Morphological and molecular evidence supports recognition of

Cithaerias cliftoni (Constantino, 1995) as a distinct from C. pireta (Stoll, 1780) and C. aurorina (Weymer, 1910)

Luis Miguel Constantino

2016 Supplementary Material: Constantino, 2016. Revision of the tribe (: Satyirnae)

Remarks on the nominal taxon cliftoni (Constantino, 1995)

Cithaerias cliftoni Constantino, 1995 is a member of the tribe Haeterini (Lepidoptera: : ) confined to the Neotropical region. The of this tribe, are for the most part, readily distinguished from all other groups of the Satyrinae by having largely transparent wings with one or two ocelli and patches of color on the hindwing margin (Constantino 1992). C. cliftoni is a good species inhabiting the rainforests of the upper Amazon basin in the eastern slopes of the Andes of Colombia, Ecuador and North of Peru, not sympatric with C. aurora (C. Felder & R. Felder, 1862) (Penz et. al. 2014). C. cliftoni was synonymized by Lamas (2004) with C. phantoma (Fassl, 1922) from Manicoré at Rio Madeira, Tefé and São Paulo de Olivença, Amazonas, Brazil, a very distant population from Colombia, however Lamas (1998) was not able to locate the type material of C. phantoma. As a result of this Penz et. al (2014) synonymized C. phantoma with C. aurora and reinstated C. cliftoni as a full species based on morphological differences of the genitalia. Geographical distribution C. cliftoni is found in the east slope of the Andes of Colombia, Ecuador and North of Perú. In Colombia is found in the departments of Meta, Caquetá and Putumayo at an altitudinal range of 100 up to 800 m. It is sympatric with C. aurorina (Wymer, 1910) in lowland regions of the upper Amazon (Putumayo, Orteguaza and Caqueta rivers). In Ecuador is present in the provinces of Napo, Sucumbios and Pastaza and in Perú in the department of Loreto (Yurimaguas). Diagnosis Male (Figure 2 a) HW marginal band very thin; male HW submarginal band usually thin, clearly separated from marginal band and slightly staggered; male HW postmedial band that outlines the ocellus complete, reaching vein M3, or incomplete, not reaching M3; male HW postmedial band usually thin, and usually complete across cells M3 through Cu2, forming a staggered pattern; male distance between HW submarginal and postmedial bands usually similar to the width of the cells, but variable between cells; male HW rose scaling more diffuse than C. aurora, less than C. pireta, usually entering discal cell. Female (Figure 2b) with much wider HW brown bands than male, forming arches in each cell; female HW scaling usually limited to postmedial area but sometimes reaching the

1 Supplementary Material: Constantino, 2016. Revision of the tribe Haeterini (Lepidoptera: Satyirnae) medial area, varying in color from white to rose. In both sexes, FW brown bands vary from incomplete (below discal cell only) to nearly absent. The female of C. aurorina can be confused with the female of C. cliftoni. In Penz et. al. 2014) figures 3e and 3h correspond to the female of C. aurorina.. Male genitalia (Figure 1g): in lateral view the tall valva is narrow, and in ventral view it lacks an inner projection; in dorsal view the lateral edges of uncus plus tegument are rounded; in ventral view the triangular shape of the weakly sclerotized subscaphium bears small spines—a key difference between C. cliftoni and C. aurora (Penz et. al. 2014).

Taxonomic position of Cithaerias cliftoni in the tribe Haeterini

The rose colored Cithaerias species were recently revised by Penz, Alexander and DeVries (2014) due to the confusion generated in the checklist of Lamas (2004). Lamas (2004) placed incorrectly most of the rose colored Cithaerias with C. pireta (Stoll, 1780), despite that Constantino (1995) showed that C. pireta (previously known as C. menander) was morphologically different from C. aurorina (Figure 1g), observation corroborated by Penz et. al. 2014).

Based on the revision of Constantino 1994 and Penz et. al. 2014 the tribe Haeterini is composed by the following genera and species:

DULCEDO d´Almeida, 1951. D. polita (Hewitson, 1869). Central America to W. Colombia and W. Ecuador. PARADULCEDO Constantino, 1992. P. mimica (Rosenberg & Talbot, 1914). W. Colombia. = Callitaera mimica Rosenberg & Talbot, 1914. = Cithaerias gilmouri Okano, 1986. PSEUDOHAETERA F.M. Brown, 1943. P. hypaesia (Hewitson, 1854). Andes of , Colombia to Bolivia. Fabricius, 1807. H. macleannania H. W. Bates, 1865. Central America to W. Colombia and W. Ecuador.

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H. piera (Linnaeus, 1775). Amazon basin. a) H. piera piera (Linnaeus, 1775). Brasil (Amazon basin). b) H. piera diaphana Lucas, 1857. Brasil (Bahia). c) H. piera negra C. Felder & R. Felder, 1862. Peru. d) H. piera pakitza Lamas, 1998. Peru. e) H. piera unocellata Weymer, 1910. Bolivia. f) H. piera sanguinolenta Constantino & Salazar, 2007. Colombia.

CITHAERIAS Hubner, 1819 =Callitaera Butler, 1868. C. andromeda (Fabricius, 1775). a) C. andromeda andromeda (Fabricius, 1775). Surinam, Brazil. b) C. andromeda azurina (J. Zikan, 1942). Colombia. C. bandusia Staudinger, 1887. Brasil (Amazonas). C. esmeralda (Doubleday, 1845). Brasil (Pará). = C. rubina (Fassl, 1922). C. pireta (Stoll, 1780). = Papilio menander Drury, 1782. a) C. pireta pireta (Stoll, 1780). Central America. b) C. pireta magdalenensis Constantino, 1995. Central Colombia (Rio Magdalena). C. cliftoni (Constantino, 1995). Colombia (Meta, Caquetá, Putumayo), Ecuador (Napo). = C. ereba Clifton M/S. Nomen nudum. C. aurora (C. Felder & R. Felder, 1862). Peru (Huanuco, Junin, Satipo). = Callitaera phantoma Fassl, 1922. = Callitaera pireta aura Langer, 1943. = Cithaerias juruaënsis [sic] D’Almeida, 1951. a) C. aurora aurora (C. Felder & R. Felder, 1862). Peru. b) C. aurora tambopata Lamas, 1998. Peru. C. aurorina (Weymer, 1910). Upper Amazon (Colombia, Peru, Brasil). = C. merolina (J. Zikan, 1942). C. pyritosa (Zikán, 1942). Brasil, Colombia, Peru (Amazonas).

3 Supplementary Material: Constantino, 2016. Revision of the tribe Haeterini (Lepidoptera: Satyirnae)

= Cithaerias similigena D’Almeida, 1951. = Callitaera pireta amaryllis Bryk, 1953. C. pyropina (Salvin & Godman, 1868). a) C. pyropina pyropina (Salvin & Godman, 1868). Peru. b) C. pyropina songoana (Langer, 1944). Bolivia.

Table 1. DNA analyses of specimens of Cithaerias used in molecular phylogenetic analyses and their GenBank (NCBI) accession numbers for the genes sequenced.

Marker Taxon Sequence Reference COI C. pireta pireta GenBank 1 tacattatac tttatttttg gaatttgggc cggtatagta ggaacttccc ttagtcttat KP848792.1 (Panama) 61 tattcggata gaattaggaa accccagatt tttaattgga gatgatcaaa tttataatac 121 tattgttact gcccatgcat ttattataat tttttttata gtaataccta ttataattgg 181 ggggtttggc aattggcttg tccctcttat attaggagcc cctgatatag ctttccctcg Basset et. al. 241 cataaataac ataagatttt gacttctccc cccctcttta attttactaa tttcaagtag 2015 301 tatcgtagaa aatggggcag gtacaggatg aacagtgtac cctccacttt catctaatat 361 tgcccataga ggttcttcag ttgatttagc aattttttca ttacatttag ctggtatctc 421 ctctatttta ggagcaatta attttattac tacaattatt aatatacgaa ttaatataat 481 atcttatgat caaatacccc tattcgtttg atctgttgga atcacagcat tacttcttct 541 tctttcactt cctgttttag caggagctat tactatactt ttaactgatc gtaatctaaa 601 cacctctttt tttgatccag cagggggagg agatcctatt ctttatcaac atttattt

Cytb C. pireta 1 attcaacctg aatgatattt tttatttgcc tatgcaattt tacgttctat tcctaataaa GenBank 61 ttaggaggag taattgcctt aattatatcc atttttattt taattattct accttttacc EU660001.1 magdalenensis 121 ttcaataaaa aaattcaagg tatgcaattt tatcctttaa atcaattctt attttgaatt Constantino, 181 tttatcgtta ctatttttct tttaacatga attggcgctc gccccgtaga aatattatat 1995 241 gtaattacaa gtcaattttt aactttattc tattttttat attttattat caacccaatt Marin et. al. 301 ttaaataaat tttgagatac attaattttt aaattttaat taatgagctt gtttaagcat 2009 (Colombia) 361 ttgttttgaa aacttaagaa agaactttat tctattaatt taattttttt ttaatactaa 421 attaatttta tagtataaaa ttaattttac accaataaaa aataataaat aatttaaaga 481 aataggcaaa tatctttttc aacataaata tattaattta tcata

COI C. aurora 1 tcacctgata tagcattccc ccgtataaat aatataagat tttgacttct gcctcctgct GenBank 61 ttaattttat taatttcaag tagtatcgtc gaaagtgggg caggaaccgg atgaacagtg AY508517.1 (Peru) 121 taccccccac tctcatctaa cattgcccac agaggttctt cagttgattt agcaattttt 181 tcgttacatt tagctggtgt ctcctctatt ttaggagcaa ttaattttat tacaacaatt 241 attaatatac gagttaataa aatatcctat gatcaaatac ccttatttgt ttgatctgtg Murray &. 301 ggaattacag cattacttct tcttctttct cttcctgttt tagcaggagc tatcactata Powell, 2005 361 cttttaactg atcgtaactt aaacacttct ttttttgacc ccgcaggagg aggagaccct 421 attctttatc aacatttatt ttgatttttt ggtcaccctg aagtttatat tttaatttta 481 ccaggatttg gaataatttc tcatattatt tctcaagaaa gtggtaaaaa agaaactttt 541 ggttgtttag gaataattta tgcaatacta gctattggtt tattaggatt tattgtatga 601 gctcatcata tatttaccgt tggtatagat attgataccc gagcttattt tacttccgct

4 Supplementary Material: Constantino, 2016. Revision of the tribe Haeterini (Lepidoptera: Satyirnae)

COI C. aurorina 1 tgagcaggta taattgggac atcccttagt ctcattattc ggatagaatt aggtaaccct GenBank 61 agttttttaa ttggagatga ccaaatttat aatactattg ttactgctca tgcatttatt (Peru) DQ338756.1 121 ataatctttt ttatagtaat acctattata attggaggat ttggtaattg acttgtcccc 181 cttatacttg gagcccctga tatagctttc ccccgtataa ataatataag attttgactt 241 ttacctcctg ctttaatttt attaatttca agtagtatcg tcgaaagtgg agcaggaacc Peña et. al. 301 ggatgaacag tgtacccccc actctcatct aatattgctc acagaggttc ttcagttgat 2006 361 ttagcaattt tttcattgca tttagctggt atttcctcta tcttgggagc aattaatttt 421 attacaacaa ttattaacat acgagttaat aatatatcct atgatcaaat acccctattt 481 gtttgatctg ttgggattac agcattaatt cttcttcttt cccttcctgt tttagcggga 541 gctattacta tacttttaac tgatcgtaat ttaaatactt ctttttttga cccagcagga 601 ggaggggatc ctattcttta ccaacnnnnn nnnnnnnnnn nnnnnnnnnn nnnnnnntat

Figure 1. Male genitalia. a. polita b. Paradulcedo mimica c. Pseudohaetera hypaesia d. negra e. Haetera macleannania f. pireta g. Cithaerias cliftoni h. Cithaerias aurorina i. Cithaerias pyritosa.

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Figure 2. Cithaerias. a-b. C. cliftoni ♂ ♀ (Putumayo, Colombia) c-d. C. pyritosa ♂ ♀ (Amazonas, Colombia) e-f. C. aurorina ♂ ♀ (Rio Amazonas, Colombia) g. C. aurora (Huanuco, Perú). h. C. pireta pireta ♂ (Chocó, Colombia).

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Figure 3. a. C. pireta pireta ♀ (Chocó, Colombia) b. C. pireta magdalenensis (Rio Porce, Antioquia, Colombia) c. C. pyropina (Junin, Peru). d. C. andromeda andromeda (Manaus, Brazil) e. C. bandusia (Amazonas, Brazil) f. C. esmeralda (Pará, Brazil). g. Paradulcedo mimica (Chocó, Colombia) h. Dulcedo polita (Rio Anchicayá, Valle, Colombia).

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Figure 4. a. Pseudohaetera hypaesia (Valle, Cordillera Occidental, Colombia) b-c. Haetera macleannania ♂ ♀ (Rio Raposo, Valle, Colombia) d-e. Haetera piera negra (Puerto Nariño, Amazonas, Colombia) f-g. H. piera sanguinolenta ♂ ♀ (Serrania de La Macarena, Meta, Colombia). h. Haetera diaphana (Bahia, Brazil).

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REFERENCES

Basset, Y., Barrios, H., Segar, S., Srygley, R.B., Aiello, A., Warren, A.D., Delgado, F., Coronado, J., Lezcano, J., Arizala, S., Rivera, M., Perez, F., Bobadilla, R., Lopez,Y., Ramirez, J.A. 2015. The Butterflies of Barro Colorado Island, Panama: Local Extinction since the 1930s. PLoS ONE 10 (8), E0136623.

Constantino, L. M. 1992. Paradulcedo, a new of Satyrinae (Nymphalidae) from western Colombia. Journal of the Lepidopterists’ Society 46:44-53.

Constantino, L. M. 1993. Notes on Haetera from Colombia, with description of the immature stages of Haetera piera (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera 4:13-15.

Constantino, L. M. 1995. Revisión de la tribu Haeterini Herrich-Schäffer, 1864 en Colombia (Lepidoptera: Nymphalidae, Satyrinae). SHILAP Revista de Lepidopterología 23:49-76

D’Almeida, R.F. 1951. Ligeiras observações sôbre o gênero Cithaerias Hübner, 1819 (Lep. Satyridae). Arquivos de Zoologia do Estado de São Paulo, 7 (10): 493–505.

Fassl, A. 1922. Zwei neue Callitaera-Formen (Lep.). Entomologische Zeitschrift, 36 (6): 22.

Felder, C., Felder, R. 1862. Specimen faunae lepidopterologicae riparum fluminis Negro superioris in Brasilia septentrionali. Wiener entomologische Monatschrift 6 (6): 175–192.

Lamas, G. 1998. Lista sinonímica de los géneros Cithaerias Hübner y Haetera Fabricius (Lepidoptera: Nymphalidae: Satyrinae), con la descripción de dos subespecies nuevas del Perú. Revista peruana de Entomología, 40: 133–138.

Lamas, G. 2004. Nymphalidae. Satyrinae. Tribe Haeterini. In: Lamas, G. (Ed.), Checklist: Part 4A. Hesperioidea - . In: Heppner, J. B. (Ed.), Atlas of Neotropical Lepidoptera. Volume 5A. Association for Tropical Lepidoptera; Scientific Publishers, Gainesville, pp. 205–206.

Marin ,M.A., Lopez,A., Lucci Freitas,A.V., Uribe ,S.I. 2009. Caracterización molecular de Euptychiina (Lepidoptera:Satyrinae) del norte de la cordillera central de los Andes. Revista Colombiana de Entomologia 35(2):235-244.

Murray, D., Prowell, D.P. 2005. Molecular phylogenetics and evolutionary history of the neotropical Satyrine Subtribe Euptychiina (Nymphalidae: Satyrinae) Molecular Phylogenetics and Evolution 34 (1): 67-80.

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Penz, C., Alexander, L., DeVries, P. 2014. Revised species definitions and nomenclature of the rose colored Cithaerias butterflies (Lepidoptera, Nymphalidae, Satyrinae). Zootaxa 3873(5): 541-559.

Peña, C., Wahlberg, N., Weingartner, E., Kodandaramaiah, U., Nylin, S., Freitas, A.V., Brower, A.V. 2006. Higher level phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data. Molecular Phylogenetics and Evolution 40:29-49.

Weymer, G. 1910. 4. Familie: Satyridae. In: Seitz, A. (Ed.), Die Gross-Schmetterlinge der Erde. A. Kernen, Stuttgart, pp. 173–283.

Cite as:

Constantino, L. M. 2016. Morphological and molecular evidence supports recognition of Cithaerias cliftoni (Constantino, 1995) as a species distinct from C. pireta (Stoll, 1780) and C. aurorina (Weymer, 1910) Supplementary material. Document 1: 1-10.

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