'Species' from two different butterfly genera combined into one: description of a new genus of Euptychiina (Nymphalidae: Satyrinae) with unusually variable wing pattern

André Victor Lucci Freitas Eduardo Proença Barbosa Keith Richard Willmott Niklas Wahlberg Gerardo Lamas About the authors

ABSTRACT

Sepona Freitas and Barbosa, gen. nov. is proposed for the Neotropical satyrine butterfly species Euptychia punctataWeymer, 1911Weymer, G., 1911. 4. Familie: Satyridae. In: Seitz, A. (Ed.), Die Gross-Schmetterlinge der Erde 5. A. Kernen, Stuttgart, pp. 1–37. and its junior subjective synonyms Euptychia griseolaWeymer, 1911Weymer, G., 1911. 4. Familie: Satyridae. In: Seitz, A. (Ed.), Die Gross-Schmetterlinge der Erde 5. A. Kernen, Stuttgart, pp. 1–37. and Taygetis indecisa Ribeiro, 1931. The new genus has a distinctive wing pattern and shape of the valvae in the male genitalia, the latter being a unique autapomorphy within the subtribe Euptychiina. Based on molecular data, this genus is not sister to any other single euptychiine genus, instead appearing as the sister to all remaining genera in the Taygetis clade. The present paper illustrates the complexity of the taxonomy of Euptychiina, and the importance of using different sources of evidence in taxonomic studies.

Keywords:
Harjesia; Integrative taxonomy; Pseudodebis; Satyrini; Yphthimoides

Introduction

In recent years, the highly diverse butterfly subfamily Satyrinae has been subject to several studies attempting to clarify its internal relationships and taxonomy (Murray and Prowell, 2005Murray, D.L., Prowell, D.P., 2005. Molecular phylogenetics and evolutionary history of the neotropical satyrine subtribe Euptychiina (Nymphalidae: Satyrinae). Mol. Phylogenet. Evol. 34, 67-80.; Peña et al., 2006Peña, C., Wahlberg, N., Weingartner, E., Kodandaramaiah, U., Nylin, S., Freitas, A.V.L., Brower, A.V.Z., 2006. Higher level phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data. Mol. Phylogenet. Evol. 40, 29-49., 2010Peña, C., Nylin, S., Freitas, A.V.L., Wahlberg, N., 2010. Biogeographic history of the butterfly subtribe Euptychiina (Lepidoptera, Nymphalidae, Satyrinae). Zool. Scripta 39, 243-258.; Marín et al., 2011Marín, M.A., Peña, C., Freitas, A.V.L., Wahlberg, N., Uribe, S.I., 2011. From the phylogeny of the satyrinae butterflies to the systematics of Euptychiina (Lepidoptera: Nymphalidae): history, progress and prospects. Neotrop. Entomol. 40, 1-13.; Matos-Maravi et al., 2013Matos-Maravi, P.F., Peña, C., Willmott, K.R., Freitas, A.V.L., Wahlberg, N., 2013. Systematics and evolutionary history of butterflies in the "Taygetis clade" (Nymphalidae: Satyrinae: Euptychiina): towards a better understanding of Neotropical biogeography. Mol. Phylogenet. Evol. 66, 54-68.; Siewert et al., 2013Siewert, R.R., Zacca, T., Dias, F.M.S., Freitas, A.V.L., Mielke, O.H.H., Casagrande, M.M., 2013. The "Taygetis ypthima species group" (Lepidoptera, Nymphalidae, Satyrinae): taxonomy, variation and description of a new species. Zookeys 356, 11-29.; Seraphim et al., 2014Seraphim, N., Marín, M.A., Freitas, A.V.L., Silva-Brandão, K.L., 2014. Morphological and molecular marker contributions to disentangling the cryptic Hermeuptychia hermes species complex (Nymphalidae: Satyrinae: Euptychiina). Mol. Ecol. Resour. 14, 39-49.). These studies have revealed many non-monophyletic genera, and a number of complexes of cryptic species waiting to be disentangled, especially in the predominantly lowland, largely Neotropical subtribe Euptychiina (e.g. Peña et al., 2010Peña, C., Nylin, S., Freitas, A.V.L., Wahlberg, N., 2010. Biogeographic history of the butterfly subtribe Euptychiina (Lepidoptera, Nymphalidae, Satyrinae). Zool. Scripta 39, 243-258.; Freitas et al., 2012aFreitas, A.V.L., Wahlberg, N., Matos-Maravi, P.F., Marin, M.A., Mielke, O.H.H., 2012a. Euptychia boulleti (Le Cerf) n. comb. (Lepidoptera: Nymphalidae: Satyrinae), a Rare and Endangered Butterfly from Southeastern Brazil. Neotrop. Entomol. 41, 461-467.,bFreitas, A.V.L., Kaminski, L.A., Mielke, O.H.H., Barbosa, E.P., Silva-Brandão, K.L., 2012b. A new species of Yphthimoides (Lepidoptera: Nymphalidae: Satyrinae) from the southern Atlantic forest region. Zootaxa 3526, 31-44.; Matos-Maravi et al., 2013Matos-Maravi, P.F., Peña, C., Willmott, K.R., Freitas, A.V.L., Wahlberg, N., 2013. Systematics and evolutionary history of butterflies in the "Taygetis clade" (Nymphalidae: Satyrinae: Euptychiina): towards a better understanding of Neotropical biogeography. Mol. Phylogenet. Evol. 66, 54-68.; Zacca et al., 2013Zacca, T., Mielke, O.H.H., Pyrcz, T.W., Casagrande, M.M., Freitas, A.V.L., Boyer, P., 2013. Stegosatyrus, a new genus of Euptychiina from the grasslands of neotropical realm (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa 3682, 331-350.; Siewert et al., 2013Siewert, R.R., Zacca, T., Dias, F.M.S., Freitas, A.V.L., Mielke, O.H.H., Casagrande, M.M., 2013. The "Taygetis ypthima species group" (Lepidoptera, Nymphalidae, Satyrinae): taxonomy, variation and description of a new species. Zookeys 356, 11-29.; Seraphim et al., 2014Seraphim, N., Marín, M.A., Freitas, A.V.L., Silva-Brandão, K.L., 2014. Morphological and molecular marker contributions to disentangling the cryptic Hermeuptychia hermes species complex (Nymphalidae: Satyrinae: Euptychiina). Mol. Ecol. Resour. 14, 39-49.).

Including 10 genera, the "Taygetis clade" is one of five major groups of Euptychiina (Peña et al., 2010Peña, C., Nylin, S., Freitas, A.V.L., Wahlberg, N., 2010. Biogeographic history of the butterfly subtribe Euptychiina (Lepidoptera, Nymphalidae, Satyrinae). Zool. Scripta 39, 243-258.); a preliminary phylogeny for this clade (Matos-Maravi et al., 2013Matos-Maravi, P.F., Peña, C., Willmott, K.R., Freitas, A.V.L., Wahlberg, N., 2013. Systematics and evolutionary history of butterflies in the "Taygetis clade" (Nymphalidae: Satyrinae: Euptychiina): towards a better understanding of Neotropical biogeography. Mol. Phylogenet. Evol. 66, 54-68.) showed that four genera, namely Harjesia Forster, 1964, Pseudodebis Forster, 1964, Forsterinaria R. Gray, 1973 and Taygetis Hübner [1819], are polyphyletic, requiring some revised generic combinations and the description of new genera. The genus Harjesia, as then conceived, included species placed in three different clades within the "Taygetis clade" (Matos-Maravi et al., 2013Matos-Maravi, P.F., Peña, C., Willmott, K.R., Freitas, A.V.L., Wahlberg, N., 2013. Systematics and evolutionary history of butterflies in the "Taygetis clade" (Nymphalidae: Satyrinae: Euptychiina): towards a better understanding of Neotropical biogeography. Mol. Phylogenet. Evol. 66, 54-68.). In that phylogeny, Harjesia griseola (Weymer, 1911Weymer, G., 1911. 4. Familie: Satyridae. In: Seitz, A. (Ed.), Die Gross-Schmetterlinge der Erde 5. A. Kernen, Stuttgart, pp. 1–37.) appeared as sister to the entire "Taygetis clade" (Matos-Maravi et al., 2013Matos-Maravi, P.F., Peña, C., Willmott, K.R., Freitas, A.V.L., Wahlberg, N., 2013. Systematics and evolutionary history of butterflies in the "Taygetis clade" (Nymphalidae: Satyrinae: Euptychiina): towards a better understanding of Neotropical biogeography. Mol. Phylogenet. Evol. 66, 54-68.), suggesting that it should be placed in a new genus.

Ongoing research into the phylogenetic relationships of another euptychiine genus, Yphthimoides Forster, 1964, showed that this genus is clearly polyphyletic, with several species that should be reassigned to other genera (Freitas et al., 2012bFreitas, A.V.L., Kaminski, L.A., Mielke, O.H.H., Barbosa, E.P., Silva-Brandão, K.L., 2012b. A new species of Yphthimoides (Lepidoptera: Nymphalidae: Satyrinae) from the southern Atlantic forest region. Zootaxa 3526, 31-44., Barbosa et al., 2015Barbosa, E.P., Silva, A.K., Paluch, M., Azeredo-Espin, A.M., Freitas, A.V.L., 2015. Uncovering the hidden diversity of the Neotropical butterfly genus Yphthimoides Forster (Nymphalidae: Satyrinae): description of three new species based on morphological and molecular data. Org. Divers. Evol. 15, 577-589., EPB and AVLF, in prep.). One species in particular, Yphthimoides punctata (Weymer, 1911Weymer, G., 1911. 4. Familie: Satyridae. In: Seitz, A. (Ed.), Die Gross-Schmetterlinge der Erde 5. A. Kernen, Stuttgart, pp. 1–37.), is quite distinct from all other described Yphthimoides, and further morphological studies revealed that Y. punctata and H. griseola are similar enough to be considered subjective synonyms.

This study presents evidence based on an integrative taxonomic approach (e.g., Dayrat, 2005Dayrat, B., 2005. Towards integrative taxonomy. Biol. J. Linn. Soc. 85, 407-415.; Yeates et al., 2011Yeates, D.K., Seago, A., Nelson, L., Cameron, S.L., Joseph, L., Trueman, J.W.H., 2011. Integrative taxonomy, or iterative taxonomy? Syst. Entomol. 36, 209-217.; Pante et al., 2015Pante, E., Schoelinck, C., Puillandre, N., 2015. From integrative taxonomy to species description: one step beyond. Syst. Biol. 64, 152-160.) using both morphological and molecular data for the synonymy of Y. punctata and H. griseola, and describes a new genus to harbor the resulting single species.

Material and methods

Adult specimens were studied in a number of American and European collections, and the following acronyms are used here: AWLW – Allan & Lesley Wolhuter collection, United Kingdom; DZUP – Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil; FLMNH – Florida Museum of Natural History, Gainesville, FL, USA; KWJH – Keith R. Willmott & Jason P. W. Hall collection, Gainesville, FL, USA; LBCB: L. & C. Brévignon collection, French Guiana; MNHN – Muséum National d'Histoire Naturelle, Paris, France; MNRJ – Museu Nacional do Rio de Janeiro, Rio de Janeiro, Brazil; MOBE: Mohamed Benmesbah collection, France; MUSM – Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru; MZUJ – Muzeum Zoologiczne Uniwersytetu Jagiellónskiego, Kraków, Poland; NHMUK – The Natural History Museum, London, United Kingdom; YUGA – Yuvinka Gareca colletion, Santa Cruz, Bolivia; ZSM – Zoologische Staatssammlung München, München, Germany; ZUEC – Museu de Zoologia da Universidade Estadual de Campinas, Campinas, São Paulo, Brazil; ZUEC-AVLF – André V. L. Freitas Collection, Universidade Estadual de Campinas, Campinas, São Paulo, Brazil.

Morphology

Dissections were made using standard techniques. Legs, palpi and abdomens were soaked in hot 10% potassium hydroxide for nearly 10 minutes before dissection, and dissected parts were stored in glycerol. In order to see the venation, wings were diaphanized by soaking them in alcohol and NaClO solution (bleach). Taxonomic nomenclature follows Lamas (2004aLamas, G., 2004a. Checklist: Part 4A. Hesperioidea-Papilionoidea. In: Heppner, J.B. (Ed.), Atlas of the Neotropical Lepidoptera, vol. 5A. Association for Tropical Lepidoptera/Scientific Publishers, Gainesville.,b)Lamas, G., 2004b. Euptychina. In: Lamas, G. (Ed.), Checklist: Part 4A. Hesperioidea – Papilionoidea. Heppner, J.B. (Ed.), Atlas of the Neotropical Lepidoptera, vol. 5A. Association for Tropical Lepidoptera/Scientific Publishers, Gainesville, pp. 217–223., modified by Peña et al. (2006)Peña, C., Wahlberg, N., Weingartner, E., Kodandaramaiah, U., Nylin, S., Freitas, A.V.L., Brower, A.V.Z., 2006. Higher level phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data. Mol. Phylogenet. Evol. 40, 29-49. and Wahlberg et al. (2009)Wahlberg, N., Leneveu, J., Kodandaramaiah, U., Peña, C., Nylin, S., Freitas, A.V.L., Brower, A.V.Z., 2009. Nymphalid butterflies diversify following near demise at the Cretaceous/Tertiary boundary. Proc. R. Soc. Lond. B: Biol. Sci. 276, 4295-4302.. Drawings and measurements of wings, legs and palpi were made using a Leica® MZ7.5 stereomicroscope equipped with a micrometric scale and a drawing tube. Photographs of the male and female genitalia were taken using a Zeiss Discovery V20 Stereomicroscope. The following abbreviations are used: (FW) forewing, (HW) hind wing, (D) dorsal, (V) ventral.

Phylogenetic inference

Genomic DNA was extracted from two legs of adults by using the DNeasy Blood & Tissue Kit protocol (QIAGEN, Düsseldorf, Germany). DNA was stored in TE buffer at -20 °C. The mitochondrial gene cytochrome c oxidase I (Cox1, ca. 658 bp, corresponding to the 'DNA barcode' region) for all specimens and the nuclear genes GAPDH for one specimen (YPH-0240) and RpS5 for the outgroups were amplified, purified and sequenced using standard techniques (see Silva-Brandão et al., 2005Silva-Brandão, K.L., Freitas, A.V.L., Brower, A.V.Z., Solferini, V.N., 2005. Phylogenetic relationships of the New World Troidini swallowtails (Lepidoptera: Papilionidae) based on COI, COII, and EF-1 alpha genes. Mol. Phylogenet. Evol. 36, 468-483.; Wahlberg and Wheat, 2008Wahlberg, N., Wheat, C.W., 2008. Genomic outposts serve the phylogenomic pioneers: designing novel nuclear markers for genomic DNA extractions of Lepidoptera. Syst. Biol. 57, 231-242.). The sequences of nuclear gene RpS5 for Harjesia griseola were obtained from GenBank.

All the sequences were aligned by eye with sequences obtained previously and available on GenBank by using BioEdit v. 7.2.4 (Hall, 2013Hall, T., 2013. Biological Sequence Alignment Editor Software. Ibis Biosciences, Carlsbad, CA, pp. 92008., available at http://www.mbio.ncsu.edu/bioedit/bioedit.html#downloads). The final matrix comprised 32 specimens from species of 10 genera (including 11 specimens from the new genus Sepona) and three species used as outgroups, namely Hermeuptychia maimoune (A. Butler, 1870), Paryphthimoides grimon (Godart [1824]) and Splendeuptychia doxes (Godart [1824]) (see Table 1 for the sequence codes).

Table 1
Species of Euptychiina with code, sampling site data, and GenBank accession numbers for sequenced genes.

The phylogenetic relationships of the new species were estimated using maximum likelihood. Analyses were run using RAXML (Stamatakis et al., 2008Stamatakis, A., Hoover, P., Rougemont, J., 2008. A rapid bootstrap algorithm for the RAxML web-servers. Syst. Biol. 75, 758-771.) with 1000 rapid bootstrap replicates and a search for the maximum likelihood topology on the CIPRES portal (Miller et al., 2010Miller, M.A., Pfeiffer, W., Schwartz, T., 2010. Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: Proceedings of the Gateway Computing Environments Workshop (GCE), 14 Nov. 2010, New Orleans, LA, pp. 1–8.). The data were modeled according to the GTR + G model for each partition independently.

Sepona Freitas & Barbosa, gen. nov.

Type species: Euptychia punctataWeymer, 1911Weymer, G., 1911. 4. Familie: Satyridae. In: Seitz, A. (Ed.), Die Gross-Schmetterlinge der Erde 5. A. Kernen, Stuttgart, pp. 1–37., here designated.

Diagnosis

Molecular data (Peña et al., 2010Peña, C., Nylin, S., Freitas, A.V.L., Wahlberg, N., 2010. Biogeographic history of the butterfly subtribe Euptychiina (Lepidoptera, Nymphalidae, Satyrinae). Zool. Scripta 39, 243-258.) place this genus within the "Taygetis clade" of the satyrine subtribe Euptychiina. In terms of wing shape and pattern, adults are similar to those of some species of Harjesia, Pseudodebis and Taygetina Forster, 1964, which all have undulate rather than straight dark discal and postdiscal lines on the ventral surface, but this genus can be distinguished from all other euptychiines by the unique shape of the valvae in the male genitalia (Fig. 3A). The valvae bear a long, thin, inwardly curved projection arising abruptly from the otherwise rounded main body of the valva. See Table 2 for comparisons of additional morphological characters of Sepona punctata with representatives of other genera of the "Taygetis clade". The relationships of Sepona to other euptychiina genera, and justification for its recognition as a monotypic genus, are addressed further below under 'Discussion'.

Fig. 1
Adult male of Sepona punctata – Jaru, Rondônia, Brazil. Dorsal above, ventral below.
Fig. 2
Morphological characters of Sepona punctata. A, male wing venation – forewing above and hind wing below; B, female wing venation – forewing above and hind wing below; C, male palpus; D, male foreleg; E, male midleg; F female foreleg.
Fig. 3
Male and female genitalia of Sepona punctata. A, male genitalia in lateral view; B, male genitalia in dorsal view; C, male genitalia in ventral view; D, male aedeagus (lateral view); E, female genitalia in ventral view; F. female genitalia: detail of the signa in corpus bursae. (sa) saccus, (te) tegumen, (un) uncus, (va) valva, (bu) corpus bursae, (st) sterigma.

Table 2
Comparisons of Sepona punctata with exemplar species from related genera in "Taygetis clade". For aedeagus and saccus width, the value refers to length/width ratio in the medial portion of each structure; higher numbers therefore represent a narrower structure.

Etymology

Sepona is an arbitrary combination of letters, derived from the Latin transitive verb "sepono", meaning to put aside, separate, or remove, in reference to the isolated position of the genus in comparison with other members of the "Taygetis clade". It should be treated as a feminine noun.

Sepona punctata (Weymer, 1911Weymer, G., 1911. 4. Familie: Satyridae. In: Seitz, A. (Ed.), Die Gross-Schmetterlinge der Erde 5. A. Kernen, Stuttgart, pp. 1–37.) comb. nov.

Euptychia punctata Weymer, 20 April 1911: 205. Type Locality: Brazil, Minas Gerais. Syntype(s), not located.

=Euptychia griseola Weymer, 20 July 1911: 211, n. syn. Type Locality: Bolivia [La Paz], Mapiri. Lectotype female (here designated): 'Original! / nun aber heissen griseola // Mapiri // Collection / v.Rosen // grisea / Weym. // Typus Nr. / Euptychia griseola Stgr. i. l. / Weymer / Zoologische / Staatssammlung / München', deposited in ZSM (examined).

=Taygetis indecisa Ribeiro, 1931: 33, n. syn. Type Locality: Brazil [Rondônia, Rio Jamari]. Holotype female: 'Comm.Rondon / No. 43. 19. 6. 14 / Coll. Stolle // HOLOTIPO // N 053/645 // Holótipo Taygetis / indecisa/Ribeiro, 1931 / Mielke & Casa-grande det. 1985', deposited in MNRJ (examined).

Taygetis indecisa: May, 1933May, E.G.L., 1933. Notas sobre a collecção de lepidopteros do Estado de Matto Grosso feita pela commissão Rondon. Boletim do Museu nacional do Rio de Janeiro 9, 119-126.: 120; Euptychia griseola: May, 1933May, E.G.L., 1933. Notas sobre a collecção de lepidopteros do Estado de Matto Grosso feita pela commissão Rondon. Boletim do Museu nacional do Rio de Janeiro 9, 119-126.: 120; D'Abrera, 1988D'Abrera, B., 1988. Butterflies of the Neotropical region. Part V. Nymphalidae (Conclusion) & Satyridae. Hill House, Victoria, pp. 679–877.: 789; Pseudodebis griseola: Forster, 1964Forster, W., 1964. Beiträge zur Kenntnis der Insektenfauna Boliviens XIX. Lepidoptera III. Satyridae. Veröffentlichungen der zoologischen Staatssammlung München 8, 51-188.: 77; Lamas, 1983Lamas, G., 1983. Adiciones y correcciones a la lista de mariposas de la Reserva de Tambopata, Perú. Revista de la Sociedad mexicana de Lepidopterología 8, 13-24.: 16; Robbins et al., 1996Robbins, R.K., Lamas, G., Mielke, O.H.H., Harvey, D.J., Casagrande, M.M., 1996. Taxonomic composition and ecological structure of the species rich butterfly community at Pakitza, Parque Nacional del Manu, Peru. In: Wilson, D.E., Sandoval, A. (Eds.), Manu: The Biodiversity of Southeastern Peru. Smithsonian Institution Press, Washington, DC.: 230; Freitas, 2003Freitas, A.V.L., 2003. Description of a new genus for "Euptychia" peculiaris Nymphalidae: Satyrinae). J. Lepid. Soc. 57, 100-106.: 103; Euptychia punctata: D'Abrera, 1988D'Abrera, B., 1988. Butterflies of the Neotropical region. Part V. Nymphalidae (Conclusion) & Satyridae. Hill House, Victoria, pp. 679–877.: 789; Taygetis griseola: D'Abrera, 1988D'Abrera, B., 1988. Butterflies of the Neotropical region. Part V. Nymphalidae (Conclusion) & Satyridae. Hill House, Victoria, pp. 679–877.: 756; Harjesia griseola: Lamas, 2004: 219; Brévignon, 2008Brévignon, C., 2008. Inventaire des Satyrinae de Guyane Française (Lepidoptera: Nymphalidae). In: Lacomme, D., Manil, L. (Eds.), Lépidoptères de Guyane. Tome 3. Rhopalocères 2. Association des Lépidoptéristes de France, Paris, pp. 62–94.: 68; Brévignon and Benmesbah, 2012Brévignon, C., Benmesbah, M., 2012. Complément à l’inventaire des Satyrinae de Guyane (Lepidoptera: Nymphalidae). In: Lacomme, D., Manil, L. (Eds.), Lépidoptères de Guyane. Tome 7. Nymphalidae. Association des Lépidoptéristes de France, Paris, pp. 36–52.: 39; Matos-Maravi et al., 2013Matos-Maravi, P.F., Peña, C., Willmott, K.R., Freitas, A.V.L., Wahlberg, N., 2013. Systematics and evolutionary history of butterflies in the "Taygetis clade" (Nymphalidae: Satyrinae: Euptychiina): towards a better understanding of Neotropical biogeography. Mol. Phylogenet. Evol. 66, 54-68.: 54; Yphthimoides punctata: Lamas, 2004: 223.

Redescription

Male (Figs. 1, 2A-C-D-E, 4A-B-D). Eyes reddish brown, covered with sparse black hairs. Palpus 1.5 times as long as head, brown with light brown hairs (Fig. 2C). Antenna of males 9.0–10.0 mm in length with 36 antennomeres, extending to mid-costa; shaft rust-brown dorsally, orange brown ventrally, sparsely scaled dorsally; club not conspicuously developed, including eleven segments, with apical portion (last five segments) dark brown. Forewing length 23–25 mm (n = 6); hindwing length 19–20 mm (n = 6). HW outer margin slightly undulate. Male wing venation shown in Fig. 2A. Wings with dorsal ground color dark brown with few markings, restricted to a suffused dark brown outer margin on DFW, and to dark double marginal line, and a submarginal line following contours of marginal line on DHW. Ventral wings light brown; VFW crossed by two thin zigzag dark brown lines, extending from costa to 2A, first line one-third distance from wing base to apex; second line extending from costa to 2A at two-thirds distance from wing base to apex; a conspicuous lighter outer band is adjacent to second line, followed by a darker ocellar region (see next); a thin dark brown zigzag submarginal line with single black dots in vertices and a brown regular marginal line extending from costa to 2A; four dark ocelli in spaces R5–M1 (ocellus 1), M1–M2 (2), M2–M3 (3) and M3–CuA1 (4). VHW crossed by two thin dark brown lines from costa to anal margin, in similar position to those on forewing; a conspicuous lighter outer band is adjacent to second line, followed by a darker ocellar region (see next); a dark brown zigzag submarginal line with single black dots in vertices and a brown regular marginal line extending from costa to 2A; a series of five dark ocelli can be found in cells Rs–M1 (ocellus 1), M1–M2 (2), M2–M3 (3), M3–CuA1 (4) and CuA1–CuA2 (5). Details about ocelli size and shape discussed further below. No conspicuous androconial scales observed.

Fig. 4
Variation in wing pattern of Sepona punctata (all from Brazil). A, Abunã, Rio Madeira, Rondônia; B, Porto Velho, Rondônia; C, Estação Ecológica do Alto Acre, Acre; D, Porto Acre, PAD Humaitá, Acre; E, Conceição do Mato dentro, Minas Gerais; F, Parque Municipal do Trabijú, Pindamonhangaba, São Paulo (A, B, D – males; C, E, F – females).

Male genitalia (Fig. 3AE). Saccus elongate; tegumen rounded and short; gnathos long and pointed, projecting upwards above uncus; uncus elongated, with lateral expansions in dorsal view, giving an arrowhead appearance; valvae elongated, ending in a bump with a long thin pointed process; aedeagus curved; cornuti absent; juxta sclerotized, linking both valvae together.

Female (Fig. 2B, F, 4C-E-F). Forewing length 24–26 mm (n = 2); hindwing length 20–22 mm (n = 2). Antenna 11.0 mm in length, with 35 antennomeres, extending to mid-costa. General color and pattern very similar to those of males. Female genitalia as in Fig. 3E, F. Ductus bursae partially sclerotized, corpus bursae rounded; a pair of conspicuous signa present.

Taxonomy and variation

Weymer (1911)Weymer, G., 1911. 4. Familie: Satyridae. In: Seitz, A. (Ed.), Die Gross-Schmetterlinge der Erde 5. A. Kernen, Stuttgart, pp. 1–37. described Euptychia punctata based on an unstated number of specimens from Minas Gerais, Brazil. Several pages later, he described Euptychia griseola based also on an unstated number of specimens from Mapiri, Bolivia. Later, Ribeiro (1931)Ribeiro, V. de M., Lepidópteros de Matto Grosso, 1931. Material colligido pelos senhores General Cândido Rondón, Prof. Alípio de Miranda-Ribeiro e Emil Stolle. Boletim do Museu nacional do Rio de Janeiro 7, 31-52. described a third taxon as Taygetis indecisa, based on one female from Brazil, Rio Jamari; this taxon was promptly synonymized with Euptychia griseola by May (1933)May, E.G.L., 1933. Notas sobre a collecção de lepidopteros do Estado de Matto Grosso feita pela commissão Rondon. Boletim do Museu nacional do Rio de Janeiro 9, 119-126.. Descriptions of both E. punctata and E. griseola were also based on female specimens, and although this cannot be determined unambiguously from their original descriptions, the fact that the types are females in both cases suggests that males were unknown to the authors. No type specimen(s) of punctata has been found, but there is a single female in ZSM identified by Forster (1964)Forster, W., 1964. Beiträge zur Kenntnis der Insektenfauna Boliviens XIX. Lepidoptera III. Satyridae. Veröffentlichungen der zoologischen Staatssammlung München 8, 51-188. as the "Typus" of griseola, and we accept that it indeed represents a syntype (which we designate herein as lectotype), since this particular specimen (examined) matches precisely the illustration provided subsequently by Weymer (1911: pl. 47g, fig. [7])Weymer, G., 1911. 4. Familie: Satyridae. In: Seitz, A. (Ed.), Die Gross-Schmetterlinge der Erde 5. A. Kernen, Stuttgart, pp. 1–37.. The female holotype (examined) of Taygetis indecisa Ribeiro is deposited in MNRJ. Although appearing rather different in wing pattern, the names punctata and griseola apparently represent extremes of geographical variation within a single species. Variation on the dorsal wing surfaces is practically absent and obvious seasonal variations have not been detected. The ventral surface of both wings, however, shows much variation, especially in the number and size of the ocelli. Most individuals from central and southeastern Brazil and eastern Bolivia ("punctata" phenotype) have the ocelli reduced to small black dots, sometimes with a tiny white pupil on the VHW; they also present a more homogeneous ventral pattern (Fig. 4E, F). Conversely, individuals from western Amazonia and Guianas (the "griseola" phenotype) have more developed ocelli circled by yellowish cream scales and with a white pupil on both wings, and a conspicuous banded pattern on the ventral wings (Fig. 4A, B). Intermediate phenotypes between "punctata" and "griseola" are known from Acre and Rondônia in Brazil, and from Bolivia, and are usually more similar to the "griseola" phenotype (Fig. 4C, D). To our knowledge, no two of these three phenotypes ("punctata", "griseola" and intermediate) have been recorded in sympatry. The two names were published several months apart, and we thus treat E. griseola as a junior subjective synonym of S. punctata n. syn.

Specimens examined (34 ♂, 27 ♀): Bolivia. – Beni: Prov. Mamoré, San Ramón, Estancia S/Lorenzo [13°26'9.26"S, 64°36'2.79"W], 13 Oct 2003, T. V. Bosque de Galeria, T17, Yuvinka Gareca leg. (YUGA); La Paz: 'Madidi' [14°3'55"S,68°50'49"W] (Aliaga, W.), 4 Oct 2005, 1 ♀ (MUSM), 5 Oct 2005, 1 ♂ (MUSM); Mapiri [15°19'27"S,68°6'50"W], 1 ♂ [LT griseola; "Original! / nun aber heissen griseola // Mapiri // Collection / v.Rosen // grisea / Weym. // Typus Nr. / Euptychia griseola Stgr. i. l. / Weymer / Zoologische / Staatssammlung / München"] (ZSM); Santa Cruz: Buenavista [17°27'S,64°40'W], 750 m (Steinbach, J.), Aug 1907–Apr 1908, 1 ♂ [BMNH-E-1204766] (NHMUK). Brazil. – Acre: 50 km NW Bujari [9°32'53"S,68°18'9"W], 200 m (Mielke, O., M. Casagrande), 25 Sep 2003, 1 ♀ (DZUP); Assis Brasil, Estação Ecológica do Alto Acre [11°3'S,70°16'W], 300 m (Brown, K. S.), 19 Jul 2005, 1 ♂ [DNA voucher YPH-0346] (ZUEC-AVLF), 21 Aug 2005, 1 ♀ (ZUEC-AVLF), Aug 2005, 1 ♂ [DNA voucher YPH-0502] (ZUEC-AVLF); Marechal Thaumaturgo, Reserva Extrativista do Alto Juruá, Boca do Caipora [9°9'16"S,72°40'21"W] (Brown, K. S. & A. V. L. Freitas), 1 Sep 1997, 1 ♂ [ZUEC LEP-9231], 1 ♀ [ZUEC LEP-9232] (ZUEC); Marechal Thaumaturgo, upper Juruá River, Foz do [Rio] Tejo [8°59'1"S,72°43'W] (Freitas, A.V.L.), 20–27 Aug 1997, 1 ♀ (ZUEC-AVLF); PAD Humaitá, Porto Acre [9°35'29"S,67°32'20"W], 200 m (Mielke, O., M. Casagrande), 8 Oct 2004, 1 ♂ (DZUP); Espírito Santo: Colatina [19°31'55"S,40°40'42"W], Aug–Sep 1937, 3 ♂ [24/579, 52/439, 52/438], Sep 1937, 1♀ [24/689] Coll. E. May (MNRJ); Cor.[rego] do Sabiá, Est.[rada] N.[ova] Venecia [19°8'20"S,40°37'8"W], Collatina [sic], Oct 1936, 1♀, E. May leg. [52/336] (MNRJ); Mato Grosso do Sul: Bonito [21°7'34"S,56°29'25"W] (Araújo, P. F.), 15 Oct 2014, 1 ♀ [ZUEC LEP-9234; DNA voucher YPH-0507] (ZUEC), 5 Nov 2014, 1 ♂ [ZUEC LEP-9235; DNA voucher YPH-0506] (ZUEC); Mato Grosso: 11–18 km N Ribeirão Cascalheira [12°50'47"S,51°46'59"W] (Mielke, O.), 21–23 Aug 1997, 1 ♀ (DZUP); C[oron]el. Rio Branco [15°14'24"S,58°6'49"W], 400 m (Buzzi, Mielke, Elias & Casagrande), 19 Sep 1984, 1 ♂ [DZ-5465; Proj.[eto] Polonoroeste] (DZUP) (Mielke, O., K. S. Brown), 3 Jul 1972, 1 ♂ (DZUP); Diamantino, Rio Arinos, Faz.[enda] S.[ão] João [14°21'18"S,56°9'2"W], 300–400 m (Ebert, H. & H. D.), 5 May 1978, 1 ♂ (DZUP); Chapada dos Guimarães, Buriti [15°23'31"S, 55°53'11"W], 1 Jan 1969, 1 ♂ [ZUEC LEP-9246; Slide no 1762, ♂ genitália, Lee D. Miller] (ZUEC); Minas Gerais: Conceição do Mato Dentro [19°2'S,43°25'31"W] (Ramos, G.), 11 Sep 2013, 1 ♀ [ZUEC LEP-9233; DNA voucher YPH-0494] (ZUEC); no specific locality (Boullet), 1913, 2 ♂ (MNHN); Rio de Janeiro: Itabapoana [21°8'S,41°41'W], 1 ♀ [BMNH-E-1204794] (NHMUK); Rondônia: 67 km S Ariquemes, 5 km S of Cacaulândia on linha C-10 at Rio Pardo off B-65 [10°23'15"S,62°54'53"W] (Austin, G.T.), 14 Oct 1993, 1 ♂ (FLMNH), 8 Oct 1993, 1 ♀ (FLMNH) (Gomes, O.), 15 May 1994, 1 ♀ (FLMNH); 67 km S Ariquemes, 5 km S of Cacaulândia on linha C-10 [10°23'13"S,62°54'12"W], 170 m (Gomes, O.), 1 Aug 1993, 1 ♂ (FLMNH); Jaru [10°27'S,62°30'W] (Brown, K. S.), 30 Sep 1975, 2 ♀ (FLMNH), 5 Oct 1975, 1 ♀ (FLMNH) (Callaghan, C. J.), 10 Aug 1976, 1 ♂ [genitalia vial AVF 7841], 1 ♀ [genitalia vial AVF 7840] (FLMNH); Pôrto Velho [8°45'S,63°53'W], 31 Jan 2010, 1 ♂ [ZUEC LEP-9236; DNA voucher YPH-0511] (ZUEC); Rio Jamari [10°12'S,63°15'W], 1 ♂ [HT indecisa; "'Comm.Rondon / No. 43. 19. 6. 14 / Coll. Stolle // HOLOTIPO // N 053/645 // Holótipo Taygetis / indecisa /Ribeiro, 1931/ Mielke & Casa-grande det. 1985"] (MNRJ); Upper Rio Madeira, Abunã [9°42'S,65°21'W] (Nunes, R. V.), 1 May 2013, 1 ♂ [ZUEC LEP-9230; DNA voucher YPH-0240] (ZUEC); São Paulo: Pindamonhangaba, Reserva Natural Municipal do Trabijú [22°50'18"S,45°31'23"W] (Rosa, A. H. B.), 22 Jul 2014, 1 ♂ [ZUEC LEP-9237; DNA voucher YPH-0503] (ZUEC). Ecuador. – Napo: Río Jatunyacu, Pimpilala [1°4'31"S,77°56'13"W], 800 m (Jasinski, A.), 28 May 1997, 1 ♀ (MZUJ); Orellana: [c. 13 km W Coca], Río Payamino [0°26'18"S,77°6'46"W] (Wolhuter, A. & L.), 28 Oct 2009, 1 ♀ (AWLW) (Wolhuter, A. & L., pers. comm. and CD of images); Reserva Biológica Río Bigal, 8 de Diciembre [0°32'15"S,77°25'18"W], 975 m (T. & M. García), Oct 2009 (photograph live specimen); Pastaza: Puyo, Chifa Restaurant [1°29'13"S,78°0'W], 1000 m (K. Willmott & J. Hall), 25 Aug 1993, 1 ♂ (KWJH); 13 km N Puyo [1°23'26"S,77°58'35"W], 1100 m (Emmel, T. C.), 8 Sep 1969, 1 ♀ (FLMNH). French Guiana.Saint-Laurent-du-Maroni: Maroni river, 1 ♀ (FLMNH). Peru. – Cuzco: Echarate, Alto Manugali [12°27'S,73°02'W], 812 m (Valencia, G.), 5 Apr 2009, 1 ♀ (MUSM); Río Urubamba, Saringabeni [12°11'05"S,72°49'48"W], 442 m (Cerdeña, J., J. Farfán, R. Gutiérrez), 31 Aug 2007, 1 ♀ [fish bait trap] (MUSM); San Martín-3 Camp [11°47'S,72°42'W], 475 m (Valencia, G.), 11 Apr 1997, 1 ♀ (MUSM); Loreto: 'Cavallo Cocha' [=Caballococha] [3°55'S,70°31'W], 90 m (Mathan, M. de), May-Jul 1884, 1 ♂ [BMNH-E-786155], 1 ♂ [BMNH-E-786156] (NHMUK); Madre de Dios: Boca Río La Torre [12°50'S,69°17'W], 300 m (Covell, C.V.), 22 Oct 1983, 1 ♀ (MUSM) (Lamas, G.), 15 Feb 1982, 1 ♀ (MUSM); Parque Manu, Pakitza [11°56'40"S,71°16'58"W], 340 m (Mielke, O.), 1 Oct 1991, 1 ♂ (DZUP), 24 Sep 1991, 1 ♂ (DZUP), 27 Sep 1991, 1 ♂ (DZUP), 7 Oct 1991, 1 ♂ (DZUP); Parque Manu, Pakitza [11°53'S,70°58'W], 400 m (Lamas, G.), 16 Oct 1990, 1 ♂ (MUSM), 20 Oct 1990, 1 ♂ (MUSM), 7 Oct 1990, 1 ♂ (MUSM) (Rowe, W.), 3 Nov 1990, 1 ♀ (MUSM); Reserva Tambopata [12°50'S,69°17'W], 300 m (Lamas, G.), 31 Oct 1990, 1 ♀ (MUSM); Quebrada Agua Negra [12°53'S,69°17'W], 200 m (Williams, H.), 16 Sep 1995, 1 ♀ (MUSM); Río Madre de Dios, Albergue Amazonia [12°52'S,71°23'W], 500 m (Lamas, G.), 24 Oct 2013, 1 ♀ (MUSM), 28 Sep 2014, 1 ♂ (MUSM), 29 Sep 2014, 1 ♀ (MUSM); Salvación [12°50'S,71°21'W], 500 m (Matthews, M.J.), 6 Aug 1982, 1 ♂ (MUSM); Río Azul [13°3'S,69°55'W], 850 m (Peña, C.), 1 Oct 2010, 1 ♂ [voucher CP24-01] (MUSM), 23 Sep 2010, 1 ♂ (MUSM); Río Azul, Reserva Comunal Amarakaeri [12°49'S,71°6'W], 507 m (Vílchez, M.), 11 Oct 2010, 1 ♀ (MUSM).

Other records: French Guiana. – Cayenne, Roura, Cacao [4°35'N,52°28'W] (Damico, R.), Feb 1996, 1 ♀ (LBCB) (Brévignon, 2008Brévignon, C., 2008. Inventaire des Satyrinae de Guyane Française (Lepidoptera: Nymphalidae). In: Lacomme, D., Manil, L. (Eds.), Lépidoptères de Guyane. Tome 3. Rhopalocères 2. Association des Lépidoptéristes de France, Paris, pp. 62–94.: 68, pl. 7, fig. 81, 82); Saint-Laurent-du-Maroni, Saül [3°37'N,53°12'W], 9 Oct 2011, 1 ♂ (MOBE) (Brévignon & Benmesbah, 2012Brévignon, C., Benmesbah, M., 2012. Complément à l’inventaire des Satyrinae de Guyane (Lepidoptera: Nymphalidae). In: Lacomme, D., Manil, L. (Eds.), Lépidoptères de Guyane. Tome 7. Nymphalidae. Association des Lépidoptéristes de France, Paris, pp. 36–52.: 46, pl. 3, fig. 11 [adult wings], 11a [male genitalia]).

Biology and distribution

Sepona punctata is known from eastern Ecuador to southeastern Brazil (Espírito Santo, Rio de Janeiro, Minas Gerais and São Paulo). There are also records from two sites in French Guiana (Brévignon, 2008Brévignon, C., 2008. Inventaire des Satyrinae de Guyane Française (Lepidoptera: Nymphalidae). In: Lacomme, D., Manil, L. (Eds.), Lépidoptères de Guyane. Tome 3. Rhopalocères 2. Association des Lépidoptéristes de France, Paris, pp. 62–94.; Brévignon and Benmesbah, 2012Brévignon, C., Benmesbah, M., 2012. Complément à l’inventaire des Satyrinae de Guyane (Lepidoptera: Nymphalidae). In: Lacomme, D., Manil, L. (Eds.), Lépidoptères de Guyane. Tome 7. Nymphalidae. Association des Lépidoptéristes de France, Paris, pp. 36–52.) (Fig. 6). In eastern Ecuador the species is extremely rare, and the few known localities are in the Andean foothills on the types of sandstone soils that frequently support stands of bamboo. Adults are usually scarce and rare in collections, although they were sometimes common in areas with large bamboo patches in the upper Juruá River, in Acre, Brazil (AVLF and K. S. Brown Jr., pers. obs.). The species is usually associated with forested habitats, but some populations in SE Brazil (the "punctata" phenotype) are known from riparian forests in the cerrado. The immature stages and hostplants are unknown.

Fig. 5
Relationships among Sepona punctata and selected species in the "Taygetis clade" and several outgroups inferred with maximum likelihood. Numbers near branch nodes are bootstrap branch support. Names in parentheses for Sepona punctata refer to the phenotype of the voucher specimens (see text).
Fig. 6
Recorded localities of Sepona punctata (based on all examined material; see text for details). Black dots = "griseola" phenotype; white dots = "punctata" phenotype; gray dots = intermediate phenotypes.

Discussion

The position of Sepona punctata as a well-supported sister to the remaining genera in the "Taygetis clade", and the polyphyletic nature of Harjesia as illustrated by Matos-Maravi et al. (2013)Matos-Maravi, P.F., Peña, C., Willmott, K.R., Freitas, A.V.L., Wahlberg, N., 2013. Systematics and evolutionary history of butterflies in the "Taygetis clade" (Nymphalidae: Satyrinae: Euptychiina): towards a better understanding of Neotropical biogeography. Mol. Phylogenet. Evol. 66, 54-68. and in the present paper (Fig. 5), clearly shows that this species is not part of Harjesia (which has as its type species Taygetis blanda Möschler, 1877). The reasons for erecting a new genus for this taxon are therefore clear: unless all species in the "Taygetis clade" are lumped into a single genus, an undesirable option given the morphological variation and taxonomic diversity within the clade, there is no way to circumscribe monophyletic genera in the clade without making this taxon a monobasic genus. In addition to its phylogenetic position, the male genitalia of S. punctata is quite distinct from all known species of Harjesia (Forster, 1964Forster, W., 1964. Beiträge zur Kenntnis der Insektenfauna Boliviens XIX. Lepidoptera III. Satyridae. Veröffentlichungen der zoologischen Staatssammlung München 8, 51-188. and unpublished results of the authors), presenting several unique characters, including the extremely thin and curved aedeagus and the unique shape of the valvae (see Fig. 3A and Table 2).

The known wing patterns of S. punctata are highly variable, but although specimens from western Amazonia and Guianas are quite divergent from those from southeastern Brazil, individuals with intermediate wing pattern are known from eastern Bolivia, and Acre and Rondônia, Brazil. In addition, these differences are not related to seasonal forms and, based on the few known individuals, there is low variation within populations, including in the sites where intermediate populations are known. These reasons were considered sufficient to not recognize subspecific taxa within this species.

The above-described variation in wing patterns throughout the distribution of S. punctata easily explains why this taxon was described as three different species, twice by the same author (Weymer, 1911Weymer, G., 1911. 4. Familie: Satyridae. In: Seitz, A. (Ed.), Die Gross-Schmetterlinge der Erde 5. A. Kernen, Stuttgart, pp. 1–37.), in three different genera (see the synonymic list above). This situation is a perfect example of how complex is the taxonomy of Euptychiina, where most of the large genera are non-monophyletic, with species spread in two or more different clades (as is the case of Splendeuptychia Forster, 1964, Cissia Doubleday, 1848, and Paryphthimoides Forster, 1964, see Peña et al., 2010Peña, C., Nylin, S., Freitas, A.V.L., Wahlberg, N., 2010. Biogeographic history of the butterfly subtribe Euptychiina (Lepidoptera, Nymphalidae, Satyrinae). Zool. Scripta 39, 243-258.).

Hopefully, forthcoming studies combining morphological and molecular data will help to disentangle the complex and species-rich clade which constitutes the subtribe Euptychiina, providing a well resolved phylogeny that will serve as a framework for future studies focusing on diversification patterns of Neotropical butterflies.

Acknowledgements

We would like to thank Keith S. Brown Jr., Blanca Huertas, Marcio Uehara-Prado, Lee D. Miller, Jacqueline Y. Miller, Debra Murray, Jason Hall, Axel Hausmann, Carla Penz, Gláucia Marconato, Yuvinka Gareca and Andreas Segerer for their help in diverse phases of the manuscript. We also thank Poliana F. Araujo and Augusto H. B. Rosa for providing fresh specimens, Luiza M. Magaldi for helping with the DNA extraction and Tamara M. C. Aguiar for helping spread the specimens. Lee D. Miller and Jacqueline Y. Miller gave great inspiration to the first author to work on Satyrinae systematics. EPB acknowledges FAPESP (2012/03750-8) for a graduate fellowship. AVLF acknowledges support from FAPESP (Biota-Fapesp – grants 2011/50225-3, 2012/50260-6, 2013/50297-0), from the Brazilian Research Council – CNPq (fellowship 302585/2011-7). This publication is part of the RedeLep 'Rede Nacional de Pesquisa e Conservação de de Lepidópteros' SISBIOTA-Brasil/CNPq (563332/2010-7), of the project "Identificação Molecular de Biodiversidade de Invertebrados Terrestres" (grant 564954/2010-1) included in the "Rede Nacional de Identificação Molecular da Biodiversidade—BR-BoL" (MCT/CNPq/FNDCT 50/2010), the collaborative grant "Dimensions US-Biota-São Paulo: Assembly and evolution of the Amazon biota and its environment: an integrated approach", US NSF, NASA, and FAPESP (2012/50260-6), and from the National Science Foundation (DEB-1256742).

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Publication Dates

  • Publication in this collection
    Apr-Jun 2016

History

  • Received
    08 May 2015
  • Accepted
    12 Jan 2016
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