Zaretis elianahenrichae, 2019

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 : 176-179

ZARETIS ELIANAHENRICHAE DIAS SP. NOV.

( FIGS 14A–D, 15L, 17I, J, 18C, D, 21; SUPPORTING INFORMATION, APPENDIX S6)

[u r n:l s i d: z o o b a n k. o r g: a c t: 4 3 0 E 1 A D 4 - B D 4 8 - 4F5B-A906-30B2A47FD113]

Type material

Male holotype with the following labels: / HOLOTYPUS / HOLOTYPE Zaretis elianahenrichae Dias det. 2016 / Voucher: D.H. Janzen & Winnie Hallwachs caterpillar ( Lepidoptera ) DB: http://janzen.sas.upenn. edu Área de Conservación Guanacaste, Costa Rica [Guanacaste, Área de Conservación Guanacaste, Sector Pitilla, Sendero Orosilito , 900 m (10.98332, −85.43623 [10°58′60″N, 85°26′10″W]), 8-XI-2003, Rios leg. ex larvae in Casearia arborea ], 03-SRNP-21709 / LEGS AWAY FOR DNA / DZ 30.203 / (DZUP). Allotype female with the following labels: / ALLOTYPUS / ALLOTYPE Zaretis elianahenrichae Dias det. 2016 / Voucher: D.H. Janzen & Winnie Hallwachs caterpillar ( Lepidoptera ) DB: http://janzen.sas.upenn.edu Area de Conservación Guanacaste, Costa Rica [Guanacaste, Área de Conservación Guanacaste, Sector Del Oro, Sendero Puertas, 400 m (11.01087, −85.48817 [11°0′39″N, 85°29′17″W]), 18-VI-2003, Garcia leg. ex larvae in Casearia sylvestris ], http://janzen.sas.upenn. edu 03-SRNP-15898 /LEGS AWAY FOR DNA / DNA FER036 wg, EF1a, CAD /DZ 30.113 / (DZUP).

Paratypes: COSTA RICA, Alajuela, Área de Conservación Guanacaste, San Cristobal , Buenos Aires , 400 m, 1 #m, 24.XI.2006, Cano leg., 06-SRNP-9573 ( UPENN) ; San Cristobal, Puente Palma , 460 m, 1 #m, 28.XI.2006, Córdoba leg., 06-SRNP-9681 ( UPENN) ; Guanacaste, Área de Conservación Guanacaste, Brasilia, Morales , 440 m, 1 #m, 10.XI.2011, Carmona leg., 11-SRNP-66261 ( UPENN) ; Del Oro, Sendero Puertas , 400 m, 1#m & 2 #f, 19.IX.2003, Rios leg., 03-SRNP-28921, 03-SRNP-15897, 04-SRNP-27147 ( UPENN) ; Del Oro, Sendero Puertas , 400 m, 1 #f, 19.IX.2003, Rios leg., 03-SRNP-28921 ( UPENN) ; 1 #f, 26.IX.2005, Cantillano leg., 05-SRNP-24281 ( UPENN) ; Pitilla, Bullas , 440 m, 1 #f, 29. VI.2012, Calero leg., 12-SRNP-71521 ( UPENN) ; Pitilla, Ingas , 580 m, 1 #f, 5.VII.2012, Rios leg., 12-SRNP-31086 ( UPENN) ; Pitilla, Loaiciga , 445 m, 1 #m, 29. VI.2004, Rios leg., DZ 30.112 ( DZUP) ; 1 #f, 21.VII.2003, Moraga leg., 03-SRNP-20263 ( UPENN) ; Pitilla, Medrano , 380 m, 1 #m & #f, 11.IX.2012, Calero leg., 12-SRNP-72120, 12-SRNP-72121 ( UPENN) ; 1 #m, 4.VII.2012, Calero leg., 12-SRNP-71601 ( UPENN) ; 1 #m, 4.VII.2012, Martínez leg., 12-SRNP-71603 ( UPENN) ; 1 #m, 5.VII.2012, Calero leg., 12-SRNP-71664 ( UPENN) ; Pitilla, Pasmompa , 440 m, 1 #m, 30.III.2003, Cantillano leg., 03-SRNP-1845 ( UPENN) ; Pitilla, Quebradona , 475 m, 1 #f, 24.X.2010, Calero leg., 10-SRNP-73168 ( UPENN) ; Pitilla, Sendero Evangelista , 660 m, 1 #m, 15.X.2004, Rios leg., 04-SRNP-55792 ( UPENN) ; Pitilla, Sendero Memos , 740 m, 1#m & 1 #f, 21.IV.2005, Rios leg., DZ 30.209 ( DZUP) , 05-SRNP-31614 (UPENN); San Cristobal , 460 m, 1 #m, 4.IX.2006, Sihezar leg., 06-SRNP-7337 ( UPENN) ; MEXICO, Chiapas, La Independencia, San Antonio Buena Vista , 1 #m, 1.VIII.1992, Ibarra leg., AIV194 ( UNAM) .

Diagnosis

Zaretis elianahenrichae sp. nov., as with most species of the genus, is somewhat variable and similar to its congeners. This species is likely to be sympatric with Z. ellops , Z. pythagoras , Z. delassisei , Z. crawfordhilli sp. nov. and Z. mirandahenrichae sp. nov. Zaretis elianahenrichae sp. nov. strikingly resembles Z. ellops , but the male genitalia are distinct; the uncus is longer, thinner and curved, with a small distal callus; and the dorsal and ventral halves of the gnathos are about the same size ( Fig. 15L). Males ( Fig. 14A, B) can be externally distinguished from Z. crawfordhilli sp. nov. and Z. mirandahenrichae sp. nov. by the generally much lighter and speckled coloration, with post-median and submarginal bands usually present in both uppersides of the wings, especially the HWD post-median band, which usually runs to the tornal projection at 2A; additionally, Z. elianahenrichae sp. nov. can be distinguished from Z. crawfordhilli sp. nov. by the fully developed emargination of the inner margin of the FW at the tornus; and from Z. pythagoras and Z. delassisei by the absence of a purplish sheen on the FWD when viewed obliquely. Additionally, it can be distinguished from Z. pythagoras by the lighter colour of the upperside of the wings, pale orange to orange; and from Z. delassisei by the FWD pale orange to orange coloration not restricted to the basal and part of the post-median areas and the HW tornal projection at 2A longer; and from Z. ellops by the shape of the FW apex, moderately to strongly falcate; coloration of the submarginal area near the apex and the marginal area, solid pale orange to orange; bands of the wings upperside generally darker and developed, orange to brown; and HW tornal projection at 2A shorter and wider. Females ( Fig. 14C, D) can be distinguished from Z. crawfordhilli sp. nov. and Z. mirandahenrichae sp. nov. by the generally lighter and homogeneous coloration of basal, post-median and submarginal areas along the outer margin of the upper and undersides of the wings; from Z. pythagoras by the shape of the FW apex, considerably less falcate, and the coloration of the post-median and submarginal areas along the outer margin of the FW, lighter and never suffused with scales of different colour; and from Z. ellops by the coloration of the the submarginal area near the apex and the marginal area, solid brown to dark brown; FWD and HWD dissimilar in colour, basal and post-median areas and submarginal area along the outer margin darker in HW; and HW tornal projection at 2A longer and thinner; females of Z. delassisei are unknown.

Description

Head: Eyes reddish brown and naked; labial palpus creamy white ventrally, pale orange to orange dorsally and at the tip; antennal length about one-third of the forewing length, segments orange with some ventral creamy white scaling; club slender and elongated, tip pale orange to orange. Female as in male, but usually lighter in colour, pale yellow to yellow.

Thorax: Dorsally pale orange to orange with scattered brownish and greenish scaling; ventrally pale orange to orange, with area between legs creamy white; forelegs with creamy white scales in the tarsus; mid-leg femora, tibiae and tarsi and hindleg tibiae and tarsi creamy white, speckled with pale orange to reddish brown scales. Female as in male, but usually lighter in colour, pale yellow to yellow.

Wing size and shape: Forewing length, medium. Forewing costal margin convex; apex pointed and falcate; outer margin sinuous, slightly crenulated to crenulated; inner margin straight, emargination before tornus developed, about one-third the length of the inner margin. Hindwing with slight emargination at Sc–Rs; outer margin rounded, smooth to slightly crenulated, with a developed projection at 2A, long and thin; inner margin almost straight. Female larger than male, and FW shape rounder and more falcate; emargination before tornus developed; hindwing proportionately larger than male; emargination at Sc–Rs developed; apex projected at Rs; outer margin straighter than male; inner margin emarginated near the tornus.

Wing colour and pattern, upper side: Ground colour of both wings pale orange to orange with light brown to brown markings, fore- and hindwings of similar ground colour. Forewing basal, median and submarginal areas along the outer margin about the same colour, but post-median band slightly lighter; median and post-median bands faint; submarginal area near the apex and marginal area coalesced, brown to dark brown, along the outer margin to the apex; discal spot of the same colour, usually faint; presence of hyaline areas in M 3 –CuA 1 and CuA 1 –CuA 2 on the median band variable, but usually present. Hindwing areas of the same colour, orange to reddish brown; median, post-median and submarginal bands faint, more noticeable near the costal margin; discal spot absent; border ocelli faint or absent; anal fold lighter in colour; tornal projection at 2A usually darker with some creamy white scaling. Female forewing basal, post-median and submarginal area along the outer margin usually coalesced, pale yellow to yellow; discal spot, submarginal area near the apex and marginal area brown to dark brown, the two latter coalesced; median and post-median bands faint or absent. Hindwing marginal area usually darker, pale orange to orange, lighter towards the wing base; median, post-median and submarginal bands faint, more noticeable near the costal margin; discal spot absent; border ocelli faint or absent; anal fold lighter in colour; tornal projection at 2A usually darker with some creamy white scaling.

Wing colour and pattern, underside: Ground colour of both wings reddish brown to brown, with random speckles of scales lighter and darker than the ground colour in a ripple pattern. Forewing areas of similar colour, post-median and submarginal areas along the outer margin lighter, with yellowish scaling; all bands noticeable, slightly darker than the ground colour, but submaginal band from the inner margin to CuA 1 darker and more distinct; border ocelli faint, formed by dark brown and creamy white scales, scattered near the apex. Hindwing areas similar in colour to the FW; median and post-median areas lighter, with yellowish scaling; all bands noticeable, slightly darker than the ground colour, but median and post-median bands and the posterior part of umbra darker and more distinct. Female forewing basal area pale yellow to yellow; post-median and submarginal areas along the outer margin pale yellow; marginal and submarginal areas near the apex darker than other areas, orange to brown; all bands noticeable, orange to brown. Hindwing basal area darker, pale yellow to yellow; median, post-median, submarginal and marginal areas pale yellow; all bands noticeable, orange to brown, but median and post-median bands and the posterior part of umbra darker and more distinct; border ocelli faint, formed by dark brown and creamy white scales; tornal projection at 2A usually with some creamy white scaling.

Abdomen: Dorsally uniform pale orange to orange; ventrally reddish brown to brown. Female dorsally pale yellow to yellow and ventrally darker, pale yellow to yellow.

Male genitalia ( Fig. 15L): Tegumen trapezoidal in lateral view, dorsally wider, 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 about the same size as the tegumen, semitubular, thin and lightly curved, with a well-developed median dorsal ridge, distally hooked and with a developed ventral callus; gnathos laterally slightly curved, dorsally thin, produced ventrad, arms parallel, ventral part of the gnathos bar shaped and fused medially; valva externally covered with short setae; costa long and curved, developed anteriad, with two developed projections, one between the costa and the harpe, and another at the end of the harpe; sacculus triangular, ampulla developed and rounded; aedeagus longer than the length of the tegumen and uncus combined, cylindrical and bifid distally, without cornuti; manica inserted slightly anterior to the half of the aedeagus; fultura inferior thin, bar shaped.

Female genitalia ( Fig. 17I, J): Tergum VIII triangular, ventrally attached to the sides of the lamella postvaginalis, and dorsally to the lamella antevaginalis by a dorsally projected loop, thickened after the loop; papilla analis rounded and with short setae, projecting the apophysis posterioris; lamella antevaginalis assymetrical, connected to the sides of the lamella postvaginalis by wide sclerotized projections; left side wider and larger than the right; lamella postvaginalis longer than wide, anterior area with a small membranous area, and posterior edge bilobed; seminal duct close to the base of the ductus bursae; corpus bursae laterally compressed, about the half the length of the ductus bursae, bearing two parallel signa, these are thin and long, formed by minute sclerotized bumps.

Discussion

The description of Z. elianahenrichae sp. nov. is based on 26 barcoded specimens, 14 male and 11 female specimens from Costa Rica (Guanacaste and Alajuela) deposited in UPENN, two male and two female specimens, deposited at the DZUP, and one male deposited at the UNAM; only barcoded specimens are designated as paratypes. This species is strikingly similar to Z. ellops , although easily and reliably distinguished from it by examining the genitalia. Zaretis elianahenrichae sp. nov., according to the data presented by Janzen & Hallwachs (2017), is not as common as Z. ellops and Z. crawfordhilli sp. nov. Similar to Z. crawfordhilli sp. nov. and Z. mirandahenrichae sp. nov., it occurs on the Atlantic slopes of Costa Rica. This species was reared multiple times by Janzen & Hallwachs (2017) ( Fig. 18C, D), on C. arborea , C. sylvestris and one record on Zuelania guidonia . Most immature stages use C. arborea (~56% of the records) and C. sylvestris (40% of the records) as host plants. The former is the preferred host plant of both Z. mirandahenrichae sp. nov. and Z. crawfordhilli sp. nov., but the latter is at best secondary to all species of Zaretis occurring in Costa Rica, although frequently cited as host plants of Z. strigosus and Z. ellops elsewhere.

Distribution

Probably widespread in Central America, except the Antilles, and Trans-Andean South America; however, it is currently known to occur only in forest habitats from 380 to 900 m in Costa Rica, Mexico (Chiapas) and western Ecuador ( Fig. 21). It may occur in the remaining countries of Central America, except the Antilles, and other countries in Trans-Andean South America.

Etymology

Zaretis elianahenrichae sp. nov. is named for Eliana Henrich, in recognition of her mother Charlotte Hill’s generous support of taxonomic resolution of the caterpillar parasitoids of this and many other species of Lepidoptera inhabiting the Área de Conservación Guanacaste rain forest.

Examined material

See Supporting Information, Appendix S1.

List of unavailable names and nomina nuda not assigned to any taxon

Zaretes [sic] isidora vulpina lutulenta Talbot, 1923: p. 52; infrasubspecific, nom. nud.

Zaretes [sic] isidora vulpina f. polcarea Talbot, 1923: p. 52; infrasubspecific, nom. nud.