A cladistic analysis of the genus
The biodiversity on earth is so enormous that it would not be possible to study it, if it were not classified [
This genus is well known for producing an audible clicking sound, an ability that has been reported for many but not all species. It has been suggested that this sound is used in courtship, defense, and territoriality [
In contrast to most of other nymphalid butterflies, this genus perches on tree trunks with the wings open against the substrate and the head downward [
The confirmed host plants of all
At present,
223 specimens were analyzed:
The species
Male genitalia were treated with a 10% potassium hydroxide digestion process. To analyze them a stereomicroscope Nikon model 1064565 and dissection scissors where employed, then the material was stored in glycerol vials in the Insect Museum of the University of Costa Rica.
The character matrix is composed of 66 characters (all binary), of which 41 were derived from the external morphology and color pattern of the wings, 2 from the color pattern of the body, 2 from the spiral organ morphology inside the fore-wing subcostal vein, and 21 from the male genitalia. Also, three phylogenetic attributes where mapped on the resulting tress: one ethological, one biogeographical, and one physiological.
The terminology of the adult external morphology follows Scoble [
The first three analyses were done to test if
The identification key to the species was prepared using characters that are easy to distinguish with the naked eye butterfly specimen. The key follows a different logic of characteristics than the one employed by Jenkins [
The analyses show the phylogenetic relationships obtained between these four closely related genera using
Majority rules consensus compromise from two primary trees obtained from a heuristic analysis of 66 characters for 23 species (tree length =168, CI = 0.39, RI = 0.63). Numbers above tree branches represent consensus compromises.
Single tree obtained from a heuristic analysis of 66 characters for 23 species (tree length = 161, CI = 0.40, and RI = 0.65).
Majority rules consensus tree obtained from two primary trees from a heuristic analysis of 66 characters for 23 species (tree length =168, CI = 0.39, RI = 0.63). Numbers above tree branches represent consensus compromises.
The first analysis used
Single tree obtained from a heuristic analysis of 66 characters for 21 species (tree length = 126, CI = 0.50, and RI = 0.69).
Trees in Figures
Biogeography of the species mapped on the tree obtained from analyses of the Figure
Palatability of the species mapped on the tree obtained from analyses of the Figure
Human audible click noise production mapped on the tree obtained from analyses of the Figure
The characters found to form the different species groups (Figure
Characters justifying the grouping of species and genera. WinClada ver. 1.00.08 was used to map character changes on the three in Figure
Clade 1: ( |
(2 : 1) Absence of two submarginal ocelli on DHW |
(22 : 1) Absence of submarginal ocelli on DHW |
Clade 2: ( |
(3 : 1) Existence of sex patch on DHW |
(4 : 1) Presence of red submarginal spots in VHW |
(18 : 1) Ventral wings with blue iridescent reflectance |
(29 : 1) Existence of red spots around humeral vein in VHW |
(33 : 1) Presence of white in dorsal forewings of females only in the diagonal medial band |
(42 : 1) Red markings on ventral area of thorax |
Clade 3: ( |
(7 : 1) Forewings with M3-Cu2 veins extended more than the others |
(8 : 1) DFW with two red maculae inside discal cell |
Clade 4: ( |
(44 : 0) Spiral organ on forewings sub coastal vein short and thick |
(51 : 1) Vinculum of male genitalia does not have anterior projection |
Clade 5: ( |
(46 : 1) Distal point of the arm of the gnathos pointing to the uncus |
Clade 6: ( |
(56 : 1) Uncus overlaps with clasper |
Clade 7: ( |
(35 : 0) Absence of double concentric ocelli on DHW from Cu1 to Cu2 |
Clade 8: ( |
(49 : 1) Point of the arm of the gnathos not acute |
(50 : 1) Arm of the gnathos finishing in a thick triangle |
(64 : 0) Posterior point of the saccus thicker than the rest |
Clade 9: ( |
(39 : 1) No sexual dimorphism present in the way that discal cell close on the Forewings |
Clade 10: ((( |
(15 : 1) Existence of color white or gray inside DFW discal cell |
(20 : 1) Hindwings distal border serrated |
(21 : 1) VHW with white marginal spots between veins |
(25 : 1) VHW submarginal ocelli intercalated large ones with smaller ones |
(35 : 1) DHW with two or three concentric ocelli in sub-margins from Cu1 to Cu2 |
(36 : 1) Presence of a depression followed by an outward curve on the costal vein at the end of the discal-cell of forewings |
(37 : 1) Females without diagonal wide and regular white band on DFW |
Drawings of different structures which are important for the interpretation of the species key (taxonomic characters) are shown in Figures
Hypothetical
Three different patterns in the distal area of the discal cell employed in the
Following a single cladistic methodology, four different analyses produced four similar trees, although, some important dissimilarities can be pointed out in order to understand the evolutionary relationships of the groups. The CI (Consistency Index) of all the resulting trees is between 0.39 and 0.50 which is considered appropriate under the scenario presented by [
The species groups supported by the strongest evidence based on the results of this paper (because they are identical in the four analyses) are as follows.
The diagnosis of
Consistently, all the trees obtained in this study grouped these three species apart from all others, and when other related genera (
Under the scenario where
Another well-supported species group is the one composed by
All these species are distinguished by having orange color of the ventral surface of the hind wings, a character probably lost as an autapomorphy in
These two species are the sister group of the last species group. They evolved from a continental ancestor which was distributed throughout practically all of the Neotropical region.
The close relation between
The analyses shown in Figures
In all the analyses done in this work,
Jenkins [
Also, in Figure
More phylogenetic analyses in which early-stage characters are included, in addition to molecular analyses, may provide more evidence as to whether
The tree in Figure
This analysis is the most parsimonious of all the analyses done because it is the one in which the early stages and host plants of
The palatability to predators in the genus
The capacity of the studied species to produce human audible clicking noise is also mapped. This ethological attribute is consistent with the unpalatability one. The basal taxa:
The key presented in this paper uses new characters obtained from the careful analysis of the color patterns which helps in the identification of the species; some of the characters are taken from the key presented by Jenkins [
(a) Submarginal ocelli in DHW from M3-Cu2 circular or tear shaped—2 (b) Submarginal ocelli in DHW from M3-Cu2 absent—19 (a) Submarginal ocelli in DHW enclosing only one blue spot—3 (b) Submarginal ocelli in DHW enclosing other ocelli or moon-shaped spots of any color—6 (a) VFW with a white spot or macula in R3-R4 anterior of the bifurcation of R4-R5—4 (b) VFW with no white spot or macula in R3-R4 just anterior of the bifurcation of R4-R5— (a) DFW with a white wide median band—5 (b) DFW without the white wide median band— (a) DFW in Cu1-Cu2 with the white macula of the median band enlarged until reaching the point of the bifurcation of M3-Cu1— (b) DFW in Cu1-Cu2 with the white macula of the median band finishing two mm distally from the point of the bifurcation of M3-Cu1— (a) VHW with Submarginal ocelli absent in M2-M3— (b) VHW with Submarginal ocelli or white point present in M2-M3—7 (a) VHW with four consecutive ocelli of different sizes from M1 to Cu2—8 (b) VHW with four consecutive ocelli of similar size from M1 to Cu2—12 (a) VHW ocelli from M1 to Cu2, the one in M2-M3 absent or if present then less than a half the diameter of the one in Cu1-Cu2—9 (b) VHW ocelli from M1 to Cu2, the anterior one large, and the next three small, reduced to white points or absent—11 (a) VHW in M2-M3 with a small ocelli (sometimes rudimentary) and without a white marginal macula, but if present never as large as the ocelli—10 (b) VHW in M2-M3 with a small ocelli and a white marginal macula as large or larger than the ocelli— (a) DFW with three rose-red bars or bands on proximal half— (b) DFW without three rose-red bars or bands on proximal half— (a) DHW ocellus with a black external ring— (b) DHW ocellus with the external ring any color but black— (a) VHW ocelli composed of a black ring with light center—13 (b) VHW ocelli composed of a complete of incomplete rust-red ring, surrounding a black-crescent-shaped moon with white center—16 (a) DFW with a white Submarginal ocellus in Cu1-Cu2 all surrounded by black—14 (b) DFW without any a white Submarginal ocelli in Cu1-Cu2 all surrounded by black—15 (a) VFW in R3-R4 with the white Submarginal spot as large as or larger than the marginal spot— (b) VFW in R3-R4 without the white Submarginal spot, but if present then not larger than the marginal spot— (a) DHW ocelli form M3 to Cu2 circular or somewhat tear shaped— (b) DWH ocelli from M3 to Cu2 dorsoventrally compressed— (a) DHW ocelli in M3-Cu2 dorsoventrally compressed, with a white internal line reaching more than the 70% of the width between the veins— (b) DHW ocelli in M3-Cu2 not dorsoventrally compressed, with a white internal line reaching no more than the 50% of the width between the veins—17 (a) DFW showing distal cross bar of discal cell completely separated in two clear bands by a thin dark line from level of Radial vein to level of Cubital vein. Figure (b) DFW showing distal cross bar of discal cell not completely separated in two clear bands from level of Radial vein to level of Cubital vein (Figures (a) DFW showing distal cross bar in M2-M3 width, with a constriction at the origin of M2, and becoming width again in R5-M1 (Figure (b) DFW showing distal cross bar in M2-M3 wide, and finishing at the origin of M2, and not becoming wide again in R5-M1 (Figure (a) VHW with red bars or spots in anal cell— (b) VHW without red bars or spots in anal cell—20 (a) DHW with reddish anterior costal markings— (b) DWH with no reddish anterior costal markings— (a) DFW in M2-M3 with only two blue spots— (b) DFW in M2-M3 with four or five blue spots—22 (a) DHW in Rs-M1 with three blue spots— (b) DHW in Rs-M1 with two blue spots—
Costa Rica: Guapiles, Q. Molinete, 400 m. 5-X-2006, col. per. Santa Clara, 400 m. Rec: T. Assmann, col. per. Heredia, Horquetas, 600 m., 8-VIII-1993, rec. I. Chacón, MNC. Heredia, Horquetas, 600 m., 11-VIII-1993, rec. I. Chacón, MNC. Limón, Guápiles, 100 m., 30-III-1985, rec. A. Solís, MNC. Puntarenas, Carara, 50 m. 29-V-1990, rec. I. Chacón, MNC. Puntarenas, Golfito, 200 m., 18-II-1982, rec. R. Canet, MNC. Limón, Guápiles, 100 m., 26-III-1978, rec. P. J. DeVries, MNC. Heredia, Horquetas, 600 m., 2-V-1993, rec. I. Chacón, MNC. Heredia, Horquetas, 600 m., 21-VI-1993, rec. I. Chacón, MNC. San José, Mora, El Rodeo, 700 m., 11-VII-1982, rec. R. Canet, MNC. Guanacaste, Volcán Santa Maria, 800 m., 8-VII-1978 rec. P. J. DeVries, MNC. San José, P. N. Braulio Carrillo, 700 m., 6-VII-1989, rec. R. Canet, MNC.
Peru: Loreto, Iquitos, 100 m. Ene. 2001, col. per. Loreto, Iquitos, 100 m. Ene. 2001, col. per.
Costa Rica: Guanacaste, La Cruz, Santa Cecilia, 300 m., 18-VI-2009, rec. G. Vega, MNC. Guanacaste, La Cruz, Santa Cecilia, 300 m., 18-VI-2009, rec. J. Solano, MNC, Alajuela, Florencia, 6-VI-1995, rec. W. Haber, MNC. Guanacaste, La Cruz, Santa Cecilia, 300 m., 18-VI-2009, rec. J. Solano, MNC. Sin Datos, col. per.
Peru: Manu, Madre de Dios, Feb. 2001, col. per. Manu, Madre de Dios, Feb. 2001, col. per.
Venezuela: Patanemo, 50 m. Ene. 2006, Rec: E. Lorenzoni col. per. Patanemo, 50 m. Ene. 2006, Rec: E. Lorenzoni col. per. Patanemo, 50 m. Ene. 2006, Rec: E. Lorenzoni col. per. Patanemo, 50 m. Ene. 2006, Rec: E. Lorenzoni col. per. Pijiguaos, 700 m. Oct. 2005, Rec: E. Lorenzoni col. per. Pijiguaos, 700 m. Dic. 2005, Rec: E. Lorenzoni col. per. Costa Rica: Puntarenas, Río Barú, 29-XII-1998, Rec: L. R. Murillo-Hiller, col. per. Puntarenas, Villa Nelly, 500 m., 11-X-1976, rec. P. J. DeVries, MNC. Puntarenas, Carara, 50 m. 1-X-1983, rec. R. Canet, MNC. San José, P. N. Braulio Carrillo, 700 m., 10-IV-1984, rec. I. Chacón, MNC. San José, P. N. Braulio Carrillo, 700 m., 20-V-1984, rec. I. Chacón, MNC. Limón, Guápiles, 100 m., 21-V-1976, rec. P. J. DeVries, MNC. Alajuela, San Miguel, 450 m., 13-XII-1987, rec. R. Canet, MNC. Heredia, La Selva, 50 m., 25-VIII-1979, Rec. P. J. DeVries, MNC. San José, P. N. Braulio Carrillo, 700 m., 1-VI-1984, rec. A. Chacón, MNC. San José, P. N. Braulio Carrillo, 700 m., 10-IV-1984, rec. A. Chacón, MNC. Puntarenas, Carara, 50 m. 1-X-1983, rec. I. Chacón, MNC. Heredia, Horquetas, 600 m., 28-IV-1985, rec. I. Chacón, MNC. Heredia, Horquetas, 600 m., 18-III-1993, rec. I. Chacón, MNC. Alajuela, San Carlos, 150 m., 21-VIII-1983, rec. E. Corea, MNC. Puntarenas, Corcovado, 0 m., 7-VI-1986, rec. I. Chacón, MNC. San José, Mora, El Rodeo, 700 m., 11-X-1998, rec. P. Gloor, MNC. Puntarenas, Aguirre, Savegre, 100 m., 20-II-2002, rec. G. Vega, MNC. Puntarenas, Corcovado, 0 m., 17-I-1986, rec. R. Canet, MNC. San José, P. N. Braulio Carrillo, 700 m., 24-IV-1984, rec. A. Chacón, MNC. San José, P. N. Braulio Carrillo, 700 m., 15-V-1984, rec. A. Chacón, MNC. San José, P. N. Braulio Carrillo, 700 m., 25-IV-1984, rec. A. Chacón, MNC. Heredia, Horquetas, 600 m., 12-IV-1994, rec. J. Solano, MNC. San José, Puriscal, 500 m., 18-I-1979, rec. F. Wolg, MNC. Heredia, Magsaysay, 150 m., 15-VIII-1981, rec. R. Canet, MNC. San José, P. N. Braulio Carrillo, 700 m., 2410-V-1984, rec. A. Chacón, MNC. San José, P. N. Braulio Carrillo, 700 m., 27-V-1984, rec. A. Chacón, MNC. Heredia, Horquetas, 600 m., 25-V-1993, rec. I. Chacón, MNC. Heredia, Horquetas, 600 m., 22-V-1993, rec. I. Chacón, MNC. Heredia, Horquetas, 600 m., 18-III-1993, rec. I. Chacón, MNC. Heredia, Magsaysay, 150 m., 8-VIII-1981, rec. R. Hesterberg, MNC. San José, P. N. Braulio Carrillo, 700 m., 10-IV-1984, rec. A. Chacón, MNC. Puntarenas, Corcovado, 0 m., 30-IV-1985, rec. R. Canet, MNC. San José, P. N. Braulio Carrillo, 700 m., 14-IV-1984, rec. A. Chacón, MNC. Puntarenas, Corcovado, 0 m., 23-VI-1986, rec. I. Chacón, MNC. San José, Mora, El Rodeo, 700 m., 25-VII-1997, rec. J. Solano, MNC. San José, Mora, El Rodeo, 700 m., 30-VIII-1998, rec. P. Gloor, MNC. San José, Dota, 800 m., 22-VII-1998, rec. G. Vega, MNC. San José, Mora, El Rodeo, 700 m., 2-VII-1997, rec. G. Vega, MNC.
Brazil.
Brazil: Pará. Pará, Obidos, XI-1992, Manuel Ortíz col. per. VENEZUELA: Pijiguaos, 700 m. Feb. 2009, E. Lorenzoni col. per.
Costa Rica: Heredia, Horquetas, 600 m., rec. I. Chacón, MNC. Limón, Bordón, 500 m., 27-IV-1995, rec. G. Vega y A. Valerio, MNC. Heredia, Sarapiquí, 50 m., 12-VII-1981, rec. R. Canet, MNC. San José, P. N. Braulio Carrillo, 500 m., 4-VI-1980, rec. P. J. DeVries, MNC. Heredia, Sarapiquí, 50 m., 12-VII-1981, rec. R. Canet, MNC. Limón, Guápiles, 400 m., 28-IV-1988, rec. I. Chacón, MNC.
COSTA RICA: Guanacaste, Rincón de la Vieja, 1100 m. 16-IX-1997, Rec: L. R. Murillo-Hiller. Alajuela, La Gúacima, 800 m., 27-IX-2008, Rec: L. R. Murillo-Hiller. Guanacaste, Rincón de la Vieja, 1100 m. 16-IX-1997, Rec: L. R. Murillo-Hiller. Guanacaste, Rincón de la Vieja, 1100 m. 16-IX-1997, Rec: L. R. Murillo-Hiller. Guanacaste, Liberia, 150 m. 6-VII-1999, Rec: L. R. Murillo-Hiller. San José, Mora, El Rodeo, 700 m., 31-ago-1994, rec. G. Vega y J. Solano, MNC. Guanacaste, Santa Rosa, 100 m., 15-VI-1978, rec. P. J. DeVries, MNC. Guanacaste, Santa Rosa, 100 m., 1-XII-1976, rec. P. J. DeVries, MNC. Puntarenas, Quepos, 50 m., 27-II-1982, rec. R. Hesterberg, MNC. Guanacaste, Palo Verde, 50 m., 30-I-1982, rec. I. Chacón, MNC. Guanacaste, Santa Rosa, 100 m., 3-XII-1979, rec. D.H. Janzen, MNC. Alajuela, Atenas 800 m., 27-XI-1980, rec. R. Canet, MNC. Guanacaste, Santa Rosa, 100 m., I-XII-1976, rec. P. J. DeVries, MNC. Guanacaste, Palo Verde, 50 m., 30-I-1982, rec. I. Chacón, MNC. San José, Dota, 800 m., 22-XII-1998, rec. G. Vega, MNC. Guanacaste, Palo Verde, 50 m., 30-I-1982, rec. I. Chacón, MNC. Guanacaste, Santa Rosa, 100 m., 5-VII-1978, rec. P. J. DeVries, MNC. Guanacaste, Santa Rosa, 100 m., 5-VII-1978, rec. P. J. DeVries, MNC. Guanacaste, Palo Verde, 50 m., 30-I-1982, rec. I. Chacón, MNC. Guanacaste, Palo Verde, 50 m., 30-I-1982, rec. I. Chacón, MNC. Puntarenas, Barranca, 200 m., 26-XII-1984, rec. R. Canet, MNC. Puntarenas, Isla San Lucas, 0 m., 14-IX-2005, rec. G. Vega, MNC. Puntarenas, Isla San Lucas, 0 m., 14-IX-2005, rec. G. Vega, MNC. Puntarenas, Barranca, 200 m., 23-II-1982, rec. R. Canet, MNC. Puntarenas, Barranca, 200 m., 21-I-1984, rec. R. Canet, MNC. Guanacaste, Santa Rosa, 100 m., 4-XII-1976, rec. P. J. DeVries, MNC. Guanacaste, Palo Verde, 50 m., 30-I-1982, rec. I. Chacón, MNC. Guanacaste, Cerro el Hacha, 18-IX-1987, rec. I. Chacón, MNC. Guanacaste, Palo Verde, 50 m., 30-I-1981, rec. R. Campos, MNC. Guanacaste, Palo Verde, 50 m., 30-I-1981, rec. R. Campos, MNC.
Costa Rica: Guanacaste, Liberia, 150 m., 7-VII-1999, Rec: L. R. Murillo-Hiller. Guanacaste, Liberia, 150 m., 6-VII-1999, Rec: L. R. Murillo-Hiller.
Brazil: Maranhao, Sao Luis, 12-VIII-1961, col. per.
Bolivia: Cochabamba, Limbo, 2000 m. Dic. 2008 col. per. Cochabamba, Limbo, 2000 m. Dic. 2008 col. per. Cochabamba, Limbo, 2000 m. Dic. 2008 col. per. Cochabamba, Limbo, 2000 m. Dic. 2008 col. per. ARGENTINA: Misiones, Dorado, 18-II-1996, M. Ortiz col. per.
Costa Rica: Alajuela, La Guácima, 800 m. VII-2008 col. per. Alajuela, La Guácima, 800 m. IX-2008 col. per. San José, Mora, El Rodeo, 19-VI-2009 col. per. San José, Mora, El Rodeo, 19-VI-2009 col. per. Puntarenas, San Vito, 1150 m., 19-IX-1977, rec. G. B. Small, MNC. Alajuela, Atenas 800 m., 17-XII-1977, rec. P. J. DeVries, MNC. Puntarenas, San Vito, 1150 m., 17-III-1978, rec. P. J. DeVries, MNC. Puntarenas, Coto Brus, 1500 m., 11-II-1991, rec. A. Solís, MNC. Heredia, San Rafael, 1350 m., 20-VII-1981, rec. R. Canet, MNC. Puntarenas, Coto Brus, 1500 m., 2-II-1991, rec. H. Sparrow, MNC. San José, Mora, El Rodeo, 700 m., 10-VIII-1982, rec. R. Canet, MNC. Puntarenas, Coto Brus, 1500 m., 9-X-1995, rec. I. Chacón, MNC. San José, Acosta, Palmichal, 1400 m., 20-XII-2003, rec. M Alfaro, MNC. Puntarenas, Santamaría de Pittier, 6-IX-1984, rec. A. Solís, MNC. San José, Mora, El Rodeo, 700 m., 5-V-1997, rec. G. Vega y J. Solano, MNC. Puntarenas, San Vito, 1150 m., 27-V-1985, rec. R. Canet, MNC. Alajuela, Atenas, 800 m., 17-XII-1977, rec. P. J. DeVries, MNC. San José, Mora, El Rodeo, 700 m., 9-III-1996, rec. G. Vega, MNC. San José, Mora, El Rodeo, 700 m., 2-IX-1996, rec. G. Vega y J. Solano, MNC. San José, Mora, El Rodeo, 700 m., 19-VII-1995, rec. G. Vega, MNC. San José, Mora, El Rodeo, 700 m., 2-IX-1997, rec. G. Vega, MNC. San José, Mora, El Rodeo, 700 m., 19-XII-1995, rec. G. Vega, MNC. San José, Mora, El Rodeo, 700 m., 2-X-1995, rec. G. Vega, MNC. San José, Acosta, Palmichal, 1400 m., 22-IX-2003, rec. M Alfaro, MNC.
Costa Rica: Alajuela, La Guácima, 800 m., VII-2008 col. per. Alajuela, La Guácima, 800 m., VII-2008 col. per. San José, Mora, El Rodeo, 700 m., 31-ago-1994, rec. G. Vega y J. Solano, MNC. Puntarenas, Carara, 50 m. 28-May-1983, rec. R. Hesterberg, MNC. San José, Puriscal, 500 m., 18-May-1979, rec. P. J. DeVries, MNC. San José, Mora, El Rodeo, 700 m., 3-ago-1978, rec. P. J. DeVries, MNC. San José, Mora, El Rodeo, 700 m., 7-jul-1988, rec. I. Chacón, MNC. San José, Mora, El Rodeo, 700 m., 11-VII-1982, rec. R. Canet, MNC. Puntarenas, Aguirre, Savegre, 100 m., 15-I-2002, rec. G. Vega, MNC. San José, Mora, El Rodeo, 700 m., 19-V-1995, rec. G. Vega y A. Valerio, MNC. VENEZUELA: Pijiguaos, 700 m., II 2009 E. Lorenzoni col. per.
Venezuela: Caura, Ago-2004, E. Lorenzoni col. per.
Venezuela: Pijiguaos, 700 m., Dic. 2005 E. Lorenzoni col. per. Caura, Ago-2004, E. Lorenzoni col. per. Sta. Elena, Vía Paují, 1000 m. Oct. 2002, E. Lorenzoni col. per. BOLIVIA: Cochabamba, Limbo, 2000 m., Ene-2008. Cochabamba, Limbo, 2000 m., Ene-2008. BRASIL: Pará, Obidos, XI-1992, M. Ortíz col. per.
Costa Rica: Alajuela, La Guácima, 800 m. Jul. 2008, rec. L. R. Murillo-Hiller. Guanacaste, Rincón de la Vieja, 800 m., 8-VIII-2008, rec. L. R. Murillo-Hiller. Puntarenas, Corcovado, 0 m., 19-XI-1976, rec. P. J. DeVries, MNC. Heredia, La Selva, 50 m., 1-VI-1970, Rec. P. J. DeVries, MNC. San José, Mora, El Rodeo, 700 m., 7-X-1984, rec. R. Canet, MNC. Guanacaste, Santa Rosa, 100 m., 6-VII-1979, rec. D. H. Janzen, MNC. San José, Mora, El Rodeo, 700 m., 15-VII-1998, rec. G. Vega, MNC. Alajuela, Grecia, 900 m., 23-XII-1983, rec. R. Campos, MNC. Guanacaste, Santa Rosa, 100 m., 22-VI-1978, rec. P. J. DeVries, MNC.
Colombia: Bojará, Muzo, Ene. 1993. M. Ortiz col. per.
Brazil: Maranhao, Sao Luis, 12-VIII-1961, col. per. Sta. Catarina, Bombinhas, 20-IX-2003, rec: L. R. Murillo-Hiller. Pará, Obidos, XI-1992, M. Ortíz col. per. VENEZUELA: Hda. Bucarito, Montalban, 700 m., 15-V-1983, rec: L. D. Otero. Urb. Guapazo, Valencia, 500 m., 20-V-1983, rec: L. D. Otero. Urb. Guapazo, Valencia, 500 m., 20-V-1983, rec: L. D. Otero. Pijiguaos, 700 m., feb 2009, E. Lorenzoni col. per. Caura, Km. 20, Mar-2003, E. Lorenzoni col. per.
Costa Rica: San José, Mora, El Rodeo, 800 m., 19-VI-2009, rec: L. R. Murillo-Hiller. Guanacaste, Liberia, 100 m. 1968, rec: P. Kazan. Alajuela, La Guácima, 2008, rec: L. R. Murillo-Hiller. Alajuela, La Guácima, 2008, rec: L. R. Murillo-Hiller. Alajuela, La Guácima, 2008, rec: L. R. Murillo-Hiller. Alajuela, La Guácima, 2008, rec: L. R. Murillo-Hiller. Guanacaste, Liberia. 100 m., 14-IX-1980, rec: Y. Chavarri, col: MIUCR.
Puntarenas, Corcovado, 0 m., 28-VI-1986, rec. I. Chacón, MNC. Puntarenas, Aguirre, Savegre, 100 m., 20-II-2002, rec. G. Vega, MNC. Puntarenas, Corcovado, 0 m., 27-I-1980, rec. R. Canet, MNC.
Costa Rica: Guanacaste, Liberia, 100 m. rec: T. Assmann. Guanacaste, Liberia, 100 m., 13-V-1980, col. MIUCR. Puntarenas, Barranca, 200 m., 27-II-1982, rec. R. Canet, MNC. Puntarenas, Barranca, 200 m., 5-VII-1981, rec. R. Canet, MNC. Guanacaste, Palo Verde, 50 m., 30-I-1982, rec. I. Chacón, MNC. Guanacaste, Río Corobicí, 90 m., 7-V-1976, rec. P. J. DeVries, MNC. San José, Tibás, 1200 m., 12-XII-1977, rec. P. J. DeVries, MNC. Guanacaste, Santa Rosa, 100 m., 27-VI-1977, rec. P. J. DeVries, MNC. Guanacaste, Palo Verde, 50 m., 30-I-1982, rec. I. Chacón, MNC. Guanacaste, Santa Rosa, 100 m., 12-VI-1977, rec. P. J. DeVries, MNC. Guanacaste, Palo Verde, 50 m., 30-I-1982, rec. I. Chacón, MNC. Guanacaste, Santa Rosa, 100 m., 26-VI-1978, rec. P. J. DeVries, MNC. Heredia, San Rafael, 1250 m., 25-VIII-1981, rec. R. Canet, MNC.
Costa Rica: San José, Mora, El Rodeo, 1995, rec: L. R. Murillo-Hiller. San José, Mora, El Rodeo, 1995, rec: L. R. Murillo-Hiller. Alajuela, La Guácima, 800 m., 13-IX-2008, rec: L. R. Murillo-Hiller. Alajuela, La Guácima, 800 m., 13-IX-2008, rec: L. R. Murillo-Hiller. Guanacaste, Rincón de la Vieja, 800 m. 8-VIII-2008, rec: L. R. Murillo-Hiller. Alajuela centro, 800 m., 19-XI-1986, rec: A. Ureña. Guanacaste, Rincón de la Vieja, 800 m. 16-VII-1997, rec: L. R. Murillo-Hiller. San José, Escazú, 1300 m., 3-Ago-1978 L. R. Murillo-Hiller col. per. Alajuela, La Guácima, 800 m., 10-VIII-2008, rec: L. R. Murillo-Hiller. Puntarenas, Sta. Teresa, 0 m., 21-XIII-2008, L. R. Murillo-Hiller col. per. Alajuela, La Guácima, 800 m., Jul-2008, rec: L. R. Murillo-Hiller. Alajuela, La Guácima, 800 m., Jul-2008, rec: L. R. Murillo-Hiller. BRASIL: Rio Grande do Sul, Porto Alegre, 22-IV-1940. Pará, Obidos, XI-1992, M. Ortíz col. per. VENEZUELA: Pijiguaos, 400 m., 2005, E. Lorenzoni col. per. Urb. Guapazo, Valencia, 500 m., 28-V-1983, rec: E. Carabobo. COLOMBIA: Boyacá, Muzo, I-1993, M. Ortiz col. per. Guanacaste, Santa Rosa, 100 m., 5-VII-1976, rec. P. J. DeVries, MNC. Puntarenas, Barranca, 200 m., 27-II-1982, rec. R. Canet, MNC. Guanacaste, Palo Verde, 50 m., 30-I-1982, rec. I. Chacón, MNC. Guanacaste, Santa Rosa, 100 m., 1-XII-1976, rec. P. J. DeVries, MNC.
Brazil: Rio Grande do Sul, Porto Alegre, X-2003, rec: L. R. Murillo-Hiller. BOLIVIA: Cochabamba, Limbo, Ene-2008, L. R. Murillo-Hiller col. per.
See Table
Predominant dorsal wings color any except black (0), black (1). Presence of at least two submarginal ocelli on DHW (0), absence of ocelli (1). Absence of sex patch on DHW (0), existence of such structure (1). Absence of red submarginal spots on VHW (0), presence of such pattern (1). Spots or maculae inside the discal cell of VHW (0), devoid of such a pattern (1). DHW submarginal ocelli without red-rust scales (0), presence of such scales (1). Hind wings veins from M3-Cu2 not longer than the rest (0), such veins longer than the rest (1). DFW without two red maculae inside the discal cell (0), with such a pattern (1). VHW without red maculae inside the discal cell (0), with such a pattern (1). Female with a wide diagonal white band on DFW (0), devoid such a pattern (1). Male with a wide diagonal white band on DFW (0), devoid such a pattern (1). Wings color no sexual dimorphic (0), wings color sexually dimorphic (1). On males Forewings the discal cell closes right on the bifurcation of M3-Cu1 (0), the discal cell close anteriorly of such bifurcation (1). Forewing apex closer to the body than distal border (0), Forewing apex more distant to the body than distal border (1). Absence of white or gray inside the discal cell of DFW (0), presence of such a pattern (1). Females DFW with turnus macula not split in two (0), such macula completely divided in two (1). DFW discal cell solid color or with dots (0), with lines or maculae (1). Ventral wings without iridescent reflectance (0), with iridescent reflectance (1). Females DFW with a white submarginal macula from R5 to M1 (0), devoid of such a pattern (1). Hing wings distal border not serrated (0), serrated (1). Without a white spot on distal border of the VHW between each vein (0), with such a spot (1). Presence of submarginal ocelli in DHW (0), devoid of such a pattern (1). Absence of tear shaped submarginal ocelli on DHW (0), presence of such a pattern (1). Presence of dorsoventrally compressed submarginal ocelli on DHW (0), absence of such a pattern (1). VHW submarginal ocelli all similar size (0), presenting small ones between big ones (1). VHW not presenting proximal half orange-red color (0), presenting orange-red color (1). Absence of a white marginal macula on VFW from m2 to m3 and another from cu1 to cu2 (0), presence of both macula (1). Absence of a white submarginal macula form r3 to r4 on DFW (0), presence of such a macula (1). Absence of red spots at the side of the humeral vein on VHW (0), presence of such spots (1). Absence of a row of submarginal ocelli on DHW (0), presence of that pattern (1). Females presenting a submarginal white macula on VFW from r5 to m1 (0), devoid of such a pattern (1). Absence of a proximal red spot on VHW from Sc + r1 to r5 (0), presence of such a spot (1). Females present white coloration in other parts of the DFW besides the white diagonal band (0), devoid of such a pattern (1). Presence of a submarginal ocelli on DHW from m2-Cu2 (0), devoid of such a pattern (1). Absence of a double concentric ocelli on DHW from Cu1 to Cu2 (0), presence of such ocelli (1). Absence of a coastal depression at the distal end of the discal cell followed by a bump on the Forewings (0), presence of such a depression (1). Females DFW with the diagonal white band not fragmented (0), diagonal white band fragmented in many parts or absent (1). Forewing with distal border straight (0), distal border convex (1) (Modification of character no. 1 of [ No sexual dimorphism present in the way that discal cell close on the Forewings (0), sexual dimorphism present in such a character (1). Cryptic coloration on VHW (0), aposematic (1). Rammi presented (0), rammi absent (1). Ventral thorax sides without red markings (0), with red markings (1). Ventral thorax not completely orange (0), completely orange (1) (Modification of character no. 24 of [ Spiral organ on Forewings short and thick (0), long and thin (1) [ Absence of the spiral organ (0), presence of the spiral organ (1). Distal point of the arm of the gnathos do not point to the distal point of the uncus (0), distal point of the arm of the gnathos pointing to the uncus (1). Arm of the gnathos as thick or less than the uncus (0), thicker than the uncus (1). Arm of the gnathos with elbow of more than 90° (0), less than 90° (1). Point of the arm of the gnathos acute (0), devoid of such a pattern (1). Arm of the gnathos finishing in any shape except a thick triangle (0), finishing in a thick triangle (1). Vinculum with an anterior projection (0), devoid of such a pattern (1). Anterior projection of vinculum closer to the saccus than to the tegumen (0), closer to the tegumen (1). Anterior projection of vinculum round pointing (0), acute (1). Uncus without setae dorsally (0), with long setae (1) ([ Uncus same long or longer than clasper (0), shorter than clasper (1). Uncus do not overlaps with clasper (0), overlapping (1). Medial part of the uncus as thick as the posterior part (0), less thick (1). Uncus without dorsal depression (0), with depression (1). Valva without dorsal callus (0), with dorsal callus (1). Valva twice thicker than the saccus (0), valva three times thicker than the saccus (1). Valva length less than a half of the length of the saccus (0), valva length longer than a half of the saccus (1). Saccus longer than the rest of the genitalia (0), same or shorter than the rest of the genitalia (1). Saccus as thick or less than the arm of the gnathos (0), saccus thicker (1). Posterior point of the saccus thicker than the rest (0), devoid of such a pattern (1). In lateral view, anterior portion of tegumen extremelly proiected (0), devoid of such a pattern (1) ([ Anterior portion of tegument less or same long than the projection of the vinculum (0), longer than the projection of the vinculum (1).
See Figures
See Figures
The author thanks James Coronado for important discussion of the paper, J. Monge-Nájera and Paul Hanson for providing helpful literature and also for providing critical comments and a review of the paper, Enrique Lorenzoni and Manuel Ortiz for providing important specimens from their collections, Axel Retana for sharing methodology comments, and German Vega for allowing access to the Museo Nacional de Costa Rica collection.