Zaretis isidora, (CRAMER, 1779)

Dias, Fernando Maia Silva, Janzen, Daniel, Hallwachs, Winnie, Chacón, Isidro, Willmott, Keith, Ortiz-Acevedo, Elena, Mielke, Olaf Hermann Hendrik & Casagrande, Mirna Martins, 2019, DNA barcodes uncover hidden taxonomic diversity behind the variable wing patterns in the Neotropical butterfly genus Zaretis (Lepidoptera: Nymphalidae: Charaxinae), Zoological Journal of the Linnean Society 185, pp. 132-192 : 150-156

ZARETIS ISIDORA ( CRAMER, 1779)

( FIGS 7E–H, 15B, 16C, D, 20; SUPPORTING INFORMATION, APPENDIX S6)

Type material

Male lectotype of Papilio isidora Cramer, 1779 , here designated, with the following labels: / N. 35 ISIDORA Cr [amer]. III.235.A.B.E.F/ Surinam [e] Coll[ection]. [C. van] Len[n]ep/ Felder Coll[ectio]n./ Papilio isidora Cramer det. R.I. Vane-Wright, 1972/? SYNTYPE #m/ (NHMUK).

Male holotype of Siderone zethus Westwood, 1850 with the following labels: /Type/ Para [Pará, Brazil]/ B.M. TYPE No. Rh.10379 Siderone zethus , #m Hew[itson]./ Siderone zethus Westwood det. R.I. Vane-Wright, 1972 HOLOTYPE #m/ zethus Westwo [od]G[eneraof].D[iurnal]. L[epidoptera]. [volume] 5 [page] 321/ (NHMUK).

Male lectotype of Siderone isidora var. cacica Staudinger, 1887 , here designated, with the following labels: /Origin / Chanchamayo[, Peru] Thamm [leg.]/ cacica Stgr. [Staudinger]/ Sid [erone]. spec[ies]. [illegible] ign[ota]./ (ZMHU).

Zaretes [sic] isidora isidora f. bisaltina 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: / Type / Type / SYN-TYPE/ Fruhstorfer Coll. B.M. 1987-285. /Obidos [Pará, Brazil] Amazonenstrom [Amazon river] VIII-IX [18]99 ex coll. H. Fruhstorfer/ isidora #f forma bisaltina Fruhst [orfer]./ (NHMUK).

Male lectotype of Zaretes [sic] isidora leopoldina Fruhstorfer, 1909 , here designated, with the following labels: / Paratype / Fruhstorfer Coll. B.M. 1987-285. / Espírito Santo Brasil ex coll. Fruhstorfer/ Zaretes [sic] isidora leopoldina Fruh [storfer]. det. R.I. Vane-Wright, 1972 #m SYNTYPE./ leopoldina Fruhst [orfer]./ (NHMUK).

Zaretis itys itys forma monops Bryk, 1953 does not have a formal type because it is an infrasubspecific entity. However, the specimen described by Bryk (1953) is a male with the following labels: / Zaretis itys itys f[orm]a. monops m[ale] F. Bryk det. 194[blank]/ Sv. Amaz[on]. Exp[editon]. Roman/ Manáos [Manaus, Amazonas, Brazil]/ 24.nov[ember]/ (RMS).

The male holotype of Siderone isidora naama Brévignon, 2006 was not examined, but has the following label data: French Guiana, Saint-Georges de l’Oyapock , 13.IV.1984, Number 48 (Trap captured) (Bévignon, 2006) ( LCB) .

Diagnosis

Zaretis isidora , as with most species of the genus, is extremely variable and similar to its congeners. This species is sympatric with Z. itys , Z. strigosus , Z. falcis and Z. hurin sp. nov. in the Amazon basin and the Guianas, and with Z. strigosus and Z. itylus in Atlantic forests. Males ( Fig. 7E, F) can be distinguished from Z. itys by the well-developed emargination of the inner margin at the FW tornus; from Z. falcis by the FW apex, which is considerably less falcate, and by the FW outer margin being less crenulated; from Z. hurin sp. nov. by the absence of a purplish sheen on the FWD of Z. isidora when viewed obliquely; from Z. itylus by the shape of the outer margin of the FW, which is sinuous, and HW not strongly emarginated in Sc–Rs and projected at Rs at the apex as in Z. itylus ; and from Z. strigosus by the generally darker and more homogeneous coloration, with post-median and submarginal bands absent or weakly developed on both wings on the upperside. Owing to the extreme variation of males of Z. isidora and Z. strigosus , the identification of the males can be confirmed only by examining the genitalia; in Z. isidora , the uncus is longer, thinner and curved, with a distal callus, and the gnathos is not enlarged at the dorsal half ( Fig. 15B). Females of Z. isidora ( Fig. 7G, H) can be distinguished from females of Z. itys and Z. itylus by the shape of the outer margin of the FW, which is sinuous, and by the upperside of the wings’ basal and post-median areas, never reddish brown and beige to pale yellow, as in Z. itys and Z. itylus , repectively; from Z. falcis by the FW apex, which is considerably less falcate, by the outer margin not being crenulated, and by the FW post-median band, which is usually weakly developed; from Z. hurin sp. nov. by the generally lighter coloration, especially at the wing base, and by the upperside of the wings bands being usually more weakly developed; and from Z. strigosus by the two-tone pattern of the wing underside, which is variably darker in the basal, submarginal area near the apex and marginal areas, in contrast to lighter post-median and submarginal areas along the outer margin. Nevertheless, owing to the extreme variation of females of Z. isidora and the latter two species, identification with respect to Z. strigosus can be confirmed only by the examination of the genitalia; in Z. isidora , the lamella postvaginalis is longer than wider, with the edge slightly indented medially ( Fig. 16C, D); identification with respect to Z. hurin sp. nov. can be reliably confirmed only with molecular data, although there may be slight differences between the genitalia of the species.

Discussion

Zaretis isidora is the type species of the genus Zaretis , selected by subsequent designation by Scudder (1875). Therefore, the resolution of its taxonomic status is paramount. Papilio isidora Cramer, 1779 was described based on an unstated number of specimens from Suriname, known to be at least two, one male and one female [illustrated by Cramer (1779) in plate 235, fig. A, B, E, F]. Vane-Wright (1975) found two specimens, one male and one female, from Suriname similar to the illustrations, in the C. van Lennep collection at the NHMUK and labelled these specimens as potential syntypes (‘?SYNTYPE’). Van Lennep’s collection was the source of many of Cramer’s specimens; his collection was later acquired by the Felders, then by Rothschild, and finally by the NHMUK. None of these specimens could be located by us in the type collection of the NHMUK, and they must be deposited in the main collection. However, the male specimen was illustrated by Pyrcz & Neild (1996) and Warren et al. (2016) and is closely comparable with Cramer’s original illustrations and with specimens of Z. isidora from Suriname, the Guianas and the Amazon basin. Therefore, in the present study, this specimen is designated as the lectotype to reduce present and future taxonomic uncertainty. Siderone zethus Westwood, 1850 was described based on a single specimen from Pará, examined by Vane-Wright (1975) and illustrated by Warren et al. (2016). The phenotype of the holotype compares almost perfectly with specimens of Z. isidora from the Amazon basin and the Guianas, but with a rather falcate FW apex. This specimen is similar in this regard to the somewhat distinctive Z. falcis , but other characters of wing shape and coloration clearly identify it as Z. isidora .

Siderone isidora v. cacica Staudinger, 1887 was described based on a ‘small number’ of specimens from Chanchamayo, Peru ( Staudinger, 1887). This name refers to the typical phenotype of Z. isidora from certain parts of Peru and Bolivia, with suffused reddish brown on the upperside submarginal area near the apex and marginal areas, and which is larger than typical Z. isidora from the Amazon basin. Five male syntypes with the Staudinger’s typical pink ‘Origin’ label collected by Mr Thamm in Chanchamayo, Peru, were located at the ZMHU. Of these, only two agree with the species description, two are specimens of Z. strigosus , and one certainly is a mislabelled specimen of Z. itylus . Therefore, one of the specimens that agrees perfectly with the species description and was illustrated by Warren et al. (2016) is here designated as the lectotype.

Zaretes [sic] isidora isidora f. bisaltina Fruhstorfer, 1909 was described as a ‘female form’, denoting an infrasubspecific entity. Therefore, the name is invalid and without a formal type, but it corresponds to a distinct phenotype of females of Z. isidora from the lower Amazon river basin, with the anterior part of the post-median band, between M 3 and the costal margin, yellowish and separated from the rest of the post-marginal band by a dark brown bar, which connects the discal spot to the submarginal and marginal areas. Thus, it is similar in appearance to species of Doleschallia C. Felder & R. Felder, 1860 , especially Doleschallia bisaltide ( Cramer, 1777) ( Nymphalinae ). One of Fruhstorfer’s specimens, illustrated by Warren et al. (2016), is kept at the type collection of the NHMUK.

Zaretes [sic] isidora leopoldina Fruhstorfer, 1909 , was described based on an unstated number of specimens from Espírito Santo, Brazil. Only one syntype was located by Vane-Wright (1975) and by us in the NHMUK type collection. The specimen agrees perfectly with specimens of Z. isidora from the dense ombrophilous forests along the Brazilian coast, differing from the much more common Z. strigosus , as the upperside of the wings has a generally darker ground colour and the post-median and submarginal bands are absent or faint, not reaching the tornus in the hindwing dorsum (HWD). Therefore, this specimen, illustrated by Warren et al. (2016), is here designated as the lectotype.

Zaretis itys itys f. monops ( Bryk, 1953) was described from a single specimen from Manaus, Amazonas, Brazil. However, it was described denoting an infrasubspecific entity and, therefore, the name is invalid and does not have a formal type. This specimen, similar to the type of P. isidora Cramer, 1779 and specimens of Z. isidora occurring in the Amazon basin, was inadvertently described based on the presence of only one hyaline area (thus ‘ monops ’, ‘single eye’) in the post-median area of the FW, in CuA 1 –CuA 2. However, the presence or absence of hyaline areas in the FW is variable in species of Zaretis .

Siderone isidora naama Brévignon, 2006 was described based on nine specimens from French Guiana, five males and four females. The holotype is similar to the type of P. isidora Cramer, 1779 and specimens of Z. isidora from the Amazon basin and Guianas; the allotype is similar to Fruhstorfer’s infrasubspecific female form ‘ bisaltina ’ ( Fruhstorfer, 1909b), similar to specimens from the lower Amazon River basin. Therefore, this name is here considered to refer to a species of the genus Zaretis and treated as a junior subjective synonym of Z. isidora (comb. nov., syn. nov.).

Distribution

Widespread in the Amazon basin and the Guianas, usually occurring in low to mid-altitudes from sea level to 1400 m elevation in the eastern slopes of the Andes, in Bolivia, Brazil (Acre, Amazonas, Rondônia, Mato Grosso and Pará), Colombia, Ecuador, French Guiana, Guyana, Peru, Suriname and Venezuela; and, disjointly, in Atlantic forests of Brazil (Alagoas, Bahia, Espírito Santo, Distrito Federal, Minas Gerais, Rio de Janeiro , São Paulo and Pernambuco) ( Fig. 20). It may occur in northwestern Argentina (i.e. Catamarca, Jujuy, Salta and Tucumán) and further south and east in Brazil (i.e. Goiás, Maranhão, Mato Grosso do Sul, Paraná , Tocantins and Roraima).

Examined material

See Supporting Information, Appendix S1.