Splendeuptychia tupinamba Freitas, Huertas & Rosa, 2021

Rosa, Augusto H. B., Huertas, Blanca, Willmott, Keith R., Barbosa, Eduardo P., Machado, Patrícia A., Mielke, Olaf H. H., Canaan, Carlos H. P. & Freitas, André V. L., 2021, Fifty years without a name: a new species of Splendeuptychia Forster (Lepidoptera: Nymphalidae: Satyrinae), Zootaxa 5061 (1), pp. 95-114 : 97-104

Splendeuptychia tupinamba Freitas, Huertas & Rosa , sp. nov.

( Figs. 1–3 , 4 , 7 )

urn:lsid:zoobank.org:act:CD4F52CF-3EA2-4A95-B0AD-F8CF3CB76EF1

Euptychia naeli D’Abrera, 1988: 770-771 , nomen nudum.

Euptychia naeli Brown & Ebert : Kochalka et al. 1996: 212, nomen nudum.

Splendeuptychia naeli [n. sp.] Lamas, MS: Lamas, 2004: 101, nomen nudum.

Splendeuptychia ‘ naeli ’ Brown & Ebert . Huertas 2014: 127, 161–163, nomen nudum.

Magneuptychia segesta [misidentification]: Garwood et al. 2007: 119; Garwood et al. 2009: 157.

Splendeuptychia doxes [misidentification]: Klimaitis et al. 2018: 203.

Diagnosis. This species is distinguished from other species in the clade containing the type species of Splendeuptychia ( Euptychia ashna Hewitson, 1869 ) by the combination of the following characters: no distinct white discal band on the VHW (present in all other species except Nubila purusana (Aurivillius, 1929)) ; uneven yellow surrounding the VHW ocelli which is expanded basally (an even yellow ocellar ring in all other species except Splendeuptychia ambra (Weymer, [1911])) ; a distinct, complete dark discal line on the VHW (incomplete or indistinct in S. ashna and S. ambra ); elongate ocelli in cells M 2 -M 3 and M 3 -CuA 1 (more circular in S. ashna and S. ambra ) on the VHW; and the yellow ocellar rings surrounding the VHW ocelli in cells CuA 1 -CuA 2 and CuA 2 -2A separate (fused in N. purusana and Nubila moderata (Weymer, 1911)) .

Description. Male ( Figs. 1 A–B , 2 A–C , 3 A–D ): Antenna dark brown, 8–9 mm in length (n = 12), extending to mid-costa, with 34 antennomeres, 11 of which form a well-defined club. FW length 17–19 mm (mean = 18.1 mm, SD = 0.67, n = 15); HW length 13–15 mm (mean = 14.1 mm, SD = 0.60, n = 14).

VFW ground color brown with marginal and two sub-marginal dark brown lines, as well as two further uneven slightly rufous dark brown discal and sub-discal lines. VHW with two slightly rufous dark brown discal and sub-discal narrow lines, the last forming a border between a region of brown and beige or cream basal coloration.

A series of six ocelli at submarginal region in Rs-M 1 (ocellus 1), M 1 -M 2 (2), M 2 -M 3 (3), M 3 -CuA 1 (4), CuA 1 -CuA 2 (5), CuA 2 -2A (6); ocelli 1, 2, 5, and 6 black, with a tiny white pupil and bordered with yellow, ocelli 5 similar to previous one but with a double white pupil, ocelli 3 and 4 as blurred silverish lines with a more diffuse and broader yellow edge merging into edges of adjacent ocelli. One marginal and two sub-marginal dark brown lines are present, with beige basal coloration in this region. DFW is essentially plain brown. On DHW, there are one marginal and two sub-marginal dark brown lines and vestiges of central wing bands and dusky markings reflecting black-centered ocelli. DFW of females has light yellow borders towards base.

Male genitalia. ( Fig. 2 A–C ): Tegumen short and rounded in dorsal view; valva sub-triangular, with a narrow apex curved inwards, covered by long hair-like setae latero-ventrally, and short setae at inner side; uncus straight and narrow; gnathos as paired long and pointed processes, half the size of valvae; saccus short and club-shaped in dorsal view, ending in a rounded end; aedeagus straight, cylindrical, almost same length as valva; tiny cornuti present in two elongate patches, one dorsal and one ventral.

Female ( Fig. 1 C–D , 7 C–D ): Antenna dark brown, 8 mm in length (n = 3), extending to mid-costa, with 36 antennomeres, 12 of which form a well-defined club. FW length 18–19 mm (mean = 18.2 mm, SD = 0.58, n = 5); HW length 14–15 mm (mean = 14.2 mm, SD = 0.48, n = 5). In a photograph of a live female individual, eyes appeared to be bluish and antennal tips red ( Fig. 7 C–D ).

Taxonomy and variation. D’Abrera (1988) figured a specimen from the NHMUK under the name “ Euptychia naeli Brown & Ebert ”, believing it to represent a paratype of that putative name based on a label attached to that specimen. However, Brown & Ebert never published a description and so Euptychia naeli D’Abrera, 1988 becomes a nomen nudum, since it was not accompanied by any description ( Lamas 2004). There is some variation in the VW pattern of this species ( Fig. 3 A–D ), especially in the size and shape of VHW ocelli. Individuals from localities in the Amazon region tend to show larger ocelli and the cream coloration on the VHW extends basal of the dark discal line. Extreme variations, such as shown on Fig. 3C, D , appear rather distinct, but they are connected by intermediate forms (e.g., Fig. 3A, B ) and molecular data showed no substantial divergence between such forms ( Fig. 4 ).

Type material. Holotype. Male ( Fig. 1 A–B ): Deposited at ZUEC. With the following six labels separated by transverse bars: / HOLOTYPUS / Brazil, Minas Gerais, Divinópolis , -20.197765° -44.927279°, 755 m, 04.II.2018, A[ugusto]. H[enrique]. B[atista]. Rosa & E[lisa]. K[aren]. S[ilva]. Ramos leg. / ZUEC LEP 11126 / AHBR 107/ DNA voucher BPU 028 / Holotypus Splendeuptychia tupinamba Freitas, Huertas & Rosa det. 2020 /.

Allotype. Female ( Fig. 1 C–D ): Deposited at ZUEC (this specimen was one of the proposed paratypes as indicated by the Keith Brown’s handwritten note “PT” on the envelope). With the following four labels separated by transverse bars: / ALLOTYPUS / Paraopeba, M [inas] G[erais], BR[ASIL] Flona de Paraopeba ( EFLEX) 19°15’S 44°24’W 750m 11.II.1970 Brown Jr., K [eith]. S[palding]. leg. / ZUEC LEP 11132 / Allotypus Splendeuptychia tupinamba Freitas, Huertas & Rosa det. 2020 /.

Paratypes. 23 males and 8 females (* = genitalia dissected): ARGENTINA: Misiones: General Manuel Belgrano, Reserva Privada Yacutinga , 17.X.2003, 1 male, 10.X.2004, 1 male, Ezequiel O. Núñez Bustos leg., ( MACN) ; BRAZIL: Maranhão: Imperatriz , 12.VIII.1974, 1 female *, Exc. Dept. Zool. leg., DZ 5.496 ( DZUP) . Mato Grosso: Alta Floresta, Cristalino Lodge, Restaurante , 17.II.2016, 1 male *, L. L. Mota leg., ZUEC-LEP 11117 ( ZUEC) ; Barra do Garças, Km 12 Torixoréu , 19.IV.1978, 1 female, d. s. f. leg., ex. coleção Gifford, DZ 29.325 ( DZUP) ; Cáceres , 16.XI.1984, 1 female *, Buzzi, Mielke, Elias & Casagrande leg., Proj. Polonoroeste, BC-DZ Willmott 152, DZ 5.487 ( DZUP) ; Diamantino, Vale da Solidão , 6.IV.1996, 1 male, E. Furtado leg., Nº 8301 ( CEEF) , 7.IV.1996, 1 male, E. Furtado leg., Nº 8302 ( CEEF) . Distrito Federal: Brasília, Brasília Country Club , 15°56’S 47°59’W, 7.II.1970, 1 male, K. S. Brown Jr. leg., ZUEC-LEP 11135 ( ZUEC) . Goiás: Piracanjuba, Topo de morro a leste do Km 85 a sul de Goiânia, 24.V.1971, 1 male, K. S. Brown Jr. leg., ZUEC-LEP 11136 ( ZUEC) . Minas Gerais: Curvelo , 13–14.XII.1968, 1 female, no collector, Coleção H. Ebert, DZ 29.326 ( DZUP) ; Divinópolis , -20.197765° -44.927279°, 04.II. 2018, 755 m, 6 males and 1 female, A. H. B. Rosa & E. K. S. Ramos leg., ZUEC-LEP 11118, ZUEC-LEP 11119, ZUEC-LEP 11120, ZUEC-LEP 11121, ZUEC-LEP 11122, ZUEC-LEP 11123*, ZUEC-LEP 11124 ( ZUEC) , 1 male, DNA Voucher BPU 027, AHBR 106, ZUEC-LEP 11125 ( ZUEC) , 1 male, DNA Voucher BPU 029, AHBR 108, ZUEC-LEP 11127 ( ZUEC) ; Paraopeba , 19°17’S, 44°25’W, 750m, 10.II.1970, 1 male, K. S. Brown Jr. leg., 10.2.1970 , Paratype, Splendeuptychia naeli sp n Paratype Brown & Ebert; Brit. Mus. 1970-473, VIAL 8933, BMNH(E)# 784853 ( NHMUK) ; Paraopeba, Flona de Paraopeba ( EFLEX) , 19°15’S 44°24’W, 10.II. 1970, 750 m, 3 males, K. S. Brown Jr. leg., ZUEC-LEP 11128, ZUEC-LEP 11129, ZUEC-LEP 11130* ( ZUEC) , 11.II. 1970, 750m, 3 females, K. S. Brown Jr. leg., ZUEC-LEP 11131, ZUEC-LEP 11133, ZUEC-LEP 11134 ( ZUEC) (all these specimens were indicated as possible paratypes by Keith Brown’s handwritten notes “PT” on the envelopes) . Paraná: Alto Paraíso, Faz. Lupus , 23°25’19’’S 53°49’49’’W, 4.X.2012, 1 male *, LABLEP leg., BC-DZ Willmott 208, DZ 31.469 ( DZUP) . Rondônia: Cacaulândia, 67 Km S Ariquemes, Linea C-10, 5 km S Cacaulândia , 14.VIII.1993, 1 male, O. Gomes leg., TRAP # 6, 2nd growth, DNA voucher LEP-58093, FLMNH-MGCL Specimen 278932 ( FLMNH) ; Cacaulândia 17.XII.1994, 1 male *, O. Gomes leg., TRAP # 8, 2nd growth, G. T. Austin Coll. MGCL Acc 2004-5, Genitalic Vial KW-17-91 K.R. Willmott, FLMNH-MGCL Specimen 297213 ( FLMNH) . PARAGUAY: Concepción, San Carlos, ca Río Apa, bosque Ribereño , 28.I.1992, 1 individual (no abdomen), JK, CA, BB, PF leg., Euptychia naeli, Brown & Evert [sic.], Aguilar det. 1992 ( MNHNPY) . No country, no specific locality, FT 5, 1.VII.1995, 1 male, DNA voucher LEP-58092, FLMNH-MGCL Specimen 278933 ( FLMNH) .

Etymology. This new species name is proposed in tribute to the Tupinambás, one of the various Tupi ethnic Brazilian natives. Originally the Tupinambás were distributed in two regions in coastal Brazil, the first extending from São Francisco river to central Bahia (the “Recôncavo” region) and the second from Rio de Janeiro to northern São Paulo. The name comes from the Tupi Tubüb-abá, meaning “descendants of the first fathers” or “all from the Tupi family” ( Bueno 1999; Navarro 2013). The name should be treated as a neuter noun in apposition.

Geographic distribution and habitat. This species is known from various localities in the Brazilian states of Maranhão , Rondônia , Mato Grosso , Goiás , Minas Gerais , Paraná and in the Distrito Federal , and also from Paraguay (Concepción) and Argentina ( Misiones ), at elevations from 120 to 1160 m ( Figs. 5 , 6 A–F ). Most localities are in the Cerrado domain (in forest fragments), some surrounded by open savannas, but there are also records from localities in the rain forests of southwest Amazonia and in the Atlantic Forest ( Figs. 5 , 6 E , 7 A–B ). This species has been recorded in disturbed forests and in medium to small fragments surrounded by urban and agricultural areas ( Figs. 6 C–F , 7 B ), suggesting that it is relatively tolerant of disturbance.

Behavior and natural history. Based on collecting records and field observation, adults are possibly multivoltine, with records from all months of the year except September ( Table 3 ). Although the geographic distribution of S. tupinamba sp. nov. is very broad, specimens are scarce in collections, perhaps partly associated with the dull appearance of this species, with the exception of ZUEC, which has 20 individuals (see data of the type series). Moreover, abundance is not apparently associated with habitat quality; for example, S. tupinamba sp. nov. was remarkably common in a small and highly impacted forest fragment in Divinópolis (Minas Gerais) ( Figs. 6 F and 7 B ), with about 15 individuals easily observed during a field trip in February, 2018. Adults were observed perched on leaves ( Fig. 7 C–D ) and flying low, usually 50 cm above ground between grass tufts and the secondary forest vegetation. No additional behavioral data was observed (courtship, mating and oviposition), and the host plants and immature stages are unknown.

Phylogenetic relationships. Based on the Maximum Likelihood analysis using three genes (COI, GAPDH and RpS5), Splendeuptychia is paraphyletic, with Splendeuptychia tupinamba sp. nov. appearing within a clade including Splendeuptychia ashna , and several species placed in the recently described genus Nubila Viloria, Andrade & Henao, 2019 (Andrade et al., 2019), including Nubila moderata and N. purusana , with S. ashna as sister to a clade containing the three above taxa ( Fig. 4 , Table 4 ).

Figures

FIGURE 1. Splendeuptychia tupinamba sp. nov. A, B. Holotype male (dorsal, ventral view) and C, D. Allotype female (dorsal, ventral view).

FIGURE 2. Male genitalia of Splendeuptychia tupinamba sp. nov. A. Lateral view; B. Dorsal view; C. Ventral view. Codes: ae, aedeagus; sa, saccus; te, tegumen; un, uncus; va, valva.

FIGURE 3. Males of Splendeuptychia tupinamba sp. nov. showing variation in underside wing pattern. A. Divinópolis, Minas Gerais; B. Paraopeba, Minas Gerais; C. Alto Paraíso, Paraná; D. Alta Floresta, Mato Grosso.

FIGURE 4. Maximum likelihood tree showing the phylogenetic relationship of Splendeuptychia tupinamba sp. nov. with other species of Euptychiina. Numbers near the nodes are bootstrap values (<75 are not shown).

FIGURE 5. Map showing all known localities for Splendeuptychia tupinamba sp. nov. in Brazil, Paraguay and Argentina.

FIGURE 6. Orthoimages of general habitat of Splendeuptychia tupinamba sp. nov. A. Cristalino Lodge, Alta Floresta, Mato Grosso, Brazil; B. Private Reserve Yacutinga, General Manuel Belgrano, Misiones, Argentina; C. Imperatriz, Maranhão, Brazil; D. Paraopeba National Forest, Paraopeba, Minas Gerais, Brazil; E. Brasília Country Club, Brasília, Distrito Federal, Brazil; F. Divinópolis, Minas Gerais, Brazil. Orthoimages were taken from Google Earth software.

FIGURE 7. Habitat and adults of Splendeuptychia tupinamba sp. nov. A. General view of the forest at Cristalino Lodge, Alta Floresta, Mato Grosso; B. A forest fragment at Divinópolis, Minas Gerais; C. Male perched on a leaf; D. Male with open wings perched on a leaf.