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Ocampo, David, Hanauer, Rachel E., Greeney, Harold F., Londoño, Gustavo A. (2020): First description of the nest, eggs, and nestlings of White-tufted Sunbeam (Aglaeactis castelnaudii) and incubation behaviours of Shining Sunbeam (A. cupripennis) in the southeast of Peru. Journal of Natural History 54 (23-24): 1455-1463, DOI: 10.1080/00222933.2020.1811415, URL: http://dx.doi.org/10.1080/00222933.2020.1811415

Daedalma inconspicua cuencana Pyrcz, n. ssp. (Figs. 2H, 12B) Material examined: ECUADOR: HOLOTYPE male: Azuay, Barabón, 24.V.1994, F. Piñas leg., red, rectangular label saying: Holotype, MZUJ; PARATYPES (5 males): 1 male: Azuay, Gualaceo, vía Plan de Milagro Km 16, 3300–3400 m, 22.XI.1998, P. Boyer leg., TWP (1 male to be deposited in MECN) 2 males: Ecuador, Azuay, Cabogana, 3000 m, 09.II.2004, P. Boyer leg., PBF; 1 male: Azuay, Girón, 02.X.1994, F. Piñas leg., FPQ; 1 male: Azuay, Girón, 79 o 08’42’’W, 03 o 09’24’’S, 2090 m, 02.X.1994, F. Piñas leg., FPQ. Additional material: 1 male: Azuay, Tarqui, 3000 m, 26.V.2007, S. Padrón leg., SPC. Description: MALE (Fig. 2H): Head, thorax and abdomen: not differing from nominate subspecies. Wings: FW length: 22–24 mm, mean: 22.75 mm, n=6. The subspecies differs from nominate and other races in its considerably smaller size; dorsal surface of wings grey-brown as in nominate, considerably lighter than in orientalis; FWV has no reddish median suffusion apparent in nominate; lighter, pale yellow patch in distal border of discal cell is disconnected from wide lighter chestnut postdiscal area in cells M3-Cu1 and Cu1-Cu2, contrary to nominate, variegata and orientalis; HWV ground colour is considerably lighter than in orientalis, which has a noticeable chocolate brown pattern, and similar to nominate. Genitalia (Fig. 12B): Uncus length of tegumen dorsum, stout, slightly arched; gnathos half length of uncus, stout basally, gradually narrowing to a sharp tip, similar to onorei and orientalis; saccus about same depth as palacio and tapichalaca; valvae about length of tegumen + uncus, slender, gradually narrowing from middle to apex, similar to onorei; aedeagus not differing from other subspecies. FEMALE: Unknown. Etymology: This subspecies is named after the Cuenca valley, where it occurs. Remarks: This subspecies, recognised immediately from other races of D. inconspicua by its smaller size, is apparently endemic to the environs of Cuenca, where it has been recorded on both east and west Andean slopes. It occurs in the uppermost forest, where it is most frequently observed while hilltopping in the company of Junea doraete and Steremnia species. A single male specimen collected by Sebastián Padrón in the hills southeast of Río Tarqui, just south of Cuenca, is tentatively associated with this taxon. This specimen differs from typical cuencana in having the pale markings of the FWV a pale orange colour, rather than pale grey. In addition, the pale dusting of scales at the distal end of the discal cell and subapex of the FWD seen in the figured cuencana appears in this specimen orange, rather than white., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on pages 35-36

Daedalma eliza Pyrcz & Willmott, n. sp. (Figs. 4C, 4D, 9A, 13D) [Daedalma adamsi d’Abrera; Salazar, 2004: 255 (misidentification).] Material examined: ECUADOR: HOLOTYPE male: Carchi, Las Golondrinas, 2400 m, 19.VI.1999, T. Pyrcz & J. Wojtusiak leg., red, rectangular label saying: Holotype, MZUJ; PARATYPES (19 males and 2 females), 1 male: same data as the holotype, TWP; 1 male: same data but 20.VI.1999, 2400–2500 m, BMNH; 1 male: same data but 22.VI.1999, 2200 m, PUCE; 1 male: same data but 2250 m, TWP; 1 male: same data but 2450 m, TWP; 2 males: same data but 01.VII.1999, 2250 m, TWP; 1 male: same data but 2350 m, TWP; 1 male: same data but 02.VII.1999, 2400 m, TWP; 1 male: same data but no altitude, 19.VI.1999, TWP; 4 males: same data but 2100 m, 27.xi.1996, K. Willmott leg., KWJH (1 to be deposited in MECN, 1 to be deposited in FLMNH); 2 males: Carchi, above Las Juntas, 1600–2000 m, VIII.2001, I. Aldaz leg., MBLI; 1 female: same data as the holotype 2450 m, TWP; 1 female: same data as the holotype but 15.IX.1997, K. Willmott leg., KWJH; COLOMBIA: 1 male: Nariño, Cumbitará, 23.IX.1996, G. Rodríguez leg., GRM. Additional material: 1 male: Pichincha, km 13 Nanegalito-Quito rd., Reserva El Pahuma, 2400 m, 26.X.1997, K. Willmott leg., KWJH. Diagnosis: This species and D. parvomaculata are both recognised from D. dinias by the small size of the FWD postmedian patch in males, orange in D. parvomaculata and dark crimson in D. eliza. In the females of D. dinias and D. eliza the FWD orange patch is wide and extends to the costa, whereas in D. parvomaculata it ends in mid-discal cell. The female of D. parvomaculata has no trace of orange or red on the HWD contrary to D. eliza and D. dinias, which have a well marked patch along the costa and at the apex, while in D. eliza some orange markings are also apparent in the median area (cell M2-M3). Daedalma eliza is also similar in appearance to the central Peruvian D. adamsi, whose FWD patch is the same colour as in D. eliza, but is situated more distally, and does not enter the discal cell. Description: MALE (Fig. 4C): Head: Eyes dark brown, covered with short and dense setae; palpi, twice length of head, light brown covered with chestnut hair; antennae 2/5 length of costa, slender, club only slightly thicker than shaft, chestnut ventrally, dark brown dorsally. Thorax: Dorsally and ventrally blackish, legs light brown. Abdomen: Dorsally and laterally blackish, ventrally lighter dull brown. Wings: FW (length: 27–30 mm; 28,16 mm; n=12) costa slightly arched, outer margin protruded at vein M1 and slightly wavy. HW costa protruded at apex, outer margin undulate with two tail-like extensions at veins Cu1 and Cu2. FWD blackish; a faint dark crimson oval patch with inner 1/3 third inside discal cell and outer 2/ 3 in cell M3-Cu1, slightly extending in some individuals into cell Cu1-Cu2. HWD uniform blackish. FWV blackish grey-brown; a lighter grey median patch in same position as upperside crimson patch but extending further across discal cell and reaching costal vein; a whitish subapical elongate patch with scaling and a dark brown spot distally; a marginal band in apical area composite of olive and shades of brown. HWV ground colour chocolate brown with a complex mosaic of silver, milky white and brown patches typical for genus; postdiscal milky white patch in cell M2-M2 triangular. Genitalia (Fig. 9A): Uncus stout, curved downwards in middle; gnathos thin, 2/3 length of uncus; saccus deep and widening in basal area; valvae elongate with a smooth ampulla, and nearly as wide at apex as in median part, with a small apical tip curved upwards; aedeagus straight and smooth, shorter than valva + saccus. FEMALE (Fig. 4D): Head: Eyes chocolate brown covered with dense, short setae; palpi twice length of head, pale yellow covered with sandy yellow hair; antennae 2/5 length of costa, slender, orangeish dorsally and ventrally, club only slightly thicker than shaft, slightly darker. Thorax: dorsally and ventrally medium brown, legs sandy yellow. Abdomen: dorsally and laterally medium brown, ventrally beige. Wings: FW (length: 32 mm) and HW shape same as in male. FWD ground colour dark brown, slightly lighter in basal area; an oblique, elongate orange patch across discal cell and vein Cu1, whitish along costal cell; three connected whitish subapical patches along costa, in M1-M2 and M2-M3; fringes alternately milky white and brown. HWD ground colour dark brown; a diffuse postdiscal patch extending from costa to vein Cu1, enclosing darker brown patches in M1-M2 and M2-M3; fringes yellowish in costal area, orange on “tails”. FWV colour pattern reflected from dorsal surface but lighter; subapical and apical area sandy yellow. HWV pattern typical for genus consisting of a complex mosaic of yellow, chestnut and chocolate brown elements; ground colour sandy yellow; darker, chocolate brown in median area and in cell M2-M3 along distal margin; magenta scaling along zigzagging brown median line. Genitalia (Fig. 13D): Sinus vaginalis small and shallow. V-shaped batten on lamella postvaginalis large, and heavily sclerotized. Hump in middle of lamella postvaginalis sharp in lateral view, pointing posteriorly. Heavy sclerotization of eighth segment limited to dorsal most part of tergum and to lateral slats connecting with sinus vaginalis. Posterior part of lamella postvaginalis slightly sclerotized and wrinkled. Ductus bursae with delicate parallel wrinkles, gradually narrowing toward posterior. Antrum very short, and bent, opens to sinus vaginalis at its very bottom. Colliculum absent. Papillae anales flattened, separated on ventral side by wide, slightly wrinkled wall of tuba analis. Strongly sclerotized cuticular slat connects eighth tergite with lamella postvaginalis. Apophyses posteriores reduced. Bursa copulatrix asymmetric, ductus bursae gradually narrowing towards ostium, with parallel wrinkles, bent at site where it meets with ductus seminalis. Two ribbon-like signa in middle of bursa slightly bent. Etymology: This species is dedicated to Miss María Eliza Manteca Oñate, the president of the Las Golondrinas Foundation, which established the reserve where most of the type specimens were collected. Remarks: Daedalma eliza is most closely related to D. parvomaculata. The two species share common features of the male genitalia, in particular a noticeably stout uncus. In D. parvomaculata, sexual dimorphism is slight, and the female barely differs from the male except for a larger and better marked FW orange patch, whereas D. eliza is strongly sexually dimorphic, as are D. boliviana or D. fraudata, and the female has a wide oblique reddish orange patch on the FW. Given that these differences in wing pattern are much more marked than others that separate other sympatric species (e.g. D. boliviana, D. fraudata and D. rubroreducta), we believe D. eliza is best treated as a distinct species. Daedalma eliza is endemic to the west Andean Cordillera of Ecuador (Carchi) and Colombia (Nariño). Its northern distributional limit appears to be the Río Patía valley, while specimens have been collected from as far south as Pichincha province in western Ecuador. Quantitative sampling with baited traps showed a narrow altitudinal range, 2250–2450 m, occasionally as low as 2100 m. Other data, indicating collections below 2000 m, are unreliable. It is not uncommon in its habitat, undisturbed mid-elevation montane forests, being encountered most often along ridge tops., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on pages 20-21

Daedalma drusilla krugeriana Pyrcz, n. ssp. (Figs. 7E, 7F, 16B) [Daedalma drusilla Hewitson; Pyrcz, 1999: 228] Material examined: COLOMBIA: HOLOTYPE female: Tolima, Albania, Central Cordillera, East slopes, 14.VIII.1916, 2800 m, E. Krüger leg., red, rectangular label saying: Holotype, MIIZ; ECUADOR: PARATYPE male: Carchi, Tulcán-Maldonado km 40–50, 2800–3200 m, 07.V.1999, P. Boyer leg., PBF. Description: MALE (Fig. 7E): Head, thorax and abdomen: not differing from nominate subspecies. Wings (FW length: 26 mm): FWD blackish brown, a shade lighter in distal one-third, as compared to uniform blackish brown in nominate. FWD uniform blackish brown, lustrous, same as in nominate. FWV ground colour blackish brown; patches in distal part of discal cell and submarginal area barely lighter than ground colour, same as in some specimens of nominate, somewhat variable in this respect. HWV with more prominent chestnut and chocolate brown pattern than in other subspecies, especially in subapical and marginal area; submarginal milky white patches slightly larger than in nominate, similar as in tinta; three subapical-apical ocelli in Sc-Rs, Rs-M1 and M1-M2 form an arch pointing distally at costa, similarly to tinta and contrary to the nominate whose ocelli form a straight line. Genitalia: Not examined. FEMALE (Fig. 7F): Head, thorax and abdomen: not differing from nominate subspecies. Wings: FW length: 27.5 mm; FWD dark brown in basal half, light brown along outer margin; an irregular yolk-yellow patch in distal 2/3 of discal cell connected through base of cell M3-Cu1 to a large lighter yellow patch covering most of wing surface from postmedian to submarginal area except for an incision of brown along vein M3, gradually turning into light brown in subapical area, enclosing a faint brownish spot in M1-M2; fringes alternately brown and yellow, dark brown at vein ends. HWD medium brown, slightly darker in basal one-third; traces of rich yellow forming faint submarginal streaks in cells M1-M2 and M2-M3; fringes yellow in apical area, turning orange towards apex, vein ends brown. FWV ground colour slightly lighter brown and duller than on upperside; yellow markings as on upperside except that discal cell patch is disconnected from yellow postmedian area; subapical and apical area dusted with milky white scales, and different shades of brown; two dark brown subapical ocelli in R5-M1 and M1- M2; a zigzagging subapical line gradually turning into a wider area of brown in M2-M3. HWV ground colour light brown with a nondescript pattern of brown, chestnut and milky white lines and bands, similar to other taxa, with a somewhat better marked dark brown median diffused band and a milky white roughly triangular submarginal patch contiguous to vein M3. Genitalia (Figs. 16B): This subspecies differs from nominate by significantly longer ductus bursae and narrowly bent central part of batten on lamella postvaginalis. Sinus vaginalis with narrow entrance, flattened anteriorly, with wrinkles on lateral and ventral walls. Antrum opens to sinus vaginalis posteriorly, not at its base. Ductus bursae connects with bursa copulatrix centrally. Central part of lamella postvaginalis widely concave. Coliculum well developed at posterior part of ductus bursae. Narrow central part of a batten on lamella postvaginalis well pronounced. Outer walls of papillae anales at base of reduced apophyses posteriores set off as well marked bumps. Cuticule around anus with pronounced wrinkles. Bursa copulatrix with two ribbon-like, parallel signa consisting of minute teeth. Remarks: This subspecies of D. drusilla is described based on a female, which bears some resemblance to the "dora type " form of the nominate subspecies, with its wide yellow markings on the FWD. However, contrary to that form and other females of D. drusilla, there is no trace of yellow on the HWD. The only known individual was collected by Edwin Krüger at Albania in Tolima on the eastern slopes of the Colombian Central Cordillera (Pyrcz, 1999). Available data indicate that Central and Eastern Cordillera populations of most species of Pronophilina, especially inhabiting high elevations, belong to separate subspecies (Adams, 1986), and Pyrcz & Rodríguez (2007) confirm that the valley of Magdalena is an important zoogeographic barrier. Given the distinctive wing pattern of the only known female and biogeographic data from other pronophilines, we feel justified in describing it here as a new subspecies. Curiously, Krüger (1924) lists a male of " D. dora " (= D. drusilla), collected in the area of Bogotá, and states that he did not collect any individual of D. drusilla, yet mentions no female of either. Daedalma drusilla is apparently a rare species in the central departments of the Central Cordillera, as it was not collected by Adams (1986) who sampled extensively in Tolima, and it was not listed as occurring in Tolima or Caldas by Vargas & Salazar (2004)., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on pages 30-31, {"references":["Adams, M. J. (1986) Pronophiline butterflies (Satyridae) of the three Andean Cordilleras of Colombia. Zoological Journal of the Linnean Society, 87, 235 - 320.","Pyrcz, T. W. & Rodriguez, G. (2007) Mariposas de la tribu Pronophilina en la Cordillera Occidental de Colombia. Shilap, 35, 455 - 489.","Kruger, E. (1924) Beitrage zur Kenntnis der columbischen Satyriden. Entomologische Rundschau, 41, 7, 9 - 10, 16, 19 - 20, 23 - 24, 27 - 28, 31 - 32, 35, 38 - 39, 41 - 42, 46 - 47.","Vargas, J. I. & Salazar, J. A. (2004) Mariposas Colombianas VII. Breve nota sobre las especies de los generos Junea Hemming 1964, Daedalma Hewitson 1858 y Thiemeia Weymer 1912 depositadas en algunas colecciones colombianas (Lep., Sat.). Boletin Cientifico Centro del Museo de Historia Natural de la Universidad de Caldas, 8, 247 - 259."]}

Daedalma eliza Pyrcz & Willmott, n. sp. (Figs. 4C, 4D, 9A, 13D) [Daedalma adamsi d’Abrera; Salazar, 2004: 255 (misidentification).] Material examined: ECUADOR: HOLOTYPE male: Carchi, Las Golondrinas, 2400 m, 19.VI.1999, T. Pyrcz & J. Wojtusiak leg., red, rectangular label saying: Holotype, MZUJ; PARATYPES (19 males and 2 females), 1 male: same data as the holotype, TWP; 1 male: same data but 20.VI.1999, 2400–2500 m, BMNH; 1 male: same data but 22.VI.1999, 2200 m, PUCE; 1 male: same data but 2250 m, TWP; 1 male: same data but 2450 m, TWP; 2 males: same data but 01.VII.1999, 2250 m, TWP; 1 male: same data but 2350 m, TWP; 1 male: same data but 02.VII.1999, 2400 m, TWP; 1 male: same data but no altitude, 19.VI.1999, TWP; 4 males: same data but 2100 m, 27.xi.1996, K. Willmott leg., KWJH (1 to be deposited in MECN, 1 to be deposited in FLMNH); 2 males: Carchi, above Las Juntas, 1600–2000 m, VIII.2001, I. Aldaz leg., MBLI; 1 female: same data as the holotype 2450 m, TWP; 1 female: same data as the holotype but 15.IX.1997, K. Willmott leg., KWJH; COLOMBIA: 1 male: Nariño, Cumbitará, 23.IX.1996, G. Rodríguez leg., GRM. Additional material: 1 male: Pichincha, km 13 Nanegalito-Quito rd., Reserva El Pahuma, 2400 m, 26.X.1997, K. Willmott leg., KWJH. Diagnosis: This species and D. parvomaculata are both recognised from D. dinias by the small size of the FWD postmedian patch in males, orange in D. parvomaculata and dark crimson in D. eliza. In the females of D. dinias and D. eliza the FWD orange patch is wide and extends to the costa, whereas in D. parvomaculata it ends in mid-discal cell. The female of D. parvomaculata has no trace of orange or red on the HWD contrary to D. eliza and D. dinias, which have a well marked patch along the costa and at the apex, while in D. eliza some orange markings are also apparent in the median area (cell M2-M3). Daedalma eliza is also similar in appearance to the central Peruvian D. adamsi, whose FWD patch is the same colour as in D. eliza, but is situated more distally, and does not enter the discal cell. Description: MALE (Fig. 4C): Head: Eyes dark brown, covered with short and dense setae; palpi, twice length of head, light brown covered with chestnut hair; antennae 2/5 length of costa, slender, club only slightly thicker than shaft, chestnut ventrally, dark brown dorsally. Thorax: Dorsally and ventrally blackish, legs light brown. Abdomen: Dorsally and laterally blackish, ventrally lighter dull brown. Wings: FW (length: 27–30 mm; 28,16 mm; n=12) costa slightly arched, outer margin protruded at vein M1 and slightly wavy. HW costa protruded at apex, outer margin undulate with two tail-like extensions at veins Cu1 and Cu2. FWD blackish; a faint dark crimson oval patch with inner 1/3 third inside discal cell and outer 2/ 3 in cell M3-Cu1, slightly extending in some individuals into cell Cu1-Cu2. HWD uniform blackish. FWV blackish grey-brown; a lighter grey median patch in same position as upperside crimson patch but extending further across discal cell and reaching costal vein; a whitish subapical elongate patch with scaling and a dark brown spot distally; a marginal band in apical area composite of olive and shades of brown. HWV ground colour chocolate brown with a complex mosaic of silver, milky white and brown patches typical for genus; postdiscal milky white patch in cell M2-M2 triangular. Genitalia (Fig. 9A): Uncus stout, curved downwards in middle; gnathos thin, 2/3 length of uncus; saccus deep and widening in basal area; valvae elongate with a smooth ampulla, and nearly as wide at apex as in median part, with a small apical tip curved upwards; aedeagus straight and smooth, shorter than valva + saccus. FEMALE (Fig. 4D): Head: Eyes chocolate brown covered with dense, short setae; palpi twice length of head, pale yellow covered with sandy yellow hair; antennae 2/5 length of costa, slender, orangeish dorsally and ventrally, club only slightly thicker than shaft, slightly darker. Thorax: dorsally and ventrally medium brown, legs sandy yellow. Abdomen: dorsally and laterally medium brown, ventrally beige. Wings: FW (length: 32 mm) and HW shape same as in male. FWD ground colour dark brown, slightly lighter in basal area; an oblique, elongate orange patch across discal cell and vein Cu1, whitish along costal cell; three connected whitish subapical patches along costa, in M1-M2 and M2-M3; fringes alternately milky white and brown. HWD ground colour dark brown; a diffuse postdiscal patch extending from costa to vein Cu1, enclosing darker brown patches in M1-M2 and M2-M3; fringes yellowish in costal area, orange on “tails”. FWV colour pattern reflected from dorsal surface but lighter; subapical and apical area sandy yellow. HWV pattern typical for genus consisting of a complex mosaic of yellow, chestnut and chocolate brown elements; ground colour sandy yellow; darker, chocolate brown in median area and in cell M2-M3 along distal margin; magenta scaling along zigzagging brown median line. Genitalia (Fig. 13D): Sinus vaginalis small and shallow. V-shaped batten on lamella postvaginalis large, and heavily sclerotized. Hump in middle of lamella postvaginalis sharp in lateral view, pointing posteriorly. Heavy sclerotization of eighth segment limited to dorsal most part of tergum and to lateral slats connecting with sinus vaginalis. Posterior part of lamella postvaginalis slightly sclerotized and wrinkled. Ductus bursae with delicate parallel wrinkles, gradually narrowing toward posterior. Antrum very short, and bent, opens to sinus vaginalis at its very bottom. Colliculum absent. Papillae anales flattened, separated on ventral side by wide, slightly wrinkled wall of tuba analis. Strongly sclerotized cuticular slat connects eighth tergite with lamella postvaginalis. Apophyses posteriores reduced. Bursa copulatrix asymmetric, ductus bursae gradually narrowing towards ostium, with parallel wrinkles, bent at site where it meets with ductus seminalis. Two ribbon-like signa in middle of bursa slightly bent. Etymology: This species is dedicated to Miss María Eliza Manteca Oñate, the president of the Las Golondrinas Foundation, which established the reserve where most of the type specimens were collected. Remarks: Daedalma eliza is most closely related to D. parvomaculata. The two species share common features of the male genitalia, in particular a noticeably stout uncus. In D. parvomaculata, sexual dimorphism is slight, and the female barely differs from the male except for a larger and better marked FW orange patch, whereas D. eliza is strongly sexually dimorphic, as are D. boliviana or D. fraudata, and the female has a wide oblique reddish orange patch on the FW. Given that these differences in wing pattern are much more marked than others that separate other sympatric species (e.g. D. boliviana, D. fraudata and D. rubroreducta), we believe D. eliza is best treated as a distinct species. Daedalma eliza is endemic to the west Andean Cordillera of Ecuador (Carchi) and Colombia (Nariño). Its northern distributional limit appears to be the Río Patía valley, while specimens have been collected from as far south as Pichincha province in western Ecuador. Quantitative sampling with baited traps showed a narrow altitudinal range, 2250–2450 m, occasionally as low as 2100 m. Other data, indicating collections below 2000 m, are unreliable. It is not uncommon in its habitat, undisturbed mid-elevation montane forests, being encountered most often along ridge tops.

Daedalma inconspicua inconspicua Butler, stat. rev. (Figs. 5C, 5D, 11E, 16D) Material examined: ECUADOR: 1 male: white, square label: Quito, coll. By M. Bourcier, 50-111, white square label: B.M. Type, No. Rh. 70-64, red and white rounded label: Daedalma inconspicua Butler, type, [LECTOTYPE of Daedalma inconspicua Butler, herein designated], BMNH; 1 male: Pichincha, Quito vers Chiriboga km 12–15, 3300–3400 m, 01.II.2004, P. Boyer leg., PBF; 1 male: Pichincha, Chillogallo vía a Chiriboga, 3000 m, 10.I.1998, I. Aldaz leg., PBF; 1 male: Pichincha, Volcán Pichincha, San Juan-La Victoria, 3300–3350 m, II.2002, I. Aldaz leg., MZUJ; 2 males: Pichincha, Quito-La Victoria rd., Loma La Palmira, 3300–3500 m, 25.VIII.1999, K. Willmott leg., KWJH; 1 male: Pichincha, Volcán Pichincha, Yanacocha, 3500 m, 18.IX.1997, K. Willmott leg., KWJH; 1 male: Pichincha, nr. Amaguaña, Volcán Pasochoa, 3500 m, 7.X.1997, K. Willmott leg., KWJH; 1 female: Ecuador, Pichincha, Chillogallo-San Juan, vía La Victoria, 3300–3350 m, 05.II.2002, T. Pyrcz leg., MZUJ; 1 male: Pichincha, Pasochoa, 3200 m, no, 9078, J-C. Petit leg, JCP. Redescription: MALE (Fig. 5C): Head: chocolate brown; labial palpi chestnut covered with dark brown hair; antennae dorsally dark brown, ventrally lighter, chestnut brown, club dark brown. Thorax: black; legs chestnut. Abdomen: dorsally and laterally black, ventrally chestnut. Wings: FW length: 24–27 mm, mean: 25.3 mm, n=4; FW and HWD medium greyish brown, same as in blancae n. ssp., slightly lighter than in orientalis n. ssp. and tapichalaca n. ssp.; FWV lighter area in cells M3-Cu1 and Cu1-Cu2, variably or without any reddish suffusion, similar to variegata n. ssp. and blancae but in contrast to onorei n. ssp.; FWV lighter area in cells M3-Cu1 and Cu1-Cu2 widely connected to similarly coloured bar in discal cell, similar to orientalis, in contrast to nominate, onorei and tapichalaca, in which discal cell pale marking is small and restricted to distal edge of discal cell. HWV ground colour varying between chocolate brown and chestnut. Genitalia (Fig. 11E): Uncus slightly shorter than in nominate; gnathos similar to other subspecies; saccus similar in length and width to nominate; valvae slightly thinner than in blancae, with a slightly irregular dorsal surface; aedeagus not differing from other subspecies. FEMALE (Fig. 5D): FW length: 32 mm; with faint, light reddish orange markings on FWD, in contrast to orientalis, which is only patterned in lighter and darker shades of grey-brown, and to variegata, which has extensive and intense light orange patches in distal half (females of onorei and blancae unknown). Genitalia (Fig. 16D): Lamella postvaginalis large, concave in middle. Bottom part of sinus vaginalis delicately ribbed. V-shaped batten on lamella postvaginalis sharply bent in middle. Additional, smaller group of setae at ventral side of papille anales complementing main group of setae. Sinus vaginalis shallow and wide. Antrum long, slightly shorter that ductus bursae. Combined length of antrum and ductus bursae same as length of bursa copulatrix. This subspecies differs from other subspecies by the lack of colliculum, very shallow sinus vaginalis and by having ductus bursae and antrum of same length. In all other subspecies antrum is much shorter than ductus bursae. Sclerotized lateral parts of eighth segment that connect tergum with sinus vaginalis narrow. Remarks: The identity of the nominate population of D. inconspicua is crucial for the systematics of this highly polytypic species. The type locality specified in the original description is “Quito, Chimborasso”. “Quito” often referred, in the 19 th century, to any Ecuadorian locality (as did “Bogotá” for any Colombian locality), not necessarily the capital city situated in the inter-Andean valley in the north of the country. In our experience, however, most specimens labelled “Quito” come from the western slopes of the Andes in Pichincha, central Ecuador. The locality “Chimborasso”, however, suggests the slopes of Volcán Chimborazo, or, more broadly, the province of Bolívar, further south. An examination of the syntype specimen, simply labelled “Quito,” shows that it corresponds with individuals known from both these regions, such as specimens collected along the Guaranda-Balzapamba road, south-west of Chimborazo, or those from the south-western slopes of Volcán Pichincha in the Quito area. Unfortunately, these localities are apparently inhabited by two separate subspecies of D. inconspicua, differing slightly in colour patterns but distinguished mostly by biogeographical factors, namely the presence of a third subspecies in between their respective ranges, onorei n. ssp. Although it is impossible to unequivocally associate the syntype with either population, the original label stating “Quito”, and the fact that the syntype's collector, M. Bourcier, lived in Quito and would therefore have had greater access to its environs than to regions further south, suggest the name should be applied to the Pichincha population, which is considered herein as topotypical., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on pages 31-32

Daedalma inconspicua onorei Pyrcz, n. ssp. (Figs. 5B, 11D) Material examined: ECUADOR: HOLOTYPE male: Cotopaxi, Pilaló, Carretera Nueva, 3000–3050 m, 03.IX.2004, T. Pyrcz leg., red, rectangular label saying: Holotype, MZUJ. Description: MALE (Fig. 5B): Head, thorax and abdomen: not differing from nominate subspecies. Wings: FW length: 28 mm; HW costal lobe more prominent than in nominate and other subspecies; FWD and HWD dark greyish brown, same as in orientalis, lighter than in other west Andean subspecies; FWV lighter postdiscal area in M3-Cu1, Cu1-Cu2 connected to a small lighter discal cell patch by a narrow neck, similar to nominate and cuencana n.ssp. a faint reddish suffusion in median area along discal cell vein, somewhat reminiscent of nominate subspecies; HWV ground colour distinctively olive green, differing from other subspecies, with a wide, rather uniform median area, devoid of any pattern of black or brown lines apparent in other subspecies. Genitalia (Fig. 11D): Uncus gently arched, about the length of the dorsum of tegumen; gnathos half the length of uncus; saccus wide, shallower than in other subspecies except in orientalis; valvae the length of tegumen + uncus, ampulla smooth, a prominent apical spine-like process; aedeagus not differing from other subspecies. FEMALE: Unknown. Etymology: This subspecies is dedicated to Giovanni Onore, an Italian entomologist and professor of entomology and director of the Museo de Entomología for many years at the Universidad Católica in Quito, a great promoter of conservation of natural Andean environments, and a friend of the authors. Remarks: Daedalma inonspicua onorei is known so far only from the type specimen collected above Pilaló. Despite that, in our opinion it is justified to consider this specimen as representing a separate subspecies in view of apparently important differences in the wing colour pattern, especially the highly distinctive olive green HWV, not found in any other subspecies., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on page 34

Daedalma inconspicua petitia Pyrcz, n. ssp. (Figs. 6H, 12E) Material examined: ECUADOR: HOLOTYPE male: Loja, Cord. de Lagunillas, 3000 m, XI.2000, I. Aldaz leg., red, rectangular label saying: Holotype, MZUJ; PARATYPES (3 males): 1 male: Loja, Cord. de Lagunillas, 3000 m, XI.2000, I. Aldaz leg., MBLI; PERU: 1 male: Cajamarca, Santuario Natural Tabaconas-Namballe, Lagunas Arrebiatadas, S 05��14��05" W 79��17��18", 3122 m, 03.X.2009 E. Huamani leg., MHN-SA; 1 male: Cajamarca, Sanctuario Nacional Tabaconas-Namballe, Sector Miraflores, S 05��10��02" W 79��12��06", 2983 m, 19.X.2009 E. Huamani leg., MHN-SA. Description: MALE (Figs. 6H): Head, thorax and abdomen: not differing from nominate subspecies. Wings: FW length: 25���26 mm, n=4, mean: 25.8 mm, n=2. FWD postdiscal band dark orange with a brown suffusion along a diffused distal edge, compared to milky white in palacio with a sharp distal edge. HWD with a faint, irregular postdiscal dark orange line, milky white in nominate. FWV postdiscal band light orange, milky white in nominate, otherwise similar. HWV not differing noticeably from other subspecies. Genitalia (Fig: 12E): Uncus slightly longer than in tapichalaca, similar in length and shape to palacio; gnathos approximately half length of uncus, thinner than in palacio but longer than in tapichalaca; saccus slightly longer than in other subspecies (however, length of saccus is generally subject to noticeable individual variation); valvae elongated, narrowing gradually towards apex, most similar to onorei, apical hook prominent; aedeagus not differing from other subspecies. FEMALE: Unknown. Etymology: This subspecies is dedicated to Jean-Claude Petit, a French lepidopterist, in recognition of his efforts in studying the butterflies of Sangay National Park in Ecuador. Remarks: Daedalma inconspicua petitia is so far known exclusively from the type locality, the Lagunillas massif, on the Ecuador-Peru border. It is replaced in the Nudo de Sabanillas portion of Podocarpus National Park by ssp. tapichalaca. Two individuals were collected while hilltopping on higher bushes at the edge of p��ramo (I. Aldaz, pers. comm.). Among other species of Pronophilina occurring in the same habitat are Pedaliodes arturi Pyrcz & Viloria, Lasiophila palades limes Pyrcz, Lymanopoda hazelana summa Pyrcz, Willmott & Hall, and Steremnia umbracina misella Thieme., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on pages 38-39

Daedalma dinias leticia Pyrcz, n. ssp. (Figs. 1C, 8C) Material examined: COLOMBIA: HOLOTYPE male: Depto. de Cauca, between Leticia & Puracé, E. slopes, Cent. Cordillera, 2650 m, 10.VIII.1979, red, rectangular label saying: Holotype, BMNH; PARATYPES (2 males): 1 male: same data as the holotype but 2650–2700 m, BMNH; 1 male: same data but 11.VIII.1979, BMNH. Description: MALE (Fig. 1C): Head, thorax and abdomen: not differing from nominate subspecies. Wings: FW (length: 27–28 mm, mean: 27,3 mm, n=3). FWD ground colour blackish brown; oblique patch medium orange with a light reddish shade, same colour as in salmoni, but shorter, not reaching costal vein, ending in mid-discal cell. HWD uniform blackish brown. FWV median patch orange, slightly lighter than on dorsal surface with a diffused upper edge. HWV not differing noticeably from nominate subspecies or salmoni. Genitalia (Fig. 8C): Uncus slightly shorter than in other subspecies; saccus extremely long, longer than in other subspecies, even though long saccus is a distinctive character of D. dinias; valvae with a prominent apical hook curved upwards similarly to salmoni and nominate, while same structure points distally in emma; otherwise similar to other subspecies. FEMALE: So far unknown. Etymology: This subspecies is named after the town of Leticia, near the type locality. Remarks: D. dinias leticia is so far known only from the types collected by Michael Adams on the eastern slopes of the Puracé volcano massif, in the upper valley of the Río Magdalena. Elevational data indicate similar ecological preferences to salmoni. Its range likely does not extend much farther north, as in the Tolima range it is replaced by salmoni. So far, no specimens of D. dinias have been collected in the Huíla massif. The southern distributional limits remain to be determined, given the wide geographic gap between this and D. dinias emma. A more thorough sampling in the area east of Pasto and Tulcán would be most informative., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on pages 11-12

Daedalma dinias romeliana Pyrcz, n. ssp. (Figs. 1D, 8D) Material examined: COLOMBIA: HOLOTYPE male: Cauca, Parque Nacional Munchique, La Romelia vía Pico Santa Rosa; 2800 m, 18.VIII.2007, C. Prieto leg., red, rectangular label saying: Holotype, MZUJ (to be deposited in MHN-UNC). PARATYPES (5 males): 1 male: same data as the holotype, TWP; 1 male: same data, CPC; 2 males: Cauca, P. N. Munchique, La Romelia, 2760 m, 11.VI.2006, C. Prieto leg.; 1 male: same data but 17.IX.2005, CPC. Additional material: 1 male: Valle, P. N. Farallones, Pico de Loro, 2800 m, 02.VII.2006, C. Prieto & C. Muñoz leg., CPC. Description: MALE (Fig. 1D): Head, thorax and abdomen: not differing from nominate subspecies. Wings: FW length: 26–27 mm, mean: 26.5 mm, n=2. FWD ground colour blackish-brown; oblique patch orange, similar to salmoni but slightly larger, particularly wider in middle, extending to base of vein Cu2 where forming a short tip. HWD uniform blackish brown. FWV median patch orange, lighter than on dorsal surface, but similar in shape. HWV not differing noticeably from salmoni. Genitalia (Fig. 8D): Tegumen stout; uncus stout and about length of tegumen; saccus long but shorter than in other subspecies except nominate; a prominent apical tooth on valvae curved apically, contrary to other subspecies, in which it is pointing upwards. FEMALE: Unknown. Etymology: This subspecies is named after the type locality, La Romelia in Munchique National Park. Remarks: This subspecies is only known to occur in the southern part of the Colombian Western Cordillera. It has been collected in the department of Cauca, where it is found at the highest elevations (above 2600 m) of Munchique National Park (Prieto, 2003). Specimens from further north, mentioned by Pyrcz & Rodríguez (2007), are referable to salmoni but were not included in the type series. Daedalma dinias romeliana differs very slightly in facies from salmoni, mostly in the slightly wider FW orange patch. However, the Munchique and Medellín-Caldas populations cannot be considered as belonging to the same subspecies, as they are separated geographically by leticia, which is characterised by the noticeably smaller orange FWD patch., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on page 12, {"references":["Prieto, C. H. (2003) Satirinos (Lepidoptera, Nymphalidae, Satyrinae) del Parque Nacional Natural Munchique. Diversidad de especies y distribucion altitudinal. Revista Colombiana de Entomologia, 29 (2), 203 - 210.","Pyrcz, T. W. & Rodriguez, G. (2007) Mariposas de la tribu Pronophilina en la Cordillera Occidental de Colombia. Shilap, 35, 455 - 489."]}

Daedalma dinias emma Pyrcz & Greeney, n. ssp. (Figs. 1G, 1H, 8F, 13C) Material examined: ECUADOR: HOLOTYPE male: Napo, Cosanga, Estación Científica Yanayacu, 2100 m, 03.I.2004, T. Pyrcz leg., red, rectangular label saying: Holotype, MZUJ; PARATYPES (19 males and 5 females): 2 males: Napo, Cuyuja, 2300 m, 02.II.2004, P. Boyer leg. PBF; 2 males: Napo, Cerro Condijua nr. Baeza, 2500 m, 16.III.1995, K. Willmott & J. Hall leg., KWJH; 2 males: same data as preceding, FLMNH; 2 males: Napo, nr. Cosanga, Estación Científica Yanayacu, 2000 m, 24.XI.2006, K. Willmott leg., FLMNH; 1 male: Napo, Cord. Huacamayos, La Virgen, 2250 m, 12.X.1996, K. Willmott leg., KWJH; 1 female: Napo, Yanayacu, 2200 m, 19.IX.2004, T. Pyrcz leg., TWP; 1 male: Napo, Baeza, 2200 m, 22.XII.2000, F. Piñas leg., FPQ; 1 male: Napo, Quijos valley, Yanayacu Station, 2200 m, voucher n o 99-988, time: 11.00, 23.XI.1999, H. Greeney leg., FPQ; 1 male: Hacienda San Isidro, 08.XII.1996, 3:30PM, bamboo forest, sunny, T. Walla leg., n o. 1254, FPQ; 1 female: Napo, Yanayacu, 2100 m, 30.XI.2002, no. 8373, J-C. Petit leg., JCP; 1 male: Yanayacu Biological Station, 26.X.2007, H07-1336, MZUJ; 1 male: Yanayacu Biological Station, 24.X.2007, H07-1335, MZUJ; 1 male: Yanayacu Biological Station, 23.X.2007, H07-1332, MZUJ; 1 male: Yanayacu Biological Station, 25.X.2007, H07-1331, MZUJ; 1 male: Yanayacu Biological Station, 24.X.2007, H07-1340, MZUJ; 1 male: Yanayacu Biological Station, 25.X.2007, H07-1338, MZUJ; 1 female: Yanayacu Biological Station, 23.X.2007, H07-1337, MZUJ; 1 female: Yanayacu Biological Station, 29.X.2007, H07-1334, MZUJ; 1 female: Yanayacu Biological Station, 23.X.2007, H07-1339 (eclosed), MZUJ. Description: MALE (Fig. 1G): Head, thorax and abdomen: not differing from nominate subspecies. Wings: FW length: 23–25 mm, mean: 24 mm, n=7. FWD ground colour blackish brown; reddish orange patch much smaller than in nominate subspecies, romeliana and salmoni, oval, entering discal cell and touching vein Cu2; HWD ground colour blackish brown, same as in other subspecies. FWV median brick red patch narrower than on the upperside, faint and overcast with a dark brown, in some specimens heavy, suffusion, similarily to D. rubroreducta. HWV ground colour chocolate brown, darker than in salmoni or leticia. Genitalia (Fig. 8F): Uncus and gnathos stout, broader than in granadillas n. ssp. and leticia, particularly gnathos; saccus same length as in granadillas and romeliana, shorter than in leticia; dorsal surface of valvae towards apex slightly serrate, apical process blunt, compared to pointed process in other subspecies; aedeagus similar to most other subspecies but slightly shorter than in salmoni and nominate. FEMALE (Fig. 1H): FW length: 25–27 mm, mean: 26 mm, n=4. FWD and FWV orange patch slightly smaller, otherwise similar to other subspecies. Genitalia (Fig. 13C): Differs from the nominal subspecies by the shape of Vshaped batten on lamella postvaginalis that is more sharply bent in the middle. Anterior wall of sinus vaginalislamella antevaginalis without parallel wrinkles. Ductus bursae much shorter than the length of the bursa. Bursa copulatrix larger than in nominate subspecies. Sinus vaginalis slightly wider than in the nominal form. Early stages: Hostplant: Chusquea cf. scandens, Poaceae (2100 m, Yanayacu, Napo, Ecuador). EGGS-FOURTH INSTAR: Data not available. FIFTH INSTAR (Fig. 20A): Overall form similar to D. rubroreducta (Figs. 19A, 19B). Larval patterning similarly complex, but dark brown areas reduced, especially on lateral areas of abdomen where the dark patch in dinias is generally square rather than triangular as in D. rubroreducta. Daedalma dinias also has more extensive green markings on the body, especially dorsally on the thorax and subdorsally on the terminal abdominal segments. These give the larvae an overall “mossier” appearance. PUPA (Figs. 20B, 20C): Pupa similarly complexly sculptured as in D. rubroreducta (Figs. 19C, 19D). Most notable differences in shape are form of the head, thoracic, and abdominal projections. Head projections of dinias are slightly longer and flatter, more widely separated, curve slightly upwards, and are more strongly notched on inner margin. Thoracic projection of dinias more flattened, expanded near apex giving a clubbed appearance from side, and posterior edge of projection bears a small protrusion. Pair of abdominal projections of dinias also more flattened, projecting slightly backwards, and distinctly three-lobed. Both species show some variation in extent of metallic green patterning but overall green markings are reduced in dinias. Most notably green areas are reduced on all projections and angled green line traversing wing pads does not reach anterio-dorsal margin of wing pad. Etymology: This subspecies is dedicated to Emma Espinoza Carabali, from Quito. Remarks: Daedalma dinias emma is found on the eastern slopes of the Andes in Ecuador (Tungurahua to Napo). It is replaced further south, in Morona-Santiago, by D. d. granadillas n. ssp., described below. This uncommon subspecies is found at slightly lower elevations than its Colombian Central Cordilleran allies, generally at 2100–2200 m, where it occurs in perhumid areas of cloud forests., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on pages 12-13

Daedalma inconspicua palacio Dognin, n. stat. (Figs. 6A, 6B, 6D, 12D, 17B) Daedalma palacio Dognin, 1891: 125. Daedalma palacio Dognin; Thieme, 1906: 137; Weymer, 1912: 266; d’Abrera, 1988: 844, 845 (male, dorsal, ventral); Racheli & Racheli, 2001: 331. Material examined: ECUADOR: 2 males: Loja, Saraguro, vía Las Antenas, 3100–3200 m, 15.IX.2004, T. Pyrcz leg.; 2 males: same locality, 2900–2950 m, 02.II.2004, T. Pyrcz leg. TWP; 2 males: same locality, 1997, I. Aldaz leg., TWP; 1 male: Loja (dept.), XII.1997, I. Aldaz leg., TWP; 3 males: Loja, Loja-Cuenca Km 27, 2800 m, 26.XI.1998, P. Boyer leg., TWP; 1 male: Loja, Zenén, 2800–2850 m, IV.2002, I. Aldaz leg., TWP; 1 male: Loja, Zenén, 2900 m, XI.2000, I. Aldaz leg., MBLI; 4 males: Loja, km 27 Loja-Cuenca rd., Cerro Palma [=Cerro Viña Mano], 2800–2950 m, 7.IV.1995, K. Willmott leg., KWJH; 1 female: Loja, Loja-Cuenca Km 27, 3000 m, 3.XII.2002, P. Boyer leg., PBF; 1 female: Loja, Les Antennes, 3 km sud de Saraguro, 3000–3200 m, 10.II.2004, P. Boyer leg., PBF; 1 male: Ecuador, Loja, Lagunillas, 3200 m, V.1998, A. Jasi ṅski leg., TWP; 3 males: Loja, old road Loja-Zamora, 2500 m, XI.2000, I. Aldaz leg., ex MBLI, TWP; 1 male: Loja, Loja, road Loja-Zamora, 2500– 2700 m, II.1999, I. Aldaz leg., ex MBLI, TWP; 2 males: Loja, old road Loja-Zamora, 2600 m, XI.1999, I. Aldaz leg., ex MBLI, TWP; 1 male: environs de Loja, 1890, collection W. Schaus, USNM; 1 male: Loja, El Monje près Loja, 1893, 32.21, ex coll. Dognin, 1921, BMNH; 1 male: Loja, Jimbura-vía Laguna Negra, 3000–3200 m, 15.V.1998, A. Jasi ṅski leg., TWP. Redescription: MALE (Figs. 6A, 6B): Head, thorax and abdomen: not differing from nominate subspecies. FW length: 23–26 mm, mean: 24.3 mm, n=12; FWD dark greyish brown with a milky white oblique postmedian band extending from costa to tornus, considerably variable in size and shape, in some individuals approximately 2– 3 mm wide, in other nearly twice as wide; faint milky white subapical patches, in some individuals barely noticeable. HWD brown, a shade darker than on FW; some individuals with two faint, dirty yellow postdiscal streaks, suffused with brown, accompanied by a third in cell M2-M3. FWV ground colour lighter than on upperside; milky white band extending slightly further distally between veins Cu1 and 1A, enclosing a brown submarginal dot in Cu1-Cu2; subapical and apical area mottled with chocolate brown, whitish and chestnut; a zigzagging black subapical line; two blackish brown subapical ocelli in R5-M1 and M1-M2. HWV predominantly chocolate brown with a nondescript pattern of blackish brown, chestnut, magenta and silver, similar to other subspecies. Genitalia (Fig. 12D): Uncus stout, hooked downwards; gnathos less than half length of uncus, basally stout, and with distal half thin; saccus deep and wide, longer than uncus; valvae elongate, gradually thinner towards apical part, ampulla very slightly undulated, apical hook prominent; aedeagus marginally longer than valvae, slightly arched in middle. FEMALE (Fig. 6D): Head, thorax and abdomen: not differing from other subspecies. FW length: 28 mm; FWD dark brown, a shade darker than in male; a wide oblique yolk-yellow postmedian band reaching from costa to tornus and anal margin, extending further distally than in male between veins Cu1 and 1A; two yolk-yellow small, elongated postdiscal patches. HWD dark brown, lighter brown towards distal and anal margin; a yolk-yellow band in postmedian to submarginal area gradually overcast with brown from apex, fading away before reaching vein Cu2, enclosing a series of dark brown oval or rounded dots well marked between apex and vein M2, others faint and merging with background colour; fringes yellow and dark brown along vein ends. FWV patterned as on the upperside, except that subapical and apical area olive green with two, milky white patches and two small blackish ocelli with white pupils in R5-M1 and M1-M2. HWV patterned as in male but all white or silver elements replaced with sandy yellow, whereas brown elements with olive green. Genitalia (Fig. 17B): Sinus vaginalis small and rounded; ductus bursae straight pointing dorso-ventrally; antrum short, its wall at bottom wrinkled; coliculum well pronounced, concave at area where ductus seminalis leaves ductus bursae; lateral parts of eighth segment large and wide; lamella postvaginalis wide, grooved at centre; left and right arms of V-shaped batten on lamella postvaginalis twisted and rounded in central part; papillae anales with bumps on their outer surface opposite to base of rudimentary apophyses posteriores; strongly developed setae on edges of anterior part of papillae analis pointing toward themselves; ductus bursae smaller than length of bursa; bursa copulatrix with two ribbon-like, parallel signa consisting of minute teeth. Female genitalia differs from those of D. inconspicua variegata primarily by bent ductus bursae so that bursa copulatrix points ventrally. Remarks: This taxon was originally described as a separate species and, indeed, the FWD pattern characterised by the wide median pale band is markedly different from nominate D. inconspicua. However, similiarities in the HWV colour pattern, wing shape, and male genitalia, coupled with ecological similarities and allopatric distribution, strongly favour considering palacio as yet another subspecies of the highly polytypic D. inconspicua. The subspecies palacio occurs in uppermost cloud forest, locally quite common at 2800–3200 m. Males can be observed in the late morning perching on higher trees, and occasionally keeping territories where they may fight with conspecifics. Females are only rarely seen in the field. Daedalma i. palacio is the western subspecies of D. inconspicua in south-western Ecuador, and typical individuals occur from Saraguro to Jimbura. In at least two areas palacio possibly locally hybridizes with the eastern subspecies, as individuals presenting intermediate phenotypes were found. It is relatively easy to detect a possible hybrid between palacio and tapichalaca n. ssp. or petitia n. ssp. because of their sharply different FWD colour patterns, marked by the presence of a wide whitish band in palacio. An individual collected in El Monje (south of Loja) (Fig. 6C) does not have a complete FW band but a series of disconnected pale yellow patches heavily suffused with brown. It is phenotypically intermediate between tapichalaca n. ssp. and palacio, and until additional specimens are available for study it is considered as a subspecific hybrid. Further south, one specimen collected in the Cordillera de Lagunillas (above Jimbura) (Fig. 6G) has a faint, dirty orange FW band and is phenotypically intermediate between palacio and petitia., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on pages 36-37, {"references":["Weymer, G. (1912) Satyridae [publ. 1910 - 1912]. in A. Seitz (ed.), Die Gross-Schmetterlinge der Erde, 2; Exotische Fauna, 5, 173 - 283.","D'Abrera, B. (1988) Butterflies of the Neotropical Region, part V, Nymphalidae (Concl.) & Satyridae. pp. 680 - 887, colour plates throughout. Hill House. Victoria, Australia.","Racheli, T. & Racheli, L. (2001) An annotated list of Ecuadorian butterflies (Lepidoptera: Papilionidae, Pieridae, Nymphalidae). Fragmenta entomologica, 33 (2), 213 - 380."]}

Daedalma vertex milleri Pyrcz, n. ssp. (Figs. 3G, 3H, 10B) Material examined: PERU: HOLOTYPE male: Huánuco, Carpish, 2700–3000 m, 08.VI.1995, 0943/7606, G. Lamas leg., red, rectangular label saying: Holotype, MUSM; PARATYPES (9 males and 1 female): 3 males: Huánuco, Carpish, 2800–3000 m, 26.X.2004, P. Boyer leg., 2 PBF, 1 TWP; 1 male: same locality but 19.II.2003, TWP; 1 male: same data but 19.II.2003, PBF; 3 males: Huánuco, Carpish, 2700–2800 m, 08.VI.1995, G. Lamas leg., MUSM; 1 male: Huánuco, au dessus de Huanacaure, km 43 de Pachachupan, est de Acomayo, 3000–3100 m, P. Boyer leg., PBF; 1 female: Huánuco, 2 km au dessus de Huanacaure, route Pachachupan-Huanacaure km 42, 3000 m, 22.X.2009, P. Boyer leg., PBF. Description: MALE (Fig. 3G): Head, thorax and abdomen: not differing from nominate subspecies. Wings: FW length: 26–27 mm (mean: 26.2 mm, n=7), shorter than in nominate subspecies (mean: 28,6 mm); FWD ground colour blackish brown; orange patch a shade darker and smaller than in nominate subspecies, only slightly shorter distally but nearly half as wide; blackish postmedian patch in cell Cu1-Cu2 situated at edge of orange area, rather than enclosed by orange as in nominate subspecies. HWD uniform blackish brown, same as in nominate subspecies. New subspecies differs from nominate subspecies on FWV in shape and size of patch, and brick red colour, markedly darker in milleri. HWV pattern similar to nominate subspecies but ground colour chocolate brown instead of blackish brown. Genitalia (Fig. 10B): As compared to nominate subspecies, uncus same length but stouter, especially in basal part; gnathos similar; valvae narrowing gradually towards apex; saccus and aedeagus not differing noticeably from nominate subspecies. FEMALE (Fig. 3H): FWD patch wider than in the male; a shade lighter orange. HWD with orange reddish patches along costal margin to apical area, gradually darkening basally, some faint reddish markings in the submarginal and marginal area of M3-Cu1 and Cu1-Cu2. FWV similarly patterned as in the male except for more prominent orange markings. HWV patterned as in the male but with more prominent sandy yellow basal, postbasal and postdiscal markings, a dark brown median band, and olive scaling denser in postmedian and marginal areas. Genitalia: not examined. Etymology: This subspecies in dedicated to Lee D. Miller, eminent American entomologist, who contributed significantly to the knowledge of Pronophilina. Remarks: Daedalma vertex milleri is apparently confined to central Peru, where it occurs in Huánuco, and most probably in Pasco and Junín. Daedalma vertex milleri apparently occurs at slightly lower elevations than the nominate subspecies (Pyrcz, 2004), with available data indicating an altitudinal range of 2600–3100 m., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on page 24, {"references":["Pyrcz, T. W. (2004) Pronophiline butterflies of the highlands of Chachapoyas in northern Peru: faunal survey, diversity and distribution patterns (Lepidoptera, Nymphalidae, Satyrinae). Genus, 15 (4), 455 - 622."]}

Daedalma dinias Hewitson Daedalma dinias Hewitson [1858]: [85], pl. 1[43], figs. 1–3. Remarks: This is a polymorphic species occurring in all three Colombian Cordilleras and on the eastern slopes of the Andes in Ecuador. Its subspecies can be recognised immediately by the size, shape and ventral expression of the FW orange patch. It is generally parapatric above the similarly marked D. fraudata and D. rubroreducta n. ssp. in eastern Ecuador, from which it is recognised by the fact that the FW orange patch never extends into the postdiscal area (cell M2-M3)., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on page 9, {"references":["Hewitson, W. C. (1858) Satyridae. Daedalma. Illustrations of New Species of Exotic Butterflies, 2, 79 - 80."]}

Daedalma inconspicua tapichalaca Pyrcz & Willmott, n. ssp. (Figs. 6E, 6F, 12C) Material examined: HOLOTYPE male: Ecuador, Zamora-Chinchipe, Yangana-Valladolid rd., Reserva Tapichalaca, Loma Cruz Grande, 2650m, 4°29.22'S, 79°7.49'W, 14.IX.2007, J. Radford leg., FLMNH / MGCL# 143210, red, rectangular label saying: Holotype, FLMNH (to be deposited in MECN). PARATYPES (5 males, 1 female): 2 males: same data as HT except 01.XII.2005, K. Willmott leg., FLMNH / MGCL# 111174, 111175, FLMNH; 1 male, same data as HT except 13.IX.2007, JR (to be deposited in MZUJ); 1 male, 1 female: Ecuador, Zamora-Chinchipe, Yangana-Valladolid rd., Reserva Tapichalaca, trail to páramo (sector Ventanillas), 2770 m, 4°29.16'S, 79°8.96'W, 18.IX.2007, J. Radford leg., FLMNH / MGCL# 143213, 143214, FLMNH; 1 male, same data as previous except 19.IX.2007, JR. Additional material (8 males and 1 female): 1 male: Ecuador, Zamora-Chinchipe, Loja-Zamora Old Road, 2500 m, 31.V.2000, S. Attal leg., MZUJ; 5 males: Old road Loja-Zamora, 2500 m, XI.2000, I. Aldas leg., 3 TWP, 2 MBLI; 1 male: same data but XI–XII.1999, MBLI; 1 male: same data but 2600 m, XI.1999, MBLI; 1 female: Zamora-Chinchipe, old Loja-Zamora rd., 2640 m, 23.IX.2007, K. Willmott leg., FLMNH. Description: MALE (Fig. 6E): Head, thorax and abdomen: not differing from nominate subspecies. Wings: FW length: 24–27 mm, mean: 25.7, n=4. FW and HWD ground colour dark grey-brown, similar to orientalis or onorei; FWV with a light, milky white submarginal oblique, elongated patch, and a smaller patch of same colour in discal cell, thus differing from orientalis and other subspecies in which these patches are pale greyish, discal cell patch narrowly connected to pale grey postmedian area in cells M3-Cu1 and Cu1-Cu2; HWV variable in overall appearance, ranging from olive brown (as in palacio), to rufous orange (as in the holotype), to variegated dark brown, black and rufous, as in orientalis. Genitalia (Fig. 12C): Uncus slightly curved downwards; gnathos roughly 2/3 length of uncus, markedly stouter than in other subspecies; saccus long, similar to palacio; valvae length of tegumen + uncus, narrowing gradually from middle to apex, similar to cuencana or orientalis; aedeagus not differing from other subspecies. FEMALE (Fig. 6F): Head, thorax and abdomen: not differing from other subspecies. Wings: FW length: 29 mm. Dorsal ground colour dark brown with tinge of rufous scales, similar to orientalis; FW with dark reddish orange irregular discal band from costal edge discal cell to near tornus, considerably broader in cells Cu2-Cu1 and Cu1-M3, tapering in discal cell and in cell 2A-Cu2, similar in shape to palacio, paler at costa, black dot in middle of band in cell Cu2-Cu1; FW with dark reddish orange subapical band from costa to vein M3 c. 2mm in width, paler at costa; HW with dark reddish orange irregular postdiscal band enclosing single black spot in each cell from costa to vein Cu2, similar in shape to palacio but more strongly marked in cells Cu2-M3; HW "tails" medium brown; ventral surface similar to palacio, except for colour of FW discal band, rufous in cells 2A-M3 and posterior half discal cell, cream in anterior half discal cell, pale subapical band much broader than in palacio and cream, not orange. Etymology: This subspecies is named for the type locality, the Reserva Tapichalaca, established by the Fundación de Conservación Jocotoco (www.fjocotoco.org) to protect the faunistically unique forests of the Valladolid region. Remarks: A series of specimens from the upper Zamora valley, collected along the old Loja-Zamora road, is currently associated with this subspecies but excluded from the type series. These specimens share with those from the type locality the broad and pale FWV subapical band and pale discal band in the discal cell, thus being distinct from orientalis to the north (uniform greyish discal and subapical bands) and palacio to the west. In addition, the only known female is similar to that from the type locality in having much more extensive pale dorsal markings than orientalis, but these markings are orange, similar to palacio. It shares the broad FWV subapical band with the female from the type locality, thus differing from both orientalis and palacio. The phenotypic characteristics of the female and the variation observed among males from the Zamora-Loja road, and the fact that this region is not isolated by any geographic barrier from sites where typical palacio occurs, suggest that some introgression might occur in this region between palacio and tapichalaca., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on pages 37-38

Daedalma rubroreducta Pyrcz & Willmott, n. sp. (Figs. 3C, 3D, 9C, 14C, 18A–E, 19A–E, 23B) Material examined: ECUADOR: HOLOTYPE male: Morona-Santiago, Macas, Loma Tigrillo, 1700–1750 m, X.2002, I. Aldaz leg., red, rectangular label saying: Holotype, MZUJ; PARATYPES (9 males and 2 females), 4 males: same data as the holotype, TWP; 1 female: Ecuador, no further data, (prep. genit. 07- 01.07.2005), TWP; 1 female: Napo, Yanayacu, 2000 m, H. Greeney (ex larva), TWP; 1 male: same locality data as preceding but 24.XI.2006, K. Willmott leg., FLMNH; 1 male: Napo, Baeza-Tena km 10, "1600" [=1850] m, 12.XII.1996, P. Boyer leg., PBF; 1 male: Sucumbíos, km 2 La Bonita-Tulcán rd., 2100 m, 01.I.2002, K. Willmott leg., KWJH; 1 male: Tungurahua, Río Topo [mislabeled], ex. Grose-Smith, 1910, BMNH; 1 male: Sarayacu, Eastern Side Ecuador [mislabeled], C. Buckley 1879, ex. Oberthür coll., 1927-3, BMNH. Diagnosis: Differs from D. dinias by the FW orange patch extending into postdiscal area, similar to D. boliviana and D. fraudata, but differing from those species in the patch being slightly smaller and darker. On the underside, the FW orange patch is considerably fainter than in D. boliviana or D. fraudata, in most individuals obsolete, heavily overcast with brown and barely marked, in this respect being most similar to D. dinias emma. Description: MALE (Fig. 3C): Head: Eyes chocolate brown covered with dense setae; palpi twice the length of head, yellowish covered with chocolate brown hair; antennae 2/5 length of costa, dorsally brown, ventrally yellowish, club darker than shaft. Thorax: Dorsally and ventrally blackish, legs sandy yellow. Abdomen: Dorsally and laterally blackish, ventrally lighter dull brown. Wings: FW (length: 25-27 mm; 25.6 mm; n=5) costa slightly arched, outer margin protruded at vein M1 and slightly undulate. HW costa protruded at apex, outer margin undulate with two tail-like extensions at veins Cu1 and Cu2. Wings: FWD ground colour blackish brown; orange patch oval, smaller and slightly darker than in D. fraudata, with faint edges. HWD uniform blackish brown. FWV patch faint and diffuse, reddish, overcast with brown to such an extent that it is barely noticeable in some individuals. HWV overall darker than in D. fraudata with a heavier chocolate brown overcast, milky white postdiscal and submarginal patches smaller than in D. fraudata or obsolete. Genitalia (Fig. 9C): Similar to D. fraudata except for the thinner uncus, considerably broader valvae in distal half, and a slightly deeper saccus. FEMALE (Fig. 3D): Differs from D. fraudata in fainter and slightly smaller FW orange patch, suffused along edges with some brown scaling. Genitalia (Fig. 14C): Papillae anales small, with prominent bump on lateral side, slightly wrinkled wall of tuba analis. Lamella postvaginalis as a concave plate with a small protrusion pointing posteriorly. Bursa copulatrix pear-like, with two narrow and parallel signa consisting of minute teeth. Ductus bursae narrow, same length as the length of bursa copulatrix, connected with ductus seminalis in the middle. Lamella antevaginalis concave and shallow. Coliculum absent. Apophyses posteriores reduced. Early stages: Host plant: Chusquea cf. scandens Poaceae (2100 m, Yanayacu, Napo, Ecuador). EGGS: Data on morphology are not available, but newly emerged first instars were found in the field (see below) along a small road cut at the edge of an extensive patch of Chusquea (ca. 2 ha). The remains of the clutch (Fig. 18B) were on the underside of a mature (fully expanded) leaf located at the tip of a small isolated shoot draping over the road. FIRST INSTAR (Figs. 18A): Head capsule shining black, nearly round but slightly broader at base; epicranial suture weak; body elongate, parallel-sided with little or no constriction behind the head, roughly round in cross section; body colour dull olive-green with a red-brown cast anteriorly and posteriorly, becoming all red-brown later in instar; body with sparse, pale setae, prothoracic shield poorly developed and dark brown. First instars originally rested in a tightly packed group on the underside of the food plant leaf, near where the eggs were laid. Later, prior to molt, larvae aggregate at the apex of the skeletonized food plant leaf, rearing back and regurgitating when disturbed. The first instar lasted at least 5 days. SECOND INSTAR (Figs. 18C, 18F): Early 2 nd second instar head capsule similar to first instar, but with a pair of small, rounded, bump-like scoli anterio-dorsally; immediately after molting head capsule dark olive-green, paler dorsally, but darkening to shiny black later in instar; head and body with sparse dark setae; body nearly parallel sided, tapering slightly posteriorly to small, dark-brown bifid tail, roughly square in cross section; dorsum redbrown, sides and venter dark brown to black; a faint mid-dorsal stripe extends from T1 to around A3, T1 extended subdorsally into two individual fleshy lobes. Late 2 nd Instar: Overall colour pattern similar but shape becoming more elongate and nearly round in cross section, fleshy protuberances on T1 disappearing; a pair of thin creamcoloured stripes develop dorsolaterally and extend back from T1, gradually fading posteriorly; bifid tail and margins of anal plate become cream coloured. Second instars remain aggregated along the skeletonized mid-vein of the leaf. The larvae measured 8 mm at premolt. The second instar lasted 9 days. THIRD INSTAR (Figs. 18D): Early 3 rd instar head capsule dull black with scoli becoming more prominent and light brown to cream coloured, appearing as an extension of dorsolateral cream stripes on T1. Otherwise similar in colour to late second instars, but with bifid tail more prominent, and setae becoming slightly denser and longer, especially on head scoli. Late 3 rd Instar: As described for early third instar, but abdomen laterally developing a complex pattern of cream coloured stripes surrounding narrow red-brown areas. This pattern fades anteriorly and sides of thorax remain dark brown, spiracles, especially posteriorly develop small green areas surrounding them. Overall, appearing like a dead stick. Third instar larvae remain along the skeletonized leaf mid-vein. The larvae measured 13 mm at premolt. The third instar lasted 10 days. FOURTH INSTAR: As described for late third instar but with head scoli and bifid tails becoming more prominent and paler brown; anterior portion of head becoming dark red-brown with indistinct pale crescent-shaped spots on either side of clypeal suture; lateral patterning becoming more distinct and dorsolateral cream stripes becoming less distinct, especially posteriorly. Fourth instar larvae break into groups of 2–5 individuals but remain fairly clumped on the host plant. The larvae measured 20.5 mm at premolt. The fourth instar lasted 11–12 days. FIFTH INSTAR (Figs. 19A, 19B): Overall appearance much like a mossy stick, body roughly square in cross section, narrowing slightly from A1-A6, T2-T3 and A3 with small fleshy triangular protuberances dorsolaterally; larvae develop many small swellings at setal bases, especially on head and bifid tails; head patterning becoming stronger; early in instar larvae are patterned in various shades of brown with well defined pair of dark brown triangles subdorsally on A3; sides of thorax and A3-A4 dark brown with abdominal brown patch tapering posteriorly and supraspiracularly to A5 forming a distinct and roughly triangular patch laterally; dorsum of T1, A4–A5, and bifid tails becoming pale brown; late instar larvae develop mossy green highlights, especially on dorsal thorax, dorsum of A3, and around spiracles. Fifth instar larvae disperse and do not remain aggregated. The larvae measured a maximum of 35 mm before the prepupal stage (estimated from photographs). The fifth instar lasted 14–16 days. PUPA (Figs. 19C, 19D). All larvae were removed from the field before pupation and it is unknown where pupation occurs in the wild. Pupa heavily sculptured with a distinct forward curved thoracic keel, abdomen with subdorsal pairs of protrusions, progressively smaller towards cremaster; thorax laterally projected into two pairs of roundly triangular keels; head with a pair of dorso-ventrally flattened projections; overall colour dark brown with large bright metallic-green patches. The pupal length was not measured. The pupal stage lasted 28–30 days. Freshly emerged female (Fig. 19E). Etymology: The epithet of this species is an allusion to the diagnostic reduction of the reddish FW patch. Remarks: This species is known to occur in the area between Macas (Morona-Santiago) and La Bonita (Sucumbíos), along the eastern slopes of the Andes in Ecuador. We initially considered it as a subspecies of D. fraudata, from which it differs mostly in the expression of the FW orange patch, but the two taxa, if not strictly sympatric, have ranges that overlap over a broad region, in Tungurahua and Morona-Santiago, and both appear to occur in the Pastaza valley. Although the single specimen of D. rubroreducta from "Río Topo" is probably mislabelled and was presumably collected at higher elevations, the scant available elevational data do suggest that D. rubroreducta may in general inhabit slightly lower elevations than D. fraudata, which occurs in the Pastaza valley around and above Baños, as high as 2500 m. Other specimens of D. rubroreducta come from elevations slightly below 2000 m. Daedalma rubroreducta and D. boliviana are apparently allopatric but there is no good reason to regard them as conspecific, since the characters that distinguish D. rubroreducta from D. fraudata all apply to an even greater extent to distinguishing D. rubroreducta from D. boliviana., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on pages 18-20

Daedalma drusilla Hewitson Daedalma drusilla Hewitson, 1858: 86, pl. 1, fig. 7. Colombia, New Granada. Remarks: This is a polytypic species, whose subspecies are best separated by their females, which differ in the contrasting, orange, yellow or reddish upperside colour patterns. The males have few, if any, distinctive colour patterns, all being blackish brown on the upperside and differing, slightly, in the FWV subapical markings. Comparative material from Colombia is, unfortunately, poor, particularly for females. Moreover, most examined females from Cundinamarca do not bear precise collecting locality data, hindering our understanding of the distribution of D. drusilla subspecies. The systematic arrangement presented below should therefore be considered preliminary. Adams (1986) claimed that D. drusilla is restricted to the Colombian Eastern Cordillera, but this is incorrect, as this species is found in other Colombian ranges as well, certainly in the Central Cordillera and, possibly, also in the Western Cordillera (Pyrcz & Rodríguez, 2007). It also occurs in northern Ecuador and south-western Venezuela (Pyrcz & Viloria, 2007). Daedalma drusilla is closely related to D. inconspicua and the two are locally sympatric and/or parapatric along an elevational gradient in the Colombian Central Cordillera and northern Ecuador, supporting their separate species status. D. drusilla is generally smaller than D. inconspicua, with males darker, blackish brown on the upperside, and with shorter HW “tails”., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on page 27, {"references":["Hewitson, W. C. (1858) Satyridae. Daedalma. Illustrations of New Species of Exotic Butterflies, 2, 79 - 80.","Adams, M. J. (1986) Pronophiline butterflies (Satyridae) of the three Andean Cordilleras of Colombia. Zoological Journal of the Linnean Society, 87, 235 - 320.","Pyrcz, T. W. & Rodriguez, G. (2007) Mariposas de la tribu Pronophilina en la Cordillera Occidental de Colombia. Shilap, 35, 455 - 489.","Pyrcz, T. W. & Viloria, A. L. (2007) Erebiine and pronophiline butterflies of the Serrania del Tama, Venezuela-Colombia border (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera, 15 (1 - 2), 18 - 52."]}

Daedalma inconspicua Butler Daedalma inconspicua Butler, 1866: 77. Daedalma inconspicua Butler; Butler, 1868: 183; Kirby, 1871: 107; Thieme, 1906: 137; Weymer, 1912: 266. Remarks: Daedalma inconspicua is very similar to D. drusilla, and characters useful for distinguishing the two species are discussed under the latter species. Daedalma inconspicua is a polytypic species. Males of the various subspecies can be recognised by size and by the amount and shape of the pale yellow on the FWV. Females differ more markedly by their dorsal reddish or yellow patterns, similar to the females of D. drusilla. Locally occurring subspecies distributed throughout Ecuador, extreme northern Peru and Colombia show intricate distributional patterns. Pacific and Amazonian slope faunas of cloud forest butterflies, including populations of Daedalma, are geographically widely isolated in northern and central Ecuador by the Andes, with ridges rising above 4000–5000 m covered with páramo grassland vegetation and a dry inter-Andean valley at 2500–3000 m. However, in southern Ecuador the Andean Cordillera is lower and there is no inter-Andean plateau, but instead a complex system of relatively low and narrow ranges descending below 2500 m at some of the lowest passes. The orography is highly complicated and it is difficult to identify the main Andean ridge. The lowest point between the watersheds of the Amazon and the Pacific Ocean some kilometres south of Loja is situated at merely 2450 m. Such a topography affects the distribution patterns of Daedalma taxa, with otherwise isolated populations potentially coming into contact in southern Ecuador. Such contact is facilitated by the behaviour of Daedalma species, which are less sedentary than most other pronophilines. They may sometimes fly over long distances, with some specimens having been collected over páramo, some distance from the nearest cloud forest. Occasional dispersal may lead to gene flow between western and eastern slopes populations, and indeed there are specimens whose phenotypes suggest that hybridisation at the subspecific level does occur. This phenomenon is particularly clear in D. inconspicua, an uppermost forest species., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on page 31, {"references":["Weymer, G. (1912) Satyridae [publ. 1910 - 1912]. in A. Seitz (ed.), Die Gross-Schmetterlinge der Erde, 2; Exotische Fauna, 5, 173 - 283."]}

Daedalma dognini Pyrcz, n. sp. [Daedalma drusilla n. ssp. Lamas; Lamas et al., 2004: 208.] Remarks: This species is currently comprised of two recognised subspecies, occurring in the south-eastern Peruvian Andes. The dorsal surface is grey-brown and similar to that of D. inconspicua, but the wing shape and ventral colour patterns are more reminiscent of D. adamsi.

Daedalma rubroreducta Pyrcz & Willmott, n. sp. (Figs. 3C, 3D, 9C, 14C, 18A–E, 19A–E, 23B) Material examined: ECUADOR: HOLOTYPE male: Morona-Santiago, Macas, Loma Tigrillo, 1700–1750 m, X.2002, I. Aldaz leg., red, rectangular label saying: Holotype, MZUJ; PARATYPES (9 males and 2 females), 4 males: same data as the holotype, TWP; 1 female: Ecuador, no further data, (prep. genit. 07- 01.07.2005), TWP; 1 female: Napo, Yanayacu, 2000 m, H. Greeney (ex larva), TWP; 1 male: same locality data as preceding but 24.XI.2006, K. Willmott leg., FLMNH; 1 male: Napo, Baeza-Tena km 10, "1600" [=1850] m, 12.XII.1996, P. Boyer leg., PBF; 1 male: Sucumbíos, km 2 La Bonita-Tulcán rd., 2100 m, 01.I.2002, K. Willmott leg., KWJH; 1 male: Tungurahua, Río Topo [mislabeled], ex. Grose-Smith, 1910, BMNH; 1 male: Sarayacu, Eastern Side Ecuador [mislabeled], C. Buckley 1879, ex. Oberthür coll., 1927-3, BMNH. Diagnosis: Differs from D. dinias by the FW orange patch extending into postdiscal area, similar to D. boliviana and D. fraudata, but differing from those species in the patch being slightly smaller and darker. On the underside, the FW orange patch is considerably fainter than in D. boliviana or D. fraudata, in most individuals obsolete, heavily overcast with brown and barely marked, in this respect being most similar to D. dinias emma. Description: MALE (Fig. 3C): Head: Eyes chocolate brown covered with dense setae; palpi twice the length of head, yellowish covered with chocolate brown hair; antennae 2/5 length of costa, dorsally brown, ventrally yellowish, club darker than shaft. Thorax: Dorsally and ventrally blackish, legs sandy yellow. Abdomen: Dorsally and laterally blackish, ventrally lighter dull brown. Wings: FW (length: 25-27 mm; 25.6 mm; n=5) costa slightly arched, outer margin protruded at vein M1 and slightly undulate. HW costa protruded at apex, outer margin undulate with two tail-like extensions at veins Cu1 and Cu2. Wings: FWD ground colour blackish brown; orange patch oval, smaller and slightly darker than in D. fraudata, with faint edges. HWD uniform blackish brown. FWV patch faint and diffuse, reddish, overcast with brown to such an extent that it is barely noticeable in some individuals. HWV overall darker than in D. fraudata with a heavier chocolate brown overcast, milky white postdiscal and submarginal patches smaller than in D. fraudata or obsolete. Genitalia (Fig. 9C): Similar to D. fraudata except for the thinner uncus, considerably broader valvae in distal half, and a slightly deeper saccus. FEMALE (Fig. 3D): Differs from D. fraudata in fainter and slightly smaller FW orange patch, suffused along edges with some brown scaling. Genitalia (Fig. 14C): Papillae anales small, with prominent bump on lateral side, slightly wrinkled wall of tuba analis. Lamella postvaginalis as a concave plate with a small protrusion pointing posteriorly. Bursa copulatrix pear-like, with two narrow and parallel signa consisting of minute teeth. Ductus bursae narrow, same length as the length of bursa copulatrix, connected with ductus seminalis in the middle. Lamella antevaginalis concave and shallow. Coliculum absent. Apophyses posteriores reduced. Early stages: Host plant: Chusquea cf. scandens Poaceae (2100 m, Yanayacu, Napo, Ecuador). EGGS: Data on morphology are not available, but newly emerged first instars were found in the field (see below) along a small road cut at the edge of an extensive patch of Chusquea (ca. 2 ha). The remains of the clutch (Fig. 18B) were on the underside of a mature (fully expanded) leaf located at the tip of a small isolated shoot draping over the road. FIRST INSTAR (Figs. 18A): Head capsule shining black, nearly round but slightly broader at base; epicranial suture weak; body elongate, parallel-sided with little or no constriction behind the head, roughly round in cross section; body colour dull olive-green with a red-brown cast anteriorly and posteriorly, becoming all red-brown later in instar; body with sparse, pale setae, prothoracic shield poorly developed and dark brown. First instars originally rested in a tightly packed group on the underside of the food plant leaf, near where the eggs were laid. Later, prior to molt, larvae aggregate at the apex of the skeletonized food plant leaf, rearing back and regurgitating when disturbed. The first instar lasted at least 5 days. SECOND INSTAR (Figs. 18C, 18F): Early 2 nd second instar head capsule similar to first instar, but with a pair of small, rounded, bump-like scoli anterio-dorsally; immediately after molting head capsule dark olive-green, paler dorsally, but darkening to shiny black later in instar; head and body with sparse dark setae; body nearly parallel sided, tapering slightly posteriorly to small, dark-brown bifid tail, roughly square in cross section; dorsum redbrown, sides and venter dark brown to black; a faint mid-dorsal stripe extends from T1 to around A3, T1 extended subdorsally into two individual fleshy lobes. Late 2 nd Instar: Overall colour pattern similar but shape becoming more elongate and nearly round in cross section, fleshy protuberances on T1 disappearing; a pair of thin creamcoloured stripes develop dorsolaterally and extend back from T1, gradually fading posteriorly; bifid tail and margins of anal plate become cream coloured. Second instars remain aggregated along the skeletonized mid-vein of the leaf. The larvae measured 8 mm at premolt. The second instar lasted 9 days. THIRD INSTAR (Figs. 18D): Early 3 rd instar head capsule dull black with scoli becoming more prominent and light brown to cream coloured, appearing as an extension of dorsolateral cream stripes on T1. Otherwise similar in colour to late second instars, but with bifid tail more prominent, and setae becoming slightly denser and longer, especially on head scoli. Late 3 rd Instar: As described for early third instar, but abdomen laterally developing a complex pattern of cream coloured stripes surrounding narrow red-brown areas. This pattern fades anteriorly and sides of thorax remain dark brown, spiracles, especially posteriorly develop small green areas surrounding them. Overall, appearing like a dead stick. Third instar larvae remain along the skeletonized leaf mid-vein. The larvae measured 13 mm at premolt. The third instar lasted 10 days. FOURTH INSTAR: As described for late third instar but with head scoli and bifid tails becoming more prominent and paler brown; anterior portion of head becoming dark red-brown with indistinct pale crescent-shaped spots on either side of clypeal suture; lateral patterning becoming more distinct and dorsolateral cream stripes becoming less distinct, especially posteriorly. Fourth instar larvae break into groups of 2–5 individuals but remain fairly clumped on the host plant. The larvae measured 20.5 mm at premolt. The fourth instar lasted 11–12 days. FIFTH INSTAR (Figs. 19A, 19B): Overall appearance much like a mossy stick, body roughly square in cross section, narrowing slightly from A1-A6, T2-T3 and A3 with small fleshy triangular protuberances dorsolaterally; larvae develop many small swellings at setal bases, especially on head and bifid tails; head patterning becoming stronger; early in instar larvae are patterned in various shades of brown with well defined pair of dark brown triangles subdorsally on A3; sides of thorax and A3-A4 dark brown with abdominal brown patch tapering posteriorly and supraspiracularly to A5 forming a distinct and roughly triangular patch laterally; dorsum of T1, A4–A5, and bifid tails becoming pale brown; late instar larvae develop mossy green highlights, especially on dorsal thorax, dorsum of A3, and around spiracles. Fifth instar larvae disperse and do not remain aggregated. The larvae measured a maximum of 35 mm before the prepupal stage (estimated from photographs). The fifth instar lasted 14–16 days. PUPA (Figs. 19C, 19D). All larvae were removed from the field before pupation and it is unknown where pupation occurs in the wild. Pupa heavily sculptured with a distinct forward curved thoracic keel, abdomen with subdorsal pairs of protrusions, progressively smaller towards cremaster; thorax laterally projected into two pairs of roundly triangular keels; head with a pair of dorso-ventrally flattened projections; overall colour dark brown with large bright metallic-green patches. The pupal length was not measured. The pupal stage lasted 28–30 days. Freshly emerged female (Fig. 19E). Etymology: The epithet of this species is an allusion to the diagnostic reduction of the reddish FW patch. Remarks: This species is known to occur in the area between Macas (Morona-Santiago) and La Bonita (Sucumbíos), along the eastern slopes of the Andes in Ecuador. We initially considered it as a subspecies of D. fraudata, from which it differs mostly in the expression of the FW orange patch, but the two taxa, if not strictly sympatric, have ranges that overlap over a broad region, in Tungurahua and Morona-Santiago, and both appear to occur in the Pastaza valley. Although the single specimen of D. rubroreducta from "Río Topo" is probably mislabelled and was presumably collected at higher elevations, the scant available elevational data do suggest that D. rubroreducta may in general inhabit slightly lower elevations than D. fraudata, which occurs in the Pastaza valley around and above Baños, as high as 2500 m. Other specimens of D. rubroreducta come from elevations slightly below 2000 m. Daedalma rubroreducta and D. boliviana are apparently allopatric but there is no good reason to regard them as conspecific, since the characters that distinguish D. rubroreducta from D. fraudata all apply to an even greater extent to distinguishing D. rubroreducta from D. boliviana.

Daedalma dognini mirianae Pyrcz, n. ssp. (Fig. 4F, 11B) Material examined: PERU: HOLOTYPE male: Cuzco, P. N. Manu, Acjanaco, 3200–3450 m, 05.VII.1991, M. Medina leg., red, rectangular label saying: Holotype, MUSM; PARATYPES (3 males): 2 males: same data as HT but 07.VII.1991, MUSM; 1 male: Cuzco, Abra Acjanaco vers Pillcopata km 10, 3000–3100 m, 24.II.2005, P. Boyer leg., PBF. Description: MALE (Figs. 4F): Head, thorax and abdomen not differing from nominate subspecies. FW length 27 mm. FWD and HWD dark brown, a shade darker than in nominate subspecies; FWV differs immediately from that of nominate subspecies in totally lacking any trace of red suffusion in wide, light brown postmedian patch. HWV ground colour lighter brown with an olive shade, otherwise similar to nominate. Genitalia (Fig. 11B): Uncus longer than in nominate; gnathos and saccus similar; valvae also longer than in nominate with a wider apical half, apical spine-like process slightly shorter; aedeagus longer than in nominate, otherwise similar. FEMALE: Unknown. Etymology: This subspecies is dedicated to Mirian Medina Hay-Roe, Peruvian entomologist and first collector of this taxon. Remarks: D. dognini mirianae is known so far exclusively from the upper valley of Kosñipata, but its range is most probably much wider. The parallel and more southerly valleys of Marcapata and Ollachea have barely been sampled for butterflies., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on page 27

Daedalma inconspicua tapichalaca Pyrcz & Willmott, n. ssp. (Figs. 6E, 6F, 12C) Material examined: HOLOTYPE male: Ecuador, Zamora-Chinchipe, Yangana-Valladolid rd., Reserva Tapichalaca, Loma Cruz Grande, 2650m, 4°29.22'S, 79°7.49'W, 14.IX.2007, J. Radford leg., FLMNH / MGCL# 143210, red, rectangular label saying: Holotype, FLMNH (to be deposited in MECN). PARATYPES (5 males, 1 female): 2 males: same data as HT except 01.XII.2005, K. Willmott leg., FLMNH / MGCL# 111174, 111175, FLMNH; 1 male, same data as HT except 13.IX.2007, JR (to be deposited in MZUJ); 1 male, 1 female: Ecuador, Zamora-Chinchipe, Yangana-Valladolid rd., Reserva Tapichalaca, trail to páramo (sector Ventanillas), 2770 m, 4°29.16'S, 79°8.96'W, 18.IX.2007, J. Radford leg., FLMNH / MGCL# 143213, 143214, FLMNH; 1 male, same data as previous except 19.IX.2007, JR. Additional material (8 males and 1 female): 1 male: Ecuador, Zamora-Chinchipe, Loja-Zamora Old Road, 2500 m, 31.V.2000, S. Attal leg., MZUJ; 5 males: Old road Loja-Zamora, 2500 m, XI.2000, I. Aldas leg., 3 TWP, 2 MBLI; 1 male: same data but XI–XII.1999, MBLI; 1 male: same data but 2600 m, XI.1999, MBLI; 1 female: Zamora-Chinchipe, old Loja-Zamora rd., 2640 m, 23.IX.2007, K. Willmott leg., FLMNH. Description: MALE (Fig. 6E): Head, thorax and abdomen: not differing from nominate subspecies. Wings: FW length: 24–27 mm, mean: 25.7, n=4. FW and HWD ground colour dark grey-brown, similar to orientalis or onorei; FWV with a light, milky white submarginal oblique, elongated patch, and a smaller patch of same colour in discal cell, thus differing from orientalis and other subspecies in which these patches are pale greyish, discal cell patch narrowly connected to pale grey postmedian area in cells M3-Cu1 and Cu1-Cu2; HWV variable in overall appearance, ranging from olive brown (as in palacio), to rufous orange (as in the holotype), to variegated dark brown, black and rufous, as in orientalis. Genitalia (Fig. 12C): Uncus slightly curved downwards; gnathos roughly 2/3 length of uncus, markedly stouter than in other subspecies; saccus long, similar to palacio; valvae length of tegumen + uncus, narrowing gradually from middle to apex, similar to cuencana or orientalis; aedeagus not differing from other subspecies. FEMALE (Fig. 6F): Head, thorax and abdomen: not differing from other subspecies. Wings: FW length: 29 mm. Dorsal ground colour dark brown with tinge of rufous scales, similar to orientalis; FW with dark reddish orange irregular discal band from costal edge discal cell to near tornus, considerably broader in cells Cu2-Cu1 and Cu1-M3, tapering in discal cell and in cell 2A-Cu2, similar in shape to palacio, paler at costa, black dot in middle of band in cell Cu2-Cu1; FW with dark reddish orange subapical band from costa to vein M3 c. 2mm in width, paler at costa; HW with dark reddish orange irregular postdiscal band enclosing single black spot in each cell from costa to vein Cu2, similar in shape to palacio but more strongly marked in cells Cu2-M3; HW "tails" medium brown; ventral surface similar to palacio, except for colour of FW discal band, rufous in cells 2A-M3 and posterior half discal cell, cream in anterior half discal cell, pale subapical band much broader than in palacio and cream, not orange. Etymology: This subspecies is named for the type locality, the Reserva Tapichalaca, established by the Fundación de Conservación Jocotoco (www.fjocotoco.org) to protect the faunistically unique forests of the Valladolid region. Remarks: A series of specimens from the upper Zamora valley, collected along the old Loja-Zamora road, is currently associated with this subspecies but excluded from the type series. These specimens share with those from the type locality the broad and pale FWV subapical band and pale discal band in the discal cell, thus being distinct from orientalis to the north (uniform greyish discal and subapical bands) and palacio to the west. In addition, the only known female is similar to that from the type locality in having much more extensive pale dorsal markings than orientalis, but these markings are orange, similar to palacio. It shares the broad FWV subapical band with the female from the type locality, thus differing from both orientalis and palacio. The phenotypic characteristics of the female and the variation observed among males from the Zamora-Loja road, and the fact that this region is not isolated by any geographic barrier from sites where typical palacio occurs, suggest that some introgression might occur in this region between palacio and tapichalaca.

Daedalma dognini dognini Pyrcz (Figs. 4G, 4H, 11A, 14D) Material examined: PERU: HOLOTYPE male: Cuzco, Ollantaytambo-Alfamayo, Qda. San Lu��s, Carrizales, 3100���3150 m, 20.V.2003, T. Pyrcz leg., red, rectangular label saying: Holotype, MZUJ (to be deposited in MUSM); PARATYPE female: same data but 3200���3250 m, TWP. Diagnosis: Recognised from the only sympatric congener, D. boliviana, by the uniform brown upperside; very similar to the Colombian species D. drusilla, except for the slightly lighter upperside, wider wings and a reddish suffusion of the FWV. Description: MALE (Fig. 4G): Head: eyes blackish brown, hairy; labial palpi covered with dense, short, brown hair; antennae extending to 2/5 length of costa, orange, shaft slender, club twice as thick and flattened. Thorax: blackish brown; legs light beige. Abdomen: blackish brown. Wings: FW length 27 mm; outer margin sharply produced at vein M1; fringes yellowish. HW with two tail-like extensions along veins Cu1 and Cu2, former slightly longer; fringes very short, brown, except at apex where yellowish and slightly longer. FWD: uniform medium brown, with a very faint, barely noticeably lighter postdiscal area reflecting shape of reddish patch from underside. HWD: uniform medium brown. FWV: ground colour medium brown; a faint reddish suffusion in postbasal area, and a red-brown, irregular patch in discal cell; a wide red-light brown area covering most of postdiscal to submarginal area, reddish suffusion fading away towards subapical area giving place to light brown, and some white scaling along costa, with a deep notch pointing distally along its basal edge between veins M1 and M3; subapical area chocolate brown with some silver and magenta scaling forming a zigzagging submarginal line extending as far as vein M3; three minute subapical ocelli in R4-R5, R5-M1 and M1-M2. HWV: ground colour chocolate brown with some red-brown and milky white scaling forming a complex, nondescript mosaic typical of genus; two milky white submarginal patches in M2-M3 and M3-Cu1, latter noticeably bigger and reminiscent in shape and position of that of D. drusilla. Genitalia (Fig: 11A): Uncus stout, longer than dorsum of tegumen; gnathos 2/3 length of uncus, gradually thinner from a wide base to a fine apical part; saccus deep, about length of uncus; pedunculus prominent, especially compared to D. inconspicua; valvae wide and solid in basal half, gradually narrowing towards apex, terminated with a rather short hook curved upwards; aedeagus about length of valvae, smooth, slightly arched in middle. FEMALE (Fig. 4H): FW length 29 mm; FWD and HWD lighter than in male, with lighter patches showing more conspicuously from underside; FWV reddish suffusion lighter and much fainter, concentrated along basal edge of light orange-brown postmedian area. HWV colour pattern lighter, otherwise similar to male. Genitalia (Fig. 14D): Sinus vaginalis wide and shallow. Marginal parts of V-shaped batten on lamella postvaginalis consisting of heavily sclerotized slats that narrow slightly laterally. Papillae anales posteriorly close to each other. Apophyses posteriores as short triangular plates sharply pointing anteriorly. Posterior wall of lamella postvaginalis concave, with delicate wrinkles. Ductus bursae curved, with few pronounced wrinkles, twice length of antrum. Posteriorly to well developed colliculum, another ring-like cuticular thickening on wall of antrum. Hump in middle of lamella postvaginalis very small, triangular. Bursa copulatrix with two ribbon-like and narrow, parallel signa consisting of minute teeth. Etymology: This species is dedicated to Paul Dognin, the French naturalist who, by the end of 19th century, had described several species and subspecies of Pronophilina, including Daedalma (inconspicua) palacio. Remarks: This species superficially resembles the northern Andean D. drusilla and D. inconspicua, particularly in the uniform dorsal brown pattern. However, their ventral patterns, showing some distinctive characters such as the wide reddish FW patch (conspicuous in the nominate subspecies), and the male genitalia, are sufficiently distinct to treat D. dognini as a species. Furthermore, any closer association of D. dognini with D. inconspicua would be unjustified from a biogeographical standpoint, considering that their ranges are widely disjunct. D. dognini has no known close relatives in central Peru. D. adamsi, which occurs at similar elevations in central Peru, has very different male genitalia, which rules out any closer affinity (see under D. adamsi). Available elevational data (3100���3250 m) indicate that D. dognini is an uppermost forest species occurring near the treeline., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on page 26

Daedalma dognini Pyrcz, n. sp. [Daedalma drusilla n. ssp. Lamas; Lamas et al., 2004: 208.] Remarks: This species is currently comprised of two recognised subspecies, occurring in the south-eastern Peruvian Andes. The dorsal surface is grey-brown and similar to that of D. inconspicua, but the wing shape and ventral colour patterns are more reminiscent of D. adamsi., Published as part of Pyrcz, Tomasz W., Greeney, Harold F., Willmott, Keith R. & Wojtusiak, Janusz, 2011, 2898, pp. 1-68 in Zootaxa 2898 on page 25, {"references":["Lamas, G., Viloria, A. L. & Pyrcz, T. W. (2004) Subtribe Pronophilina, in: G. Lamas ed., Atlas of Neotropical Lepidoptera, Checklist: Part 4 A, Hesperoidea - Papilionoidea. Association for Tropical Lepidoptera, Gainesville, pp. 206 - 215."]}