Zaretis mirandahenrichae, 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 : 174-176

ZARETIS MIRANDAHENRICHAE DIAS SP. NOV.

( FIGS 13E, F, 15K, 17G, H, 18G, 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: 7 3 7 B 7 B 8 3 - B 0 6 E - 41C4-A773-4A9FE3429885]

Type material

Male holotype with the following labels: / HOLOTYPUS / HOLOTYPE Zaretis mirandahenrichae Dias det. 2016 / COSTA RICA. Prov. Heredia, Sarapiquí, Est. Biol. La Tirimbina, Mirador , 167 m [10°25′6″N, 84°7′7″W], 24 ABR [IV] 2009, I. Chacón, A. Barrientos [leg.], Tp. Luz. [light trap] L_N_266268_523359 #96598/ [barcode] INB0004211690 INBIOCRI COSTA RICA / Zaretis isidora DHJ 02/ Legs away for DNA I. A. Chacón 2010/ DZ 30.206/ (DZUP). Female allotype with the following labels: / ALLOTYPUS / ALLOTYPE Zaretis mirandahenrichae Dias det. 2016 / COSTA RICA. Prov. Heredia, Sarapiquí, Starke. O a 100 m [10°26′16″N, 84°0′43″W]. 29 JUN [VI] 2010. E. Calderón, M. Cruz [leg.]. Tp. de Fruta [fruit-baited trap] #15 (Dosel [canopy]) L_N_269206_536862 #101206 /[barcode] INB0004277346 / Costa Rica-Heredia, Sarapiquí, Starke TA 15- 10°26.485′N 83° 59.684 Eduardo Calderón Espinoza & Miguel Cruz Artavia 6/29/2010 Zaretis isidora, Hembra Det. I.A. Chacón Julio 2010 / Legs away for DNA I.A. Chacón 2010/ Zaretis isidora DHJ 02 Det: I.A. Chacón feb[uary] 2013/ DZ 30.211/. (DZUP).

Paratypes: COSTA RICA, Alajuela, Área de Conservación Guanacaste, Rincón Rain Forest, Camino Río Francia, 410 m, 1 #m, 11.X.2003, Carmona leg., DZ 30.114 (DZUP); Rincón Rain Forest , Quebrada Escondida , 420 m, 2 #m, 22.III.2005, Pérez leg., DZ 30.111 (DZUP),05-SRNP-40837(UPENN); RincónRain Forest, Quebrada Guarumo , 400 m, 1 #f, 30.VIII.2001, Vargas leg., 01-SRNP-5691 (UPENN); Rincón Rain Forest, Sendero Rincón , 430 m, 1 #f, 18.X.2011, Córdoba leg., 11-SRNP-44587 (UPENN); 1#m & 1 #f, 25.XI.2002, Pérez leg., 02-SRNP-21453, 02-SRNP-21455 (UPENN); San Cristobal, Sendero Huerta , 527 m, 1 #m & 1#f 10.V.2012, Cano leg., 12-SRNP-1873, 12-SRNP-1874 (UPENN); Guanacaste, Pitilia, Sendero Paleta, 570 m, #f, 23.VIII.2004, Moraga leg., 04-SRNP-34810 (UPENN); Pitilla, Bullas , 440 m, 1 #m, 15.VI.2012, Calero leg., 12-SRNP-71413 (UPENN); Pitilla, Ingas , 580 m, #m, 7.III.2009, Moraga leg., 09-SRNP-30712 (UPENN); 1 #m, no date, Rios leg., DZ 30. 210 (DZUP); Pitilla, Loaiciga , 445 m, #m, 29.IX.2005, Espinosa leg., 05-SRNP-34151 (UPENN); Pitilla, Manguera , 470 m, #f, 4.III.2010, Moraga leg., 10-SRNP-71089 (UPENN); Pitilla, Medrano , 380 m, 1 #m & 1 #f, 8.IX.2012, Calero leg., 12-SRNP-72116, 12-SRNP-72117 (UPENN); Pitilla, Pasmompa , 440 m, 1 #m, 16.II.2012, Rios leg., 12-SRNP-30532 (UPENN); Pitilla, Sendero Evangelista , 660 m, 1 #m, 13.VII.2002, Moraga leg., 02-SRNP-28460 (UPENN); 1 #m, 28.X.2004, Moraga leg., 04-SRNP-55932 (UPENN); Pitilla, Sendero Laguna , 680 m, 1 #m, 11.VIII.2007, Rios leg., 07-SRNP-32980 (UPENN); Pitilla, Sendero Nacho , 710 m, 1 #m, 14.X.2003, Rios leg., 03-SRNP-21390 (UPENN); Pitilla, Sendero Mismo , 680 m, 1 #f, 18.VIII.2004, Rios leg., DZ 30.110 (DZUP); Heredia, Sarapiquí, Agrícola Sofia , 0–100 m, 1 #m, 6.VI.2009, Brenes & Paniagua leg., DZ 30.207 (DZUP); 2 #f, 5.VI.2009, Brenes & Paniagua leg., INB0004273125, INB0004273119 (INBIO); Sarapiquí, Estación Biológica La Tirimbina , 100–200 m, 1 #m, 5.IV.2010, Miranda & Rojas leg., INB0004278034 (INBIO); Sarapiquí, Starke , 0–100 m, 1 #f, 2.VI.2009, Calderón & Cruz leg., INB0004273143 (INBIO); 1 #m & 1 #f, 5.VI.2009, Calderón & Cruz leg., INB0004269382, INB0004273198 (INBIO); 1 #f, 9.V.2009, Calderón & Cruz leg., INB0004273139 (INBIO); ECUADOR, Esmeraldas, Río Chuchuví , 1 #m, 2010, Aldaz leg, R67 (FLMNH); San Francisco ridge, 1 #m, 2010, Aldaz leg., R11 (FLMNH); San Lorenzo-Lita rd. Tundaloma Lodge 100m, 100 m, 1 #m, 17–19.VII.2011, Willmott & Hall leg., LEP-09963 (FLMNH); Tundaloma Lodge, 100 m, 1 #m, 17–19.VII.2011, Willmott & Hall leg., LEP-17613 (FLMNH); 2, #m, 22.VII.2014, Willmott leg., LEP-14943, LEP-14947 (FLMNH); Pichincha, km 20 Pacto-Guayabillas rd., 900 m, 1#m & 1 #f, 7–8.VIII.2011, Willmott & Hall leg., LEP-17612, LEP-17614 (FLMNH); km 21 Pacto-Guayabillas rd., Sardinas, 850 m, 1 #m, 7–8.VIII.2011, Willmott & Hall leg., LEP-09961 (FLMNH).

Diagnosis

Zaretis mirandahenrichae sp. nov., as with most species of the genus, is somewhat variable and similar to its congeners. The species is sympatric with Z. ellops , Z. pythagoras , Z. delassisei , Z. elianahenrichae sp. nov. and Zaretis crawfordhilli sp. nov. Males ( Fig. 13E, F) can be distinguished from Z. crawfordhilli sp. nov. by the fully developed emargination of the inner margin at the FW tornus; from Z. phytagoras by the FW apex, which is considerably less falcate, and the outer margin, which is not crenulated; from Z. pythagoras and Z. delassisei sp. nov. by the absence of a purplish sheen on the FWD when viewed obliquely; from Z. ellops and Z. elianahenrichae sp. nov. by the FW, which is similarly falcate but less pointed; by the generally darker and homogeneous orange coloration, with post-median and submarginal bands absent or weakly developed on the upperside of both wings; FW submarginal area near the apex and marginal area along the outer margin coalesced, wider and dark brown. Females ( Fig. 13G, H) can be distinguished from females of Z. crawfordhilli sp. nov. by the upperside of the wings basal and post-median areas along the outer margin, orange to dark orange and yellow to yellowish orange, repectively, and two-tone wing underside pattern not as evident; from Z. pythagoras by the FW apex, considerably less falcate; from Z. ellops and Z. elianahenrichae sp. nov. by the wing upper and underside basal and post-median areas, and marginal and submarginal areas near the apex notably darker than other areas, with a two-tone pattern of coloration, although subtler than in Z. crawfordhilli sp. nov., variably darker in those areas.

Description

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

Thorax: Dorsally orange to dark orange with scattered brownish and greenish scaling; ventrally orange to dark 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 orange to orange.

Wing size and shape: Forewing length, medium. Forewing costal margin convex; apex pointed and falcate; outer margin sinuous, smooth to slightly 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; 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 moderatedly emarginated near the tornus.

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

Male genitalia ( Fig. 15K): 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 ventral callus; gnathos laterally curved, dorsally thin, 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, with two projections, one smaller, between the costa and the harpe, and another at the end of the harpe; sacculus triangular, ampulla developed and rounded; aedeagus as long as 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. 17G, H): Tergum VIII triangular, ventrally attached to the sides of the lamella postvaginalis and dorsally to the lamella antevaginalis by a strong dorsally projected loop, this with a small anterior projection; papilla analis rounded and with short setae, projecting the apophysis posterioris; lamella antevaginalis assymetrical, connected to the sides of the lamella postvaginalis by lightly sclerotized projections, left side much wider and larger than the right; lamella postvaginalis longer than wide, anterior area tapering and with a small membranous area, posterior edge slightly indented; seminal duct close to the base of the ductus bursae; posterior half of the ductus bursae bulbous, with corpus bursae laterally compressed, about the same length of the ductus bursae, bearing two parallel signa, which are thin and long, formed by minute sclerotized bumps.

Discussion

The description of Z. mirandahenrichae sp. nov. is based on 42 barcoded specimens, 19 males and 14 females, from Costa Rica (Alajuela, Guanacaste and Heredia), deposited at the DZUP, INBIO and UPENN, and nine barcoded specimens, eight males and one female, from Ecuador deposited at the FLMNH. Only barcoded specimens are designated as paratypes, although other specimens were examined at the INBIO and USNM. This species is similar to and frequently misidentified as Z. isidora from the Amazon basin, Guianas and Atlantic forests owing to the somewhat similar wing shape and coloration of the wing upper and undersides. This species, according to the data presented by Janzen & Hallwachs (2017), is not as common as Z. ellops and Z. crawfordhilli sp. nov. and, similarly to Z. crawfordhilli sp. nov. and Z. elianahenrichae sp. nov., it occurs on the Atlantic slopes of Costa Rica. This species was reared multiple times by Janzen & Hallwachs (2017), on C. arborea , C. corymbosa and C. sylvestris . Most immature stages used C. arborea (~82% of the records), which is likewise the preferred host plant of Z. elianahenrichae sp. nov. and Z. crawfordhilli sp. nov, but it is never used by Z. ellops in Costa Rica.

Distribution

Probably widespread in Central America, except the Antilles, and Trans-Andean South America, in northwestern and western Colombia, Costa Rica, western Ecuador, Mexico and Panamá; occurring in forest habitats from low to mid-elevations ( Fig. 21). It may occur in the remaining countries of Central America, except the Antilles, western Peru and northwestern Venezuela.

Etymology

Zaretis mirandahenrichae sp. nov. is named for Miranda Henrich in recognition of her mother Charlotte Hill’s generous support for 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.