Euptychia stigmatica Godman, 1905

Nakahara, Shinichi, Kleckner, Kaylin, Barbosa, Eduardo P., Lourenco, Giselle M., Casagrande, Mirna M., Willmott, Keith R. & Freitas, Andre V. L., 2023, Reassessment of the type locality of Euptychia stigmatica Godman, 1905, with the description of two new sibling species from Amazonia (Lepidoptera, Nymphalidae, Satyrinae, Satyrini), ZooKeys 1167, pp. 57-88 : 57

= Euptychia stigmatica Godman, 1905

Fig. 2C-F

Euptychia stigmatica : Godman 1905: 186, pl. X[10], fig. 2 [original description]; Weymer 1911: 218; [in the synonymy of Euptychia cyanites Butler, 1871]; Riley and Gabriel 1924: 55; Gaede 1931: 444 [in the synonymy of Papilio crantor Fabricius, 1793]; D’Abrera 1988: 766 [in the synonym of E. cyanites ]; Lamas 2004: 217 [in the synonymy of E. cyanites Butler, 1871].

Type locality.

"Entre Rios, Argentina" = Três Rios, Rio de Janeiro, Brazil (see discussion below). Holotype (fixed by monotypy) (Fig. 2C-F ): male with following labels written verbatim separated by double forward slashes: //BMNH(E) 1267020//Type//Type of Species// Sept.//Entre Rios, Argentina H. H. Smith//Godman-Salvin Coll. 1904. - 1. Euptychia stigmatica , Godm.//B. M. TYPE No. Rh.3167 Euptychia stigmatica , ♂ Godm.//(NHMUK).

Systematic placement and diagnosis.

The genus Caeruleuptychia is found as a clade in the so-called Splendeuptychia clade of Euptychiina ( Espeland et al. 2019; 2023). Within Caeruleuptychia , C. cyanites is recovered as a member of the caerulea clade based on the ML tree inferred from COI barcode data ( Nakahara et al. 2022: fig. 1, as Caeruleuptychia sp. nov. 2). Despite the caerulea clade not being found as monophyletic in the present work, preceding studies (e.g., Nakahara et al. 2022) did recover this clade as a natural group. Caeruleuptychia cyanites is recovered as a sister taxon to C. harrisi sp. nov. with a strong support in the present work (Fig. 1A ), where this clade ( C. cyanites + C. harrisi sp. nov.) is found as a sister to C. coelestis (Butler, 1867), and furthermore, (( C. cyanites + C. harrisi sp. nov.) + C. coelestis ) is recovered as a sister to ( C. aemulatio sp. nov. + C. glauca ). The mean inter-specific pairwise distance between C. cyanites and C. harrisi sp. nov. was 1.6%, and mean infra-specific pairwise distance of 0.1% (n = 6) ( C. cyanites ) and 0.2% (n = 2) ( C. harrisi sp. nov.).

Female specimens are currently known only for C. cyanites among taxa discussed herein, thus the diagnostic characters provided below are restricted to male individuals. Caeruleuptychia cyanites is distinguished from C. harrisi sp. nov. by its larger forewing length: 22.8-26.1 mm, mean 24.66 mm (n = 16) for C. cyanites ; 21.0-22.4 mm, mean 21.38 mm (n = 5) for C. harrisi sp. nov. Caeruleuptychia cyanites is further distinguished from C. harrisi sp. nov. by possessing a larger scent patch in the dorsal forewing cell 2A, where this patch encompasses ca. one-third of the forewing inner margin (25-31%, mean 28.0% (n = 16)), whereas this patch encompasses less than one-fifth of the forewing inner margin (14-19%, mean 16.2% (n = 5)) in C. harrisi sp. nov. (see Table 1 ). The length of this dorsal forewing scent patch occupies ca. one-fourth of the forewing inner margin in C. aemulatio sp. nov. (21-27%, mean 23.7.4%, n = 6; Table 1 ). We created bivariate scatterplots based on these forewing measurements to graphically delineate these three species discussed above (Fig. 1B ). Caeruleuptychia cyanites is further distinguished from C. aemulatio sp. nov. by often having the dorsal hindwing cell Cu1 adorned with a black spot, whereas this spot is absent in C. aemulatio sp. nov.; by having broader dark bands on the ventral surface (most pronounced in the ventral forewing umbra); and by cell Cu2 on the ventral forewing having paler blue scaling than the remainder of the wing, with this color extending slightly into the posterior half of cell Cu1, with the middle of cell Cu2 having even paler, whitish blue scales that might represent androconial scales, whereas similar pale whitish blue scales in C. aemulatio sp. nov. are confined to cell 2A. Nevertheless, given the subtlety of these characters and slight variation, it is advisable to use these characters in combination in order to distinguish C. cyanites from C. aemulatio sp. nov. Caeruleuptychia cyanites is further distinguished from C. harrisi sp. nov. by the ventral margin of the tegumen exhibiting an upwards concavity in lateral view, as well as an apical process of the valva terminating in a broader point in lateral view. In addition, the male of C. coelestis is readily distinguishable from these three taxa discussed above by the absence of a forewing scent patch.

Variation.

Black spot in the dorsal hindwing cell Cu1 is variable in male specimens in size, namely clearly present in many individuals although reduced or apparently absent in some specimens (e.g., a pair at FLMNH; lectotype of E. cyanites ).

Examined material

(21 males and 18 females). Males: With the following labels written verbatim separated by double forward slashes: São Paulo - //Casa Br[anca]. G. (MfN); //Prov. S. Paulo Brasil mer. Littke. // Most probably ♂ of Euptychia ziza, Butl.// genitalia vial M- 9125 ♂ Lee D. Miller (MfN); Minas Gerais - //P[arque]e[stadual do]r[io]d[oce] - 05/VIII/15 Trilha 2 - Arm 3D Lagoa Bonita// BLU 778 [molecular voucher]// (ZUEC); // BRAZIL, Minas Gerais, Timoteo Pq. Est. do Rio Doce 05.VIII.2015 Lourenço G.M., Soares G.R. & Palacio T. leg.// YPH-0854 [molecular voucher]// ZUEC-LEP 12054// (ZUEC); // BRAZIL, Minas Gerais, Timoteo Pq. Est. do Rio Doce 06.X.2015 Lourenço G.M., Soares G.R. & Palacio T. leg.// ZUEC-LEP 12055// (ZUEC); // BRAZIL, Minas Gerais, Timoteo Pq. Est. do Rio Doce 04.X.2015 Lourenço G.M., Soares G.R. & Palacio T. leg.// ZUEC-LEP 12064// (ZUEC); // BRAZIL, Minas Gerais, Timoteo Pq. Est. do Rio Doce 04.X.2015 Lourenço G.M., Soares G.R. & Palacio T. leg.// ZUEC-LEP 12056// (ZUEC); // BRAZIL, Minas Gerais, Timoteo Pq. Est. do Rio Doce 14.IX.2015 Lourenço G.M., Soares G.R. & Palacio T. leg.// ZUEC-LEP 12057// (ZUEC); // BRAZIL, Minas Gerais, Timoteo Pq. Est. do Rio Doce 12.IX.2015 Lourenço G.M., Soares G.R. & Palacio T. leg.// ZUEC-LEP 12058// (ZUEC); // BRAZIL, Minas Gerais , Timoteo Pq. Est. do Rio Doce 15.IX.2015 Lourenço G.M., Soares G.R. & Palacio T. leg.// ZUEC-LEP 12059// (ZUEC); // 29-I-3-II-2003. ESTAÇÃO BIOLÓGICA DE CARATINGA, CARATINGA, MG, 400 m MIELKE & CASAGRANDE LEG.// (DZUP); Espírito Santo - // BRAZIL, Espírito Santo , Sooretama, Reserva Biologica de Sooretama , 19°02'07.0"S, 40°09'30.2"W, 18.VIII.2018, Santos J.P., Rosa A.H.B., Machado P.A. & Lopes A.C. leg.// LBR0654 [molecular voucher]// (ZUEC); // ESPÍRITO SANTO RES. ECOLÓGICA SOORETAMA 19°03'25"S, 40°08'50"W 19-26 - II - 2013 MIELKE & CASAGRANDE LEG.// DZ 40.523// BC-DZ Willmott 8 [molecular voucher]// (DZUP); BRAZIL, ESPÍRITO SANTO RES. ECOLÓGICA SOORETAMA 19°03'25"S, 40°08'50"W 19-26 - II - 2013 MIELKE & CASAGRANDE LEG.// DZ 40.372// BC-DZ Willmott 6 [molecular voucher]// (DZUP) ; BRAZIL, ESPÍRITO SANTO RES. ECOLÓGICA SOORETAMA 19°03'25"S, 40°08'50"W 19-26 - II - 2013 MIELKE & CASAGRANDE LEG.// DZ 40.373// (DZUP) ; BRAZIL: ESPIRITO SANTO: Itaguassu 1x.1971 Paulo Cesar Elias // A. C. Allyn Acc. 1971-38// Genitalic vial SN-19-13// Allyn Museum photo No. 073075-11// FLMNH 1138898// (FLMNH) ; BRAZIL, BAHIA, CAMACAN RESERVA SERRA BONITA 15°23'S, 39°33'W, 40°08'50" 3-10-XII-2016 200 m MIELKE, CARNEIRO, DIAS, DOLIBAINA & SANTOS LEG.// DZ 40.512// (DZUP) ; BRAZIL, BAHIA , CAMACAN RESERVA SERRA BONITA 15°23'S, 39°33'W, 40°08'50" 3-10-XII-2016 200m MIELKE, CARNEIRO, DIAS, DOLIBAINA & SANTOS LEG.// DZ 40.503// (DZUP); BRAZIL, BAHIA, CAMACAN RESERVA SERRA BONITA 15°23'S, 39°33'W, 40°08'50" 3-10-XII-2016 200m MIELKE, CARNEIRO, DIAS, DOLIBAINA & SANTOS LEG.// DZ 40.502// BC-DZ Willmott 9 [molecular voucher]// (DZUP); // CONCEIÇÃO BARRA ES - BRASIL 10-IX-1969 C.& C.T.Elias lg// GEN. PREP. BILOTA 1997// DZ 5268 ♂ Caeruleuptychia cyanites I. G. BILOTA DET. 1997// DZ 5.268// (DZUP); Bahia - // Rio. R. Hewitson Coll. 79-69. Euptychia Coelestis Butl. 1.// BMNH(E) 1717835// (NHMUK). Females: with the following labels written verbatim separated by double forward slashes: Rio de Janeiro - //ALCINDO GUANA-BARA-E. F.C. B. 15-IV-1940 P. SANDIG LEG// Ex Col. Gagarin // alcindo guanabara E.F.C.B. Paulo Sandig 15-4-1940 // DZ 40.381// (DZUP); // Novo Friburgo. // genitalia vial M- 9126 ♀ Lee D. Miller // coll. Sommer (MfN); Minas Gerais - BRAZIL, Minas Gerais, Timoteo Pq. Est. do Rio Doce 05.X.2015 Lourenço G.M., Soares G.R. & Palacio T. leg.// ZUEC-LEP 12063// (ZUEC); // BRAZIL, Minas Gerais, Timoteo Pq. Est. do Rio Doce 15.IX.2015 Lourenço G.M., Soares G.R. & Palacio T. leg.// ZUEC-LEP 12062// (ZUEC); // BRAZIL, Minas Gerais, Timoteo Pq. Est. do Rio Doce 06.XI.2015 Lourenço G.M., Soares G.R. & Palacio T. leg.// ZUEC-LEP 12061// (ZUEC); // BRAZIL, Minas Gerais , Timoteo Pq. Est. do Rio Doce 11.V.2016 Lourenço G.M., Soares G.R. & Palacio T. leg.// ZUEC-LEP 12060// (ZUEC); Espírito Santo - // BRAZIL, ESPÍRITO SANTO RES. ECOLÓGICA SOORETAMA 19°03'25"S, 40°08'50"W 19-26 - II - 2013 MIELKE & CASAGRANDE LEG.// (DZUP) ; BRAZIL: E. SANTO Baixo Guandu x.1971 P. C. Elias // A. C. Allyn Acc. 1971-47// Genitalic vial SN-19-14// Allyn Museum photo No. 073075-10// (FLMNH); // BRAZIL, ESPÍRITO SANTO RES. ECOLÓGICA SOORETAMA 19°03'25"S, 40°08'50"W 19-26 - II - 2013 MIELKE & CASAGRANDE LEG.// DZ 40.522// (DZUP); // BRAZIL, ESPÍRITO SANTO RES. ECOLÓGICA SOORETAMA 19°03'25"S, 40°08'50"W 19-26 - II - 2013 MIELKE & CASAGRANDE LEG.// DZ 48.799// BC-DZ Willmott 7 [molecular voucher]// (DZUP); // BRAZIL, ESPÍRITO SANTO RES. ECOLÓGICA SOORETAMA 19°03'25"S, 40°08'50"W 19-26 - II - 2013 MIELKE & CASAGRANDE LEG.// DZ 40.513// (DZUP); // BRAZIL, ESPÍRITO SANTO RES. ECOLÓGICA SOORETAMA 19°03'25"S, 40°08'50"W 19-26 - II - 2013 MIELKE & CASAGRANDE LEG.// DZ 48.819// (DZUP); // BRAZIL, ESPÍRITO SANTO RES. ECOLÓGICA SOORETAMA 19°03'25"S, 40°08'50"W 19-26 - II - 2013 MIELKE & CASAGRANDE LEG.// DZ 48.809// (DZUP); // BRAZIL, ESPÍRITO SANTO RES. ECOLÓGICA SOORETAMA 19°03'25"S, 40°08'50"W 21-25 - I - 2014 MIELKE & CASAGRANDE LEG.// DZ 40.363// (DZUP); // BRAZIL, ESPÍRITO SANTO RES. ECOLÓGICA SOORETAMA 19°03'25"S, 40°08'50"W 19-26 - II - 2013 MIELKE & CASAGRANDE LEG.// DZ 40.351// (DZUP); // Conceicao da Barra Esp [ írito]. Santo Brazil 18-VI-1969 C. & C. T. Elias leg.// DZ 40.352// (DZUP); // CONCEIÇÃO BARRA ES - BRASIL 18-X-1968 C.& C.T.Elias leg// DZ 5.269// DZ 5269 ♀ Caeruleuptychia cyanites I. G. BILOTTA DET. 1997// GEN. PREP. BILOTTA 1997// (DZUP); Bahia - // BRASIL: BA, Camacan Res. Serra Bonita 15°23'S- 39°33'W 800m, xi. 2012 V.O. Becker Col. // (VOB); Rio. Hewtson Coll. 79-69. Euptychia Coelestis Butl. 2. [ “Rio” on the underside]// BMNH(E) 1717836// (NHMUK) .

Other examined material

(3 females). With the following labels written verbatim separated by double forward slashes: São Paulo (?) - Araras (S.P.) 600 m, 11.11.65 Ebert// Colecão H.Ebert// DZ 40.371//(DZUP); //Araras (S. Paulo) 600 m 23.11.68 Ebert// Colecão H.Ebert// DZ 40.361//(DZUP); //Araras (S. Paulo) 600 m 23.11.68 Ebert// Colecão H.Ebert// DZ 40.362//(DZUP).

Distribution

(Fig. 5 ). This species is known from the states of Bahia, Espírito Santo, Minas Gerais, Rio de Janeiro, and São Paulo, all situated in southeastern Brazil. Records from Araras, São Paulo are likely erroneous.

Biology

(Fig. 6 ). Based on recent records, C. cyanites is associated with areas of well-preserved old growth Atlantic Forest. In a 12-month trap study in PERD, Minas Gerais, a total of 34 individuals were captured, 26 captured in the canopy and 8 in the understory, indicating a clear preference for the upper stratum (X2 = 9.529, P <0.0020, DF = 1). The sex could be attributed to 33 of the captured individuals, totaling 18 males and 15 females, a sex ratio not significantly different from 1:1 (1:0.83, X2 = 0.27, P = 0.6015, DF = 1). Most records (32 of 34) were concentrated from August to November/2015. In addition, most individuals (n = 20) were captured in the ecotone between forests and lakes, a habitat with high sunlight availability resulting in the formation of a "brought low canopy" (5-15 m high); the remaining individuals were captured in the forest interior (n = 13) and a single individual was captured in the forest edge. General behavior, immature stages and host plants are unknown.

Figures

Figure 1. A maximum likelihood tree (LnL = - 2678.083) showing the phylogenetic relationships among Caeruleuptychia taxa selected for this study. Numbers beside branches are UFBoot / SH-aLRT values; colored tips represent taxonomic hypotheses proposed in the present study B the differentiation of three Caeruleuptychia species (C. cyanites, C. harrisi sp. nov., C. aemulatio sp. nov.) by forewing length (mm) and scent patch percentage. Each species is denoted by a unique color and shape with a single point representing a single specimen. The holotype (HT) of each name and the lectotype (LT) of C. cyanites are denoted by a black border and the corresponding letters.

Figure 2. Lectotype of E. cyanites (NHMUK) A in dorsal view B in ventral view, with associated labels to the right. Holotype of E. stigmatica (NHMUK) and its original description C in dorsal view D in ventral view, with associated labels to the right E text of the original description by Godman (1905: 186) F illustration associated with Godman (1905: pl. 10, fig. 2).

Figure 5. Distribution map for three Caeruleuptychia species discussed herein. Key: * denotes location of Tres Rios, Rio de Janeiro (= hypothesized type locality of E. stigmatica in the present study);? denotes dubious records from Araras, Sao Paulo.

Figure 6. Habitat pictures for two Caeruleuptychia species discussed herein A interior of ecotone habitat of C. cyanites at PERD with trap pictured B general view of PERD C habitat of C. cyanites at Camacan, Bahia D habitat of two Brazilian males of C. harrisi sp. nov. (Photograph credit: Bruno Ferreira).