Zaretis, HUBNER, [1819]

ZARETIS HÜBNER, [1819] View in CoL

( FIGS 5–21; SUPPORTING INFORMATION, APPENDIX S6)

Type species: Papilio isidora Cramer, 1779 by subsequent designation ( Scudder, 1875).

Diagnosis

Zaretis View in CoL are brushfoot ( Nymphalidae View in CoL ) leafwings ( Charaxinae View in CoL ), and therefore are butterflies with reduced forelegs and sclerotized parapatagia. The tribe Anaeini View in CoL , in which Zaretis View in CoL is placed, currently does not have any clearly defined synapomorphies. Yet, species of Zaretis View in CoL can be distinguished from the Preponini View in CoL , and most species of related Anaeini View in CoL genera, by the characteristic underside wing pattern resembling dead leaves, labial palpus ( Reuter, 1896; Rydon, 1971), wing venation ( Röber, 1892; Comstock, 1961; Rydon, 1971), male genitalia ( Mielke, Mielke & Casagrande, 2004), and a number of other characters of the immature stages (e.g. Rydon, 1971; Dias, Casagrande & Mielke, 2010). Many of these characters are shared with species of Coenophlebia View in CoL (whose immature stages are unknown), Siderone View in CoL and Phantos gen nov. However, species of Zaretis View in CoL can be distinguished easily from Coenophlebia View in CoL , represented by C. archidona View in CoL , by the presence of deep emarginations in the inner margin near the tornus of the forewing (FW) and between Sc+R and M 1 of the hindwing (HW), and a developed tornal projection at 2A on the HW. Additionally, in C. archidona View in CoL , M 1 and M 2 arise from a common stalk, but separately from the discal cell in species of Zaretis View in CoL , Siderone View in CoL and Phantos gen. nov.

Zaretis View in CoL can be distinguished from Siderone View in CoL and Phantos gen. nov. by the wing shape, colour and pattern, which is variable, but always ranges from pale yellow to reddish brown in both wings, with reddish brown to dark brown markings on the upperside, and the male genitalia, with a conspicuous projection between the costa and the harpe (except for Z. syene View in CoL , in which this projection is reduced and similar to species of Siderone View in CoL and Phantos gen. nov.).

Zaretis can be distinguished from Siderone chiefly by the wing shape, colour and pattern. The wings are never as rounded at the outer margin or bear black with scarlet red bands or markings in species of Zaretis as in Siderone . Zaretis also has FW vein R 4 ending at the costal margin before the apex, marked sexual dimorphism, the head capsule scoli of the larvae lacking protrusions, and pupae without lateral indentations.

Zaretis can be distinguished from Phantos gen. nov. by the stronger development of the thorax, the wing colour and pattern, with the wings never being pearly white or yellowish white on the wing upper side, by R 1, R 2 and R 3 running free to the costal margin of the FW, by the presence of a marked discal spot (element ‘e’) on the forewing dorsum (FWD), the absence of projections at M 3 beyond the external margin of the HW in both sexes, the stronger development of the tornal projection at 2A, and by the relative size and shape of the head capsule scoli of the larvae, which are straight and of the same size or shorter than the height of the head capsule. A key to species of Zaretis is presented below.

Redescription

Head: Eyes reddish brown and naked ( Fig. 6A); labial palpus creamy white ventrally, pale orange to reddish brown dorsally and at the tip ( Figs 5D, E, 6A–C); antennal length about one-third of the forewing length, segments dark brown with some ventral creamy white scaling; club slender and elongated, tips usually lighter. Female as in male, but usually lighter in colour, pale yellow to reddish brown, depending on the species.

Thorax: Dorsally pale orange to reddish brown with scattered brownish and greenish scaling; ventrally pale orange to dark brown, with area between legs creamy white; forelegs with creamy white scales in the tarsus ( Figs 5F, G, 6D, E); mid- and hindleg femora, tibiae and tarsi and creamy white speckled with pale orange to dark brown scales, respectively ( Fig. 5H, I). Female as in male, but usually lighter in colour, pale yellow to reddish brown, depending on the species.

Wing size and shape ( Fig. 5A): Forewing length 2.4–4.6 cm. Forewing costal margin convex; apex usually pointed, rounded in one species, slightly to strongly falcate; outer margin usually sinuous, but almost straight or rounded in some species, smooth to crenulated; inner margin straight, emargination before tornus slightly to strongly developed. Hindwing with emargination at Sc–Rs, outer margin rounded, smooth to crenulated, with a long and rounded projection at 2A; inner margin slightly emarginated near the tornus. Female larger than male, and wing shape usually different; outer margin usually straighter or rounder, depending on the species; emargination before tornus always developed; hindwing proportionately larger than male; emargination at Sc–Rs always developed; apex always projected at Rs; outer margin usually straighter than male; inner margin moderately to strongly emarginated near the tornus.

Wing colour and pattern, upper side ( Fig. 5B): Ground colour of both wings beige to reddish brown with light to dark brown markings, fore- and hindwings usually of similar ground colour, but of different colours in some species. Forewing usually with three distinct bands, formed by light brown to dark brown continuous or interrupted markings: a median band, formed by the discal spot (element ‘e’) and the distal band of the central symmetry system (element ‘f ’); a post-median band, formed by the proximal band of the border symmetry system (element ‘g’); and a marginal band, formed by the parafocal element (element ‘i’). Even when the median band is absent, the discal spot is always present, light to dark brown in colour. The border ocelli and the submarginal and marginal bands (elements ‘h’, ‘i’ and ‘j’) are either indistinct or coalesced with the parafocal element. These three bands delimit four distinct areas of the forewing upperside: the basal area, from the base of the wing to the median line; the post-median area, from the median band to the post-median band; the submarginal area, from the post-median band to the submarginal band; and the marginal area, from the submarginal band to the outer margin. Marginal area is usually darker than other areas, from light to dark brown, similar in colour to the colour of the discal spot; submarginal area, when distinct, of the same colour as the post-median band posterior to M 2, along the outer margin, and coalesced with the marginal band distal to M 2, near the apex; intraspecifically variable presence of hyaline areas (although they occur more frequently in some species than others) in M 3 –CuA 1 and CuA 1 –CuA 2 on or immediately anterior to the median band. Hindwing usually with three distinct bands, similar to those on the forewing. However, in the hindwing the median band runs only to the discal cell, and the discal spot is reduced or absent; the post-median and submarginal bands run almost regularly along the outer margin to the tornal projection at 2A, frequently with rudimentary border ocelli (element ‘h’) between them; anal fold usually lighter in colour, from pale beige to reddish brown; tornal projection at 2A usually darker, with creamy white scales.The hindwing upperside areas are delimited by the bands of the hindwing underside in five distinct areas, described below: basal, median, post-median, submarginal and marginal areas. Long and thin scales of the same colour of the background, in the discal cell, at the base of CuA 1 –CuA 2, and throughout the length of 2A and 3A. Female with bands and areas of both wings as in male, but frequently different in colour, usually lighter.

Wing colour and pattern, underside ( Fig. 5C): Ground colour of both wings beige to reddish brown, with random speckles of scales lighter and darker than the ground colour (‘ripple pattern’), usually resembling a skeletonized dead leaf. Forewing pattern of bands and areas similar to the upper side, but with two additional faint bands, formed by light brown to dark brown scales darker than the ground colour: a basal band, formed by part of the central symmetry system (element ‘c’); and a discal band, formed by the proximal band of the basal symmetry system (element ‘d’). The umbra connects the median band to the post-median band at CuA 1 –CuA 2, forming a continuous line similar to the midrib of a dead leaf. Hindwing pattern of bands and areas are similar to the upperside, but with an additional faint band, the discal band, formed by the proximal band of the basal symmetry system (element ‘d’), as in the forewing. Anterior part of the median band and posterior part of the umbra form a continuous line similar to the midrib of a dead leaf; the anterior part of the umbra, the posterior part of the median band and the post-median band complete the masquerade. Part of the median band posterior to CuA 2 and marginal band are usually not as developed as the others; border ocelli rudimentary, dark brown and surrounded by creamy white scales; apical ocelli usually as scattered creamy white scales. Hindwing with five distinct areas: the basal area, from the base of the wing to the continuous line formed by the medial band and the umbra; the median area, from the medial band to the umbra; the post-median area, from the umbra to the post-median band; the submarginal area, from the post-median band to the marginal band; and the marginal area, from the marginal band to the outer marigin. Female bands and areas of both wings are as in the male, but frequently different in colour, usually lighter.

Abdomen: Dorsally uniform pale orange to reddish brown; ventrally uniform light brown to reddish brown. Female as in male, but lighter in colour, dorsally pale yellow to reddish orange, ventrally beige to reddish brown, depending on the species.

Male genitalia ( Fig. 15A–L): Tegumen trapezoidal in lateral view, dorsally wider and humped posteriad, strongly attached to the uncus, and attached to the gnathos only by membranes; appendix angularis hooked; saccus short, not projected anteriorly; dorsal projection of the saccus ‘C’ shaped and projected dorsad at a right angle. Uncus semitubular, almost straight or slightly curved, with a well-developed median dorsal ridge and distally hooked, usually with a distal callus; gnathos laterally slightly curved, dorsally thin or wide, produced ventrad, arms parallel, with ventral part of the gnathos bar shaped and fused medially; valva externally covered with short setae, costa long and curved, developed anteriad, two projections of varying shapes and development, one between the costa and the harpe, and another at the end of the harpe, sacculus triangular, ampulla developed and rounded; aedeagus varying slightly in length and width, cylindrical and bifid distally, without cornuti; manica inserted slightly anterior to the half of the aedeagus; fultura inferior thin, bar shaped.

Female genitalia ( Figs 16A–L, 17A–J): Tergum VIII triangular, ventrally attached to the sides of the lamella postvaginalis and dorsally to the lamella antevaginalis by a varying sclerotized loop; papilla analis round and with short setae, projecting the apophysis posterioris; lamella antevaginalis assymetrical and variable, connected to the sides of the lamella postvaginalis by projections of varying sclerotization; lamella antevaginalis left side usually larger than the right; lamella postvaginalis varying in width and length; posterior edge of the lamella postvaginalis straight, rounded, smooth, bilobed or medially indented; seminal duct close to the base of the ductus bursae; corpus bursae rounded or laterally compressed, half the length or shorter than the ductus bursae, bearing two parallel signa; these are thin and long, formed by minute sclerotized bumps.

Immature stages ( Fig. 18): All species with known immature stages (namely, Z. strigosus , Z. ellops , Z. crawfordhilli sp. nov., Z. mirandahenrichae Dias sp. nov. and Z. elianahenrichae Dias sp. nov.) are similar to those recently described by Dias et al. (2015) ( Müller, 1886; Muyshondt, 1973; Janzen & Hallwachs, 2017); host plants are mostly in the Salicaceae , chiefly on species of Casearia , but also on species of Laetia , Ryania , Xylosma and Zuelania ( Beccaloni et al., 2008; Dias et al., 2015; Janzen & Hallwachs, 2017); records for species of Zaretis on Rhamnaceae and Piperaceae are doubtful and need confirmation ( Beccaloni et al., 2008).

Discussion

Zaretis View in CoL was erected by Hübner ([1819]) to include Papilio isidora Cramer, 1779 and the Indo-australian nymphaline species Papilio polibete Cramer, 1779 . Inexplicably, Hübner ([1819]), placed Papilio itys Cramer, 1777 in Apatura (Hübner, [1819]) View in CoL . The type species of the genus, P. isidora Cramer, 1779 , was subsequently designated by Scudder (1875). Early authors combined and described species currently in Zaretis View in CoL in Papilio View in CoL (sometimes in the sections ‘Danaus’, ‘Festivus’ and ‘Nobilis’ of the Linnean system), Nymphalis View in CoL (once in the section ‘Gemmatus’ of the Linnean system) and Paphia View in CoL , as a genus or as a subgenus of Maniola View in CoL . However, in the middle of the 19th century, several authors, recognizing the similarities between species in Zaretis and View in CoL Siderone View in CoL , regarded the former as synonymous with the latter, an arrangement preferred by some authors to the present day (e.g. Brévignon, 2006). Nevertheless, Zaretis (Hübner, [1819]) View in CoL has priority over Siderone (Hübner, [1823]) View in CoL . The misspelling Zaretes [sic], probably introduced by Röber (1892), was widely adopted in the late 19th and beginning of the 20th century by several authors who were unaware of the name Zaretis View in CoL or who did not agree with the synonymization of Zaretis View in CoL with Siderone View in CoL . Although others (e.g. Scudder, 1875) had already noted the misspelling, the correct spelling was widely adopted after the catalogue by Stichel (1939). Comstock (1961), in his revision of species currently in Anaeini View in CoL , regarded Zaretis View in CoL as a subgenus of Anaea View in CoL . However, this arrangement was never popular, and later authors tended to lump all species of Anaeini View in CoL in Anaea View in CoL or, more frequently, regarded each one of Comstock’s (1961) subgenera as genera. The latter view was strongly backed up by the most comprehensive taxonomic study of the Charaxinae View in CoL to date, by Rydon (1971), and it is currently the most widely accepted arrangement (e.g. Lamas, 2004; Willmott & Hall, 2004). From early on, the species included here in Phantos gen. nov. were kept apart from species currently in Zaretis View in CoL by many authors, most notably Röber (1916) and Stichel (1939), who retained them in Anaea View in CoL owing to their distinctiveness.

Zaretis View in CoL is currently placed in Anaeini ( Lamas, 2004) View in CoL , but adults and immature stages are different from most of the genera included in that tribe. Even though the monophyly of Anaeini View in CoL is contested by some authors (e.g. Rydon, 1971; Pyrcz & Neild, 1996), the tribe is usually treated as monophyletic (e.g. Comstock, 1961; DeVries, 1987; Lamas, 2004), despite the lack of unambiguous synapomorphies. Species of Zaretis View in CoL , Siderone View in CoL , Coenophlebia View in CoL and Phantos gen. nov. share several common and distinct characters (particularly in the immature stages). Therefore, Rydon (1971) erected the taxon ‘Zaretidinae’ (i.e. Zaretidini), following the lead of previous authors (e.g. Röber, 1892; Reuter, 1896) in considering species of the above genera ‘transitional’ between Preponini View in CoL and Anaeini View in CoL . Rydon’s (1971) ‘Zaretidinae’ is most likely to be monophyletic, although probably sister to the remainder of Anaeini ( Wahlberg et al., 2009) View in CoL .

Distribution

Neotropical, in forest habitats from Mexico to Argentina, except most of the Antilles (occurring only on the island of Trinidad) and Chile, from sea level to 2200 m ( Figs 19–21). Of the 12 species included, five are Trans-Andean, three are widespread in Central America and Trans-Andean South America, and two occur on the western slopes of the Andes. One is restricted to middle elevations of the Andes, and the remaining six species occur in Cis-Andean South America. Of these, one is widespread in all suitable habitats, two occur exclusively in the Amazon basin and eastern slopes of the Andes, two occur in other South American forests further from the Amazon basin, and one is restricted to dense ombrophilous forests on the southeastern coast of Brazil.