Phantos, 2019

PHANTOS DIAS GEN. NOV.

( FIGS 22–27; SUPPORTING INFORMATION, APPENDIX S6)

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Type species: Nymphalis callidryas R. Felder, 1869 , here designated.

Diagnosis

Phantos gen. nov. is closest in appearance to Zaretis , and it can be distinguished from other taxa of Charaxinae by the set of characters cited above for that genus. It can be distinguished from species of Zaretis chiefly by the coloration of the upperside of the wings, which are pearly white or yellowish or greenish white, sometimes with an orangish or yellowish suffusion along the outer margin, with light to dark brown markings only on the submarginal and marginal areas; the underside ground colour is beige, brown, yellow or green, speckled with light and dark shades of these colours. Additionally, Phantos gen. nov. can be distinguished by the labial palpus, which is distally beige to light brown, speckled with creamy white scales ( Fig. 23A–C); the slender thorax; veins R 1, R 2 and R 3 partly anastomosed to Sc; discal spot (element ‘e’) absent on the upperside ( Fig. 22B); external margin of the HW at vein M 3 projected in both sexes, short and pointed in males, long and spatulate in females; tornal projection at 2A at the HW weakly to moderately developed ( Fig. 22A); male genitalia with a pit between the tegumen and uncus; uncus short, curved and evenly keeled; saccus posteriorly projected; gnathos ‘C’ shaped, with the dorsal half always much larger than the ventral; projection between the costa and the harpe weakly developed ( Fig. 25); and relative size and shape of the head capsule scoli of the larvae thin, curved posteriorly and longer than the height of the head capsule; larval colour pattern similar to species of Siderone , dorsally lighter posterior to the A2 hump. A key to species of Phantos gen. nov. is presented below.

Description

Head: Eyes reddish brown and naked; labial palpus creamy white ventrally, distally beige to light brown, speckled with creamy white scales ( Fig. 23A); antennal length about one-third of the forewing length, segments orange to yellowish orange with some ventral creamy white scaling; club slender and elongated, tip orange to yellowish orange. Female as in male.

Thorax: Slender, dorsally mostly creamy white speckled with greyish scales; ventrally beige speckled with creamy white scales; legs mostly beige speckled with creamy white scales, tibiae and tarsi creamy white or speckled with creamy white scales ( Fig. 23D, E). Female as in male.

Wing size and shape ( Fig. 22A): Forewing length, 3.1– 3.9 cm. Forewing costal margin convex; apex pointed, slightly to moderately falcate; outer margin slightly sinuous to sinuous, smooth to slightly crenulated; inner margin straight, emargination before tornus shallow and long, about one-third of the length of the inner margin. Hindwing with emargination at Sc–Rs, outer margin angled and projected beyond the outer margin at M 3, smooth to crenulated; tornus slightly to moderately projected at 2A; inner margin slightly emarginated near the tornus. Female larger than male, and wing shape different; FW outer margin rounded; HW emargination at Sc–Rs developed; apex projected at Rs; outer margin rounder, with a long, spatulated projection at M 3, crenulated; inner margin moderatedly to strongly emarginated near the tornus.

Wing colour and pattern, upper side: Bands and areas equivalent to Zaretis , but pattern much reduced; ground colour of both wings mostly pearly white or yellowish white, with or without an orangish or yellowish suffusion along the outer margin. Marginal and submarginal areas light to dark brown, formed by the coalesced part of the post-median band (element ‘g’) at the apex and the marginal band (elements ‘h’, ‘i’ and ‘j’) along the outer margin. Hindwing usually with two distinct bands, formed by interrupted light to dark brown markings; the post-median and submarginal bands run regularly along the outer margin to the inner margin before the tornal projection at 2A, frequently with rudimentary border ocelli (element ‘h’) between them; anal fold similar in colour to the rest of the HW, with some beige scaling. Female as in male, but usually lighter in colour.

Wing colour and pattern, underside ( Fig. 22B): Ground colour of both wings beige, brown, yellow or green, speckled with light and dark shades of these colours (‘ripple pattern’). Forewing pattern of bands and areas equivalent to Zaretis ; basal, discal, median (including the discal spot), post-median and marginal bands subtle, beige to brown; median and post-median bands close to each from the outer margin to CuA 1; ground colour of the different areas variable, but the basal, submarginal and marginal areas usually darker than post-median area; border ocelli in R

3

–R

4

, R

4

–R

5

and R 5 –M 1 usually greatly enlarged. Hindwing pattern of bands and areas equivalent to Zaretis ; anterior part of the median band and posterior part of the umbra forming a continuous and marked line from the costal to the inner margin, similar to the midrib of a dead leaf; discal, post-median and marginal bands and the remainder of the median band and umbra subtle, beige to brown; rudimentary border ocelli between post-median and submarginal bands. Female as in male, but usually lighter in colour.

Abdomen: Dorsally and laterally creamy white speckled with thin grey scales; ventrally beige speckled with creamy white scales. Female as in male.

Male genitalia ( Fig. 25): Tegumen trapezoidal in lateral view, dorsally wider and humped, attached to the anterior part of the uncus forming a median pit, and attached to the gnathos only by membranes; appendix angularis hooked; saccus short, projected anteriorly, dorsal projection of the saccus ‘C’ shaped and projected dorsad at about a right angle; uncus semitubular and curved, with a well-developed and even keel and distally pointed; gnathos laterally slightly curved, dorsal half wider than ventral, produced ventrad, arms parallel, ventral part of the gnathos bar shaped, slighty angled in ventral view and fused medially; valva externally covered with short setae, costa long and curved, developed anteriad, end of the harpe pointed, angled between the costa and the harpe, sacculus triangular, ampulla developed and rounded; aedeagus short, cylindrical and bifid distally; vesica without cornuti; manica inserted about the half of the aedeagus; fultura inferior thin, bar shaped.

Female genitalia ( Fig. 26): Tergum VIII triangular and thin, dosally and anteriorly projected, posteriorly attached to the lamella antevaginalis by a sclerotized loop, ventrally attached to the sides of the lamella postvaginalis; papilla analis rounded and with short setae, projecting the apophysis posterioris; lamella antevaginalis assymetrical, connected to the sides of the lamella postvaginalis by slightly sclerotized prejections; lamella antevaginalis thin, left side larger than the right; lamella postvaginalis wider than long, with a membranous posterior projection longer than the lamella postvaginalis, posterior edge of the lamella postvaginalis slightly convex to slightly bilobed; seminal duct close to the base of the ductus bursae; corpus bursae rounded, half the length of the ductus bursae, bearing two parallel signa, which are thin and long, formed by minute sclerotized bumps.

Immature stages: Generally similar to species of Zaretis and Siderone ( Lichy, 1976; Muyshondt, 1976, 2005). Early instars almost indistinguishable from species of Zaretis and Siderone , but fifth instar head capsule smoother, scoli curved backwards, longer and thinner in comparison with species of Zaretis ; projections of A2 smaller and dorsal coloration posterior to A2 lighter than from T1 to A2, and from the same area of species of Zaretis ; pupa similar to species of Zaretis ; host plant records all on the Salicaceae , with reliable records on two species of Casearia , C. sylvestris and C. nitida ( Lichy, 1976; Muyshondt, 1976; DeVries, 1987; Beccaloni et al., 2008; Janzen & Hallwachs, 2017).

Discussion

Phantos gen. nov. is erected to include two species, P. callidryas comb. nov., here designated as the type species of the genus, and P. opalina comb. nov., stat. rev., removed from synonymy; Siderone clara Staudinger, 1886 View in CoL is recognized as a junior subjective synonym of the latter name.These species are superficially similar to species of Zaretis and View in CoL , to some extent, of Siderone View in CoL . However, since the middle of the 19th century, several authors have already combined and described species here included in Phantos gen. nov. in Nymphalis View in CoL , Paphia View in CoL or Anaea View in CoL because of their peculiar morphology. The anastomosis of R 1, R 2 and R 3 with Sc in the FW, and the presence of projections in M 3 on the HW of males and females, similar to many other species of Anaeini View in CoL , certainly encouraged authors, such as Godman & Salvin (1884) and Staudinger (1887), to retain them in the catch-all genus Anaea View in CoL , together with other species of uncertain affinity, instead of including them in Zaretis View in CoL . Rydon (1971), in contrast, included the species in Zaretis View in CoL , with some doubts. Nevertheless, it is clear that the genus represents a distinct lineage of Anaeini View in CoL closely related to Zaretis and View in CoL Siderone View in CoL . Molecular analyses, as shown by the NJ analysis and by EFA Toussaint, FMS Dias, OHH Mielke, MM Casagrande, CP Sañudo-Restrepo, A Lam, J Morinière, M Balke, and R Vila (unpublished data), suggest that Phantos gen. nov. is probably sister to Zaretis View in CoL + Siderone View in CoL , rendering Zaretis, sensu Lamas (2004) View in CoL and Willmott & Hall (2004), polyphyletic. The description of a new genus to include these species is advisable to convey more information and be less disruptive to taxonomy than the alternative of synonymizing Zaretis View in CoL with Siderone View in CoL .

Etymology

The name of the new genus is derived from the greek word φαντός (‘ phantós ’), meaning ‘visible’, ‘present to the eye or mind’, treated as a singular, masculine noun in the nominative case ( ICZN 1999, Art. 11.8). The word ‘ phantós ’ is the root to several English words, such as ‘phantom’ and ‘fantasy’. The name of the new genus is intended to make reference to the popular name of the species included, ghost leafwings ( Glassberg, 2007), a name probably given because of their pale colour and swift flight, and additionally to pay tribute to new molecular methods that ‘make visible’ many cryptic species and genera.

Distribution

Neotropical, in forests from about sea level to 1200 m throughout Central America except the Antilles, from Mexico to Panama ( Fig. 27).