Lymanopoda ionius lilliput Pyrcz

Pyrcz, Tomasz W., Clavijo, Alejandra, Uribe, Sandra, Marin, Mario A., Alvarez, Carlos F. & Zubek, Anna, 2016, Páramo de Belmira as an important centre of endemism in the northern Colombian Andes: new evidence from Pronophilina butterflies (Lepidoptera: Nymphalidae, Satyrinae, Satyrini), Zootaxa 4179 (1), pp. 77-102 : 86-95

Lymanopoda ionius lilliput Pyrcz , n. ssp.

( Figs. 6 , 7 )

Type locality. Colombia, Antioquia department, municipality Belmira, vereda Río Arriba , El Morro.

Material examined. MEFLG –UN: HOLOTYPE Ƌ: municipality Belmira: Páramo El Morro, N 6°39’10.4” W75°40’17.2” 3220 m, 12.V.2012, 16.00h, BMC16489, baited trap ; ALLOTYPE ♀: pathway toward Páramo de Malvazá from La Truchera , N6°38’48.1” W75°42’48.3” 3227 m, 7.VII.2012, 13.35h, BMC17051, baited trap ; Paratypes (9 ƋƋ and 1 ♀): pathway toward Alto del Indio from La Truchera, N 6°37623’ W75°42.349’: 1 Ƌ: 3011 m, 11.XII.2011, 09.50h, BMC 11235 in pasture, net, C. F. Álvarez leg. ; 2 ƋƋ and 1 ♀: 2980 m, 5.VII.2012, 12.50 and 13.35h, BMC16684, BMC 16701 in patch of forest dominated by Tibouchina lepidota , prep. molec. 10.VIII.2013 I. C. Cadavid GSM, and BMC 16729 in patch dominated by shrubs, net, F. Restrepo and C. F. Álvarez leg.; 1 Ƌ: 3000 m, 7.VII.2012, 14.10h, BMC 17120 in forest clearing, net, A. Clavijo leg.; pathway toward Páramo de Malvazá from La Truchera , N6°38’42.3” W75°42’19.5”; 1 Ƌ male: 2966 m, 2.III.2012, 10.35h, BMC 11546 in forest clearing, net, C. F. Álvarez leg. prep. molec . 30.XI.2014 I. C. Cadavid GSM; 1 ♀: 3190 m, 4.III.2012, 13.40h, BMC11657, net, L. García leg.; Páramo de Malvazá, N 6°38’48.6” W75°42’46.0”: 1 Ƌ: 3225m, 5.III.2012, 14.15h, BMC14P75, net, A. Clavijo leg.; vereda Río Arriba , sector Montañitas, N6°36.592’ W75°39.302’: 1 Ƌ: 2820 m, 30.IX.2012, 14.22h, BMC17439, net, C. F. Álvarez leg. ; pathway toward Alto del Río, N6°40’42.92” W75°41’36.412”: 1 Ƌ: 2974 m, 20.XII.2012, 10.54h, BMC 17698 in mosaic, net, C.F. Álvarez leg.

Diagnosis. This subspecies differs immediately from all the remaining subspecies of L. ionius Westwood , [1851] found in the Colombian Western and Central Cordillera and in Ecuador by the significantly smaller size, comparable only to the size of the nominate subspecies found in the Eastern Cordillera , the latter differing in the less produced FW apex, the presence of FWD whitish subapical dots, also apparent in L. ionius excisa Weymer, 1911 , and the lighter ventral pattern expressed in the yellowish or yellow–orange FWV basal patch, instead of reddish in L. ionius lilliput , and the sulphur yellow HWV ground colour, reddish brown in L. ionius lilliput . The new subspecies differs from L. ionius excisa in the absence of FWD subapical white dots. Ventral colour patterns of the two subspecies are similar.

Description. MALE ( Fig. 6 A): Head, thorax and abdomen: similar to the nominate subspecies. FW length: 2 cm. FWD uniform chocolate brown, lustrous. HWD uniform chocolate brown, lustrous. FWV from base to median area dark red with several, small dark brown patches in discal cell and in spaces M3–Cu1 and Cu1–Cu2, distally blackish brown with a series of minute white dots three in subapical area and two in postdiscal area in M3–Cu1 and Cu1–Cu2, distal margin finely orange. HWV orange brown suffused with diffused dark brown scales in basal, postbasal and along distal margin, with a concentration in median area forming a straight oblique band running from apex to mid–inner margin.

MALE GENITALIA ( Fig. 7 ): Not differing noticeably from L. ionius excisa , dorsal process on the valva more prominent, longer than in the nominate subspecies.

FEMALE ( Fig. 6 B): Sexual dimorphism expressed in the lighter and duller dorsal brown wing colour, the presence of a reddish suffusion along HWD anal margin, and the dull, almost uniform light brown HWV colour.

FEMALE GENITALIA: Not examined.

Etymology. The epithet of this subspecies is derived from the fictional place of Lilliput, from Jonathan Swift’s novel Gulliver’s Travels, an island inhabited by tiny people, a term which currently applies to persons or objects of minute size, and is an allusion to the particularly small size of this subspecies of L. ionius .

Comments. Lymanopoda ionius is a fairly polytypic species occurring throughout the northern Andes of Ecuador and Colombia, marginally penetrating into Venezuela in the El Tamá range, and northern Peru in the Cajamarca department. The various subspecies differ in size, colour patterns and, to some extent, in male genitalia. Lymanopoda ionius ionius occurs throughout the Colombian Eastern Cordillera , except on the south–western slopes where it is replaced by L. ionius excisa which is also found in central and southern part of the Colombian Central Cordillera . Lymanopoda ionius gargantua Pyrcz & Rodríguez, 2007 is known from several localities in the Colombian Western Cordillera , whereas L. ionius browni Pyrcz, 1999 occurs in southern Ecuador and northernmost Peru. The population inhabiting the western slopes of the Andes in Ecuador possibly represents a separate, as yet undescribed subspecies. The alpha–taxonomy, ecology and phylogenetical position of L. ionius were discussed in several papers ( Adams 1986; Pyrcz 1999; Pyrcz et al. 1999; Pyrcz & Rodríguez 2007; Pyrcz & Viloria 2007; Casner & Pyrcz 2010).

Panyapedaliodes rojasi Pyrcz & Álvarez , n. sp. ( Figs. 8 , 9 , 10 )

Type locality. Colombia, Antioquia Department, Municipality of Belmira, vereda Río Arriba , El Morro.

Material examined. MEFLG –UN: HOLOTYPE Ƌ: Belmira Municipality, Páramo El Morro, N 6°39’51.412” W 75°40’6.96” 3223 m, 11.V.2012, 13.40h, BMC16418, in grassland, net, A. Clavijo leg. ; ALLOTYPE ♀: same locality as the holotype, N6°39’31.64” W75°40’20.60”, 3281 m, 12.V.2012, 14.35h, BMC16075, net, A. Clavijo leg. ; Paratypes (29 ƋƋ and 6 ♀♀): same locality as the holotype: 1 Ƌ: 3274 m, 23.I.2012, 13.05h, BMC14703, net, C. F. Álvarez leg. ; 1 ♀: 3223 m, 11.V.2012, 13.40h, BMC15120, net, A. Clavijo leg.; 16 ƋƋ: 3200–3300 m, 12/ 13.V.2012, 11.20h to 14.35h, BMC16090 and BMC 16509 in baited trap, BMC16079, BMC16085, BMC16087, BMC16113, BMC16222, BMC16417, BMC15095, BMC16424, BMC16425, BMC16427, BMC16461, BMC16465, BMC16466 and BMC 16223 in net, A. Clavijo, F. Restrepo and C. F. Álvarez leg.; 2 ƋƋ and 1 ♀: 3244 m, 14.V.2012, 11.30h, BMC15154, BMC15178 and BMC15180, net, A. Clavijo leg.; 2 males: 3223 m, 14.V.2012, 11.15h, BMC16252 and BMC16527, net, J. Duque and F. Restrepo leg.; pathway toward Alto del Indio from La Truchera, N6°38’2.9” W75°42’4.0”: 1 ♀: 2954 m, 7.VII.2012, 11.20h, BMC 17159 in grassland, net, A. Marín leg. ; 1 ♀: 2963 m, 4.III.2012, 12.50h, BMC 11872 in pasture, net, C. F. Álvarez leg.; pathway toward Alto de Malvazá from La Truchera, N6°38’42.3” W75°42’19.5”: 1 Ƌ: 2750 m, 3.VII.2012, 12.45h, BMC16973, net, F. Restrepo leg. ; vereda Río Arriba , sector Montañitas, N6°37’19.4” W75°38’43.4”: 7 ƋƋ and 1 ♀: 3120 m, 15/ 18.VI.2012, 10.45h to 14.00h, BMC15582, BMC15674 and BMC 15719 in baited trap, BMC 15398 in grassland near the lake, BMC15400, BMC 15429 in grassland, BMC 15506 in grassland and BMC 15724 in grassland, net, F. Restrepo, J. Duque and A. Marín leg. ; 1 ♀: 2789 m, 02.X.2012, 10.10h, BMC17309, net, C. F. Álvarez leg.

Diagnosis. Differs from the most similar species in the bole brown dorsal brown, medium brown in P. panyasis (Hewitson, 1862) and P. traceyannae Pyrcz & Viloria, 1999 , smaller size and more acute FW apex, larger lilac FWV subapical suffusion, the less prominent or absent submarginal sinuate yellow band, and the less prominent HWV chocolate brown overcast.

Description. MALE ( Fig.8 A): Head: eyes chocolate brown, covered with dense black hair; frons with a tuft of brown and sandy yellow hair; antennae to two–fifths the length of costa, slender, naked, light chestnut, club developed gradually, slightly thicker than shaft, made of 12 flagellomeres, ventrally orange–brown; labial palpi two times the length of head, covered with brown and sandy yellow hair, ventrally long, dorsally short. Thorax: black, dorsally covered with sparse golden brown hair, somewhat thicker on protothorax, metathorax and on the tegulae; legs brown–grey, femur covered with long, black hair, tibia and tarsus with sparse, sandy yellow scales. Abdomen: black, covered with black and golden scales, thicker ventrally, basal segments hairy. Wings: FW length 23–24, mean: 23.6mm, n=3; apex acute, outer margin straight, slightly truncate below apex; fringes intermittently sandy yellow and brown. FWD bole brown, lustrous with a large, a shade darker median androconial patch covering two–fifths of wing surface. FWV light chestnut; lilac dusting in subapical area; chocolate brown dusting along costal margin and outer margin along apex; in some individuals a faint sandy yellow, sinuate submarginal band extending from subapical area to tornus. HW rounded with undulating outer margin, the impression of scalloping sharpened by the intermittently sandy yellow intravenous, and brown fringes at vein ends. HWD bole brown, slightly darker in basal half. HWV ground colour marbled, predominantly chocolate brown with two lighter, chestnut areas, a narrow postbasal band and a wide postdiscal submarginal band with a sandy yellow basal edge and two basal incisions along veins Rs and in space Cu2–1A.

MALE GENITALIA ( Fig. 9 ): Tegumen dorsum with a shallow immersion in the middle; uncus one–fourth shorter than tegumen, slender and slightly bent downwards; subunci slender, approximately one–third the length of uncus; peduncuclus short, with a sharp tip pointing downwards; valvae oblong, with a smooth dorsum, ending with two roughly similar, short and blunt processes pointing distally; saccus deep and moderately wide, aligned with valva; aedeagus long, longer than saccus+valvae, thin and nearly straight with a small apical crest.

FEMALE ( Fig. 8 B): FW length 22 mm (n=2); FWD and HWD lighter chestnut brown; FWV subapical lilac dusting fainter, submarginal yellow, situate band not apparent; HWV considerably lighter and more uniformly patterned particularly in postdiscal to submarginal area, with a well–defined chocolate brown costal patch in apical area.

FEMALE GENITALIA ( Fig. 10 ): Papillae anales medium sized, rectangular in lateral view, more strongly sclerotized in basal part, membranous distally, were covered with dense and moderately long bristles, anterior apophysis developed, but very short; lamella postvaginalis massive, slightly sclerotized, bowl shaped; ductus bursa tubular, long, approximately as long as the length of bursa copulatrix, straight, lightly sclerotized, wider and more sclerotized at opening into bursa copulatrix, with ductus seminalis originating close to the opining of bursa; bursa copulatrix large, oval, with two prominent parallel signa.

Etymology. This species epithet comes by Hector Rojas an enthusiastic guide and conservationist dedicated to Belmira environmental protection, who contributed significantly in the butterfly inventory, from sampling areas definition to logistical support and guidance.

Comments. Panyapedaliodes rojasi is morphologically most similar to P. panyasis . The latter is a widespread species occurring throughout the Andes, including the isolated ranges such as the Sierra Nevada de Santa Marta and the Venezuelan Cordillera de la Costa. It is found in Bolivia, Peru, Ecuador and Colombia. Ecologically speaking, however, P. ro j a s i is more alike P. traceyannae because the two species occur in similar habitat, the forest—paramo ecotone. Panyapedaliodes panyasis is a mid–elevation dense cloud forest species found at 2200– 2800 m ( Adams 1986; Pyrcz & Garlacz 2012). Panyapedaliodes traceyannae occurs at 2800–3200 m (Pyrcz et al. 1999), which is about the same elevational band as P. ro j a s i. So far, neither P. panyasis nor P. traceyannae were reported from Antioquia, although considered the wide distribution of P. panyasis its presence is there is very likely. In most of southern Colombia and Ecuador P. panyasis and P. traceyannae are parapatric along an elevational gradient. Southwards, P. traceyannae is replaced by its close allopatric ally, P. mara (Thieme, 1905) . It is most probable that P. ro j a s i is a genuine endemic of the Belmira massif. It was not found so far in other areas of Antioquia at elevations around 3000 m, especially in the highest parts of the San Felix and Padre Amaya massifs, although the latter two areas support no paramo vegetation, and only pockets of subparamo. Barcode data supported this new species separate specific status relative to other Panyapedaliodes COI sequences ( Fig.15 ).

Pedaliodes nutabe Pyrcz & Álvarez , n. sp. ( Figs. 11 , 12 , 13)

Type locality. Colombia, Antioquia department, Belmira municipality, vereda El Yerbal, Alto de Malvazá.

Material examined. MEFLG –UN: HOLOTYPE Ƌ: Belmira Municipality : pathway toward Alto de Malvazá from La Truchera, Páramo Malvazá N 6°38’47.8” W75°42’52.6” 3264 m, 5.III.2012, 13.45h, BMC14947, baited trap ; ALLOTYPE ♀: vereda Río Arriba , páramo El Morro, N 6°39’19.6” W75°40’18.7”, 3240 m, 23.I.2012, 14.40h, BMC14768, baited trap, prep. genit. 04/ 26.04.2013 J. Lorenc MZUJ ; Paratypes (20 ƋƋ): same locality as the holotype: 1 Ƌ: 3291 m, 16.I.2012, 11.30h, BMC11178, net, C. F. Álvarez leg. ; 5 ƋƋ: 2856–3290 m, 2– 5.III.2012, 12.10h to 15.45h, BMC11622 and BMC 14955 in baited trap, BMC11528, BMC17074 and BMC 14979 in net, C. F. Álvarez, A. Clavijo and F. Restrepo leg.; 6 ƋƋ: 3215–3290 m, 4–7.III.2012, 14.20h to 15.00h, BMC16772, BMC16776, BMC17040, BMC17078, BMC17080 and BMC 17213 in baited trap; vereda Río Arriba , páramo El Morro, N 6°39’51.412” W 75°40’6.96”: 2 ƋƋ : 3240 m, 24.I.2012, 11.40h to 13.00h, BMC 14665 in baited trap and BMC 14670 in net, C. F. Álvarez leg.; 6 ƋƋ: 12.V.2012, 14.45h, BMC16088, BMC16094, BMC16438, BMC16491, BMC16500 and BMC 16501 in baited trap.

Diagnosis. Smaller than the most closely allied P. n e gre t i Pyrcz, 1999, with a darker, blackish brown upperside and HWV. Similar in upperside colour and size to P. pheretias (Hewitson, 1872) from which it differs in the expression of the HWV median band, wider and continuous between anal and costal margin, constricted in the middle or discontinuous in P. pheretias , and in another somewhat similar species, P. hardyi Adams, 1986 . FW apex more acute than in P. fassli Weymer, 1912 , the latter differing also in the wider HWV median band, particularly in the middle section.

Description. MALE ( Fig. 11 A): Head: eyes chocolate brown, covered with dense black hair; frons with a tuft of brown and sandy yellow hair; antennae to two–fifths the length of costa, slender, naked, dark brown, club developed gradually, slightly thicker than shaft, made of 12 flagellomeres, ventrally chestnut; labial palpi two times the length of head, covered with long, brown and sandy yellow hair ventrally, dorsally with short milky white and black hair. Thorax: black, dorsally covered with sparse golden brown hair, thicker on the tegulae; legs grey, femur covered with long, black hair, tibia and tarsus mostly naked. Abdomen: black, covered with black and golden scales, thicker ventrally, basal segments hairy. Wings: FW length 25 mm (n=3), apex subacute, outer margin straight; fringes intermittently dark brown and sandy yellow. FWD uniform blackish brown, lustrous; androconial patch small, some 2 mm wide, extending in median area from discal cell end to anal margin, compact. FWV blackish brown; a short costal streak in postdiscal area; subapical area dusted with sparse, white scales; and apical area with some chocolate brown; a faint black submarginal line extending from costa to Cu1–Cu2, barely noticeable in some individuals. HW rounded with slightly undulating outer margin; fringes intermittently brown and sandy yellow in apical area, elsewhere brown. HWD uniform blackish brown, densely hairy in median half. HWV black; a wide median band extending from costal to anal margin, from costa to M2–M3 white, then golden yellow, constricted and displaced distally between Rs and M3, gradually widening from M3 to anal margin were reaching some 4 mm in width, dusted with blackish brown scales on the yellow portion; a series of minute whitish submarginal dots, in some individuals not apparent; some white scaling along outer margin.

MALE GENITALIA ( Fig. 12 ): Dorsum of tegumen flat; uncus 3/4rd the length of tegumen, nearly straight, stout with a blunt tip; subunci slender, less than half the length of uncus; pedunculus short; valvae slender with a bumped, smooth dorsum, extending into a prominent, sharp dorsal process pointing distally, and a blunt tip; saccus moderately long and wide; aedeagus 1 1/4th longer than saccus+valva, strongly contorted and flattened dorso– ventrally with a sharp, minute tip.

FEMALE ( Fig. 11 B): Differing slightly from the male, only in the lighter brown upperside, and lighter and considerably paler HWV.

FEMALE GENITALIA ( Fig. 13): Papillae anales medium sized, rectangular in lateral view, rather well sclerotized over the whole surface, covered with dense and long bristles, anterior apophysis developed, roughly half the length of papillae from base to apex; lamella postvaginalis wide, slat–like; ductus bursa tubular, long, approximately as long as the length of bursa copulatrix, s–shaped, lightly sclerotized, gradually opening into bursa copulatrix, with ductus seminalis originating close to the opining of bursa; bursa copulatrix large, oval, with two prominent parallel signa.

Etymology. This species epithet, nutabe makes an allusion and pays tribute to an important pre–Colombian indigenous groups, farmers in essence, but also fishermen and alluvium gold miners that inhabited central Antioquia department including the research area.

Comments. Pedaliodes nutabe is, judging from its external and genital morphology, closely allied to a number of species, which possibly form a monophyletic group ( Pyrcz & Rodríguez 2007), although the astonishing scarce published molecular data in the genera (in relation to his ecological prevalence in butterfly Andean communities), strongly reduces integrative taxonomic inferences. Nevertheless, they all can be recognized by the white and yellow HWV median band, which in some species is wide and fully developed, in others constricted, in others restricted to a costal streak and an anal wedge. Nevertheless, in all these species, the male genitalia is very similar with a diagnostic sharp subapical dorsal process and a blunt apex. In geographical terms, the closest ally is P. negreti found in the southern part of the Central Cordillera and in Ecuador as far south as the Pastaza valley (Pyrcz 1999). South of it, it is replaced by P. pheretias which occurs into extreme northern Peru. In the Colombian Western Cordillera there are at least two species of this group, including P. f as s l i found only in the Farallones de Cali area, P. tatama Pyrcz & Rodríguez, 2007 occurring in the Tatamá range, and possibly one other species recently discovered in Frontino, known so far from one specimen only (Pyrcz & Rodríguez op. cit.). In the Eastern Cordillera , two species belonging to this group occur allopatrically on the western, P. ha rd y i, and eastern slopes, P. arnotti Adams,1986 . If we adopt a lumper approach, all these taxa could possibly be treated as the subspecies of one widely distributed polytypic species. However, they are geographically well separated, and apparently historically isolated for longer periods of time, thus they are best to be considered as specifically distinct until a more focused molecular based phylogenetical work is performed. Pedaliodes nutabe is not entirely endemic of the Belmira range, contrary to other two species described here ( Lymanopoda casneri and Panyapedaliodes rojasi ). It was found as well in the uppermost part of the San Felix range. All the “ Pedaliodes pheretias ” species are confined to the uppermost forest and forest–paramo ecotone but they occasionally fly in the open paramo as well (Pyrcz & Rodríguez op. cit.).

Figures

FIGURE 6. Adultd of Lymanopoda ionius lilliput n. ssp.; top: upperside, bottom: underside. A, male (paratype). B, female (paratype).

FIGURE 7. Male genitalia of Lymanopoda ionius lilliput n. ssp. (paratype): lateral view, aedeagus extracted from its natural position.

FIGURE 8. Adults of Panyapedaliodes rojasi n. sp. top: upperside, bottom: underside; A, male (paratype). B, female (paratype).

FIGURE 9. Male genitalia of Panyapedaliodes rojasi n. sp. (paratype). A, lateral view. B, aedeagus, lateral view.

FIGURE 10. Female genitalia of Panyapedaliodes rojasi n. sp. (paratype), lateral view.

FIGURE 11. Adults of Pedaliodes nutabe n. sp.; top: upperside, bottom: underside. A, male (paratype). B, female (paratype).

FIGURE 12. Male genitalia of Pedaliodes nutabe n. sp. (paratype). A, lateral view. B, aedeagus, lateral view. C, aedeagus, dorsal view.

FIGURE 15. NJ tree using the uncorrected p distance model, based on DNA barcode sequences (ca. 658 bp of COI). Above the branches bootstrap support values (> 50) are provided for each node.