Systematic revision of the Andean butterfly genus Orophila Staudinger, 1886 (Lepidoptera: Nymphalidae: Biblidinae) Author Zubek, Anna Author Pyrcz, Tomasz Author Lorenc-Brudecka, Jadwiga Author Boyer, Pierre Author Cerdeña, Jose Author Mahecha-Jiménez, Oscar Author Huertas, Blanca text Zootaxa 2017 4258 5 443 461 journal article 33125 10.11646/zootaxa.4258.5.3 31fc11cb-e4a0-4587-8312-300525996a24 1175-5326 570422 D59801DA-B5AD-42EA-9114-BE398CC84864 Orophila cardases ( Hewitson, 1869 ) ( Fig. 1 B, F) Cybdelis cardases Hewitson 1869 : 91 , pl. 45, fig. 25. Type locality: Ecuador. Syntype male, NHMUK [examined]. = Perisama diotima zurita Fruhstorfer, 1916 : 97 . Type locality: Ecuador /west/. Syntypes (3) male, NHMUK [examined]. = Perisama diotima demata Fruhstorfer, 1916 : 97 . Type locality: Macas, Ecuador. Syntype male, NHMUK [examined]. Myscelia cardases (Hewitson) ; Kirby, 1871 : 202 . Cybdelis cardases (Hewitson) ; Kirby, 1879 : 75 . Orophila cardases (Hewitson) ; Staudinger, 1886 : 112 ; Fassl, 1914 : 99 ; Neild, 1996: 54, fig. 294, 295. Perisama diotima (Hewitson) ; Röber, 1915 : 499 , pl. 102 B ( cardases considered a synonym of Perisama diotima (Hewitson) . Perisama campaspe cardases (Hewitson) ; Fruhstorfer, 1916 : 97 . Perisama cardases (Hewitson) ; Oberthür, 1916 : 100 ; d'Abrera, 1987 : 565 , 2 figs. Perisama diotima zurita Fruhstorfer ; Röber, 1924 : 1034 . Perisama diotima demata Fruhstorfer ; Röber, 1924 : 1034 . Odontosama cardases (Hewitson) ; Aurivillius, 1929: 161. Orophila cardases cardases (Hewitson) ; Lamas, 2004 : 245 . Orophila cardases zurita Fruhstorfer ; Lamas, 2004 : 245 . Diagnostic characters. Metallic blue rays on DF reach as far as the cell CuA1–CuA2. DH uniformly brown, no ashen-blue band. Basal red patch on VF usually small, limited to the discal cell, never covering the basal part of CuA1–CuA2, in some populations with additional red submarginal markings. Redescription. MALE ( Fig. 1 B): Wings: FW length: 17–23 mm (n=96) DF: ground colour blackish brown, small diffuse white subapical mark; both the two dark blue metallic rays in the basal area and the set of five irregular spots in postmedian area thicker and larger, than in O. campaspe ; fringe brown, with creamy-white marks between the veins; DH: almost uniform, blackish-brown, with long, hair-like scales covering the basal half of the wing, sometimes a few metallic blue scales in median area; VF: basal part ground colour blackish brown, getting lighter towards the anal margin; apical triangle light brown, with a thin, wavy, dark brown submarginal line; small, white spot in submarginal area, where the two ground colours meet; a series of white, light brown and red markings in postmedian area: smaller white and red ones in R5–M1, M1–M2, and M2–M3, and two larger red ones in M3–CuA1 and CuA1–CuA2; basal red patch in the majority of specimens limited to the discal cell and varying in size from almost absent to covering the whole discal cell ( Fig. 3 A–E), sometimes accompanied by blue markings, in some specimens ( Fig. 3 F–H) basal red patch expands outside the discal cell and is fused with median red markings (see also the 'Remarks' section); marginal line brownish-red, usually disappearing around the cubital veins. VH: ground colour pale brown and yellowish-brown, varying in shade throughout the wing surface; marginal line thin and red; submarginal line dark brown, with a series of curves; marginal area usually yellowish and somewhat lighter than the rest of the wing; postdiscal line irregularly wavy; costal area with white marking at the base and pale yellow marking between postdiscal and submarginal lines; little white spot in M1–M2 touching the postdiscal line; three to five diffused dark-brown submarginal dots; two or three faint, diffused white markings in submarginal area. MALE GENITALIA: similar to O. campaspe : as in the general description, except for hypandrium: fork strongly sclerotized, with a U-shaped opening and sharp, pronounced, claw-shaped, tips; collar thin, longer than body, smoothly widening at both ends; body triangular, slightly shorter and twice wider than the collar, with a deep, semi-circular proximal incision, saccus slightly longer than tegumen and uncus combined. FEMALE ( Fig. 1 F): differs from male in: DF discal metallic blue rays reduced, larger and paler median blue markings; DH and ventral pattern same as in males. FEMALE GENITALIA: papillae anales sclerotized interiorly, thin apophyses, longer than the width of the papillae, with club-shaped tips, ostium bursae wide and irregularly shaped; distal half of ductus bursae sclerotized, followed by a shorter and wider, unsclerotized part, opening smoothly into the corpus bursae; signa in a form of two small sclerotized dots, corpus bursae oval, as long as ductus bursae lamella antevaginalis expanded into a large pleated lobe, widening proximally, as long as the sclerotized part of the ductus. Remarks. Lamas (2004) synonymized O. demata with O. zurita , but not with the nominate O. cardases , most likely because of the difference in VF apical and median red patches size in type specimens. The type specimen of Cybdelis cardases ( Fig. 2 H) exhibits large basal and median red patches, while in type specimens of both, Perisama diotima demata ( Fig. 2 J) and P. d. zurita ( Fig. 2 I) the red patches are reduced. Our study revealed that among the individuals found in the same locality the discal and median red patch differs from absent to large ( Fig. 3 A–H), as a result, O. c. zurita should be considered a synonym of O. c. cardases . Orophila cardases displays considerable infraspecific variation affecting the size and the shape of the VF basal red patch. Not only some individual variation is observed within the populations where larger numbers of individuals were examined, but a geographical pattern can also be identified. Specimens from central Ecuador and southern Colombia present the smallest patch of all. In some of them, the patch is very small indeed, restricted to basal area, faint and suffused with brown scales or even totally absent. The latter specimens occur on both the western and eastern slopes of the Andes in Ecuador , however they are more frequent on the Pacific slopes in Pichincha and Cotopaxi (Boyer, pers. comm.). Towards the north and south from the distribution centre, the VF patch becomes progressively larger. This is noticeable in southern Ecuador (Morona–Santiago and Zamora– Chinchipe) ( Fig. 3 G) and northern Peru ( Cajamarca , Amazonas ) where specimens bearing particularly large patches covering most of discal cell coupled with large postdiscal patches can be observed in local populations, alongside small patched individuals. However it is on the northern and north-western peripheries of the geographic range of O. cardases , in the Colombian Western Cordillera and the northern extremity of the Central Cordillera ( Antioquia ) where large patch red pattern not only dominates but is seemingly exclusive of local populations. It covers basal two-thirds of discal cell and extends into three large postdiscal patches forming a large continuous red area ( Fig. 3 H). The size of the red patch probably misguided Salazar (2007) who upon examining the individuals from Antioquia and Risaralda considered them to represent the Peruvian—south Ecuadorian O. cecidas which exhibits the largest basal red patch among all Orophila species. The Western Cordillera and northern Central Cordillera populations could in fact represent a separate subspecies however more individuals from the entire range have to be examined, also including molecular data, in order to be conclusive about the extent of genetic divergence in O. cardases . Interestingly, however the individuals of O. cardases from the Colombian Eastern Cordillera are characterized by a rather small red patch and inversely, in O. campaspe , whose range overlaps in Cundinamarca and Boyacá with O. cardases , the patch is equally large and similarly shaped. There are very few specimens from the northern part of the Colombian Eastern Cordillera ( Fig. 3 F) and only one is known from the Venezuelan Cordillera de Mérida , and those have rather small reddish patches, therefore we cannot speculate about local individual variation. Possibly, the red patch is an expression of a recessive gene or/and is related with selective pressure. During the re-curation of the NHMUK collections two previously unidentified syntypes of Perisama diotima zurita Fruhstorfer were found. Type material. NHMUK: Syntype ♂ of Cybdelis cardases ( Fig. 2 H): SYN-/TYPE // Type // Ecuador/ Hewitson Coll./79–69./ Cybdelis cardaces1. // B.M. TYPE/No. Rh. 9350./ Cybdelis / cardases /♂ Hew. // C. Cardaces SIC // 249 // Illustrated in/"The Butterflies/of Venezuela"/A. Neild, 1996 // BMNH(E) 1670021. Syntype # 1 ♂ of Perisama diotima zurita ( Fig. 2 I): SYN-/ TYPE // Ecuador /ex coll Fruhstorfer // BMNH (E) 1718888 // SYNTYPE ♂ / Perisama diotima zurita ( Fruhstorfer, 1916 ) /Det. Zubek & Huertas , 2016 ; Syntype # 2 ♂ of Perisama diotima zurita : SYN-/ TYPE // Paratype // Ecuador /ex.coll. Fruhstorfer // diotima / zurita Fr. // Fruhstorfer Coll./B.M. 1937-285 // BMNH (E) 1670029 // SYNTYPE ♂ / Perisama diotima zurita ( Fruhstorfer, 1916 ) /Det. Zubek & Huertas, 2016; *abdomen missing . Syntype # 3 ♂ of Perisama diotima zurita : SYN-/ TYPE // Ecuador /Fruhstorfer // BMNH (E) 1718891 // SYNTYPE ♂ / Perisama diotima zurita ( Fruhstorfer, 1916 ) /Det. Zubek & Huertas, 2016. Syntype ♂ of Perisama diotima demata ( Fig. 2 J): SYN-/ TYPE // Type // Type // Macas/ Equador // diotima / demata Fruhst. // Fruhstorfer Coll./B.M. 1937-285 // BMNH (E) 1670028. Examined material. MZUJ : VENEZUELA : 1♂ : Mérida , La Culata N.P. , Sector El Playón , 2700 m , 01.II.2005 , T. Pyrcz leg. , prep. genit. 126/ A. Zubek , 22.07.2014 ; 1♂ : Táchira , Betania , El Tamá N.P. , 2300–2400 m , 13–15.IV.1996 , T. Pyrcz leg. ; 1♂ : same data but prep. genit. 86/ A. Zubek , 0 4.0 4.2013 . COLOMBIA : 1♂ : Cauca , Paramo Malvasá , 3200–3400 m , 17–23.II.1997 , T. Pyrcz leg. , prep. genit. 127/A. Zubek, 22.07.2014 ; 1♂ : Cauca , Paletara– San José de Isnos , km 55, 3000, 11.III.2013 , T. Pyrcz leg. ; 4♂ : Putumayo , Sibundoy via Mocoa , 2200 m , 23.I.2015 , C. Prieto , J. Lorenc & A. Zubek leg. ; 1♂ : Antioquia , Jardín , 1800 m , 28.VII.2002 , G. Rodriguez leg. ; 1♂ : Antioquia , Santa Rosa , 2700 m , 21.XII.2003 , prep. genit. 87/ A. Zubek , 04.IV.2013 ; 1♂ : Cauca , Munchique N. P. , La Romelia , 2600 m , 19.0 8.2007, T. Pyrcz leg. , prep. genit. 89/ A. Zubek , 04.IV.2013 ; ECUADOR : 1♀ : Azuay , Urchuloma , 26.X.1994 , F. Piñas leg. , prep. genit. 121/ J. Lorenc , 23.04.2014 .; 1♂ : Carchi , Golondrinas Forest Res. , 2600 m , 03.VII.1999 , J. Wojtusiak leg. , prep. genit. 90/A. Zubek, 04.04.2013 ; 1♂ : same data but 24.VI.1999 ; 1♂ : Carchi , El Playón , 2700 m , 01.II.2005 , T. Pyrcz leg. ; 1♀ : Cotopaxi , Pilaló , 3000 m , 03.IX.2004 , T. Pyrcz leg. ; 1♂ : Loja , West of Valladolid , 1600–2100 m , 23.IV.2007 , A. Jasiński leg. ; 4♂ : Morona– Santiago , Quebrada Tinguichaca , Guamote–Macas , 2600–2650 m , 23.I.2004 , R. Garlacz & T. Pyrcz ; 2♂ : Morona– Santiago , Gualaceo–Limon , Las Chacras , 2200–2250 m , 31.VIII.2003 , T. Pyrcz leg. ; 1♂: Morona–Santiago, Gualaceo–Limon, east slopes, 3100 m , 08.II.2002 ; J. Wojtusiak & T. Pyrcz leg. ; 1♀ : same data, prep. genit. 122/ J. Lorenc , 23.04.2014 ; 1♀ : same data but 2750 m , 21.VIII.2003 , prep. genit. 123/ J. Lorenc , 23.04.2014 ; 1♂ : Morona–Santiago , Sigsig via Chiguinda , 2900 m , 16.IX.2004 , J. Wojtusiak & T. Pyrcz leg. , prep. genit. 78/A. Zubek, 25.03.2013 ; 1♂ : Morona–Santiago , Tiguichaca , 2600 m , 23.I.2004 , T. Pyrcz leg. ; 1♂ : Napo , Rio Horituyacu , 1800 m , 08.VI.1999 , T. Pyrcz ; 1♂ : Napo , Rio Sucio , 1800 m , 16.VI.1999 , T. Pyrcz leg. ; 1♂ : Rio Sucio Near Pastaza , 1800 m , 16.VI.1999 , J. Wojtusiak & T. Pyrcz leg. ; 2♂ : Napo , Baeza–Tena , Res. Yanayacu, 2100– 2200 m , 07–09.IX.2003 , T. Pyrcz leg. ; 1♂ : Napo , Baeza , XII.1995 , prep. genit. 73/ A. Zubek , 10.09.2012 ; 1♂ : Sucumbíos , La Bonita , ex. coll. A. Jasiński ; 1♂ : same data but IX.1997 , I. Aldas leg. ; 1♂ : same data but II.1998 , prep. genit. 128/ A. Zubek , 22.07.2014 ; 1♂ : same data but 01.V.1998 , A. Jasiński leg. ; 1♂ : Sucumbíos , Quebrada Del Morro , 2100, 08.X.1990 , local dealers leg. ; 1♂ : Sucumbíos , El Playón , XI.1996 ; 1♂ : Tungurahua , Baños– El Tablón , 2600–2900 m , 07.VIII.1998 , J. Wojtusiak & T. Pyrcz leg. , prep. genit. 74/A. Zubek, 20.03.2013 ; 1♂ : Tungurahua , Runtun , 15.IX.1997 , A. Jasinski leg. ; 2♂ : Tungurahua , Chinchin , Sangay N.P. , 1600 m , 06.II.2004 , T. Pyrcz leg. ; 1♂ : Tungurahua , Chinchin , 04.II.1997 ; 1♂ : Tungurahua , Baños–Runtun , 2900 m , 22.I.2003 , J. Wojtusiak leg. ; 2♂ : Tungurahua , Arriba de Baños , El Tablón , 2600–2900 m , 07.VIII.1998 , T. Pyrcz leg. ; 1♂ : Tungurahua , Viscaya, IV .1997 , 1♂ : same data but 2150 m , 02.V.1996 , A. Jasiński leg. ; 4♂ : Tungurahua , Machay , 04.IV.1997 , ex. coll. A. Jasiński ; 3♂ : Tungurahua , Baños area , ex. coll. A. Jasiński ; 3♂ : Tungurahua , Baños , 27.XII.1999 ; 7♂ : Tungurahua , Rio Verde , 22–28.IV.1997 , ex. coll. A. Jasiński ; 1♂ : Tungurahua , Nanegalito , IV.1997 , A. Jasiński leg. ; 1♂ : Tungurahua , Leito Patate , 3300 m , 27.VI.2005 ; 1♂ : Zamora–Chinchipe , San Andres – Calderon , 2000–2600 m , 20.V.1998 , A. Jasiński leg. , prep. genit. 129/ A. Zubek , 22.07.2014 ; 3♂ : Zamora – Chinchipe , San Andres , 2000–2200 m , A. Jasiński leg. ; 1♂ : Zamora– Chinchipe , Cordillera Llagunillas , Rd. Padilla – San Andrés , 1500–2000 m , 19.V.1998 , A. Jasiński leg. ; 23♂: no data, ex coll. A. Jasiński; 1♂: no data.; RCCP : COLOMBIA : 2♂ : Cauca , Munchique , 2200 m , 28.IV.2000 , C. Prieto leg. ; 1♂ : Cauca , Munchique , La Romelia , 2760 m , 16.IX.2009 , C. Prieto leg. ; 1♂ : same data but 2300 m , 19.V.2007 ; 2♂ : Cauca , San Juan de Villalobos , 1800 m , 22.VI.2013 , C. Prieto leg. NHMUK : COLOMBIA : 3♂ : Bogota to Coachi , 2800 to 1700 m ., I. 1997 , dry season, Dr. Bürger ; 1♂ : Bogota , ex coll. Oberthür ; 1♂ : Bogota to Buenaventura, O. Thieme, 14. XII.77 – 22.II.78 ; 2♂ : Fusagasugá , 6000 ft ., Wheeler leg. ; 2♂ : Manizales ; 1♂ : Corinto , Cauca , May–July 1906 (Paine & Brinkley) ; 2♂ : Cauca , Tornel , nr. Tacueyo , 5500 ft ., end of Nov. 1906 (Paine & Brinkley) ; 2♂ : Cauca Valley, Torne , January 1907 (Paine & Brinkley) , 1♂ : same data but 01–15.II. 07 ; 5♂ : Cauca Valley, Torneé [sic], ex. coll. W. Feather ; 1♂: Bates coll.; 2♂ : Huila Dept., above San Jose de Isnos , Eastern slopes cent. Cordillera , 2600–2700 m , 6.VIII.1982 , M. Adams coll. ; 1♂ : Tolima Dept., S. above Cajamarca , 2950 m ., 26.VII.1979 , 1♂: Santa Rita, ex. coll. Feather; ECUADOR : 35♂ : Env. De Ambato R. P. , Irenee Blanc leg. , ex coll. Oberthür ; 3♂ : East , Baños , Rio Pastaza , 5000–7000 ft , M. G. Palmer leg. ; 1♂: Rio Verde, C. Buckley leg. ; 1♂ : same data but 5 0 0 0 ft, M. G. Palmer leg. ; 2♂ : East , Sarayacu, C. Buckley leg. ; 1♂: Macas, Bates coll.; 9♂: Zamora, 3000–4000 ft ., O. T. Baron leg. ; 1♂ : Rio Verde Grande , 5000 ft ., I.1912 , M. G. Falmer leg. ; 2♂ : Balsapamba ; 1♂ : no data, Fruhstorfer ; 1♂ : Yunguilla , Baños , 1800 m . 20.VIII.36 ; 1♂ : same data but V, 38–1800 m ; PERU : 2♂ : North , River Tabaconas , 6000 ft ., 1912, A.E. & F. Pratt. leg. 1 6 : Cauca , Munchique N. P. , La Romelia , 2600 m , T. Pyrcz leg. , 19.0 8.2007, prep. genit. 89/ A. Zubek , 04.04.2013 ; 4♂ : Antioquia , Frontino, T. K. Salmon ; 1♂ : Micay , August 1896 , ex. coll. Dognin ; PBF : ECUADOR : 1♂ : Azuay , Gualaceo–Limón , 3100, 09.V.1998 ; 1♀ : Cotopaxi , Latacunga vers Quevedo , 3200– 3400 m , 10.XII.1998 , P. Boyer leg. ; 1♂ : Imbabura , Route de Buenos Aires , 2600 m , 05.V.2000 , P. Boyer leg. ; 1♂ : Loja , Las Antenas , 3 km sud de Saraguro , 3000–3200 m , 10.II.2004 , P. Boyer leg. ; 1♀ : Morona–Santiago , Cebadas vers Macas via Atillo km 48, 3200–3400 m , 17.XI.2012 , P. Boyer leg. ; 1♂ : Morona–Santiago , Plan de Milagro , 2300 m , XII.2006 , P. Boyer leg. ; 1♂ : Morona–Santiago , Riobamba vers Macas via Atillo km 47–50, 3200 m , 21.V.2011 , P. Boyer leg. ; 3♂ : Pichincha , Nanegalito , 2000 m , IV.1999 , P. Boyer leg. ; 1♂ : Sucumbíos , La Bonita , 2000 m , III.1997 , P. Boyer leg. ; 1♂ : same data but XII.1996 ; 5♂ : Tungurahua , 3100 m , 29.XI.1998 , P. Boyer leg. ; 3♂ : Tungurahua , Rio Blanco , 2000 m , 23.V.1991 , P. Boyer leg. ; 1♂ : Tungurahua , Viscaya , 2000 m , XI.1996 , P. Boyer leg. ; 1♂ : Zamora–Chinchipe , Loja vers Zamora , 1800 m , 20.II.1996 , P. Boyer leg. ; 1♂ : same data but 02.VIII.1999 ; 1♂ : Valle de Cauca , Cali , no data, P. Boyer leg. ; 1♂ : same data but prep. genit. 86/ A. Zubek , 04.IV.2013 . EH: COLOMBIA : 1♂ : Caldas , Marulanda via a San Luis , 2900, 01.VI.2001 , E. Henao leg. ; 1♂ : Risaralda , Santa Rosa de Cobal Ecotermales , 17.IV.1999 , E. Henao leg. ANDES-E .: COLOMBIA : 1♂ : Cundinamarca , Páramo Guanachas , 3156 m , 23.IX.2011 , K. Martinez leg. MUSA : PERU : 1♂ : Piura , Huancabamba , Sapalache , Cerro Chinguela , 2800–3000, VII.2011 , H. Cardenas leg. , prep. genit. 146/A. Zubek, 22.0 7.2015. Additional material. MHN-UC: COLOMBIA : 1♂ : Cauca , San Sebastián–Páramo de las Papas , 3000 m , 26.III.1986 , J. Vélez leg. ; 1♂ : Huila , San Agustín , 15.VIII.1978 , E-Schmidt M. leg. ; 1♂ : same data but 18.III.1967 ; 1♂ : Nariño , Monopamba– El Socorro , 1500 m ., 26.II.1986 , J. Salazar leg. ; 2♂ : Putumayo , San Francisco –Río Putumayo , 2300 m , 31.XII.1985 , J. Vélez leg. ; 1♂ : Antioquia , Jardín–Ventanas , 2300 m , 17.IV.1987 , J. Vélez leg. ; 1♂ : Risaralda , Pueblo Rico–Río Negro , 1550 m , 18.IV.1984 , J. Vélez leg. ; 2♂ : Caldas : Manizales–Río Blanco , 2750 m , 28.I.1983 , J. Vélez leg. CSJ: COLOMBIA : 2♂ : Nariño , Monopamba– El Socorro , 1500 m , II.1986 , J. Salazar leg. Immature stages and biology. Unknown.