Neotheronia Krieger, 1899

Khalaim, Andrey I. & Ruíz-Cancino, Enrique, 2021, Darwin wasps of the subfamily Pimplinae (Hymenoptera: Ichneumonidae) of Mexico: genera Neotheronia Krieger, Nomosphecia Gupta and Xanthopimpla Saussure, Zootaxa 4950 (3), pp. 401-440 : 403-405

Genus Neotheronia Krieger, 1899

Neotheronia Krieger 1899: 119 . Type species: Theronia tolteca Cresson, 1874 , by original designation.

Synonyms:

Acrocremnus Brèthes, 1926 .

Epimecoideus Ashmead, 1900 .

References. Cresson 1874: 395 [ Theronia ; 4 new species from Mexico]. Ashmead 1900a: 52 [ Epimecoideus (description)]. Krieger 1905 [revision; key to 62 Neotropical species and forms (53 described as new)]. Morley 1914: 48 [review of 55 species (1 described as new); key]. Cushman 1920: 44 [remarks on taxonomy], 44 [ Neotheronia ; review of 2 species from North America; key]. Townes & Townes 1960: 351 [review of 4 Nearctic species of Theronia (including 2 species of Neotheronia ); key]. Townes & Townes 1966: 30 [ Theronia (Neotheronia) ; Acrocremnus (syn.); catalogue; 57 species and subspecies in Neotropical region; 7 or 8 species and subspecies in Mexico]. Carlson 1979: 347 [catalogue; 2 species in North America]. Gauld 1991: 386 [description; review of 29 species from Costa Rica (19 described as new); key]. Gauld et al. 1998: 127 [30 species in Costa Rica (1 described as new); key]. Gauld et al. 2002: 49 [review of 12 species from El Salvador; key]. Ruíz-Cancino et al. 2002: 645 [checklist; 9 species in Mexico].

Neotheronia is a large, predominantly Neotropical genus with 76 species. Four species occur in Africa and the rest are mostly in tropical America. Only two species, N. bicincta and N. septemtrionalis (= nigrolineata ), occur in the southeastern USA. Two species, N. bicincta and N. nubecula ( Cresson, 1865) , occur in Cuba; and N. fuscicornis ( Brullé, 1846) is known only from Guadeloupe (The Caribbean). In Central America, 30 species occur in Costa Rica, and N. hastata Krieger, 1905 is known only from Panama. Brazilian species N. kohli Krieger, 1905 was mentioned from Nicaragua in the catalogue by Maes (1989), but this record requires confirmation.

Twenty four species are found to occur in Mexico, including five undescribed species, and seven are new records for the country. South American species N. cristata is excluded from the Mexican fauna, see explanations in the Remarks section under this species.

The species of Neotheronia are primary parasitoids of pupae or cocooned prepupae of various Lepidoptera , or secondary parasitoids via Ichneumonidae (Hymenoptera) or Tachinidae (Diptera) parasitizing primary hosts.

Key to Mexican species of Neotheronia

1. Head, in dorsal view, with a black mark extending from lateral ocellus to margin of eye ( Fig. 20 ), AND back of head always with a strong mediodorsal notch ( Fig. 20 ).................................................................. 2

- Head, in dorsal view, with area between lateral ocellus and margin of eye yellowish, at most slightly infuscate adjacent to ocellus. Back of head with or without a mediodorsal notch in occiput............................................ 9

2. Wings strongly tinged with orange-brown, subapically infuscate. Hind femur reddish brown. Tergites 1–2(or 3) anteriorly brown or black, posteriorly yellow, following tergites more or less uniformly reddish brown. Clypeus with rounded median transverse ridge.................................................................. N. bicincta ( Cresson, 1865)

- Wings hyaline or weakly yellowish, with or without fuscous spot near apex. Hind femur usually bright yellow. Metasoma with colour pattern not as above. Clypeus without transverse ridge, more or less flat.................................... 3

3. Flagellum basally blackish, centrally yellowish or brownish, and usually more or less darkened apically. Scutellum posteriorly usually black marked ( Fig. 39 ). Head with posterior transverse blackish band on vertex (as in Figs 5 and 20 ). Scutellum with lateral longitudinal carinae only present on extreme anterior end. Mesopleuron yellow with anterior and posteroventral margins black marked. Hind tarsus more or less entirely yellow or brownish yellow, sometimes with apical tarsomere slightly infuscate. Ovipositor 2.2–2.5× as long as hind tibia, slightly upcurved.................................................... 4

- Flagellum with most proximal part brown, distally infuscate, or flagellum is entirely black. Other characters varied....... 5

4. Propodeum with anterior margin, transverse mark immediately before transverse carina and median longitudinal mark extending from insertion of metasoma towards transverse carina black ( Fig. 39 ). Upper tooth of mandible slightly shorter than lower tooth; teeth sharp, not unusually broad ( Fig. 38 ). Propodeum with median and lateral longitudinal carinae present (though sometimes incomplete) in front of transverse carina ( Fig. 39 )..................................... 19. N. rosai Gauld

- Propodeum with anterior margin and dorsal inverted U-shaped mark black. Upper tooth of mandible as long as lower tooth; teeth broad and somewhat blunt. Propodeum with median and lateral longitudinal carinae absent in front of transverse carina.................................................................................. 3. N. bostrandae Gauld

5. Lateral longitudinal carinae of propodeum present in front of transverse carina, but generally not completely enclosing area spiracularis ( Figs 31, 32 ); median longitudinal carinae usually complete in front of the transverse carina ( Figs 31, 32 ), rarely (especially in small specimens) weak or incomplete. Propodeum usually with a median longitudinal black mark extending from insertion of metasoma to transverse carina ( Fig. 32 ), but sometimes only slightly blackish or brownish posteriorly ( Fig. 31 ). 6

- Lateral longitudinal carinae of propodeum completely absent in front of transverse carina ( Figs 21, 22 ); median longitudinal carinae usually completely absent. Flagellum usually entirely black. Propodeum variously coloured, if posterior longitudinal black mark present, then it is large and triangular ( Fig. 21 )..................................................... 7

6. Scutellum entirely yellow. Hind tarsus orange with distal tarsomere blackish. Flagellum dorsally with most proximal part brown, distally infuscate. Ovipositor projecting beyond margin of subgenital plate by about 1.5× length of hind tibia; apex of upper valve smooth....................................................................... 11. N. lineata (F.)

- Scutellum posteriorly black marked. Hind tarsus entirely black. Flagellum (in Mexican specimen) dorsally entirely black. Ovipositor shorter, projecting beyond margin of subgenital plate by about length of hind tibia; apex of upper valve on its lateral sides bearing several weak transverse ridges............................................ 8. N. hespenheidei Gauld

7. Fore wing with pterostigma black. Hind tarsus extensively blackish, narrowly yellowish basally. Flagellum black. Propodeum with distinct subtriangular median black mark extending from insertion of metasoma to transverse carina, without any longitudinal carinae in front of transverse carina ( Fig. 21 ). Mesopleuron entirely yellow............ 1. N. altacima sp. nov.

- Fore wing with pterostigma pale brown or yellowish. Hind tarsus entirely yellow or yellowish orange, sometimes with only apical segment blackish ( Fig. 23 ). Flagellum sometimes extensively brown. Propodeum without subtriangular median black mark posteriorly, but sometimes with a transverse black mark along anterior margin of transverse carina ( Fig. 40 ), with median longitudinal carinae in front of transverse carina often more or less developed. Mesopleuron with or without black marks... 8

8. Mesopleuron entirely yellow. Propodeum entirely yellow, groove in front of propodeum sometimes black ( Fig. 22 ). Legs bright yellow, without brownish or black marks on coxae and femora, hind tarsus with distal tarsomere blackish ( Fig. 23 ). Wings rather strongly infumate with yellow-brown. Flagellum, in dorsal view, entirely black................ 2. N. bonita sp. nov.

- Mesopleuron yellow with black markings on anterior and posterior margins. Propodeum yellow with anterior margin black and a transverse black mark along anterior margin of transverse carina ( Fig. 40 ). Legs yellow to orange, with brownish or black dorsal stripes on hind coxa and on internal surface of hind femur; hind tarsus often entirely yellow or orange, rarely with tarsomere 5 more or less infuscate. Wings weakly yellowish. Flagellum, in dorsal view, usually more or less brownish..................................................................................... 20. N. tacubaya (Cresson)

9. Mandible very strongly tapered so distal apex is barely wider than malar space ( Fig. 27 ). Clypeus unusually small ( Fig. 27 ) 10

- Mandible weakly to moderately tapered so distal apex is at least twice as wide as malar space ( Figs 38, 41 ). Clypeus larger ( Figs 38, 41 )............................................................................................. 11

10. Propodeum and metasoma entirely yellow, without black markings. Subalar prominence weakly convex ( Fig. 29 ). Epomia reduced, only the upper subvertical part discernible, the part parallel with pronotal margin absent ( Fig. 28 ). Female: Tergite 1 with lateromedian longitudinal carinae extending back almost to, or beyond the level of spiracles. Ovipositor slightly upcurved, long, projecting beyond margin of subgenital plate by 1.4–1.5× length of hind tibia. Male: Posterior end of metasoma subcylindrical. Subgenital plate elongate. Propodeum without any trace of vestiges of median longitudinal carinae anteriorly.............................................................................. 16. N. montezuma (Cresson)

- Anterior margin of propodeum blackish, metasomal tegites 2+ with blackish transverse subbasal bands. Subalar prominence in the form of a sharp ridge ( Fig. 26 ). Epomia with a distinct lower part which is parallel to pronotal margin ( Fig. 25 ). Female: Tergite 1 with lateromedian longitudinal carinae present only anteriorly as vestiges. Ovipositor slightly decurved, shorter, projecting beyond margin of subgenital plate by 0.9–1.0× length of hind tibia. Male: Posterior end of metasoma somewhat depressed. Subgenital plate slightly transverse. Propodeum with tubercles anteriorly which represent vestiges of median longitudinal carinae................................................................... 10. N. jugaldei Gauld

11. Clypeus with distinct median transverse ridge ( Fig. 18 ). Tergite 2 of metasoma with clearly delineated, elliptical gastroceli. Sternite 1 with a strongy developed medioventral keel anteriorly............................................... 12

- Clypeus centrally more or less flat (rarely transversely creased in abnormal specimens). Tergite 2 of metasoma with weak to strong oval gastroceli. Sternite 1 without or with weak medioventral keel anteriorly................................ 14

12. Pterostigma translucent brownish yellow. Tergite 1 stout, 1.4–1.6× as long as posteriorly broad. Female with ovipositor projecting beyond apex of subgenital plate by 1.8–1.9× length of hind tibia....................... 7. N. donovani Gauld

- Pterostigma dark brown to brownihs black. Tergite 1 slender, 1.9–2.3× as long as posteriorly broad. Female with ovipositor projecting beyond apex of subgenital plate by 1.4–1.7× length of hind tibia...................................... 13

13. Hind leg yellow, tarsus pale basally and black apically, often only apical tarsomere black........... 6. N. concolor Krieger

- Hind leg yellow with both trochanters, apical 0.2 of femur and entire tarsus black ( Fig. 19 )......... 24. N. zaldivari sp. nov.

14. Back of head without a mediodorsal notch in occiput ( Fig. 37 )................................................ 15

- Back of head with a strong mediodorsal notch in occiput (as in Fig. 20 )......................................... 16

15. Propodeum anteriorly usually brownish or blackish marked. Metasoma orange with anteromedian transverse bands on tergites 2 and 3 usually brown to black, but sometimes (especially in small specimens) body is entirely orange without any dark markings. Hind tarsus orange, at most only one or two apical tarsomeres infuscate. Ovipositor projecting beyond apex of subgenital plate by 1.4–1.8× length of hind tibia....................................................... 15. N. mellosa (Cresson)

- Propodeum entirely orange-brown. Metasoma unicolorously orange-brown. Hind tarsus more or less entirely black or dark reddish brown. Ovipositor shorter, projecting beyond apex of subgenital plate by 1.2–1.4× length of hind tibia........................................................................................ 5. N. chiriquensis (Cameron)

16. Mid tarsus mostly or entirely black. Hind leg with apex of coxa, apical third of femur and tarsus black ( Figs 6 , 17 )....... 17

- All legs yellow to orange, only hind tarsus partly or entirely black or blackish.................................... 18

17. Hind leg with trochanters and tibia entirely yellow ( Fig. 17 ). Ovipositor short, not projecting beyond apex of metasoma ( Fig. 16 )................................................................................ 9. N. juanitae sp. nov.

- Hind leg with trochanters and base and apex of tibia black ( Fig. 6 ). Ovipositor projecting beyond hind margin of subgenital plate by 1.1× length of hind tibia ( Fig. 6 ).................................................. 21. N. tequila sp. nov.

18. Propodeum with median longitudinal carinae strong, extending from anterior margin back to transverse carina ( Fig. 24 ). Propodeum and metasoma entirely orange................................................................. 19

- Propodeum with median longitudinal carinae completely absent or represented by short tubercles anteriorly ( Fig. 35 ); lateral longitudinal carina completely absent anteriorly, present or absent posteriorly. Propodeum and metasoma sometimes black marked............................................................................................ 21

19. First tergite with lateromedian longitudinal carinae completely absent. Female with ovipositor long, projecting beyond hind margin of subgenital plate by 1.7–2.1× length of hind tibia.................................... 17. N. murilloi Gauld

- First tergite with lateromedian longitudinal carinae well developed at least basally. Female with ovipositor very short, not projecting beyond hind end of metasoma.................................................................. 20

20. First tergite with lateromedian longitudinal carinae very strong, reaching almost to hind margin of tergite, and for their posterior 0.4 parallel and very close to each other, separated by less than thickness of flagellum. Second tergite with very broad, large gastrocoeli, which are separated medially by a ridge which is narrower than 0.3 of the width of either gastrocoelus. Notauli very strongly impressed...................................................................... 13. N. lloydi Gauld

- First tergite with lateromedian longitudinal carinae present only anteriorly, not reaching beyond level of spiracles. Second tergite with gastrocoeli moderately impressed, medially separated by more than 0.5 of width of gastrocoelus. Notauli weakly impressed............................................................................. 4. N. charli Gauld

21. Face strongly protuberant below antennal insertion, and with a deep furrow extending between and below antennal bases. Female with ovipositor very short, barely projecting beyond apex of metasoma ( Fig. 36 )............................ 22

- Face not especially protuberant below antennal insertion. Female with ovipositor long, distinctly projecting beyond apex of metasoma.......................................................................................... 23

22. Propodeum with lateral longitudinal carinae posteriorly (behind transverse carina) present. Body generally orange, mesosoma and metasoma without black markings...................................................... 12. N. lizzae Gauld

- Propodeum with longitudinal carinae completely absent ( Fig. 35 ). Body yellow with black mark on occiput dorsally ( Fig. 33 ), stripe on posterior lower margin of mesopleuron, anterior margin of propodeum and subbasal transverse bands on tergites 1–6 ( Fig. 36 )...................................................................... 23. N. veracruzana Khalaim

23. Metasoma uniformly brownish orange, without conspicuous black banding. Propodeum uniformly orange-brown........................................................................................... 22. N. tolteca (Cresson)

- Metasoma yellow with conspicuous anteromedial transverse black bands on all tergites. Propodeum anteriorly black marked............................................................................... 18. N. nigrolineata (Brullé)

Figures

FIGURES 20–24. Neotheronia altacima sp. nov., ♀: 20—head, dorso-postero-lateral; 21—propodeum, dorsal. Neotheronia bonita sp. nov., ♀: 22—propodeum, dorsal; 23—hind tarsus, lateral. Neotheronia charli, ♀: 24—propodeum, dorsal.

FIGURES 37–41. Neotheronia mellosa, ♀: 37—head, dorso-postero-lateral. Neotheronia rosai, ♀: 38—head, antero-ventral; 39—mesosoma, dorsal. Neotheronia tacubaya, ♀: 40—propodeum, dorsal. Neotheronia tequila sp. nov., ♀: 41—head, front.

FIGURES 3–7. Neotheronia tacubaya, ♀: 3—head, front; 4—propodeum, dorso-posterior. Neotheronia tequila sp. nov., holotype, ♀: 5—head, dorso-postero-lateral; 6—habitus (without apices of antennae and wings), lateral; 7—head and mesosoma, lateral.

FIGURES 31–36. Neotheronia lineata, ♀: 31, 32—propodeum, dorsal. Neotheronia veracruzana, ♀ (Tamaulipas): 33—head, dorso-posterior; 34—mesoscutum, dorsal; 35—propodeum, dorsal; 36—posterior part of metasoma, lateral.

FIGURES 25–30. Neotheronia jugaldei, ♀: 25—pronotum, lateral; 26—mesopleuron, front. Neotheronia montezuma, ♀ (27– 29) and ♂ (30): 27—head, front; 28—pronotum, lateral; 29—mesopleuron, front; 30—metasoma, dorsal.

FIGURES 16–19. Neotheronia juanitae sp. nov., holotype, ♀: 16—apex of metasoma with ovipositor, lateral; 17—hind leg, lateral. Neotheronia zaldivari sp. nov., holotype, ♀: 18—head, antero-lateral; 19—mid and hind legs, lateral.