Optimandes eugenia transversa ( Weymer, 1911 ) Willmott & Marín & Nakahara & Pomerantz & Lamas & Huertas & Espeland & Xiao & Hall & Robinson Willmott & Freitas, 2019

Willmott, Keith R., Marín, Mario A., Nakahara, Shinichi, Pomerantz, Tatiana, Lamas, Gerardo, Huertas, Blanca, Espeland, Marianne, Xiao, Lei, Hall, Jason P. W., Robinson Willmott, James I. & Freitas, André V. L., 2019, A revision of the new Andean butterfly genus Optimandes Marín, Nakahara & Willmott, n. gen., with the description of a new species (Nymphalidae: Satyrinae: Euptychiina), Tropical Lepidoptera Research 29 (1), pp. 29-44 : 36-40

Optimandes eugenia transversa ( Weymer, 1911) , n. comb.

Figs. 1, 2C-F, 3, 4A-E, 6, 7, 8

Euptychia transversa Weymer (1911: 197 , pl. 47a) Lectotype ♂ [here designated]: “ transversa Weym. //Original?//Para-Typus Euptychia transversa Weym. //Marcap.// Peru Marcapata Staatsamml. München // Präparat Nr. SA20 Zoolog. Staatssammlung München” ( ZSM, examined). Paralectotype ♂: “Lectotype ♂ Euptychia transversa Weymer designated by: Lee D.Miller 1989//genitalia vial M-9138 ♂ Lee D. Miller// Marcapata Cuzco ” ( MNHU, examined) .

Euptychoides transversa: Forster (1964: 97 , fig. 85)

Euptychia phineus: D’Abrera (1988: 777 , ♂ V), misidentification

Euptychoides eugenia transversa: Lamas (2004: 219)

Euptychoides phineus: Piñas (2004: 22 , fig. 129, 130), misidentification

Euptychoides eugenia [ transversa ]: Murray & Prowell (2005); Espeland et al. (2019: 122)

Diagnosis and identification: Diagnostic characters of this taxon are discussed under the nominate subspecies. This taxon is superficially similar to a number of sympatric cloud forest euptychiines, particularly Euptychoides fida and Hermeuptychia species, but may be distinguished by the very reduced VHW ocelli in the middle of the wing and the undulating VHW dark discal line, in addition to other distinguishing characteristics of the genus.

Taxonomy: Weymer (1911) described this taxon based on an unspecified number of specimens from Marcapata in Peru, stating that it also occurred at Río ‘Vitaca’ (=Río Bitaco) in Valle del Cauca, Colombia. The figure of the ventral surface corresponds closely to a syntype specimen in the ZSM that is here designated as lectotype to fix the identity of the name ( Fig. 2 E); a male paralectotype from the same locality is in the MNHU. We follow Lamas (2004) in treating this taxon as a subspecies of O. eugenia since the two share very similar wing patterns, except for white scaling on the VHW, as discussed above under Diagnosis and identification. There is some variation throughout the Andes in the extent of the white scaling on the VHW and expression of the dark lines on the VHW. Specimens from western Ecuador ( Fig. 2 C, D) have pure white scaling lacking scattered brown scales, narrow discal and postdiscal lines and a broad outer submarginal line that remains broad into M 3 -M 2. East Andean specimens ( Fig. 2 E) have broader, more indistinct discal and postdiscal lines, narrower submarginal lines and more diffuse white scaling, which is reduced even further in Bolivian specimens ( Fig. 2 F) to the extent that they resemble the nominate subspecies ( D’Abrera, 1988: 777, figured a Bolivian male specimen under the name Euptychia phineus ). Deciding whether any of this variation merits taxonomic recognition requires more comprehensive material from more intervening regions; nonetheless, based on specimens we have examined, the variation appears to be insufficiently stable or marked. DNA barcode sequences from east and west Andean specimens show no significant divergence ( Fig. 1).

Distribution and natural history: In western Ecuador, this subspecies is known to date from Carchi to Pichincha. There are single records from Cañar (collected by R. de Lafebre) and Bolívar (collected by M. de Mathan) that require confirmation, since specimens of numerous other butterfly taxa with the same labels in the FLMNH and NHMUK, respectively, are evidently mislabeled. The Bolívar specimen, in particular, has a ventral wing pattern similar to east Ecuadorian specimens. The subspecies is also known in the eastern Andes from Ecuador (Napo) south to Bolivia (Cochabamba). The species presumably also occurs throughout the montane areas of Colombia and in the Venezuelan Cordillera de Mérida, although we have seen no records from these areas aside from Weymer’s (1911) report of the taxon from Río Bitaco (Colombia, Valle del Cauca). Despite being rather rare in historical collections (e.g., a total of 14 specimens in the three largest European collections, NHMUK, MNHU and ZSM), we have found it to be merely uncommon, or even locally common, in the field in Ecuador and Peru, where it occurs in cloud forest from 1000-2200 m. The rarity of specimens in collections may be due to the tendency of the species to fly high in the canopy. In Ecuador, males were frequently attracted to traps 8-10 m above the ground baited with rotting fish and were observed flying and perching 15 m above a stream in a small light gap from 13:00-14:00. Females were not attracted to baits and were therefore much more rarely encountered in the field, with one individual observed resting at 0.5 m on a stand of bamboo in a forest light gap at 12:30.

Immature stages ( Figs. 7, 8): The following description of the immature stages of E. eugenia transversa is based on two lots collected in Zamora-Chinchipe, eastern Ecuador, in July-August 2018. These lots include: 2018-JK-01, comprising four 1 st instars from the ridge E of San Roque, of which one survived until 3 rd instar and was barcoded to confirm identification; 2018- JK-06, comprising eight 3 rd instars from km 20 Los Encuentros-Zarza (3°50'14'' S, 78°35'31'' W), with four surviving until 4 th instar, of which two were barcoded to confirm identification, and with the remaining four producing 2 adult males ( Fig. 8 C), 1 female and 1 of undetermined sex (failed to completely eclose). JK-01 was collected feeding on an unidentified species of Chusquea ( Poaceae , Fig. 8 D-F, identified by Lynn Clark, pers. comm.) c. 1 m above the ground growing along the edge of forest along a ridge top road, and JK-06 was collected c. 2 m above the ground on what appears to have been the same plant species ( Fig. 8 G) growing at the edge of forest along a river.

1 st instar ( Fig. 7 A-E): The 1 st instar fed from the edge of the leaf near the leaf base, making a characteristic elongate hole along the leaf edge ( Fig. 7 A, B). Body green, slightly darker dorsally, with an indistinct dorsolateral pale line, very short caudal filaments, legs and prolegs pale green, body covered with short, club-tipped setae, and with a black head capsule bearing setae and a pair of short, rounded dorsal scoli on vertex. Molted to 2 nd instar 3 days after collection (n=4). Maximum length c. 5.3-5.9 mm, head capsule width c. 0.7- 0.9 mm.

2 nd instar ( Fig. 7 F-I): Body pale green on molting, becoming darker reddish brown in anterior half in later part of instar, with pale yellowish green dorsolateral line and three similar but thinner lateral lines, pair of stubby caudal filaments, legs and prolegs pale green, and with a black head capsule bearing a pair of longer (c. half head height), tapering, blunt scoli on vertex. Molted to 3 rd instar after 5-9 days (exact duration not recorded) (n=1). Maximum length c. 9.9-10.1 mm, head capsule width c. 1.0-1.2 mm.

3 rd instar ( Fig. 7 J-L): Body pale bluish green, with pale yellowish green dorsolateral line and three similar but thinner lateral lines, spiracles yellowish brown, pair of stubby caudal filaments, legs and prolegs pale green, and with a green head capsule bearing a pair of shorter (c. one third head height), tapering, pointed orange scoli on vertex and black stemmata. Molted to 4 th instar 5-9 days after collection (exact duration not recorded) (n=6). Maximum length c. 15.9- 16.5 mm, head capsule width c. 1.5-1.7 mm.

4 th (last) instar ( Fig. 7 M-O): Body bright green, with pale green dorsolateral line and three similar but thinner lateral lines, spiracles yellowish brown, pair of longer caudal filaments, legs and prolegs pale green, and with a green head capsule bearing a pair of short (c. one fifth head height), conical orange scoli on vertex (relatively shorter than in previous instar) and black stemmata. Duration at least 6 days (exact duration not recorded) (n=5). Maximum length c. 28.0 mm, head capsule width c. 2.4-2.5 mm.

Pupa ( Fig. 8): Short and smooth, with rounded ocular caps, bright green except for few scattered small black and brown spots on abdomen, continuing as single dorsolateral black spots onto thorax, edge of wing case delineated with thin white line bordered dorsally by thin brown line, cremaster short and broad. Adults eclosed after 11-12 days (n=4).

Overall, the immature stages are morphologically simple and similar to those of several other Euptychiina with green immatures, such as Pareuptychia , Cepheuptychia Forster, 1964 and Taydebis Freitas, 2003 ( Freitas et al., 2016a), all members of the Pareuptychia clade. In particular, the green last instar with orange head scoli and the short green pupa are remarkably similar to those of Pareuptychia ( Freitas et al., 2016a) .Although the immature stages of Euptychiina are known for relatively few taxa, an entirely green last instar is apparently uncommon, with predominantly brown larvae being far more widespread in the subtribe ( Freitas et al., 2016 a, b, 2018b and references therein). Although camouflage in Euptychiina larvae has been suggested as a major strategy for escaping from predators ( Freitas, 2017; Freitas et al., in press), it would be interesting to investigate why green larvae (and pupae) are especially prevalent in the Pareuptychia clade.

Specimens examined (38 ♂, 20 ♀): Ecuador: Carchi: Reserva Las Golondrinas, N of La Carolina , Nariz del Diablo , [0°49'39''N, 78°7'29''W], 1900 m, (Willmott, K. R.), 28 Nov 1996, ( FLMNH) ; Imbabura: km 26 Chontal Bajo-Chontal Alto , Chontal Alto , [0°17'48''N, 78°42'3''W], 1550-1650 m, (Willmott, K. R., Hall, J. P.W.), 10 Aug 2011, 1 ♀, ( INABIO) ; Pichincha: [Alluriquín], Río Toachi , [0°19'6''S, 78°57'13''W], 800 m, Sep, 1 ♀ [ FLMNH-MGCL-193438 ], ( FLMNH) ; [Salto de] Napac , [0°20'6''S, 78°53'24''W], 1000 m, (Nicolay, S. S.), 23 Sep 1975, 1 ♀ [ FLMNH-MGCL-193439 ], ( FLMNH) ; 12 km SW Las Tolas , [0°3'3''N, 78°50'18''W], 1200 m, (Willmott, K. R., Hall, J. P. W.), 1, 3 Aug 2011, 1 ♂ [ FLMNH-MGCL-157244 ], 1 ♂ [ FLMNH-MGCL-157245 ], 1 ♀ [ FLMNH-MGCL-157246 ], ( FLMNH), 1 ♀, ( INABIO) ; km 85 Quito-Sto. Domingo old rd., 1372 m, (Hyatt, J.), 2 Jul 1980, 1 ♀ [FLMNH-MGCL-193437], ( FLMNH) ; NW Quito, Alamo valley , 1800 m, (Adams, M. J. & J.), 6 Aug 1986, 1 ♀ [ BMNH(E) 1420248 ], ( NHMUK) ; Quito-Sto. Domingo old rd., Hacienda Santa Isabel , [0°18'48''S, 78°56'W], 1200 m, (Willmott, K. R.), 2 Sep 1996, 1 ♂, ( FLMNH) ; Río Alambi , Reserva Maquipucuna , [0°5'42''N, 78°38'W], 1250- 1350 m, (Willmott,K. R., Hall, J. P.W.), 16,17Aug 1993, ( FLMNH) ; Tandayapa Bird Lodge , [0°0'7''N, 78°40'41''W], 1700 m, (Willmott, K. R., Hall, J. P. W.), 2-5 Aug 2011, 1 ♂ [ FLMNH-MGCL-157247 ], 1 ♂ [ FLMNH-MGCL-157248 ], ( FLMNH) ; Bolívar: Balzapamba - (mislabeled?), [1°47'S, 79°10'W], (Mathan, M. de), Sep 1893 - Feb 1894, 1 ♂ [ NHMUK012824432 ], ( NHMUK); Cañar: Río Angas , nr. Huigra , Angas - (mislabeled?), [2°18'S, 79°3'W], 1000 m, (Lafebre, R. de), Jul 1974, 1 ♂ [ FLMNH-MGCL-193440 ], ( FLMNH) ; Napo: nr. Cosanga, Estación Científica Yanayacu , [0°35'24''S, 77°53'W], 2000 m, (Willmott, K. R.), 24 Nov 2006, 1 ♂ [ FLMNH-MGCL-112612 ], ( FLMNH) ; Pastaza: km 11 Mera-Río Anzu rd., [1°25'15''S, 78°3'8''W], 1200 m, (Hall, J. P. W., Willmott, K.R., J.C.R., J.I. R), 31 Jul 2015, 1 ♀ [FLMNH-MGCL-209655], ( FLMNH) ; Tungurahua: Reserva Cerro Candelaria, Fundación Ecominga , [1°25'30''S, 78°17'59''W], 2200 m, (Radford, J.), 19 Aug 2008 [ JR-08-350 ], ( JARA), 20 Aug 2008 [ JR-08-374 ], ( JARA), 21 Aug 2008 [ JR-08-128 ], ( JARA) ; Morona-Santiago: Cóndor Mirador, [3°37'42''S, 78°23'41''W], 1974 m, (Radford, J.), 23 Aug 2010, 1 ♀ [ CON91 ], ( FLMNH) (CULEPEX Expedition, 2010) ; Guarumales/Hidropaute, [2°34'9''S, 78°30'49''W], 1900 m, (Willmott, K. R.), 7 Nov 2010, 1 ♂ [ FLMNH-MGCL-146103 ], ( FLMNH) ; km 14 Limón-Gualaceo rd., [3°0'36''S, 78°30'W], 1900 m, (Willmott, K. R.), 11 Nov 1996, 1 ♂, ( FLMNH) ; Zamora-Chinchipe: km 10 Los Encuentros-El Panguí rd., ridge E San Roque , [3°42'11''S, 78°35'36''W], 1050 m, (Willmott, K. R., Hall, J. P. W.), 4 Aug 2009, 1 ♀ [ FLMNH-MGCL-145733 ], 18 Jul 2018, 1 individual [larva lot 2018-JK-01 , barcoded], ( FLMNH) ; km 20 Los Encuentros-Zarza rd., [3°50'14''S, 78°35'31''W], 1450 m, (Hall, J. P. W., Willmott, K. R., J. I. R), 22 Jul 2018, 2 ♂, 1 ♀ [larvae, lot 2018-JK-06 , 1 ♂ eclosed 18 Aug 2018, 1 ♂, 1 ♀ eclosed 22 Aug 2018], 2 individuals [larvae lot 2018-JK-06 , barcoded], ( FLMNH) ; km 24 Loja-Zamora rd., San Francisco, casa de Arcoiris , [3°59'18''S, 79°5'42''W], 2000-2100 m, (Aldaz, R.), 3 Oct 2006, 1 ♂ [ FLMNH-MGCL-112616 ], ( FLMNH), (Willmott, K. R.), 2 Dec 2006, 1 ♂ [ FLMNH-MGCL-112617 ], ( FLMNH), (Willmott, K. R., Aldaz, R.), 11 Oct 2006, 1 ♂ [ FLMNH-MGCL-112614 ], ( FLMNH), 1 ♂ [ FLMNH-MGCL-112613 ], 1 ♂ [ FLMNH-MGCL-112615 ], ( INABIO) ; km 4.3 San Andrés-Jimbura rd., [4°47'59''S, 79°18'18''W], 2020 m, (Willmott, K. R.), 13 Oct 2010, 1 ♀, ( INABIO) ; Not located: ' Ecuador', 1 ♂ [ BMNH(E) 1420249 ; Joicey Bequest 1934-120], ( NHMUK). Peru: Junín: 1 km S Mina Pichita , [11°5'28''S, 75°24'58''W], 2100 m, (Lamas, G.), 22 Aug 2003, 1 ♂ [ MUSM-LEP-101616 ], ( MUSM), (Peña, C.), 22 Aug 2003, 1 ♂ [ MUSM-LEP-101615 ], ( MUSM) ; 1-3 km SE Mina Pichita, [11°5'28''S, 75°24'58''W], 2100 m, (Lamas, G.), 26 Aug 1988, 1 ♂ [ MUSM-LEP-101614 ], ( MUSM) ; Quebrada Siete Jeringas, [11°12'S, 75°24'W], 1700 m, (Peña, C.), 15 Nov 2003, 1 ♂ [ MUSM-LEP-101617 ], 1 ♀ [ MUSM-LEP-101621 ], ( MUSM) ; Cuzco: Cosñipata rd., San Pedro Lodge , 1375 m, (Kinyon, S.), 22 Oct 2013, 1 ♂, ( USNM), 23 Sep 2011, 1 ♂, ( USNM) ; Cosñipata , Quebrada Quitacalzón, [13°1'S, 71°30'W], 1100 m, (Lamas, G.), 23 Oct 2010, 1 ♂ [ MUSM-LEP-101618 ], ( MUSM) ; Marcapata , [13°30'34''S, 70°53'57''W], 1800 m, 1 ♂ [' LECTOTYPE ♂ Euptychia transversa Weymer designated by: Lee D. Miller 1989//genitalia vial M-9138 ♂ Lee D. Miller//Marca-pata Cuzco'], ( MNHU), 1 ♂ ['Original?' //'Para-typus Euptychia transversa Weym. '//'marcap'//' Peru Marcapata Staatsamml. München'//' Präparat Nr. SA20 Zoolog.Staatssammlung München'//' transversa Weym. '], ( ZSM) ; San Pedro , [13°3'S, 71°33'W], 1400 m, (Lamas, G.), 10 Nov 2007, 1 ♀ [ MUSM-LEP-101624 ], ( MUSM), 13 Nov 2012, 1 ♀ [ MUSM-LEP-101626 ], ( MUSM), 23 Sep 2011, 1 ♂ [ MUSM-LEP-101619 ], 1 ♀ [ MUSM-LEP-101625 ], ( MUSM), (West, F.& A.), 4-8 Nov 2007, 1 ♀ [ MUSM-LEP-101623 ], ( MUSM) ; San Pedro , [13°3'S, 71°33'W], 1400-1650 m, (Lamas, G.), 7 Nov 2001, 1 ♀ [ MUSM-LEP-101622 ], ( MUSM) ; upper Río Urubamba, Río Vilcanota , 3000 m - (mislabeled), (Garlepp, O.), 1898, 1 ♂, ( MNHU) ; El Mirador , [13º 4'S, 71º33'W], 1720 m, (Kinyon, S.), 28 Oct 2018, 1 J, ( MUSM). Bolivia: La Paz: Río Zongo, [16°3'40''S, 68°1'2''W], 1200 m, (Garlepp), 1896, 1 ♂, ( MNHU) ; Sandillani , [16°12'S, 67°54'W], 2000 m, (Lamas, G.), 4 Dec 2007, 1 ♂ [ MUSM-LEP-101620 ], ( MUSM) ; Cochabamba : Alto Palmar, [17°9'2''S, 65°42'47''W], 1100 m, (Baumann), Nov 1960, 1 ♀, ( ZSM) ; Yungas del Espíritu Santo , [17°6'S, 65°40'W], (Germain, P.), 1888-1889, 1 ♂ [ NHMUK012824433 ], 1 ♂ [ NHMUK012824430 ], 1 ♀ [ BMNH(E) 1420246 ], 1 ♀ [ BMNH(E) 1420247 ], ( NHMUK) ; no specific locality, 1 ♂ [ NHMUK012824431 ], ( NHMUK) .

Other records (sight records and photographs of live individuals; “W & H, SR” indicates sight records by Willmott and Hall): Ecuador: Carchi: Reserva Las Golondrinas, N of La Carolina , Santa Rosa , [0°49'38''N, 78°7'42''W], 1700 m, (Willmott, K. R., Hall, J. P. W.), 4, 5 Sep 1996, (W & H, SR); Imbabura: Selva Alegre-Otavalo rd., Mina Selva Alegre , [0°17'3''N, 78°32'44''W], 2020 m, (Willmott, K. R., Hall, J. P.W.), 11Aug 2011, (W & H, SR); Zamora-Chinchipe: c. 3 km W Guayguayme Alto, ridge above San Luís , [3°55'14''S, 78°54'49''W], 1470 m, (Neild, A.), Nov 2015, 1 ♂, (photograph live specimen) (Neild, A., pers. comm. (10 Nov 2015 by email to KRW with photo)); Destacamento Paquisha Alto, [3°54'28''S, 78°29'5''W], 2100 m, (Radford, J.), 1 Sep 2010, 1 ♂ / ♀ [ PAN56 ], (sight record) (CULEPEX Expedition, 2010); km 24 Loja-Zamora rd., San Francisco, casa de Arcoiris , [3°59'18''S, 79°5'42''W], 2000- 2100 m, (Willmott, K. R.), 28 Nov 2003, (W & H, SR) .