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Morphological and molecular evidence supports recognition of Cithaerias cliftoni (Constantino, 1995) as a good species distinct from C. pireta (Stoll, 1780) and C. aurorina (Weymer, 1910) (Lepidoptera: Nymphalidae: Satyrinae)

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Cithaerias cliftoni Constantino, 1995 is a member of the tribe Haeterini (Lepidoptera: Nymphalidae: Satyrinae) confined to the Neotropical region. The butterflies 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.
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Luis Miguel Constantino
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)
Suppl emen tary Material: Constantino, 2016. Revision of the tribe Haeterini (Lepidoptera: Satyirnae)
Remarks on the nominal taxon Cithaerias cliftoni (Constantino, 1995)
Cithaerias cliftoni Constantino, 1995 is a member of the tribe Haeterini (Lepidoptera:
Nymphalidae: Satyrinae) confined to the Neotropical region. The butterflies 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 Pein 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
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Suppl emen tary 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 spinesa 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 Venezuela, Colombia to Bolivia.
HAETERA Fabricius, 1807.
H. macleannania H. W. Bates, 1865. Central America to W. Colombia and W. Ecuador.
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Suppl emen tary Material: Constantino, 2016. Revision of the tribe Haeterini (Lepidoptera: Satyirnae)
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).
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Suppl emen tary 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
Reference
COI
C. pireta pireta
(Panama)
GenBank
KP848792.1
Basset et. al.
2015
Cytb
C. pireta
magdalenensis
Constantino,
1995
(Colombia)
GenBank
EU660001.1
Marin et. al.
2009
COI
C. aurora
(Peru)
GenBank
AY508517.1
Murray &.
Powell, 2005
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Suppl emen tary Material: Constantino, 2016. Revision of the tribe Haeterini (Lepidoptera: Satyirnae)
COI
C. aurorina
(Peru)
GenBank
DQ338756.1
Peña et. al.
2006
Figure 1. Male genitalia. a. Dulcedo polita b. Paradulcedo mimica c. Pseudohaetera
hypaesia d. Haetera piera negra e. Haetera macleannania f. Cithaerias pireta pireta g.
Cithaerias cliftoni h. Cithaerias aurorina i. Cithaerias pyritosa.
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Suppl emen tary Material: Constantino, 2016. Revision of the tribe Haeterini (Lepidoptera: Satyirnae)
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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Suppl emen tary Material: Constantino, 2016. Revision of the tribe Haeterini (Lepidoptera: Satyirnae)
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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Suppl emen tary Material: Constantino, 2016. Revision of the tribe Haeterini (Lepidoptera: Satyirnae)
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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Suppl emen tary Material: Constantino, 2016. Revision of the tribe Haeterini (Lepidoptera: Satyirnae)
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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 genus 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): 493505.
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): 175192.
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: 133138.
Lamas, G. 2004. Nymphalidae. Satyrinae. Tribe Haeterini. In: Lamas, G. (Ed.), Checklist:
Part 4A. Hesperioidea - Papilionoidea. In: Heppner, J. B. (Ed.), Atlas of Neotropical
Lepidoptera. Volume 5A. Association for Tropical Lepidoptera; Scientific Publishers,
Gainesville, pp. 205206.
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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Suppl emen tary Material: Constantino, 2016. Revision of the tribe Haeterini (Lepidoptera: Satyirnae)
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. 173283.
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.
10
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Lista sinonímica de los géneros Cithaerias Hübner y Haetera Fabricius (Lepidoptera: Nymphalidae: Satyrinae), con la descripción de dos subespecies nuevas del Perú
  • G Lamas
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.