Taxonomic revision of the Neotropical stalk-eyed fly Plagiocephalus Wiedemann (Diptera, Ulidiidae, Ulidiinae)

ABSTRACT

Plagiocephalus is a genus composed of three species mainly recognized by the males with stalked-eyes. Plagiocephalus lobularis is distributed from Northeastern Brazil to North of Argentina, P. latifrons is distributed from Mexico to Bolivia, and P. intermedius occurs in Costa Rica. We review the species of the genus, providing new diagnostic characters, an identification key, and new information on the terminalia of Plagiocephalus. Also, we update the geographic distribution of the species of the genus.

Keywords:
Eyestalks; Picture-winged flies; Taxonomy; Tephritoidea

Introduction

Among the genera of the Neotropical tribe of ulidiids, Pterocallini, Plagiocephalus Wiedemann is one of the most unusual. The male of all three described species, P. lobularis (Wiedemann), P. latifrons (Hendel) and P. intermedius Kameneva, present the eyes stalked. In P. latifrons, the eyestalks can reach great proportions, extending until five times the length of the body (Grimaldi and Engel, 2005). The species of Plagiocephalus can also be recognized by the brownish body background colour, with grey and yellow microtrichia on the thorax and abdomen, pattern of dark bands on the wing, vein R₁ with setae on the apical half, vein R₂₊₃ almost straight and cell cup with a very short posteroapical lobe (Kameneva, 2004b).

Wiedemann (1830a) described the type species of the genus, Plagiocephalus lobularis, originally belonging to Achias Fabricius (Platystomatidae), another genus of stalk-eyed fly. In the same year, Wiedemann (1830b) described Plagiocephalus, considering A. lobularis as belonging to this genus. Hendel (1909a) described the second Plagiocephalus species, P. latifrons, originally on the genus Terpnomyia Hendel. Hendel (1936) created a new genus, Ophryoterpnomyia, to allocate T. latifrons, considering differences in the head while comparing it to other species of Terpnomyia. However, Ophryoterpnomyia was synonymized with Plagiocephalus by Carrera (1950). Kameneva (2004b) described P. intermedius, the last described species of Plagiocephalus.

Carrera (1950), Steyskal (1963) and Kameneva (2004b) revised the genus, Kameneva (2004b) presented a key to the species of the genus, and only Steyskal (1963) presented an illustration of a terminalia of one of the species, a male terminalia of P. latifrons. In the present paper, we provide a general description with new characters from the terminalia of Plagiocephalus. In addition, we update the geographical distribution of the species of the genus.

Material and methods

The description of the labels of the type material was obtained from Kameneva (2004b). The type material of Plagiocephalus was examined by photos. The pictures of the types were taken from the material deposited in the following colletions (curators are between parentheses): INBio: Instituto Nacional de Biodiversidad, Santo Domingo, Heredia, Costa Rica (sent by Valery Korneyev (Schmalhausen Institute of Zoology, Kyiv, Ukraine)); INTA: Instituto Nacional de Tecnología Agropecuaria, Buenos Aires (Esteban Daniel Saini), Argentina; MTD: Museum für Tierkunde, Dresden, Germany (Uwe Kallweit); MZH: Finnish Museum of Natural History, Helsinki, Finland (Pekka Vilkamaa); NHMW: Naturhistorisches Museum Wien, Vienna, Austria (Peter Sehnal); and ZMUC: Zoological Museum University of Copenhagen, Copenhagen, Denmark (Thomas Pape). Non-type material was provided by the following collections: CEUA: Colección Entomológica, Universidad de Antioquia, Medellín, Colombia (Marta Wolff); CZMA: Coleção Zoológica do Maranhão, Universidade Estadual do Maranhão, Caxias, Maranhão, Brazil (Francisco Limeira-de-Oliveira); DZUP: Coleção Entomológica Padre Jesus Santiago Moure, Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil (Claudio J. B. de Carvalho); EMUS: Utah State University, Entomological Museum, Logan, Utah, United States (Wilford J. Hanson (deceased)) (now donated to the LACM); INPA: Instituto Nacional de Pesquisas da Amazônia, Coleção Sistemática de Entomologia, Manaus, Amazonas, Brazil (Márcio Oliveira); LACM: Natural History Museum of Los Angeles County, Los Angeles, California, United States (Brian Brown); MNRJ: Museu Nacional da Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil (Márcia S. Couri); MZSP: Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (Carlos J. E. Lamas); NHMUK: National History Museum of London, London, England (Daniel Whitmore); and USNM: United States National Museum of Natural History, Department of Entomology, Washington, District of Columbia, United States (Allen L. Norrbom). The locality cited as Seara (Santa Catarina, Brazil) in the Material examined section of Plagiocephalus lobularis is the city correspondent to the district of Nova Teutônia. New records of geographic distribution are marked with an asterisk in the Distribution section of each species.

General terminology is based on Cumming and Wood (2009). The terminology of wing pattern and male terminalia follow White et al. (1999). The "anterior apical band" termed by White et al. (1999) is here named as "apical band". The female terminalia structures follow Norrbom and Kim (1988). The pictures presented in this work were taken from non-type specimens of the DZUP and USNM collections, with exception of the Plagiocephalus intermedius male and female wings (Fig. 3E, F), which were taken, respectively, from the holotype and a paratype. The photographs of the specimens and terminalia were taken with an Auto-Montage Imaging System^® Leica DFC 500 in the Taxonline (UFPR), and drawings were made with a microscope Zeiss Standard 20 coupled with a camera lucida. The map was constructed on QGIS 2.18.14 using a Latin America political shape and a satellite raster from the Google plugin installed on the QGIS.

Taxonomy

Plagiocephalus Wiedemann, 1830b

(Figs. 1–5)

Fig. 1
A–C. Plagiocephalus lobularis, male: A. Head in frontal view; B. Body in dorsal view; C. Body in lateral view. D–F. Plagiocephalus latifrons, male: D. Head in frontal view; E. Body in dorsal view; F. Body in lateral view.

Fig. 2
A–C. Plagiocephalus lobularis, female: A. Head in frontal view; B. Body in dorsal view; C. Body in lateral view. D–F. Plagiocephalus latifrons, female: D. Head in frontal view; E. Body in dorsal view; F. Body in lateral view. G–I. Plagiocephalus intermedius, female: G. Head in frontal view; H. Body in dorsal view; I. Body in lateral view.

Fig. 3
A–B. Plagiocephalus lobularis: A. Male wing; B. Female wing. C–D. Plagiocephalus latifrons: C. Male wing; D. Female wing. E–F. Plagiocephalus intermedius: E. Male wing; F. Female wing. Abbreviations: ab: apical band; db: discal band; sab: subapical band; rmb: radial-medial band.

Fig. 4
A–D. General morphology of the male terminalia of Plagiocephalus: A. Ejaculatory apodeme in lateral view (P. lobularis); B. Male terminalia in lateral view (P. lobularis); C. Epandrium in posterior view (P. latifrons); D. Hypandrium, phallapodeme, phallapodemic arms, basiphallus and distiphallus (P. latifrons). E–F. General morphology of the female terminalia of Plagiocephalus: E. Female terminalia in dorsal view (P. lobularis); F. Spermathecae (P. lobularis). Abbreviations: basiph: basiphallus; cerc: cerci; distph: distiphallus; ej apod: ejaculatory apodeme; epand: epandrium; ev memb: eversible membrane; hypd: hypandrium; lat sur: lateral surstylus; med sur: medial surstylus; ovscp: oviscape; phapod: phallapodeme; phapod arm: phallapodemic arm; prens: prensiseta; sg 8: segment 8; spmth: spermathecae; tae: taeniae.

Fig. 5
A–D. Plagiocephalus, male terminalia: A. Epandrium of P. lobularis; B. Epandrium of P. latifrons; C. Hypandrium, basi- and distiphallus of P. lobularis; D. Hypandrium, basi- and distiphallus of P. latifrons. Abbreviations: basiph: basiphallus; cerc: cercus; distph: distiphallus; epand: epandrium; hypd: hypandrium; lat sur: lateral surstylus; med sur: medial surstylus; phapod: phallapodeme; phapod arm: phallapodemic arm; prens: prensiseta.

PlagiocephalusWiedemann, 1830b: 12, 15; Westwood, 1849: 235; Osten-Sacken, 1881: 478; Hendel, 1911: 4, 52; Cresson, 1923: 258; Frey, 1926: 47; Carrera, 1950: 261; Aczél, 1951: 421; Steyskal, 1963: 511; 1964: 400; 1968: 54.21; Kameneva, 2004b: 15. Plagiocephala: Macquart, 1843: 213; Loew, 1873: 26; Hendel, 1909b: 47 (unjustified emendation); Plagiocephalas: Frey, 1926: 47 (error).

Stylophthalmyia Frey, 1926. Type species: Stylophthalmyia fascipennisFrey, 1926 (by original designation).

Ophryoterpnomyia Hendel, 1936. Type species: Terpnomyia latifronsHendel, 1909a (by original designation).

Paragoniaeola Blanchard, 1938a. Type species: Paragoniaeola tanycephalaBlanchard, 1938a (by original designation).

Eupterocerina Blanchard, 1938b. Type species: Eupterocerina conjuctaBlanchard, 1938b (by original designation).

Willineria Blanchard, 1951. Type species: Willineria orfilaiBlanchard, 1951 (by original designation).

Type species: Achias lobularisWiedemann, 1830a(by monotypy).

Diagnosis. Male with stalked eyes, frons wider than long (moderately wide in female). Thorax and abdomen brownish with pattern of yellowish-grey microtrichia; one supra-alar and two dorsocentral setae. Vein R₁ setulose on apical half; vein R₂₊₃ almost straight; crossvein r-m at apical 2/5 of d-m cell; cell cup with short posterior lobe; wing hyaline with four main dark-brown bands: discal band, radial-medial band, subapical band and apical band. Abdominal tergites 3–5 subequal in both sexes. Female with tergite 6 short, hidden under tergite 5, with several short marginal setulae; sternites 4–6 without apodemes. Male terminalia with distiphallus microtrichose at apical 2/3, elongated; ejaculatory apodeme with apical portion at least as long as its fan-like portion; epandrium elongate oval and setulose; medial surstylus bearing two subequal prensisetae with small denticles on inner surface. Female with oviscape brown, setulose, longer than the four preceding tergites together; segment 8 long; cerci oval, slightly turned upwards, dorsally with one basal, one subapical and two apical pairs of setae; three rounded, brown and smooth spermathecae.

Redescription.Head: Male (Fig. 1A, D): Wider than thorax; in frontal view wider than high (at least five times the width of the eye); eyes stalked, entirely microtrichose; eye higher than gena; inner vertical setae parallel; outer vertical, postocellar and ocellar setae divergent; orbital setae reclinate; ocellar triangle brown to black; dorsolateral and anterior portions of occiput brown; frons gold, with darker spot, wider than long and with sparsely distributed black setulae; gena with black setulae and developed vibrissa; antennal groove absent; scape, pedicel and first flagellomere yellow to gold; first flagellomere oval, about 2.5 times the length of pedicel; arista brown with very short pubescence, about 3.5 times the length of first flagellomere; clypeus with white microtrichia; palpus not extending beyond anterior oral margin, with black setulae on apex; proboscis capitate, covered with setulae. Female (Fig. 2A, D, G): Similar to male, except: in frontal view wider than high (less than four times the width of eye); eyes not stalked; frons with two brown to black spots anterior to ocellar triangle; face convex, with black spot between antennae and two transversal brown spots above clypeus; gena with C-shaped spot of brown microtrichia.

Thorax (Figs. 1B, E, 2B, E, H): Brownish black with patterns of yellow-grey microtrichia; scutum with brown microtrichose pattern; postpronotal lobe, scutellum, subscutellum and mediotergite mostly shiny brown; scutellum with a yellow-grey microtrichose stripe reaching the subscutellum; one postsutural supra-alar seta, one postalar seta, one intra-alar seta, two dorsocentral setae, and two scutellar setae; anepisternum setulose and with up to 10 setae; katepisternum with one seta.

Wing (Fig. 3A–F): Humeral break distinct; vein C between veins Sc and R₁ almost straight; cell c with brown spots; pterostigma unmodified, five to seven times as long as wide; vein R₁ setulose on apical half; vein R₂₊₃ bare and almost straight; vein R₄₊₅ bare, ending in the middle of wing apex; crossvein r-m at apical 2/5 of cell d-m; cell cup with very short posterior lobe; vein A₁ + CuA₂ reaching the posterior margin; wing hyaline, with pattern of four main bands; discal band from middle of cell sc inclined to posterior margin close to vein CuA₁; radial-medial band from apex of cell sc reaching the crossvein r-m; subapical band from vein CuA₁ crossing the wing to the 5/6 of vein C; apical band from final portion of vein C bypassing the apex. Upper calypter slightly longer than lower calypter, white and with white fringe. Halter white to yellow.

Legs (Figs. 1C, F, 2C, F, I): Unmodified, moderately setulose and with brown or yellow colouration.

Abdomen: Dark brown, with areas of yellow-grey and brown microtrichia at anterior and posterior margins of tergites; uniformly setulose; male with tergite 5 without microtrichia; female tergite 5 shorter than the 4th, and tergite 6 smaller, hidden under the 5th, without microtrichias and with 4–5 short marginal setulae; sternites 4–6 without apodemes.

Terminalia: Male (Figs. 4A–D, 5A–D): Ejaculatory apodeme with apical portion at least as long as its fan-like portion; epandrium elongate oval and setulose; medial surstylus "V" shaped, with each apex connected to lateral surstylus; two subequal prensisetae with small denticles on the inner surface, positioned subapically on medial surstylus; lateral surstylus with anterior lobe longer than the posterior; basiphallus D-shaped, connected to phallapodeme; phallapodeme Y-shaped, with apex slightly surpassing the hypandrium; phallapodemic arms connected to hypandrium, with few small setulae; phallus with microsetulae at the apical 2/3, elongated and with no glans-like structures on apex. Female (Fig. 4E, F): Oviscape brown, setulose, longer than the four preceding tergites together; segment 8 long; cerci oval, slightly turned upwards, and dorsally with one basal, one subapical and two apical pairs of setae; three spermathecae rounded, brown, with smooth surface.

Observation. The male terminalia of P. intermedius could not be analyzed, because no male specimens were found in the material provided by the collections that supported our study. The female terminalia of all three species of Plagiocephalus were analyzed.

Distribution. Neotropical (Mexico, Belize, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Trinidad, Venezuela, Colombia, Brazil, Ecuador, Peru, Bolivia, Paraguay and Argentina) (Fig. 6).

Fig. 6
Distribution map of Plagiocephalus with Costa Rica detached. Circles show distribution records from the literature. Stars show new distribution records. Yellow: P. intermedius; Red: P. latifrons; Light blue: P. lobularis.

Identification key to the species of Plagiocephalus

• 1. – Wing with radial-medial band well connected with discal band (Fig. 3A, B); male wing with three lobes on the hind margin (Fig. 3A); male face blackish (Fig. 1A)........................................................P. lobularis

• 1' – Wing with radial-medial band not connected with discal band (Fig. 3C–E); male wing without lobes on the hind margin (Fig. 3C, E); male face yellowish white (Fig. 1D).............................................................. 2

• 2. – Wing with radial-medial band with similar width along the base and apex (Fig. 3C, D); crossvein r-m almost at the level of apex of vein R₁ (Fig. 3C, D); female parafacialia black (Fig. 2D)......................................P. latifrons

• 2' – Wing with radial-medial band with base much wider than apex (Fig. 3E, F); crossvein r-m located distinctly before the apex of vein R₁ (Fig. 3E, F); female parafacialia yellow (Fig. 2G).................................. P. intermedius

Plagiocephalus lobularis (Wiedemann, 1830a)

(Figs. 1A–C, 2A–C, 3A, B, 4A, B, E, F, 5A, C)

Plagiocephalus lobularis (Wiedemann, 1830a): Wiedemann, 1830b: 15; Macquart, 1843: 213 (Plagiocephala lobularis); Hendel, 1909b: 47; Westwood, 1849: 235; Frey, 1926: 47; Carrera, 1950: 262, 265 (Plagiocephalus latifrons (misidentification; see Steyskal, 1963); Aczél, 1951: 422; Hennig, 1952: 616; Steyskal, 1963: 512, 1968: 54.21; Kameneva, 2004b: 16.

Achias lobularisWiedemann, 1830a: 555 (Lectotype male, ZMUC; Type locality: Brasilia (Brazil)) (examined by photographs); Macquart, 1835: 260.

Paragoniaeola tanycephalaBlanchard, 1938a: 371 (Holotype male, INTA; Type locality: Argentina: Misiones) (synonymized by Aczél, 1951: 399) (examined by photographs).

Eupterocerina conjunctaBlanchard, 1938b: 91 (Holotype female, unknown collection; Type locality: Argentina: Misiones, Puerto Bemberg) (synonymized by Steyskal, 1968: 54.21).

Ophryoterpnomyia zikaniCapoor, 1954: 205 (Holotype female, Instituto Oswaldo Cruz (no. 5787, 4541–4543); Type locality: Brazil: Rio de Janeiro, Itatiaia) (synonymized by Steyskal, 1963: 512).

Type material. Lectotype of Achias lobularis (designed by Kameneva, 2004b) (male): "P. lobularis / Wied. / Brazils / Lund", "Mus./Westerm.", "Syntype male / Achias lobularis / Wiedemann / 1830 / des. V. Korneyev / XII.2003", "Syntypus", "ZMUC 00025500".

Diagnosis.Plagiocephalus lobularis can be differentiated from P. latifrons and P. intermedius by the shortest eyestalks (1.42–3.94 mm) (Fig. 1A, B); female parafacialia yellow (Fig. 2A); radial–medial band well connected to discal band (Fig. 3A, B), and male wing with three posterior lobes (Fig. 3A). The species can also be distinguished by male frons with dark gold spot on anterior portion of ocellar triangle (Fig. 1B); male with purplish black spot from ventral side of the head up to face and part of the parafacialia (Fig. 1A); scape dark gold to brown (Figs. 1A, 2A); pedicel and first flagellomere gold, sometimes with darker marks (Figs. 1A, 2A); male palpus brown, and female palpus yellow; proboscis brown, with brown and yellow setulae. Wing with crossvein r-m located distinctly before apex of vein R₁ (Fig. 3A, B); cell cup yellow and, in males, closed by an almost straight vein (Fig. 3A); cells c and sc, and base of discal band with colouration darker than in the other bands (Fig. 3A, B); male discal and subapical bands forming a rounded angle (Fig. 3A). Legs brown, with tarsi lighter and fore coxa yellow; male fore femur with base yellow (Fig. 1C); female fore femur yellow, with a subbasal brown ring (Fig. 2C).

Measurements: Male: Body: 2.37–3.52 mm; Wing: 4.35–5.34 mm. Female: Body: 2.49–3.75 mm; Wing: 4.35–5.82 mm; Oviscape: 1.0–1.5 mm.

Material examined. ARGENTINA: Misiónes, Iguazú (25.6036 S, 54.5558 W), 04–10.x.1927, R. C & E. M. Shannon leg., 1 male (Det.: E. Kameneva, 2001) (USNM). BRAZIL: Ceará: Ibiapaba, Cachoeira Samambaia, 21.x.2011, Gomes & Duarte leg., 1 female (DZUP); Ubajara, Parque Nacional do Ubajara, Cachoeira do Cafundó (3.8369 S, 40.9097 W), 01–15.i.2013, F. Limeira-de-Oliveira & J. S. Pinto Júnior leg., 1 male (CZMA). Goiás: (Est. Goyaz), Campinas, i–1936, R. Spitz leg., 1 female (MNRJ) and 5 females (MZSP); Goianesia, ix.1969, H. Ebert leg., 2 females (MZSP). Mato Grosso do Sul: Aquidauana (20.4344 S, 55.6558 W), 15–26.x.2011, Lamas, Nihei & eq. leg., 1 female (MZSP); Maracaju, ii.1937, 1 female (USNM). Paraná: Antonina, Reserva Rio Cachoeira (25.3119 S, 48.6717 W), 50 m, 23–27.i.2017, A. C. Domahovski leg., 1 male and 1 female (DZUP); Curitiba, Jardim Botânico (25.4421 S, 49.2388 W), 05.iv.2015, O. Aguirre-Obando leg., 1 male (DZUP); idem, Universidade Federal do Paraná, Centro Politécnico, Mata Viva (25.4458 S, 49.2324 W), 921 m, 28.vii–11.viii.2015, L. Wendt leg., 1 male (DZUP); ibidem, 11–25.viii.2015, 2 females (DZUP); ibidem, 09–24.ix.2015, 1 female (DZUP); ibidem, 09–23.x.2015, 1 male (DZUP); ibidem, 18.xi–03.xii.2015, 1 female (DZUP); ibidem, 08–25.i.2016, 1 male and 2 females (DZUP); ibidem, 11–25.ii.2016, 1 male and 2 females (DZUP); ibidem, 26.ii–14.iii.2016, 2 males and 1 female (DZUP); ibidem, 14–28.iii.2016, 3 males and 2 females (DZUP); ibidem, 28.iii–13.iv.2016, 2 males and 2 females (DZUP); ibidem, 13–28.iv.2016, 2 males and 4 females (DZUP); ibidem, 25.viii–09.ix.2016, 1 female (DZUP); idem, 6–9.xii.2016, A. C. Vasconcelos leg., 1 female (DZUP); ibidem, 21–23.ii.2017, 1 male and 1 female (DZUP); ibidem, 28.iii–03.iv.2017, 1 male (DZUP); idem, 15.ix.2017, S. Silva leg., 3 females (DZUP); Paranaguá, Floresta Estadual do Palmito (25.5688 S, 48.5355 W), 16–17.vii.2014, Leviski, Siewert & Queiroz-Santos leg., 2 males (DZUP); São José dos Pinhais (25.6047 S, 49.1933 W), 897 m, xi.2016, A. C. Domahovski leg., 2 females (DZUP); ibidem, xii.2016, 1 female (DZUP). Pernambuco: Bonito, Cachoeira Véu da Noiva (8.5423 S, 35.715 W), 510 m, 25.iii.2015, F. Bravo leg., 1 male (DZUP). Rio de Janeiro: Angra, Japuíba "Japuhyba" (22.9949 S, 44.2920 W), i.1985, E. Dorio & T. Travasso leg., 1 female (MNRJ); Casimiro de Abreu, Reserva Biológica da União, Trilha Buracão (22.4240 S, 42.0391 W), 14.i–14.ii.2014, Eq. Col. Biota Diptera Fluminense leg., 1 female (MNRJ); Rio de Janeiro, x–xii.1937–1.1938, R. C. Shannon leg., 2 females (USNM); idem, ix.1938, 1 female (USNM); idem, x.1938, YelFevServ. MESBrazil, 9 females (Det.: G. Steyskal, 1962) (USNM); ibidem, i.1939, 1 female (USNM); idem, 08.xi.1933, H. Lopes Souza leg., "Terpnomyia latifrons", 1 female (MNRJ); idem, ix.1938, Serv. Febre Amarela MESBrazil, 1 female (USNM); idem, "Dist. Federal", x.1937, Serv. Febre Amarela MESBrazil, 1 male (USNM); idem, xii.1938, Serv. Febre Amarela MESBrazil, 1 female (USNM); ibidem, xiii.1938, 1 female (USNM); idem, ix.1938, R. C. Shannon leg., YelFevServ. MESBrazil, 1 female (USNM); Jacarepaguá, Repr. Rio Grande, iii.1968, M. Alvarenga leg., 1 female (MZSP); Nova Friburgo, Caledônia, 2219 m, ii.2001, E. & P. Grossi leg., 1 male (DZUP). idem, Sans Souci (22.2808 S, 42.5121 W), 06.i.2010, 2 males (DZUP). idem, Sítio Caturama, 1050 m, 30.xii.2008, P. Grossi leg., 1 male (DZUP); Petrópolis, Taquara (22.6185 S, 43.2301 W), 28.xii.197?, H. S. Lopes leg., 1 male (MZSP); ibidem, 13.xii.1970, 1 female (MZSP); ibidem, 28.xii.1970, 1 female (MZSP); ibidem, 01.i.1971, 1 female (MZSP); ibidem, 03.i.1971, 3 females (MZSP); ibidem, ?.i.1971, 1 female (MZSP); ibidem, 11.i.1971, 1 female (MZSP); ibidem, 14.i.1971, 1 male and 3 females (MZSP); ibidem, 06.ii.1971, 1 male and 2 females (MZSP); ibidem, 07.ii.1971, 1 male and 2 females (MZSP); ibidem, 14.ii.1972, 2 females (MZSP); ibidem, 15.ii.1972, 3 females (MZSP); ibidem, 11.vi.1972, 1 female (MZSP) and 7 females (MZSP). Santa Catarina: Florianópolis, vii.1960, Casemiro leg., 1 female (MZSP); Joinville, 27.i.1972, H. S. Lopes leg., 1 female (MZSP); Seara [=Nova Teutônia] (27.1833 S, 52.3833 W), 24.x.1936, Fritz Plaumann leg., Brit. Mus. 1936–256, A. E. Whittington (2004), 1 male (NHMUK 010862540); ibidem, 29.xi.1937, Brit. Mus. 1938–40, 2 females (NHMUK 010862538, NHMUK 010862539); ibidem, 28.iii.1938, Brit. Mus. 1938–312, 1 female (NHMUK 010862537); ibidem, ii.1967, 1 female (MZSP); ibidem, ii.1969, 1 female (MZSP); ibidem, x.1969, 2 females (MZSP); ibidem, xi.1970, 1 female (MZSP); ibidem, vii.1971, 1 female (MZSP). São Paulo: Andes, 1955, M. Carrera leg., 1 male and 2 females (MZSP); Barueri, ?.ii.1966, K. Lenko leg., 1 female (MZSP); Butantan, 02.vii.1979, H. S. Lopes leg., 1 female (MZSP); ibidem, 12.vii.1979, 1 female (MZSP); idem, Horta O. Cruz, 06.viii.1969, L.T.F. leg., 1 male (MZSP); ibidem, 08.viii.1979, 1 male (MZSP); ibidem, 11.viii.1971, 2 males and 3 females (MZSP); Cantareira, Chapadão (23.4142 S, 46.6000 W), viii.1946, Barreto leg., 1 female (MZSP); Rio Paraná, Porto Cabral, 01–25.iv.1944, Trav. Fo., Carrera & E. Dente leg., 1 male (MZSP); São José dos Campos, 15–22.viii.1997, Eurico R. de Paulo leg., 1 female (LACM); ibidem, 07–14.ix.1997, 1 female (LACM).

Distribution. Brazil (Ceará*, Pernambuco*, Goiás, Mato Grosso do Sul, Rio de Janeiro, São Paulo, Paraná* and Santa Catarina), Paraguay and Argentina (Fig. 6).

Comments.Blanchard (1938a, 1938b) deposited the holotypes of Paragoniaeola tanycephala and Eupterocerina conjuncta in his personal collection, and only the material of P. tanycephala could be tracked down (INTA).

Plagiocephalus latifrons (Hendel, 1909a)

(Figs. 1D-F, 2D–F, 3C, D, 4C, D, 5B, D)

Plagiocephalus latifrons(Hendel, 1909a): Aczél, 1951: 421; Steyskal, 1963: 512; 1964: 400; 1968: 54.21; Kameneva, 2004b: 18; Kameneva et al., 2017: 127.

Terpnomyia latifrons (Hendel, 1909a): 18 (Syntypes: 3 females, MTD; 1 female, NHMW; Type localities: Bolivia: Mapiri; and Peru: Urubambaflufs) (examined by photographs); Hendel, 1909b: 31.

Stylophthalmyia fascipennisFrey, 1926: 46 (Holotype male, MZH; Type locality: Guatemala: Barbereau) (synonymized by Steyskal, 1963: 511) (examined by photographs).

Ophryoterpnomyia latifrons: Hendel, 1936: 76 (synonymized by Carrera, 1950: 260).

Willineria orfilaiBlanchard, 1951: 32 (Holotype male, Museo de Ciencias Naturales de Salta; Type locality: Bolivia: Chapare, Yungas) (synonymized by Steyskal, 1964: 490).

Type material. Syntypes of Terpnomyia latifrons (4 females): "Bolivia – Mapiri / 21.I.03 / S. Carlos 800 m", "Terpnomyia latifrons / det. F. Hendel", "Cotypus"; "Bolivia – Mapiri / 5.III.03 / Sarampioni 700 m", "Terpnomyia latifrons / det. F. Hendel", "Cotypus"; "Peru – Urubambafl. / 10.IX.03 / Umahuankilia"; "Terpnomyia latifrons / det. F. Hendel", "Cotypus", "Staatl. Museum für / Tierkunde Dresden/Coll. W. Schnuse, 1911″ (MTD); "Peru – Urubambafl./13.IX.03/Umahuankilia", "Terpnomyia latifrons/det. F. Hendel", "Paratype", "Coll. Hendel" (NHMW).

Diagnosis.Plagiocephalus latifrons can be differentiated from P. lobularis and P. intermedius by the longest eyestalks (5.08–18.08 mm) (Fig. 1D, E); female parafacialia black (Fig. 2D); radial-medial band with base almost as narrow as the apex, at most barely touching the discal band (Fig. 3C, D). The species can also be distinguished by male frons with gold spot on anterior portion of ocellar triangle (Fig. 1E); male face yellowish white, with region close to antennae yellower (Fig. 1D); male scape, pedicel and first flagellomere entirely yellow (Fig. 1D), and female scape, pedicel and first flagellomere gold with apex sometimes darker (Fig. 2D); palpus yellow; male proboscis yellow, with yellow setulae, and female proboscis reddish yellow, with brown and yellow setulae. Male wing of normal outline, without posterior lobes (Fig. 3C); crossvein r-m almost at the level of apex of vein R₁ (Fig. 3C, D). Male fore and mid legs entirely yellow, and hind leg yellow to gold (Fig. 1F); female legs brown, with tarsi lighter, and fore femur yellowish on the apex (Fig. 2F).

Measurements: Male: Body: 2.55–3.75 mm; Wing: 4.05–6.8 mm. Female: Body: 2.85–4.00 mm; Wing: 4.75–6.48 mm; Oviscape: 1.05–1.5 mm.

Material examined. BELIZE: Stann Creek Valley, 06.iv.1979, B. Bowers leg., 4 males (USNM). BRAZIL: Acre: Cruzeiro do Sul, Rio Moa (7.6172 S, 72.7708 W), 19–28.xi.1996, J. A. Rafael, J. Vidal & R. L. Menezes leg., 1 female wing (INPA). Amapá: Serra do Navio, 13.x.1957, J. Lane leg., 1 female (MZSP). Amazonas: Barcelos, Rio Demeni Pirico (0.325 S, 62.7892 W), viii.2008, A. Silva & R. Machado leg., 1 female (INPA); Ipixuna, Rio Gregório, Com. Lago Grande (7.1699 S, 70.8195 W), 18–23.v.2011, J. A. Rafael, J. T. Câmara, R. F. Silva, A. Somavilla, C. Gonçalves leg., 1 female (INPA); idem, Rio Liberdade, Estirão da Preta (7.363 S, 71.8686 W), 11–15.v.2011, J. A. Rafael, J. T. Câmara, R. F. Silva, A. Somavilla, C. Gonçalves & A. Agudelo leg., 1 female (INPA); ibidem, J. A. Rafael, J. T. Câmara, R. F. Silva, A. Somavilla & C. Gonçalves leg., 2 females (INPA); Manaus, Campus Universitário, 23.vi.1979, J. A. Rafael leg., 1 female (INPA); idem, Reserva Ducke (2.9166 S, 59.9833 W), 20 m, 07–21.xii.1994, J. A. Rafael & J. Vidal leg., 1 female (INPA); idem, 09.viii.2000, J. Vidal & A. F. Oliveira leg., 1 male (INPA); ibidem, 12.ix.2000, 1 male (INPA); ibidem, 26.x.2000, 2 males (INPA); ibidem, 27.x.2000, 1 female (INPA); ibidem, 28.xi.2000, 1 male (INPA); ibidem, 24.v.2001, 1 male (INPA); idem, ix.2001, J. Vidal leg., 1 female (INPA); idem, ZF2 km-14 Torre (2.5892 S, 60.1153 W), 19–22.iii.2004, J. A. Rafael, C. S. Motta, F. F. Xavier F, A. Silva F & J. T. Câmara leg., 1 female (INPA); idem, 18–21.v.2004, J. A. Rafael, F. B. Baccaro, F. F. Xavier & A. Silva leg., 1 female (INPA); idem, 12–15.x.2004, J. A. Rafael, C. S. Motta, F. F. Xavier, A. Silva F & S. Trovisco leg., 1 female (INPA); idem, 17–21.viii.2017, J. A. Rafael, A. C. Vasconcelos, F. F. Xavier, T. Mahlmann, S. Lima & B. Oliveira leg., 1 male (INPA); Novo Aripuanã, Reserva Soka (5.2647 S, 60.1188 W), 28.iv–06.v.1999, R. L. Ferreira, R. A Rocha, J. Vidal & R. S. Leite leg., 2 females (INPA); idem, 17–25.viii.1999, J. Vidal & A. L. Henriques leg., 1 male and 4 females (INPA); Pq. N. Jau, Ig. Miracutu, Ig. do Gerlei (1.9500 S, 61.8167 W), 23–28.vii.1995, J. A. Rafael & J. Vidal leg., 1 female (INPA); Rio Javari, Retirão do Equador, x.1979, Alvarenga leg., 2 females (MZSP); São Gabriel da Cachoeira, Morro dos Seis Lagos, 800 m, 28.ix–6.x.1990, J. Vidal & J. A. Rafael leg., 1 male (INPA). Maranhão: Bom Jardim, REBIO-Res. Biol. Gorupi, 02–11.ix.2010, D. W. A. Marques, E. A. S. Barbosa, J. A. Silva & M. M. Abreu leg., 1 female (CZMA). Pará: Belém, APEG Forest, x.1969, T. H. G. Aitken leg., 1 female (USNM); Óbidos, Colônia São Tomé (1.8461 S, 55.0397 W), 01–11.ix.2001, J. A. Rafael & J. Vidal leg., 1 female (INPA); idem, Faz. Parujá (1.6225 S, 55.3872 W), 05–11.ix.2000, J. A. Rafael & J. F. Vidal leg., 1 male (INPA); idem, Sítio Curuó (1.7842 S, 55.1181 W), 29.viii–08.ix.2001, J. A. Rafael & J. Vidal leg., 1 female (INPA). Rondônia: 62 Km SE Ariquemes (10.2418 S, 62.5492 W), 17–24.iii.1989, W. J. Hanson leg., 4 females (LACM); idem, 15–22.iii.1991, W. Hanson & G. Bohart leg., 1 female (LACM); idem, 13–25.iv.1992, W. J. Hanson leg., 1 female (LACM); ibidem, 8–20.xi.1994, 5 females (LACM); ibidem, 7–18.xi.1995, 1 female (LACM); ibidem, 22–31.x.1997, 1 female (LACM); ibidem, 1–14.xi.1997, 1 female (LACM); Monte Negro, Fazenda Amorim (10.6683 S, 63.4833 W), 03–15.xii.2011, Amorim, Ament & Riccardi leg., 1 female (MZSP); Porto Velho, AHE Jirau, Rio Madeira (9.5981 S, 65.3667 W), 28.iii–08.iv.2011, R. R. Silva, E. Z. Albuquerque & eq. leg., 1 female (MZSP). Roraima: Ilha de Maracá, Rio Uraricoera, 19–24.vii.1987, J. A. Rafael & L. S. Aquino leg., 1 female (Det.: A. Norrbom, 1990) (INPA); Pacaraima, 5–8.iii.1988, Eq. J. A. Rafael leg., 1 female (INPA). BOLIVIA: Sta Cruz, Buena Vista, 21.ii.1999, F. D. Parker leg., 1 female (LACM). COLOMBIA: Antioquia: Mpio la Pintada, Farailon La Paz, 16.ii.2007, N. Uribe leg., 1 male (CEUA); idem, Sán Jerónimo, Parque los Tamarindos, 425 m, 24–27.iv.2006, A. Velez leg., 1 male (CEUA 38267); idem, San Roque vrd, El Topacio, VSR Rez Bosque F1, v.2013, 1 male (CEUA); Ma, Santa Marta, Puerto Mosquito Rva La Iguana Verde, bosque VSR (11.1852 N, 74.1769 W), 09.xi.2012, C. Valverde leg., 1 male (CEUA); ibidem, 10.xi.2012, 1 male (CEUA). Santander: Cimitarra, Ciénaga de Cachimberos, Hacienda San Miguel, 05–8.x.2001, M. Castaño & M. Velez leg., 2 males (CEUA 38268); idem, Hacienda El Bosque, Campamento Ecuador, Bosque Ecuador, 09–12.x.2001, 1 male (CEUA 38133); idem, Vd. Primavera, 05–8.v.2001, M. Castaño & M. Velez leg., 3 males (CEUA 38269); idem, Vereda Primavera Km 4, Hacienda El Bosque, Campamento Santa Isabel, 01–4.x.2001, 1 male (CEUA 38270); San J. Girón, Prof. Sogamoso Denso Tierra, 603 m, 10–11.v.2015, Y. Correa leg., 1 male (CEUA). COSTA RICA: Alajuela: 2 Km S Pital, 05–28.ix.1988, F. D. Parker leg., 1 female (LACM); 20 Km S Upala, 14–17.viii.1990, F. D. Parker leg., 1 female (EMUS); ibidem, 21–23.viii.1990, 1 female (EMUS); ibidem, 28–30.viii.1990, 2 females (EMUS); ibidem, 01–10.v.1990, 1 female (LACM); ibidem, vi.1990, 2 females (LACM); ibidem, 15.vii.1990, 1 female (LACM); ibidem, 29.vii.1990, 1 female (LACM); ibidem, 01.viii.1990, 1 male (LACM); ibidem, 07–09.viii.1990, 1 female (LACM); ibidem, 21–23.viii.1990, 2 females (LACM); ibidem, 28–30.viii.1990, 1 female (LACM); ibidem, 10–13.ix.1990, 1 female (LACM); ibidem, 16–25.ix.1990, 1 female (LACM); ibidem, 27.ix–18.x.1990, 1 male and 2 females (LACM); ibidem, 1–5.x.1990, 1 female (LACM); ibidem, 16.x.1990, 3 females (LACM); ibidem, 23.x.1990, 4 females (LACM); ibidem, 26–30.x.1990, 2 females (LACM); ibidem, 28.x.1990, 4 females (LACM); ibidem, 30.x.1990, 3 females (LACM); ibidem, 01.xi.1990, 7 females (LACM); ibidem, 01–20.xi.1990, 2 females (LACM); ibidem, 06.xi.1990, 5 females (LACM); ibidem, 8.xi.1990, 11 females (LACM); ibidem, 13.xi.1990, 7 females (LACM); ibidem, 20.xi.1990, 3 females (LACM); ibidem, 29.xi.1990, 3 females (LACM); ibidem, 06.xii.1990, 4 females (LACM); ibidem, 11.xii.1990, 3 females (LACM); ibidem, 13.xii.1990, 1 female (LACM); ibidem, 13.xii.1990–09.i.1991, 2 males (LACM); ibidem, 06.i.1991, 1 male (LACM); ibidem, 20.i–12.ii.1991, 1 male (LACM); ibidem, 29.i.1991, 2 females (LACM); ibidem, 05.ii.1991, 2 females (LACM); ibidem, 07.ii.1991, 1 female (LACM); ibidem, 12.ii–05.iii.1991, 1 female (LACM); ibidem, 3–9.iii.1991, 1 male and 1 female (LACM); ibidem, 05–18.viii.1991, 1 male (LACM); ibidem, 10–19.iii.1991, 3 females (LACM); ibidem, 12.iii.1991, 2 females (LACM); ibidem, 18–26.iii.1991, 1 female (LACM); ibidem, 20–26.iii.1991, 2 females (LACM); ibidem, 27–31.iii.1991, 5 females (LACM); ibidem, 1–10.iv.1991, 6 females (LACM); ibidem, 11–20.iv.1991, 2 females (LACM); ibidem, 12–30.iv.1991, 1 female (LACM); ibidem, 1–9.v.1991, 1 female (LACM); ibidem, 10–21.v.1991, 2 females (LACM); ibidem, 1–11.vi.1991, 1 female (LACM); ibidem, 03.vi.1991, 1 female (LACM); ibidem, 07.vi.1991, 1 female (LACM); ibidem, 21.vi.1991, 1 female (LACM); ibidem, 1–15.vii.1991, 1 female (LACM); ibidem, 16–24.vii.1991, 4 females (LACM); ibidem, 21–31.vii.1991, 2 females (LACM); ibidem, 1–10.x.1991, 1 female (LACM); ibidem, 11–21.x.1991, 1 female (LACM); ibidem, 22–31.x.1991, 1 female (LACM); Golfito: 22.vii.1957, Truxal & Menke leg., 2 females (Det.: Korneyev & Kameneva, 2001, Det.: Kameneva, 2001) (LACM); ibidem, 24.vii.1957, 1 male (LACM). Guanacaste: S Cañas, 11–15.iii.1989, F. D. Parker leg., 1 male (LACM); 14 Km S Cañas, 14–16.x.1989, F. D. Parker leg., 1 male (LACM); ibidem, 15–25.vii.1990, 1 female (LACM); ibidem, 24–31.viii.1990, 2 females (LACM); ibidem, 16–19.xi.1990, 1 female (LACM); ibidem, 1–22.vi.1991, 1 female (LACM); 3 Km SE R. Naranjo (10.7911 N, 85.6689 W), 15–22.x.1991, F. D. Parker leg., 1 female (LACM); ibidem, 20.xi.1991, 1 male (LACM); ibidem, 1–5.xii.1991, 1 female (LACM); ibidem, 10–23.i.1992, 1 female (LACM); ibidem, 20–31.i.1992, 1 female (LACM); ibidem, 10–19.ii.1992, 1 female (LACM); ibidem, 23–31.viii.1992, 1 female (LACM); ibidem, iv.1992, 1 female (LACM); ibidem, v.1992, 1 female (LACM); ibidem, 13.vii.1992, 1 female (LACM); ibidem, 10–20.ix.1992, 1 male (LACM); ibidem, 11–20.ix.1992, 1 male (EMUS); ibidem, 1–9.x.1992, 1 female (LACM); ibidem, 22.x.1992, 1 female (LACM); ibidem, xii.1992, 1 female (LACM); ibidem, 4–8.i.1993, 1 female (LACM); ibidem, 15–19.iii.1993, 1 male (LACM); ibidem, 15–30.iv.1993, 1 female (LACM); ibidem, 1–9.vii.1993, 1 female (LACM); ibidem, 14–16.vii.1993, 1 female (LACM); ibidem, 1–3.vii.1993, 1 female (LACM); ibidem, 25.vii.1993, 1 female (LACM); ibidem, 23.viii.1993, 1 female (LACM). Heredia: La Selva (10.4333 N, 84.0167 W), 06.ix.1999, INBio-OET, 1 female (LACM); ibidem, 10.ix–03.x.1999, 1 female (LACM); ibidem, 20.ix.1999, 3 females (LACM); La Selva Res. Sta., 24–30.viii.1988, W. J. Hanson leg., 1 female (LACM). Higuito: San Mateo CR, Pablo Schild leg., 1 male (USNM). Limon: 7 mi N Guacimo, 22.ii–3.iii.1988, F. D. Parker leg., 1 female (LACM); Prov. Guapiles, 25.vi.1965, R. J. Hamton leg., 1 male (LACM). Puntarenas: Piedras Blancas 24 Km W (8.7833 N, 83.2500 W), 200 m, xi.1990, P. Hanson leg., 1 female (Det.: Kameneva, 2001) (USNM). San Jose: Escazu, 19–24.iv.1988, F. D. Parker leg., 1 female (LACM); ibidem, 15–22.vii.1989, 1 female (LACM); ibidem, 23–24.ix.1989, 1 female (LACM); San Isidro 9 mi S (9.331 N, 83.6993 W), 31.xii.1988, F. D. Parker leg., 1 female (EMUS). Turrialba: Catie/IICA Research Station, 24–30.vii.1981, W. R. Dolling B. M. 1981–411, 1 male (NHMUK 010862541). ECUADOR: Napo: Lago Agrio 8 Km W, 28.viii.1975, Langley & Cohen leg., 1 female (USNM); Limoncocha, 15.vi.1977, P. J. Spangler & D. R. Givens leg., 3 females (USNM); Misahualli nr. Tena, 6–19.x.2001, C. Brammer leg., 1 male (LACM); Res. Ethnica Waorani 1 Km S, Onkone Gare Camp (0.6527 S, 76.4333 W), 03.vii.1995, T. L. Erwin et al. leg., 1 female (USNM); ibidem, 26.vi.1996, 1 male (USNM); ibidem, 08.ii.1996, 1 male (Det.: V. Korneyev, 2008) (USNM); Yasuni Res. Sta. (0.6667 S, 76.3833 W), 250 m, 19–30.x.1998, W. J. Hanson leg., 8 females (LACM). Sucumbios: Sacha Lodge (0.5000 S, 76.4833 W), 270 m, 20–30.ix.1994, P. Hibbs leg., 1 female (LACM). EL SALVADOR: El Impossible, 04.vii.1977, Jarger leg., 1 female (USNM). La Unión (13.3351 N, 87.8470 W), 25.i.1957, P.A.B leg., 1 male (USNM); idem, 25.i.1957, G.R.S. leg., 2 females (USNM). Santa Tecla, 23.ii.1935, P.A.B. leg., 1 male (USNM). GUATEMALA: Escuintla: Palín, 1992, J. Lopez leg., 1 male and 1 female (Det.: A. Norrbom, 1992) (USNM). Rodriguez, 1 male (Det.: E. E. Austen, 8.ix.1926: Stylophthalmyia fascipennis Frey) (NHMUK 010862536). Sta. Lucia Cotz., 02–11.v.1988, F. D. Parker leg., 1 female (LACM). HONDURAS: Atlántida, La Ceiba, 07.ix.1916, F. J. Dyer leg., 1 female (USNM). MEXICO: Chiapas: Cacahoatan, 30.viii.1967, H. Sanchez R. leg., 2 females (USNM); Tuxtla Chico, 02.viii.1967, H. Sanchez R. leg., 1 male (USNM). Sinaloa, 68 mi SE Culiacan, 23.iv.1977, Hanson & Davis leg., 1 female (LACM). NICARAGUA: Masaya: La Concha, Estrada 62–2376, 02.vii.1961, Sequeira & Leal leg., 2 males (USNM). PANAMA: Barro Colorado: CZ, iv–v.1937, J. Zetek leg., 1 female (USNM); ibidem, vii.1937, 3 females (USNM); ibidem, x–xi.1941, 1 female (USNM); ibidem, i–iii.1944, 2 males and 2 females (USNM); Canal Zone, 09.vii.1978, N. E. Woodley leg., 1 female (Det.: Terpnomyia sp. G. Steyskal, 1982) (USNM); ibidem, 14.vii.1978, 1 female (USNM); Is, xi.1937, J. Zetek leg., 2 females (USNM); idem, 10–17.v.1964, W. D. & S. S. Duckworth leg., 1 male (USNM); idem, 25–28.iii.1965, S. S. & W. D. Duckworth leg., 1 female (USNM). Cerro Campana, 11–14.vii.1967, O. S. Flint, Jr. leg., 1 female (USNM). El Cermeño, iv–v.1939, 1 female (USNM); idem, xii.1939–i.1940, J. Zetek leg., 2 males and 1 female (USNM). La Campana, i–iii.1938, J. Zetek leg., 1 male and 3 females (USNM). PERU: Dp. Junin: Chanchamayo, 11.viii.1918, J. M. Schunke leg., 1 female (USNM). Huanuco: Cochicote, 08.ix.1965, J. C. Hitchcock, Jr. leg., 1 female (Det.: G. Steyskal, 1965) (USNM); vic. Tingo Maria, 1–5.vi.1999, W. Hanson & S. Keller leg., 2 females (LACM). Iquitos, iii–iv.1932, R. C. Shannon leg., 1 female (Det.: Greene: Terpnomyia latifrons Hendel) (USNM). TRINIDAD: Simla, Arima-Blanchisseuse Rd., 20.vii.1975, J. Price leg., 1 male (USNM); Simla Res. Sta., I., 2–15.vi.1981, Hanson & Clemons leg., 1 male and 6 females (LACM). VENEZUELA: Lara, 4 Km NW de La Pastora, 2–3.iii.1978, J. B. Heppner leg., 1 male (USNM). T. F. Amaz., Cerro de La Neblina Basecamp (0.8333 N, 66.1622 W), 140 m, 4–12.ii.1984, D. Davis & T. McCabe leg., 2 females (USNM); ibidem, 1–10.iii.1984, 1 female (USNM).

Distribution. Mexico, Belize*, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Trinidad, Venezuela, Colombia, Brazil (Maranhão*, Amapá*, Pará, Roraima*, Amazonas, Rondônia* and Acre*), Ecuador, Peru and Bolivia (Fig. 6).

Comments. A female specimen with the label "Paraguay: Depto Alto Paraguay, 14–16.04.1986, Pague, Solis leg. (USNM)" was analyzed and identified by Kameneva (2004b)as P. latifrons. However, from the analysis of this specimen, it was concluded that it belongs to the genus Pterocerina Hendel.

Plagiocephalus intermedius Kameneva 2004b

(Figs. 2G–I, 3E, F)

Plagiocephalus intermediusKameneva, 2004b: 19 (Holotype: 1 male, INBio; Paratypes: 3 males and 17 females, INBio; Type locality: Costa Rica: Puntarenas, Monteverde) (examined by photographs).

Type material. Holotype (male): Costa Rica: Puntarenas: Monteverde, San Luis, Buen Amigo, 1000–1350 m, ix.1994, 3224, Fuentes leg.. Paratypes (3 males, 17 females): Costa Rica: Cartago: A. C. Amistad, P. N. Tapanti, 1150 m, i.1994, Mora leg., 1 male; Guanacaste: Rio San Lorenzo, Tierras Morenas, Z. P. Tenorio, 1050 m, i.1993, Rodriguez leg., 1 female; P. N. Guanacaste: Est. Cacao, vii.1989, GNP Biod. Sur. leg., 1 female; idem, SW side Volcan Cacao, 1000–1400 m, xi–xii.1989, Blanco, Chaves leg., 1 female; idem, Lado SO Volcan Cacao, 800–1600 m, 12–17.vii.1993, Fuentes leg., 1 female; Puntarenas: Monteverde, San Luis, Buen Amigo, 100–1350 m, ix.1994, 1 female; ibidem, xi.1994, 1 male; ibidem, 25.xi–10.xii.1996, Fuentes leg., 1 male; Est. Pittier, Sendero Pittier, 1670 m, 26.vi.1995, Angulo leg., 1 female; ibidem, 21.vi–4.vii.1995, Moraga leg., 2 females; ibidem, 25.vi–4.vii.1995, Zumbado leg., 3 females; ibidem, 1700 m, 3.vii.1995, Zumbado leg., 2 females; Buenos Aires, Est. Altamira, Sendero Los Gigantes, 1450 m, 4.i–3.ii.2000, Rubi leg., 4 females (INBio).

Diagnosis.Plagiocephalus intermedius can be differentiated by the eyestalks of male longer than in P. lobularis and shorter than in P. latifrons (3.00–7.00 mm); female parafacialia yellow (Fig. 2G); radial-medial band with base wider than apex, at most barely touching the discal band (Fig. 3E, F). The species can also be distinguished by male face yellowish white; male scape, pedicel and first flagellomere entirely yellow, and female scape, pedicel and first flagellomere gold with apex darker (Fig. 2G); palpus yellow; proboscis reddish yellow, with brown and yellow setulae. Male wing of normal outline, without posterior lobes (Fig. 3E); crossvein r-m located distinctly before the apex of vein R₁ (Fig. 3E, F); male subapical band curved and narrower when touching the apical band (Fig. 3E); male apical band wider than in P. lobularis and P. latifrons (Fig. 3E). Male fore and mid legs entirely yellow, and hind leg yellow to gold; female legs brown with tarsi lighter, and fore femur yellowish on the apex (Fig. 2I).

Measurements: Male: Body: 4.50–5.80 mm; Wing: 4.70–6.10 mm. Female: Body:

2.88–3.35 mm; Wing: 4.62–5.7 mm; Oviscape: 1.29–1.50 mm.

Material examined. COSTA RICA: La Suiza, 1961, P. Schild leg., 1 female (USNM); idem, 24.xi.1961, PablShild leg., 1 female (USNM).

Distribution. Costa Rica (Fig. 6).

Comments. Part of the INBio collection, including the type material of P. intermedius, was aggregated to the Museo Nacional de Costa Rica. The material of P. intermedius from the USNM examined in this work had been previously misidentified as P. latifrons by Kameneva (2004b).

Discussion on the terminalia of Plagiocephalus

One of the characters shared by the species of Pterocallini is the distiphallus bare or microsetulose. In other tribes of Ulidiidae, mainly from the subfamily Otitinae, the distiphallus is found spinulose, setulose, or both (Kameneva and Korneyev, 2006). In both P. lobularis and P. latifrons, the distiphallus is microsetulose on the two thirds apical. Other genera of Pterocallini, such as Cymatosus Enderlein, Paragorgopis Giglio-Tos and some species of Neoacanthoneura Hendel, have the distiphallus bare (Kameneva, 2004a, 2009). The epandrium of Plagiocephalus is elongate oval, similarly to the group of species apicalis of the genus Neoacanthoneura (Kameneva, 2012). Other genera of the tribe, such as Cymatosus, Megalaemyia Hendel and Aciuroides Hendel (Kameneva, 2009, 2012), have the epandrium short oval shaped. Plagiocephalus male terminalia can also be differentiated from other genera of the tribe by the prensisetae with small denticles on the inner surface.

We did not find any significant morphological difference between the male terminalia of P. lobularis and P. latifrons, and among the female terminalia of the species. A low variation in the morphology of the male terminalia among closely related species was also observed in other sexually dimorphic species with an ornamented head, for instance, species of Teleopsis Rondani (Diopsidae) (Földvári et al., 2007) and Richardia Robineau-Desvoidy (Richardiidae) (Wendt and Ale-Rocha, 2015). A likely explanation for the low variation of the terminalia among ornamented species is the access of the female to the male quality before copulation. The size of ornamented traits, such the eyestalks or gena processes, indicates genetic quality for the females, thus sexual selection acts primarily on these traits (Badyaev, 2004; Kelly, 2014), preventing the terminalia to be highly variable, as usually do in several dipteran taxa.

The presence of three spherical spermathecae with a smooth surface in the female of the species of Plagiocephalus is also shared by the other species of Pterocallini. The Plagiocephalus species have the female terminalia with a narrow segment 8 and cerci with an oval shape, differently from Aciuroides Hendel, Apterocerina Hendel, Coscinum Hendel, Cyrtomostoma Hendel, Elapata Hendel, Lathrostigma Enderlein, Micropterocerus Hendel, Pterocerina, and Tetrapleura Schiner, which have the segment 8 and cerci flattened and rigid (Kameneva, 2012). Based on the female terminalia and in other characters, Kameneva (2012) proposed that these last genera form a monophyletic group, excluding Plagiocephalus of the clade. Together with the lack of phylogenies for the Neotropical groups of ulidiids, only a minority of the Pterocallini species had their male and female terminalia described or illustrated, precluding the positioning of Plagiocephalus in a possible clade within the tribe.

Remarks on the geographical distribution of Plagiocephalus

Plagiocephalus is a widely distributed genus in the Neotropical region (Fig. 6). Among the species of Plagiocephalus, P. latifrons has the largest area of distribution. The distribution pattern of P. latifrons and P. lobularis seems well delimited. The distribution range of P. latifrons goes from Northwest of the Neotropical region until the diagonal line from Northern Brazil to South of Bolivia, while the range of distribution of P. lobularis remains in the Atlantic side of South America, going from Northeastern Brazil to South Brazil, and North of Paraguay and Argentina. Plagiocephalus intermedius is known only from Costa Rica. This species probably is distributed across the highest parts of Central America and do not occur in sympatry with P. latifrons, which is mostly distributed in areas of low altitude of the Neotropics (Fig. 6). However, the patterns of distribution found may have been influenced by the deficiency of appropriate collecting in the Neotropical region.

Xanthacrona Wulp, another genus of Pterocallini, has a similar area of distribution, occurring from North of the Neotropical region to South of Brazil, Paraguay and North of Argentina (Soares et al., 2018). However, the pattern of distribution of any of the five described species of Xanthacrona overlap with the distribution pattern of the Plagiocephalus species. Considering the number of species, Ulidiidae are among the most unexplored families in the Neotropical region (Amorim, 2009). Before making robust statements on distribution and diversification patterns of the ulidiids in the Neotropical region, there are still much to be done, including more collecting of specimens, taxonomic revisions and phylogenetic analyses of this still unstudied group of flies.

Acknowledgments

The authors acknowledged all curators cited in M&M that sent us the specimens and photos used in this revision, and also, Freddy Bravo and Adolfo Calor for the donation of specimens. ACOV thanks the Natural History Museum of Los Angeles County for the study award. We are grateful to Carlos J. E. Lamas, Angelo P. Pinto and Rodrigo M. Feitosa for the useful comments in a previous version of this manuscript. The photos were taken with Automontage equipment available through the project "Taxonline: Rede Paranaense de Coleções" headed by L. Marinoni (UFPR). ACOV, LDW and CJBC thank the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for the provided support (process # 133999/2016–5, process # 152527/2018-4, and process # 309873/2016–9, respectively).

References

• Aczél, M.L., 1951. Catálogo de la subfamilia americana "Pterocallinae" (Dipt., Acalypt. Otitidae). Acta Zool. Lill. 11, 397-433.
• Amorim, D.S., 2009. Neotropical Diptera diversity: richness, patterns, and perspectives. In: Pape, T., Bickel, D., Meier, R. (Eds.), Diptera Diversity: Status, Challenges and Tools. Koninklijke Brill NV, Leiden, pp. 71–97.
• Badyaev, A.V., 2004. Developmental perspective on the evolution of sexual ornaments. Evol. Ecol. Res. 6, 975-991.
• Blanchard, E.E., 1938. Descripciones y anotaciones de dipteros argentinos. An. Soc. Cient. Argent. 125-126, 345-386.
• Blanchard, E.E., 1938. Seis nuevas espécies de Otitidos argentinos. Rev. Argent. Entomol. 1, 85-96.
• Blanchard, E.E., 1951. Un extraordinario pterocallido de Los Yungas. Rev. Soc. Entomol. Argent. 15, 32-34.
• Capoor, V.N., 1954. New species of Ortalidae (Diptera) from Brasil. Rev. Bras. Entomol. 2, 205-214.
• Carrera, M., 1950. Sobre o gênero Plagiocephalus Wiedemeann, 1830 (Diptera, Otitidae, Pterocallinae). Departamento de Zoologia da Secretaria de Agricultura de São Paulo, Pap. Avulsos Zool. 9, 259-268.
• Cresson, E.T., 1923. A new species of an Achias-like fly from Nicaragua, apparently belonging to the little-known genus Plagiocephalus (Diptera, Ortalidae). Entomol. News 34, 257-260.
• Cumming, J.M., Wood, D.M., 2009. Morphology and terminology. In: Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M., Woodley, N.E., Zumbado, M.A. (Eds.), Manual of Central American Diptera, vol. 1, pp. 9–50.
• Frey, R., 1926. Eine neue mittelamerikanische Dipteren – Gattung mit gestielten Augen. Not. Entomol. 6, 44-48.
• Földvári, M., Pomiankowski, A., Cotton, S., Carr, M., 2007. A morphological and molecular description of a new Teleopsis species (Diptera: Diopsidae) from Thailand. Zootaxa 1620, 37-51.
• Grimaldi, D.A., Engel, M.S., 2005. Evolution of Insects, University Press. Cambridge, New York.
• Hendel, F., 1909. Übersicht der bisher bekannten Arten der Pterocallinen (Dipt.). Deut. Entomol. Z., 1-84.
• Hendel, F., 1909b. Diptera, Fam. Muscaridae, Subfam. Pterocallinae. In: Wytsman, P. (Ed.), Genera Insectorum, Bruxelles, Fasc., vol. 96, p. 50 + 4 taf.
• Hendel, F., 1911. Diptera, Fam. Muscaridae, Subfam. Richardiinae. In: Wytsman, P. (Ed.), Genera Insectorum. Bruxelles, Fasc., vol. 113, p. 56 + 3 taf.
• Hendel, F., 1936. Ergebnisse einer zoologischen Sammelreise nach Brasilien, insbesondere in das Amazonas gebiet, ausgefuhrt von Dr H. Zerny.. X. Teil. Diptera: Muscidae acalyptratae (excl. Chloropidae). Ann. Nat. His. Mus. Wien. 47, 61-106.
• Hennig, W., 1952. Bemerkenswerte neue Acalyptraten in der Sammlung des Deutschen Entomologischen Institutes (Diptera: Acalyptrata). Beitr. Entomol. 2, 604-618.
• Kameneva, E.P., 2004. New records of picture-winged flies (Diptera: Ulidiidae) of Central America including the description of new taxa. Stud. Dipterol. 10, 609-652.
• Kameneva, E.P., 2004. A new species of the genus Plagiocephalus (Diptera, Ulidiidae) from Central America. Vestn. Zool. 38, 15-22.
• Kameneva, E.P., 2009. A new species of the genus Cymatosus (Diptera, Ulidiidae) from Central America, with a key to species. Vestn. Zool. 43, 267-273.
• Kameneva, E.P., 2012. Revision of Aciuroides Hendel and Neoacanthoneura Hendel (Diptera: Ulidiidae: Pterocallini). Zootaxa 3227, 1-33.
• Kameneva, E., Korneyev, V., 2006. Myennidini, a New Tribe of the Subfamily Otitinae (Diptera: Ulidiidae), with Discussion of the Suprageneric Classification of the Family. Isr. J. Entomol. Biotaxonomy of Tephritoidea 35–36, 497-586.
• Kameneva, E.P., Korneyev, V.A., Ramos-Pastrana, Y., 2017. A new genus, new species and new records of Ulidiidae (Diptera, Tephritoidea) from Colombia. Vestn. Zool. 51, 125-136.
• Kelly, C.D., 2014. Sexual selection, phenotypic variation, and allometry in genitalic and non-genitalic traits in the sexually size-dimorphic stick insect Micrarchus hystriculeus Biol. J. Linnean Soc. 113, 471-484.
• Loew, H., 1873. The family Ortalidae. In: Monographs of the Diptera of North America. Part III. Smithson. Misc. Collect. 11 (256), vii + 351 + xiii +4 pls.
• Macquart, J.P.M., 1835. Histoire naturelle des insectes. Dipteres. Tome deuxieme. N.E. Roret, Paris. iv + 703
• Macquart, J.P.M., 1843. Dipteres exotiques nouveaux ou peu connus. Mem. Soc. Aci. Agric. Lille., 162–460 + 36 pls.
• Norrbom, A.L., Kim, K.C., 1988. Revision of the schausi group of Anastrepha Schiner (Diptera: Tephritidae), with a discussion of the terminology of the female terminalia in the Tephritoidea. Ann. Entomol. Soc. Am. 81, 164-173.
• Osten-Sacken, C.R., 1881. Enumeration of the Diptera of the Malay Archipelago collected by Prof. Odoardo Beccari, Mr. L.M. D'albertis and others. Ann. Mus. Civ. Stor. Nat. Genova 16, 393-492.
• Soares, M.M.M., Santiago, A.S., Ale-Rocha, R., 2018. New records of Xanthacrona Wulp, 1899 (Diptera, Ulidiidae) from Brazil. Check List 14, 771-778.
• Steyskal, G.C., 1963. The genus Plagiocephalus Wiedemann (Dipt, Otitidae). Stud. Entomol. 6, 511-514.
• Steyskal, G.C., 1964. The synonymy of Willineria orfilai Blanchard (Diptera, Otitidae). Stud. Entomol. 7, 490.
• Steyskal, G.C., 1968. Family Otitidae (Ortalidae; including Pterocallidae, Ulidiidae). In: Vanzolini, P.E., Papavero, N. (Eds.), A Catalogue of the Diptera of the Americas South of the United States, vol. 54. Departamento de Zoologia da Secretaria de Agricultura de São Paulo, pp. 54.1–54.31.
• Wendt, L.D., Ale-Rocha, R., 2015. Antlered richardiid flies: new species of Richardia (Tephritoidea: Richardiidae) with antler-like genal processes. Entomol. Sci. 18, 153-166.
• Westwood, J.O., 1849. Diptera nonnula exotica descripta. Trans. Entomol. Soc. Lond. 5, 231-236.
• White, I.M., Headrick, D.H., Norrbom, A.L., Carrol, L.E., 1999. Glossary. In: Aluja, M., Norrbom A.L. (Eds.), Fruit Flies (Tephritidae): Phylogeny and Evolution of Behavior. CRC Press, Boca Raton, pp. 881–924.
• Wiedemann, C.R.W., 1830. Aussereuropäische zweiflügelige Insekten. Als Fortsetzung des Meigenschen Werks. Zweiter Theil. Hamm. Shulz 2, xii + 684 + 5 pls.
• Wiedemann, C.R.W., 1830. Achias dipterorum genus a Fabricio condituum; illustratum novisque speciebus auctum et conventui physicorum Germanorum oblantum. Kiliae Holsatorum, C.F. Mohr., 16 + 2 pls.

Publication Dates

• Publication in this collection
  Jan-Mar 2019

History

• Received
  9 July 2018
• Accepted
  19 Dec 2018