Diversidade de padrão das asas em Brassolini (Nymphalidae, Satyrinae)

Penz, Carla Maria; Mohammadi, Neda

doi:10.1590/S1676-06032013000300020

Figuras (9)
Tabelas (2)

Figure 1. Male wings in ventral view plus a schematic drawing showing color-coded pattern elements. Scale bars: 1 cm. A, Bia actorion, Suriname; B, Narope cyllastros, no data; C, Brassolis sophorae, French Guiana; D, Aponarope sutor, Brazil, Rondônia; E, Dynastor napoleon, Brazil, Santa Catarina; F, Dynastor macrosiris macrosiris, French Guiana.

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Figure 2. Male wings in ventral view plus a schematic drawing showing color-coded pattern elements. Scale bars: 1 cm. A, Dasyophthalma rusina, South Brazil; B, Opoptera syme, Brazil, São Paulo; C, Caligopsis seleucida, Peru, Madre de Dios; D, Opoptera aorsa, Brazil, Paraná; E, Eryphanis automedon, Ecuador, Sucumbios; F, Eryphanis bubocula, Costa Rica, Guanacaste.

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Figure 3. Male wings in ventral view plus a schematic drawing showing color-coded pattern elements. Scale bars: 1 cm. A, Caligo eurilochus, Ecuador, Sucumbios; B, Caligo atreus, Colombia, Antiochia; C, Caligo martia, Brazil, Santa Catarina; D, Selenophanes cassiope, Peru, Chanchamayo; E, Penetes pamphanis, Brazil, Paraná; F, Catoblepia xanthus, Brazil, Pará.

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Figure 4. Male wings in ventral view plus a schematic drawing showing color-coded pattern elements. Scale bars: 1 cm. A, Catoblepia orgetorix championi, Panama, Chiriqui; B, Catoblepia berecynthia, Central Peru; C, Mielkela singularis, Mexico, Chiapas; D, Orobrassolis ornamentalis, Brazil, São Paulo; E, Blepolenis bassus, Brazil; F, Opsiphanes sallei, Peru, Pasco.

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Figure 5. Wings in dorsal view (except for A, left side) indicating color-coded pattern elements and trailing bands. Mostly males, except when indicated. Scale bars: 1 cm. A, Opoptera syme, Brazil, São Paulo, ventral view on the left; B, Bia actorion, Suriname; C, Brassolis sophorae, French Guiana; D, Narope cyllastros, no data; E, Dynastor napoleon, Brazil, Santa Catarina; F, Dasyophthalma rusina, female on the left and male on the right, both from South Brazil.

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Figure 6. Wings in dorsal view (except for G, left side) indicating color-coded pattern elements and trailing bands. Mostly males, except when indicated. Scale bars: 1 cm. A, Caligopsis seleucida, male on the left and female on the right, both from Peru, Madre de Dios; B, Eryphanis zolvizora, Bolivia; C, Eryphanis bubocula, Costa Rica, Guanacaste; D, Caligo eurilochus, male on the left and female on the right, both from Ecuador, Sucumbios; E, Caligo atreus, Colombia, Antiochia; F, Caligo martia, Brazil, Santa Catarina; G, Caligo oileus, male on the left (in ventral and dorsal views) and female on the right, from Peru, Huanuco and Ayacucho, respectively; H, Caligo arisbe, Brazil; I, Caligo beltrao, Brazil, Santa Catarina.

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Figure 7. Male wings in dorsal view (except for E and H, left side) indicating color-coded pattern elements and trailing bands. Scale bars: 1 cm. A, Selenophanes cassiope, Peru, Chanchamayo; B, Selenophanes supremus, Peru, Chanchamayo; C, Penetes pamphanis, Brazil, Paraná; D, Catoblepia xanthus, Brazil, Pará; E, Mielkela singularis, Mexico, Chiapas, ventral view on the left; F, Orobrassolis ornamentalis, Brazil, São Paulo; G, Blepolenis bassus, Brazil; H, Opsiphanes sallei, Peru, Pasco, ventral view on the left; I, Opsiphanes cassina, Ecuador, Sucumbios; J, Opsiphanes boisduvallii, Mexico.

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Figure 8. A, Subordinate groundplan for the Brassolini ventral and dorsal wing surfaces. Continuous lines (or a large circle for element h) indicate pattern elements or trailing bands that are usually present, and small dots or dotted lines denote those that are often absent. The wing outline corresponds to Opoptera syme. B, Annotated Brassolini phylogeny on which selected wing color characters have been optimized.

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Figure 9. Sample of brassoline species showing sexual dimorphism and color convergence. Scale bar for all images: 1 cm. A, Bia actorion male, Suriname and female, Ecuador, Sucumbios; B, Opoptera syme male, Brazil, São Paulo and female (no data); C, Dasyophthalma rusina male on the left and female on the right, both from South Brazil; D, Caligopsis seleucida male on the left and female on the right, both from Peru, Madre de Dios; E, Caligo oileus male on the left and female on the right, from Peru, Huanuco and Ayacucho, respectively; F, Caligo illioneus male, Ecuador Sucumbios; G, Eryphanis automedon male on the left and female on the right, from Colombia, Cali and Paraguay, San Juan Caballero, respectively; H, Catoblepia orgetorix championi male, Panama, Chiriqui and C. orgetorix magnalis female Ecuador, Pastaza; I, Caligo atreus male, Colombia, Antiochia; J, Opoptera staudingeri male, Costa Rica, Heredia; K, Mielkela singularis male on the left and female on the right, both from Mexico, Chiapas; L, Opsiphanes cassina male on the left and female on the right, both from Ecuador, Sucumbios.

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Table 1. General description of ventral patterns (all links last accessed 1 April 2013).

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Table 2. General description of dorsal patterns (all links last accessed 1 April 2013).

Genus	VFW pattern elements	VHW pattern elements	Trailing bands	Ripple pattern	Notes
Bia http://fs.uno.edu/cpenz/bia.html Figure 1A	h present in both sexes, usually 4 oceli, that below M1 being the largest; females may also show c, d and f	h, i, j present in both sexes, females may also show f; 3-4 oceli, that below Cu1 elongated and near wing edge and tail	on the VFW of females between i and j; on the VHW of both sexes between i and j	striated; widespread on both the VFW and VHW	females may have more sparse VFW ripple pattern thus showing more yellow color and visible pattern elements than males
Narope and Aponaropehttp://fs.uno.edu/cpenz/narope.html http://fs.uno.edu/cpenz/aponarope.html Figure 1B and D	b to i may be present in both sexes, depending on the species; 4-7 small oceli; pattern elements usually highly fragmented or blurred, but better defined in N. denticulatus and N. anartes (not illustrated)	c to i may be present in both sexes; typically 7 small oceli; pattern elements usually highly fragmented or blurred, but better defined in N. denticulatus and N. anartes (not illustrated)	visible on both wings when present; variable: delicate (Figure 1B), or well-developed (Figure 1D); considering the group as a whole, most conspicuous in N. nesope and N. denticulatus (not illustrated)	granulated, sparse; widespread on both the VFW and VHW	pattern elements simpler and less developed in females than in males
Brassolis http://fs.uno.edu/cpenz/brassolis.html Figure 1C	c, d, f, h, i and j present in both sexes; the absence of i and j is a diagnostic character for B. haenschi (not illustrated); 4 oceli, that below M1 being the largest	c and h present in both sexes; 3 oceli, anterior and posterior slightly exceeding cell width	on VFW of both sexes, distal to d and f	minute, regular dots; widespread on both the VFW and VHW	pattern similar between the sexes
Dynastorhttp://fs.uno.edu/cpenz/dynastor.html Figure 1E and F	c, d and f vestigial or absent; h present in both sexes; 4-5 oceli with the three anterior near the wing margin and the posterior more proximally placed, the oceli can be small (D. darius) or large (D. macrosiris)	h present in both sexes; usually 3 oceli, 5 in D. macrosiris stryx	on the VFW distal to f, and in D. m. macrosiris and D. m. stryx these are also found distal to h	granulated or striated; widespread on both the VFW and VHW	pattern similar between the sexes; species broadly different from each other ventrally due to the appearance of the ripple pattern and oceli size
Dasyophthalma http://fs.uno.edu/cpenz/dasyophthalma.html Figure 2A	d and f vestigial or absent; h visible in both sexes, 1 or 2 well developed oceli (below R5 and below M1); i vestigial, blurry, more conspicuous in females (D. rusina, Figure 2A, and D. geraensis, not illustrated) or absent (D. creusa and D. vertebralis, not illustrated)	h present in both sexes as 2 or 3 well developed oceli, that below M1 is well developed in D. creusa and D. vertebralis (not illustrated); i vestigial, blurry, or absent	on the VFW distal to f and proximal to i; on the VHW proximal to i, but vestigial in D. creusa (not illustrated)	striated; widespread on both the VFW and VHW	pattern similar between the sexes
Opopterahttp://fs.uno.edu/cpenz/opoptera.html Figure 2B and D	b to i present in both sexes; 4 oceli, that below M1 being the largest; j present or absent	c, d, f, h, i and sometimes j present in both sexes; usually 4 oceli, with those below M1 and M2 being rudimentary	on the VFW, present between b and c, c and d, sometimes within d, distal to f, proximal to i, and sometimes distal to i; on the VHW, distal to d, proximal to i, distal to i and j	striated; limited between f and h on the VFW	pattern similar between the sexes; female paler than male
Caligopsis http://fs.uno.edu/cpenz/caligopsis.html Figure 2C	b to j present in both sexes, c and d composed of multiple twisted lines inside the discal cell; 5 oceli, those below M1 and M2 being the largest	d, e, f, h, i and j present in both sexes; i and j located far from the wing margin; 3 oceli, with the 2 posterior ones being interconnected; f is adjacent to the anterior and posterior oceli	on the VFW proximal to d, between d and f, distal to i, distal to j; on the VHW proximal to d, between h and i, distal to i	striated; limited between f and h on the VFW; the region between the anterior and posterior VHW oceli lacks ripple pattern and, as a result, d is visible inside the discal cell	pattern similar between the sexes; female paler than male
Eryphanishttp://fs.uno.edu/cpenz/eryphanis.html Figure 2E and F	b to j present in both sexes, c and d composed of multiple twisted lines inside the discal cell; g absent or present; 5 oceli, well-developed or rudimentary	d, e, f, h, i and j present in both sexes; i and j located far from the wing margin; 3 oceli, with the 2 posterior ones being interconnected (the small ocelus below M3 may be absent); f is adjacent to the anterior and posterior oceli	on the VFW proximal to d, between d and f, distal to i and distal to j in some species; on the VHW proximal to d, between h and i, distal to i and j in some species	striated; limited between f and h on the VFW; the region between the anterior and posterior VHW oceli lacks ripple pattern and, as a result, d is visible inside the discal cell	pattern similar between the sexes; females paler than males; pattern elements thinner in the automedon-group and wider in the zolvizora-group
Caligohttp://fs.uno.edu/cpenz/caligo.html Figure 3A, B and C	b to j present in both sexes, c and d usually composed of multiple twisted lines inside the discal cell, but sometimes vestigial (Figure 3A); g absent or vestigial; 4 oceli, 1-3 well-developed, but element h may be expressed across the entire wing (e.g., Figure 3C)	d, e, f, h, i and j present in both sexes; i and j located far from the wing margin; 2-3 oceli, small ocelus below M1 sometimes vestigial (Figure 3B) or absent; f may be visible adjacent to the anterior and posterior oceli in some species	sometimes absent or faded (Figure 3A); on the VFW, may be visible distal to c, distal to f, distal to i; on the VHW, proximal to d, distal to i and j	striated; on the VFW, variable from widespread (Figure 3A) to localized between f and h (Figure 3B) to nearly absent (Figure 3C); on the VHW, variable from tight striations that are widespread (Figure 3A, B) to loose striations absent from the submedial area (Figure 3C)	pattern similar between the sexes; females paler than males
Selenophanes http://fs.uno.edu/cpenz/selenophanes.html Figure 3D	b to j present in both sexes, c and d composed of multiple twisted lines inside the discal cell; g absent or vestigial; 4-6 oceli, that below M1 being the largest and most complex	d, h, i and j present in both sexes; anterior ocelus round (S. josephus, not illustrated) or sickle-shaped (Figure 3D); 2-6 oceli, anterior and posterior well-developed	on the VFW trailing colors interspersed with c and d, and a subtle shade distal to f	striated-granular, present on the distal half of the VFW and throughout the VHW	female colors similar to male but VHW h series may be more visible
Penetes http://fs.uno.edu/cpenz/penetes.html Figure 3E	pattern elements extremely reduced in both sexes; f, g and h diffuse, barely noticeable	pattern elements extremely reduced in both sexes; h diffuse and barely visible	on the VFW, distal to f (weak)	absent	pattern similar between the sexes; females paler than males
Catoblepiahttp://fs.uno.edu/cpenz/opsiphanes.html Figs 3F, 4A and B	quite variable among species; b to j present in both sexes of some species, but e, f and g vestigial or absent in others; c and d may be simple or intricate inside the discal cell; 4 oceli, that below M1 being the largest	in both sexes c, d, f, h, i, and j present and f may be vestigial (Figure 3F); 2-6 oceli, depending on the species (compare Figure 4A and B); pattern elements quite variable among species	quite variable among species; on the VFW of C. xanthus (Figure 3F) trailing colors are subdued distal to f and proximal to i, and are diffuse on the VHW; in C. orgetorix (Figure 4A) they are prominent on the VFW between b and c, c and d, distal to f, and proximal to i and are diffuse on the VHW	striated to granular, variable in contrast from faded to bold; present on the VFW discal cell and space between f and h, and throughout the VHW	pattern similar between the sexes; females may be paler than males
Mielkella http://fs.uno.edu/cpenz/mielkella.html Figure 4C	b to j present in both sexes; 4 oceli, that below M1 being the largest	b, c, d, e, f, h, i, j present in both sexes; i and j fragmented; 2-3 oceli, the anterior being the largest, and that below M2 may be vestigial or absent	on the VFW, visible between c and d, distal to f and proximal to i; poorly developed on the VHW	striated; limited to a narrow stripe between f and h on the VFW, present throughout the VHW except for a localized region near the wing base	pattern similar between the sexes; females paler than males
Orobrassolis http://fs.uno.edu/cpenz/orobrassolis.html Figure 4D	b, c, d, e, f, h, i present in both sexes; 3-4 oceli, that below M1 being the largest	b, c, d, e, f, h, i present in both sexes, but f barely noticeable and i fragmented; 2 oceli, the anterior being the largest	on the VFW, visible between b and c, c and d, distal to f; and distal to i; on the VHW, visible between b and c, and c and d	striated; limited to a narrow stripe between f and h on the VFW, present throughout the VHW	pattern similar between the sexes; females may be slightly paler than males
Blepolenis http://fs.uno.edu/cpenz/blepolenis.html Figure 4E	b to j present in both sexes; h particularly well developed across the entire wing, forming a dark band that is unique to Blepolenis among all brassoline genera; 4-5 oceli, that below M1 being the largest	c, d, e, f, h, i, j present in both sexes; j fragmented; 2 oceli of approximately the same size	on the VFW, visible between b and c, c and d, and distal to f; on the VHW, visible between b and c, distal to f, proximal and distal to i	striated; limited to a triangle between g and h on the VFW, present throughout the VHW	pattern similar between the sexes; females paler than males
Opsiphanes http://fs.uno.edu/cpenz/opsiphanes.html Figure 4F	b to j visible in both sexes; 3-4 oceli, that below M1 being the largest	b, c, d, e, f, h, i, j present in both sexes, but f and e vestigial or absent, and i and j fragmented; 2 oceli, the anterior usually larger	variable among species from being nearly absent (O. cassiae, not illustrated) to quite conspicuous (e.g., O. cassina, not illustrated); on the VFW, visible distal to f below M3 or Cu1, and may also be present distal to d (Figure 4F); usually absent or very faint on the VHW (except for O. blythekitzmillerae, not illustrated)	striated; between f and h on the VFW, present throughout the VHW, but sometimes absent from the VHW base	pattern similar between the sexes; females paler than males

Genus	DFW pattern elements	DHW pattern elements	Trailing bands	Iridescence	Sexual dimorphism
Bia http://fs.uno.edu/cpenz/bia.html Figure 5B	difficult to identify due to dark brown background; h present as three anterior oceli near wing outer edge	difficult to identify due to dark brown background	on the DFW, one trailing band distal to f; absent on the DHW	Male: depending on the species, a strongly iridescent patch near DFW tornus, a strongly iridescent patch occupying most of the DHW, or no visible iridescence	coloration clearly dimorphic due to differences in iridescence; wing shape similar between the sexes
Bia http://fs.uno.edu/cpenz/bia.html Figure 5B		difficult to identify due to dark brown background		Female: depending on the species, a large, diffuse iridescent patch in the posterior portion of the DFW medial area, accompanied or not by a small diffuse iridescent patch centrally on the DHW
Narope and Aponaropehttp://fs.uno.edu/cpenz/narope.html http://fs.uno.edu/cpenz/aponarope.html Figure 5D	some species are homogeneously brown, so pattern elements could not be identified (e.g., N. cyllastros, not illustrated); d to j (minus g) are visible in some species (e.g., N. cyllarus, Figure 5D); N. guilhermei (not illustrated) is exceptional in which all pattern elements seem to be present dorsally, especially in the female	some species are homogeneously brown, so pattern elements could not be identified; when pattern elements are visible, typically e, i and j only can be identified	absent from DFW and DHW; we hypothesize that the orange patches present in N. guilhermei (not illustrated) constitute background coloration not covered by the bold dorsal pattern elements	absent from both sexes	some species clearly dimorphic, others mildly dimorphic in color; wing shape dimorphic
Brassolis http://fs.uno.edu/cpenz/brassolis.html Figure 5C	d to j (minus g) are visible	i and j	on the DFW, distal to d and distal to f, amalgamated but f sometimes visible at the edge of the discal cell; on the DHW, distal to i	absent from both sexes	mildly dimorphic in color; wing shape dimorphic
Dynastor http://fs.uno.edu/cpenz/dynastor.html Figure 5E	possibly f to j (minus g) given that the distal portion of the wing is darker than the basal to medial area	i only, or i and j faintly visible	on the DFW, a broken band distal to f, and a continuous band distal to j; on the DHW, distal to i and distal to j	absent from both sexes	mildly dimorphic in color; wing shape slightly dimorphic
Dasyophthalma http://fs.uno.edu/cpenz/dasyophthalma.html Figure 5F	difficult to identify from the brown background, h barely visible in female D. rusina only (Figure 5F, left image)	i and j faintly visible, slightly darker brown than the rest of the wing	on the DFW, bands distal to f and between f and i; on the DHW, a band proximal to i	present in the post-basal and posterior DFW and medial area of the DHW of both sexes of D. rusina and D. geraensis, females having smaller patches	dimorphic in color mostly due to the anterior trailing band distal to f present in females, but the band between f and i is more tilted diagonally in males; wing shape slightly dimorphic
Opoptera http://fs.uno.edu/cpenz/opoptera.html Figure 5A	f to j (minus g) are visible in some species (e.g., O. symeFigure 5A), but in others the wing background is dark and only h and sometimes i can be identified	i only, or i and j	on the DFW, bands distal to f and proximal to i; on the DHW, a band distal to i and sometimes one distal to j	absent from males; a patch present on the DHW of O. syme females; widespread and very faint iridescence present on both wings of female O. aorsa and O. hilaris	species vary in being monomorphic (females only slightly paler in O. sulcius and O. fruhstorferi) to mildly dimorphic; wing-shape similar between males and females
Caligopsis http://fs.uno.edu/cpenz/caligopsis.html Figure 6A	d to j (minus g; e absent or present); h present as anterior oceli becoming a bold band posteriorly	difficult to identify from the brown background, but i and j can be seen in the female in Figure 6A	on the DFW, bands distal to d, distal to f, proximal to i, proximal to j are visible in the female, but only the latter two can be identified in the male; on the DHW, a faint band is visible distal to j on the female	faint iridescence on the medial to basal area of both wings in the female; males of some locations display iridescence that is stronger than in females	dimorphic in color; wing-shape similar between males and females, but male forewing posterior margin bowed
Eryphanishttp://fs.uno.edu/cpenz/eryphanis.html Figure 6B and C	difficult to identify from the brown background; h present as three to five anterior oceli	difficult to identify from the brown background; i only, or i and j are sometimes visible	on the DFW, bands distal to f, proximal to i and distal to j are absent or present depending on the species; on the DHW, a band distal to i is absent or present depending on the species	variable between the sexes and between species; when clear blue iridescence is present in both sexes, it is stronger in the male than in the female sex; in some species the male has a sheen that gives out a faint insinuation of iridescence (Figure 6B), while the females clearly have pale blue iridescent areas on both wings	dimorphic in color; wing shape similar between males and females
Caligohttp://fs.uno.edu/cpenz/caligo.html Figure 6D to I	d to j or h to j (minus g; e absent or present); in some species h is present as anterior oceli becoming a bold band posteriorly	difficult to identify from the brown background, but i and j can be seen in some species	on the DFW, a band distal to f is present in several species, proximal to i in some, and distal to d in C. martia (Figure 6F); two species have a band distal to i on the DHW (C. atreus, Figure 6E; and C. uranus, not illustrated), and that of C. oberthurii (not illustrated) seems to be located proximal to i; some species are hypothesized to have light or colorful background that produces a similar appearance to trailing bands	in both sexes, absent or present in one wing only or both	mildly dimorphic in color; wing shape similar between males and females
Selenophaneshttp://fs.uno.edu/cpenz/selenophanes.html Figure 7A and B	difficult to identify from the brown background, presumably d to j or h to j (minus g); h forms a long series of spots in S. supremus (Figure 7B)	difficult to identify from the brown background, but i and j can be seen in some cases, especially S. supremus (Figure 7B)	on the DFW, a band distal to f only, or combined with one distal to d; on the DHW, bands distal to i and distal to j present or absent	absent from both sexes	monomorphic to mildly sexually dimorphic in color; wing shape similar between males and females
Penetes http://fs.uno.edu/cpenz/penetes.html Figure 7C	difficult to identify from the brown background	difficult to identify from the brown background	on the DFW, a discontinuous band distal to f and another between c and d; on the DHW, a spot near the anterior margin that presumably corresponds to the band distal to i	absent from both sexes	monomorphic in color, but females slightly paler than males; wing shape similar between males and females
Catoblepia http://fs.uno.edu/cpenz/catoblepia.html Figs 7D	difficult to identify from the brown background, presumably d to j or f to j (minus g); h usually appears as white spots above M1 and an ocelus below this vein, more spots are occasionally present below M2	difficult to identify from the brown background; i and j, or i only, presumably present	on the DFW, a continuous or broken band distal to f combined with one proximal to i; on the DHW, a band distal to i that may or may not reach the wing margin	absent from both sexes	wing shape similar between males and females but variable between species
Mielkella http://fs.uno.edu/cpenz/mielkella.html Figure 7E	difficult to identify from the brown background, presumably d to j (minus g); h usually appears as small spots above M1 and a faint ocelus below this vein	difficult to identify from the brown background; i and j presumably present	on the DFW, a band distal to f combined with one proximal to i; on the DHW, a band distal to i	absent from both sexes	sexual dimorphism in wing color and shape
Orobrassolis http://fs.uno.edu/cpenz/orobrassolis.html Figure 7F	c to i (minus g); h appears as small white spots above M1 an ocelus below this vein and a white spot below M2	e and i	difficult to identify due to the light background	absent from both sexes	monomorphic, but females slightly paler; wing shape similar between males and females
Blepolenis http://fs.uno.edu/cpenz/blepolenis.html Figure 7G	d to j (minus g); h appears as small white spots above M1 a faint brown ocelus below this vein and one or two white spots below M2	i and j	difficult to identify due to the light background; it is possible that the diffuse dark orange edges of the blended pattern elements could be a trailing band	absent from both sexes	mildly sexually dimorphic in color; wing shape similar between males and females, but female wings slightly broader
Opsiphaneshttp://fs.uno.edu/cpenz/opsiphanes.html Figure 7H to J	for most species, difficult to identify from the brown background, presumably d to j or f to j (minus g); h usually appears as white spots above M1 and a faint ocelus below this vein; h only in O. boisduvallii, h and i in O. blythekitzmillerae	for most species, difficult to identify from the brown background; i and j presumably present; all pattern elements absent in O. boisduvallii, i only present in O. blythekitzmillerae	for most species, on the DFW, a band distal to f possibly blended with one proximal to i; on the DHW, a band distal to i is usually present; absent from O. boisduvallii and O. blythekitzmillerae	absent from both sexes	mildly sexually dimorphic in color; wing shape dimorphic

Diversidade de padrão das asas em Brassolini (Nymphalidae, Satyrinae)

Resumo