Blepolenis Röber, 1906

Penz, Carla M., Mohammadi, Neda & Wahlberg, Niklas, 2011, Neotropical Blepolenis butterflies: wing pattern elements, phylogeny, and Pleistocene diversification (Lepidoptera, Nymphalidae), Zootaxa 2897, pp. 1-17 : 4-7

Blepolenis Röber, 1906

Fig. 1–2

Redescription. HEAD: nearly as wide as thorax; eyes glabrous, outlined with off-white scales; antenna red-brown, narrowly clubbed; labial palp approximately one-third longer than head, pale yellow with a broad brown ventral line. THORAX: coated dorsally with orange ( batea , catharinae ) or yellow ( bassus ) flat and piliform scales that match the color of the wing base; ventrally, both thoracic and leg scale cover are dark brown mixed with pale yellow ( batea , catharinae ) or pale yellow only ( bassus ), also matching predominant ventral wing colors. Mid and hind leg femur and tibia with numerous spines on the outer-lateral surface (small on femur, large on tibia; referred to as ‘dorsal spines’ in Penz 2007), one pair of apical tibial spurs present on mid and hind legs. Wing shape uniform between males and females; forewing triangular-shaped, distal margin straight or only slightly concave; hind wing oval-shaped, margins slightly scalloped. Dorsally, wings orange ( batea , catharinae ) or yellow ( bassus ) from basal to medial area, postmedial to marginal areas dark brown (broader in batea and catharinae than bassus ) with faint or distinct pale yellow markings along wing margins; wing orange/yellow areas usually allow visualization of ventral pattern, especially on the hind wing; three large forewing white subapical spots (usually small in Opsiphanes ), small additional subapical spots often present. Ventrally, forewing discal cell markings clearly defined, submedial/ medial areas solid yellow below the forewing discal cell, broad brown forewing postmedial band extended from anterior eyespots to posterior region of the wing – unique to this genus among all Brassolini , submarginal lines thin and wavy. The two ventral hind wing eyespots are bordered on each side by light-colored bars (wider and more visible in bassus than in batea and catharinae ). Wing pattern elements as defined by Nijhout (1991) are identified in a separate section below. Venation: forewing R 4 -R 5 fork deep; forewing crossveins m 1 -m 2 and m 2 -m 3 somewhat aligned, producing a truncate aspect to the distal end of the discal cell; forewing Cu 1 originates close to Cu 2 than to M 3. Androconial organs: ventral forewing androconial scales above 1 A+ 2 A present ( batea ) or absent ( bassus , catharinae ); dorsal hind wing discal cell hair pencil and pad of androconial scales adjacent to Cu 2 present ( batea ) or absent ( catharinae , bassus ), anal male hair pencil present ( batea , catharinae ) or absent ( bassus ). ABDOMEN: coated dorsally with orange ( batea , catharinae ) or yellow ( bassus ) flat and piliform scales, ventral scale cover dark brown mixed with pale yellow ( batea , catharinae ) or pale yellow only ( bassus ), both dorsal and ventral abdomen colors match colors of the wings. Abdominal androconial organ present on pleural membrane and above the spiracle of segment 4, large (occupying the entire length of segment 4) and oval-shaped, surrounded by an area of bare cuticle, scales of androconial organ broad, pale orange-yellow, oily in appearance. Male genitalia: tegumen large and domed; uncus evenly curved; gnathos paired, large, hook-shaped, with membranous articulation to tegumen; saccus almost as long as valva, cylindrical and narrowly constricted anteriorly; valva long, basally broad but narrowing distally to form a rounded tip, dorsal margin concave with a cluster of dorso-distal spines that vary in shape and size within and between species; phallus long and cylindrical, without basal coecum, tapered distal tip. Female genitalia: sterigma broader than long, delicate; posterior section (lamella postvaginalis) well-developed, smooth, and with prominent central lobe; anterior section (lamella antevaginalis) with two very thin arms curving inward that do not meet centrally but are, instead, separated by a membranous area; lateral section thin, extended dorsally to edge of tergum 8; ductus bursae long, distally narrow ( batea , bassus ) or broad ( catharinae ), corpus bursae ovalshaped; signa parallel, shorter than corpus bursae, and composed of small spines; posterior edge of papillae anales curved, covered with setae of approximately even length.

Comparison to Opsiphanes . Röber (1906) listed eleven characters in his original description of Blepolenis . The original German terminology for venation is given in Fig. 5 A, and the Comstock-Needham system is used in Fig. 5 B. These characters consist of: (1) forewing discal cell broader than in Opsiphanes ; (2) Costalis completely independent (“free-running”); (3) first Subcostalis branched at a larger distance from the end of the cell than in Opsiphanes ; (4) upper (vordere) Discocellularis longer than in Opsiphanes ; (5) mid (mittlere) Discocellularis completely straight, in the same direction with lower Discocellularis; (6) the latter interrupted in the center by a spur (weaker spur in Opsiphanes ); (7) origin of the mid Median at shorter distance of the upper Median than in Opsiphanes ; (8) pre-costal cell larger than Opsiphanes ; (9) lower (hintere) Discocellularis dividing the hind wing cell atrophied; (10) palpus slimmer than Opsiphanes ; (11) body less robust than Opsiphanes .

We examined these characters for all species and multiple specimens of Blepolenis , and compared them to Opsiphanes quiteria (also examined by Röber 1906), O. sallei Doubleday, 1849 (type species of Opsiphanes ), O. cassina C. Felder & R. Felder, 1862 , O. invirae , O. tamarindi , O. boisduvallii Doubleday, 1849 and O. cassiae (Linnaeus, 1758) (see Appendix 1). Röber’s character (2) is not informative, given that the costalis (costal vein) is independent in all brassolines. Three characters are incorrect (compare diagrams in Fig. 5): character (6), a spur is not present in the lower Discocellularis (crossvein m 2 -m 3) of either Blepolenis or Opsiphanes , a corresponding fold on the wing membrane might have misled Röber; character (7) appears to have been written incorrectly given that in Blepolenis the mid Median (Cu 1) originates close to the lower Median (Cu 2), and not the upper Median (M 3); finally, the hind wing lower Discocellularis (m 2 -m 3) is not atrophied in Blepolenis or any other brassoline, contradicting character (9). Five characters vary continuously within and between species of Blepolenis and Opsiphanes , and appear to be correlated with body size (3, 4, 8, 10, and 11). For example, although the body of B. bassus is delicate, that of B. batea and B. catharinae are comparable to Opsiphanes of similar wing size (character 11). Characters (1) and (5) refer to the forewing discal cell which indeed seems slightly broader and generally more truncated distally in Blepolenis than Opsiphanes species. Finally, Röber (1906) overlooked the fact that R 4 -R 5 fork is deeper in Blepolenis than Opsiphanes .