Nivaliodes Pyrcz & Mahecha-J. & Boyer & Lachowska-Cielik & Cerdeña & Farfán & Garlacz & Lorenc-Brudecka & Bálint & Fåhraeus & Zając-Garlacz & Espeland 2024, gen. nov.

Nivaliodes Pyrcz & Espeland gen. nov. Nivaliodes Pyrcz & Espeland: LSIDurn:lsid:zoobank.org:act: E2E63ED0-6829-439F-8948-186DAC61B112 Type species: Nivaliodes negrobueno Pyrcz, Fåhraeus & Boyer sp. nov. Diagnosis and description Adults of medium size, with a FW with costa length between 21 and 28 mm, with brown or blackish brown FWD and HWD, except for a white postdiscal costal patch in the FW, and with an almost uniform underside with some faint white scaling, blackbrown FW and black HW freely sprinkled by snow-white scales, HWV median costal patch whitish. No androconial patch, except for a few vestigial androconial scales detected under the scanning electron microscope in one of the three known species, Nivaliodes puriq (Figs. 7, 8). Wing venation similar to other genera of the Pedaliodes clade (see: Pyrcz et al. 2023). Eyes densely hairy; palpi two times the length of head; antennae reaching two-fisshs the length of costa, slender, mostly naked, with a club formed gradually, composed of 11 flagellomeres, flattened dorsoventrally. Male genitalia: With stout uncus and subunci and, contrary to the species of Pedaliodes, the valva is devoid of any dorsal process, instead with a humped, either slightly irregular or serrate dorsum, and with a short, straight, and wide, tubular aedeagus, differing from the contorted and usually considerably longer aedeagus of Pedaliodes, bearing a sharp apical ending and a widened proximal opening. Female genitalia: Papillae anales prominent, postvaginal lamella wide and strongly sclerotized with a smooth surface, enclosing a spacious antrum; ductus bursae mildly sclerotized, medium long and very wide, opening gradually into a large, pear-like corpus bursae, with a smooth surface, no signa, unlike the species of Pedaliodes, which invariably have two parallel signa on corpus bursae. Scale morphology Uppersides of wings are overlaid with two thickly packed layers of flat, cover scales, the top layer with predominantly elongated scales with dentate apices, and the bottom layer with scarcer, wide scales with gently undulating apices, visible only when top layer scales are removed; long, hairy scales are rather sparse and concentrate along outer areas of wings, fringes composed of elongated brushy scales. There are no specialized androconial scales. Scale ultramorphology is homogeneous, with the body of the individual scales being structured by longitudinal ridges and cross ribs connected by trabeculae, with no apparent lower-level lamina (Figs 7, 8). Etymology A compound noun: ‘ Nival- ’, root of snowy in Latin, - liodes, an allusion to Pedaliodes. Noun grammatical gender is considered masculine. Phylogeny and radiation The genus Nivaliodes is fully supported and sister to the genus Pherepedaliodes Forster, 1964 (type species: Pedaliodes pheretiades Grose-Smith & Kirby, 1894), with the two forming a fully supported clade sister to Pedaliodes + the monotypic Dangond Adams & Bernard, 1979 (type species: Dangond dangondi Adams & Bernard, 1979), within the diverse ‘ Pedaliodes clade’ of the subtribe Pronophilina (Nymphalidae: Satyrinae: Satyrini) (Fig. 1). The topology of Nivaliodes shows a fully supported Nivaliodes viracocha sp. nov. sister to N. puriq sp. nov. + N. negrobueno sp. nov. The latter clade is not entirely resolved, with a monophyletic N. negrobueno sp. nov. but a paraphyletic N. puriq owing to the position of the sample DLC127. The latter comes from a different locality from the remaining N. puriq, from which only two specimens are available now, and only one was sequenced. The divergence of Nivaliodes started in the late Pliocene, ~3 Mya, and continued through the Pleistocene. The split between Nivaliodes and Pherepedaliodes occurred at the break of the Miocene and Pliocene some 5 Mya, and was preceded marginally by the separation of the Pherepedaliodes + Nivaliodes clade from Pedaliodes, which occurred some 5.6 Mya (Fig. 2).