Zaretis strigosus, (GMELIN, [1790])

ZARETIS STRIGOSUS (GMELIN, [1790]) View in CoL

( FIGS 8A–D, 15C, 16E, F, 20; SUPPORTING INFORMATION, APPENDIX S6)

Type material

Female lectotype of Papilio strigosus Gmelin, [1790] with the following labels: / 55 strigosus, Gmel [in] type specimen/ Lectotype / Papilio strigosus Gmelin det. R.I. Vane-Wright, 1971 LECTOTYPE #f/ (NMI).

Male lectotype of Siderone isidora var. strigosa Staudinger, 1886 , here designated, with the following label: / Blumenau [18]85 Mull[er]/ ( ZMHU) .

Male lectotype of Zaretes [sic] isidora russeus Fruhstorfer, 1909 , here designated, with the following labels: / Type/ Type/ Fruhstorfer Coll. B.M. 1937-285./ Columbien [ Colombia] ex coll. H. Fruhstorfer/ russeus Fruhst [orfer]./ Zaretes isidora russeus Fruh [storfer]. det. R.I. Vane-Wright, 1972 #m SYNTYPE / (NHMUK).

Male lectotype of Zaretes [sic] isidora vulpecula Fruhstorfer, 1909 , here designated, with the following labels: / Type/ Type / Fruhstorfer Coll. B.M. 1937-285./ Bahia Brasilia [ Brazil] Fruhstorfer/ isidora vulpecula Fruhst [orfer]./ Zaretes isidora vulpecula Fruh [storfer]. det. R.I. Vane-Wright, 1972 #m SYNTYPE / (NHMUK).

Female lectotype of Zaretes [sic] isidora vulpina Fruhstorfer, 1909 , here designated, with the following labels: / Type/ Type / Fruhstorfer Coll. B.M. 1937- 285./ Paraguay ex coll. H. Fruhstorfer/ Zaretes isidora vulpina Fruh [storfer]. det. R.I. Vane-Wright, 1972 #f SYNTYPE / (NHMUK).

Zaretes [sic] isidora isidora f. foliacea Fruhstorfer, 1909 does not have a formal type because it is an infrasubspecific entity. However, the specimen described by Fruhstorfer (1909b) is a female with the following labels: / SYN-TYPE/ Type / Type / Fruhstorfer Coll. B.M. 1937-285./ Obidos [Pará] Amazonenstrom VIII-IX. 99 ex. coll. H. Fruhstorfer/ foliacea 18–70 Fruh[storfer]./ (NHMUK).

The male holotype of Siderone ellops mellita Brévignon, 2006 was not examined, but has the following label data: French Guiana, Saint-Georges de l’Oyapock , 30. V .1984, Number 46 ( Brévignon, 2006) (LCB).

Diagnosis

Zaretis strigosus is the most variable species of the genus and, as with other species of Zaretis , it is extremely similar to many of its congeners. This species is sympatric with Z. itys , Z. isidora , Z. falcis and Z. hurin sp. nov. in the Amazon basin and the Guianas, and with Z. isidora , Z. itylus and Z. hurin sp. nov. in the Atlantic forest. However, the male genitalia of this species are distinct ( Fig. 15C), comparable only to Z. ellops , a Trans-Andean species. Differing from all the above cited species, in Z. strigosus the uncus is almost straight, short, thick, medially keeled and without a distal callus, and the dorsal half of the gnathos is larger than the ventral. Males ( Fig. 8A, B) can be externally distinguished from Z. itys by the developed emargination of the inner margin at the tornus; from Z. falcis by the FW apex, only slightly to moderately falcate, outer margin not crenulated, and discal spot, marginal area and submarginal area near the apex never as developed and dark brown; from Z. itylus by the shape of the outer margin of the FW, sinuous, and HW not strongly emarginated in Sc–Rs and projected at Rs at the apex; from Z. isidora , throughout its distribution, and from Z. hurin sp. nov., in Atlantic forests, by generally much lighter and speckled coloration, with post-median and submarginal bands usually developed in both uppersides of the wings, especially the HWD post-median band, which usually runs to the tornal projection at 2A; and from Z. hurin sp. nov., in the Amazon basin and Guianas, by the absence of a purplish sheen on the FWD when viewed obliquely. Females ( Fig. 8C, D) can be distinguished from Z. itys and Z. itylus by the shape of the FW, sinuous, and upperside of the wings basal and post-median areas, never reddish brown and beige to pale yellow, respectively, and the lighter and homogeneous coloration of basal, post-median area and submarginal areas along the outer margin in both wings upper and undersides; from Z. falcis by the FW apex, considerably less falcate; and from Z. hurin sp. nov. and Z. isidora by the generally lighter and homogeneous coloration of basal, post-median and submarginal areas in both wings upper and undersides. Owing to the extreme variation of females of Z. strigosus and the latter two species, the identification only be confirmed can by the examination of the genitalia; in Z. strigosus , the lamella postvaginalis is wider than long, and its posterior edge is bilobed ( Fig. 16E, F).

Discussion

The name Z. strigosus was evaluated by Vane-Wright (1975) and briefly discussed by Dias et al. (2015). The species it represents is the most taxonomically problematic of the genus. The lectotype of Papilio strigosus Gmelin, [1790] , designated by Vane-Wright (1975), is a heavily worn female specimen with several repairs; the left antenna was replaced by a bristle. According to Kirby (1879), that specimen was described by Zschach (1788) as ‘species 55’ of the Museum Leske and afterwards properly named by Gmelin ([1790]). The exact origin of this specimen was unknown by both Zschach and Gmelin (cited as ‘ Exoticus ’ and ‘ extra Europam ’). However, as discussed by Vane-Wright (1975), it probably came from Suriname, the generally accepted locality for the Leske material. The lectotype agrees perfectly with specimens of Z. strigosus from the Amazon basin and the Guianas, with the upperside of the wings having a homogeneous pale orange to orange ground colour.

Siderone isidora v. strigosa Staudinger, 1887 was described based on an unstated number of specimens from Brazil (‘ Rio de Janeiro to Rio Grande do Sul’). Staudinger (1887) was unsure whether S. isidora v. strigosa Staudinger, 1887 was equivalent to P. strigosus Gmelin, [1790] , and suggested that Gmelin’s poorly described species might not even belong in the genus, wrongly assuming that the type was missing. Vane-Wright (1975) correctly recognized S. isidora v. strigosa Staudinger, 1887 as a subsequent secondary homonym of P. strigosus Gmelin, [1790] . No specimens can be recognized unequivocally as pertaining to the type series of S. isidora v. strigosa Staudinger, 1887 , as none of the potential syntypes at the ZMHU (i.e. one female from Rio de Janeiro , two males from São Paulo , two males and one female from Santa Catarina, one male, collected by Mr Bescke, and one female, collected by Mr Sommer, from unknown localities in Brazil) have Staudinger’s typical pink ‘Origin’ label. However, he frequently did not include this label on many of his ‘varieties’ (G. Lamas, personal communication). One of the potential syntypes, from Casa Branca, São Paulo , Brazil, collected by Mr Garbe in 1887 and illustrated by Warren et al. (2016), is labelled ‘SYNTYPUS’. It is improbable that this specimen is part of the type series of S. isidora v. strigosa Staudinger, 1887 , as it seems unlikely that this specimen could have been collected by Mr Ernst Garbe and sent to Europe early in 1887, reaching Staudinger in time to prepare and publish that part of the series, presented in mid-March 1887 ( Lamas, 2017). At least one of the potential syntypes, a female collected in ‘Brazil’ by Mr Sommer, does not belong to Z. strigosus as here conceived. A male specimen from Blumenau, Santa Catarina, Brazil, collected in 1885 by Mr F. Müller, is here designated as the lectotype.

Zaretes [sic] isidora russeus Fruhstorfer, 1909 View in CoL was described based on an unstated number of specimens from ‘Colombia’. Only one syntype was located by Vane-Wright (1975) and by us at the NHMUK type collection. This specimen was not dissected, but agrees perfectly with specimens of Z. strigosus from the Amazon basin and Guianas, being generally darker than specimens from the Atlantic forests, but having the post-median and submarginal bands usually developed on the upperside of both wings, especially the HWD post-median band, which runs to the tornal projection at 2A. This specimen, illustrated by Warren et al. (2016), is here designated as the lectotype.

Zaretes [sic] isidora isidora f. foliacea Fruhstorfer, 1909 View in CoL was described as a ‘female form’ denoting an infrasubspecific entity; therefore, the name is invalid and without a formal type. Nevertheless, it corresponds to a distinct phenotype of females of Z. strigosus from the Amazon basin, with rounded and crenulated outer margins. One of these specimens, illustrated by Warren et al. (2016), is kept at the NHMUK type collection.

Zaretes [ sic] isidora vulpecula Fruhstorfer, 1909 was described based on four specimens (three males and one female) from Bahia, Brazil, collected by Mr Haensch. All four specimens were located by Vane-Wright (1975) at the NHMUK, but we had access to only one male. This specimen, which agrees perfectly with the pale orange phenotype of specimens of Z. strigosus from the northeastern states of Brazil, is here designated as the lectotype.

Zaretes [sic] isidora vulpina Fruhstorfer, 1909 View in CoL was described based on an unstated number of specimens from Paraguay. Fruhstorfer (1909b) claimed he had access to a large quantity of specimens, but Vane-Wright (1975) found only eight syntypes (seven males and one female) at the NHMUK. Only two of these (one male and one female) were located by us. The male specimen could not be dissected, but it is rather different from the most common phenotype of Z. strigosus in Paraguay, even though the female certainly represents Z. strigosus . There is a strong possibility that the type series contains multiple species. Nevertheless, the species description and the picture and description ( Fruhstorfer, 1909b: 167–168, fig. 4) of the male genitalia clearly agree with Z. strigosus , one of the few species of the genus that can be identified reliably by the examination of the genitalia. Therefore, the above cited female specimen, which agrees perfectly with specimens of Z. strigosus from Paraguay, deposited in the NHMUK type collection, is designated here as the lectotype.

Siderone ellops melita Brévignon, 2006 was described based on nine specimens from French Guiana, two males and seven females. The male paratype illustrated ( Brévignon, 2006: p. 300, figs 9, 10) is unlike most specimens of Z. strigosus from the Amazon basin and Guianas, but the species description and the plates of the male genitalia unequivocally identify this name as Z. strigosus . The female specimen illustrated is similar to the type of Z. strigosus and to specimens from the Amazon basin and Guianas, with pale orange to orange ground colour on the upperside of the wings. Therefore, this name is treated here as a junior subjective synonym of Z. strigosus (comb. nov., syn. nov.).

The immature stages of Z. strigosus are typical of the genus and were described by Müller (1886) and Dias et al. (2015) from Santa Catarina, Rio de Janeiro and Paraná View in CoL and Santa Catarina in Brazil, respectively. Several authors recorded host plants for what is assumed to be Z. strigosus , the most common species of the Zaretis View in CoL in Cis-Andean South America. Most authors cite host plants in the Salicaceae View in CoL : unidentified species of Casearia View in CoL (F. Müller, 1878; Otero & Marigo, 1990; Brown, 1992; Freitas apud Beccaloni et al., 2008) and Casearia sylvestris View in CoL ( Müller, 1886; Mabilde, 1896; Costa-Lima, 1928, 1936; Ronna, 1933; Biezanko, Ruffinelli & Carbonell, 1966; Silva et al., 1968; Hayward, 1969; Biezanko, Ruffinelli & Link, 1974; Bizarro apud Beccaloni et al., 2008); records of species of Colubrina View in CoL and Colubrina glandulosa View in CoL ( Rhamnaceae View in CoL ), provided by Silva et al. (1968) and Zikán & Zikán (1968), are uncertain and in need of confirmation.

Distribution

Widespread in the Amazon basin and the Guianas, usually in low to mid-altitudes, occurring from about the sea level to 1400 m on the eastern slopes of the Andes, in Argentina, Bolivia, Brazil (Acre, Amazonas, Rondônia, Maranhão, Mato Grosso, Mato Grosso do Sul and Pará), Colombia, Ecuador (Orellana and Morona-Santiago), French Guiana, Guyana, Paraguay, Peru, Suriname, Uruguay and Venezuela; and in Atlantic forests of Brazil (Alagoas, Bahia, Ceará, Espírito Santo, Distrito Federal, Goiás, Minas Gerais, Rio de Janeiro , Rio Grande do Sul, São Paulo , Santa Catarina, Paraíba and Pernambuco) ( Fig. 20). It may occur in northwestern Argentina (i.e. Catamarca, Jujuy, Salta and Tucumán) and in all remaining states of Brazil.

Examined material

See Supporting Information, Appendix S1.