Optimandes Marín, Nakahara & Willmott, 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 : 30-33

Optimandes Marín, Nakahara & Willmott , new genus

Type species: Neonympha eugenia C. Felder & R. Felder, 1867

Diagnosis and identification: Genetic sequence data show that Optimandes n. gen. is a member of the Pareuptychia clade ( Murray & Prowell, 2005), where it is sister to Nhambikuara Freitas, Barbosa & Zacca, 2018 ( Espeland et al., 2019) ( Fig. 1), albeit with only moderate ultrafast bootstrap support (83%). Aside from Nhambikuara cerradensis Freitas, Barbosa & Zacca, 2018, the type of Nhambikuara and related species forming the sister clade to Optimandes all have distinctive, oval submarginal ocelli on the VHW with pale scaling offset from the center of each ocellus. In contrast, both Optimandes species have circular or slightly oval submarginal VHW ocelli with a single, well-marked central white pupil in each, in addition to two well-developed subapical ocelli on the VFW in cells M 2 -M 1 and M 1 -R 5 (the latter reduced in O. e. eugenia and barely visible or absent in Nhambikuara). The combination of the two subapical ocelli in the VFW apex, and single-pupilled VHW ocelli, which fill cells Cu 2 -Cu 1 and M 2 -M 1 on the VHW, distinguish this genus from most other euptychiines. Magneuptychia tiessa (Hewitson, 1869) has these characteristics also, and is also a member of the Pareuptychia clade based on genetic data, but the species is sister to Satyrotaygetis satyrina (Bates, 1865) and both are more closely related to Pareuptychia than they are to Optimandes . The male genitalia of M. tiessa likewise shows no obvious similarities to those of Optimandes .

Description: MALE (Fig. 2,3,4): Forewing length 23-24 mm (n=5). Wings: FW triangular, distal and anal margins straight and almost perpendicular, vein R 2 arising just basally of origin of vein R 3+4+5 ( Fig. 3); HW approximately triangular, distal margin rounded, anal margin slightly indented basal of tornus. Dorsal surface: Ground color gray-brown to reddish brown, slight darker blackish brown marginal line on both wings, and indistinct submarginal line on DHW. Ventral surface: Ground color grayish to reddish brown, a little paler than dorsal surface, with scattered to solid white scaling in posterior basal half of VHW, discal area, and surrounding more anterior ocelli in one taxon. VFW with a discal and a postdiscal line/band, darker brown, latter bordered distally with paler ground scales followed by darker ground scales and two to four postdiscal ocelli in cells Cu 1 -M 3, with those in M 2 -R 5 black spots encircled by clear yellowish brown rings with a single central white pupil in each ocellus, remaining ocelli smaller but similar, or simple yellow rings, or obsolete; two broad, dark brown submarginal lines, more basal line wavy and bordered basally by band of paler ground scaling, more distal line straighter, wing margin lined with black. VHW with dark brown discal and postdiscal lines from costa to anal margin, postdiscal line bordered distally with paler ground scaling; five to six postdiscal ocelli, black with yellow ring and central white pupil in cells 2A-Cu 2 and M 1 -Rs (half width of cell) and Cu2 -Cu 1 and M 2 - M 1 (almost filling cell), ocelli in cells Cu 1 -M 2 similar to others but reduced in size, or more indistinct and lacking central pupil, or obsolete; two broad, dark brown submarginal lines, more basal line wavy and bordered basally by band of paler grayish scaling, wing margin lined with black. Head: eyes brown with dense, long setae; antennae with c. 33 antennomeres, distal c. 10 comprising club (n=4), dark brown with sparse white scales at ventral base of antennomeres in basal part of antennae; labial palpi ( Fig. 3 B) dark brown with long dark brown hair-like scales ventrally, with O. eugenia having cream-colored scales laterally; head covered with dark brown scales and hair-like scales. Thorax: thorax, forelegs ( Fig. 3 C), mid- and hindlegs dark brown (mid- and hindlegs ventrally with sparse cream-colored scales in O. eugenia ), mid- and hindlegs with pair of tibial spurs. Abdomen: dark brown dorsally, paler brown ventrally, eighth tergite unsclerotized except for narrow anterior band and slightly broader posterior patch; eighth sternite appearing as single plate, in contrast to several species currently placed in Euptychoides (e.g., E. pseudosaturnus Forster, 1964 ). Genitalia ( Fig. 4): notable features in comparison with other Euptychiina include aedeagus relatively broad, curving upwards anteriorly and containing two conspicuous elongate patches of teeth-like cornuti; uncus with broad basal half in lateral view with a prominent dorsal ‘keel’; brachia directed upwards with respect to uncus; valva tapering with distal tip pointed.

FEMALE (known only for O. eugenia ) (Fig. 2,3,5): Forewing length 25 mm (n=1). Wings: similar to male except more rounded, paler. Head, thorax, abdomen: similar to male except with sparse cream-colored hair-like scales among dark brown hair-like scales on ventral palpi, foreleg tarsus not reduced ( Fig. 3 E), covered with cream-colored scales and hair-like scales. Genitalia ( Fig. 5): inter-segmental membrane between 7 th and 8 th abdominal segments pleated and expandable, as in many other euptychiines; lamella postvaginalis forming broad lateral plates extending close to ( E. e. eugenia , Fig. 5 A) or fused with (E. e. transversa ) anterioventral edge of eighth tergum; posteriorlydirected, ‘scoop’-like plate ventral of ostium bursae ( Fig. 5 A, B); no sclerotized antrum, ductus bursae relatively long and broadening gradually into poorly defined corpus bursae that bears two elongate spiny signa ( Fig. 5 D), ductus seminalis joins ductus bursae dorsally very close to ostium bursae.

Etymology: The generic name is derived from the masculine Latin noun optimas, an aristocrat, combined with ‘Andes’, to which mountain range the genus is largely endemic, and it is considered masculine. The name is in reference to O. eugenia, whose specific name means ‘well-born’, as well as being a feminine given name; the Felders may have named the species after Empress Elisabeth Amalia Eugenia of Wittelsbach (“Sissi”) (1837-1898).

Relationships and taxonomy: Genetic sequence data show that Optimandes n. gen. is a member of the Pareuptychia clade ( Murray & Prowell, 2005), where it is strongly supported as monophyletic (100%) and moderately well supported (83%) as sister to Nhambikuara ( Espeland et al., 2019) ( Fig. 1). This result was also confirmed in ongoing analyses aimed at generating as comprehensive a phylogeny as possible for the Euptychiina , incorporating a much larger dataset including additional unpublished sequences, comprising>2000 sequences and representing>420 species (Espeland, unpublished data). This dataset includes the type species of 59 of the 61 available generic names for Euptychiina , and species closely related (based on morphology) to the remaining two type species.

As discussed above under Diagnosis and identification, Optimandes and Nhambikuara do not closely resemble one another in wing pattern or morphology, and the two seem ecologically rather distinct, with Nhambikuara predominantly a lowland genus and Optimandes confined to cloud forest. We therefore believe that they are best treated as distinct genera. The type species of Euptychoides , Euptychia saturnus Butler, 1867 (= Euptychoides laccine (C. Felder & R. Felder, 1867)) , is also a member of the Pareuptychia clade, but it is not close to Optimandes , instead being more closely related to Satyrotaygetis Forster, 1964 and Pareuptychia . Although a comprehensive phylogenetic anlaysis of the Pareuptychia clade based on morphological data has not been published, E. laccine does not show any obvious characters in the genitalia that might suggest a close relationship to Optimandes eugenia n. comb. The male genitalia of E. laccine differ from those of O. eugenia in having a much narrower uncus (in lateral view), with its dorsal edge more smoothly aligned with the dorsal edge of the tegumen, narrower valvae with a small dorsal projection just basal of the posterior tip, and the aedeagus becoming narrower anteriorly. These characters are also shared with E. pseudosaturnus and Euptychoides albofasciata (Hewitson, 1869) , consistent with molecular data that show a close relationship among these species ( Fig. 1, and Marín, unpublished data).

Optimandes eugenia has had a somewhat unstable history of generic classification. Butler (1877) considered O. eugenia eugenia (as ‘ Euptychia phineus ’) as a member of his ‘ E. harmonia group’, which included Euptychia gulnare Butler, 1870 , E. harmonia Butler, 1867 , Neonympha yphthima C. Felder & R. Felder, 1867 , E. nebulosa Butler, 1867 , and E. oreba Butler, 1870 . These five species are currently placed in five different genera. Weymer (1911) placed O. e. transversa in his ‘Saturnus Group’, which also included three species later placed in Euptychoides by Forster (1964) and Lamas (2004), namely Euptychoides laccine (referred to as ‘ E. saturnus ’ by Weymer), Euptychoides fida ( Weymer, 1911) and Graphita griphe (C. Felder & R. Felder, 1867) . However, Weymer (1911) placed O. eugenia eugenia (as ‘ E. phineus ’) in his ‘Harmonia Group’, corresponding to Hermeuptychia Forster, 1964 , of later authors, and evidently confused it with several other true Hermeuptychia occurring throughout the eastern Andes. Forster (1964) placed O. e. eugenia in Yphthimoides Forster, 1964 , and O. e. transversa in Euptychoides , apparently influenced by the presence or absence of white on the VHW. This character seems to perhaps be involved in mimicry, as we discuss further below, and based on molecular and morphological data we regard the two taxa as conspecific, following Lamas (2004).

Distribution and natural history: The genus is known from the Cordillera de la Costa in northern Venezuela, from both Andean slopes of Ecuador, and south of Ecuador along the eastern Andes to Bolivia ( Fig. 6). Both known species are uncommon to rare inhabitants of cloud forest, ranging from 1000-2200 m in elevation. The immature stages of O. eugenia transversa are described below.