The name is a feminine noun in the nominative singular, formed from the type species name: [auro] Capilla (Zhang et al., 2023a).

Helias aurocapilla Staudinger, 1876, currently a junior subjective synonym of Hesperia musculus Burmeister, 1875.

The subgenus Capilla is phenotypically more diverse when compared to other subgenera of Staphylus. In most species, the harpe exceeds the ampulla. Usually, the fultura superior bears laterally two dense clusters of small spines (Figs 3B, 3E, 3F, 3H, 5A, 5F, 5J, 5N, 9B, 9F, 9J), which Evans (1953) cited as vinculum brushes. In females, the lamella postvaginalis is usually a plate with a central division on the distal margin, bearing microtrichia (Fig. 10), except in Staphylus (C.) insignis, where it is a large and rectangular, heavily sclerotized plate (Fig. 10E), in S. (C.) neideae sp. nov., where it is a small triangular plate (Fig. 10M), and in S. (C.). nicoleae sp. nov., where it is a horizontal, rectangle-shaped plate (Fig. 10L).

Head: Brown with some yellow or golden scales dorsally; white or yellow ventrally, third segment of the palp brown. Antenna brown dorsally, ventrally with small yellow dots at the joints on the side. Nudum with 10–11 segments. Thorax: Brown, sometimes with scarce golden or yellow scales dorsally, with white or cream hair-like scales ventrally. Legs brown, with some yellow scales and white hair-like scales. — FW length: 1.0–1.5 cm. HW length and shape: 0.7–1.2 cm. Outer margin undulated. DFW: Ranging from reddish brown to dark brown. Costal fold present. Two or three small white dots on the subapical area, between R₂-R₃, R₃-R₄ and R₄-R₅, can be present or absent. Hyaline dots in the cell, in M₃-CuA₁, in CuA₁-CuA₂ and in CuA₂-lA can be present or absent. Fringe concolorous, ranging from paler brown to brown, or bicolorous, brown and white. VFW: Ranging from light brown to brown, with markings as in DFW present or absent. DHW: Ranging from reddish brown to dark brown. VHW: Ranging from light brown to brown or with tornal area white. Genitalia: Tegumen rounded proximally, laterally straight, rounded or concave, usually tapering towards the apex, distally concave or with a small protuberance at the insertion of the uncus. Saccus broad. Uncus very variable, rectangular, tubular, or conical in shape, base dorsally bearing a tuft (sometimes absent) of short to long hair-like scales or setae. Fenestra developed or absent. Gnathos as two plates, ranging from heavily to weakly sclerotized. Valva: sacculus broad, variably shaped, sometimes rounded, other triangle-shaped, bearing short setae or a dense set of socketed spines, or at least thin hair-like scales internally; harpe broad, variably-shaped, rounded, straight or concave at the distal margin, usually concave at the proximal margin, may exceed ampulla, bearing external hair-like scales or socketed, stout, short to long spines; ampulla variable, broad or small, rounded, or with a straight distal margin, or as a rectangular plate, which may exceed the harpe, bearing external hair-like scales or socketed, stout, short to long spines. Aedeagus ranging from much shorter than valva to approximately the same length of valva, which may bear small denticles dorsally on the distal portion. Fultura superior as a plate which may bear two dense clusters of small to large, thin or stout, lateral spines.

The females differ from the males as follows: FW length: 1.1–1.5 cm. HW length: 0.8–1.1 cm. Wings: Usually lighter than males, with bands and hyaline white dots more distinct. Genitalia: The lamella postvaginalis is variable, usually plate with a central division on the distal margin, bearing microtrichia, or a small funnel-shaped or horizontal rectangle or quadrate plate. Membranous or at least less sclerotized area linking the postvaginalis and the antevaginalis lamellae. Lamella antevaginalis as a plate; the proximal margin around the ostium bursae variable, with external margins usually as pointed projections. Ductus bursae membranous, with a proximal very slightly sclerotized funnel-shaped area at the insertion with the lamella antevaginalis. Corpus bursae ranging from about 1/3 to 1/4 the length of the ductus bursae.

Species included:

Staphylus (Capilla) ascalon (Staudinger, 1876) (Helias)

= anginus Schaus, 1902

= dissimilis Hayward, 1937 (Pholisora)

Staphylus (Capilla) azteca (Scudder, 1872)

= machuca (Schaus, 1913) (Bolla)

= quadratus (Dyar, 1926) (Thanaos)

= inconstans Bell, 1932 (Staphylus)

Staphylus (Capilla) buena (Williams & Bell, 1940) (Pholisora)

Staphylus (Capilla) caribbea (Williams & Bell, 1940) (Pholisora)

Staphylus (Capilla) corumba (Williams & Bell, 1940) (Pholisora)

Staphylus (Capilla) eryx Evans, 1953

Staphylus (Capilla) insignis O. Mielke, 1980

Staphylus (Capilla) lizeri (Hayward, 1938) (Pholisora)

a) Staphylus (Capilla) lizeri lizeri (Hayward, 1938) (Pholisora)

= cruza (Williams & Bell, 1940) (Pholisora)

= imperspicua (Hayward, 1940) (Pholisora) new synonym

b) Staphylus (Capilla) lizeri album Steinhauser, 1989

Staphylus (Capilla) melangon (Mabille, 1883) (Hesperia)

a) Staphylus (Capilla) melangon melangon (Mabille, 1883) (Hesperia)

= melangon epicaste Mabille, 1903 new synonym

= rileyi (Hayward, 1937) (Pholisora)

b) Staphylus (Capilla) melangon stanga Evans, 1953

Staphylus (Capilla) melius Steinhauser, 1989

Staphylus (Capilla) musculus (Burmeister, 1875) (Hesperia)

= aurocapilla (Staudinger, 1876) (Helias)

= similis Bell, 1932

Staphylus (Capilla) perna Evans, 1953

Staphylus (Capilla) tucumanus (Plötz, 1884) (Nisoniades)

= cordovanus Draudt, 1923

= fascia Hayward, 1933

Staphylus (Capilla) nicoleae Lemes, new species

Staphylus (Capilla) neivae Lemes, new species

Staphylus (Capilla) ricardoi Lemes, new species

Staphylus (Capilla) neideae Lemes, new species

Staphylus (Capilla) tyro (Mabille, 1878)

Staudinger (1876) described Helias ascalon in the same description of Helias ascalaphus Staudinger, 1876. The name ascalon was subsequently placed in the genus Pholisora by different authors. Mabille (1903) cited it as Staphylus ascalon for the first time, being followed by most subsequent authors. Schaus (1902) described Staphylus anginus, treated as Pholisora anginus by Shepard (1935). Hayward (1933) cited Hesperia musculus Burmeister, 1875 as a synonym of Staphylus ascalon. Hayward (1937) also described Pholisora dissimilis. Evans (1953) synonymized S. anginus Schaus, 1902 and P. dissimilis Hayward, 1937 with Staphylus ascalon and revalidated Staphylus musculus (Burmeister, 1875). Subsequent authors have mentioned this species in taxonomic comments, faunistic studies and catalogues.

Helias ascalon Staudinger, 1876 was described based on the male [holotype] from Novo [sic] Friburgo, [Rio de Janeiro State], Brazil, deposited in Sommer collection. Currently, the holotype is deposited in MfN and contains the following labels: / Origin/ N. Friburgo/ Coll. Sommer/ aut. ascalon Stgr./ ascalon Stgr./ GEN. PREP., MIELKE 1996/.

Staphylus anginus Schaus, 1902 was described based on type No. 5985 deposited in USNM. However, some of the Schaus’s descriptions are based on more than one specimen with the same type-number label. For example, for Papias sobrinus Schaus, 1902, also deposited in USNM, there are two specimens labelled as Type No. 6034. Therefore, to stabilize the correct nomenclature, the male syntype from Novo [sic] Friburgo, [Rio de Janeiro State], deposited in USNM, is hereby designated as the lectotype, with the following labels: /Nova Friburgo, Brazil./ Collection W. Schaus/ Staphylus anginus type Sch./ Type No. 5985 U.S.N.M./ Lectotypus/ Lectotypus Staphylus anginus Schaus, 1902, Lemes, O. Mielke, Casagrande, Siewert, A. Warren, 2025.

Pholisora dissimilis Hayward, 1937 was described based on the female holotype from Puerto Bemberg (currently Puerto Liberdad), Misiones, Argentina, December 1933, Hayward leg., deposited in Hayward collection. The holotype is currently deposited in IML, with the following labels: /Bemberg, Misiones, Ab. 1933, Hayward/ Pholisora dissimilis Hayw. ♀, Holotipo/ Tipus/ H. 56/ N.° 210/.

Staphylus (C.) ascalon (Fig. 2A–D) externally resembles S. (C.) lizeri album (Fig. 2S, T), some specimens of S. (C.) melangon melangon (Fig. 4E–H), S. (C.) melangon stanga (Fig. 4I, J) and S. (C.) neideae sp. nov. (Fig. 7U–X) due to a white VHW tornus. Among these species, S. (C.) ascalon has unique male genitalia (Fig. 3A–D), with the uncus somewhat rectangular at the base and a tapered apex, and with ampulla broad, exceeding the harpe. The female genitalia (Fig. 10A) differ from the other species due to the lamella postvaginalis being very excavated at the center of the distal margin, bearing microtrichia, with the proximal part funnel-shaped and without microtrichia.

Fig. 2A, B. Head: Brown with golden scales dorsally, white cream ventrally. FW length: 1.2–1.3 cm. HW length: 0.9–1 cm. DFW: Brown. Three white dots in the subapical area, two hyaline dots in the cell and one in CuA₁-CuA₂. Fringe brown. VFW: Same white markings as on DFW. VHW: Lighter brown with tornal area white. Genitalia (Fig. 3A–D): Base of uncus somewhat rectangular, longer than wide, dorsally bearing a tuft of long hair-like scales, and with a tapered apex. Fenestra developed. Valva: sacculus rounded, bearing some hair-like scales internally; harpe rounded at the distal margin, excavated on its proximal-internal margin, not exceeding ampulla; ampulla broad, rounded, exceeding harpe, bearing long, thin setae at the distal margin. Aedeagus shorter than valva, bearing small denticles dorsally on the distal portion. Fultura superior bearing two dense clusters of small, thin spines laterally.

Fig. 2C, D. The females differ from the males in the following characters: FW length: 1.3–1.5 cm. HW length: 1–1.1 cm. Wings. Additional white dots can be present or absent: two in the cell, one in M₂-M₃, one in M₃-CuA₁ and one in CuA₁-CuA₂. Genitalia (Fig. 10A): The lamella postvaginalis is a plate with a deep central division on the distal margin, bearing microtrichia, with the proximal part funnel-shaped, without microtrichia. Membranous or at least less sclerotized area linking the lamellae postvaginalis and antevaginalis. Lamella antevaginalis as a plate with reticulations; the proximal margin around the ostium bursae not fused. Ductus bursae membranous, except on the proximal part that has a sclerotized semi-ring.

Fig. 11. Brazil: Espírito Santo, Goiás, Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul (Ritter et al. 2011; Piovesan and Morais 2015) and São Paulo; Paraguay (Hayward 1948); Argentina: Buenos Aires (Burmeister 1878, Hayward 1948), Córdoba (Núñez-Bustos 2015), Misiones, Santiago del Estero (Hayward 1934).

Staphylus (C.) ascalon is recorded throughout the year, except June.

Fonseca et al. (2006) recorded this species visiting the flowers of Stachytarpheta cayennensis (Rich.) (Verbenaceae).

Staudinger (1876) did not provide an etymology for the name ascalon. Schaus (1902) did not provide an etymology for the name anginus. Although Hayward (1937) did not provide an etymology for Pholisora dissimilis, he cites that the species is very near Staphylus similis Bell, 1932; that is presumably the reason why he named the species as dissimilis.

The genitalia of the holotype of Pholisora dissimilis is partially destroyed, lacking its corpus and ductus bursae (O. Mielke, 1975).

BRAZIL: Espírito Santo – Baixo Gandu, 26.VII.1966, 3♂, O. Mielke, Brown & Elias leg., DZ 370*, DZ 44.762, DZ 44.763 (DZUP); 16.IX.1966, 1♂, Claudinor Elias leg., DZ 846* (DZUP); 11.X.1971, 1♂, C. & C. T. Elias leg., DZ 40.851* (DZUP); Santa Teresa, 4–7.III.1973, 1♀, C. Callaghan leg., A. C. Allyn Acc. 1973-24, Genit. Prep. SRS-1521* (MGCL). Goiás – Formosa, Salto da Itiquira, 15.V.1977, 1♀, Forer leg., DZ 46.836* (DZUP). Minas Gerais – Barbacena, 17.II.1966, 1♀, O. Mielke leg., OM 8.078* (OM). Brumadinho, 900 m, 5.I.2002, 1♀, O.-C. Mielke leg., OM 55.735* (OM). Camanducaia, Monte Verde, 1.650 m, 6.IX.1967, 1♀, H. Ebert leg., Coleção H. Ebert, DZ 46.724* (DZUP). Caratinga, Estação Ecológica de Caratinga, 400 m, 29.I–3.II.2003, 2♂, O. Mielke & Casagrande leg., DZ 44.775, DZ 44.776* (DZUP). Catas Altas, 1300 m, 6.II.2003, 2♀, O. Mielke & Casagrande leg., OM 59.153, OM 59.369 (OM). Conceição dos Ouros, Serra Grande, 1.365 m, 8.II.2003, 4♀, O. Mielke & Casagrande leg., OM 58.766, OM 58.883*, OM 58.920, OM 59.028 (OM). Ibirité, 800 m, 11.III.1993, 1♂, O. Mielke & Machado leg., OM 37.533* (OM). Juiz de Fora, 500–800 m, 10.XII.1953, 1♂, H. Ebert leg., Coleção H. Ebert, DZ 46.753* (DZUP); Recanto dos Lagos, 25.II.2010, 3♂, N. Tangerini leg., Ex Coleção Nirton Tangerini, DZ 46.657*, DZ 46.744*, DZ 46.745 (DZUP). Passa Quatro, Paiolinho, Fazenda Serra Fina, 1.600 m, 16.II.2010, 2♂, 3♀, O. Mielke & Casagrande leg., DZ 46.811*, DZ 46.812*, DZ 46.813, DZ 46.814, DZ 46.815* (DZUP). Poços de Caldas, 1.250 m, 2.III.1964, H. Ebert leg., Coleção H. Ebert, DZ 46.719* (DZUP); 6.III.1964, 1♂, 2♀, H. Ebert leg., Coleção H. Ebert, DZ 46.819*, DZ 46.754, DZ 46.755* (DZUP); Morro do Ferro, 1.600 m, 27.III.1965, 1♂, O. Mielke leg., OM 11.288 (OM); 1400 m, 27.III.1965, 1♂, O. Mielke leg., OM 11.286 (OM); 30.III.1965, 1♂, O. Mielke leg., OM 11.287* (OM); Serra da Mantiqueira, 1.100 m, 5.IX.1992, 2♂, H. Ebert leg., Coleção H. Ebert, DZ 46.756*, DZ 46.757 (DZUP). Santos Dumont, 850 m, 10.XII.1953, 1♂, H. Ebert leg., Coleção H. Ebert, DZ 46.752* (DZUP). São Gonçalo do Rio Abaixo, Peti, 560 m, 30.XI.2002, 1♂, N. G. Fonseca leg., DZ 5.320* (DZUP); 5–8.XII.2002, 2♂, O. Mielke leg., DZ 44.774, DZ 44.777 (DZUP). Varginha, III.1972, 1♂, Alvarenga leg., DZ 46.817* (DZUP). Virgínia, 1.500 m, 13–15.II.2010, 2♂, O. Mielke & Casagrande leg., DZ 46.809*, DZ 46.810* (DZUP). Paraná – Ponta Grossa, Vila Velha, 8.XII.1961, 1♂, O. Mielke leg., DZ 44.752 (DZUP); 3.II.1966, 1♂, no collector, OM 10.225* (OM); 1.V.1966, 2♂, O. Mielke & Laroba leg., DZ 841*, DZ 44.753 (DZUP); 3.IV.1971, 4♂, 1♀, Moure & O. Mielke leg., DZ 44.743, DZ 44.744, DZ 44.745, DZ 44.747, DZ 915* (DZUP); 16.III.1972, 2♂, O. Mielke leg., DZ 44.746, DZ 44.754 (DZUP); 5.III.1973, 1♂, 1♀, O. Mielke leg., DZ 44.755, DZ 44.759 (DZUP); 07.III.1973, 1♂, O. Mielke leg., DZ 44.748 (DZUP); 2.III.1974, 1♀, O. Mielke leg., DZ 44.760 (DZUP); 31.III.1974, 1♀, O. Mielke leg., DZ 44.761 (DZUP); 7.III.1975, 1♂, O. Mielke leg., DZ 44.756 (DZUP); 4.IV.1981, 1♀, O. Mielke leg., DZ 44.757 (DZUP); 21.III.2002, 1♂, 2♀, O. Mielke leg., DZ 44.749, DZ 44.750, DZ 44.751 (DZUP). São Pedro do Ivai, Reserva Particular do Patrimônio Natural (RPPN) da Fazenda Barbacena, 380m, 23°48’34.7”S, 51°55’45.4”W. 8.II.2016, 1♂, 1♀, Andrew D. Warren (ADW); Rio de Janeiro – No specific locality, 5.VIII.1966, 1♂, H. Ebert leg., Coleção H. Ebert, DZ 46.747* (DZUP); Guapimirim, 1.III.1970, 1♂, O. Mielke leg., DZ 849* (DZUP). Petrópolis, Alto da Serra, 18.X.1964, 1♂, O. Mielke leg., OM 6.092* (OM); 28.XI.1965, 2♂, 1♀, O. Mielke leg., OM 7.603, OM 7.604, OM 7.605 (OM). São Paulo – No specific locality, 25.X.1962, 1♀, H. Ebert leg., Coleção H. Ebert, DZ 46.728 (DZUP). Campos do Jordão, 1.200 m, I.1966, 1♂, Ebert leg., Coleção H. Ebert, DZ 45.146* (DZUP). Cordeirópolis, 600 m, 28.XI.1965, 1♀, H. Ebert leg., Coleção H. Ebert, DZ 46.730 (DZUP). Ribeirão Pires, 800 m, 22.IV.1963, 1♂, H. Ebert leg., Coleção H. Ebert, DZ 46.748 (DZUP). Teodoro Sampaio, Morro do Diabo, 300–600 m, 22–25.X.1987, 2♂, OM 17.996*, OM 17.998* (OM). — ARGENTINA: Misiones – Puerto Iguazú, 19.I.1998, 1♂, Nuñez leg., DZ 15.404* (DZUP).

Scudder (1872) described Pholisora azteca, which was later placed in the genera Nisoniades and Bolla by different authors. Godman and Salvin (1896) cited it as Staphylus aztecus [sic]. Schaus (1913) described Bolla machuca, which was placed in the genera Staphylus and Pholisora by different authors. Dyar (1926) described Thanaos quadratus, which was placed in the genera Erynnis and Staphylus by different authors. Bell (1932) described Staphylus inconstans, which was placed in the genus Pholisora by different authors. Evans (1953) treated the name azteca as a subspecies, together with Staphylus azteca tyro (Mabille, 1878). He also synonymized unifascia Mabille, 1889, machuca Schaus, 1913 and inconstans Bell, 1932 under Staphylus a. azteca. Steinhauser (1975) reinstated the specific status for Staphylus azteca and Staphylus tyro. Most subsequent authors followed Evans (1953) and Steinhauser (1975), and have mentioned this species in taxonomic comments, faunistic studies and catalogues.

Pholisora azteca Scudder, 1872 was described from an unspecified number of specimens from Tehuantepec, [Oaxaca, Mexico]. One male syntype was found in the MCZ. To stabilize the nomenclature, this syntype is hereby designated as the lectotype with the following labels: / TYPE 15319/ Tehu./ Pholisora azteca Type Scudd./ Lectotypus/ Lectotypus Pholisora azteca Scudder, 1872, Lemes, O. Mielke, Casagrande, Siewert, A. Warren, 2025.

Bolla machuca Schaus, 1913 was described from an unspecified number of males from San Mateo, Costa Rica, deposited in NHMUK. Two male syntypes were found, one in NHMUK and the other in USNM. To stabilize the nomenclature, the syntype in NHMUK is designated as the lectotype, with the following labels: / H 761 [round type label underside]/ Bolla machuca Schaus/ Costa Rica. San Mateo CR 1912-503 W. Schaus/ [label with genitalia attached]/ BMNH(E) 1669536/ Lectotypus/ Lectotypus Bolla machuca Schaus, 1913, Lemes, O. Mielke, Casagrande, Siewert, A. Warren, 2025/. The paralectotype has the following labels: / June/ SanMateo CR/ Bolla machuca type Schs/ Type no 16814 U.S.N.M./ Paralectotypus/ Paralectotypus Bolla machuca Schaus, 1913, Lemes, O. Mielke, Casagrande, Siewert, A. Warren, 2025/

Thanaos quadratus Dyar, 1926 was described from a [female] [holo]type, deposited in USNM, with the following labels: / Colima Colima Mexico/ RMuller Collector/ May 1925/ Thanaos quadratus type Dyar/ Type No. 39988 U.S.N.M./ GENITALIA NO. X-42 87/ J.M.Burns 1997/.

Staphylus inconstans Bell, 1932 was described based on the male holotype from Orotina, Costa Rica, 7.IX.1931, O. Fulda leg., deposited in Bell collection, currently in AMNH, with the following labels: /Orotina Costa Rica IX.7.31 2700 ft./ G594/ TYPE/ Staphylus inconstans Bell Holotype ♂/.

Males of Staphylus (C.) azteca are most similar to S. (C.) tyro due to the rectangular uncus (Fig. 8I–L), but in S. (C.) azteca (Fig. 8E–H) the distal part of the uncus ends with two small, lateral, sharply pointed tips, and ventrally with a center carina. Also, S. (C.) azteca occurs in Mexico and Central America, while S. (C.) tyro is known from Colombia and Venezuela (Fig. 15).

Fig. 7E, F. FW length: 1.2–1.5 cm. HW length: 0.9–1.1 cm. Genitalia (Fig. 8E–H): Uncus rectangular, slightly concave laterally, with two small, lateral, sharply pointed tips at the distal margin, and ventrally with a central longitudinal carina. Gnathos as two lateral plates, very sclerotized and distally expanded, bearing microtrichia. Valva with ampulla at distal margin bearing denticles, without a finger-like projection at the ventral margin, externally bearing tufts of large hair-like scales. Fultura superior without cluster of spines.

Fig. 7G, H. The females differ from the males in the following characters: FW length: 1.4–1.6 cm. HW length: 1.0–1.1 cm. Genitalia (Fig. 10J): The lamella postvaginalis appears as a funnel-shaped plate.

Fig. 15. Mexico: Chiapas, Colima (Dyar 1926; Freeman 1977), Guerrero, Jalisco, Michoacán, Morelos, Nayarit (Freeman 1977), Oaxaca, Puebla (C. Hoffmann 1941), San Luis Potosí, Sinaloa (Godman and Salvin 1896), Tabasco (Godman and Salvin 1896; C. Hoffmann 1941), Tamaulipas and Veracruz. Guatemala: San Marcos and Zacapa. Honduras (Godman and Salvin 1896; Bell 1932). El Salvador: Ahuachapán, La Libertad, San Miguel and San Salvador. Nicaragua: Carazo, León and Matagalpa. Costa Rica: Alajuela, Guanacaste, Limón, Puntarenas, San José. Panama: Chiriqui, Granada (Anderson 2007), Managua (Anderson 2007) and Panamá.

Staphylus (C.) azteca has been recorded throughout the year.

According to Janzen and Hallwachs (2009), the following parasitoid species emerged from Staphylus azteca: Hymenoptera: Braconidae: Meteorus alejandromasisi Zitani, 1998; and Diptera: Tachinidae: Houghia aurifera Fleming & Wood, 2014.

According to Janzen and Hallwachs (2009), larvae of Staphylus azteca feed on the following plants: Amaranthaceae: Achyranthes aspera L.; Celosia nitida Vahl.; Alternanthera laguroides (Standl.) Standl.; Alternanthera pubiflora (Benth.) Kuntze; Amaranthus spinosus L.; Chamissoa altissima (Jacq.) Kunth; Cyathula achyranthoides (Kunth) Moq; and Caryophyllaceae: Polycarpaea nivea (Ait.) Webb.

Although Scudder (1872) did not mention so in the description, he certainly proposed the epithet in honor of the Aztec civilization, a population that flourished in Mexico, the type locality of Staphylus (C.) azteca. Schaus (1913) did not provide an etymology for the name machuca. Dyar (1926) did not provide an etymology for the name quadratus. Bell (1932) did not provide an etymology for the name inconstans.

Before this study, there was no illustration of the female genitalia of Staphylus (C.) azteca in the literature. Currently, the names machuca and quadratus are maintained as synonyms of azteca because we have not analyzed their genitalia nor assessed their DNA sequences. Thus, it is suggested that future researchers study the types of these taxa, as they could represent distinct species or be misplaced in the synonymy of Staphylus.

MEXICO: Chiapas – Comitán de Domínguez, 1♀, VII–VIII.2010, OM 76.585 (OM). El Aguacero, 2300’, 3.IX.1989, 2♂, John Kemner leg., Allyn Museum Acc. 1989-7, MGCL/FLMNH 22999, MGCL/FLMNH 23000 (MGCL). Escuintla, 2000’, 8.IX.1974, 1♂, R. Wind leg., A. C. Allyn Acc. 1974-24, Genit. Prep. SRS-1410*, MGCL/FLMNH 23010 (MGCL). San Jerónimo, 14.VII.1975, 1♂, no collector, MGCL Accession #2016-40 – E. C. Knudson-Knudson/Bordelon (MGCL); 17.VII.1978, 1♂, Welling leg., OM 45.033* (OM); 28.VII.1978, 1♀, Welling leg., DZ 45.085* (DZUP). San Quintin, 8.VII.1975, 1♂, no collector, MGCL Accession #2016-40 – E. C. Knudson-Knudson/Bordelon (MGCL). Guerrero – Pueblo Madero (El Playón), 5 mi. N., 2.IX.1967, 1♂, Allyn Found. Acc. 1970-1, Genit. Prep. SRS-1429*, MGCL/FLMNH 23008 (MGCL). Tierra Colorada, 6 mi. S., Rio Papagayo, 180 m, 28.VIII.1967, 5♂, 2♀, no collector, Allyn Museum Acc. 1973-16, MGCL/FLMNH 22994, MGCL/FLMNH 22995, MGCL/FLMNH 22996, MGCL/FLMNH 22997, MGCL/FLMNH 22998, MGCL/FLMNH 23011, MGCL/FLMNH 23012 (MGCL). Zihuatanejo, 28.III.1986, 1♂, 1♀, D. L. Lindsley leg., D. L. Lindsley colln.-MGCL Accession #2008-20 (MGCL). Jalisco – La Manzanilla, 21.XII.2000, 1♂, D. L. Lindsley leg., D. L. Lindsley colln.-MGCL Accession #2008-20 (MGCL); 22.XII.2000, 2♂, D. L. Lindsley leg., D. L. Lindsley colln.-MGCL Accession #2008-20, GEN. PREP.* (MGCL); 31.XII.2000, 2♂, D. L. Lindsley leg., D. L. Lindsley colln.-MGCL Accession #2008-20, GEN. PREP.* (MGCL). Michoacán – Uruapan, P. H. Cupatitzio, 4.V.1991, 1♀, L. González-Cota leg., MGCL Accession #2016-40. – E. C. Knudson – Knudson/Bordelon (MGCL); 5.III.1995, 1♂, A. D. Warren leg., MGCL Accession #2016-40. – E. C. Knudson – Knudson/Bordelon (MGCL). Morelos – Cuernavaca, II.1991, 1♀, no collector, J. Brenner coll.-MGCL Accession #2015-33 (MGCL). Oaxaca – Candelaria Loxicha, 14.VI.1972, 1♂, Welling leg., DZ 45.084 (DZUP); 13.VI.1989, 1♂, J. Kemner leg., G. T. Austin colln.-MGCL Acc. 2004-5, Genit. Vial GTA-1445*, MGCL/FLMNH 37004 (MGCL); San Luis Potosí – Ciudad Valles, 13.VI.1972, 2♂, H. A. Freeman leg., A. C. Allyn Acc. 1973-37, MGCL/FLMNH 22981, MGCL/FLMNH 22983 (MGCL); 2.VI.1973, 1♂, 1♀, H. A. Freeman leg., A. C. Allyn Acc. 1973-37, Genit. Prep. SRS-1529*, MGCL/FLMNH 22992, MGCL/FLMNH 23015; 14.VI.1973, 1♂, H. A. Freeman leg., A. C. Allyn Acc. 1973-37, MGCL/FLMNH 22986 (MGCL); 15.VI.1973, 1♂, H. A. Freeman leg., A. C. Allyn Acc. 1973-37, MGCL/FLMNH 22982 (MGCL); 19.VI.1973, 1♂, H. A. Freeman leg., A. C. Allyn Acc. 1973-37, MGCL/FLMNH 22984 (MGCL); 21.VI.1971, 4♂, H. A. Freeman leg., A. C. Allyn Acc. 1973-37, MGCL/FLMNH 22985, MGCL/FLMNH 22987, MGCL/FLMNH 22989, MGCL/FLMNH 22993 (MGCL); 24.VI.1973, 1♀, H. A. Freeman leg., A. C. Allyn Acc. 1973-37, Genit. Prep. SRS-1437*, MGCL/FLMNH 23017 (MGCL); 25.VI.1973, 1♂, H. A. Freeman leg., A. C. Allyn Acc. 1973-37, MGCL/FLMNH 22988 (MGCL); 16.VI.1977, 1♂, H. A. Freeman leg., A. C. Allyn Acc. 1973-37, MGCL/FLMNH 22990 (MGCL); 25.VI.1977, 1♂, H. A. Freeman leg., A. C. Allyn Acc. 1973-37, MGCL/FLMNH 22991 (MGCL). Matlapa, 24.V.1962, 1♂, G. W. Rawson leg., Allyn Museum Acc. 1981-11, MGCL/FLMNH 23004 (MGCL). Tamaulipas – No specific locality, Galeana Canyon, 16.VIII.1967, 1♀, no collector, Allyn Museum Acc. 1987-11, MGCL/FLMNH 23014 (MGCL). Gomez Farias, 2.IX.1967, 1♀, no collector, Allyn Museum Acc. 1973-16, MGCL/FLMNH 23013 (MGCL); 10.IX.1967, 2♂, Lee D. Miller leg., Allyn Museum Acc. 1973-16, MGCL/FLMNH 23005, MGCL/FLMNH 23006 (MGCL); 20.II.1969, 2♂, L. D. & J. Y. Miller leg., A. C. Allyn Acc. 1969-1, Genit. Prep. SRS-1362*, MGCL/FLMNH 23007, MGCL/FLMNH 23009 (MGCL). Veracruz – Catemaco, 470 m, VI–VII.2012, 1♂, G. Nogueira leg., OM 76.125* (OM); Dos Amates, 17.V.1972, 1♂, no collector, MGCL Accession #2016-40 – E. C. Knudson – Knudson/Bordelon (MGCL); La Victoria, VI–VIII.2013, 1♂, G. Nogueira leg., OM 77.252 (OM); Posolapan, 490 m, VI–VII.2012, 2♂, 1♀, G. Nogueira leg., OM 77.231, OM 77.161, OM 76.687 (OM); VI–VIII.2013, 2♀, G. Nogueira leg., OM 77.041, OM 77.101* (OM). La Candelaria, 500 m, VII–VIII.2013, 1♂, G. Nogueira leg., OM 77.172* (OM). Presidio, IX.1939, 1♂, T. Escalante leg., A. C. Allyn Acc. 1973-48, MGCL/FLMNH 23002 (MGCL); VI.1940, 1♂, T. Escalante leg., A. C. Allyn Acc. 1973-48, MGCL/FLMNH 23003 (MGCL). Tecolapán, 450 m, VII–VIII.2013, 3♂, 1♀, OM 76.811, OM 76.831*, OM 76.881, OM 76.901 (OM). Veracruz, 31.IX.1967, 1♂, no collector, Allyn Museum Acc. 1981-11, MGCL/FLMNH 23001 (MGCL). GUATEMALA: San Marcos – Finca los Andes, XI.1974, 1♂, M. Simon leg., A. C. Allyn Acc. 1984-31 (MGCL). Zacapa – Santa Cruz, 18.IX.1993, 1♂, D. & J. Lindsley leg. (MGCL). — EL SALVADOR: Ahuachapán - Ahuachapán, River El Molino, 680 m, 22.IV.1971, 2♂, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23 (MGCL). La Libertad – La Libertad, 10 m, 15.VIII.1970, 1♂, 2♀, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23 (MGCL); 8.VI.1971, 1♂, M. Serrano leg., A. C. Allyn Acc. 1971-25 (MGCL); 6.XI.1971, 2♀, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23, Genit. Prep. SRS-223* (MGCL); 27.XI.1971, 2♀, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23 (MGCL); 28.XI.1971, 1♂, 3♀, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23 (MGCL); 5.II.1972, 1♀, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23, Genit. Prep. SRS-1511* (MGCL); 19.III.1972, 2♂, 2♀, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23 (MGCL). Santa Tecla, 900 m, 20.V.1967, 1♀, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23 (MGCL); 15.VII.1970, 1♀, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23 (MGCL); 29.VII.1970, 1♂, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23, MGCL 1112133 (MGCL); 18.II.1972, 1♀, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23 (MGCL); 13.V.1972, 1♂, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23 (MGCL); 23.V.1972, 1♀, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23 (MGCL); 25.V.1972, 1♂, 1♀, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23, Genit. Prep. SRS-236* (MGCL); 3.VI.1972, 1♂, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23 (MGCL); 16.VI.1972, 1♀, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23 (MGCL); 20.VI.1972, 1♀, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23 (MGCL); 23.VI.1972, 1♂, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23 (MGCL); 25.VI.1972, 1♀, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23 (MGCL); 13.VII.1972, 1♀, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23 (MGCL); Los Chorros, 8.VII.1971, 1♂, M. Serrano leg., A. C. Allyn Acc. 1971-42 (MGCL). San Miguel – Ciudad Barrios, 550 m, 26.IV.1972, 1♀, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23 (MGCL). San Salvador – San Salvador, 10.VI.1971, 1♀, M. Serrano leg., A. C. Allyn Acc. 1971-25 (MGCL); 1.VII.1971, 1♂, M. Serrano leg., A. C. Allyn Acc. 1971-25 (MGCL); 15.IX.1971, 1♀, M. Serrano leg., A. C. Allyn Acc. 1971-48 (MGCL); Cerro San Jacinto, 800–900 m, 27.V.1972, 1♂, S. & L. Steinhauser leg., A. C. Allyn Acc. 1973-23, MGCL 1112091 (MGCL). — NICARAGUA: Carazo – Diriamba, 2000’, 1.VIII.1971, 1♀, T. Taylor leg., A. C. Allyn Acc. 1971-40, Genit. Prep. SRS-1353* (MGCL). León – León, Rio Pochote, 25.XII.1994, 4♂, 4♀, L. R. Hernandez & J. M. Maos leg., Allyn Museum Acc. 1995-21, Genit. Vial No. SRS-4691*, Genit. Vial No. SRS-4692*, Genit. Vial No. SRS-4696*, Genit. Vial No. SRS-4685*, Genit. Vial No. SRS-4687* (MGCL); Ponelya Beach, 2.I.1995, 1♀, L. R. Hernandez & J. M. Maos leg., Allyn Museum Acc. 1995-21, Genit. Vial No. SRS-4693* (MGCL). Matagalpa – Matagalpa, Fuente Pura, 28.XII.1994, 1♂, L. R. Hernandez leg., Allyn Museum Acc. 1995-21 (MGCL). — COSTA RICA: Alajuela – Guayabal, 26.IX.1987, 1♀, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial SRS-2996* (MGCL). Upala, 22.IX.1987, 1♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL). San Antonio, Ojo de Agua, 21.IX.1986, 1♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL). Guanacaste – Cañas, 20.V.1972, 1♂, H. L. King leg., A. C. Allyn Acc. 1972-5 (MGCL); Finca Taboga, 18.IX.1986, 1♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial SRS-2971* (MGCL). Hacienda Naranjo, Área de Conservación Guanacaste, Rio Gongora, 6.VII.1997, 2♂, 2♀, D. & J. Lindsley leg., D. L. Lindsley colln.-MGCL Accession #2008-20, GEN. PREP* (MGCL); Santa Rosa Plateau, 12.VII.1997, 1♂ D. & J. Lindsley leg., D. L. Lindsley colln.-MGCL Accession #2008-20, GEN. PREP* (MGCL). Playa Del Coco, Cerro Ceiba, 23.IX.1987, 1♂, G. & A. Austin leg., G. T. Austin colln. MGCL Acc. 2004-5, Genit. Vial SRS-2983*, MGCL 1112103 (MGCL). Limón – Not specific locality, Germana, 15.IV.1989, 1♂, G. & A. Austin leg., G. T. Austin colln. MGCL Acc. 2004-5 (MGCL). Puntarenas – No specific locality, Rio Pita, 18.IX.1987, 2♂, 1♀, G. & A. Austin leg., G. T. Austin colln. MGCL Acc. 2004-5, Genit. Vial SRS-3002* (MGCL). Buenos Aires, 13.IX.1987, 1♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial SRS-2961* (MGCL). Mata de Limón, 15.III.1986, 6♂, 1♀, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial SRS-2985*, MGCL 1043694 (MGCL). Miramar, 16.IX.1986, 1♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL). San José – Colón, 13.VI.1990, 1♂, Pagels leg., OM 26.260* (OM); 20.VI.1990, 1♀, Pagels leg., OM 26.302* (OM); Rodeo, 5.I.1992, O. Posels leg., OM 32.765* (OM). Desamparados, Calle Fallas, 20.IX.1986, 2♀, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, MGCL 1043724, Genit. Vial SRS-3021*, Genit. Vial SRS-3022* (MGCL). Escazú, 1.VII.1990, 1♂, Pagels leg., OM 26.179* (OM). San Isidro de El General, 14.3 Km N., 3.IV.1989, 1♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, MGCL 1043688 (MGCL). — PANAMA: Chiriqui – Boquete, Caldera, 400 m, 22.VII.1997, 1♂, O. Mielke & Casagrande leg., OM 46.429* (OM). Panama – No specific locality, Bayano Lake Bridge, 8–10 Km SE of, 9°10’496"N, 78°44’568"W, 29.V.2009, 1♂, 1♀, C. V. Covell Jr. leg., C. V. Covell colln.-MGCL-Acc. 2009-19 (MGCL); Rio Manoni, 9°11’947"N, 79°4’374"W, 23.V.2009, 1♀, Dennis Strenge leg., C. V. Covell colln.-MGCL-Acc. 2009-19 (MGCL).

Williams and Bell (1940) described Pholisora buena. Evans (1953) cited it as Staphylus buena, as have most subsequent authors that mentioned this species in taxonomic comments, faunistic studies and catalogues.

Pholisora buena Williams & Bell, 1940 was described from the male holotype from Buenavista, 75 km northwest of Santa Cruz, Bolivia, deposited in ANSP, currently in CMNH, with the following labels: / Type 7188 BUENA Will-Bell/ Buenavista 75 Km. N. W. of Santa Cruz, Bolivia/ Pholisora buena Williams + Bell Holotype ♂/ 450 m alt. 1926–27 (Jose Steinbach)/ Genitalia slide No. 2294/.

Staphylus (C.) buena is readily distinguished from other species of Staphylus (Capilla) by a combination of characters of the male genitalia (Fig. 3E, F): sacculus bearing a dense set of small spines internally and ampulla produced at distodorsal margin, bearing a dense set of small, socketed spines, proximally concave with a very produced dentate area on the ventral-internal margin.

Fig. 2E, F. Head: Brown with golden scales dorsally; presumably yellow ventrally. FW length: 1.3 cm. HW length: 1 cm. DFW: Brown, two subapical white dots. VFW: Same white markings as on DFW. VHW. Brown. Genitalia (Fig. 3E, F). Dorsal view of uncus unknown, apparently conical, curving downwards in lateral view, tapering distally, bearing a tuft of short hair-like scales at the base. Valva: sacculus scalene, triangular-shaped, bearing a dense set of small spines internally; harpe rounded at distal margin, not exceeding ampulla; ampulla broad, produced at distodorsal margin, bearing a dense set of small, socketed spines, proximally concave with a very produced dentate area on the ventral-internal margin. Aedeagus approximately the same length of valva, bearing small denticles on the distal portion. Fultura superior bearing two dense clusters of apparently large, stout spines laterally.

Unknown.

Fig. 11. Bolivia: Buena Vista; Argentina: Salta (Hayward 1973), Santa Fé (Hayward 1973) and Tucumán (Hayward 1973).

Unknown.

Although not mentioned in the description, the epithet of Staphylus (C.) buena clearly is in reference to the type locality, Buenavista (Bolivia).

As the holotype genitalia was not studied, all the morphological interpretations are from the drawing of the genitalia in the original description (Fig. 3E). In the original drawing, the dorsal view of the genitalia is not illustrated, also there is no indication of the saccus. It is assumed that the saccus is triangular and broad, as the other species of the subgenus. In the costa region of the valva, there is a set of spines. However, the species of Staphylus (Capilla) do not have a well-defined costa. These species usually have a fultura superior developed as a plate bearing two dense clusters of small stout spines laterally. Therefore, in this study, the spines represented in the costa region of the original drawing are treated as the spines of the fultura superior.

Williams and Bell (1940) described Pholisora caribbea. Evans (1953) cited it as Staphylus caribbea for the first time, being followed by other authors that mentioned this species in taxonomic comments, faunistic studies and catalogues.

Pholisora caribbea Williams & Bell, 1940 was described based on the male holotype from Barro Colorado Island, Canal Zone, Panama, deposited in AMNH, with the following labels: /Panama XI-16.’30/ Barro Colorado Canal Zone/ Collection E.I.Huntington/ G988/ Pholisora caribbea Williams + Bell Holotype ♂/ Holotype/.

Staphylus (C.) caribbea has a brown VHW tornal area (Fig. 2G–J) and unique male genitalia (Fig. 33), with the uncus in dorsal view resembling a guitar. The female genitalia (Fig. 47B) are unique due to the proximal margin of lamella antevaginalis as two rounded lobes.

Fig. 2G, H. Head: Brown with some yellow scales dorsally; white ventrally. FW length: 1.1–1.3 cm. HW length: 0.8–1.0 cm. DFW: Dark brown. Three small faint white dots in the subapical area can be present or absent, the middle one slightly smaller. Fringe paler brown. VFW: Same white markings as on DFW. VHW: Lighter brown. Genitalia (Fig. 3G–J): Uncus longer than wide, guitar-shaped in dorsal view, dorsally bearing a tuft of short hair-like scales. Fenestra developed. Valva: sacculus scalene triangle-shaped, bearing on its dorsal angle a dense set of small, thin and socketed spines; harpe rounded at distal margin, bearing a line of somewhat malleable spines at the distal margin, exceeding ampulla; ampulla broad, rounded at distal margin, bearing medium-sized thin setae at distal margin, ventral-proximal margin with a claw-like process, very pointed and with irregular margins. Aedeagus almost same length as valva, without denticles. Fultura superior bearing two dense clusters of small, thin spines laterally.

Fig. 2I, J. The females differ from the males in the following characters: FW length: 1.2–1.3 cm. HW length: 0.9–1.1 cm. Wings: Two extra dots can be present or absent in M₃-CuA₁ and CuA₁-CuA₂. Genitalia (Fig. 10B): The lamella postvaginalis as a plate very excavated at the center of distal margin, bearing microtrichia, produced at the proximal part as tubular process that almost reaches the lamella antevaginalis. Absence of a membranous or at least less sclerotized area linking the lamellae postvaginalis and antevaginalis. The proximal margin of lamella antevaginalis around the ostium bursae as two rounded lobes, not fused.

Fig. 11. Nicaragua: Rivas (Gauthier and Maes 2018, 2019) and South Caribbean Autonomous Region (Anderson 2007; Erenler et al. 2013); Costa Rica: Alajuela, Cartago, Heredia, Limón, Puntarenas and San José; Panama: Canal Zone, Capira and Colón; Colombia: Cudinamarca and Tolima; Ecuador: Chimborazo (Williams and Bell 1940), Cotopaxi, Napo, Pichincha (Evans 1953) and Santo Domingo de los Tsáchilas.

Staphylus (C.) caribbea is recorded throughout the year.

Although not mentioned in the description, the epithet of Staphylus (C.) caribbea is clearly in reference to the type locality (Panama), which comprises part of the Caribbean region

Before this study, there was no illustration of the female genitalia of Staphylus (C.) caribbea in the literature.

COSTA RICA: No specific locality, 26.VII.1968, 1♂, no collector, OM 36.078* (OM). Alajuela – San Ramón, Bajo Rodriguez, 25.III.1990, 1♂, 1♀, Pagels leg., OM 26.476, OM 26.464 (OM); 1.IV.1990, 1♂, Pagels leg., OM 26.140* (OM); 1.XI.1992. 1♂, no collector, OM 37.887 (OM). Upala, 18.3 Km S., 22.IX.1987, 1♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL). Cartago – No specific locality, Finca la Lola, 6–8.VII.1965, 1♀, J. C. Downey leg., Allyn Museum Acc 1985-14, Genit. Vial No. SRS-5630* (MGCL). Turrialba, 15.VII.1986, 1♀, no collector, OM 36.298 (OM). Heredia – Chilamate, Finca Selva Verde, 24.IX.1986, 2♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL). Puerto Viejo, Finca la Selva, 14.III.1986, 2♂, 1♀, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL); 25.IX.1986, 3♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial SRS-2969* (MGCL); Santa Clara, 3.8 Km N., 5.X.1987, 1♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL). Sarapiquí, La Selva, 50 m, 28.III.1972, 2♂ Becker leg., DZ 46.206*, DZ 46.217 (DZUP). Limón – No specific locality, Rio Victoria, 6.X.1987, 1♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL); 12.IX.1989, 1♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL). Bananito Sur, 5.4 Km SW., 1.IX.1987, 2♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL). Herediana, between Herediana and Francia, Rio Herediana, road to Guapiles, 11.IX.1986, 2♀, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL). Road to Guapiles, 10.III.1986, 2♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL). Germania, 12.IX.1986, 3♂, 1♀, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial SRS-2965*, MGCL 1112078 (MGCL); 15.IV.1989, 1♀, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL); Road to Guapiles, 10.III.1986, 2♂, 2♀, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial SRS-2968*, Genit. Vial SRS-3012* (MGCL). Guapiles, Rio Blanco, 11.III.1986, 1♂, 1♀, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL); 12.IX.1986, 1♀, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL). La Bomba, 2.IX.1987, 3♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial SRS-2980*, MGCL 1112082 (MGCL). Limón, Playa Bananito, 9.III.1986, 2♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL); 13.IX.1986, 1♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL). Puerto Viejo de Talamanca, 13.IX.1986, 2♂, 2♀, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial SRS-3011*, Genit. Vial SRS-3017* (MGCL); 11.IV.1989, 1♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL). Rio Blanco, 4.IX.1987, 2♂, 1♀, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL). Siquirres, 200 m, 27.I.1973, Becker leg., DZ 46.207* (DZUP); Freehold, 8.III.1986, 2♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL); Perla de Siquirres, 4.IV.1985, 3♂, Miguel leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial SRS-2966*, Genit. Vial SRS-2967* (MGCL). Puntarenas – Buenos Aires, 13.IX.1987, 1♀, G. & A. Austin leg., Genit. Vial SRS-3020* (MGCL). La Unión, E of San Vito, 4.IV.1989, 1♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL). San José – Colón, 5 Km S., 17.IX.1987, 1♀, G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL); 26.II.1989, 1♂, no collector, OM 35.221* (OM). — PANAMA: Canal Zone – Madden Forest, 8.IX.1962, 1♀, G. B. Small leg., A. C. Allyn Acc. 1973-23 (MGCL). Farfan [Beach], 12.II.1963, 1♂, S. S. Nicolay leg., A. C. Allyn Acc. 1973-23, MGCL 1112079 (MGCL). Capira – Campana, 12.I.1968, 1♂, Small leg., DZ 45.595* (DZUP). Colón – No specific locality, Frijoles, 20.VIII.1977, 1♂, Robbins leg., DZ 45.181* (DZUP). Soberanía National Park, Gamboa Pipeline Rd., 9.1272N, 79.7148W, 5.II.2014, 1♂, W. H. Russell leg., MGCL Acc #2015-39 W. Russel, GEN. PREP. LEMES 2021-66* (MGCL). — COLOMBIA: Cundinamarca – La Mesa, 1.600 m, 17.XII.2014, 2♂, 1♀, O. Mielke, Callaghan & Henao leg., DZ 45.516, DZ 45.586*, DZ 45.576 (DZUP). Tolima – No specific locality, Rio Chili, 900–1000 m, 7.XII.1975, 1♀, L. M. Steinhauser leg., A. C. Allyn Acc. 1976-3, Genitalia Slide SRS-125* (MGCL); 700–800 m, 21.II.1976, 1♀, S. & L. Steinhauser leg., A. C. Allyn Acc. 1976-3, Genit. Vial SRS-2018* (MGCL). — ECUADOR: Cotopaxi – San Francisco de los Pampas, 1.600 m, 18–19.VII.1987, 3♂, O. Mielke leg., OM 15.528, OM 15.529, OM 15.668* (OM). Napo – Cotundo, IV.1979, Lafebre leg., DZ 45.166* (DZUP). Santo Domingo de los Tsáchilas – La Unión del Toachi, 850 m, 18–19.VII.1987, 13♂, 1 ♀, OM 15.267*, OM 15.676*, OM 15.695, OM 15.696, OM 15.697, OM 15.699, OM 16.068, OM 16.070, OM 16.144, OM 16.145, OM 16.146, OM 16.177, OM 16.188, OM 15.698 (OM). San José de Alluriquin, 1000 m, 18.VII.1981, 2♀, O. Mielke & Casagrande leg., DZ 46.582*, DZ 46.654* (DZUP); 20.VII.1981, 1♀, O. Mielke & Casagrande leg., DZ 45.168 (DZUP); 800 m, 8.II.2001, 10♂, 1♀, O. Mielke & Casagrande leg., OM 52.437, OM 52.458, OM 52.486*, OM 52.549*, OM 52.556, OM 52.570*, OM 52.591, OM 52.612, OM 52.745*, OM 52.850, OM 52.759 (OM). Santo Domingo de los Colorados, 19.VIII.1976, 1♂, no collector, G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-586* (MGCL); 12 Km E., Hotel Tinalandia, 750–850 m, 8.V.1988, 3♂, G. & A. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-524*, Genit. Vial GTA-528* (MGCL); 9.V.1988, 7♂, G. & A. Austin leg., G. T. Austin colln. MGCL Acc. 2004-5, Genit. Vial GTA-536*, Genit. Vial GTA-538*, Genit. Vial GTA-539* (MGCL); 36 Km W., XII.1938, 1♀, R. de Lafebre leg., A. C. Allyn Acc. 1969-2, Genit. Prep. SRS-1384* (MGCL); 36 Km W., Napac, 26.VI.1980, 2♂, 1♀, J. Hyatt leg., G. T. Austin colln. – MGCL Acc. 2004-5, GEN. PREP. LEMES 2021-39*, GEN. PREP. LEMES 2021-38* (MGCL).

Williams and Bell (1940) described Pholisora corumba. Evans (1953) treated this taxon as a subspecies of Staphylus lizeri. Lamas (1994) reinstated the specific status of Staphylus corumba, being followed by most subsequent authors that mentioned this species in taxonomic comments, faunistic studies and catalogues.

Pholisora corumba Williams & Bell, 1940 was described based on the male holotype from Corumbá, Mato Grosso do Sul, Brazil, deposited in ANSP, currently in CMNH, with the following labels: / Type 7183 CORUMBA Will-Bell/ Pholisora corumba Williams + Bell Holotype ♂/ Corumbá/ March/ Genitalia slide No. 1226/ 2457/.

Staphylus (C.) corumba (Fig. 2O–R) is most similar to Staphylus (C.) lizeri lizeri (Fig. 2U–X) but can be differentiated due to the absence of white dots in M₃-CuA₁ and in CuA₁-CuA₂, combined with the ampulla with a very produced dentate area on the internal-ventral margin, with “teeth” of different sizes (Fig. 5B), while in S. (C.) lizeri lizeri, the produced area is not so developed, and the “teeth” are all about same size (Fig. 5J). The female genitalia differ from other species due to the lamella antevaginalis with a central transverse darker carina (Fig. 10C).

Fig. 2O, P. Head: Brown with some yellow scales dorsally; yellow and white ventrally. FW length: 1.0–1.2 cm. HW length: 0.8–0.9 cm. DFW: Dark brown. Two or three white dots can be present or absent in the subapical area. Fringe brown. VFW: Same white markings as on DFW. VHW: Lighter brown. Genitalia (Fig. 5A–D): Uncus longer than wide, with the same width along the entire length, except at the tapered apex, curving ventrally, dorsally bearing a tuft of short hair-like scales at base. Fenestra developed. Valva: sacculus scalene triangular-shaped, bearing thin and socketed spines internally; harpe rounded at distal margin, exceeding ampulla, bearing thin and short setae at distal margin; ampulla broad, rounded at distal margin, not exceeding harpe, bearing a dense set of medium-sized, stout and socketed spines, concave at proximal margin, with a very produced dentate area on the internal-ventral margin, with “teeth” of different sizes. Aedeagus shorter than valva, without denticles. Fultura superior bearing two dense clusters of small, stout spines laterally.

Fig. 2Q, R. The females differ from the males in the following characters: FW length: 1.2–1.4 cm. HW length: 0.9–1.0 cm. Genitalia (Fig. 10C): The lamella postvaginalis as a plate is very excavated at the center of distal margin, bearing microtrichia. Membranous or at least less sclerotized area linking the lamellae postvaginalis and antevaginalis. Lamella antevaginalis as a plate with a central transverse darker carina; the proximal margin not fused with the ostium.

Fig. 12. Brazil: Bahia (Evans 1953), Mato Grosso do Sul (Williams and Bell 1940) and Pará; Peru: Cusco (Lamas 2003), Loreto (Lamas et al. [1997]) and Madre de Dios (Lamas 1994).

Staphylus (C.) corumba is recorded throughout the year, except September and October.

Although not mentioned in the description, the epithet of Staphylus corumba is clearly in reference to the type locality, Corumbá, Mato Grosso do Sul, Brazil.

Before this study, there was no illustration of the female genitalia of Staphylus (C.) corumba in the literature.

BRAZIL: Pará – Santo Antônio do Tauá, Reserva Sonho Azul, 2.XII.2004, 1♂, P. Jauffret leg., DZ 44.977 (DZUP); 10.XII.2004, 1♀, P. Jauffret leg., DZ 44.978* (DZUP); 8.I.2005, 1♂, P. Jauffret leg., DZ 44.979 (DZUP); 14.I.2005, 1♀, P. Jauffret leg., DZ 44.980 (DZUP); 21.I.2005, 1♂, 1♀, P. Jauffret leg., DZ 44.981, DZ 44.982 (DZUP); 26.I.2005, 1♀, P. Jauffret leg., DZ 44.983* (DZUP); 3.II.2005, 1♂, P. Jauffret leg., DZ 44.984* (DZUP); 9.II.2005, 1♀, P. Jauffret leg., DZ 44.985 (DZUP); 13.II.2005, 1♂, P. Jauffret leg., DZ 44.986 (DZUP); 15.II.2005, 1♂, P. Jauffret leg., DZ 44.987 (DZUP); 16.II.2005, 1♂, 1♀, P. Jauffret leg., DZ 44.988, DZ 44.989 (DZUP); 20.II.2005, 1♀, P. Jauffret leg., DZ 44.990 (DZUP); 21.II.2005, 1♂, P. Jauffret leg., DZ 44.991 (DZUP); 22.II.2005, 1♀, P. Jauffret leg., DZ 44.992* (DZUP); 23.II.2005, 1♂, P. Jauffret leg., DZ 44.993 (DZUP); 26.II.2005, 1♂, 1♀, P. Jauffret leg., DZ 17.178*, DZ 44.994 (DZUP); 28.II.2005, 1♀, P. Jauffret leg., DZ 44.995 (DZUP); 2.III.2005, 1♂, P. Jauffret leg., DZ 44.996 (DZUP); 3.III.2005, 1♂, P. Jauffret leg., DZ 44.997 (DZUP); 4.III.2005, 1♀, P. Jauffret leg., DZ 44.998 (DZUP); 5.III.2005, 1♂, 2♀, P. Jauffret leg., DZ 45.000, DZ 44.999, DZ 45.001 (DZUP); 12.III.2005, 1♂, 3♀, P. Jauffret leg., DZ 45.002, DZ 45.003, DZ 45.004, DZ 45.005 (DZUP); 16.III.2005, 1♀, P. Jauffret leg., DZ 45.006 (DZUP); 19.III.2005, 1♂, 1♀, P. Jauffret leg., DZ 45.007, DZ 45.008 (DZUP); 21.III.2005, 1♂, P. Jauffret leg., DZ 45.009 (DZUP); 24.III.2005, 2♂, 1♀, DZ 45.011, DZ 45.012, DZ 45.010 (DZUP); 25.III.2005, 1♂, 3♀, P. Jauffret leg., DZ 45.015, DZ 45.013, DZ 45.014, DZ 45.016 (DZUP); 26.III.2005, 1♂, 1♀, P. Jauffret leg., DZ 45.018, DZ 45.017 (DZUP); 29.III.2005, 2♂, P. Jauffret leg., DZ 45.019*, DZ 45.021, DZ 45.020 (DZUP); 31.III.2005, 1♀, P. Jauffret leg., DZ 45.022 (DZUP); 1.IV.2005, 1♂, 2♀, P. Jauffret leg., DZ 45.023, DZ 45.024, DZ 45.025 (DZUP); 2.IV.2005, 1♂, P. Jauffret leg., DZ 45.026 (DZUP); 11.IV.2005, 1♂, P. Jauffret leg., DZ 45.027 (DZUP); 12.IV.2005, 1♀, P. Jauffret leg., DZ 45.028 (DZUP); 13.IV.2005, 1♀, P. Jauffret leg., DZ 45.029 (DZUP); 17.IV.2005, 1♂, P. Jauffret leg., DZ 45.030* (DZUP); 20.IV.2005, 1♂, P. Jauffret leg., DZ 45.031 (DZUP); 21.IV.2005, 3♂, 4♀, DZ 45.033, DZ 45.037, DZ 45.038, DZ 45.032, DZ 45.034, DZ 45.035, DZ 45.036 (DZUP); 24.IV.2005, 1♂, P. Jauffret leg., DZ 45.039* (DZUP); 28.IV.2005, 1♂, P. Jauffret leg., DZ 45.040 (DZUP); 9.V.2005, 1♂, P. Jauffret leg., DZ 45.041 (DZUP); 10.V.2005, 1♀, P. Jauffret leg., DZ 45.042 (DZUP); 11.V.2005, 1♂, P. Jauffret leg., DZ 45.043* (DZUP); 10.VI.2005, 2♂, P. Jauffret leg., DZ 45.044, DZ 45.045 (DZUP); 11.VI.2005, 1♀, P. Jauffret leg., DZ 45.046 (DZUP); 17.VI.2005, 1♂, P. Jauffret leg., DZ 45.047 (DZUP); 28.VI.2005, 2♂, P. Jauffret leg., DZ 45.048, DZ 45.049 (DZUP); 6.VII.2005, 1♀, P. Jauffret leg., DZ 45.050 (DZUP); 8.VII.2005, 1♂, P. Jauffret leg., DZ 45.051 (DZUP); 9.VII.2005, 1♀, P. Jauffret leg., DZ 45.052 (DZUP): 10.VII.2005, 1♂, 1♀, P. Jauffret leg., DZ 45.054, DZ 45.053 (DZUP); 11.VII.2005, 1♂, 1♀, P. Jauffret leg., DZ 45.055*, DZ 45.056 (DZUP); 14.VII.2005, 1♂, P. Jauffret leg., DZ 45.057 (DZUP); 20.VII.2005, 1♂, P. Jauffret leg., DZ 45.058 (DZUP); 3.VIII.2005, 1♂, P. Jauffret leg., DZ 45.059 (DZUP); 24.VIII.2005, 1♀, P. Jauffret leg., DZ 45.060 (DZUP); 12.XI.2005, 1♂, P. Jauffret leg., DZ 45.061 (DZUP); 27.XI.2005, 1♂, P. Jauffret leg., DZ 45.062 (DZUP); 1.XII.2005, 1♂, P. Jauffret leg., DZ 45.063* (DZUP); 27.XII.2005, 1♂, P. Jauffret leg., DZ 45.064 (DZUP); 22.III.2006, 1♀, P. Jauffret leg., DZ 45.065 (DZUP); 15.V.2006, 1♀, P. Jauffret leg., DZ 45.066 (DZUP); 10.I.2007, 1♂, P. Jauffret leg., DZ 45.067 (DZUP); 15.III.2007, 1♀, P. Jauffret leg., DZ 45.068 (DZUP).

Evans (1953) described Staphylus eryx. According to Evans, Plötz (1884) erroneously cited this species as Nisoniades auricapilla Staudinger. Other authors followed Evans (1953) and mentioned this species in catalogues.

Staphylus eryx was described based on the male [holo]type from Santarém, Amazonas, [Pará, Brazil], H. H. Smith leg., deposited in NHMUK, with the following labels: / eryx Evans [“type” written on the label back]/ SANTAREM H. H. SMITH 1470. 47./ mab n° 17/ Santarem, Amazons, H. H. Smith./ ♂/ Godman-Salvin Coll. 1912.-23./ 436 / [label with genitalia attached]/ BMNH(E) 1669832/.

Staphylus (C.) eryx is differentiated from other species of the Staphylus (Capilla) due to its head with golden scales and unique male genitalia (Fig. 3K–N). The ampulla, at the distal margin, bears short and socketed spines that increase in size from the dorsodistal margin downwards, and a pronounced rounded lobe at the proximal-ventral margin, bearing small spines on the distal margin. The female genitalia (Fig. 10D) are different from other species due to its lamella postvaginalis as a broad plate slightly excavated at the distal margin and with a central longitudinal carina.

Fig. 2K, L. Head: Brown with golden scales dorsally; pure-white ventrally. FW length: 1.1–1.2 cm. HW length: 0.8–1.0 cm. DFW: Brown. Two or three small faint white dots in the subapical area. Two white dots in the dorsal-distal end of cell. One white dot in M₃-CuA₁, other in CuA₁-CuA₂, and two in CuA₂-A2, the upper one smaller. Fringe paler brown. VFW: Same white markings as on DFW. VHW: Lighter brown than DHW. Genitalia (Fig. 3K–N): Uncus oval at the base, dorsally bearing a tuft of medium sized hair-like scales, distally tapered and cylindrical towards the apex. Fenestra developed. Valva: sacculus somewhat quadrate, bearing hair-like scales internally; harpe rounded at distal margin, approximately as long as ampulla, with a flattened, rounded lobe on its dorsal margin overlapping ampulla internally; ampulla broad, rounded at distal margin, bearing a set of dense, short, socketed spines at distal margin that increase in size from the dorsodistal margin downwards, with a pronounced rounded lobe at proximal-ventral margin, bearing small spines on the distal margin of the lobe. Aedeagus approximately the same length as valva, without denticles. Fultura superior without spines.

Fig. 2M, N. The females differ from the males in the following characters: FW length: 1.2 cm. HW length: 0.9 cm. Genitalia (Fig. 10D): The lamella postvaginalis is a broad plate, bearing microtrichia, slightly excavated at the distal margin and with a central longitudinal carina, with proximal margin as funnel without microtrichia. Membranous or at least less sclerotized area linking the lamellae postvaginalis and antevaginalis. Lamella antevaginalis as a plate; the proximal margin reticulated around the ostium bursae and not fused.

Fig. 12. Brazil: Amazonas and Pará.

Staphylus (C.) eryx is recorded only in the following months: May, July and November.

Evans (1953) did not provide an etymology for the name eryx.

Before this study, there was no illustration of the female genitalia of Staphylus (C.) eryx in the literature.

BRAZIL: Amazonas – Not specific locality, Diamantina, 2 Km E. of Maruru, 23.XI.1973, 1♂, C. Callaghan leg., A. C. Allyn Acc. 1974-3 (MGCL). Pará – Santarém, V.1968, 1♂, Kesselring leg., DZ 46.566* (DZUP); 27.VII.1978, 1♂, 1♀, C. Callaghan leg., Allyn Museum Acc. 1985-8, Genit. Vial SRS-1723*, Genit. Vial SRS-1754*, MGCL 1112109 (MGCL); no data, 1♂, 1♀, no collector, ex coll. Le Moult, A. C. Allyn Acc. 1968-1, MGCL 1112109 (MGCL).

O. Mielke (1980) described Staphylus insignis and other authors have mentioned this species in taxonomic comments and catalogues.

Staphylus insignis O. Mielke, 1980 was described based on the male holotype (Fig. 29G, H), from Pelotas, Rio Grande do Sul, Brazil, 28.X.1967, L. M. Biezanko leg., deposited in DZUP, with the following labels: / Holótipo/ Pelotas, 1.X.1955 C. Biezanko leg./ Staphylus insignis Mielke, O Mielke det. 1977 Holótipo/ DZ 1410/ Gen. prep. Mielke 1977/.

Staphylus (C.) insignis can be readily separated from the other species of Staphylus (Capilla) due to a very short ampulla, as a rectangular plate (Fig. 6B). The female genitalia (Fig. 10E) are different from all other species of the subgenus due to the lamella antevaginalis as a somewhat quadrate sclerotized plate and lamella postvaginalis with very sclerotized V-shaped process.

Fig. 4K, L. Head: Brown with some yellow scales dorsally; pale-yellow ventrally. FW length: 1.1 cm. HW length: 0.8 cm. DFW: Brown. Two white dots in subapical area. Fringe brown. VFW: Same white markings as on DFW. VHW: Brown. Genitalia (Fig. 6A–D): Uncus with base semi-rounded, dorsally bearing a tuft of short hair-like scales, tapering towards the apex. Fenestra absent. Valva: sacculus longer than wide, rectangular-shaped, apparently bearing thin, socketed spines internally; harpe rounded at distal margin, exceeding ampulla, bearing a dense set of stout, socketed spines; ampulla very short, as a rectangular plate. Aedeagus approximately the same length as valva, bearing four denticles at the distal portion. Fultura superior bearing two dense clusters of large and stout spines laterally.

Fig. 4M, N. The females differ from the males in the following characters: FW length: 1.3–1.4 cm. HW length: 1.0–1.1 cm. Genitalia (Fig. 10E): The lamella postvaginalis as a large and rectangular, very sclerotized plate, and the lamella antevaginalis as a sclerotized plate fused with the proximal margin of lamella postvaginalis, with a very sclerotized V-shaped process.

Fig. 12. Brazil: Minas Gerais and Rio Grande do Sul.

Staphylus (C.) insignis is only recorded from February–April and October.

Although O. Mielke (1980) did not provide an etymology for insignis, the name refers to “insignificance”, due to the simplicity of the color pattern of the species.

BRAZIL: Minas Gerais – Cambuquira, illegible data, 1♂, Ebert leg., Coleção H. Ebert, DZ 46.667* (DZUP); 7.II.1969, 1♂, Ebert leg., Coleção H. Ebert, DZ 46.660* (DZUP); 15–16.III.1981, 1♂, Ebert leg., Coleção H. Ebert, DZ 46.655* (DZUP); 7.II.1985, 3♂, O. Mielke & Casagrande leg., DZ 46.957*, DZ 46.958*, DZ 46.959* (DZUP). Sabará, 13–18.IV.1981, 2♂, C. Elias leg., DNA Voucher JRAL-COI-84 José R. A. Lemes, DZ 45.250, DZ 45.251* (DZUP). Rio Grande do Sul – Pelotas, 1.X.1955, 1♂, C. Biezanko leg., Holotype, DZ 1.410* (DZUP); 28.X.1977, 1♀, L. M. Biezanko leg., Allotype, DZ 859* (DZUP).

Hayward (1938) described Pholisora lizeri and Williams and Bell (1940) described Pholisora cruza. Evans (1953) synonymized P. cruza with P. lizeri lizeri, and recognized Staphylus lizeri corumba Williams & Bell, 1940 as a subspecies. Steinhauser (1989) described Staphylus lizeri album. Hayward (1940) described Pholisora imperspicua, which Evans (1953) cited as Staphylus imperspicua for the first time. Most subsequent authors followed Evans (1953) when mentioning these taxa in taxonomic comments, faunistic studies and catalogues.

Pholisora lizeri Hayward, 1938 was described based on the male holotype from Caranaví, La Paz, Bolivia, deposited in Hayward collection, currently in IML, with the following labels: / Pholisora lizeri Hayw. Holotipo ♂/ Tipus / Prep. 131 / H. 97/ Caranavi, Bolivia/.

Pholisora cruza Williams & Bell, 1940 was described based on the male holotype from Buenavista, 75 km. northwest of Santa Cruz de la Sierra, Santa Cruz, Bolivia, Steinbach leg., deposited in ANSP, currently in CMNH, with the following labels: / TYPE 7187 CRUZA WILL.-BELL./ Buenavista 75 Km. N. W. of Santa Cruz, Bolivia/ 450 m. alt. 1926–27 (Jose Steinbach)/ Genitalia slide No. 2219/.

Pholisora imperspicua Hayward, 1940 was described based on the male holotype from Rio Topo, Ecuador, Macintyre leg., deposited in Hayward collection, currently in IML, with the following labels: / Ecuador / Rio Topo, Oriente, Ecuador, C. Macintyre / Prep. 455/ Pholisora imperspicua Hayw. Holotype ♂/ Typus/.

Staphylus (C.) lizeri lizeri (Fig. 2U–X) is most similar to Staphylus (C.) corumba (Fig. 2O–R) but can be differentiated due to the absence of white dots in M₃-CuA₁ and CuA₁-CuA₂, combined with the ampulla with a very produced dentate area on the internal-ventral margin, with “teeth” of different sizes (Fig. 5B), while in S. lizeri lizeri the produced area is not so developed, and the “teeth” are about the same size (Fig. 5J). The male genitalia of Staphylus (C.) lizeri lizeri (Fig. 5I–L) is like S. (C.) l. album (Fig. 5E–H), but S. (C.) l. album has a white VHW tornal area (Fig. 2T), while S. (C.) l. lizeri has a brown VHW tornal area (figs 2V, X). The female genitalia (Fig. 10F) differ from other species due to a lamella antevaginalis with a central transverse darker carina with small denticles.

Fig. 2U, V. Head: Brown with some yellow scales dorsally; white with yellow scales ventrally. FW length: 1.2–1.4 cm. HW length: 0.9–1.0 cm. DFW: Brown. Three white dots in the subapical area, one in M₃-CuA₁ and one in CuA₁-CuA₂. Fringe brown. VFW: Same white markings as on DFW. VHW: Lighter brown. Genitalia (Fig. 5I–L): Uncus longer than wide, the same width along the entire length, except at the tapered apex, curving ventrally, dorsally bearing a tuft of short hair-like scales at base. Fenestra developed. Valva: sacculus scalene triangular-shaped, bearing thin, socketed spines internally; harpe rounded at distal margin, exceeding ampulla, bearing thin setae at the distal margin; ampulla broad, rounded at distal margin, not exceeding harpe, bearing a dense set of medium-sized setae distally, proximal margin concave with a produced dentate area on the internal-ventral margin, with the “teeth” about the same size. Aedeagus shorter than valva, without denticles. Fultura superior bearing two dense clusters of small, stout spines laterally.

Fig. 2W, X. The females differ from the males in the following characters: FW length: 1.1–1.3 cm. HW length: 0.8–1.0 cm. Genitalia (Fig. 10F): The lamella postvaginalis is a very excavated plate at the center of the distal margin, bearing microtrichia. Membranous or at least less sclerotized area linking the lamellae postvaginalis and antevaginalis. Lamella antevaginalis as a plate with a central transverse darker carina with small denticles; the proximal margin reticulated and fused; laterally with external margins usually as pointed projections.

Fig. 14. Colombia: Atlántico (Murillo et al. 2018), Meta and Valle del Cauca; Brazil: Acre, Amazonas, Pará and Rondônia; Ecuador: Carchi, Chimborazo, Napo, Orellana, Pichincha and Sucumbíos; Peru: Amazonas, Callao (Evans 1953), Carabaya (Evans 1953), Chanchamayo (Evans 1953), Cusco, Huánuco, Junín, Madre de Dios, Maynas (Evans 1953), Mariscal (Evans 1953), Moyobamba (Evans 1953), Oxapampa (Williams and Bell 1940), Puno, Quispicanchi (Evans 1953), San Martin, Sandia (Evans 1953), Satipo and Ucayali; Bolivia: Cercado (Evans 1953), José Ballivián (Evans 1953), La Paz, Santa Cruz and Yungas (Evans 1953).

Staphylus (C.) lizeri lizeri is recorded throughout the year.

According to Hayward (1938), Staphylus (C.) lizeri was named in honor of his friend and colleague Carlos A. Lizer y Trelles. Although Williams and Bell (1940) did not provide an etymology for Pholisora cruza, they were clearly making reference to Santa Cruz de la Sierra, Santa Cruz, Bolivia, the type locality. Hayward (1940) did not provide an etymology for the name imperspicua.

The illustration of the male genitalia provided by Williams and Bell (1940) of Pholisora cruza is very similar to the genitalia of Staphylus lizeri. Also, the authors mentioned that the ventral wings are brown. Considering that S. (C.) lizeri lizeri also occurs in Bolivia, Pholisora cruza is herein confirmed as a synonym of Staphylus (C.) lizeri lizeri. The male genitalia of the holotype of Pholisora imperspicua is identical to specimens of S. (C.) lizeri lizeri, and the type locality (Ecuador) also corresponds to recorded specimens of S. (C.) lizeri lizeri. Therefore, aiming to stabilize the taxonomy, Pholisora imperspicua is herein considered a new synonym of S. (C.) l. lizeri. Before this study, there was no illustration of the female genitalia of Staphylus (C.) lizeri lizeri in the literature.

COLOMBIA: Meta – Villavicencio, 600 m, 14.V.1972, 2♂, DZ 46.807*, DZ 46.808 (DZUP); Bosque Bavaria, 12.XII.2014, 9♂, 2♀, O. Mielke, Carneiro & Henao leg., DZ 45.256, DZ 45.266, DZ 45.286, DZ 45.296*, DZ 45.316, DZ 45.326*, DZ 45.784, DZ 45.814, DZ 45.824, DZ 45.276*, DZ 45.346 (DZUP); 14.XII.2014, 5♂, 5♀, O. Mielke, Carneiro & Henao leg., DZ 45.366, DZ 45.396, DZ 45.426, DZ 45.496 (in copula), DZ 45.566, DZ 45.486, DZ 45.506 (in copula), DZ 45.526, DZ 45.536*, DZ 45.546 (DZUP); La Cuncia, 13.XII.2014, 5♂, 2♀, O. Mielke, Carneiro & Henao leg., DZ 45.406, DZ 45.416, DZ 45.436*, DZ 45.446, DZ 45.466*, DZ 45.356, DZ 45.476 (DZUP); La Únion, 4°03’43.3 N, 73°42’04.6 W, 500 m, 12.X.2014, 1♀, G. Nielsen leg., DZ 45.376 (DZUP); 25.XI.2014, 1♀, G. Nielsen leg., DZ 45.774 (DZUP). Valle del Cauca – Buenaventura, 250 m, 16.VII.1981, 1♀, O. Mielke & Casagrande leg., DZ 46.668 (DZUP). — BRAZIL: Acre – Bujari, 50 Km NO, 200 m, 18–20.IX.2003, 1♂, O. -C. Mielke & Casagrande leg., DZ 45.625 (DZUP); 20–23.VII.2004, 2♂, 1♀, O.-C. Mielke leg., OM 65.746*, OM 64.672, OM 65.823 (OM). Cruzeiro do Sul, Rio Juruá, 200 m, no data, 2♂, H. Ebert leg., Coleção H. Ebert, DZ 46.671, DZ 46.672 (DZUP); III.1975, 1♂, H. Ebert leg., Coleção H. Ebert, DZ 46.676 (DZUP); 9–10.VII.1975, 1♀, H. Ebert leg., Coleção H. Ebert, DZ 46.673 (DZUP); 10–15.II.1976, 2♂, 1♀, H. Ebert leg., Coleção H. Ebert, DZ 46.670*, DZ 46.675, DZ 46.669 (DZUP); 28.IX.1977, 1♂ H. Ebert leg., Coleção H. Ebert, DZ 46.674* (DZUP). Porto Acre, Reserva Humaitá, 200 m, 16.VII.2004, 2♂, 1♀, O.-C. Mielke leg., OM 64.980, OM 65.345, OM 65.359 (OM). Santa Rosa do Purus, 6,4 Km E, 3–5.VIII.2008, 2♂, O. Mielke & Carneiro leg., DZ 15.624*, DZ 16.025*; 11,9 Km SE, 11.VIII.2008, O. Mielke & Carneiro leg., DZ 16.109* (DZUP). Amazonas – Fonte Boa, 2°33’22.23"S, 66°03’56.72"W, 80 m, 11–23.VIII.2011, 1♂, 1♀, Eduardo de Oliveira Emery leg., Ex Coleção Eduardo Emery, DZ 46.903* (DZUP). São Paulo de Olivença, no data, 1♂, no collector, OM 8.586* (OM). Pará – Belém, Parque Estadual do Utinga, 4–6.VIII.1992, 1♂, O. Mielke & Casagrande leg., OM 30.218 (OM). Benevides, Neópolis, 1–3.VIII.1992, 2♂, O. Mielke & Casagrande leg., OM 32.430, OM 32.448* (OM); 19.IV.1993, 1♂, O. Mielke leg., OM 34.403* (OM). Marabá, 8.I.1961, 1♂, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20 (MGCL). Rondônia – Cacaulândia, Fazenda Cerejeira, 18.VII–I.1996, 4♂, 1♀, O. Mielke & Miers leg., OM 42.681, OM 42.729*, OM 42.897, OM 42.889*, OM 42.689 (OM); Fazenda Rancho Grande, 7.X.1987, 1♂, 1♀, C. Mielke leg., OM 15.965*, OM 15.967 (OM); 17.III.1989, 1♂, G. T. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-251* (MGCL); 17–20.III.1989, 10♂, 12♀, O. Mielke leg., OM 20.448, OM 20.449, OM 20.450*, OM 20.903, OM 21.454, OM 22.309*, OM 22.315*, OM 22.321*, OM 22.327*, OM 22.345*, OM 20.709, OM 21.404, OM 21.425, OM 21.426, OM 21.430, OM 21.440, OM 21.451, OM 21.476, OM 21.500, OM 22.285, OM 22.291, OM 22.303 (OM); 20.III.1989, 1♂, G. T. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-261* (MGCL); 24.III.1989, 1♂, G. T. Austin leg., G. T. Austin colln. MGCL Acc. 2004-5, Genit. Vial GTA-263* (MGCL); 21.X.1989, 1♂, G. T. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-254* (MGCL); 25.X.1989, 1♂, G. T. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-684 (MGCL); 27.X.1989, 5♂, G. T. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-256*, Genit. Vial GTA-258*, Genit. Vial GTA-265*, Genit. Vial GTA-266*, Genit. Vial GTA-300* (MGCL); 28.X.1989, 1♂, G. T. Austin leg., G. T. Austin colln. MGCL Acc. 2004-5, Genit. Vial GTA-262* (MGCL); 30.X.1989, 1♂, G. T. Austin leg., G. T. Austin colln. MGCL Acc. 2004-5, Genit. Vial GTA-560* (MGCL); 2.XI.1989, 1♂, G. T. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-257* (MGCL); 5.XI.1989, 1♂, G. T. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-438* (MGCL); 6.XI.1989, 1♂, G. T. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-267* (MGCL); 2.IX.1990, 3♂, G. T. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-2107*, Genit. Vial GTA-2108*, Genit. Vial GTA-2113* (MGCL); 3.IX.1990, 1♂, G. T. Austin leg., G. T. Austin colln. MGCL Acc. 2004-5, Genit. Vial GTA-2117* (MGCL); 6.IX.1990, 1♂, G. T. Austin leg., G. T. Austin colln. MGCL Acc. 2004-5, Genit. Vial GTA-2114* (MGCL); 31.X.1990, 2♂, G. T. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-1360*, Genit. Vial GTA-1362* (MGCL); 6.XI.1990, 1♂, G. T. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-2115* (MGCL); 9.XI.1990, 1♂, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20, GEN. PREP. (MGCL); 10.XI.1990, 2♂, 1♀, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20 (MGCL); 14.XI.1990, 1♂, 1♀, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20 (MGCL); 17.XI.1990, 1♀, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20 (MGCL); 8.XII.1990, 1♂, G. T. Austin leg., G. T. Austin colln. MGCL Acc. 2004-5, Genit. Vial GTA-1008* (MGCL); 11.XII.1990, 1♂, G. T. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-998* (MGCL); 12.XII.1990, 1♂, G. T. Austin leg., Austin coll., Allyn Mus. Acc. 1991-4, Genit. Vial GTA-1010* (MGCL); 14.XII.1990, 1♂, G. T. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-1011* (MGCL); 15.XII.1990, 1♂, G. T. Austin leg., G. T. Austin colln. MGCL Acc. 2004-5, Genit. Vial GTA-1009* (MGCL); 16.III.1991, 1♂, G. T. Austin leg, G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-1361* (MGCL); 10-22.VII.1991, 1♂, C. Mielke & Miers leg., OM 27.380 (OM); 17.IX.1991, 1♂, T. Schmitz leg., G. T. Austin colln. MGCL Acc. 2004-5, Genit. Vial GTA-2116* (MGCL); 12.XI.1991, 1♀, J. & F. Preston leg., G. T. Austin colln. MGCL Acc. 2004-5 (MGCL); 1♀, G. T. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL); 14.XI.1991, 1♂, 1♀, O. Mielke leg., OM 27.472*, OM 27.442 (OM); 15.XI.1991, 1♀, O. Mielke leg., OM 27.535 (OM); 17.XI.1991, 2♂, T. Schmitz leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL); 18.XI.1991, 1♂, O. Mielke leg., OM 27.697 (OM); 19.XI.1991, 2♂, G. T. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-2109*, (MGCL); 1♂, O. Mielke leg., OM 27.783 (OM); 27.XI.1991, 1♂, O. Mielke leg., OM 28.025* (OM); XI–XII.1991, 1♂, S. Kohler leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-3116* (MGCL); 21.IV.1992, 1♂, G. T. Austin leg., G. T. Austin colln. MGCL Acc. 2004-5 (MGCL); 23.IV.1992, 1♂, L. Harris leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL); 21.VIII.1992, 1♂, 1♀, O. Gomes leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL); 25.IX.1992, 1♂, G. T. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-3117* (MGCL); 10.VI.1993, 2♀, no collector, G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL); 10.IV.1994, 1♂, O. Gomes leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL); 8–19.XI.1994, 3♂, 5♀, O. Mielke leg., OM 36.346, OM 38.541*, OM 39.257*, OM 37.596, OM 38.366, OM 38.488, OM 38.638, OM 39.137 (OM); 9–19.XI.1994, 1♂, O. Mielke leg., OM 38.074* (OM); 13.XI.1994, 2♂, G. T. Austin leg., G. T. Austin colln. – MGCL Acc. 2004-5, Genit. Vial GTA-6625*, Genit. Vial GTA-6626* (MGCL); 3.V.1995, 1♀, G. T. Austin leg., G. T. Austin colln. MGCL Acc. 2004-5, associated with Eciton burchelli, Genit. Vial GTA-7568* (MGCL); 14.V.1995, 1♂, O. Gomes leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL); 8.VII.1995, 2♀, O. Gomes leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL); 30.VII.1995, 1♂, O. Gomes leg., G. T. Austin colln. MGCL Acc. 2004-5 (MGCL); 9.IX.1995, 1♀, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20 (MGCL); 8–19.XI.1995, 1♂, 1♀, O. Mielke leg., OM 38.641*, OM 38.134 (OM); 9.XI.1995, 3♂, D. & J. Lindsley leg., D. L. Lindsley colln. MGCL Accession #2008-20 (MGCL); 10.XI.1995, 1♂, 1♀, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20 (MGCL); 12.XI.1995, 3♂, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20, GEN. PREP.* (MGCL); 17.XI.1995, 1♂, D. & J. Lindsley leg., D. L. Lindsley colln. MGCL Accession #2008-20 (MGCL); 8–19.XI.1996, 2♂, 2♀, O. Mielke leg., OM 39.095*, OM 39.239*, OM 39.035, OM 39.185 (OM); 9–19.XI.1996, 1♀, O. Mielke leg., OM 39.221 (OM); 7–10.XII.1996, 5♂, 4♀, O. Mielke leg., OM 44.233, OM 44.249, OM 44.265, OM 44.338, OM 44.346, OM 44.241, OM 44.257, OM 44.273*, OM 44.298 (OM); 19–20.XII.1996, 1♂, 1♀, O. Mielke & Miers leg., OM 43.288, OM 43.471 (OM); 1.III.1997, 1♂, O. Gomes leg., G. T. Austin colln. – MGCL Acc. 2004-5 (MGCL). — ECUADOR: No specific locality, Oriente, Sadzayacu, XI.1968, 1♂, R. de Lafebre leg., A. C. Allyn Acc. 1969-11, Genit. Vial SRS-2039* (MGCL). Carchi – No specific locality, Rio Ambi, 23.I.1979, 650 m, 1♂, M. Simon leg., Allyn Museum Acc. 1982-20, Genit. Prep. SRS-1499* (MGCL). Chimborazo – Pallatanga, 3200 m, VIII.1979, 1♂, 3♀, Lafebre leg., DZ 46.539, DZ 46.537, DZ 46.538, DZ 46.540 (DZUP). Napo – No specific locality, Pusuno, Rio Napo, 10.IX.1996, 4♂, 2♀, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20, GEN. PREP.*, GEN. PREP.* (MGCL). Archidona, 20.VII.1987, 1♂, O. Mielke leg., OM 15.470* (OM). Puerto Misahuallí, Rio Napo, 11.IX.1996, 1♂, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20 (MGCL); 5.IX.2000, 1♀, John J. Bowe leg., FSCA Florida State Collection of Arthropods (MGCL). Tena, 21.VII.1986, 1♂, Lafebre leg., OM 15.882* (OM). Zona Camping Cascada de Latas, II.1971, 4♂, 1♀, R. de Lafebre leg., A. C. Allyn Acc. 1971-12, Genit. Prep. SRS-1372*, Genit. Prep. SRS-1373*, Genit. Prep. SRS-1377* (MGCL). Yasuni National Park, Yasuni Scientific Research Station, Rios Tivacuno & Tiputini, 0°38’S, 76°36’W, 250 m, 18.X.1998, 1♂, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20, GEN. PREP.* (MGCL); 19.X.1998, 2♂, 1♀, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20 (MGCL); 23.X1998, 1♂, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20 (MGCL); 00°40’26.7"S, 76°23’53.7"W, 260–300 m, 30.VIII.2000, 1♂, Peter J. Eliazar leg. (MGCL). Orellana – Coca, Rio Napo, 850’, 23.VI.1980, 2♂, Charles M. Stevens leg., FSCA Florida State Collection of Arthropods (MGCL). Pichincha – Quito, Rio Pilatón, XI.1970, 1♂, R. de Lafebre leg., A. C. Allyn Acc. 1970-44 (MGCL). San Juan, 3200 m, V.1980, 1♂, Lafebre leg., DZ 46.541* (DZUP). Pichincha [=Santo Domingo de los Tsáchilas] – Alluriguin [=Alluriquín], La Palma [=Las Palmas], 1000 m, 20.VII.1981, 1♂, O. Mielke & Casagrande leg., DZ 46.584 (DZUP). Tinalandia, 18.IX.1996, 1♂, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20, GEN. PREP.* (MGCL); 21.IX.1996, 2♂, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20 (MGCL). Santo Domingo de los Colorados, 900 m, V.1969, 1♀, R. de Lafebre leg., A. C. Allyn Acc. 1969-11 (MGCL). Sucumbíos – Reserva Biológica Limóncocha, 2.IX.1972, 2♂, T. C. Emmel leg., FSCA Florida State Collection of Arthropods, GEN. PREP. LEMES 2021-37* (MGCL); 5.IX.1972, 1♂, 2♀, T. C. Emmel leg., FSCA Florida State Collection of Arthropods (MGCL); 6.IX.1972, 2♂, T. C. Emmel leg., FSCA Florida State Collection of Arthropods (MGCL); 9.IX.1972, 1♂, T. C. Emmel leg., FSCA Florida State Collection of Arthropods (MGCL); 24–27.VII.1980, 1♀, C. V. Covell Jr. leg., C. V. Covell colln. – MGCL-Acc. 2005-7 (MGCL); Rio Napo, 24.VI.1980, 1♂, Jim P. Brock leg., MGCL Acc. #2015-37 J. Brock (MGCL). — PERU: Amazonas – Santa Maria de Nieva, 4°35’58" S, 77°51’40" W, 200 m, X.2016, 1♂, J. Pintado leg., OM 81.452* (OM). Cusco – Cosñipata Valley, Chontachaca, 12°56’ S, 71°24’ W, 950 m, 1.IV.2016, 2♀, O. Mielke leg., OM 79.640, OM 79.680 (OM). Patria, 675 m, 25.IV.2015, 1♂, O. Mielke leg., OM 77.863* (OM). Pilcopata, 2°54’ S, 71°22’ W, 600 m, 7.XI.2012, 5♂, 2♀, O. Mielke leg., OM 72.980, OM 72.970*, OM 73.040*, OM 73.160, OM 73.410, OM 73.090, OM 73.110* (OM); Villa Carmen, 12°54’ S, 71°24’ W, 540 m, 8♂, 3♀, O. Mielke leg., OM 74.257, OM 74.317, OM 74.408*, OM 74.428, OM 74.438*, OM 74.448, OM 74.458, OM 74.387, OM 74.267, OM 74.277, OM 74.418 (OM). Quebrada Quitacalzón, 13°01’S, 71°30’W, 1.100 m, 27.X.2018, 1♂, O. Mielke leg., OM 87.847 (OM). Quebrada San Pedro, 13°03’ S, 71°30’ W, 1.100 m, 27.I.2013, 1♂, O. Mielke leg., OM 74.388* (OM); 1.400 m, 1–3.II.2013, 1♂, O. Mielke leg., OM 74.398* (OM); 4.II.2013, 12, O. Mielke leg., OM 74.337*, OM 74.357* (OM); 5.II.2013, 1♂, 1♀, O. Mielke leg., OM 74.297, OM 74.347 (OM). Santa Isabel, 13°02’S, 71°31’W, 1.200 m, 6.XI.2018, 1♂, O. Mielke leg., OM 87.647 (OM). Quispicanchi, Quince Mil, VIII.1962, 2♂ Peña leg., OM 29.967*, OM 29.997* (OM). Vale Rio Cosñipata, San Pedro, 1.300–1.500 m, 5–10.XI.2001, 2♂, O. Mielke leg., OM 57.954*, OM 57.963 (OM). Huánuco – Tingo Maria, XII.1981, 2♂, Schunke leg., DZ 45.515*, DZ 46.993* (DZUP); X.1996, 1♂, M. Büche leg., OM 47.382* (OM); 4 Km N., Parque National las Cuevas de la Lechuza, 24.XI.1964, 2♂, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20, GEN. PREP.*, GEN. PREP.*(MGCL); 25.XI.1964, 1♂, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20, GEN. PREP.* (MGCL); 28.III.1981, 3♂, J. Y. Miller leg., Allyn Museum Acc. 1981-12, Genit. Prep. SRS-1459* (MGCL); 30.III.1981, 2♂, J. Y. Miller leg., Allyn Museum Acc. 1981-12, Genit. Vial SRS-1462*, Genit. Vial SRS-1991* (MGCL); 6 Km S., Cueva de las Pavas, 9°22’24.9"S, 75°58’29.0"W, 21.VI.2001, 10♂, 3♀, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20 (MGCL); IV.2016, 1♂, J. J. Ramirez H. leg., OM 81.152* (OM); 12 Km S., Santa Rosa de Queseda, 9°22’45.6"S, 75°57’15.8"W, 2700’, 18.VI.2001, 13♂, 1♀, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20 (MGCL); 13 Km S., Tres de Mayo, 23.VI.2001, 1♂ D. L. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20, GEN. PREP.* (MGCL); 19.VI.2001, 9♂, 1♀, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20 (MGCL); 19 Km S, Puente Chincavita, 9°30’46.5"S, 75°56’37.4"W, 17.VI.2001, 11♂, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20 (MGCL); 19 Km S., Puente Cayumbe, Rio Derrepente, 9°29’37.7"S, 75°57’31.2"W, 2900’, 20.VI.2001, 2♂, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20 (MGCL); Las Palmas, 26.XI.1964, 1♂, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20, GEN. PREP.* (MGCL); 22.VII.1981, 2♂, O. Mielke & Casagrande leg., DZ 46.652*, DZ 46.653* (DZUP); Rondos, 650 m, 23.VII.1981, 2♂, O. Mielke & Casagrande leg., DZ 45.178, DZ 45.179 (DZUP). Junín – La Merced, 10 Km W Foz Rio Colorado, 1000 m, 20.X.1989, 7♂, O. Mielke & Casagrande leg., OM 23.012, OM 23.018*, OM 23.034, OM 23.125, OM 23.131*, OM 23.155*, OM 23.161* (OM). San Ramón, Hacienda Naranjal, 15–18.X.1989, 12♂, 1♀, O. Mielke & Casagrande leg., OM 23.021*, OM 23.026*, OM 23.097*, OM 23.103, OM 23.115, OM 23.152, OM 23.164, OM 24.640*, OM 24.688*, OM 24.951*, OM 24.957*, OM 24.975, OM 24.694 (OM); 21.X.1989, 2♂, O. Mielke & Casagrande leg., OM 23.173, OM 23.197 (OM); Rio Tulumayo, 1100 m, 5 Km SE Vítoc, 19.X.1989, 4♂, O. Mielke & Casagrande leg., OM 23.560, OM 23.679*, OM 23.691, OM 23.697 (OM). Madre de Dios – No specific locality, Rio Alto Madre de Dios, ca. 15 river km, 420 m, 5.VII.1980, 2♂, J. F. Douglass leg., Allyn Museum Acc. 1980-19, Genit. Prep. SRS-1296* (MGCL); 8.VII.1980, 1♂, 1♀, J. F. Douglass leg., Allyn Museum Acc. 1980-19, Genit. Prep. SRS-1298* (MGCL). Amozonia Lodge, 12°52’S, 71°22’W, 500 m, 23–28.X.2013, 3♂, 1♀, O. Mielke leg., OM 75.625, OM 75.818*, OM 75.878, OM 76.581* (OM); 29.IV–1.V.2015, 3♂, O. Mielke leg., OM 77.415*, OM 77.445, OM 77.637* (OM). Puerto Maldonado, 300 m, 14–15.X.1983, 1♂, O. Mielke & Casagrande leg., DZ 45.385*; 0–2 Km W., 250 m, 6.VIII.1981, 1♂, Lee D. Miller leg., Aronheim Exp. Allyn Museum Acc. 1982-1 (MGCL); 7.VIII.1981, 2♂, Lee D. Miller leg., Aronheim Exp. Allyn Museum Acc. 1982-1, Genit. Vial SRS-1304* (MGCL); 8.VIII.1981, 1♂, Lee D. Miller leg., Aronheim Exp. Allyn Museum Acc. 1982-1 (MGCL); 14.VIII.1981, 2♀, Lee D. Miller leg., Aronheim Exp. Allyn Museum Acc. 1982-1 (MGCL); 0–3 Km SW, Rio Tambopata, 250 m, 4.VIII.1981, 1♀, Lee D. Miller leg., Aronheim Exp. Allyn Museum Acc. 1982-1 (MGCL); 18 Km NE, Rio Madre de Dios, 250 m, 11.VIII.1981, 1♂, Lee D. Miller leg., Aronheim Exp. Allyn Museum Acc. 1982-1, Genit. Vial SRS-2787* (MGCL); Comunidad Infierno, 300 m, 16–17.X.1983, 2♂, O. Mielke & Casagrande leg., DZ 45.435*, DZ 45.465 (DZUP). Shintuya, Cerro Pantiacolla, 460 m, 16.VII.1980, 1♂, J. F. Douglass leg., Allyn Museum Acc. 1980-19 (MGCL); 3.3 Km NE Albergue Pantiacolla, 12°39’21"S, 71°13’53"W, 400 m, 4–8.XI.2017, 1♂, O. Mielke leg., OM 85.566* (OM); 30.X–2.XI.2018, 1♂, O. Mielke leg., OM 88.037* (OM). Tambopata National Reserve, 12°50’S, 69°17’W, 300 m, 25.X.1991, 1♂, R. Robbins leg., DZ 46.176 (DZUP); 1♂, M. Casagrande leg., DZ 46.177 (DZUP); 26.X.1991, 1♂, O. Mielke leg., DZ 46.178 (DZUP); 1♂, R. Robbins leg., DZ 46.179 (DZUP); 27.X.1991, 4♂, O. Mielke leg., DZ 46.181, DZ 46.182, DZ 46.183, DZ 46.184 (DZUP); 1♂, R. Robbins leg., DZ 46.180 (DZUP); 28.X.1991, 4♂, O. Mielke leg., DZ 46.185, DZ 46.186, DZ 46.187, DZ 46.188 (DZUP); 29.X.1991, 14♂, 4♀, O. Mielke leg., DZ 9.667*, DZ 9.668, DZ 46.189, DZ 46.190, DZ 46.191, DZ 46.192, DZ 46.193, DZ 46.194, DZ 46.195, DZ 46.196, DZ 46.197, DZ 46.198, DZ 46.199, DZ 46.200, DZ 45.265, DZ 45.275*, DZ 45.285, DZ 45.295 (DZUP). Puno – Sandia, 800 m, XII.2014, 9♂, 2♀, C. Tello leg., OM 76.967*, OM 76.997*, OM 77.007*, OM 77.048, OM 77.058*, OM 77.107, OM 77.137*, OM 77.147*, OM 77.197, OM 77.117, OM 77.227 (OM). San Martin – Huingoyacu, 300 m, 8.III.2012, 1♂, P. Calderón leg., OM 71.279* (OM). Juanjui, 300 m, 21.XII.2002, 1♂, no collector, OM 63.695* (OM); Rio Huayabamba, 400 m, II.2007, 2♂, J. A. Tafur Novoa leg., OM 68.471*, OM 68.632* (OM). Moyobamba, 800 m, XII.2014, 4♂, C. Tello leg., OM 76.796*, OM 76.816*, OM 76.826, OM 76.846 (OM); 27.X.2015, 2♂, O. Mielke leg., OM 79.221*, OM 79.281* (OM). Rioja, El Afluente, 5°39’48" S, 77°41’41" W, 1.300 m, IV.2017, 3♂, J. Pintado leg., OM 86.124*, OM 83.189, OM 86.246* (OM); Nascientes del Rio Negro, 06°05’ S, 77°16’ W, 900 m, 29.X.2015, 2♂, 1♀, O. Mielke leg., OM 79.241*, OM 79.251, OM 79.291 (OM); San Agostin, 05°44’34" S, 77°30’42" W, 1.050 m, 26.X.2015, 2♂, O. Mielke leg., OM 79.101*, OM 79.211 (OM). Shapaja, 2 Km L, 400 m, 21.X.2015, 1♂, O. Mielke leg., OM 79.181* (OM); 23.X.2015, 2♂, O. Mielke leg., OM 79.021*, OM 79.081* (OM). Satipo – Satipo, I.1982, 2♂, 1♀, Schunke leg., DZ 45.495*, DZ 45.565*, DZ 45.555* (DZUP). Ucayali – Atalaya, 1 Km NE, 12°52’ S, 71°22’ W, 500 m, 7–11.XI.2012, 4♂, 1♀, O. Mielke leg., OM 73.150, OM 73.170, OM 73.390, OM 73.020, OM 73.010* (OM). — BOLIVIA: La Paz – Caranavi, VIII.1989, 2♂, C. Tello leg., OM 24.209*, OM 24.229* (OM); XI.2004, 3♂, C. Tello leg., OM 66.410*, OM 66.567* OM 66.745* (OM); I.2005, 2♂, C. Tello leg., OM 66.830*, OM 66.851* (OM); 15 Km NE., 5.III.2000, 15°47’43.7"S, 67°30’49.9"W, 2500’, 2♀, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20 (MGCL). Rio Selva Resort, 16°12’14.2"S, 67°47’38.9"W, 2500’, 8.III.2000, 2♂, 1♀, GEN. PREP* (MGCL); 9.III.2000, 4♂, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20, GEN. PREP.* (MGCL); 11.III.2000, 1♂, D. & J. Lindsley leg., D. L. Lindsley colln. – MGCL Accession #2008-20 (MGCL).

Staphylus lizeri album Steinhauser, 1989 was described based on the male holotype from Sete Lagoas, Minas Gerais, Brazil, deposited in DZUP, with the following labels: / HOLOTYPE ♂ Staphylus lizeri album S. R. Steinhauser/ Sete Lagoas, MG BRASIL – 720 m 8.III.1974 V. O. Becker col./ Allyn Museum Photo No. 880729B/21,22/ Genit. Vial SRS-2021/ Allyn Museum Acc. 1977-4/.

Staphylus (C.) l. album (Fig. 2S, T) externally resembles some specimens of S. (C.) melangon melangon (Fig. 4E–H), S. (C.) melangon stanga (Fig. 4I, J) and S. (C.) neideae sp. nov. (Fig. 7U–X) due to the tornal white area on VHW. Among these species, S. (C.) l. album has a unique male genitalia morphology (Fig. 5E–H), with a tubular uncus, harpe exceeding ampulla and without a produced expansion at the distodorsal margin. The male genitalia of S. (C.) l. album is like that of S. l. lizeri (Fig. 5I–L), but the tornal VHW area of the latter is brown (Fig. 2V–X).

(Fig. 2S, T). Similar to Staphylus (C.) lizeri lizeri, but lacking the white dots in M₃-CuA₁ and in CuA₁-CuA₂, and with the VHW tornal area white. FW length: 1.2–1.4 cm. HW length: 0.8–1.1 cm.

Unknown.

Fig. 14. Brazil: Minas Gerais.

Staphylus (C.) lizeri album is recorded from January–February, May and October–November.

Although not mentioned in the description, the epithet of Staphylus (C.) lizeri album is clearly in reference to the shining white underside the tornal area of the hindwing.

BRAZIL: Minas Gerais – Carmo do Rio Claro, 9.II.1959, 1♂, O. Mielke leg., OM 2.363 (OM); 20.II.1959, 1♂, O. Mielke leg., OM 2.788 (OM); 18.XI.1959, 1♂, O. Mielke leg., OM 2.789* (OM); 20.XI.1959, 2♂, O. Mielke leg., OM 2.787*, OM 2.790 (OM). Matosinhos, Fazenda Jaguara, 18.V.1995, 1♂, O. Mielke & A. Machado leg., OM 39.643* (OM); 6.I.2002, 1♂, O.-C. Mielke leg., OM 55.435 (OM). Paraopeba, 750 m, 11.X.1969, 1♂, H. Ebert leg., Coleção H. Ebert, DZ 46.656* (DZUP).

Mabille (1878) described epicaste without a genus, consequently a nomen nudum, and in 1883 correctly described it as Hesperia melangon, which was cited by different authors in the genera Staphylus and Pholisora. Later, Mabille (1903) cited the name epicaste in the genus Staphylus. Hayward (1937) described Pholisora rileyi, which was cited in the genus Staphylus by different authors. Evans (1953) synonymized P. rileyi and Staphylus punctiseparatus Hayward, 1933 with Staphylus epicaste Mabille, 1903. Evans (1953) proposed subspecies for this taxon: Staphylus epicaste melangon, S. e. epicaste, and the newly described Staphylus e. stanga. As the name epicaste was described without a genus name, violating the Art. 11.9.3 of ICZN, O. Mielke and Casagrande (1998) corrected the subspecific names, with melangon, epicaste and stanga considered subspecies of Staphylus melangon. Other authors mentioned this subspecies in taxonomic comments, faunistic studies and catalogues.