Research Article |
Corresponding author: Andrew F.E. Neild ( andrew.neild@blueyonder.co.uk ) Corresponding author: Shinichi Nakahara ( snakahara@ufl.edu ) Academic editor: Carlos Peña
© 2015 Andrew F.E. Neild, Shinichi Nakahara, Thamara Zacca, Steven Fratello, Gerardo Lamas, Jean-François Le Crom, Diego R. Dolibaina, Fernando M.S. Dias, Mirna M. Casagrande, Olaf H.H. Mielke, Marianne Espeland.
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Citation:
Neild AFE, Nakahara S, Zacca T, Fratello S, Lamas G, Le Crom J-F, Dolibaina DR, Dias FMS, Casagrande MM, Mielke OHH, Espeland M (2015) Two new species of Euptychia Hübner, 1818 from the upper Amazon basin (Lepidoptera, Nymphalidae, Satyrinae). ZooKeys 541: 87-108. https://doi.org/10.3897/zookeys.541.6297
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Two new species of Euptychia Hübner, 1818 are described from the upper Amazon basin: E. attenboroughi Neild, Nakahara, Fratello & Le Crom, sp. n. (type locality: Amazonas, Venezuela), and E. sophiae Zacca, Nakahara, Dolibaina & Dias, sp. n. (type locality: Acre, Brazil). Their unusual facies prompted molecular and phylogenetic analyses of one of the species resulting in support for their classification in monophyletic Euptychia. Diagnostic characters for the two species are presented based on wing morphology, wing pattern, presence of androconial patches on the hindwing, and genitalia. Our results indicate that the projection of the tegumen above the uncus, previously considered a synapomorphy for Euptychia, is not shared by all species in the genus. The adults and their genitalia are documented, and distribution data and a map are provided.
Resumo
Duas novas espécies de Euptychia Hübner, 1818 são descritas da bacia do alto Rio Amazonas: E. attenboroughi Neild, Nakahara, Fratello & Le Crom, sp. n. (localidade tipo: Amazonas, Venezuela), E. sophiae Zacca, Nakahara, Dolibaina & Dias, sp. n. (localidade tipo: Acre, Brasil). Suas aparências não usuais levaram a análises molecular e filogenética de uma das espécies que sustentam sua classificação como Euptychia. São apresentados caracteres diagnósticos para as duas espécies baseados na morfologia e no padrão de coloração de suas asas, presença de manchas androconiais nas asas posteriores e suas genitálias. Os resultados indicam que a projeção do tegume sobre o unco, previamente considerada uma sinapomorfia para Euptychia, não é compartilhada por todas as espécies do gênero. São ilustrados os adultos e suas genitálias, bem como dados e mapa de distribuição.
Resumen
Se describen dos nuevas especies de Euptychia Hübner, 1818 para la cuenca alta del Río Amazonas: E. attenboroughi Neild, Nakahara, Fratello & Le Crom, sp. n. (localidad típica: Amazonas, Venezuela) y E. sophiae Zacca, Nakahara, Dolibaina & Dias, sp. n. (localidad típica: Acre, Brasil). Sus apariencias inusuales motivaron el análisis molecular y filogenético de una de las especies, lo cual sustenta su clasificación como Euptychia. Se presentan caracteres diagnósticos para las dos especies basados en la morfología y patrón de coloración alar, presencia de parches androconiales en las alas posteriores y sus genitales. Los resultados indican que la proyección del tegumen sobre el uncus, previamente considerada una sinapomorfía para Euptychia, no es compartida por todas las especies del género. Se ilustran los adultos y sus genitales, y se proporcionan datos y mapa de distribución.
Euptychiina , Neotropics, South America, Brazil, Colombia, Peru, Venezuela, mitochondrial DNA COI barcode
The nymphalid subtribe Euptychiina (Satyrinae: Satyrini) is one of the most poorly known butterfly groups. More than 400 predominantly Neotropical species in some 50 genera are recognised within the subtribe and many taxa remain undescribed (
Comparison of the morphology of the two new species was made with other Euptychia specimens in the collections listed below (museum acronyms are from
AN Andrew Neild collection, London, UK
BMNH
The
MGCL
McGuire Center for
MUSM
Type specimens of Euptychia species in the BMNH were also checked, as well as photographs of additional taxa on the website “Butterflies of America” (available online in
The following abbreviations are used:
FW forewing
HW hindwing
D dorsal
V ventral
A leg from the Brazilian male paratype (DZ 29.578) of E. sophiae sp. n. was used to obtain a DNA sequence for cytochrome c oxidase subunit I (COI) and elongation factor-1 alpha (ef1a). We extracted DNA using Qiagen’s DNeasy Blood & Tissue Kit, following the protocol and using a final elution volume of 50 µl. Primers LCO_nym (forward, 5'TTTCTACAAATCATAAAGATATTGG 3’) and HCO_nym (reverse, 5'TAAACTTCAGGGGTGACCAAAAAATCA 3’) were used to amplify COI. Elongation factor-1 alpha was amplified by using primer pairs Ef44 (forward, 5'GCYGARCGYGARCGTGGTATYAC 3’), and EfrcM4 (reverse, 5'ACAGCVACKGTYTGYCTCATRTC 3’); however, since this primer pair failed to amplify ef1a, the following primer pair was used to amplify short fragments of ef1a: Ef44 (forward, 5'GCYGARCGYGARCGTGGTATYAC 3’) and Monica (reverse, 5'CATRTTGTCKCCGTGCCARCC 3’) (
Genus | Species | Voucher code | COI | EF1a |
---|---|---|---|---|
Euptychia | enyo | CP06-73 | GQ357205 | GQ357275 |
Euptychia | sp. 2 | CP01-33 | DQ338794 | DQ338937 |
Euptychia | westwoodi | DNA96-005 | AY508543 | AY509069 |
Euptychia | boulleti | PM17-01 | JQ639284 | JQ639285 |
Euptychia | sp. 6 | CP04-55 | DQ338796 | DQ338939 |
Euptychia | sophiae | DZ 29.578 | KR818703 | KR818706 |
Euptychia | picea | DNA99-036 | AY508542 | AY509068 |
Euptychia | sp. 5 | CP01-53 | DQ338795 | DQ338938 |
Euptychia | sp. 7 | CP02-58 | GQ357206 | DQ338940 |
Magneuptychia | fugitiva | CP01-18 / DNA99-008 | GU205845 | AY509078 |
Papilio | machaon | BC ZSMLep 27060 / NA | GU707119 | EF485106 |
Sequences were aligned using MAFFT v. 7.107 (
Partitioning schemes and substitution models determined by Partitionfinder.
Partition | Best model MrBayes | Subset partitions BEAST | Best model RAxML | Subset partitions RAxML |
---|---|---|---|---|
1 | GTR+I | COI position 1 | GTR | COI position 1 |
2 | HKY+I | COI position 2 | GTR | COI position 2 |
3 | HKY+G | COI position 3 | GTR | COI position 3 |
4 | GTR | EF1a position 1 | GTR | EF1a position 1 |
5 | HKY | EF1a position 2 | GTR | EF1a position 2 |
6 | GTR+G | EF1a position 3 | GTR | EF1a position 3 |
Phylogenies were inferred using maximum likelihood (ML) and Bayesian inference on the concatenated data set of 1041bp. ML analyses were run using RAxML v 8.1.11 (
According to ML and Bayesian inference based on barcoding region and EF1a, E. sophiae sp. n. is sister to the clade consisting of E. picea Butler, 1867 and two undescribed species. Unlike
Bayesian phylogeny of Euptychia based on one mitochondrial (COI) and one nuclear (EF1-a) gene. Posterior probabilities are listed above and bootstrap values below branches. A dash denotes bootstrap support lower than 50%. (Euptychia attenboroughi is not included in the analysis – see text for details.)
Diagnostic morphological characters seen in these two new species which appear to be unique among the genus Euptychia include: (i) presence of a single signum in female E. attenboroughi sp. n. (no female is known for E. sophiae sp. n.); (ii) lamella antevaginalis unsclerotised around the ostium bursae of female E. attenboroughi sp. n. (sclerotised in all other Euptychia examined), except on the ventral surface around the neck of the ostium bursae; (iii) reduced third segment of labial palpus of female E. attenboroughi sp. n. (Fig.
Holotype male with the following labels (separated by transverse bars): HOLOTYPE / VENEZUELA – Amazonas: San Carlos de Rio Negro to Solano track, km. 3 [approximately 1°55'N 67°1'W], 5-17 III 94, 100m elev., Andrew Neild Collection / Brit. Mus. 1994-298 / BMNH(E) #1054424 (BMNH).
VENEZUELA – Amazonas: 1 female, same locality data as holotype / (Neild Prep. Genital Vial No. 274) (AN); BRAZIL – Amazonas: 1 female, Janarete [= Jauaretê] (approximately 0°36'N 69°11'W), IX, 1943 / W. Praetorius Coll. Donor Frank Johnson (
Differs from males of its congener E. sophiae sp. n. as follows (no females of E. sophiae sp. n. are known, but we expect characters indicated with an asterisk (*) will serve to differentiate this sex): (1) FW more produced apically, with outer margin straighter or more concave, and outer margin at an angle away from the central line of the body (nearly parallel in E. sophiae sp. n.); (2) dorsally with prominent orange scaling on the posterior DHW on and near the tornus* (E. sophiae sp. n. lacks this orange scaling); (3) VHW submarginal band does not increase in width at the tornus (increases substantially for E. sophiae sp. n.); (4) small ocellus at the anal margin of the VHW median band larger*; (5) androconial patch on VHW pale and barely visible, whereas prominent on E. sophiae sp. n.; (6) gnathos projecting nearly parallel to the uncus (projecting vertically in E. sophiae sp. n.); (7) distal one-fourth of the valva broad in lateral view (narrow in E. sophiae sp. n.); (8) aedeagus straighter, less curved.
The female resembles several species in the genus Euptychia, especially E. marceli Brévignon, 2005, but is distinguishable on the VHW through the unique combination of a very large and ovoid (not round) ocellus in Cu1-Cu2 bordered on its tornal side by orange scaling, and by the presence of a tiny ocellus on the anal margin at the posterior end of the median brown band (this last found rarely on some specimens of E. marceli Brévignon, 2005). It can also be differentiated from all other known Euptychia females by the three characters (i, ii, and iii) elucidated at the beginning of the Taxonomy section. The male somewhat resembles a few taxa currently in Chloreuptychia Forster, 1964 (see
MALE (Fig.
Forewing length. 17.0–18.0 mm (n = 3) (holotype = 18 mm).
Head. Frons brownish; postgenal area light brown.
Antennae. Naked, orange-brown, darker dorsally, clubs browner with orange tip, 7–8 mm long.
Eyes. Dark brown, sparsely hairy; creamy-grey scales dorsally and laterally along posterior edge of eyes.
Palpi. Covered by long creamy-grey hair-like modified scales dorso-laterally, ventrally with long fine hair-like modified scales projecting like a Mohican, mostly black along outer margin, but interior wall of modified scales creamy-grey. Mohican highest in centre, gradually reducing anteriorly and posteriorly, and anteriorly reduced to a pointed tuft. First segment covered with black scales dorsally, black and white hair-like modified scales ventrally, second segment covered with short white hair-like modified scales and white scales laterally, black scales distal one-third of dorsal surface, ventrally adorned with long black and white hair-like modified scales 3–4 times as long as segment width, second segment slightly longer than eye diameter, third segment covered with black scales dorsally and ventrally, creamy-white hair-like scales laterally, about one-seventh of second segment in length.
Thorax. Covered in long light grey hair-like modified scales.
Legs. Greyish. Foreleg tarsus about 2/5 of tibia in length, femur about 2/3 of tibia in length; tibial spurs absent on midleg and hindleg.
Abdomen. Eighth tergite and sternite well developed, apparently as equally sclerotized as other tergites and sternites, but weakly sclerotized towards posterior end.
Androconia patches. Modified wing scales, presumed to be androconia, present on either side of vein 2A on the HW at the base of the dark median band on the dorsal and ventral surface; visible on the dorsal surface as two tiny ovoid pale greyish-brown patches approximately 0.5 mm long, and on the ventral surface as a strip approximately 2.0 mm long in space 2A–3A; these patches, best viewed using backlighting, are homologous with the dorsal and ventral androconia patches of E. sophiae sp. n.
Wing venation. FW recurrent vein present, approximately 1.75 mm long; FW vein Cu not swollen at base; HW with humeral vein barely visible, very short (approximately 0.6 mm), curved anteriorly towards the costal lobe.
Wing shape. FW costa gently convex to apex, apex relatively pointed, outer margin straight, or slightly concave between M3 and Cu2, and angling about 20 degrees away from the central line of the body, inner margin almost straight; HW costa lobed in basal area then gently concave to apex at Sc+R1, apex rounded, outer margin scalloped, anal margin concave near tornus, basally convex.
DFW. Both wings slightly and variably translucent with greyish-brown to chestnut brown ground colour; fringes greyish-brown to brown; four diffuse dark brown to chestnut bands crossing from the costal to the anal margin, the first basal and barely visible, mostly ghosting through the slightly translucent wing from the ventral surface, centrally wide but tapering to a point anteriorly and posteriorly, the second submedian, wider, nearly straight and better defined, crosses the mid discal cell from mid costa to four tenths along the inner margin, the third median slightly wavy strongly defined and the widest, crosses from the costa near the discocellular veins, which it traverses, reaching to seven tenths distance along the inner margin, the fourth begins near the apex where it is narrow and very sinuous, runs parallel to the outer margin and curved in each interspace down to M3 and then angles without curves in towards the body and widens reaching the submarginal area of Cu1, then again running parallel to the outer margin down to the inner margin near the tornus; the margin with a very fine dark brown line running parallel to the outer margin, beginning at the apex, incurved in each interspace to M3, then straight to the tornus; a white-pupilled black subapical ocellus in the centre of M1-M2, touching M2 but not quite reaching M1.
DHW. Four diffuse dark brown to chestnut bands crossing from the costal to the anal margin, the first basal, mostly ghosting through the slightly translucent wing from the ventral surface, the second submedian also ghosting through, nearly straight, crosses the mid discal cell from mid costa to half distance along the anal margin, the third median and slightly wavy, in some specimens (two of the three males) curved inwards in the distal discal cell, better defined with dorsal scaling (less ghosting) and the widest, crosses from the costa two-thirds towards the apex, almost reaching 2A two-thirds along its length, and not passing to the anal margin, the fourth submarginal begins near the apex where it is narrow, runs parallel to the outer margin (curved in each interspace), gradually thickening, widest in M2-M3 (where the basal edge points inward) and M3-Cu1, and then thinning gradually to just reach 2A near the tornus; the margin with a very fine clearly defined dark brown line running parallel to the outer margin, beginning at Sc+R1, incurved in each interspace to the tornus and entering the anal lobe to 3A; three ventral ocelli visible through the slightly translucent wings, the dark circular areas showing through from the ventral surface in cells Cu1-Cu2 and M1-M2 covered with small spheroid areas of diffuse very dark brown dorsal scaling, the latter very small, the former roughly half the diameter of the black ventral “iris”; orange scaling on the distal side of the large ocellus in space Cu1-Cu2 continues to the outer margin and in the same areas of Cu2-2A.
VFW. Both wings slightly translucent with pale greyish-brown ground colour; fringes greyish-brown; one very thin well-defined submarginal dark brown to chestnut band and four more diffuse dark brown to chestnut basal bands, submedian, median and postmedian crossing from the costal to the anal margin, as described for the dorsal surface; a silvery-white-pupilled black subapical ocellus in the centre of space M1-M2, touching M2 but not quite reaching M1, circled by a gold ring with a thin grey-brown outer edge which enters R5-M1; in M2-M3 the grey-brown edge of the ocellar ring breaks open and the yellow area spills posteriorly into the centre of the interspace; in one specimen (MGCL) there is an additional tiny ocellus in Cu1-Cu2 with a reddish-brown “iris” (of the same colour as the transverse bands), a tiny pupil (or merely missing scales?), and an outer ring of the same colour as the background pale-grey-brown, surrounded by an indistinct scattering of brownish scales that define the edge of the outer pale ring.
VHW. One very thin well-defined marginal dark brown band and four more diffuse dark brown to chestnut basal bands, submedian, median, and marginal crossing from the costal to the anal margin, as described for the dorsal surface, but the submarginal and marginal bands continue from the tornus along the anal margin to the base of the chestnut median band, while the marginal band (only) continues to the base of the submedian band; a very small ocellus on the anal margin at the base of the median brown band composed of a large black subovoid “iris”, a narrow golden outer ring, and a narrow dark brown border; three postmedian ocelli, composed of a single small silvery-white pupil, a large black spheroid or ovoid “iris”, and a narrow golden outer ring enclosed in a narrow grey-brown border; the smallest of these ocelli is spheroid, entirely within Rs-M1, not touching either vein, the second, nearly twice as large, spheroid and occupying the full width of M1-M2, with the grey-brown outer border just touching M1, and the black “iris” just spilling over into M2-M3, and in the same way as on the VFW, with the outer posterior border broken open in M2-M3 and the golden outer ring protruding into the centre of the interspace, where a poorly defined wide suffused band of greyish brown scales links this and the third largest ocellus, ovoid, over three times the diameter of the first at its widest point (parallel to the outer margin), with the black “iris” fully occupying Cu1-Cu2, spilling into M3-Cu1 but just touching Cu2, the golden outer ring double the width of the other two ocelli, reaching the centre of M3-Cu1, and entering Cu2-2A, surrounded by a grey brown ring with indistinct edges; yellow-orange scaling on the distal side of the large ocellus in space Cu1-Cu2 continues slightly into M3-Cu1 and fully to the submarginal band of Cu1-Cu2 with more extensive orange scaling in the distal quarter of Cu2-2A just spilling over 2A onto the anal lobe.
Male genitalia (Fig.
Euptychia attenboroughi sp. n. Genitalia of holotype male: a dorsal view b dorso-lateral view c lateral view (“G” indicates fused gnathos) d ventral view e aedeagus, dorsal view, and f lateral view. BMNH collection. Photos by Andrew Neild, Trustees of the Natural History Museum, London. Scale bar: 0.5 mm.
FEMALE (Fig.
Forewing length 16.0–17.0 mm (n = 3). Similar to male except as follows: FW shape not elongate, subtriangular, with distinctly convex outer margin; HW shape similar to male, but slightly less elongate; dorsal surface with all the same dark semi-translucent bands but the basal, submedian, and median bands with more scales present dorsally; the DHW dark circular areas showing through from the ventral surface in cells Cu1-Cu2 and M1-M2 almost covered with large spheroid areas of diffuse very dark brown, nearly black, dorsal scaling; the large subtornal ocellus entirely encircled by diffuse orange scaling, and distally by a half-circle of golden scales homologous with the ventral outer ring of this colour; ventrally there are no obvious differences.
Female genitalia (Fig.
We name this butterfly to honour the great English naturalist, author, and TV presenter, Sir David Attenborough, in gratitude for opening the eyes and hearts of millions to the natural world through his inspiring and edifying work. To prevent any future ambiguity, the name attenboroughi is considered to be a Latinised male noun in the genitive case.
The six specimens known to date were all collected within 500 kms of each other in the north-west of the upper Amazon basin, representing a very restricted distribution. It is impossible with such a small sample size to draw any concrete conclusions, but we hypothesise that this species is restricted to suitable habitat to the north of the Amazon river, and that its sibling species occurs only to the south, although a limited area of sympatry may exist. One of the senior authors first collected specimens of this new species in 1994 while conducting field work for the Butterflies of Venezuela book series (
Unknown.
Holotype male with the following labels (separated by transverse bars): /Holotypus/ Brasil, Acre, Mâncio Lima, P[ar]q[ue] Nac[ional] Serra Do Divisor, Porção Norte, 7°26'50"S 73°39'52"W 200-400 m 10-21-IX-2011, D. Dolibaina & D. Moura Leg. / DZ 29.579/ (
1 male, same data as holotype, except: DZ 29.578 (
See relevant section for E. attenboroughi.
MALE (Fig.
Forewing length 18–19 mm (n = 3) (holotype = 18 mm).
Head. Brown. Postgenae with creamy-grey scales.
Antennae. Naked, brown, darker dorsally, clubs browner with orange tip.
Eyes. Dark brown, sparsely hairy. Creamy-grey scales dorsally and laterally along posterior edge of eyes.
Palpi. Covered by long creamy-grey hair-like modified scales dorso-laterally, ventrally with long fine hair-like modified scales projecting like a Mohican, mostly black along outer margin, but interior wall of modified scales creamy-grey. Mohican highest in centre, gradually reducing anteriorly and posteriorly, and anteriorly reduced to a pointed tuft. First segment covered with brown scales dorsally, black and creamy hair-like modified scales ventrally, second segment covered with short creamy hair-like modified scales and light brown scales laterally, brown scales distal one-third of dorsal surface, ventrally adorned with long black and creamy hair-like modified scales 3–4 times as long as segment width, second segment slightly longer than eye diameter, third segment covered with brown scales dorsally and black scales ventrally, creamy hair-like scales laterally, about one-seventh of second segment in length.
Thorax. Uniformly covered by dark grey-brown hair-like scales.
Legs. Greyish. Foreleg tarsus more than half-length of tibia, femur about 2/3 of tibia in length; tibial spurs absent on midleg and hindleg.
Abdomen. Eighth tergite and sternite well developed, apparently as equally sclerotized as other tergites and sternites, but weakly sclerotized towards posterior end.
Androconial patches. Two small (< 1.0 mm) pale grey androconial patches on DHW, barely separated by 2A, located at its distal one third; patch in cell 2A–3A prominent; patch in cell Cu1-Cu2 restricted to width of median band and located at juncture of this band with 2A. A black and short (approximately 1.5–2.0 mm) androconial patch at the distal third of 2A on the VHW.
Wing shape. FW triangular, costal margin convex, apex rounded, outer margin gently convex from apex to Cu1, tornus rounded, anal margin straight. VW costa slightly convex, apex rounded, outer margin crenulated, anal margin concave near tornus, remaining convex.
DFW. Greyish brown, darker along the costal and outer margins with a narrow ochre area on the first fourth of the costal margin length. Four dark brown to rufous bands, the former basal, dark brown (approximately 0.1 mm width) following the radial vein on its distal edge, the second submedian, dark rufous brown, from near the origin of R1 to 2A, crossing the discal cell near the middle, the third median, rufous and slightly concave, crossing the cell end from the origin of R3 across the base of Cu1 to 2A, curving distally near the inner margin, and the final band submarginal, dark brown, narrower and slightly crenulated from R4 to M3, rufous and posteriorly enlarged from M3 to 2A. Ocellus of the VFW observable through transparency.
DHW. Greyish brown. Four dark brown to rufous straight bands, the first basal, dark brown, short and tapered, the second submedian, dark rufous brown, from costal margin to anal margin, crossing the discal cell near the centre, the third median, rufous brown, from costal margin to anal margin, crossing the discal cell in its distal quarter, the final band submarginal, rufous and crenulated in each cell from Rs to anal margin, except Cu1-Cu2, following the contour of the outer margin, but widened and curved inwards in M2-M3 and to a lesser degree in M3-Cu1, wide and reddish orange from Cu1 to 2A, thinned from 2A to anal margin where it nearly reaches the median rufous band. Ocelli of the VHW observable through transparency.
VFW. Light greyish brown, bands similar to DFW. One developed black ocellus from M1 to anterior sixth in M2-M3, with a white pupil at the centre and a broad and yellow external ring, surrounded by a greyish brown area that extends posteriorly to the posterior half of M3-Cu1. Submarginal line dark brown, from R4 to anal margin, crenulated in R4-M3, remainder and straight, distally surrounded by a thin yellow line. Fringes dark brown.
VHW. Light greyish brown, bands similar to DHW. Three postmedian black ocelli, the anterior the smallest in Rs-M1, the second twice as wide as the first, from M1 to anterior third of M2-M3, and the posterior bigger, about three times wider than the second, ovoid, from the posterior third in M3-Cu1 to the edge of Cu2, all three with a white pupil at the centre and a broad and yellow external ring. A fourth minute black ocellus, with a yellow outer ring but with no pupil, located on the anal margin at the base of the postmedian brown band. Marginal dark brown, thin, and crenulated line from Sc+R1 to 3A. Fringes dark brown.
Male genitalia (Fig.
FEMALE: Unknown.
The specific epithet honours T. Zacca’s niece, Laura Sophia. To prevent any future ambiguity, the name sophiae is considered to be a Latinised modern female noun in the genitive case.
This species is only known from the type locality in Serra do Divisor National Park (SDNP), Acre, in the extreme west of Brazil, and from across the border in the neighbouring department of Loreto, in north-eastern Peru. The Brazilian specimens of E. sophiae were collected in forest characterized as submontane dense ombrophilous forest in a landscape of “terra firme” forest with patches of seasonally inundated areas with a predominance of palms in the genus Mauritia Linnaeus f. (Arecaceae) at about 200 m a.s.l. (see Figs
Unknown.
Males of E. attenboroughi and E. sophiae, especially E. attenboroughi, exhibit external wing pattern elements and wing morphology that are atypical for the genus, superficially recalling certain species in the genus Chloreuptychia. It was therefore important to support our generic classification for these two new taxa using more objective genetic analyses. Both new species are described in this genus on the basis of ML and Bayesian analyses of E. sophiae, reinforced by the ML analysis performed in
The authors wish to express their gratitude to the following for their generous assistance: Christian Brévignon (French Guiana) for helpful discussion; André Freitas (Brazil) and Mario Marín (Brazil) for valuable discussion regarding systematic placement for these taxa; Suzanne Green, David Grimaldi, and Lesley Thayer (all