Zaretis crawfordhilli, 2019

ZARETIS CRAWFORDHILLI DIAS SP. NOV.

( FIGS 13A–D, 15J, 17E, F, 18A, B, 18H, 21; SUPPORTING INFORMATION, APPENDIX S6)

[ urn:lsid:zoobank.org:act:A68CBEB3-C98B-4590-BB4D-8F7B67E1102E ]

Type material

Male holotype with the following labels: / HOLOTYPUS / HOLOTYPE Zaretis crawfordhilli Dias det. 2016 /Voucher: D.H. Janzen & Winnie Hallwachs caterpillar ( Lepidoptera ) database, Area de Conservación Guanacaste, Costa Rica [Alajuela, Área de Conservación Guanacaste, Sector Rincón Rain Forest, Vado Rio Francia, 400 m, (10.90093, −85.28915) [10°54′3″N, 85°17′21″W], 4-VII-2001, Pérez leg. ex larvae in Casearia arborea ], http://janzen.sas.upenn. edu 01-SRNP-5303 /LEGS AWAY FOR DNA / DZ 30.200 / (DZUP). Allotype female with the following labels: / ALLOTYPUS / ALLOTYPE Zaretis crawfordhilli Dias det. 2016 /Voucher: D.H. Janzen & Winnie Hallwachs caterpillar ( Lepidoptera ) database, Area de Conservación Guanacaste, Costa Rica [Alajuela, Área de Conservación Guanacaste, Sector Rincón Rain Forest, Sendero Rincón , 430 m, (10.8962, −85.27769 [10°53′46″N, 85°16′40″W]), 12-X-2002, Pérez leg. ex larvae in Casearia arborea ], http://janzen.sas.upenn. edu 02-SRNP-21246 /LEGS AWAY FOR DNA /DZ 30.204 / (DZUP).

Paratypes: COSTARICA, Alajuela, Área de Conservación Guanacaste, Rincón Rain Forest, Camino Rio Francia, 410 m, 1 m # & 1 #f, 19.VIII.2005, Pérez leg., 05-SRNP-42266, 05-SRNP-42267 (UPENN); 1 #m, 19.VIII.2005, Carmona leg., DZ 30.201 (DZUP); Rincón Rain Forest, Finca Hugo , 540 m, 1 #m, 21.VI.2011, Pérez leg., 11-SRNP-42929 (UPENN); Rincón Rain Forest , Quebrada Guarumo , 400 m, 1 #f, 20.VIII.2001, Pérez leg., 01-SRNP-5693 (UPENN); 1 #f, 8.X.2012, Calderon leg.,12-SRNP-85888 (UPENN); Rain Forest, Río Francia Arriba, 400 m, 1 #m, M 8.XI.2012, Pérez leg., 12-SRNP-86533 (UPENN); Rincón Rain Forest , San Lucas , 320 m, 1 #m, 30.VI.2011, Córdoba leg., 11-SRNP-43136 (UPENN); 1 #m, 26.VI.2012, Córdoba leg., 12-SRNP-43156 (UPENN); Rincón Rain Forest, Selva , 410 m, 29.IV.2010, Briceño leg., 10-SRNP-69622 (UPENN); 1 #m, 8.V.2010, Briceño leg., 10-SRNP-69643 (UPENN); Rincón Rain Forest, Sendero Juntas , 400 m, 1 #m, 8.XI.2006, Araya leg., 06-SRNP-44414 (UPENN); Rincón Rain Forest, Sendero Llano , 400 m, 1 #f, 10.IX.2007, Carmona leg., 07-SRNP-42457 (UPENN); Rincón Rain Forest, Sendero Parcelas , 375 m, 1 #m, 07.XII.2005, Carmona leg., 05-SRNP-41764 (UPENN); 375 m, 1 #f, 12.VII.2005, Carmona leg., 05-SRNP-41763 (UPENN); Rincón Rain Forest, Sendero Rincón , 430 m, 2 #m, 8.IV.2003, Pérez leg., 03-SRNP-10614, 03-SRNP-10615 (UPENN); 1 #m, 16.I.2005, Pérez leg., 05-SRNP-40138 (UPENN); 1 #m, 16.I.2005, Pérez leg., 05-SRNP-40139 (UPENN); 1 #m, 9.V.2012, Córdoba leg., 12-SRNP-41969 (UPENN); 1 #f, 3.X.2005, Pérez leg., 05-SRNP-42926 (UPENN); 1 #f, 12.I.2007, Pérez leg., 07-SRNP-40155 (UPENN); 1 #f, 5.IX.2011, Umaña leg., 11-SRNP-44034 (UPENN); Rincón Rain Forest, Sendero Rincón , 410 m, 1 #m, 29.IV.2010, Briceño leg., 10-SRNP-69621 (UPENN); San Cristóbal, Puente Palma , 460 m, 1 #f, 21.IX.2006, Cano leg., DZ 30.205 (DZUP); San Cristóbal, Sendero Huerta , 527 m, 1 #m, 3.X.2006, Cano leg., 06-SRNP-8208 (UPENN); 1 #m & 1 #f, 17.IV.2007, Sihezar leg., 07-SRNP-1798, 07-SRNP-1797 (UPENN); 1 f#, 4.IV.2012, Cano leg., 12-SRNP-1391 (UPENN); Guanacaste, Área de Conservación Guanacaste, Pitilla, Pasmompa , 440 m, 1 #m, 15.X.2003, Rios leg., 03-SRNP-21420 (UPENN); 1 #f, 9.X.2010, Rios leg., 10-SRNP-32187 (UPENN); Pitilla, Sendero Evangelista , 660 m, 1 #f, 29.IX.2011, Rios leg., 11-SRNP-32971 (UPENN); Pitilla, Sendero Mismo , 680 m, 1 #m, 22.IX.2003, Rios leg., 03-SRNP-21183 (UPENN); Heredia, Sarapiquí, Agrícola Sofia , 0–100 m, 2 #m, 3.VII.2010, Brenes & Paniagua leg., DZ 30.106 (DZUP), INB00042773401 (INBIO); 1 #m, 5.IV.2009, Brenes & Paniagua leg., INB0004269668 (INBIO); Sarapiquí, Estacíon Biológica La Tirimbina , 169 m, 1 #m, 6.VII.2010, Miranda & Rojas leg., DZ 30.202 (DZUP); 230 m, 1 #m, 6.VII.2010, Miranda & Rojas leg., INB0004277310 (INBIO); Sarapiquí, Starke , 0–100 m, 2 #m, 3.VIII.2009, Calderón & Cruz leg., INB0004269384, INB0004269662 (INBIO); 1 #m, 29.VI.2010 Calderón & Cruz leg., INB0004277350 (INBIO); 1 #f, 30.VI.2010, Calderón & Cruz leg., DZ 30.208 (DZUP); Limón, Veragua, Rainforest Restaurant, 400–440 m, 1 #f, 14.XI.2008, Villalobos leg., DZ 30.107 (DZUP); MEXICO, Chiapas, Ocosingo, Chajul , 200 m, 1 #f, 20.VIII.1996, Ibarra leg., AIV193 (UNAM); PANAMA, Panamá Oeste, Barro Colourado Island, 150 m, 1 #f, 15.V.2012, Bobadilla leg., YB-BCI56036 (STRI); 1 #f, 25.VI.2012, Rivera leg. YB-BCI68924 (STRI).

Diagnosis

Zaretis crawfordhilli 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 Z. mirandahenrichae sp. nov. Males ( Fig. 13A, B) can be distinguished from all above cited species by the less well-developed emargination of the inner margin at the FW tornus, which is different in comparison to that of Z. itys . Females ( Fig. 13C, D) can be distinguished from the above species by the shape of the FW, the outer margin always being smooth and rounded, and slightly falcate at the apex; and by the colour and pattern. Forewing dorsum basal area dark orange to reddish brown, post-median area beige to pale yellow, submarginal area beige to pale yellow along the outer margin and dark brown, coalesced with the marginal area near the apex; marginal area dark brown; post-median band usually reddish brown, between post-median and submarginal areas; forewing ventrum (FWV) similar to the upperside, but basal, submarginal areas near the apex almost solid brown to dark brown, post-median and submarginal areas along the outer margin pale beige to beige; FWV basal and discal bands usually indistinct, post-median band as in the upperside; hindwing dorsum (HWD) basal area reddish brown, and median area, submarginal and marginal areas reddish brown suffused with dark brown; hindwing ventrum similar to upperside, with basal, submarginal and marginal areas almost solid brown to dark brown, median area suffused with pale beige to beige from the inner margin, post-median area brown to light brown, usually lighter than basal, submarginal and marginal areas; females of Z. delassisei are unknown.

Description

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

Thorax: Dorsally orange to dark orange with scattered brownish and greenish scaling; ventrally orange to reddish brown, 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, orange to dark 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 underdeveloped, about one-quarter of 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 rounded and more falcate at the apex; emargination before tornus developed; hindwing proportionately larger than male; emargination at Sc–Rs developed; apex projected at Rs; outer margin rounded; inner margin 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 areas along the outer margin usually of the same colour; 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; 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 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 pale beige to pale yellow; 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, orange to dark orange. Hindwing basal and marginal areas usually darker than median, post-median and submarginal areas, orange to dark orange; median, post-median and submarginal areas pale yellow to yellowish orange; median band usually developed from the costal margin to the discal cell; the discal spot absent; post-median and submarginal bands faint, more noticeable near the costal margin; 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 brown to reddish 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; 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 areas darker than other areas and similar in colour, reddish brown to dark brown; post-median and submarginal areas along the outer margin beige to pale yellow; all bands noticeable, orange to brown. Hindwing basal and marginal areas darker, reddish brown to dark brown; median area beige to pale yellow; post-median and submarginal areas lighter than basal and marginal areas, but darker than medial area; 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 brown to reddish brown. Female dorsally orange to dark orange; ventrally reddish brown to dark brown.

Male genitalia ( Fig. 15J): Tegumen trapezoidal in lateral view, dorsally wider and posteriorly bulged, 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, slightly shorter than the tegumen, slightly curved, with a well-developed median dorsal ridge, distally hooked and with a ventral callus; gnathos laterally slightly curved, dorsally slightly larger than ventrally, 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, two projections, one smaller projection 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. 17 E, F): Tergum VIII triangular, ventrally attached to the sides of the lamella postvaginalis and dorsally to the lamella antevaginalis by a strong 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 wide sclerotized projections, with left side larger than the right; lamella postvaginalis wider than long, anterior area with a small membranous area, and posterior rounded to slightly bilobed; seminal duct close to the base of the ductus bursae; corpus bursae rounded, about half the length of ductus bursae, bearing two parallel signa, which are thin and long, formed by minute sclerotized bumps.

Discussion

The description of Z. crawfordhilli sp. nov. is based on 48 barcoded specimens, 29 male and 16 female specimens from Costa Rica (Alajuela, Guanacaste and Heredia) deposited at the DZUP, INBIO and UPENN, one female from México (Chiapas) deposited at the UNAM, and two females from Panamá, deposited at the STRI. Only barcoded specimens are designated as paratypes, although several other specimens were examined at the INBIO and USNM. This species is similar to and has been frequently identified as the Amazonian Z. itys , which it resembles in the weakly developed emargination of the FW at the tornus, and the somewhat similar wing shape and coloration of the female. Nevertheless, Z. crawfordhilli sp. nov. is not recovered as closely related to Z. itys in any of our analyses. Additionally, the shape of the underdeveloped emargination of the inner margin at the FW tornus is clearly different for these two species, being shorter but similarly deep in Z. crawfordhilli sp. nov., whereas it is shorter and considerably shallower in Z. itys . The female wings are broader, and the coloration of the upperside is usually lighter than in Z. itys . This species, according to the data presented by DeVries (1987) and Janzen & Hallwachs (2017), is rather common, second only to Z. ellops in Costa Rica, but occurring mostly on the Atlantic slopes, whereas Z. ellops occurs mostly on the Pacific slopes. DeVries (1987) found this species to be most common in baited traps during the dry season.

DeVries (1987) described the immature stages of Z. crawfordhilli sp. nov. (identified as Z. itys ; Fig. 18A, B), indicating several differences between that species and Z. ellops : ‘thoracic hump [A2 enlargement] and the rhomboid shapes on the dorsum highlighted in green; last segment [A9 + 10] is splayed into a short fan; head horns [head capsule scoli] curve inward toward each other; and the entire head is dotted in green granules’. It is not certain whether he was using the latter name to refer to Z. elianahenrichae sp. nov. or Z. mirandahenrichae sp. nov., because he was unaware of the existence of further species in Costa Rica. However, it is safe to assume that Z. crawfordhilli sp. nov. is the species under consideration, given his species description and figures ( DeVries, 1987; pl. 14, figs 12–14), and by the fact that this species is the second most common species of Zaretis in Costa Rica, whereas Z. elianahenrichae sp. nov. and Z. mirandahenrichae sp. nov. are much scarcer. Several authors recorded host plants for what is assumed to be Z. crawfordhilli sp. nov. Most citations are on Salicaceae : on undetermined species of the family ( Marquis, 1991; Dyer, 1995), undetermined ( DeVries, 1987; Mallet apud Beccaloni et al., 2008) and determined species of Casearia : C. arborea ( DeVries, 1986; Gentry & Dyer apud Beccaloni et al., 2008), C. arguta ( DeVries, 1986) , C. sylvestris (DeVries apud Beccaloni et al., 2008), and species of Laetia and Ryanea ( DeVries, 1987), on Laetia procera ( DeVries, 1986; Gentry & Dyer apud Beccaloni et al., 2008) and Ryanea speciosa ( DeVries, 1986) . The record for Piper trigonun ( Piperaceae ) provided by DeVries (1986) is doubtful and needs confirmation. This species was frequently reared by Janzen & Hallwachs (2017) on C. arborea , C. corymbosa and C. sylvestris . Most immature stages used C. arborea (~78% of the records) and C. sylvestris (~10% of the records), which are likewise used by Z. elianahenrichae sp. nov. and Z. mirandahenrichae sp. nov., but never or rarely used by Z. ellops .

Distribution

The species occurs in forest habitats from low to mid-elevations and is probably widespread in Central America and Trans-Andean South America, in Belize, northwestern and western Colombia, Costa Rica, western Ecuador, Guatemala, Honduras, southern Mexico, Nicaragua and Panamá; it occurs in forest habitats from low to mid-elevations ( Fig. 21). It may occur in the remaining countries of Central America, except the Antilles, western Ecuador and northwestern Peru and northwestern Venezuela.

Etymology

Zaretis crawfordhilli sp. nov. is named in honour of Crawford Hill of Philadelphia, USA, in recognition of his decades of teaching biodiversity to high school students and his serious support and encouragement for exposing those same students to new locations and especially, to Área de Conservación Guanacaste, where Z. crawfordhilli sp. nov. is prominent.

Examined material

See Supporting Information, Appendix S1.