Johnsonita auda ( Hewitson, 1867 ) d’Abrera & Bálint, 2001

Bálint, Zsolt, Boyer, Pierre, Cerdeña, José, Larico, Jackie Farfán, Brudecka, Jadwiga Lorenc-, Prieto, Carlos & Pyrcz, Tomasz W., 2021, Contributions to the knowledge of Neotropical Lycaenidae: taxonomy of Johnsonita Salazar & Constantino, 1995 with descriptions of seven new species (Theclinae Eumaeini), Zootaxa 4963 (1), pp. 11-57 : 16-19

Johnsonita auda ( Hewitson, 1867) d’Abrera & Bálint, 2001

( Figs 1A , 3 , 16 , 17A–B , 20A–C , 24G , 26 , 31 )

Theca auda – Hewitson 1867: 8; Draudt 1919 -1921: 754; d’Abrera 1995: 1131.

Owda auda (Hewitson) – Johnson, Kruse & Kroenlein 1997: 19 (lectotype designation).

Johnsonita auda (Hewitson) – d’Abrera & Bálint 2001: 195 (new combination); Lamas 2003: 49; Bálint 2003: 48; Robbins 2004b: 120.

Type material ( Fig. 26 ). The species was described on the basis of an unstated number of female specimens from New Granada ( Colombia). Johnson, Kruse & Kroenlein (1997) designated the lectotype indicating that it was a male and referring the specimen as “ Thecla auda, Hew (5)” from the Hewitson collection. Although this specimen is not a male, but a female, the labels obviously testify that it is the specimen selected, and only its sex was mistaken: (1) “ New Granada. [//] Hewitson Coll. [//] 79—69. [//] Thecla [//] auda, Hew. (5) ”(white label, printed and handwritten (Kirby) in black), (2) “Type” (confetti-type label with red frame, printed), (3) “B.M. TYPE [//] No.Rh. 570 ” (white label, printed and handwritten in black), (4) “B.M.(N.H.) [//] Rhopalocera [//] Slide V No. [//] 3491 ” (white label printed in red, handwritten in black), (5) “ f#” [//] SYNTYPE [//] Thecla auda [//] Hewitson [//] London, VI. 22. [//] Zs. Bálint, 2000” (red label with black frame, printed and handwritten (Bálint) black), (6) “ BMNH (E) # 266749” (white label, printed) .

Material examined (n=88; ten dissections).

COLOMBIA (n=18; three dissections): Risaralda, Pereira, La Florida, La Suiza, Ucuman, Vereda El Cedral, 2300 m, 4.X.1993, G. Andrade C. ( RCCP: female) ; Risaralda, Pereira, La Florida, La Suiza, Parque Ucumán, 2545 m, 29.IX–8.X.1993, G. Andrade C. ( RCCP: female) ; Caldas, Manizales, Rio Bamba y Rio Campoalegra , 3000 m, 24.X.1993, Cardos leg. ( HNHM: female, gen. prep. 894/ Bálint) ; Caldas, Rio La Miel , 3000 m, 16. V.1994, F. Henao leg. ( HNHM: male, gen. prep. 1627/ Bálint) ; Caldas, Cerro Ingrumá , 2270 m, 19.VII.1994, J. Salazar leg. ( HNHM: female) ; Caldas, El Aquila, 3.III.1996, J. Salazar leg. ( HNHM: female) ; Bogotá, Quebrada San Francisco, 2000–2100 m, 22.IX.1997, A. Jasiński ( CEP – MZUJ: male, gen. prep. 369/ J. Lorenc); Valle, Cali , Alto Aguaca- tal– Brisas , 1970 m, 18.VIII.2005, C. Prieto leg. ( HNHM: male) ; Cundinamarca, Páramo de Guasca, Peña Negra , 17.III.2006, T. T. Pyrcz leg. ( CEP – MZUJ: female) ; Cauca, PNN Muchique, La Romelia, 2640 m, 16. VI.2006, C. Prieto ( RCCP: female) ; Putumayo, Mocoa, Río Mocoa , 600 m, 15.X.2012, C. Prieto ( RCCP: female) ; Cauca, San Juan de Villalobos , 1800 m, 22. VI.2013, C. Prieto ( RCCP: female) ; Putumayo, Mocoa, las Antenas , 24.I. 2017, P. Boyer ( CB: 2 females) ; Antioquia, via Uramita, Peque, 2000 m, 3.I.2018, C. Prieto ( RCCP: female); Valle , Cali , La Elvira , 1900 m, 1.VIII.2018, C. Prieto ( RCCP: male, 2 females); Valle, Cali, Cerro Brisas, 1850 m, 6.VIII.2018, C. Prieto ( RCCP: male) .

ECUADOR (n=45; three dissections): [no data], A. Jasiński ( CEP – MZUJ: 2 males); Prov. Zamora, Rio Capi- chalaca via Zumba, 15.XII.1988, A. Jasiński ( CEP – MZUJ: female); Prov. Zamora, Quebrada Molinos prox (?) de Zamora, 8.XII.1989, A. Jasiński ( CEP – MZUJ: female); Prov. Zamora, Quebrada Tapichoclea (?), 2300 m, 21.X.1989, A. Jasiński ( CEP – MZUJ: female); Prov. Tungurahua, Viscaya, 2150 m, 2.V.1996, A. Jasiński ( CEP – MZUJ: 2 males); Prov. Tungurahua, Banos, 1800-2000 m, 9.V.1996, A. Jasiński ( CEP – MZUJ: female, gen. prep. 837/ J. Lorenc); Prov. Zamora, Tambo 1500 m; 20. XI.1996 ( CB: female); Prov. Zamora, Loja –Zamora km25, 2700 m; 21. XI.1996 ( CB: female); Prov. Tungurahua. Baños 1900 m, 12. XII. 1996 ( CB: male); Prov. Loja –Zamora, I. 1997 ( CB: male); Prov. Zamora, Loja –Zamora km30, 1800 m; 20 XII. 1996 ( CB: female); Prov. Pastaza, Chinchín, IV.1997 ( CB: female); Prov. Loja, Quebrada Los Monos, 2000–2100 m, 26.IV.1997, A. Jasiński ( CEP-MZUJ: fe- male); Prov. Tungurahua, Río Topo, 1200–1400 m, III. 1998 ( CB: 2 males); Prov. Morona–Santaigo, Macas 9 de Octubre, 1800 m, 3.III.1998, P. Boyer ( CEP – MZUJ: male); Prov. Loja, Cordillera las Llagunillas, road San Andrés–Calderón, 2000–2600 m, 20.V.1998, A. Jasiński ( CEP-MZUJ: female); Prov. Loja, Loja –Zumba km95-100, 2600 m, 27. XI. 1998 ( CB: female); Prov. Zamora, Santa Cruz de Nambija , cordillera de Nanguipa, 1600-2000 m, 30.XI.1998 ( CB: female); Prov. Morona–Santiago, San Juan Bosco, 11.VIII.2001, J. Wojtusiak ( CEP – MZUJ: 6 males, gen. prep. 840/ J. Lorenc, 2 females); Prov. Morona–Santiago, 9 de Octubre-Macas, 1700 m, VII.2001, J. Wojtusiak ( CEP – MZUJ: male, gen. prep. 368/ J. Lorenc); Prov. Morona–Santiago, Macas, 1700 m, X.2001, J. Wojtusiak ( CEP – MZUJ: 5 males, gen.prep. 839/ J. Lorenc, female); Prov. Tungurahua, Tungurahua volcano, Baños – Pondoa road, 3200 m, 17.II. I.2002, J. Wojtusiak & Garlacz ( CEP – MZUJ: male); Prov. Morona / Santiago, Chiguinda, 1750 m, I.2002, Aldas ( CEP – MZUJ: female); Prov. Bolivar, Balzabamba, 1200 m, 5.IX.2004, J. Wojtusiak & Pyrcz ( CEP – MZUJ: female); Prov. Loja. Valladoid, no date, A. Jasiński ( CEP – MZUJ: female); Prov. Morona – Santiago, Guarumales, hidropaute 2100–2300 m, 11. VIII. 2011 ( CB: female); Prov. Loja, Yangana, route de Cerro Toledo 2600–2800 m, 12.VIII.2017 ( CB: female); Prov. Loja, Yangana, route de Cerro Toledo 3100 m, 12.VIII.2017, P. Boyer ( CB: female); Prov. Loja, Yangana, route de Cerro Toledo 2600–2800 m, 12.VIII.2017 ( CB: female); Prov. Zamora, NW San Andrés, Zumba – Jimbura 2100 m, 14.VIII.2017, P. Boyer ( CB: female); Morona – Santiago, Guarumales, 2000–2200 m, no data ( CB: female). Prov. Zamora, NW San Andres, Loja – Jimbura 2000–2100 m, 14. VIII. 2017 ( CB: female) .

PERU (n=12; one dissection). Dept. Cuzco, Machu Picchu, 1.VIII.1984, via K. Johnson ( HNHM: female; gen. prep. 893/Bálint); Dept. Amazonas, la Sonada, Pomacocha, 1500–3000 m, IX. 2000 ( CB: female); Dept. Amazonas, Pedro Ruiz Gallo, II. 2002 ( CB: female); Dept. Amazonas, Cedro de Pirruro, Rodriguez de Mendoza , 2000 m, IV.2002 ( CB: female); Dept. Amazonas, Rodriguez de Mendoza , 1400 m, I. 2003, B. Calderon ( HNHM: male); Dept. San Martin –Amazonas, Jorge Chavez, 1450-1800 m, V.2003 ( CB: female); Dept. San Martin –Amazonas, Jorge Chavez, 1450-1800 m, XI. 2003 ( CB: male); Dept. Pasco, Santa Cruz, Oxapampa–Villavicencio km10, 2100– 2200 m, 5.XII.2003 ( CB: female); Dept. San Martin –Amazonas, Jorge Chavez, 1450–1800 m, II. 2004 ( CB: female); Dept. Pasco, SE Oxapampa 2400–2600 m, 16.X.2004 ( CB: female); Dept. Pasco, la Antena, SE Oxapampa, 2400–2600 m, 29.X.2004 ( CB: male, female); Dept. Pasco, la Suiza, Oxapampa 2200 m, XI.2006 ( CB: female) .

VENEZUELA (n=13; one dissection). Edo. Mérida, La Cascasa, 900 m, VIII.1966, F. Romero ( HNHM: female), Edo. Aragua, Choroní, la Casita, 1400 m. X. 1977, Romero ( HNHM: male); Edo. Barinas, San Isidro, 1700 m, 3.IV.1986, F. Romero Jr. ( HNHM: female); Edo. Mérida, Monte Zerpa, Parc National La Culata, 2500 m, 27.III.1992, T. Pyrcz ( CEP – MZUJ: female); Edo. Mérida, Monte Zerpa, Parc National La Culata , 2300 m, 25.IV.1992, T. Pyrcz ( CEP – MZUJ: female, gen. prep. 838/ J. Lorenc); Edo. Trujillo Qda el Tonto, Cachara–Humaca Alto km10, 1900 m, 10. XII. 2005 ( CB: female); Edo. Trujillo, Carache–Humocaro, Quebrada El Torito, 2000 m, 11.XII.2005, T. Pyrcz leg. ( CEP – MZUJ: female); Edo. Trujillo Escuque, Paramito, 1600 m, 15.IV.2006 ( CB: male); Edo. Mérida, Monte Zerpa, 2300 m, 5.II.2007, T. Pyrcz ( CEP – MZUJ: female); Edo. Aragua, Colonia Tovar–Puerto Maya km1–2, 2100–2200 m, 23.IV.2006 ( CB: male, female); Edo. Barinas, near San Isidro, 1400 m, 2.II.2008, Zs. Bálint and B. Benedek ( HNHM: male); Edo. Barinas, Barinitas-Santo Domingo, Minas de San Isidro , 1400–1450 m, 31.VII.2009, T. Pyrcz and A. Zubek ( CEP – MZUJ: female; gen. prep. 370/ J. Lorenc); Edo. Mérida, Estanques- Molino via Tusta, 1750 m, 26.III.2011, T. Pyrcz ( CEP – MZUJ: female) .

Redescription. Wings. FW costa length: 11–17mm (n=36, CEP – MZUJ material), apical area of costa and outer margin is not straight resulting in a somewhat rounded apex; male dorsal wing surface blue colouration bright blue; dorsal FW surface black border inner margin reaching the middle of vein M3; HW dorsal surface discal area structurally coloured; ventral wing surface medial pattern composed of light ruptive dashes; ventral FW androconia below vein Cubitus 2 absent but deep blue reflector present (cf. Faynel & Bálint 2012) ( Fig. 23H ); ventral FW submedian and median pattern running parallel; ventral HW submarginal pattern indistinct, postmedial region with submedial colouration, inner margin medial area with orange scaling. Genitalia. Male valva with rounded costal margin flip and slightly curved and pointed valval terminus (somewhat shorter than valval length), gnathos tip with a bulbous formation before pointed terminus. Female ductus with parallel edges, no lower plate, upper plate symmetric and roundish, no spines or bristles. No androconia.

Diagnosis. This species can be identified on the basis of the hind wing inner margin orange scalings in both sexes, and the lack of androconia. The male possesses ventral FW reflector. The similarly orange scaled J. turquisca sp. n. has a different (turquoise blue) male colouration, it has androconia but no reflector, HW ventral surface submarginal pattern is composed of gleaming scales, and the FW apex is pointed.

Distribution. Geographical: COLOMBIA (Bogotá, Caldas, Cali, Cauca, Cundinamarca, Manizales, Putumayo, Risaralda, Trujillo, Valle); ECUADOR (Bolívar, Morona–Santiago, Loja, Tungurahua, Zamora), PANAMA; PERU (Amazonas, Cusco, Huánuco, Pasco, San Martin); VENEZUELA (Barinas, Mérida) ( Fig. 31 ); spatial: recorded between elevations 600 and 3200 m a.s.l; temporal: recorded in all months, except June and November.

Note. This is the most widely distributed species of the genus ranging from Panama throughout the Venezuelan and Colombian mountain forests via Ecuador to southern Peru ( Lamas 2003) inhabiting tropical mountain forests. The Panama record given by Draudt (1919 –1921) is confirmed by the documentation of a recent photographic field-guide ( Glassberg 2018). In the NHM there are specimens from Bolivia, but these are old and insufficiently documented, most probably mislabelled. The Aragua ( Venezuela) and Cusco ( Peru) populations present some colour pattern differences. At present we do not have enough material for comparison to take more precise taxonomic decision.

Figures

FIGURE 1. Johnsonita male forewing venation. A: J. auda left forewing membrane dissected and digitised; B: J. turquisca right forewing documented under optical miscroscope (arrow indicates the subcubital wrinkle) (in same magnification, scale as indicated).

FIGURE 3. Johnsonita auda (Hewitson, 1867) imagines (left column: recto, right column: verso). A–B: male (Ecuador: Morona–Santiago), C–D: female (Venezuela: Mérida), E–F: male (Venezuela: Aragua), G– H: female (Venezuela: Aragua) (CEP– MZUJ specimens) (scale bar: 10 mm).

FIGURE 16. Johnsonita auda male genitalia armatures, entire organs in lateral (left column) and ventral (right column) views. A–B: Ecuador, Morona Santiago, Macas; C–D: Ecuador, Morona Santiago, San Juan Bosco; E–F: Ecuador, 9 de Octubre, Macas; G–H: Venezuela, Aragua (CEP–MZUJ specimens) (scale as indicated).

FIGURE 17. Johnsonita genitalia armatures, entire male organs in lateral (left column) and ventral (right column) views.A–B: J. auda, Ecuador, Morona-Santiago, 9 de Octubre-Macas; C–D: J. carpia, Ecuador, Napo, Res. Yanayacu; E–F: J. catadupa, Ecuador, Loja, Yangana; G–H: J. chlamydem, Peru, Pasco, Oxapampa, La Antena (CEP–MZUJ specimens) (scale as indicated).

FIGURE 20. Entire Johnsonita female organs in lateral (left column), ductus in lateral (central column) and ventral (right column) views. A–C: J. auda, Venezuela, Edo. Barinas, Minas de San Isidro; D–F: J. carpia, Ecuador, Prov. Carchi, Res. Forest Golondrinas; G–I: J. catadupa, Peru, Dept. Amazonas, Molinopampa, Tingo Ocol (CEP–MZUJ specimens) (scale as indicated).

FIGURE 23. Johnsonita mail trait ventral forewing subcubital scent patch indicated by arrow. A: J. carpia, B: J. catadupa, C: J. chlamydem, D: J. ianusca, E: J. johnsoni, F: J. pardoa, G: J. turquisca, H: J. zubkova.

FIGURE 24. Johnsonita mail traits in dorsal hindwing (left column) and ventral forewing subcubital patches (right column) indicated by arrow. A–B: J. oxalida, C–D: J. johnbanksi, E–F: J. subcunicula, G: J. auda.

FIGURE 26. Thecla auda Hewitson, 1867. NHMUK lectotype specimen and labels (scale bar: 10 mm).

FIGURE 31. Johnsonita distributions based on museum material examinded. J. auda = red circle, J. iacinta = blue diamond, J. subcunicula = green square, J. turquisca = yellow trinagle, J. zubkova = brown trinagle.