrae

35. Body partly reddish brown, black areas only on mesoscutum and partially on mesopleuron ( Figs 15–24 View FIGURES 15–20 View FIGURES 21–24 , 123–132 View FIGURES 123–128 View FIGURES 129–132 ); mesoscutellum usually with strong invagination postero-centrally ( Figs 22 View FIGURES 21–24 , 130 View FIGURES 129–132 ); cubital cell with different infuscate patches ( Figs 25 View FIGURES 25–28 , 133 View FIGURES 133–136 ). .................................................................................................. 36

35’. Body dark with big black areas ( Figs 103–112 View FIGURES 103–108 View FIGURES 109–112 , 118–119 View FIGURES 118–122 ); mesoscutellum rounded posteriorly ( Figs 111 View FIGURES 109–112 , 119 View FIGURES 118–122 ); spots in cubital cell forming a line parallel to Rs ( Figs 113 View FIGURES 113–116 , 118 View FIGURES 118–122 )............................................................ 37

36. Median mesoscutal line present (Fug. 130); mesoscutellum posteriorly rounded with a small median invagination ( Figs 130– 131 View FIGURES 129–132 ); metasomal terga smooth, without micropunctures ( Fig. 134 View FIGURES 133–136 )........................................... bulla

36’. Median mesoscutal line absent ( Fig. 22 View FIGURES 21–24 ); mesoscutellum posteriorly bilobate, with deep posteromedial invagination ( Fig. 22 View FIGURES 21–24 ); second metasomal tergum with band of micropunctures posteriorly and subsequent terga with delicate micropunctures ( Fig. 26 View FIGURES 25–28 ).......................................................................................... aggregata 37. Head more or less rounded in frontal view, gena broadened behind eyes ( Fig. 122 View FIGURES 118–122 )............................ bulbulus

37’. Head transversally quadrangular, gena not broadened behind eyes ( Fig. 103 View FIGURES 103–108 )................................ bulboides

38. Pronotum coriaceous at least in part, without striae or carinae but sometimes with very weak linear elements between coriaceous sculpture ( Figs 282 View FIGURES 282–285 , 430 View FIGURES 430–433 ); speculum alutaceous ( Figs 282 View FIGURES 282–285 , 430 View FIGURES 430–433 )................................................ 39

38’. Pronotum with some rugae laterally or with short transverse parallel striae along posterior edge ( Figs 241 View FIGURES 241–244 , 481 View FIGURES 481–484 , 578 View FIGURES 578–581 ); speculum smooth ( Figs 241 View FIGURES 241–244 , 481 View FIGURES 481–484 , 578 View FIGURES 578–581 )............................................................................ 40

39. Head transversally ovate in frontal view ( Fig. 424 View FIGURES 424–429 ); mesoscutellum with smooth foveae ( Fig. 432 View FIGURES 430–433 ); lateral propodeal area without piliferous points ( Fig. 433 View FIGURES 430–433 ); central propodeal with some delicate irregular rugae ( Fig. 433 View FIGURES 430–433 ); radial cell with a spot ( Fig. 434 View FIGURES 434–437 ).......................................................................................... pomifera

39’. Head more or less triangular in frontal view ( Fig. 276 View FIGURES 276–281 ); mesoscutellar foveae with rugose bottom ( Fig. 284 View FIGURES 282–285 ); lateral propodeal area with piliferous points ( Fig. 285 View FIGURES 282–285 ); central propodeal without rugae ( Fig. 285 View FIGURES 282–285 ); radial cell hyaline ( Fig. 286 View FIGURES 286–289 ).... emergens

40. Mesoscutum without piliferous points or very indistinct at least in the posterior area between notauli ( Fig. 579 View FIGURES 578–581 ); mesoscutellum slightly pointed distally ( Fig. 578 View FIGURES 578–581 ); Rs strongly angulate distally ( Fig. 582 View FIGURES 582–585 ); mesoscutellar foveae with some irregular rugae ( Fig. 580 View FIGURES 578–581 )...................................................................................... subnigra

40’. Mesoscutum with conspicuous piliferous points ( Figs 242 View FIGURES 241–244 , 482 View FIGURES 481–484 ); mesoscutellum with shallow invagination posterocentrally, sometimes inconspicuous ( Figs 242 View FIGURES 241–244 , 482 View FIGURES 481–484 ); Rs angulate but not strongly ( Figs 245 View FIGURES 245–248 , 485 View FIGURES 485–488 ); mesoscutellar foveae smooth, without rugae ( Figs 243 View FIGURES 241–244 , 482–483 View FIGURES 481–484 ).............................................................................. 41

41. Body densely pubescent with large black areas including lower face below toruli ( Figs 475 View FIGURES 475–480 , 481 View FIGURES 481–484 ); pronotum rugose, with short irregular rugae posteriorly ( Fig. 481 View FIGURES 481–484 ); Rs distinctly angulate, curved distally ( Fig. 485 View FIGURES 485–488 )........................ simulatrix

41’. Body scarcely pubescent, with black areas reduced to mesoscutum stripes ( Figs 235 View FIGURES 235–240 , 241 View FIGURES 241–244 ); pronotum with transverse parallel striae along posterior edge ( Fig. 241 View FIGURES 241–244 ); Rs uniformly arcuate ( Fig. 245 View FIGURES 245–248 )...................................... deceptrix

Below all known Atrusca species are listed in alphabetical order.