Modica Zacca, Casagrande & Willmott, 2023

Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), Zoological Research 2023, pp. 1-73 : 42-67

Modica Zacca, Casagrande & Willmott , genus novum.

Type species — Euptychia confusa Staudinger, 1887 , by present designation.

Zoobank registration: https://zoobank.org/Nomenclatural Acts/D3CE81CE-5587-41D5-AFB6-934343C11D6F

Systematic placement and diagnosis. Modica gen.n. is a member of the ‘ Splendeuptychia clade’ ( Figure 10 ), in which its monophyly is strongly supported (FULL dataset SH-aLRT =100, UFB = 100). Its relationships to other members of the clade, however, are not clear; the genus is placed as sister to Parypthimoides , but support is inconclusive (FULL dataset SH-aLRT 100, UFB 43), and is further found within the same strongly supported clade that also contains Deltaya gen.n. (see under that genus), Scriptor , Emeryus , Colombeia and Malaveria (FULL dataset SH-aLRT 100, UFB 95). As with Deltaya gen.n., no morphological synapomorphies were identified for Modica gen.n., but these two genera share the same somewhat distinctive characters within Euptychiina (see under the former genus). Overall, the genus is best distinguished from phenotypically similar genera by the pale pupils of the HW ocellus in Cu 2 -Cu 1 being visible on the dorsal surface, by the VHW postdiscal ocelli in cells Cu 1 -M 3 and M 3 -M 2 being of similar size, round (not elongate) and each containing two enlarged, elongated silver pupils, and by the marginal line not widening in the VHW tornus ( Figures 37 and 38 ).

The genitalia of both sexes ( Figure 39 ) are described below, and, as with Deltaya gen.n., they are broadly similar to those of a number of more or less distantly related euptychiine genera, in other words lacking obvious synapomorphies. The eighth abdominal segment of the female is only slightly pleated and expandable, similar to Scriptor and some related species but differing from others in which it is fully pleated and expandable, with this character varying within genera (e.g. within Paryphthimoides and Deltaya gen.n.). The female genitalia also lacks a sclerotized lamella antevaginalis and there is no sclerotized plate present on the ventral intersegmental membrane of the seventh and eighth abdominal segments, unlike Scriptor and some species of Deltaya gen.n. Characters that differ among the genera within the clade in which Modica gen.n. is placed are summarized in Table 1 .

Etymology. The generic name is a feminine Latin adjective treated as a noun in the nominative singular, meaning something that is modest, ordinary, average, in reference to the ‘typical’ euptychiine morphology of this genus and its lack of obvious distinguishing characters.

Description ( Figures 37 – 39 ). Some notable characters include: eyes setose; pterothoracic legs dorsally slightly darker, tibia with two principal longitudinal rows of spines ventrally, pair of spurs of similar length at distal end of tibia, first tarsomere with three principal longitudinal rows of spines ventrally, remaining tarsomeres with four principal longitudinal rows of spines ventrally. Medium-sized Euptychiina (FW length typically 21 – 28 mm), FW triangular and rather rounded at apex, HW rounded. No strong sexual dimorphism: Dorsal wings dark brown to greyish brown, pale pupils of ocellus in cell Cu 2 -Cu 1 visible on DHW, no androconial scales present. Ventral wings greyish brown to yellowish brown; relatively broad, dark brown to reddish brown discal and postdiscal lines traversing both wings; VFW with three postdiscal ocelli in cells Cu 1 -M 3, M 3 -M 2, M 2 -M 1, anterior ocellus more clearly marked than remainder, lying within a broad dark brown band (umbra), with this band and adjacent area basally in middle of wing tinged yellowish in M. myncea comb.n. and M. confusa comb.n.; VHW similar to VFW but with five postdiscal ocelli between Cu 2 and Rs, those in cells Cu 2 -Cu 1 and M 2 -M 1 typically slightly larger, black-centred with two silver dots in each ocellus as pupils, ocellus in M 1 -Rs similar but much smaller, and those in cell Cu 1 -M 2 with dark brown centres and elongate silver pupils; marginal line thin and even throughout, not thickening at tornus. Male 8th abdominal tergite reduced dorsally, leaving a sclerotized strip along anterior edge and usually an isolated sclerotized patch in posterior portion. Male genitalia with uncus longer than tegumen, brachia approximately parallel with uncus and about two-thirds its length; valvae elongate with dorsal edge straight or with slight projection; aedeagus adorned with scattered small spines in some species, and with or without cornuti. Female genitalia has 8th tergite reduced to a posterior sclerotized patch about two-thirds width of segment, intersegmental membrane between seventh and eighth abdominal segments only somewhat expandable with no strongly sclerotized plate ventrally, eighth segment with large irregular lateral sclerotized plate extending further dorsally at anterior edge, lamella antevaginalis and antrum unsclerotized, ductus bursae unsclerotized, corpus bursae small, oval and with two narrow sub-parallel signa.

Distribution and natural history ( Figure 40 ). Modica gen.n. contains five described species and several undescribed species (Zacca et al., unpublished data), which occur in rainforest from sea level to 1300 m, ranging from southern Mexico to western Ecuador and throughout the Amazon and Guianas to south-eastern Brazil. The genus reaches its peak diversity in the western Amazon, where both sexes may be common throughout the understory of both disturbed and undisturbed forest, with some species also occurring along forest edges and in overgrown, shady plantations. Males of some species perch from 1 to 3 m in the forest understory in the morning and late afternoon, sometimes on hilltops, and both sexes are attracted to rotting fruit ( DeVries, 1987; Zacca et al., pers. obs.). Notes on the immature stages of M. myncea comb.n. and M. confusa comb.n. were provided by Singer et al. (1983), with hostplants (natural and in captivity) including Cyperaceae, Palmae , Poaceae and Marantaceae (see also Beccaloni et al., 2008; Singer & Ehrich, 1993).

Discussion. The type species for this genus, Euptychia confusa , was described by Staudinger (1884) based on an unstated number of specimens from Chiriquí, Panama, and a lectotype at the MfN was designated by Singer et al. (1983). The female illustrated in Staudinger (1884: pl. 80) agrees with the description, although it was incorrectly labelled as ‘ Euptychia myncea ’. We chose to designate Euptychia confusa as the type species for this genus since DNA barcode data suggest the possibility of cryptic species within Modica myncea comb.n. that remain to be resolved.

Although strongly supported by the molecular data, Modica gen.n. lacks any clear morphological synapomorphies and it is thus not surprising that a close relationship among all of its constituent species was unnoticed until recently; M. confusa comb.n., M. myncea comb.n. and M. maripa comb.n. were hitherto placed in Cissia (e.g. Forster, 1964 [under the name Argyreuptychia ]; Singer et al., 1983; Lamas, 2004; Brévignon, 2005) and M. fugitiva comb.n. and M. kamel comb.n. were placed in Magneuptychia ( Benmesbah et al., 2018; Forster, 1964; Lamas, 2004). As discussed by Zacca, Casagrande, et al. (2018), most species placed in Cissia prior to that paper were presumably considered to be related because of their possession of a yellowish patch on the VFW, but that character is clearly homoplasious, and the Cissia of Lamas (2004) are now placed in six genera in two clades: Cissia , Vanima , Megisto, Modica , Paryphthimoides , and Vareuptychia . Benmesbah et al. (2018) noted that M. myncea comb.n., M. maripa comb.n., M. fugitiva and M. kamel comb.n. were likely closely related, based on morphology and preliminary molecular data, and suggested that a new genus might be needed to accommodate these species. Our phylogenetic analysis failed to strongly resolve the relationships of Modica gen.n. to other clades. These species therefore cannot reasonably be accommodated in any described genus without combining at least five described genera, along with Modica gen.n. and Deltaya gen.n., into a single, large genus of morphologically and ecologically dissimilar species, which would not correspond to any author ’ s prior taxonomic hypothesis.

Modica Zacca, Casagrande & Willmott , gen.n.

confusa (Staudinger, 1887) , comb.n., was Cissia

fugitiva (Lamas [1997]) , repl. name, comb.n., was Magneuptychia

= helle (Cramer, 1779) , preocc. (not [Denis & Schiffermüller], 1775), comb.n., was Magneuptychia

kamel ( Benmesbah & Zacca, 2018) , comb.n., was Magneuptychia [ Benmesbah et al. (2018, Zootaxa, 4425(1): 115-145)]

maripa ( Brévignon, 2005) , comb.n., was Cissia [ Brévignon (2005, Lambillionea, 105(3)(1): 393-404)]

myncea (Cramer, 1780) , comb.n., was Cissia

= myncena (Stoll, 1782), missp., comb.n., was Cissia

= crantor (Fabricius, 1793) , comb.n., was Cissia

= clerica (Herrich-Schäffer, 1865), repl. name, comb.n., was Cissia

= pytheus (MÖschler, 1883) , comb.n., was Cissia

= isolata (Kaye, 1921), comb. n., was Cissia [Kaye (1921, Memoirs of the Department of Agriculture of Trinidad and Tobago, 2: i-xii, 13-163, 1 pl.)]

Figures

FIGURE 10 Phylogeny of the ’Splendeuptychia clade’ based on the FULL dataset. Interpretation of support values can be found in the legend. The scale bars under the butterfly images equal 1 cm.

FIGURE 37 Modica confusa comb.n. (a) male wing venation; (b and c) male (FLMNH-MGCL-269550) dorsal (b) and ventral (c); (d and e) female dorsal (d) and ventral (e). Scale bars 1 cm.

FIGURE 38 Modica gen.n. representative species. (a) M. myncea comb.n (neotype); (b) M. fugitiva comb.n (neotype); (c) M. maripa comb.n (MB-1708-13); (d) M. kamel comb.n (KW-081111-67). Scale bars 1 cm.

FIGURE 39 Modica confusa comb.n. (a–d) male genitalia (dissection KW-21-64), lateral (a) with posterior view juxta, dorsal (b), aedeagus lateral (c) and aedeagus dorsal (d); (e–h) female genitalia (dissection KW-21-65), lateral view exterior tip abdomen (e), ventral view exterior tip abdomen (f), dorsal view interior abdomen (g), corpus bursae perpendicular to signa (h). Scale bars 1 mm.

FIGURE 40 Modica gen.n. Species diversity mapped on a 2 degree grid. Colours ranging from dark green to red represent increasing diversity.