Incisus incisus ( Mabille, 1878 )

Lemes, José Ricardo Assmann, Siewert, Ricardo Russo, Mielke, Olaf Hermann Hendrik, Casagrande, Mirna Martins & Warren, Andrew David, 2024, Taxonomic revision of the Neotropical skipper genus Incisus Grishin, 2019 (Lepidoptera, Hesperiidae, Pyrginae), with the description of a new species, Zootaxa 5437 (1), pp. 64-86 : 75-78

Incisus incisus ( Mabille, 1878)

( Figs 1 , 2G–L , 3I–N , 4I–L , 5C , 6 )

Antigonus incisus Mabille, 1878 . Pet. Nouv. Ent. 2 (188): 198; Pará, Brazil [Brasilia on the label of the lectotype —herein designated] and Cayenne, [ French Guiana].—Shepard, 1934. Lep. Cat. 64, p. 183; syn.: aterea .—Shepard, 1936. Lep. Cat. 74, p. 628.

Nisoniades aterea Plötz, 1884 (Herrich-Schäffer in litt.). Jahrb. nass. Ver. Nat., Wiesbaden, 37: 39; Rio [de Janeiro, Brazil].

Systasea incisa [sic]; Godman, 1907. Ann. & Mag. Nat. Hist. (7)20: 149; syn.: aterea .—Draudt, 1922, in Seitz. Gross-Schmett. Erde 5, p. 904, pl. 176c (d); syn.: aserea .— Spitz, 1932. Rev. Mus. Paulista 17: 872.—F. Hoffmann, 1934. Ent. Rdsch., Stuttgart, 51 (3): 63.

Systasea aserea [sic]; Draudt, 1922, in Seitz. Gross-Schmett. Erde 5, p. 904; as synonym of Systasea incisa .

Staphylus incisus ; Evans, 1953. Cat. Amer. Hesp. 3, p. 95, pl. 38 (male gen.); syn.: variegatus , aterea , dentivittata , ordinarius .— Biezanko, Ruffinelli & Carbonell, 1966. Bol. Fac. Agron. Montevideo 91: 29.—Biezanko & Ruffinelli, 1967. Bol. Fac. Agron. Montevideo 96: 11.—O. Mielke, 1968. Atas Soc. Biol. Rio de Janeiro 12: 75.— Ebert, 1969. Jour. Lep. Soc. 23, Suppl. 3: 39.—Biezanko & O. Mielke, 1973. Acta biol. paranaense 2: 69.— Hayward, 1973. Op. Lill. 23: 45; syn.: variegatus , dentivittata .—Biezanko, Ruffinelli & Link, 1978. Rev. Centro Ciências Rur., Sta. Maria, 8, Supl., p. 14.— Bridges, 1983. Lep. Hesp. 1, p. 11, 33, 56, 123; 2, p. 34; syn.: aterea , dentivittata , variegatus .—K. Brown, 1987. An. Prim. Simp. Pantanal, Brasília, p. 167.—Bridges, 1988. Cat. Hesp. 1, p. 18, 53, 89, 195; 2, p. 57; syn.: aterea , dentivittata , variegatus .—K. Brown, 1992, in Morellato. Hist. nat. Japi, p. 177, fig. 16 (male, female v).— Martins, 1993. Trop. Zool. 6: 121; ecol.—Bridges, 1994. Cat. Fam.-Group, Gen.-Group, Sp. Group Nam. Hesp. (Lep.) World 8, p. 21, 63, 107, 234; 9, p. 64; syn.: variegatus , aterea , dentivittata .—O. Mielke & Casagrande, 1998. Revta bras. Zool. 14 (4): 991.—K. Brown & Freitas, 2000. Bol. Mus. Biol. Mello Leitão, n. s., Sta. Teresa, 11/12: 111.— Canals, 2003. Marip. Misiones, p. 72, 457, figs (d, v).— Iserhard & Romanowski, 2004. Revta bras. Zool. 21 (3): 656.—O. Mielke, 2004. Hesperioidea, p. 55, in Lamas (ed.). Checklist: Part 4A, Hesperioidea—Papilionoidea, in Heppner (ed.). Atlas Neotrop. Lep. 5A; syn.: variegatus , aterea , aserea , dentivittata , ordinarius . — O. Mielke, 2005. Cat. Amer. Hesperioidea 1, p. 8; 3, p. 728; syn.: variegatus , aterea , incisa , aserea , dentivittata , ordinarius , variegata, dentivittatus.—Fonseca; Kumagai & O. Mielke, 2006. Revta bras. Ent. 50 (3): 402.—Carneiro, O. Mielke & Casagrande, 2008. Shilap Revta. lepid. 36 (142): 267.—Núñez B., 2008. Trop. Lep. Res. 18 (2): 80.—Rodrigues & Bononi (ed.), 2008. Diret. Cons. Rest. Biodiv. São Paulo, p. 219.—Bentancur-Viglione, 2009. Shilap Revta. lepid. 37 (145): 26.—Núñez B., 2009. Trop. Lep. Res. 19 (2): 77.—Francini et al., 2011. Revta bras. Ent. 55 (1): 67.—Dolibaina, O. Mielke & Casagrande, 2011. Biota Neotrop. 11 (1): 345.— Soares et al., 2011. Trop Lep. Res. 21 (2): 72.—Giovenardi et al., 2013. Rev. colomb. Ent. 39 (2), Supl., p. 12.—Volkmann & Núñez B., 2013. Marip. Serran. Arg. Central. 2, Nymph. Hesp., p. 102, figs (d); syn.: variegatus , aterea , dentivittata , ordinarius .—Garcia-Salik et al., 2014. Shilap Revta. lepid. 42 (166): 274.— Beltrami et al., 2014. Trop. Lep. Res. 24 (1): 34.— Thiele et al., 2014. Biota Neotrop. 14 (2): 4.—Piovesan et al., 2014. Scientia Plena 10 (9): 14.—Núñez B., 2015. Patrim. Nat. Cult. Área Infl. Emb. Yacyretá, Argentina, p. 110.—Piovesan & Orlandin, 2016, in Orlandin et al., Borb. Marip. S. Catarina, p. 80.—Núñez B., 2016. Trop. Lep. Res. 26 (1): 54.— Lavinia et al., 2017. PLoS ONE 12 (10) (e0186845): Suppl. Fig. S1 , S 2 , Table A–F.—Palo Jr., 2017. Borb. Brasil 3, p. 1907, 1908, figs (male, female d); p. 2306.—Klimaitis et al., 2018. Marip. Argentinas, p. 233, figs. (male d, v, female d, v, map).—Zamoner, 2021. Borboletas de Curitiba, 1ª ed., p. 51.

(no genus) incisus ; Beattie, 1976. Rhop. Direct., p. 167.

(no genus) aterea ; Beattie, 1976. Rhop. Direct., p. 87.

Bolla atahuallpai [misidentification]; Garwood et al., 2007. Butt. South. Amaz., p. 198, fig. (v).

Incisus incisus ; Cong et al., 2019. Insecta Mundi 731: 11, 49, fig. 4 (genomic tree).—Zhang et al., 2022. Insecta Mundi 921: 29, 122, fig. 5 (genomic tree). Silva et al., 2022. Shilap 50 (198): 378.—Lemes et al., 2023. An. Acad. Bras. Ciênc. 95(suppl. 2): 9, fig. 8.

Taxonomic history. Mabille (1878 c) described Antigonus incisus , which has been listed as Systasea incisa by some authors. Plötz (1884) described Nisoniades aterea . Evans (1953) treated Mabille’s species as Staphylus incisus for the first time and also synonymized it with Antigonus variegatus Mabille, 1878 , Nisoniades aterea Plötz, 1884 , Staphylus dentivittata Hayward, 1933 and Staphylus ordinarius Hayward, 1933 . Most other authors have followed Evans’ (1953) arrangement. Cong et al. (2019), through a phylogenomic analysis, transferred this species to the new genus Incisus .

Types. Antigonus incisus Mabille, 1878 was described from at least one male from Pará, Brazil, and one female from Cayenne [ French Guiana], currently deposited in NHMUK. The male syntype ( Fig. 2G–I ) carries the following labels: / H 757 Type HT [written on the label back]/ Brasilia/ incisus Mab. / aterea Plotz / R. Oberthür Coll. Brit. Mus. 1931-136/ [label with genitalia attached]/ 506/ BMNH(E) 1669532/ and is herein designated as lectotype to ensure the correct identification of the species. The following two labels will be added to it: / Lectotypus / Lectotypus Antigonus incisus Mabille, 1878 Lemes det. 2022/. The female syntype ( Fig. 2J–L ) carries the following labels: / H 758 [“Type HT” written on the label back]/ 20 5A/ ant. variegatus P. Mab. / R. Oberthür Coll. Brit. Mus. 1931- 136/ BMNHE 1669533/ is herein designated as paralectotype and two labels will be added to it: / Paralectotypus / Paralectotypus Antigonus incisus Mabille, 1878 Lemes det. 2022/; this female specimen was erroneously regarded by Evans (1953: 95) as the “type” of variegatus Mabille, 1878 . Although the syntypes are described from Pará and Cayenne [ French Guiana], their labels indicate Brasilia for the male; the female lacks locality information.

Nisoniades aterea Plötz, 1884 (Herrich-Schäffer in litt.) was described based on an unspecified number of specimens from Rio [de Janeiro, Brazil]. Considering that no syntypes were found in the museums where the Plötz types could be (MfN and SNSB), a neotype is designated herein to confirm the identity of the species. The neotype designated is the same specimen proposed as lectotype of Antigonus incisus Mabille, 1878 , and to it will be attached the following two labels: / Neotypus / Neotypus Nisoniades aterea Plötz, 1884 Lemes, Siewert, Mielke, Casagrande & Warren det. 2022/ ( Fig. 2G–I ). The neotype has the same characteristics of the original description, is from Brazil, and is deposited in a scientific institution, the NHMUK, in accordance with the ICZN (1999).

Diagnosis. Males of I. incisus can be easily separated from the other species due to the presence of two hyaline spots on the apical area of the FW ( Fig. 3I–J ). The male genitalia is most similar to that of I. fasciatus ( Fig. 4E–F ), however the process between ampulla and harpe is larger and oval. Also, I. incisus is the only species of the genus that lacks a cornutus on the aedeagus ( Fig. 4I–J ). Females of I. incisus can usually be distinguished by their much more evident darker and lighter bands on the wings ( Fig. 2K–N ).

Description. Male ( Fig. 3I–J ) — FW length: 13 mm to 15 mm. DFW: Presence of two hyaline spots on the apical area. HW length: 8 mm to 11 mm. VHW: Presence of cream-white scales most abundant near the marginal area. Genitalia ( Fig. 4I–L ): Uncus tapering to the distal region. Valva somewhat oval in lateral view; sacculus slightly longer than wide; harpe approximately four times longer than wide, with similar width throughout its extension and making a curve upwards at its distal portion; ampulla with general aspect similar to harpe, with a large and oval process with micotrichia on its distal portion, between ampulla and harpe. Aedeagus without cornutus. Female ( Fig. 3K–N )—Differ from the male as follows: FW length: 13 mm to 15 mm. HW length: 9 mm to 13 mm. DFW: Brown with lighter and darker transverse bands, in some individuals the lighter bands are much more evident than in males ( Fig 3K–L ), and others they are similar to the males ( Fig. 3M–N ). Presence of two or three small white spots in the apical area; when three points are present, the lower is smaller. Genitalia ( Fig. 4C ): Lamella antivaginalis triangular in ventral view. Lamella postvaginalis as a semi-rounded plate. Distal third of ductus bursae not enlarged, with a narrow signum.

Biology. Martins (1993) cites Editha magnifica ( Perty, 1834) ( Hymenoptera , Sphecidae ) as a predator of Incisus incisus . Fonseca et al. (2006) documented the presence of I. incisus visiting the blossoms of Stachytarpheta cayennensis (Rich.) Vahl ( Verbenaceae ) in a remaining fragment of the Atlantic Forest in Minas Gerais, Brazil, during December. Núñez B. (2016) observed aggregations of mud-puddling behavior in I. incisus along riverbanks, streams, puddles, and muddy areas within the Atlantic Forest of Misiones, Argentina, and Canals (2003) cites it as a nectarivorous species. Additionally, Canals (2003) notes that I. incisus has a jumping flight and rests in sunny places, with the wings unfolded. Mielke & Casagrande (1998) regard it as a common butterfly found in primary forests.

Hostplants. No records.

Distribution ( Fig. 6 ). Brazil: Mato Grosso (K. Brown 1987), Minas Gerais, Espírito Santo, Rio de Janeiro, São Paulo, Paraná, Santa Catarina , Rio Grande do Sul. Bolivia ( Spitz 1932). Argentina: Entre Ríos ( Hayward 1941), Formosa ( Hayward 1973), Misiones (Núñez B. 2008), Salta ( Hayward 1939), Tucumán ( Hayward 1941). Uruguay ( Evans 1953). Paraguay: Alto Paraná, Guairá.

Temporal distribution. Incisus incisus has been collected throughout the year.

Etymology. From the Latin masculine noun incisus meaning incision ( Glare 1968). Although Mabille (1878) did not provide an etymology, it is possible that he was referring to the conspicuous undulate (‘incised) margins of the wings of Incisus incisus .

Remarks. Antigonus variegatus Mabille, 1878 has been considered a synonym of Incisus incisus ( Mabille, 1878) ; however, after studying a syntype deposited in the MfN, labeled as “Origin”, it is concluded that it is a new synonym of Nisoniades macarius (Herrich-Schäffer, 1870) syn. nov. Nisoniades macarius was described from Venezuela and Suriname, although the specimen has a label stating “ Brasilien ” [ Brazil].

Staphylus dentivittata Hayward, 1933 has been considered a synonym of Incisus incisus ( Mabille, 1878) ; however, after studying the holotype currently deposited in the MLP, is concluded that it is a new synonym of Staphylus musculus (Burmeister, 1875) syn. nov.

Examined material. BRAZIL. Minas Gerais —Barbacena, Serra da Mantiqueira , 9.XII.1952, 1♂, H. Ebert leg., ex H. Ebert Collection, DZ 46.341 ( DZUP) ; Cambuquira, 3.X.1968, 1♂, 1♀, H. Ebert leg., ex H. Ebert Collection, DZ 46.309, DZ 46.311 ( DZUP) ; 10.XII.1968, 1♂, H. Ebert leg., ex H. Ebert Collection, DZ 46.314* ( DZUP) ; 18.XII.1968, 1♂, H. Ebert leg., ex H. Ebert Collection, DZ 46.312 ( DZUP) ; Carmo do Rio Claro, 20.II.1959, Mielke leg., 1♂, OM 2.769 * ( OM) ; Conceição dos Ouros, 25.IX.2000, 1♀, Pereira leg., OM 58.890 ( OM) ; 15.X.2000, 2♂, Pereira leg., OM 58.872 , OM 58.881 * ( OM) ; IV.2017, 1♂, 1♀ Pereira leg., OM 83.550 , OM 83.590 ( OM) ; Serra Grande , 1.365 m , 8.II.2003, 3♂, Mielke & Casagrande leg., OM 58.839 , OM 58.856 , OM 59.010 ( OM) ; 14.VI .2005 , 1♂, Almeida leg., OM 66.967 (OM); Ibituruna, 800 m , 15.IV.1954, 1♂, H. Ebert leg., ex H. Ebert Collection, DZ 46.342 ( DZUP) ; Poços de Caldas, 1200 m , 2.III.1964, 1♀, H. Ebert leg., ex H. Ebert Collection, DZ 46.735 ( DZUP) ; 1250 m, 18.XI.1966, 1♂, H. Ebert leg., ex H. Ebert Collection, DZ 46.345 ( DZUP) ; 14.XII.1966, 1♂, H. Ebert leg., ex H. Ebert Collection, DZ 46.344 ( DZUP) ; 16.XII.1966, 1♂, H. Ebert leg., ex H. Ebert Collection, DZ 46.343* ( DZUP) ; Morro do Ferro , 1600 m , 27.III.1965, 1♂, O. Mielke leg., OM 11.289 ( OM) ; 31.III.1965, 1♂, Mielke leg., OM 3.114 ( OM) ; Sabará, 18.IV.1981, 1♂, C. Elias leg., DZ 46.340 ( DZUP) ; São Gonçalo do Rio Abaixo, Peti , 560 m , 5–8.XII.2002, 1♂, Mielke leg., DZ 46.348 ( DZUP) ; Espírito Santo —Aracruz, near Santa Cruz, 8–10.III.1973, 1♂, C. Callaghan leg., Forest Area , A. C. Allyn Acc. 1973-24 ( MGCL) ; Baixo Gandu, 25.IV.1970, 1♂, 1♀, C. & C. T. Elias leg., DZ 46.223*, DZ 46.224 ( DZUP) ; 11.VI .1971 , 1♀, C. & C. T. Elias leg., DZ 46.225 (DZUP); Linhares, 6. VI .1972, 1♂, C. Elias leg., DZ 46.226 ( DZUP) ; 14.IX.1972, 1♂, C. Elias leg., DZ 46.227 ( DZUP) ; 15.VI .1973 , 2♂, C. Elias leg., DZ 46.228, DZ 46.229 (DZUP); 25–30.VI .1973 , 1♂, C. Elias leg., DZ 46.230 (DZUP); 10–15.VI .1974 , 1♀, C. Elias leg., DZ 46.231 (DZUP); 4.VII.1974, 1♂, C. Elias leg., DZ 46.232 ( DZUP) ; 2.IX.1974, 1♂ C. Elias leg., Allyn Musum Acc. 1987-5, MGCL 1043714 ( MGCL) ; I.1978, 1♀ C. Elias leg., DZ 46.220 ( DZUP) ; XI.1978, 1♂, Elias leg., DZ 46.219 ( DZUP); V .1981, 1♀, C. Elias leg., DZ 46.221 ( DZUP); V .1982, 1♂, C. Elias leg., DZ 46.222* ( DZUP) ; Santa Teresa, 4–7.III.1973, 3♂, 1♀, C. Callaghan leg., A. C. Allyn Acc. 1973-24 ( MGCL) ; São João de Petrópolis , 19–22.VIII.1996, 1♂, C. Elias leg., DZ 864* ( DZUP) ; Rio de Janeiro —Angra dos Reis, Japuíba , 1♂, 26.VI.1976, C. Callaghan, ex C. Callaghan Coll., Allyn Museum Acc. #1976-11, GEN. PREP. LEMES 2021-64 * (MGCL); Cachoeiras do Macacu, Boca do Mato , 11.II.2005, 1♂, N. Tangerini leg., Ex Coleção Nirton Tangerini, DZ 46.338 ( DZUP) ; 15.V .2005 , 1♂, N. Tangerini leg., Ex Coleção Nirton Tangerini, DZ 46.300 (DZUP); Duque de Caxias, Imbariê , no data, 1♀ H. Ebert leg., ex H. Ebert Collection, DZ 46.313 ( DZUP) ; Guapimirim, 50 m, 18.II.1958, 1♂, H. Ebert leg., Coleção H. Ebert, DZ 46.310 ( DZUP) ; Itatiaia, Parque Nacional de Itatiaia , 1350 m , 23.VII.1959, 1♂, Ebert leg., ex H. Ebert Collection, DZ 46.296 ( DZUP) ; 1000–1200 m, 7.IX.1964, 1♂, H. Ebert leg., ex H. Ebert Collection, DZ 46.288 ( DZUP) ; 4.VIII.1966, 1♂, N. Tangerini leg., Ex Coleção Nirton Tangerini, DZ 46.292 ( DZUP) ; 15.I.1967, 1♂, Mielke leg., DZ 46.286 ( DZUP) ; 1300–1700 m, 21.I.1969, 1♂, Mielke & Brown leg., DZ 46.283 ( DZUP) ; 15.I.1967, 1♂, N. Tangerini leg., ex Coleção Nirton Tangerini, DZ 46.290 ( DZUP) ; 200 m, 7.I.1973, 1♂, Mielke leg., DZ 46.277 ( DZUP) ; 1400 m, 12.I.1973, 2♂, Mielke leg., DZ 46.279, DZ 46.280 ( DZUP) ; Magé, Suruí, 5.VI .1971 , 1♂, C. Callaghan leg., A. C. Allyn Acc. 1971-35 (MGCL); Surui, 3 Km N. 22.V .1971 , 1♂, C. Callaghan leg., A. C. Allyn Acc. 1971-17 (MGCL); Petrópolis, 25.VIII.1951, 1♂, D’Almeida leg., ex Coll. D’Almeida, DZ 46.265* ( DZUP) ; 25.VIII.1957, 1♀, D’Almeida leg., ex Coll. D’Almeida, DZ 46.273 ( DZUP) ; 30.I.1966, 1♂, Ebert leg., ex H. Ebert Collection, DZ 46.305* ( DZUP) ; 8.I.1969, 1♂, O. Mielke leg., DZ 46.303 ( DZUP) ; 9.I.1971, 1♂, Mielke leg., DZ 46.304 ( DZUP) ; 900 m, 14.I.1971, 1♂, C. Callaghan leg., A. C. Allyn Acc. 1971-9 ( MGCL) ; 900 m, 24.I.1971, 3♂, C. Callaghan leg., A. C. Allyn Acc. 1971-9, Genit. Prep. SRS-1504* ( MGCL) ; 900 m, 5.II.1971, 2♂, C. Callaghan leg., A. C. Allyn Acc. 1971-9 ( MGCL) ; 950 m, 9.III.1971, 1♂, C. Callaghan leg., A. C. Allyn Acc. 1971-14 ( MGCL) ; 950 m, 17.III.1971, 5♂, C. Callaghan leg., A. C. Allyn Acc. 1971-14 ( MGCL) ; 950 m, 2.IV.1971, 2♂, C. Callaghan leg., A. C. Allyn Acc. 1971-14 ( MGCL) ; 9.IV.1971, 1♀, Ebert leg., ex H. Ebert Collection, DZ 46.306 ( DZUP) ; 900 m, 16.I.1972, 3♂, C. Callaghan leg., A. C. Allyn Acc. 1972-13 ( MGCL) ; 1500 m, 29.I.1972, 1♂, C. Callaghan leg., A. C. Allyn Acc. 1972-13 ( MGCL) ; 900 m, 3.III.1972, 1♂, C. Callaghan leg., A. C. Allyn Acc. 1972-13 ( MGCL) ; 30.III.1974, 1♀, C. Callaghan leg., Allyn Museum Acc. 1978-18 C. Callaghan ( MGCL) ; Alto da Serra , 15.IX.1963, 2♀, Mielke leg., OM 5.395 *, OM 5.396 ( OM) ; 1.IX.1964, 1♀, Mielke leg., OM 5.988 ( OM) ; Independência, 1.XII.1966, 1♂, N. Tangerini leg., Ex Coleção Nirton Tangerini, DZ 46.301 ( DZUP) ; 800 m, 27.I.1969, 1♂, 1♀, Mielke leg., 46.299, DZ 46.298; 31.I.1969, 1♂, N. Tangerini leg., Ex Coleção Nirton Tangerini, DZ 46.302 ( DZUP) ; 20 Km. E. Petrópolis, 7.III.1971, 1♀, C. Callaghan leg., A. C.Allyn Acc. 1971-14, Genit. Prep. SRS-1535* ( MGCL) ; Rio de Janeiro, Jacarepaguá, 13.II.1971, 1♂, C. Callaghan leg., A. C. Allyn Acc. 1971-9 ( MGCL) ; 3.IV.1971, 1♀, C. Callaghan leg., A. C. Allyn Acc. 1971-4 ( MGCL) ; 14.VII.1963, 1♀, D’Almeida leg., ex Coll. D’Almeida, DZ 46.266 ( DZUP) ; Gavea Pequena, 04.I.1972, 1♂, C. Callaghan leg., A. C. Allyn Acc. 1972-13 ( MGCL) ; Covanca, 2.XI.1932, 1♀, D’Almeida leg., ex Coll. D’Almeida, DZ 46.268 ( DZUP) ; 7.IV.1947, 1♀, D’Almeida leg., ex Coll. D’Almeida, DZ 46.269 ( DZUP) ; 25.VIII.1957, 1♂, Mielke leg., OM 1.505 ( OM) ; 10.V .1957 , 1♀, D’Almeida leg., ex Coll. D’Almeida, DZ 46.274 (DZUP); 18.VIII.1957, 1♂, D’Almeida leg., ex Coll. D’Almeida, DZ 46.272 ( DZUP) ; 19.II.1958, 1♀, D’Almeida leg., ex Coll. D’Almeida, DZ 46.275 ( DZUP) ; 6.IV.1958, 1♂, Mielke leg., OM 1.917 ( OM) ; 29.VII.1959, 1♀, Mielke leg., ex Coll. D’Almeida, DZ 46.276* ( DZUP) ; 29.V .1960 , 1♂, Mielke leg., OM 3.276 (OM); 8.IV.1962, 1♀, Mielke leg., OM 1.418 ( OM) ; 19.VIII.1962, 1♂, Mielke leg., OM 4.604 * ( OM) ; 25.IV.1962, 1♂, 3♀, Mielke leg., OM 4.357 , OM 4.358 , OM 4.360 , OM 4.361 ( OM) ; 28.IV.1962, 1♀, D’Almeida leg., ex Coll. D’Almeida, DZ 46.271* ( DZUP) ; 25.III.1964, 1♂, D’Almeida leg., ex Coll. D’Almeida, DZ 46.267* ( DZUP) ; 29.III.1964, 5♂, 2♀, Mielke leg., OM 5.546 , OM 5.547 , OM 5.548 , OM 5.549 , OM 5.551 , OM 5.550 , OM 5.551 ( OM) ; 18.IV.1964, 1♀, Mielke leg., OM 5.711 ( OM) ; Três Rios, 18.IX.1960, 1♀, D’Almeida leg., ex Coll. D’Almeida, DZ 46.270 ( DZUP) ; Paineiras, 23.I.1955, 1♂, H. Ebert leg., ex H. Ebert Collection, DZ 46.308 ( DZUP) ; Restinga da Tijuca , 1100 m , 22.VI .1952 , 1♀, H. Ebert leg., ex H. Ebert Collection, DZ 46.307 (DZUP); Sumaré, 8.XI.1966, 1♂, Brown leg., DZ 46.297 ( DZUP) ; Teresópolis, Km 16 Rio-Teresópolis Hwy, 1.VII.1972, 4♂, 1♀, C. Callaghan leg., A. C. Allyn Acc. 1972-47 ( MGCL) ; São Paulo — Anhembi (Fazenda Barreiro Rico), 500 m , 6.II.1978, 1♂, Ebert leg., ex H. Ebert Collection, DZ 46.242 ( DZUP) ; Araras, 600 m , 15.I.1966, 1♀, Ebert leg., ex H. Ebert Collection, DZ 46.260 ( DZUP) ; Campos do Jordão, 1600–2000 m , 8–12.II.1982, 3♂, Mielke & Casagrande leg., DZ 46.502, DZ 46.504, DZ 46.505 ( DZUP) ; Caraguatatuba, 10 m, no data, 1♀, Ebert leg., ex H. Ebert Collection, DZ 46.262 ( DZUP) ; 5.II.1963, 1♂, Ebert leg., ex H. Ebert Collection, DZ 46.257* ( DZUP) ; 8.IV.1963, 1♀, Ebert leg., ex H. Ebert Collection, DZ 46.245 ( DZUP) ; 4.IX.1968, 1♂, Ebert leg., ex H. Ebert Collection, DZ 46.255 ( DZUP) ; Cordeirópolis, 600 m , 25.XI.1962, 2♂, 1♀, Ebert leg., ex H. Ebert Collection, DZ 46.241, 46.263, DZ 46.256 ( DZUP) ; 11.IV.1963, 2♂, Ebert leg., ex H. Ebert Collection, DZ 46.238*, DZ 46.240* ( DZUP) ; Jundiaí, Japi, 5.V .1990 , 1♀, Brown leg., OM 25.347 (OM); Rio Claro, 600 m , 9.XII.1962, 1♂, Ebert leg., ex H. Ebert Collection, DZ 46.239 ( DZUP) ; 24.II.1963, 1♀, Ebert leg., ex H. Ebert Collection, DZ 46.258 ( DZUP) ; 24.IV.1966, 1♀, Ebert leg., ex H. Ebert Collection, DZ 46.244 ( DZUP) ; 20.XI.1966, Ebert leg., ex H. Ebert Collection, DZ 46.259 ( DZUP) ; Santana de Paranaíba, Represa Edgard de Souza , 4.VII.2000, 1♂, Bizarro leg., OM 50.972 * ( OM) ; 15.I.2004, 1♂, Bizarro leg., DZ 46.247 ( DZUP) ; São Paulo, 15.VII.1960, 1♂, D. L. Lindsley leg., D. L. Lindsley Coll. MGCL Accession #2008-20 ( MGCL) ; 13.VIII.1960, 1♂, D. L. Lindsley leg., D. L. Lindsley Coll. MGCL Accession #2008-20 ( MGCL) ; 12.IX.1960, 1♂, D. L. Lindsley leg., D. L. Lindsley Coll. MGCL Accession #2008-20 ( MGCL) ; 12.XI.1960, 1♂, D. L. Lindsley leg., D. L. Lindsley Coll. MGCL Accession #2008-20 ( MGCL) ; 3.XII.1960, 1♀, D. L. Lindsley leg., D. L. Lindsley Coll. MGCL Accession #2008-20 ( MGCL) ; 15.IV.1961, 1♀, D. L. Lindsley leg., D. L. Lindsley Coll. MGCL Accession #2008-20( MGCL) ; 29.IV.1961, 4♂, D. L. Lindsley leg., D. L. Lindsley Coll. MGCL Accession #2008-20 ( MGCL) ; Ilha Seca , 3.XI.1938, 1♂, D’Almeida leg., ex Coll. D’Almeida, DZ 46.237 ( DZUP) ; Pico do Jaraguá , 11.IX.1960, 1♂, D. L. Lindsley leg., D. L. Lindsley Coll., MGCL Accession #2008-20 ( MGCL) ; Túnel da Mata Fria , 800 m , 8.II.1985, 2♂, Mielke & Casagrande leg., DZ 46.243, DZ 46.254 ( DZUP) ; Teodoro Sampaio, Parque Estadual Morro do Diabo , 300–600 m , 22–25.X.1987, 5♂, 1♀, Mielke & Casagrande leg., OM 18.002 , OM 18.003 , OM 18.004 , OM 18.005 , OM 18.001 ( OM) ; 22–26.X.1987, 1♂, O. Mielke leg., OM 51.624 ( OM) ; 17–19.VIII.1989, 2♂, Mielke & Casagrande leg., DZ 46.252, DZ 46.253* ( DZUP) ; 5–8.XII.1989, 2♂, Mielke & Casagrande leg., DZ 46.250, DZ 46.251* ( DZUP) ; 250 m, 20–23.II.1990, 1♂, 1♀, Mielke & Casagrande leg., DZ 46.249, DZ 46.248 ( DZUP) ; Ubatuba, 10 m , 19.XII.1963, 1♂, Ebert leg., ex H. Ebert Collection, DZ 46.261 ( DZUP) ; Paraná — Antonina, Cacatu , 20 m , 25.IV.1973, 2♂, Mielke leg., DZ 46.484, DZ 46.485 ( DZUP) ; Cândido de Abreu, 10.XII.1994, 1♂, Mielke & Casagrande leg., OM 40.610 ( OM) ; Diamante do Norte, Estação Ecológica do Caiuá , 22º36’24’’S, 52º52’14’’W , 16.V .2009 , 1♂, D. Dolibaina & E. Carneiro leg., DZ 46.462 (DZUP); Fênix, 300 m , 23.XI.1986, 2♂, DZ 46.467, DZ 46.468 ( DZUP) ; 29.IV.1987, 1♂, 1♀, Mielke & Casagrande leg., DZ 46.470, DZ 46.469 ( DZUP) ; 3–4.X.1987, 1♀, Mielke & Casagrande leg., DZ 46.471 ( DZUP) ; Horto Florestal, 3.X.1986, 1♂, 2♀, DZ 46.473, DZ 46.472, DZ 46.474* ( DZUP) ; Foz do Iguaçu, Parque Nacional do Iguaçu , 4.XII.1966, 2♀, D. Z. U. F. P. leg., DZ 860*, DZ 46.496 ( DZUP) ; 7.XII.1966, 3♀, D. Z. U. F. P. leg., DZ 46.497, DZ 46.498, DZ 46.499 ( DZUP) ; 8.XII.1966, 1♂, D. Z. U. F. P. leg., DZ 46.500* ( DZUP) ; 17.II.1969, 1♂, Moure & Mielke leg., DZ 46.501 ( DZUP) ; 6.IX.1985, 1♂, 1♀, Mielke & Casagrande leg., DZ 46.494, DZ 46.493 ( DZUP) ; 21–24.IV.1995, 1♀, Mielke & Casagrande leg., DZ 46.495 ( DZUP) ; 29.V .2014 , 1♂, R. Greve leg., DZ 46.492 (DZUP); Parque Nacional do Iguaçu, Trilha da Boa Vista 12–13.I.2015, 2♂, R. Greve leg., DZ 44.221*, DZ 44.241* ( DZUP) ; Guaratuba, Limeira , 10 m , 21.IV.2000, 1♀, Mielke leg., OM 51.932 ( OM) ; Jussara, Horto Companhia Melhoramentos Norte do Paraná, 500 m , 16.XI.1975, 1♂, Mielke & Rosado leg., DZ 46.465 ( DZUP) ; 12.XII.1975, 1♂, Moure, Mielke & Wedderhoff leg., DZ 46.466 ( DZUP) ; Londrina, 10.X.1982, 1♂, Mielke leg., DZ 46.477 ( DZUP) ; 600 m, 3.I.1985, 1♀, Mielke leg., DZ 46.478 ( DZUP) ; Manoel Ribas, 450– 600 m , 12.X.2010, 1♀, Mielke, Dolibaina, Carneiro & M. Maia leg., DZ 46.461 ( DZUP) ; Morretes, Morro Alto , 400 m , 11.IV.2000, 2♂, Mielke leg., OM 51.788 , OM 51.795 ( OM) ; 300 m, 4.IV.2009, 1♂, 1♀, O. Mielke leg., DZ 46.455, DZ 46.456; 3.IV.2021, 2♂, 1♀, O. & El. Mielke leg., DZ 55.013, DZ 55.023, DZ 55.003 ( DZUP) ; Paranaguá, Alexandra, 3.X.1968, 1♀, Mielke leg., DZ 46.328 ( DZUP) ; 29.III.1971, 1♀, O. Mielke leg., DZ 46.476 ( DZUP) ; Ponta Grossa, Olarias, VIII.1949, 1♂, no Collector, ex Coleção F. Justos Jor 2019, DZ 46.489 ( DZUP) ; Parque Estadual de Vila Velha , 900 m , 8.III.1971, 1♂, 1♀, Mielke leg., DZ 46.490, DZ 46.491 ( DZUP) ; Porto Amazonas, 3.II.1971, 1♂, O. Mielke & M. Lauterjung leg., DZ 46.463 ( DZUP) ; Roncador, 15 Km SW, 550–600 m , 11.X.2010, 1♀, Mielke, Dolibaina, Carneiro & M. Maia leg., DZ 46.458 ( DZUP) ; São João do Ivaí, 20.IX.1973, 1♀, Furtado leg., DZ 46.464 ( DZUP) ; São Jorge do Ivaí, Copacabana, 16.IX.1970, 1♂, E. Furtado leg., DZ 46.481 ( DZUP) ; 550 m, 15.XI.1975, 2♂, Mielke & Rosado leg., DZ 46.482*, DZ 46.483 ( DZUP) ; São Pedro do Ivaí, RPPN Barbacena , 300 m , 8.X.2010, 1♂, Mielke, Dolibaina, Carneiro & M. Maia leg., DZ 46.457 ( DZUP); RPPN Roncador, 300 m , 8.X.2010, 1♂, 1♀, Mielke, Dolibaina, Carneiro & M. Maia leg., DZ 46.460, DZ 46.459 ( DZUP) ; Terra Boa, 650 m , 16.XI.1975, 1♂, Mielke & Rosado leg., DZ 46.488 ( DZUP) ; Companhia Melhoramento do Norrte do Paraná, 650 m , 11.XII.1975, 1♂, 1♀, Moure, Mielke & Wedderhoff leg., DZ 46.486, DZ 46.487* ( DZUP) ; Tuneiras do Oeste, REBIO das Perobas, 53º50’49’’S, 52º44’18’’W , 6.X.2012, 1♂, LABLEP leg., DZ 46.480* ( DZUP) ; Ventania, 12,5 Km N, 1000 m , 29–30.I.2006, 1♀, O. Mielke leg., DZ 46.475 ( DZUP) ; Santa Catarina —Brusque, 28.XII.1967, 1♂, V. Becker leg., DZ 46.359 ( DZUP) ; 8.I.1968, 1♂, V. O. Becker leg., DZ 45.197 ( DZUP) ; 28.XII.1969, 1♀, Becker leg., DZ 46.352 ( DZUP) ; 10.VII.1971, 2♂, V. O. Becker leg., Allyn Museum Acc. 1977-4, MGCL 10443656 ( MGCL) ; VII.1971, V. O. Becker leg., Allyn Museum Acc. 1977-4, ( MGCL) ; Florianópolis, Lagoa do Peri , 1.XI.2004, 1♀ E. Santos leg., DZ 46.351 ( DZUP) ; Garuva, 31.VIII.1985, 1♂, Mielke leg., DZ 46.350* ( DZUP) ; Joinville, 28.X.1967, 4♂, Mielke & Miers leg., DZ 46.353*, DZ 46.354, DZ 46.356, DZ 46.357 ( DZUP) ; 10–200 m, 3.II.1973, 1♂, Mielke & Miers leg., DZ 46.355 ( DZUP) ; 22.IV.1988, 1♀, O.-C. Mielke leg., OM 51.456 ( OM) ; São Bento do Sul, Rio Natal , 500 m , 13.III.1987, 1♀, Mielke & Rank leg., OM 14.402 ( OM) ; Rio Vermelho , 800 m , 15.I.1871, 1♀, Rank leg., DZ 46.361 ( DZUP) ; 850 m, 2.II.1973, 1♂, Rank leg., DZ 46.363 ( DZUP) ; 850 m, 13.III.1973, 1♂, Rank leg., DZ 46.360 ( DZUP) ; 850 m, 4.VI .1974 , 1♀, Rank leg., DZ 46.454 (DZUP); 850 m, 2.VII.2003, 1♀, Rank leg. OM 63.005 ( OM) ; Rio Natal , 500 m , 3.X.1984, 2♂, Thomas C. Emmel leg., ex FSCA Florida State Collection of Arthropods ( MGCL) ; Seara, Nova Teutônia , 450–700 m , II.1969, 1♀, F. Plaumann leg., DZ 46.362 ( DZUP) ; III.1971, 1♂, F. Plaumann leg., DZ 46.358 ( DZUP) ; II.1982, 1♂, Fritz Plaumann leg., DZ 46.479 ( DZUP) ; Taió, 29.I.1966, 1♂, Mielke leg., OM 7.881 * ( OM) ; Rio Grande do Sul —Gramado, 26.I.1973, 1♀, H. Ebert leg., ex H. Ebert Collection, DZ 46.336 ( DZUP) ; Iraí, 25.XI.1969, 3♂, Schaal leg., DZ 46.333*, DZ 46.334, DZ 46.335 ( DZUP) ; Morro Reuter, Walachei , 200–500 m , 30.III.2001, 1♂, A. Moser leg., DZ 46.337 ( DZUP) . PARAGUAY: Alto Paraná —Ciudad del Este, 6.XI.2002, 1♂, D. &. J. Lindsley leg., ex D. L. Lindsley Coll., MGCL Accession #2008-20 ( MGCL) ; Guairá — General Dias, 400 m , 15–20.I.1980, 8♂, 5♀, O.-C. Mielke & Miers leg., DZ 46.315*, 46.316, 46.317, 46.318, DZ 46.319, DZ 46.320*, DZ 46.321, DZ 46.322, DZ 46.323, DZ 46.324, 46.325, 46.326, DZ 46.327 ( DZUP) . ARGENTINA: Misiones — no specific locality, no Collector , 1♂, DZ 863* ( DZUP) ; Dos de Mayo, 13.II.1989, 1♂, no Collector, OM 21.199 ( OM) ; Eldorado, I.1940, 1♀, no Collector, DZ 866* ( DZUP) ; General Belgrano, Almirante Brown, Reserva Yacutinga, 2–5.III.2007, 3♂, 1♀, Mielke & Casagrande leg., DZ 46.329, DZ 46.330, 46.331, 46.332 ( DZUP) ; Puerto Iguazú, Parque Nacional Iguazú, 8.XI.2002, 1♀, D. &. J. Lindsley leg., ex D. L. Lindsley Coll. MGCL Accession #2008-20 ( MGCL) .

Figures

FIGURE 1. Relationships of species of Incisus Grishin, 2019 and other species of Carcharodini based on COI sequences obtained by Maximum likelihood inferred with IQTREE. The blue rectangle represents the genus Incisus. The type species of different genera used in the analysis are bold highlighted. The new species is red highlighted. Support values were considered sufficient when SH-aLRT ≥ 80, aBayes ≥ 0.5 and UFBoot ≥ 95.

FIGURE 2. Types of Incisus. A–C. Male holotype of Pholisora angulata Bell, 1937 (AMNH): (A) dorsal view, (B) ventral view, (C) labels. D–F. Male holotype of Staphylus fasciatus Hayward, 1933 (MLP): (D) dorsal view, (E) ventral view, (F) labels. G–I. Male lectotype herein designated of Antigonus incisus Mabille, 1878 and neotype herein designated of Nisoniades aterea Plötz, 1884 (NHMUK): (G) dorsal view, (H) ventral view, (I) labels. J–L. Female paralectotype herein designated of Antigonus incisus Mabille, 1878 (NHMUK): (J) dorsal view, (K) ventral view, (L) labels. M. Illustration of the male genitalia of Staphylus ordinarius Hayward, 1933 (Hayward, 1934). Scale bar = 1 cm.

FIGURE 3. Species of Incisus in dorsal and ventral views. A–D. I. angulata (Bell, 1937): (A–B) male (Brazil, Minas Gerais, Conceição dos Ouros, OM 83.262), (C–D) female (Brazil, Rio de Janeiro, Resende, Parque Nacional do Itatiaia, DZ 46.281). E–H. I. fasciatus (Hayward, 1933): (E–F) male [neotype of Staphylus ordinarius Hayward, 1933] (Argentina, Salta, Aguas Blancas, DZ 46.164), (G–H) female (Argentina, Jujuy, Calilegua, MGCL 1092134). I–N. Incisus incisus (Mabille, 1878): (I–J) male (Brazil, São Paulo, Campos do Jordão, DZ 46.504), (K–L) female (Brazil, Santa Catarina, São Bento do Sul, Rio Vermelho, DZ 46.454), (M–N) female (Brazil, Paraná, Foz do Iguaçu, Parque Nacional do Iguaçu, DZ 46.495). O–R. I. mineirus sp. nov.: (O–P) male holotype (Brazil, Minas Gerais, Corinto, DZ 46.940), (Q–R) female allotype (Brazil, Minas Gerais, Corinto, DZ 46.947). Scale bar = 1 cm.

FIGURE 4. Male genitalia of Incisus species in lateral, internal, dorsal and ventral views, respectively. A–D. Incisus angulata (Bell, 1937) (Brazil, Minas Gerais, Virgínia, Fazenda dos Campos, DZ 46.339). E–H. Incisus fasciatus (Hayward, 1933) (Brazil, Paraná, Guaíra, Parque Nacional de Sete Quedas, DZ 44.840). I–L. Incisus incisus (Mabille, 1878) (Brazil, Santa Catarina, Joinville, DZ 46.353). M–P. I. mineirus sp. nov. (Brazil, Minas Gerais, São Francisco, DZ 46.559). Scale bar = 500 µm.

FIGURE 5. Female genitalia of Incisus species, in ventral view. A. Incisus angulata (Bell, 1937) (Brazil, Rio de Janeiro, Itatiaia, Parque Nacional do Itatiaia, DZ 42.282). B. Incisus fasciatus (Hayward, 1933) (Argentina, Jujuy, Calilegua, RT. 83— 7.5km W of Rt 34, MGCL 1092134). C. Incisus incisus (Mabille, 1878) (Brazil, Paraná, Fênix, DZ 46.474). D. Incisus mineirus sp. nov. (Brazil, Minas Gerais, Corinto, DZ 46.942). Scale bar = 1 mm.

FIGURE 6. Distribution of Incisus species. The divisions of Brazilian states are represented. Solid circles represent collections studied records and open circles represent literature records of Incisus incisus.