Ormiophasia Townsend, 1919

Ormiophasia Townsend, 1919: 164 . Type species: Ormiophasia busckii Townsend, 1919, by original designation.

Pseudoneoptera Séguy, 1926b: 19 . Type species: Pseudoneoptera morardi Séguy, 1926b, by monotypy; Séguy (1926b: 20, key to genera); Séguy (1927a: 423, key to genera; 424, catalog); Townsend (1931: 82, synonymy with Ormiophasia); Sabrosky (1953: 181, as Ormiophasia, catalog); Tavares (1964: 38, comments on synonymy).

Plagiatormia Séguy, 1926b: 19 (as Plagiotormia, incorrect original spelling). Type species: Plagiatormia obscura Séguy, 1926b, by monotypy; Séguy (1926b: 20, key to genera); Séguy (1927a: 423, as Plagiotormia, misspelling, key to genera; 424, catalog); Townsend (1931: 82, synonymy with Ormiophasia); Sabrosky (1953: 181, as Ormiophasia, catalog); Tavares (1964: 38, comments on synonymy).

Pseudormia Séguy, 1927b: 262 . Type species: Pseudormia inflata Séguy, 1927b, by monotypy; Séguy (1925: 375, nomen nudum, comparison to Ormia; 376, key to genera); Séguy (1926a: 5, nomen nudum, comparison to Ormia; 9, as Peudormia, misspelling, key to genera); Séguy (1926b: 20, nomen nudum, key to genera); Séguy (1927a: 424, nomen nudum, key to genera, catalog); Townsend (1931: 82, synonymy with Ormiophasia); Sabrosky (1953: 181, as Ormiophasia, catalog); Tavares (1964: 38, comments on synonymy).

Diversity and distribution. The ranges of the 16 species currently ascribed to Ormiophasia cover an area from Southeast Mexico to northern Argentina.

References. Townsend (1919: 164, description of Ormiophasia and of the type species O. busckii); Aldrich (1922: 5, comments on Ormia, treated O rmiophasia as synonym of Ormia); Townsend (1927: 223, key to genera of Neotropical Muscoidea [sensu Townsend]); Malloch (1929: 279, as Ormia, comments on validity of Ormiophasia); Townsend (1931: 82, synonymized Plagiatormia, Pseudormia and Pseudoneoptera with Ormiophasia); Townsend (1936: 100, comments on larval characters; 101, key to genera of Ormiini, comments on synonymies of Plagiatormia, Pseudormia and Pseudoneoptera); Townsend (1938: 236, redescription, description of larva); Townsend (1942: 325, illustration of larva); Sabrosky (1953: 171, key to genera of New World Ormiini; 172, key; 181, catalog, comments on synonymies of Plagiatormia, Pseudormia and Pseudoneoptera); Thompson (1963: 455, comments on larvae being related to Euphasiopteryx and Ormia); Tavares (1964: 37–52, taxonomic revision, comments on synonymies of Plagiatormia, Pseudormia and Pseudoneoptera); Tavares (1965a: 14, key to genera of Neotropical Ormiini); Guimarães (1971: 22; catalog); Tschorsnig (1985: 55, illustration of male ejaculatory apodeme; 97, description of male terminalia of Ormiini); Gramajo (1997: 96, record of Ormiophasia, as Ormiaphasia, misspelling); Toma & Nihei (2006: 243, catalog of type material); Wood & Zumbado (2010: 1410, as Ormia Robineau-Desvoidy; treated Ormiophasia as synonym of Ormia); Evenhuis et al. (2015: 197, catalog of Townsend’s genera); Nihei (2016: 914, catalog of Colombiam Tachinidae); O’Hara (2018: 55, catalog).

Diagnosis. Ormiophasia resembles Ormia, sharing many characters with this genus, such as: occiput extremely concave, especially in females; presternum with a pair of pits on anterior surface (Fig. 1); anterior surface of distiphallus membranous (Figs 38–42); hypoproct with a lateral patch of very short, spiniform setae (Fig. 3B); and female tergite six with a row of strong, posteriorly curved setae. However, although many authors have considered Ormiophasia and Ormia as synonyms, Ormiophasia can be clearly distinguished by its darker body color, varying between brownish-yellow, brown and dark brown (Figs 9–14) ( Ormia species generally have a pale yellow body color); ocelli well developed (vestigial or missing in Ormia); one pair of presutural acrostichal setae (two to three pairs in Ormia); tegula of same color as body ( Ormia usually with tegula black, contrasting with body color); male wing without callosities on veins C and R 2+3 ( Ormia usually with callosities on C and R 2+3) (see Tavares 1962, 1965a, 1965b); male terminalia with apex of cerci broad, at least 1/3 of total width of cerci (Figs 38–42) (in Ormia the cerci are tapered apically, less than 1/3 of total width of cerci) (see Tavares 1962, 1965a, 1965b); and female terminalia with hypoproct bare, without setulae (Fig. 3B, setulose in Ormia). Additionally, Ormiophasia also differs from other Ormiini genera by having head hemispherical in profile, with genal length not exceeding 0.2 times length of head; face developed, at least 1.1 times width of facial ridge; facial ridge broad but not exceeding 2.9 times width of parafacial; and male terminalia with cerci completely fused.

Genus characterization. Both sexes. Head hemispherical in profile; oral axis shorter than antennal axis. Eye bare. Ocelli well developed. Antenna inserted in the middle of the eye, reaching halfway between lunule and lower facial margin; scape reduced, with row of marginal setulae; pedicel dorsally setulose, with one dorsal preapical seta; first flagellomere oblong; arista arising basally on dorsal surface, bare or weakly plumose, base pubescent and lightcolored, becoming tapered and darker distally. Fronto-orbital plate setulose to level of first anterior frontal seta. Parafacial bare. Facial ridge broad and subequal to facial length, setulose below vibrissa. Face slightly concave at vibrissal angle, with lower facial margin not visible in profile. Gena about 1/10 of head height, genal setae 4–5. Genal dilation setulose. Vibrissae strong and crossed, arising far above lower facial margin; supravibrissal setae 1–2, about 1/3 length of vibrissa; subvibrissal setae 3–4. Mouthparts well developed; clypeus globose; palpus developed, about 1.5 times length of prementum, clavate, covered with appressed setae from median area to apex; prementum setulose; labella padlike. Occiput covered with silver setulae, with row of short postocular setae; lower occipital area strongly concave.

Thorax densely covered with black setulae. Presternum bilobed (Fig. 1), each lobe with a pit on anterior surface. Basisternum bare, inflated; ventral surface grooved medially. Prosternal tympanal membrane corrugated, with corrugation equally distributed across entire membrane. Acrostichal setae 1+2–3; dorsocentral setae 2+3; intra-alar setae 1+2; supra-alar setae 1+2 (first postsutural seta weak, about 1/2 length of second postsutural seta). Postpronotal setae 3, with inner seta weaker. Notopleural setae 2, equal-sized. Postalar setae 2, equal-sized. Proepisternal setae 1, strong, upcurved, with 1–3 weaker setae near base. Katepisternal setae 2, divergent. Anepimeral seta 1. Katepimeron, katatergite, anatergite and postalar wall bare. Scutellar setae strong, convergent; basal pair subequal to subapical pair; lateral pair 2/3 length of subapical pair, closer to basal pair; apical pair absent; discal pair half length of subapical pair, more widely separated than subapical setae. Wing. Tegula setulose, with 1–2 inner marginal setae. Cells sc, c and r 1 with light yellow infuscation. Vein C ending just after vein M 1, at wing apex. Cell r 4+5 open; length of opening subequal to or shorter than crossvein r-m. Vein M 1 arched towards apex; bend of vein M angular or rounded, sometimes with a short stem. Legs. Fore coxa with an outer row of 3–4 anterior setae followed by 4–6 marginal setae. Fore tibia with 1 posterodorsal preapical seta, 2 posteroventral median setae, 1 preapical seta and 1 ventral preapical seta. Mid femur with 1 strong anteromedian seta and 2–3 strong posterodorsal preapical setae. Mid tibia with 1 strong anterodorsal postmedian seta, 1 apical seta, 1 posterodorsal apical seta, 2 posteromedian setae, 1 weak apical seta, 1 posteroventral apical seta, 1 weak ventral apical seta and 1 strong anteroventral apical seta. Hind femur with row of 3–4 posterodorsal basal setae and row of 11–12 weak posterodorsal setae from middle to apex. Hind tibia with 2 anterodorsal median setae, 1 preapical seta, 1 weak ventral apical seta, 2–3 weak anteroventral median setae and 1 weak apical seta. Pulvilli light yellow. Claws brown, with apex black.

Abdomen globose and widely connected with thorax, wider than long; densely covered with black setulae. Middorsal depression on syntergite 1+2 extending to hind margin of syntergite. Syntergite 1+2 and tergite 3 with one pair of marginal lateral setae. Tergite 4 with row of 8–10 marginal setae. Tergite 5 with row of 6–8 discal setae. Sternites visible, each with a weak row of marginal setae.

Male. Head holoptic (Figs 5–6). Ocellar triangle constricted, tubercle-shaped. Dorsal ommatidia larger than peripheral ones. Thorax. Lateral cervical sclerite with external lobe oblong (e.g., Fig. 15E). Basisternum less inflated than in female. Prosternal tympanal membrane reduced. Posterior spiracle with posterior lappet shaped as an operculum. Terminalia (Figs 3A, 38–42). Sternite 5 with posterior margin bilobed, setulose posteriorly. Epandrium with dorsal surface covered with strong, upcurved setae. Surstylus not fused with epandrium; strongly arcuate mediad; posterior surface with a slight distal, median ridge; articulation with bacilliform sclerite rounded; articulation with epandrium tapered and narrow; inner surface covered with microtrichia. Connection of bacilliform sclerite to hypandrium broad, becoming tapered towards surstylus. Anterior margin of hypandrium slightly longer than phallapodeme; concave; posterior area weakly sclerotized; hypandrial arms not fused with each other or with aedeagus. Basiphallus connected directly to distiphallus at a 90 o angle; distiphallus simple, smooth, with anterior surface grooved. Phallapodeme flat, slender. Ejaculatory apodeme narrow, with posterior apex slightly broader. Pregonites flat, bare, slightly concave, not fused with each other and not fused with hypandrium; anterior margin broad, reaching lower posterior margin of hypandrium. Postgonite long, bare, rod-like, with rounded apex, almost reaching connection between basiphallus and distiphallus. Cerci fused, with apex broad and at least 1/3 of total length of cerci; covered with setae.

Female. Head dichoptic (Figs 7–8). Ocellar triangle level with eyes, not protruding as a tubercle. Ommatidia all of the same size, not differentiated. One pair of inner vertical setae, strong and crossed, subequal to vibrissa. One pair of outer vertical setae, same size as inner vertical setae, divergent. One pair of upper orbital setae, divergent, half size of vertical setae. Two pairs of proclinate orbital setae. Thorax. Lateral cervical sclerite with external lobe extended and tapered at margin (e.g., Fig. 15F). Basisternum very inflated, much more than in male. Prosternal tympanal membrane broad. Posterior spiracle with posterior lappet stylus- or plume-shaped (except operculum-shaped in O. lanei). Terminalia (Fig. 3B). Tergite 6 broad, voluminous, setulose, separated into two equal-sized sclerites, bearing spiracles 6 and 7; inner margin with very strong, inward-curved setae. Tergite 7 absent. Tergite 8 developed, bare, separated into two equal-sized, concave sclerites. Epiproct (tergite 10), when present, represented by one pair of setae in membrane. Sternites 6, 7 and 8 each with a row of weak marginal setae. Hypoproct (sternite 10) bare; lateral surface with patch of very short, spiniform setae; ventral surface with longitudinal median row of short setae; posterior margin with row of slender, weak setae. Cerci free, not fused, with weak, slender marginal setae. Three equal-sized spermathechae, spherical, smooth.

Remarks. The first four species included in Ormiophasia were based mainly on female specimens, which probably caused most of the confusion about the validity of the genus. The male and female terminalia had not been studied and described at that time, but they include important diagnostic characters for the genus. Aldrich (1922) and Curran (1934) treated Ormiophasia as a junior synonym of Ormia, without justification. Only Malloch (1929) made some considerations based on characters such as the width of the female frontal vitta, which is narrow in Ormiophasia . Townsend (1919: 164) had already mentioned this character when describing Ormiophasia, and Sabrosky (1953: 172) also used it to differentiate Ormia and Ormiophasia in his key. Ormia punctata Robineau- Desvoidy, examined by Malloch, clearly has frontal vitta broader than in other Ormiophasia species, but there are also other species of Ormia with a narrower frontal vitta. Townsend did not know males of Ormiophasia for a long time after the original description (Townsend 1927: 223, “ ♂ desconhecido [unknown]”), until he included descrip- tions of male terminalia in his Manual of Myiology (Townsend 1936, 1938). These descriptions were superficial and incomplete, however, and relative to all Ormiini . In this context, the contributions and impact of the publications of Tavares (1962, 1964, 1965a, 1965b) on the taxonomy of New World Ormiini should be highlighted, since the male terminalia of a comprehensive number of species were described in detail for the first time. Furthermore, Tavares was the first to diagnose these genera based on male terminalia (Tavares 1965a: 14). However, he described his species based only on males, without associating them with their respective females. The association of females with males and the dissections of female terminalia made in the present study have provided additional and valuable characters to differentiate Ormiophasia and Ormia .

Some comments about Tavares’s holotypes are also warranted. The right wing, antenna and male terminalia of his type material were mounted on slides and linked with their respective specimens by a code (see Fig. 19E). However, during the Brazilian military government (1964–1985), an event known as the “Massacre de Manguinhos” took place (Jurberg 1993; Costa et al. 2008). This episode, which occurred on April 1st, 1970, greatly impacted the history of science in Instituto Oswaldo Cruz, Unidade Manguinhos, Rio de Janeiro, where Omar Tavares had worked and where his holotypes are deposited. The term “Massacre de Manguinhos” was coined by helmintologist Herman Lent (Jurberg 1993), who worked for the same institute. The “Manguinhos” entomological collection is one of the most important historic scientific collections in Brazil. During the “Massacre de Manguinhos”, scientists were politically persecuted and the scientific collection was dismantled and transferred to another building inside the institute. The relocation of the Instituto Oswaldo Cruz collection resulted in the loss of much of the material.Although Tavares’s holotypes are still in good condition, their respective slides were lost, probably during this episode. Only the slides of the male terminalia of the paratypes, housed in MZSP and USNM, are available for study.

The type material of all nominal species of Ormiophasia was examined during this study. All of Séguy’s and Townsend’s holotypes are in good conditions of preservation (Figs 16, 24, 29, 30). The type specimens of Tavares (Figs 18, 20, 22, 25, 27) have damaged abdomens due to their dissection. The examination of type material allowed for a reliable identification of the material included in this study. Furthermore, as Sabrosky (1953: 173) remarked, Ormiophasia specimens exhibit a significant variation in body coloration, which ranges from brownish-yellow to almost black. The color and pruinosity of the head and thorax, and the male cerci and surstylus, were the most important characters supporting the description of the eight new species.

So far, there are no known host records for any Ormiophasia species.

Key to species of Ormiophasia

Although Ormiophasia (and Ormiini) species are sexually dimorphic, the key provided below aims to identify both male and female specimens (except for O. obscura, O. buoculus sp. nov. and O. townsendi sp. nov., of which only males are known); when necessary, characters exclusive to one sex are included in couplets. The identification of female specimens, however, has to be done carefully, since the females of several species are very similar (e.g., O. costalimai and O. inflata or O. causeyi and O. chapulini sp. nov.). The male terminalia are the most reliable source of information allowing to confirm species’ identity.

Tavares (1964) provided the first key to Ormiophasia species, but some characters used in his key fail when large series are examined, due to variation (e.g., of body color or of number of setae). Furthermore, there are some errors in the key, which are commented on in the Remarks under the respective species in the present paper. Some important characters used by Tavares are maintained here for some species (e.g., those of the male terminalia of O. lanei). Séguy (1926b) also provided a key to his monotypic genera Plagiatormia, Pseudoneoptera and Pseudormia, based on the length of the opening of cell r 4+5, the shape of the bend of vein M, the length of the antenna and the shape of the face. Although these characters can be seen in Séguy’s holotypes, they are variable in the additional material examined (except in O. obscura, so far known only from the holotype).

1 Basicosta broad, twice width of tegula (Fig. 4B); female wing with strong brown infuscation around all veins except A 1 +CuA 2 (male wing with weak infuscation at the tip of vein R 2+3 and around veins M 1 and dm-cu, see Figs 33G, 34F), and with veins R 2+3, R 4+5, M 1 and CuA 1 thickened (Figs 33H, 34G); thorax dark brown or brownish-yellow.......................... 2

- Basicosta subequal in width to tegula (Fig. 4A); female and male wings hyaline (Fig. 15G) or infuscated only around veins R 1, R 2+3, M 1 and dm-cu (Fig. 28G), but without thickened veins; thorax brownish-yellow, brown or dark brown.............. 3

2 Thorax and abdomen dark brown (Figs 10C, 14B); male wing with section of vein M between crossvein dm-cu and M 1 slightly curved (Fig. 33G); apex of male cerci long and about 2/5 length of cerci in posterior view (Fig. 41A) (North Brazil, Costa Rica, French Guiana and Venezuela).......................................................... O. crassivena sp. nov.

- Thorax brownish-yellow, contrasting with dark brown abdomen (Figs 10D, 14C); male wing with section of vein M between crossvein dm-cu and M 1 straight (Fig. 34G); apex of male cerci short and about 1/3 length of cerci in posterior view (Fig. 41B) (South and Southeast Brazil)......................................................... O. manguinhos sp. nov.

3 Wing with strong brown infuscation around veins R 1 and R 2+3 (Fig. 28G) and weak brown infuscation around veins M 1 and dm-cu; thorax and abdomen dark brown or brownish-yellow................................................... 4

- Wing hyaline ( O. cruzi may have a very weak brown infuscation around veins M 1 and dm-cu, see Fig. 21G); thorax and abdomen dark brown, brown or brownish-yellow................................................................ 7

4 Thorax and abdomen dark brown; fronto-orbital plate and parafacial gray with silver pruinosity....................... 5

- Thorax and abdomen brown or brownish-yellow; fronto-orbital plate and parafacial brownish-yellow with yellow pruinosity 6

5 Male head with dorsal ommatidia larger than ocelli (Fig. 6G); ocellar triangle very constricted, not visible in profile (Venezuela)................................................................................ O. buoculus sp. nov .

- Male head with dorsal ommatidia not as large as above, about 0.6 times smaller than ocelli (Fig. 5G); ocellar triangle constricted, visible as a tubercle in profile (North Brazil, Ecuador, French Guiana, Guyana, Panama and Peru)................................................................................................... O. morardi (Séguy)

6 Male head with dorsal ommatidia larger than ocelli (Fig. 6H); ocellar triangle very constricted, not visible in profile; thorax and abdomen brownish-yellow; surstylus slender, with outer posterior surface covered with weak setae in upper two-thirds (Fig. 42C); apex of male cerci about 1/3 width of cerci in posterior view ( North Brazil)................. O. townsendi sp. nov .

- Male head with dorsal ommatidia not as large as above, subequal to ocelli size (Fig. 6B); ocellar triangle constricted, but visible as a tubercle in profile; thorax brown, abdomen brownish-yellow; surstylus stout, with outer posterior surface entirely covered with strong setae (Fig. 40C); apex of male cerci about 1/2 width of cerci in posterior view (Bolivia and Peru)................................................................................................... O. seguyi sp. nov.

7 Fronto-orbital plate with silver pruinosity clearly contrasting with yellow pruinosity of lower parts of head (both sexes, but more obvious in females, see Fig. 7F); thorax and abdomen brown; surstylus slender in posterior view.................. 8

- Head pruinosity entirely silver or yellow, without contrast between fronto-orbital plate and rest of head; thorax and abdomen brown, dark brown or brownish-yellow; surstylus stout in posterior view......................................... 9

8 Clypeus darker than frontoclypeal membrane (Figs 5F, 7F); female posterior spiracle with posterior lappet shaped as an operculum; surstylus strongly inflected inward in posterior view, outer posterior surface covered with strong setae in upper two-thirds (Fig. 39C); apex of male cerci subquadrate and abruptly constricted in posterior view (Southeast Brazil)... O. lanei Tavares

- Clypeus of same color as frontoclypeal membrane (Fig. 5D); female posterior spiracle with posterior lappet plume-shaped; surstylus not inflected inward as above, covered with weak setae in upper two-thirds (Fig. 39A); apex of male cerci subquadrate and gradually constricted in posterior view (South and Southeast Brazil, Panama and Paraguay).......... O. cruzi Tavares

9 Clypeus darker than frontoclypeal membrane (Fig. 6E); thorax and abdomen dark brown........................... 10

- Clypeus of same color as frontoclypeal membrane (Fig. 5A); thorax and abdomen brown or brownish-yellow ( O. guimaraesi sp. nov. has a dark brown scutum, but the rest of the thorax is brown, see Figs 10A, 12A)........................... 12

10 Thorax and abdomen entirely dark brown (Figs 10E, 14D); anteroventral epandrial process extending well beyond ventral epandrial margin (Fig. 41C); dorsal surface of epandrium with posterior margin at same level as anterior margin (Fig. 41C, lateral view); apex of male cerci about 1/2 width of cerci in posterior view, abruptly constricted (Fig. 41C) (Colombia, Costa Rica, Panama and Venezuela)................................................................. O. tavaresi sp. nov.

- Thorax and abdomen dark brown, with postpronotal lobe brown (Figs 9B, 13B); anteroventral epandrial process continuous with ventral epandrial margin (Fig. 42A); dorsal surface of epandrium with posterior margin higher than anterior margin (Fig. 42A, lateral view); apex of male cerci 1/3 (Fig. 38B) or more than 1/2 width of cerci in posterior view (Fig. 42A), gradually constricted.......................................................................................... 11

11 Arista weakly plumose (Fig. 17 A–B); apex of male cerci narrow, 1/3 width of cerci and rounded in posterior view (Fig. 38B) (North Brazil, Guyana, Peru, Trinidad and Tobago and Venezuela)............................... O. causeyi Tavares

- Arista bare (Fig. 36 A–B); apex of male cerci broad, more than 1/2 width of cerci and subquadrate in posterior view (Fig. 42A) (Costa Rica and Mexico)............................................................... O. chapulini sp. nov.

12 Thorax with scutum dark brown and lateral surface brown (Figs 10A, 12A); apex of male cerci broad, subrectangular in posterior view (Fig. 40B); surstylus stout in posterior view, with outer posterior surface densely covered with strong setae and inner posterior surface covered with short setae medially (Colombia and Costa Rica)................... O. guimaraesi sp. nov.

- Thorax entirely brown or brownish-yellow; apex of male cerci broad or narrow, rounded in posterior view; surstylus not as stout as above (Figs 38A, 38C, 39B), with inner posterior surface bare.............................................. 13

13 Thorax and abdomen entirely brown..................................................................... 14

- Thorax and abdomen entirely brownish-yellow............................................................. 15

14 Ocellar triangle bare, without setulae (Fig. 15 C–D); head yellow-pruinose (Figs 5A, 7A) (Panama).... O. busckii Townsend

- Ocellar triangle setulose (Fig. 30A); head silver-pruinose (Fig. 5H) (Argentina)..................... O. obscura (Séguy)

15 Female head subtrapezoidal in frontal view, with gena about 0.15 times length of head (Figs 7E, 23B); apex of male cerci broad (Fig. 39B), 3/5 width of cerci and rounded in posterior view, and gradually constricted ( North Brazil, French Guiana, Trinidad and Tobago and Venezuela)............................................................... O. inflata (Séguy)

- Female head elliptic in frontal view, with gena about 0.12 times length of head (Figs 7C, 19B); apex of male cerci narrow (Fig. 38C), 1/3 width of cerci and rounded in posterior view, and abruptly constricted (North Brazil, Colombia, Ecuador, Guyana, Peru, Suriname and Venezuela)......................................................... O. costalimai Tavares