Yphthimoides patricia ( Hayward, 1957 )

Barbosa, Eduardo P., Siewert, Ricardo R., Mielke, Olaf H. H., Lamas, Gerardo, Willmott, Keith R. & Freitas, André V. L., 2018, Redescription of Yphthimoides patricia (Hayward, 1957), with taxonomic notes on the names Euptychia saltuensis Hayward, 1962 and Yphthimoides manasses (C. Felder & R. Felder, 1867) (Nymphalidae: Satyrinae), Zootaxa 4422 (4), pp. 537-557 : 540-550

Yphthimoides patricia ( Hayward, 1957)

Euptychia patricia Hayward, 1957: 115 , figure 6 (♂ genitalia). Holotype ♂, Chulumani, Sur Yungas, Bolivia, Padre Bridarolli leg., I-1948, collection Instituto de Historia Natural “ Sánchez Labrador ”, San Miguel , Buenos Aires; genitalia preparation Nº MSS 11. [ Supposedly deposited at MACN but not found].

Euptychia saltuensis Hayward, 1962: 106 , figure 2 (♂ genitalia). Holotype ♂, no locality data, probably from Argentina, Paraguay or southeast Brazil, IML; genitalia preparation Nº SAT 252 (examined). New synonym.

Yphthimoides patricia: Forster, 1964: 102 , 103; Lamas, 2004: 223.

Yphthimoides manasses [misidentification]: Freitas, 2004: 10, figure 3 (♂ ventral); Silva et al., 2012: 294, 295, tab. 1; Silva et al., 2015: 6, tab. 1.

Yphthimoides sp. 1: Freire-Júnior & Diniz, 2015: 1209, 1210, tab. 3.

Diagnosis. Yphthimoides patricia ( Fig. 1 ) differs from Y. manasses ( Fig. 2 ) in the following characters: wings usually more rounded in Y. manasses than in Y. patricia ; DHW with ocelli in Cu1-Cu2 and Cu2-1A+2A with single pupils in Y. patricia , but usually double (when present) in Y. manasses . Males of Y. patricia have a patch of dark androconial scales on the middle of the DFW that is absent in Y. manasses . The morphology of the male genitalia is quite distinctive, as can be seen in figure 3 (A–D for Y. patricia and E–H for Y. manasses ). The valvae of the male genitalia have two series of small 'teeth' on the posterior inside edge in Y. patricia ( Fig.3C ), which are absent in Y. manasses ( Fig. 3G ), and in fact in all other Yphthimoides species.

Redescription of adults. Male ( Figs. 1 , 3–7 , 10C ). FW length: 16–20 mm (average 17.6 mm, SD = 1.17, n = 11); HW length: 12–15 mm (average 13.1 mm, SD = 0.88, n = 11). Eyes glabrous, entirely yellowish ochre ( Fig. 4A ). Male palpi ( Figs. 4A and 5D ) length approximately twice the head height, covered with cream scales and black and cream hairs. Antennae 9–10 mm in length, with 43–45 antennomeres extending to mid-costa of FW; shaft rust-brown, with white scales at base of each antennomere ( Fig. 4B ), club not conspicuously developed, with 13–14 antennomeres, yellowish brown, with the four distal-most antennomeres darker ( Fig. 4C ). DW ground color brown with a few markings; marginal and submarginal lines light brown and wavy, especially conspicuous in the HW. DFW with a broad dark androconial area extending from the inner margin to Radial vein ( Fig. 1A ). A wellmarked narrow dark irregular stripe following the dcs to dcm transverse discal veins. This dark area is not easily viewed unless with direct light, especially in darker wet season forms. VFW crossed by two lines, submedian faint orange and wavier, across discal cell, extending from Sc to 1A+2A, at one-third from the wing base; and postdiscal dark orange and straight extending from costa to 1A+2A, at two-thirds from the wing base. VFW also with a light cream, broad postdiscal stripe extending from costa to 1A+2A distad of dark orange postdiscal line. Two dark brown ocelli on DHW, outlined by a yellow ring and with silver pupil: one larger, between Cu1 and Cu2 and a smaller one between Cu2 and 1A+2A. VHW ground color light brown with several markings: in both wings marginal line straight and submarginal line wavy; VHW crossed by two orange lines extending from costa to anal margin, a submedian faint and slightly wavy line, at one-third from the wing base; and a postdiscal dark and straight line, at two-thirds from the wing base. A light cream postdiscal stripe, broader than that on the VFW, extending from costa to anal margin distad of dark orange postdiscal line. In wet season individuals, a series of six dark ocelli with white pupil and outlined by a ring of yellowish-creamy scales is present in VHW in wing cells Rs- M1 (ocellus 1), M1-M2 (2), M2-M3 (3), M3-Cu1 (4), Cu1-Cu2 (5) and Cu2-1A+2A (6). Ocelli 2 and 5 larger than others. In some individuals ocelli 3 to 6 have double pupils. HW outer margin slightly wavy. Male wing venation shown in Fig. 5A . Legs covered by cream scales ( Fig. 4D ); Male foreleg shown in Fig. 5C .

Male genitalia ( Figs. 3 and 6 ). Saccus short and triangular in dorsal view ( Fig. 3A–B ); tegumen rounded ( Fig. 3B ); gnathos long and pointed, almost half size of the valva length ( Fig. 2B ); uncus elongated with apex truncated distally ( Figs. 3B, C ; 6A); valvae elongated, trapezoidal, ending in a tapered apex in lateral view ( Fig. 3A ), internal margin with two series of small 'teeth', one series in apex and other series more internal ( Figs. 3B, C ; 6B); aedeagus straight ( Fig. 3D ); cornuti absent; fultura inferior membranous.

Female ( Figs. 1C–D and 7G–L , 10D ). FW length: 19–22 mm (average 20.5 mm, SD = 1.22, n = 6); hind wing length: 16–20 mm (average 17.5 mm, SD = 1.52, n = 6). Eyes glabrous, entirely brown. Palpi length approximately twice the head height, beige with long brown hairs. General color and pattern very similar to, but in general paler than, that of males. Wing venation shown in Fig. 5B . Foreleg with five tarsomeres, as shown in Fig. 5E .

Remarks on color variation ( Figs. 1 and 7 ). Two seasonal forms are known, affecting both sexes, with differences observed only on the underside. The wet season form is darker with larger and more conspicuous ocelli, especially on the VHW; based on museum specimens and unpublished data (AVLF pers. obs.), this form was reported as being dominant from November to April in Central Brazil ( Figs. 1E , 7A, B, C, G, H ). The dry season form is lighter, often yellowish, with reduced ocelli on both wings, occasionally presenting a whitish postdiscal stripe distad of the postdiscal line crossing the wings ventrally; based on Museum specimens and on unpublished data (AVLF pers. obs.), this form was reported as being dominant from May to September ( Figs. 1A–D, F , 7D, E, F, K, L ). Intermediates ( Figs. 7I, J ) were observed in transitions between seasons, from February to April and in October–November in Central Brazil (AVLF pers. obs.).

Description of immature stages. The morphological description and measurements of the immature stages below are based on material reared from January to April 1999, obtained from three wild caught females from Itirapina, São Paulo, southeastern Brazil.

Egg. Barrel shaped, flattened at base, pinkish cream, with a reticle of 19–20 thin longitudinal ridges forming a pattern of irregular pentagonal and hexagonal cells visible under light microscope. Height and diameter 1.0 mm (n = 5). Duration 6 days (n = 15).

First instar. Head capsule width 0.62–0.64 mm; head scoli 0.08–0.10 mm (n = 5). Head capsule black, with enlarged chalazae, bearing a pair of short scoli on vertex, each with two long narrow setae, P1 and P2 respectively. Third stemma larger than other stemmata. Body cream, smooth, with red longitudinal stripes; caudal filaments very short. Setae dark, elongated, several dorsal and subdorsal clubbed at tip. Maximum length 6 mm. Duration 9–10 days (n = 15).

Second instar ( Fig. 8A ). Head capsule width 0.90 mm; head scoli 0.16–0.18 mm (n = 5). Head dark brown with two diverging short scoli on vertex. Body beige, with a broad dorsal dark stripe and several lateral longitudinal weakly marked stripes; caudal filaments short. Maximum length 9 mm. Duration 7–8 days (n = 10).

Third instar ( Fig. 8B ). Head capsule width 1.22–1.30 mm; head scoli 0.24–0.26 mm (n = 5). Head brown, with two diverging very short scoli on vertex. Body brown with a broad dorsal longitudinal stripe and several lateral longitudinal weakly marked intersecting brown stripes; caudal filaments short. Maximum length 13 mm. Duration 6–8 days (n = 5).

Fourth instar ( Fig. 8C ). Head capsule width 1.80–1.98 mm; head scoli 0.28–0.36 mm (n = 10). Very similar to third instar. Maximum length 21 mm. Duration 9–11 days (n = 5).

Fifth (last) instar ( Fig. 8C,D ). Head capsule width 3.08–3.23 mm; head scoli 0.55–0.58 mm (n = 4). Head brown, with two diverging short scoli on vertex. Body brown with a broad dark dorsal stripe and several lateral longitudinal weakly marked intersecting brown stripes; ventral region dark brown; legs and prolegs light brown; caudal filaments short. Maximum length 25 mm. Duration 17–19 days (n = 10).

Pupa ( Fig. 8E, F, G ). Short and smooth; mostly dark brown with a mottled pattern of cream dots; short pointed ocular caps with white ridges; cremaster dark in ventral portion; dorsal abdomen with a pair of short bumps laterally in each segment. Total length 12 mm (n = 5). Duration 13–15 days (n = 6).

Behavior and Natural History. Oviposition behavior was not observed in the field and the natural host plant is unknown. In captivity, eggs were laid on provided host plant leaves and/or scattered on the inside of the rearing container (plastic bag). In the laboratory, larvae quickly accepted three common grasses: the African Guinea grass Panicum maximum Jaq. (“capim colonião”), the Molasses grass Melinis minutiflora P. Beauv. (“capim gordura”) and the Indian goose grass Eleusine indica (L.) Gaertn (“capim pé-de-galinha”). These three species are introduced in Brazil and no native grass species used by Y. patricia are known. Adults fly near the ground from early morning to afternoon. Both sexes were observed feeding on decaying fruits lying on the ground.

Distribution and habitat ( Figs. 9 and 10 ). Based on field observations, museum records and literature, the species is distributed in the Brazilian states of Pará, Maranhão, Bahia, Mato Grosso, Mato Grosso do Sul, Goiás, Minas Gerais, São Paulo and in the Distrito Federal; and in the Bolivian departments of La Paz and Santa Cruz ( Fig. 9 ). This species is strongly associated with open habitats, including the Brazilian savannas (the cerrado biome, Figs. 9 , 10A,B ), where it occurs in several environments, including the narrow gallery forests adjacent to the cerrado (Silva et al. 2015). It can be locally abundant, and is the dominant satyrine species in some cerrado sites (Silva et al. 2015, Freire-Júnior & Diniz 2015, AVLF pers. obs.). It was also recorded in the rocky montane fields of Minas Gerais (known locally as “campos rupestres”) and in the Amazonian “canga” vegetation (iron-rich rock outcrops with associated vegetation that grows on thin soil) in the region of Carajás (south Pará State). The species has never been recorded in the Brazilian semiarid “caatingas”. In Bolivia, Y. patricia has been recorded in three localities from 1400 to 1600 m in the region of Yungas near La Paz and in the region of Santa Cruz ( Fig. 9 ).

Genetic distances ( Fig. 11 ). Distances within species ranged from 0 % to 3.1 %, and interspecific distances ranged from 4.2 % to 11.0 % (mean genetic distances among 14 Yphthimoides species in Table 2), overlapping with intraspecific distances of 5.3 % and 6.9 %, respectively, between specimens of the Y. renata "complex" from Brazil and Colombia, and from Brazil and Costa Rica. Genetic distances among individuals of Y. patricia and Y. manasses ranged from 8.8 to 9.7 %. In conjunction with the morphological differences, these results and a species phenogram ( Fig. 12 ) support Y. patricia and Y. manasses as distinct species.

Taxonomic comments. Hayward (1957) described ten new taxa of Bolivian Satyrinae , assigning all of them to the genus Euptychia Hübner, 1818 . The holotypes of six of those taxa ( E. susanna , E. pamela , E. patricia , E. phares boliviana , E. proxima and E. stella ) were stated to be deposited in the collection of the Museo del Instituto de Historia Natural "Sánchez Labrador" of San Miguel, Buenos Aires, Argentina. Decades later, part of the Lepidoptera collection of the Instituto was transferred to the Museo Argentino de Ciencias Naturales (MACN), Buenos Aires, Argentina (A. Roig, pers. comm. to GL). GL examined that collection in 2005 and found the holotypes of E. susanna and E. proxima . For E. patricia and E. phares boliviana , only the labels were found. No trace could be discovered regarding the holotypes of E. pamela and E. stella . Presumably, the holotypes of these last four names are lost. Hayward described both E. patricia and E. saltuensis ( Hayward, 1962) based mostly on the wing color pattern, although drawings of male genitalia were also provided for both taxa. Despite the genitalia drawings being somewhat rough, they are detailed enough as to show the two series of teeth that make the valvae of Y. patricia so distinctive from all other species of Euptychiina , and very similar to those illustrated in the original description of Euptychia saltuensis . It is quite possible that variation in wing pattern explains why Hayward proposed two names for the same species. The holotype of E. saltuensis is very similar to wet season forms of Y. patricia ( Fig. 1E ), while the holotype of Y. patricia (based on the text in the original description) appears to have been be more similar to dry season forms, as are the three individuals from Yungas, Bolivia deposited in the NHMUK (one of these, the neotype below designated, is shown in Fig. 1F ). Lamas (2004) listed both Y. patricia and E. saltuensis in Yphthimoides , with E. saltuensis as a nomen dubium and possible subjective synonym of Y. manasses . By comparing Hayward's male genitalia drawings of Y. patricia and E. saltuensis with dissections of several Yphthimoides species, in conjunction with DNA barcode analysis to determine the range of intraspecific wing pattern variation within species, we treat E. saltuensis as a junior synonym of Y. patricia (new synonym).

Remarks. After GL examined the specimens in the collection of the MACN (Buenos Aires, Argentina) and found only the labels of Euptychia patricia , we concluded that the holotype is lost. Therefore, a neotype of Euptychia patricia is hereby designated to clarify the taxonomic status of the species and ensure nomenclatural stability.

Neotype ♂ ( Fig. 1F ). In order to fix the identity of the name, one male deposited in the NHMUK collection, which was collected in the same general area as the holotype, is herein designated as the Neotype of Euptychia patricia Hayward, 1957 ; this specimen has the following four labels separated by transverse bars: \ BOLIVIA Adams, M.J. & Bernard, G.I BMNH(E) 1985-477 See Book L. Rh. A.32 \ Adams Series NE of La Paz: Above Chairo 1900m. 11.viii.85 \M85/630 \ BMNH(E) 1204805; three other labels will be added later:\ Neotype \ Neotype of Euptychia patricia Hayward, 1957 \ Barbosa and Freitas det. 2017 \.

Examined material. ARGENTINA. Holotype ♂ of Euptychia saltuensis , no data, SAT 252 ( IML).

BRAZIL. No locality — 1 ♂, 05.XI.2011 ( DNA voucher YPH-0049) , ZUEC LEP 9837 (ZUEC); São Paulo: Brotas— Campo Alegre , 750 m, 5 ♂ (1 without abdomen), 6 ♀ (2 without abdomen), 13-I-1963, H. Ebert leg., DZ 28.947 to 28.957 ( DZUP) . Itirapina— 750 m, 2 ♀, 15-VIII-1964, 2 ♀, 18-VIII-1968, H. Ebert leg., DZ 28.958 to

DZ 28.961 (DZUP), Estação Ecológica de Itirapina , 1 ♂, 19-VII-2002, K. S. Brown & A. V. L. Freitas leg. , ZUEC LEP 9836 ( ZUEC), 3 ♂, 19-VII-2002, K. S. Brown and A. V. L. Freitas leg. (ZUEC-AVLF), Cerrado da Graúna , 2 ♀, 26.IX.1990, 4 ♀, 21.I.1999, A. V. L. Freitas leg. (ZUEC-AVLF), 1 ♂ and 1 ♀, A-213, I-IV.1999 (ex-larva), 2 ♂ and 2 ♀, A-214, I-IV.1999 (ex-larva), 1 ♀, A-215, I-IV.1999 (ex-larva), (ZUEC-AVLF). Rio Claro— 1 ♀, 14- X-1962, DZ 28.962 ( DZUP). Santa Rita do Passa Quatro— Parque Estadual da Vassununga , Gleba Pé do Gigante , 1 ♀, 13-15.III.2003 , B786, A. V. L. Freitas leg. (ZUEC-AVLF). Minas Gerais: São Roque de Minas— Parque Nacional da Serra da Canastra 1 ♂, 14-VIII-1999, 1 ♂, 19-VIII-1999, 1 ♀, 16-X-1 999, O. Marini-Filho leg. ( UFMG). Barbacena— 1.100m , 1 ♀, 9-VIII-1951, 1 ♂ (without abdomen), 11-VIII-1951, 1 ♂ (without abdomen), 14-VIII-1 951, 1 ♂, 1 ♀, 16-VIII-1951, 3 ♂, 17-VIII-1951, 1 ♀, 17-VIII-1951, 1 ♀ 28-VIII-1951, 1 ♂, 2 ♀ (without abdomens), 29-VIII-1951, 2 ♂, 1 ♀ 1-IX-1951, 1 ♀ (without abdomen), 2-IX-1951, 1 ♂, 1 ♀, 3-IX-1951, 1 ♀ 30- IX-1951, 1 ♂ 15-VIII-1952, 1 ♂, 1 ♀, 18-VIII-1952 ,, H. Ebert leg., DZ 28.880 to 28.902 (DZUP). Barroso— 1 ♀, 27-VIII-1 952, H. Ebert leg., DZ 28.903 (DZUP). Corinto— 2 ♂, 7-IX-1979, C. Elias leg., DZ 28.904, DZ 28.905; Itumirim— 900m, 1 ♀, 7- II-1 964, Ebert leg., DZ 29.385 (DZUP). Itutinga— 11 ♀ (3 without abdomen), beginning - II-1956, 1 ♂, 3 ♀, 16-IX-1956, H. Ebert leg., DZ 28.906 to 28.920 ( DZUP). Lagoa Santa— 1 ♂, 2 ♀, 31-VIII-1 956, Becker leg. , DZ 28.921 to DZ 28.923 ( DZUP). Ouro Branco— 2 ♂ (without abdomens), 8-VIII- 1954, H. Ebert leg. , DZ 28.924, DZ 28.925 ( DZUP). Ouro Preto— Fazenda Barcellos , 1.100m, 2 ♀, 14-IX-1952, 1 ♀, middle (?) VII (?) — 1953, H. Ebert leg. , DZ 28.926 to 28.928 ( DZUP). Poços de Caldas— 1 ♂, II-1963, 1 ♀, III-1964, 3 ♂, 2 ♀, VIII-1964, 2 ♂, 17-XII-1966, H. Ebert leg. , DZ 28.929 to DZ 28.937 (DZUP), Serra do Cristo, 1 ♀, 14.II.1999, A. V. L. Freitas leg. (ZUEC-AVLF). Santana do Riacho— Serra do Cipó , 1.000-1.300m , 1 ♂, 30- I-1985, Mielke & Casagrande leg., DZ 28.938 (DZUP), 1 ♂, 10-VIII-2011 ( DNA voucher YPH-0279), 1 ♀, 19-I- 2012 ( DNA voucher YPH-0280), 1 ♀, 23-X-2012 ( DNA voucher YPH-0281), 1 ♀, 17-I-2012 ( DNA voucher YPH-0282), 1 ♀, 10-VIII-2011 ( DNA voucher YPH-0283), Marina Beirão leg., ZUEC LEP 9831 to ZUEC LEP 9835 ( ZUEC). São João Del Rei— 800 - 1.000 m, 3 ♂, 31-VII-1951, 3 ♂, 14-VII-1954, 1 ♀, 16-II-1956, 1 ♀, 2-IX- 1956, H. Ebert leg., DZ 28.939 to DZ 28.946 ( DZUP). Belo Horizonte – APE Manancial Cercadinho , 1 ♀, 09-II- 2007 ( DNA voucher YPH-0158), A. R. M. Silva leg , ZUEC LEP 9830 ( ZUEC), 1 ♀, 24.I.2007, 1 ♀ 23.I.2007, 1 ♀, 26.II.2007, 1 ♀, 02.IX.2007, A. R. M. Silva leg. (ZUEC-AVLF). Nova Lima— APE Manancial Mutuca, Parque Estadual da Serra do Rola Moça , 20°1’1”S 43º58’23”W , 1 ♂, 30.I.2008 ( DNA voucher PM10-08), A. R. M. Silva leg. (ZUEC-AVLF); Mato Grosso: Barra dos Bugres— km 363 estrada Cuiabá-Porto Velho , 650 m , 3 ♂, 5- VII-1972, Mielke & Brown leg., DZ 28.821, DZ 28.822, DZ 28.823 ( DZUP). Barra do Garças— Royal Base Camp, Serra do Roncador , 1 ♂, 1 ♀ VII-1968, Laroca & Azevedo leg., DZ 28.824, DZ 28.825 ( DZUP). Nova Xavantina— Bacaba , UNIMAT, 3 ♀, 17-19-VIII-1997, Mielke leg. , DZ 28.805, DZ 28.826, DZ 28.827 ( DZUP). Chapada dos Guimarães— 700m, 2 ♀, 27-VIII-1978 , H. Ebert leg., DZ 28.828, DZ 28.829 (DZUP), 2 ♂, 23/ 26- XI-2013 (DNA vouchers YPH-0486, YPH-0487), L. A. Kaminski leg., ZUEC LEP 9828, ZUEC LEP 9829 ( ZUEC). Diamantino— Fazenda São João , 300-400m, 1 ♀, 1-8- VIII-1974, 1 ♀, 28-VII-1975, H. & H. D. Ebert leg. , DZ 28.830, DZ 28.831 (DZUP), 1 ♀, 4-IX-1978, 1 ♀, 8-IX-1978, Mielke & Furtado leg., DZ 28.832, DZ 28.833 (DZUP), 1 ♀, 3-IX-1979, Furtado leg., DZ 28.834 ( DZUP). Alto Araguaia— 1 ♂, 05-XII-2009 ( DNA voucher YPH-0071), 1 unknown sex (abdomen destroyed), 05-XII-2009 ( DNA voucher YPH-0016), Dell’Erba R. & Zillmer E. D. leg. , ZUEC LEP 9826, ZUEC LEP 9827 ( ZUEC); Goiás: Goiás— 3 ♂, XII-1975, 1 ♀ 4-IX-1976, 2 ♂, 3 ♀, XII-1976, 2 ♂, I-1972 , D. Gifford leg., DZ 28.835 to DZ 28.845 ( DZUP), Serra Dourada , 1 ♀, 6-II-1980, 16°03’ S , 50°06 W, DZ 28.846 ( DZUP). Cristalina— 20 km East , 2 ♀, 24-I-1976 , DZ 28.847, DZ 28.848 (DZUP). Goiânia— 1 ♀, 31-VIII-1969, Ebert leg., DZ 28.849 ( DZUP). Rio Quente— Pousada do Rio Quente , 2 ♀, 12-II-1979, A. Raw leg. , DZ 28.850, DZ 28.851 ( DZUP); Distrito Federal: Brasília— 1 ♀, 8-IX-1965, 1 ♀, 19- IX-1965, 1 ♀, 11-IX-1968, 1 ♀, 12-IX-1968, 1 ♂, 20-IX-1968, 1 ♀, 31-VIII-1969 ,, Ebert leg., DZ 28.852 to 28.857 (DZUP), 1 ♂, XII-1971, Stawiarsky leg., DZ 28.858 (DZUP), 1 ♂, 11-VII-1978, 1 ♂, 9-VI-1980, Gifford leg., DZ 28.859, DZ 28.860 (DZUP), 1 ♀, IX-1978, Raw leg., DZ 28.861 ( DZUP), Fazenda Água Limpa , 3 ♂, 11- I-1976, 1 ♀, 16-I-1976, 2 ♀, 16-VIII-1 976, 1 ♀, 7-IX-1976, 4 ♂, 1 ♀, 29-VII-1978, 1 ♂, 16-VIII-1 976, 1 ♀, 18- IX-1976 , Gifford leg., DZ 28.862 to 28.875 ( DZUP), Fazenda Água Limpa, Universidade de Brasília , 1000 m.a.s.l., 7 ♂, IX-2005 (DNA vouchers YPH-0019, YPH-0024, YPH-0026, YPH-0030, YPH-0040, YPH-0041, YPH-0042), ZUEC LEP 9814, ZUEC LEP 9815, ZUEC LEP 9816, ZUEC LEP 9817, ZUEC LEP 9818, ZUEC LEP 9819, ZUEC LEP 9820 (ZUEC), 2 ♀, IX-2005 (DNA vouchers YPH-0018, YPH-0028), E. O. Emery leg., ZUEC LEP 9821, ZUEC LEP 9822 (ZUEC), 2 ♂, IX-2005 (genitalia prepared AVLF0007), 2 ♀, IX-2005, E. O. Emery leg. (ZUEC-AVLF), 1 ♂, 18.VII.2005, A. V. L. Freitas leg. (ZUEC-AVLF), Parque do Gama , 1.000 m, 1 ♂, 18-VI-1972, 2 ♂, 20-VI-1972 , Mielke & Brown leg., DZ 28.876, DZ 28.877, DZ 28.804 (DZUP). Planaltina— 1.000 m, 15°35’ S, 47°42 W, 2 ♂, 4-XII-1982, Mielke & Becker leg., DZ 28.878, DZ 28.879 (DZUP), Parque Nacional de Brasília , 1180 m.a.s.l., 15°36’57,17’’S 47°57’33,02’’W, 2 ♂, 22-XI-2009 ( DNA vouchers YPH-0116, YPH-0123), E. O. Emery leg , ZUEC LEP 9823, ZUEC LEP 9824 (ZUEC), Estação Ecológica Jardim Botânico , 1 ♂, 18-XII-2010, ( DNA voucher YPH-0159), A. V. L. Freitas leg , ZUEC LEP 9825 ( ZUEC), 3 ♂, 18-XII-2010, 2 ♀, 18-XII-2010, A. V. L. Freitas leg. (ZUEC-AVLF); Bahia: Palmeiras— Morro Pai Inácio , 800 - 1.200 m, 1 ♀, 2- XII-1997, Mielke & Casagrande leg., OM 47.355 ( DZUP). Mucugê— Trilha horizontal, 1.012 m, 13°3′12.24″S, 41°20′45.96″W , 1 ♀, 01-X-2017, E.P. Barbosa, T. Zacca & R.R. Ramos leg., ZUEC LEP 10497, Trilha vertical, 962m. a.s.l., 13°2′45.24″S, 41°20′54.6″W, 1 ♀, E.P. Barbosa, T. Zacca & R.R. Ramos leg. , ZUEC LEP 10498, Trilha da Cachoeira do Cardoso , 1.043 m, 13°2′50.64″S,41°20′ 26.88″W, 1 ♀, 02-X-2017, E.P. Barbosa, T. Zacca & R.R. Ramos leg., ZUEC LEP 10499 ( ZUEC). Rio de Contas— Campo do Queiroz , 1.450 m, 13°31’13.08’’S, 41°56’7.44’’W , 1 ♀, 29-IX-2017, E.P. Barbosa, T. Zacca & R.R. Ramos leg., ZUEC LEP 10500 ( ZUEC); Maranhão: Feira Nova de Maranhão— Retiro, 28 km W [30 km W Fortaleza dos Nogueiras ], 470 m, 3 ♀, 18-23- I-2010 , O. Mielke leg., DZ 28.815, DZ 28.816, DZ 28.817 ( DZUP), Fazenda Forquilha dos Brejos , 28 km E Feira Nova do Maranhão , 2 ♀, 14-21-VIII-2011, DZ 28.818, DZ 28.819 ( DZUP). Imperatriz— Excursão do Departamento de Zoologia , 1 ♀, 18-VII-1974, DZ 28.820 ( DZUP); Pará: Carajás— Serra dos Carajás , 1 ♀, 6-IX- 1978 , Gifford leg., DZ 28.814 ( DZUP). No state: B. P. (unknown locality), 4 ♀, II-1968 , Ebert leg., DZ 28.963 to 28.965, DZ 29.384 ( DZUP). — BOLIVIA. Santa Cruz— Chiquitos, Serranía de Santiago , 1 ♂, 9.VIII.2001 , Y. Gareca & T. Riegel leg. ( MUSM). La Paz— Chairo, NE La Paz above Chairo , 1 ♀, 11-VIII-1985 , 1600 m,1 ♀, 11- VIII-1985, 1900 m, Adams series, Adams, M.J. & G. L. Bernard leg., 1985-477 (see Book L. Rh. A 32), M85/629, M85/633, BMNH(E) 1204803, BMNH(E) 1204804 (NHMUK).

Figures

FIGURE 1. Yphthimoides patricia. A–B—Male (dorsal, ventral), Brasília, Distrito Federal, 20.VI.1972 (DZ 28.804); C–D— Female (dorsal, ventral), Nova Xavantina, Mato Grosso, 17–19.VIII.1997 (DZ 28.805). E—Male holotype of Euptychia saltuensis (dorsal left, ventral right); no locality data (IML). F– Male Neotype (here designated) of Y. patricia from Chairo, La Paz, Bolivia, 11.VIII.1985 - BMNH(E) 1204805 (NHMUK) (dorsal left, ventral right). Scale bar = 1 cm. E. Photo by L. A. Kaminski.

FIGURE 2. Yphthimoides manasses. A—Male (dorsal, ventral), Ibateguara, Alagoas (ZUEC-LEP 10476); B—Female (dorsal, ventral), Ibateguara, Alagoas (ZUEC-LEP 10475); C—Male (dorsal, ventral), Camaragibe, Pernambuco (DZUP); D—Female (dorsal, ventral), São José de Mipibu, Rio Grande do Norte (DZUP). Scale bar = 1 cm.

FIGURE 3. Male genitalia of Yphthimoides patricia (A–D) and Yphthimoides manasses (E–H). Y. patricia: A—Lateral view; B—Dorsal view; C—Ventral view (Detail of valvae apex and uncus); D—Aedeagus (lateral view). Y. manasses: E—Lateral view; F—Dorsal view; G—Dorsal view (Detail of valvae apex and uncus); H—Aedeagus (lateral view). Abbreviations: ae, aedeagus; gn, gnathos; sa, saccus; te, tegumen; va, valva; un, uncus.

FIGURE 4. Yphthimoides patricia: A—Eye and palpus details; B—detail of antennal shaft; C—Antennal club; D –Midleg. All from a male specimen.

FIGURE 5. Yphthimoides patricia. A—Male wing venation; B—Female wing venation; C—Male foreleg; D—Male palpus; E—Female foreleg.

FIGURE 6. Details of the male genitalia of Yphthimoides patricia in scanning electron microscopy. A—ventral-anterior view of uncus apex; B—detail of valva tip in ventral view showing the distinctive 'teeth'.

FIGURE 7. Males (A–F) and females (G–L) of Yphthimoides patricia showing variation in ventral wing pattern. A—Brasília, Distrito Federal, 18.XII.2010; B—Brasília, DF, 18.XII.2010; C—Itirapina, SP; II.1999; D–F—Itirapina, São Paulo, 19.VII.2002; G—Itirapina, São Paulo, II.1999; H—Brasília, Distrito Federal, 18.XII.2010; I—Santa Rita do Passa Quatro, São Paulo, 13.III.2003; J—Belo Horizonte, Minas Gerais, 26.II.2007; K—Itirapina, São Paulo, IX. 1990; L—Brasília, Distrito Federal, IX.2005; A, B, C, G, H—wet season forms; D, E, F, K, L = dry season forms, I, J = intermediate forms. Scale bar = 1 cm.

FIGURE 8. Yphthimoides patricia. A—Second instar, dorsal; B—Third instar, dorsal; C—Fourth (paler) and Fifth (darker) instars, dorsal; D—Two fifth (last) instar larvae, dorsal; E–F—Pupae (two different individuals), lateral; G—Pupa, ventral.

FIGURE 9. Map showing localities (solid circles) for Yphthimoides patricia in Brazil and Bolivia. The gray shaded area corresponds to the limits of the cerrado vegetation.

FIGURE 10. Habitat and live adults of Yphthimoides patricia. A—General view of the habitat in Brasília, Distrito Federal; B—Close view of the cerrado in Itirapina, São Paulo; C–D—Male and female, respectively, from Brasília, Distrito Federal.

FIGURE 11. Frequency distribution of pairwise individual genetic distances within (grey) and between (black) fourteen species of Yphthimoides.

FIGURE 12. Limits among fourteen species of Yphthimoides based on DNA barcode sequences (ca. 658bp of cox1) obtained by a Neighbor-Joining analysis. Numbers above branches are bootstrap values. Low support values were not showed.