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Revised species definitions and nomenclature of the rose colored Cithaerias butterflies (Lepidoptera, Nymphalidae, Satyrinae)

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This study provides updated species definitions for five rose-colored Cithaerias butterflies, starting with a historical over-view of their taxonomy. Given their mostly transparent wings, genitalia morphology yielded the most reliable characters for species definition and identification. Genitalic divergence is more pronounced when multiple species occur in sympa-try than between parapatric taxa. Cithaerias aurorina is granted full species status, C. cliftoni is reinstated as a full species, and one new combination is proposed, i.e. C. aurora tambopata. Two new synonyms are proposed, Callitaera phantoma and Callitaera aura = Cithaerias aurora.
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Accepted by C. Prieto: 3 Oct. 2014; published: 20 Oct. 2014
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2014 Magnolia Press
Zootaxa 3873 (5): 541
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http://dx.doi.org/10.11646/zootaxa.3873.5.5
http://zoobank.org/urn:lsid:zoobank.org:pub:05BD334C-493D-4688-92E8-602943ECF57D
Revised species definitions and nomenclature of the rose colored Cithaerias
butterflies (Lepidoptera, Nymphalidae, Satyrinae)
CARLA M. PENZ
1
, LAURA G. ALEXANDER
2
& PHILIP J. DEVRIES
3
Department of Biological Sciences, University of New Orleans, 2000 Lakeshore Dr. New Orleans, LA 70148, USA.
E-mail:
1
cpenz@uno.edu;
2
lgalexan@uno.edu;
3
pdevries@uno.edu
Abstract
This study provides updated species definitions for five rose-colored Cithaerias butterflies, starting with a historical over-
view of their taxonomy. Given their mostly transparent wings, genitalia morphology yielded the most reliable characters
for species definition and identification. Genitalic divergence is more pronounced when multiple species occur in sympa-
try than between parapatric taxa. Cithaerias aurorina is granted full species status, C. cliftoni is reinstated as a full species,
and one new combination is proposed, i.e. C. aurora tambopata. Two new synonyms are proposed, Callitaera phantoma
and Callitaera aura = Cithaerias aurora.
Key words: pireta, menander, aurorina, cliftoni, aurora, aura, phantoma, pyritosa
Introduction
Some of the most visually striking Neotropical butterflies belong to the genus Cithaerias Hübner (Satyrinae,
Haeterini), which inhabit sea level to mid-elevation rainforests from Mexico through Central and South America.
A characteristic of all Cithaerias species is their mostly transparent wings with the distal portions of the hind wing
overlaid with partially lustrous rose, purple or blue scales. These butterflies glide through the understory along the
forest floor (Alexander & DeVries 2012), and it is during flight that the hind wing colors are most conspicuous.
Considered as a whole Cithaerias can be roughly divided into three color groups (based on similarity, not
necessarily common ancestry). The list below is based on the checklist by Lamas (2004), but some of this
nomenclature is revised here:
(1) blue group: small body size, includes the type species C. andromeda andromeda (Fabricius) and its
subspecies C. andromeda azurina (J. Zikán), C. andromeda bandusia Staudinger, and C. andromeda esmeralda
(Doubleday);
(2) purple/rose group: large body size, includes C. pyropina pyropina (Salvin & Godman), and its subspecies
C. pyropina songoana (Langer);
(3) rose group: medium body-size, includes C. pyritosa (J. Zikán); C. phantoma (Fassl); plus C. pireta pireta
(Stoll), and its subspecies C. pireta aura (Langer), C. pireta aurora (C. Felder & R. Felder), C. pireta aurorina
(Weymer), C. pireta magdalenensis Constantino, and C. pireta tambopata Lamas.
Although they are well represented in collections, historically there has been considerable confusion with
respect to species identification of the rose colored Cithaerias. This stems primarily from similarity among taxa,
intra-specific variation in color patterns, lost type specimens (or corresponding dissections), and dubious or
unreported type localities. This study seeks to clarify some of the confusion in several ways. First we use historical
information and comparative morphology to demonstrate that C. pireta as currently defined (Lamas 2004) includes
more than one species. Genitalia morphology provided the main character set for species delimitation, allowing us
to identify intra-specific variation in wing color that helps explain difficulties in species identification. Second we
provide illustrations, diagnoses and geographical distributions for each rose colored Cithaerias species, propose
new synonyms, and illustrate female genitalia for the first time. We also discuss the importance of genitalia
morphology and precise collecting locality for identification of these butterflies.
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Material and methods
Examined specimens are from the McGuire Center for Lepidoptera and Biodiversity, University of Florida (UFL),
the Carnegie Museum of Natural History (CMNH), Natural History Museum of Los Angeles County (LACM), and
collections of P. DeVries and L. Alexander (PJD, University of New Orleans).
Specimen examination followed standard techniques. Male and female abdomens were prepared by soaking in
warm 5% KOH solution, and stored in a 3:1 solution of glycerol and 70% ethanol. Illustrations of genitalia were
made using a stereomicroscope equipped with a camera lucida. Terminology follows Kristensen (2003), except for
venation. A distribution map was generated using Google Maps (dubious localities, or those that could not be
found do not appear in the map). Abbreviations used throughout are: D, dorsal; V, ventral; FW forewing; HW, hind
wing; CMP, Carla M. Penz, LGA, Laura G. Alexander, PJD, Philip J. DeVries.
Results
Historical overview
Except for the distinctive C. pyritosa, other rose colored Cithaerias have a convoluted taxonomic history, and
several taxa were originally described under the genus Callitaera Butler (a synonym of Cithaerias; Lamas 2004).
The information below provides a historical backdrop for the revisions proposed here. World-wide-web links to
referenced images of type specimens correspond to dorsal view only, and a complete set of images can be found at
the Butterflies of America webpage http://www.butterfliesofamerica.com/L/t/Cithaerias_a.htm (last accessed 22
June 2014).
(1) Caspar Stoll completed the publication of Cramer’s “De uitlandische kapellen voorkomende in de drie
Waereld-Deelen, Asia, Africa en America (1775–1791) which includes an illustration of his C. pireta (Plate
CCCXV, A; reproduced in Fig. 1a). The text (p 52) does not mention a type locality, and Lamas (1998) suggested
the type material is likely lost. Despite the poor quality of the illustration in Cramer’s volume, the broad brown
band at the hind wing edge and the continuous postmedial brown line (not staggered) match C. pireta specimens
from Central America, Colombia and Ecuador very well. Some specimens examined by us also have a rose blush
that reaches the hind wing discal cell, and display two strong brown lines across the forewing (e.g., Fig. 2g). The
new species Papilio menander Stoll was also included in Cramer’s volumes, a riodinid now named Menander
menander (Stoll).
(2) Dru Drury (1782) described his Papilio menander Drury (preoccupied by Stoll 1780, see above) from
Jamaica (type locality) where Cithaerias does not occur. Lamas (1998) suggested the type material is likely lost.
(3) Cajetan Felder and Rudolf Felder (1862) described “Haetera pireta Cram. var. aurora” (verbatim; note
reference to Cramer instead of Stoll) from Colombia and northern Brazil. The title of their publication, “Specimen
faunae lepidopterologicae ripaurm fluminis Negro superioris in Brasilia septentrionali”, indicates that the focal
species were from northern Brazil. Moreover, the description of aurora states that two male specimens were
received from Henry W. Bates (who did not collect in Peru), and several others came from the Rio Negro region.
We, therefore, find it unlikely that the type locality of aurora is Peru (Huallaga valley) as ascertained by Lamas
(1976) and upheld by Constantino (1995) and Lamas (1998). The type specimen is in the Natural History Museum,
UK (note the genitalia are missing) http://www.butterfliesofamerica.com/L/ih/n_cithaerias0025_i.htm (last
accessed 22 June 2014).
(4) In Adalbert Seitz’s, “Die Gross-Schmetterlinge der Erde”, Gustav Weymer (1910) described Callitaera
aurora form aurorina from Colombia and upper Amazon (original illustration reproduced in Fig. 1d). Lamas
(1998) designated a lectotype that closely resembles the original illustration, and is housed at the Zoologisches
Museum, Berlin, Germany (the label ‘Magdalena’ suggests Colombian origin) http://
www.butterfliesofamerica.com/L/ih/n_cithaerias0027_i.htm (last accessed 22 June 2014). Note that the postmedial
and submarginal hind wing bands of the lectotype (image posted online) are less arched than those in Fig. 5f
(Colombia, Rio Loretoyacu). Weymer’s (1910) species accounts plus male and female illustrations of menander
(Central America, Colombia) appear to correspond to the single Cithaerias species present in Central America, and
his description and illustrations of pireta (from Upper Amazon, Ecuador) resemble the Ecuadorean specimens
examined here, especially the females (compare Fig. 1b with 2d–e).
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(5) Anton H. Fassl (1922) described Callitaera phantoma from Manicoré at Rio Madeira, Tefé and São Paulo
de Olivença, Amazonas, Brazil. Lamas (1998) was not able to locate the type material.
(6) Josef F. Zikán (1942) described Callitaera pyritosa from São Gabriel [da Cachoeira], Rio Negro,
Amazonas, Brazil. Male genitalia were illustrated in lateral view, but the phallus is missing and the saccus is
incomplete—both of which are key features for species identification. The lectotype designated by Mielke &
Casagrande (1987) is in the Instituto Oswaldo Cruz, Rio de Janeiro, Brazil http://www.butterfliesofamerica.com/L/
ih/n_cithaerias0037_i.htm (last accessed 22 June 2014). In the same article, Zikán also described Callitaera
merolina Zikán from Iauareté, Amazonas, Brazil (near the border with Colombia). Male genitalia were illustrated
in lateral view (similar to Fig. 7b). The lectotype designated by Mielke & Casagrande (1987) is in the Instituto
Oswaldo Cruz, Rio de Janeiro, Brazil http://www.butterfliesofamerica.com/L/ih/n_cithaerias0029_i.htm (last
accessed 22 June 2014).
(7) Focusing on Ecuadorian taxa only, Frederick M. Brown (1942) illustrated hind wing characters and the
male genitalia of C. menander in lateral view (that match Fig. 5a), plus hind wing characters and valva of C. pireta
(similar to Fig. 5b). Not surprisingly his accounts of C. menander and C. pireta correspond to the illustrations of
such species in the Seitz catalogue (Weymer 1910), a standard reference at the time. Nonetheless, he regarded both
aurora and aurorina as subspecies of C. pireta, thus broadening the definition of this species (see also items (9)
and (12)).
(8) G. Langer (1943) described Callitaera aura and listed the type locality as Brazil, São Paulo [de Olivença,
possibly] (one male) and Bahia (locality likely incorrect, one female). The lectotype designated by Lamas (1998) is
in the Zoologische Sammlung des Bayerischen Staates (ZSBS), München, Germany (genitalia slide not found, U.
Buschbaum pers. comm. March 2014) http://www.butterfliesofamerica.com/L/ih/n_cithaerias0021_i.htm (last
accessed 22 June 2014). Although Lamas (1998) clearly indicated that a lectotype was being designated for
Callitaera aura, the specimen in the ZSBS is labeled as a holotype.
(9) Romualdo F. D’Almeida (1951) described Cithaerias similigena D’Almeida from Rio Içana, São Joaquim
[Mirin], Amazonas, Brazil. Photographs of the male genitalia were provided (similar to Fig. 7b), and the holotype
is at the Universidade Federal do Paraná, Brazil http://www.butterfliesofamerica.com/L/ih/n_cithaerias0039_i.htm
(last accessed 22 June 2014). He also described Cithaerias juruaënsis [sic] D’Almeida from Porto Walter, Acre,
Brazil from a single specimen, and such holotype is at the Universidade Federal do Paraná, Brazil http://
www.butterfliesofamerica.com/L/ih/n_cithaerias0023_i.htm (last accessed 22 June 2014). In an attempt to clarify
the menander-pireta confusion, D’Almeida (1951) examined original descriptions and illustrations from several
sources, including those in Cramer (1779–1782). Despite the poor quality of the C. pireta illustration, he
considered C. menander and C. pireta to be the same taxon based on a specimen from Maroni River, French
Guiana in his collection that matched Cramer’s figure of C. pireta (see Fig. 1a and item (1) above). D’Almeida
suggested that C. aurora and C. phantoma might correspond to the same species, and provided a checklist in which
he synonymized C. menander with C. pireta and granted C. aurorina full species status.
(10) Felix Bryk (1953) described Callitaera pireta amaryllis Bryk from São Gabriel [da Cachoeira, probably],
Rio Negro, Amazonas, Brazil. The holotype is at Naturhistoriska Riksmuseet, Stockholm, Sweden (note extended
phallus) http://www.butterfliesofamerica.com/L/ih/n_cithaerias0041_i.htm (last accessed 22 June 2014).
(11) Bernard D’Abrera (1988) used the names C. ereba ereba (from Colombia) and C. ereba browni (from
Ecuador) from an unpublished manuscript by Michael Clifton, and therefore both have become nomen nudum. The
photographs of both taxa in D’Abrera (1988:740) resemble the illustration of C. pireta by Weymer (1910;
reproduced in Fig. 1c).
(12) Luis Constantino (1995) described Cithaerias pireta cliftoni Constantino from Meta, Villavicencio,
Colombia, and the holotype is in the Museo de Historia Natural, Universidad de Caldas, Manizales, Colombia
http://www.butterfliesofamerica.com/L/ih/n_cithaerias0019_i.htm (last accessed 22 June 2014). Constantino
named this subspecies after Michael Clifton in recognition of the fact that he was the first to recognize this taxon
from Colombia (his ereba, see item (11) above). The holotype of C. pireta cliftoni, the illustration of C. pireta by
Weymer (1910), and Fig 1d are notably similar. In the same study Constantino also described Cithaerias pireta
magdalenensis from Caldas, Rio Guarinó, Colombia, and the holotype is in the Museo de Historia Natural,
Universidad de Caldas, Manizales, Colombia (note discrepancy between published type locality and holotype
label) http://www.butterfliesofamerica.com/L/ih/n_cithaerias0031_i.htm (last accessed 22 June 2014). Finally,
Constantino (1995) pointed to inconsistencies in the identification and nomenclature of Cithaerias, and made
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helpful clarifications by providing photographs of both sexes and illustrations of male genitalia of Colombian
species. He followed D’Almeida (1951) by treating C. menander as a synonym of C. pireta, and Brown (1942) by
treating aurora and aurorina as subspecies of C. pireta.
(13) Gerardo Lamas (1998) described Cithaerias pireta tambopata from Boca del rio La Torre (Zona
Reservada de Tambopata), Madre de Dios, Peru, and the holotype is in the Museo Nacional de Historia Natural,
Universidad Mayor de San Marcos, Lima, Peru http://www.butterfliesofamerica.com/L/ih/n_cithaerias0033_i.htm
(last accessed 22 June 2014). He noted that Papilio menander Drury (a species of Cithaerias) was a junior
homonym of Papilio menander Stoll (Riodinidae), and proposed five new synonyms for the rose colored
Cithaerias: C. pireta cliftoni = C. phantoma, C. juruaënsis = C. pireta aura, C. merolina = C. pireta aurorina, C.
similigena = C. pyritosa, and C. pireta amaryllis = C. pyritosa.
(14) Lamas (2004) provided a complete checklist for Cithaerias, including synonyms and formal recognition
that name menander Drury is preoccupied.
Character systems
Wings. Given the profound absence of scales in all Cithaerias the two forewing brown bands are expressed on the
wing membrane itself, and may be well-developed, weak, or absent. Although individuals from the same locality
are usually similar, we observed intraspecific, geographical variation, and the forewing bands do not seem
particularly useful for species identification (compare individuals of the same species in each of the figures in this
study). The hind wing brown bands are also expressed on the wing membrane, and their alignment, width, and
absence from certain cells can be used for species diagnoses. On the dorsal hind wing surface, rose scales cover
these bands slightly, but the brown bands are quite visible on the ventral side (which lacks rose scaling). The
female brown bands on the hind wing are wider than in males. The dorsal hind wing rose scaling varies in hue,
density, and magnitude between species and subspecies, but it also varies among conspecifics from the same
locality, fading as they age. Although the rose scaling provides some information for species identification,
different species overlap in color pattern. The hind wing ocellus and additional spots that correspond to the border
ocelli (pattern element) are expressed on both wing surfaces and are formed by colored scales only (not by
melanization of the wing membrane). The size of this ocellus varies between individuals of the same locality.
Genitalia. Based on our experience (Nymphalidae in general, and Satyrinae in particular), the divergence in
genitalia morphology among subspecies of C. pireta sensu Lamas (2004) is markedly strong and justifies their
revised status. The morphology and size of the components of male genitalia are species-specific. All species have
elaborate subscaphium, which is composed of a sub-anal plate with minute microtrichia plus an adjacent
moderately sclerotized area that bears larger microtrichia (located between the sub-anal plate and the anellus). The
shape, size and sclerotization of the latter are particularly important for identifying two cryptic species. The female
sterigma morphology and size vary among species. In most rose colored taxa the post-sterigmal area contains a
small, round, concave, and heavily sclerotized indentation. When the genitalia are prepared, the post-sterigma
region becomes limp and this indentation may fold inward (behind the sterigma). In most of the specimens we
dissected a film-like coating detached from the sterigma when the abdomen was soaked in KOH. Although the
shape and size of this structure varied between specimens, it formed a cast of the sterigma and ostium bursae,
suggesting that it functions as a mating plug. This was also observed in dry specimens (Fig. 8f). The corpus bursae
is extremely delicate, and both the antrum and ductus bursae varied in size corresponding to variation in male
phallus length.
Taxonomy
Revised checklist for rose colored Cithaerias
(---- indicates a synonym, - indicates a subspecies)
Cithaerias pireta (Stoll, 1780)
----Papilio menander Drury, 1782
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-Cithaerias pireta magdalenensis Constantino, 1995
Cithaerias cliftoni Constantino, 1995 STAT. REV.
Cithaerias aurora (C. Felder & R. Felder, 1862) STAT. REV.
----Callitaera phantoma Fassl, 1922 NEW SYN.
----Callitaera pireta aura Langer, 1943 NEW SYN.
----Cithaerias juruaënsis [sic] D’Almeida, 1951
-Cithaerias aurora tambopata Lamas, 1998 NEW COMB.
Cithaerias aurorina (Weymer, 1910) STAT. REV.
----Callitaera merolina Zikan, 1942
Cithaerias pyritosa (Zikán, 1942)
----Cithaerias similigena D’Almeida, 1951
----Callitaera pireta amaryllis Bryk, 1953
FIGURE 1. a, Illustration of C. pireta in Cramer’s “De uitlandische Kapellen voorkomende in de drie Waereld-Deelen, Asia,
Africa en America. bd, Illustrations of C. menander, pireta and aurorina in Seitz’s “Die Gross-Schmetterlinge der Erde”.
Cithaerias Hübner
Callitaera Butler
Type species: Papilio andromeda Fabricius, designated by Scudder (1875).
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Cithaerias pireta (Stoll)
(Figs 2, 6a, 8a, 9)
Papilio menander Drury, preocc. Stoll.
Type material: Likely lost (Lamas 1998).
Distribution. Central America, plus Colombia and Ecuador west of the Andes, pacific slope. See map (Fig. 9)
for examined specimens.
Diagnosis. This species is defined based on the following combination of characters (numbered in Fig. 2): (1)
male HW submarginal and marginal bands forming even arches and often amalgamated to produce a wide brown
outline distally on the wing; (2) male HW postmedial band that outlines the ocellus usually complete across cells
Rs through M2; (3) male HW postmedial band approximates an arch across cells M3 through Cu2 (more so than
any other species); (4) male HW rose scaling diffuse, often reaching the edge of the discal cell; (5) female similar
to male but with paler scaling and wider HW brown bands. Male genitalia (Fig. 6a): in lateral view, note the
sharply bent uncus; in dorsal view, the angular lateral edges of uncus plus tegument; in ventral view, the inner
projection of the valva. Phallus short. Subscaphium triangular-shaped and bearing small spines. The subscaphium
is not evident given it is weakly sclerotized. Female genitalia (Fig. 8a): note short sterigma, which is less
sclerotized than those of other species. Post-sterigmal area weakly sclerotized, lacking the discrete heavily
sclerotized indentation present in other species.
FIGURE 2. C. pireta; dorsal view on the left, ventral on the right. Collecting localities and genitalia dissection codes appear below
each specimen.
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Variation in wing pattern (Fig. 2). In both males and females FW and HW medial brown bands may be well-
developed, incomplete or absent (compare Fig. 2a and g). HW postmedial and marginal bands can vary in width,
being extremely thin in some specimens. Male and female HW rose scaling usually does not reach discal cell, but
these scales may enter the discal cell in some specimens. Male HW white spot in cell M2 present or absent, but
normally present in females. Some examined specimens showed wing wear (fewer scales), and the scales of a few
old collection specimens appear to have faded.
Subspecies. Cithaerias pireta magdalenensis Constantino; holotype in the Museo de Historia Natural,
Universidad de Caldas, Manizales, Colombia, http://www.butterfliesofamerica.com/L/ih/n_cithaerias0031_i.htm
(last accessed 22 June 2014).
Remarks. This species is distinct from all others in both wing and genitalia characters. We concur with
D’Almeida (1951) that Cramer’s figure seems to correspond to C. pireta (compare Figs 1a and 2) and that C.
menander should be considered a synonym of C. pireta. Consequently we are also confident that the illustrations of
C. menander in Weymer (1910) and Brown (1942) correspond to C. pireta. Male genitalia from specimens
collected close to the type locality of C. pireta magdalenensis match those in specimens from Central America and
Ecuador (also noted by Constantino 1995). We note that the defining wing pattern characters of C. pireta
magdalenensis are not unique to specimens of the region for which this taxon has been described (Caldas, Rio
Guarinó, Colombia; Constantino 1995). According to the original description, these are: (1) the FW brown
transversal lines are faded, and so is the HW transversal line (also in specimens from Nicaragua and Costa Rica,
Fig. 2a and c); (2) the HW show a marginal line, in addition to the submarginal, which is not present in C. pireta
pireta males (note such line in specimens from Costa Rica and Panama, Fig. 2c and d); (3) the submarginal line [of
the HW] is narrower and more faded (this characters varies within localities and is also present in the specimen in
Fig. 2d); (4) the HW rose blush ranges from the wing margin to M2, not entering the cell (this is true for most
specimens of C. pireta); (5) the same as in the male, the female shows a well defined white spot between HW veins
M2 and M3 (also present in specimens from Central America and other areas, Fig. 2b and e). Despite this, we
recognize that the type locality is isolated from the remainder of the range of C. pireta, which justifies the
recognition of this taxon. Specimens from that region were not available to us for examination. The use of DNA
sequence to compare specimens from various localities would be of interest.
Material examined: MALE: 1M Honduras, no date (CMNH); 1M Honduras, Lancetilla, 7 November 1930
(CMNH); 1M Nicaragua, Zelaya Dept. Nueva Guinea, 26 October 1976, dissection CMP 13-08 (UFL); 1M
Nicaragua, Zelaya Dept. Nueva Guinea, 27 October 1976 (UFL); 1M Costa Rica, Puntarenas, Finca Las Cruces,
San Vito de Java, 29 September 1986 (UFL); 1M Costa Rica, Turrialba, 25 August 1973 (UFL); 1M Costa Rica,
Puntarenas, San Vito, Las Alturas, 1600 m 15 September 1983 (UFL); 1M Costa Rica, Limon, Ruta 32, 18.9 km W
Compiles (sic), 12 September 1986 (UFL); 2M Costa Rica, Heredia, Sarapiqui, Chilamate, 30 March and 1 April
1989 (UFL); 1M Costa Rica, Limon, Tortuguero, Agua Fria, 50 m, 17 August 1983 (UFL); 1M Costa Rica, San
Jose Carillo, Fila Rio Sucio, 700 m, 21 April 1983 (UFL); 1M Costa Rica, Turrialba, 15 July 1973 (UFL); 1M
Costa Rica, Cartago, Moravia de Chiripo, 1200 m, 11 February 1983 (UFL); 15M Costa Rica, Heredia, Sarapiqui,
Tirimbina Biological Station, various dates in 2009, 2010 and 2011, three dissected LGA 12-01, 12-02 and 12-03
(PJD); 1M Costa Rica, no date (CMNH); 1M Costa Rica, Limon, Guapiles, 9 August 1963 (CMNH); 1M Costa
Rica, Limon, no date (CMNH); 3M Costa Rica, Sixola Riv. (sic), no date (CMNH); 3M Costa Rica, Juan Viñas, 14
and 15 March 1908 (CMNH); 1M Costa Rica, Esperanza, no date (CMNH); 1M, Costa Rica, Turrialba, 26 July
1909 (CMNH); 1M Costa Rica 6 January 1892 (CMNH); 2M Panama, Canal Zone, Piña, 27 July 1970 and July
1970 (UFL); 1M Panama, Gatun, Canal Zone, August 1915 (CMNH); 2M Panama, Madden Dam, 19 and 22 April
1966 (UFL); 1M Panama, Panama, Cerro Campana, 10 January 1973 (UFL); 1M Panama, Darien, Gorgas Bayano
Sta., 27 December 1975, dissection CMP 13-09 (UFL); 1M Panama, Chriqui, no date (CMNH); 1M Colombia 5
December 1889 (CMNH); 1M Colombia, Bogotá, no date (CMNH); 2M, Colombia, Carmen de Yacopi, April
1914 M. Clifton dissection slides 106, 107 (CMNH); 1M Colombia, Valle Calima, 19 March 1988 (UFL); 1M
Colombia, Bajo Calima Valle 300 m, 20 January 1988, dissection CMP 13-11 (UFL); 2M Colombia (UFL); 3M
Ecuador, Huigra, March 1911 (CMNH); 1M Ecuador, Dos Puentes, March 1980 (CMNH); 2M Ecuador,
Tinalandia 21–25 July 1984, one dissected LGA 12-07 (UFL); 1M Ecuador, Pichincha, 12 km E of Santo Domingo
de Los Colorados, 750–850 m 13 May 1988 (UFL); 1M Ecuador, Pichincha, 12 km E of Santo Domingo de Los
Colorados, 750–850 m 5 May 1988, dissection LGA 12-08 (UFL); 1M Ecuador, Pichincha, Loriquin 20 July 1968
dissection LGA 12-09 (UFL); 1M Ecuador, Pichincha, Rio Tachi 22 July 1968 (UFL); 1M Ecuador, El Oro, Los
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Monos February 1973 (UFL); 1M Ecuador, Napo, Limoncocha (likely mislabeled) 26 September 1974 dissection
CMP 13-10 (UFL). FEMALE: 1F Nicaragua, Zelaya Dept. Nueva Guinea, 26 October 1976 (UFL); 2F Costa
Rica, Turrialba, 28 July 1973 and 15 January 1981 (UFL); 1F Costa Rica, Limon, Cerro Guabas, 400 m, 20 January
1979 (UFL); 2F Costa Rica, Heredia, Sarapiqui, Chilamate, 30 March and 1 April 1989 (UFL); 1F Costa Rica,
Alajuela, Rion Angel, La Virgen del Socorro, 9 April 1989 (UFL); 1F Costa Rica, San Jose, Parque Nacional
Carillo, Fila Rio Sucio, 700 m, 20 February 1983 (UFL); 1F Costa Rica, Limon, Paraiso, 3 September 1987 (UFL);
2F Costa Rica, Puntarenas, Finca Las Cruces, San Vito de Java, 29 and 30 September 1986 (UFL); 3F Costa Rica,
Heredia, Sarapiqui, Tirimbina Biological Station, 10 May and 2 September 2010 and 22 February 2011, 1 dissected
CMP 13-38 (PJD); 1F Costa Rica, Sixola Riv. (sic), no date (CMNH); 3F Costa Rica, Juan Viñas, 11 March 1908
(CMNH); 2F Costa Rica, Limon, Guapiles, 9 August 1963 (CMNH); 1F, Costa Rica, Guapiles, no date (CMNH);
1F Costa Rica, Alajuela, Florencia, 13 July 1963 (CMNH); 1F Costa Rica, Puntarenas, Golfito, 17 August 1963
(CMNH); 2F Panama, Chiriqui, no date (CMNH); 1F Panama, no date (CMNH); 1F Panama, Madden Forest, 21
April 1966 (UFL); 2F Panama, Canal Zone, Piña, 2 July and July 1970 (UFL); 1F Panama, Cerro Campana, 2500
[m] 6 November 1964 (UFL); 3F Panama, Darien, Gorgas Bayano Sta., 25 December 1975, 1 dissected CMP 13-
28 (UFL); 1F Colombia (UFL); 1F Colombia, Environs of Bogota, 1913 (likely mislabeled) (UFL); 1F Peru,
Loreto, Iquitos (likely mislabeled) (UFL); 1F Ecuador, Napo, Limoncocha (likely mislabeled) 1 July 1980 (UFL);
1F Ecuador, Pichincha, Rio Tachi 22–25 July 1968 (UFL); 1F Ecuador, El Oro, Los Monos February 1973 (UFL);
2F Ecuador, Pichincha, 12 km E of Santo Domingo de Los Colorados, 750–850 m 13 May 1988 (UFL); 2F
Ecuador, Pichincha, Rio Tachi 21–25 July 1968 and 17 July 1968 (UFL); 1F Ecuador, Pichincha, Alluriquin 13
August 1972 (UFL).
Cithaerias cliftoni Constantino, STAT. REV.
(Figs 3, 6b, 8b–c, 9)
Type material: Holotype is in the Museo de Historia Natural, Universidad de Caldas, Manizales, Colombia http://
www.butterfliesofamerica.com/L/ih/n_cithaerias0019_i.htm (last accessed 22 June 2014).
Distribution. Colombia, Ecuador and Peru, east of the Andes. Available collection records suggest that this
species is not sympatric with C. aurora. See map (Fig. 9) for examined specimens.
Diagnosis. We define this species based on the following combination of characters (numbered in Fig. 3): (1)
male HW marginal band very thin; (2) male HW submarginal band usually thin, clearly separated from marginal
band and slightly staggered; (3) male HW postmedial band that outlines the ocellus complete, reaching vein M3, or
incomplete, not reaching M3; (4) male HW postmedial band usually thin, and usually complete across cells M3
through Cu2, forming a staggered pattern; (5) male distance between HW submarginal and postmedial bands
usually similar to the width of the cells, but variable between cells; (6) male HW rose scaling more diffuse than C.
aurora, less than C. pireta, usually entering discal cell; (7) female with much wider HW brown bands than male,
forming arches in each cell; (8) female HW scaling usually limited to postmedial area but sometimes reaching the
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. Male genitalia (Fig. 6b): 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. Female genitalia (Fig. 8b–c): note the rounded sterigma (slightly variable in shape and
width between specimens), and the heavily sclerotized post-sterigma indentation.
Variation in wing pattern (Fig. 3). In both sexes HW postmedial, submarginal and marginal bands show
some variation in width, but this variation is more pronounced in females. Male HW rose scaling may show some
fuchsia overtones and varies from being limited to the postmedial area (denser scaling pattern, resembling females)
to reaching or entering the discal cell (scaling highly diffuse). Female HW scaling varies in color from off-white to
rose.
Subspecies. None.
Remarks. The illustrations of C. pireta in Weymer (1910) and Brown (1942) correspond to C. cliftoni.
Although the male HW submarginal and postmedial bands are usually thin, in some specimens we found they were
wider and more arched, appearing as female-like males. Male genitalia from a specimen collected in Villavicencio
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(type locality; Fig. 3a) match those from Ecuadorean, Colombian and Peruvian specimens that have more diffuse
rose scaling (Fig. 6b). Although we observed some variation in the valva, tegumen plus uncus, and saccus among
specimens, the morphology of the subscaphium was uniformly consistent and we found no intermediate specimens
between C. cliftoni (Fig. 6b) and C. aurora (Fig. 6c–e).
FIGURE 3. C. cliftoni; dorsal view on the left, ventral on the right—except when indicated (a). Collecting localities and
genitalia dissection codes appear below each specimen.
Material examined: MALE: 1M British Guiana, Demerara River, no date, dissection CMP 13-17 (UFL); 1M
Colombia, no date dissected by M. Clifton as part of a paratype-to-be series, slide 649, labeled as C. ereba, very
similar to the type specimen of C. pireta cliftoni Constantino (CMNH); 1M Colombia, Rio Orteguaza, Rastrojo,
August–September 1947, studied by M. Clifton as part of a paratype-to-be series, labeled as C. ereba medea
(CMNH); 1M Colombia, Putumayo, Orito Rd. 400 m, 3 July 1981, dissection CMP 13-36 (UFL); 1M Colombia,
Boyaca, Villavicencio, Rio Negro, 3 January 1976, dissection CMP 13-12 (UFL); 2M Ecuador, Yasuni National
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Park, 8 September 2002, dissection LG 12-04, and 4 October 1998 (UFL); 1M Ecuador, Oriente, Rio Pumagaco
(sic) 700 m, 5 September 1971, dissection LG 12-05 (UFL); 1M Ecuador, Napo, Coca 300 m, 4 March 1971
dissection LG 12-06 (UFL); 1M Ecuador, Oriente, Rio Latas, 10 September 1971 (UFL); 1M Ecuador, San Jose
Julio 800 m, 1–30 July 1990 (UFL); 2M Ecuador, Napo, Puerto Misahualli, 6 November 1983 (UFL); 1M Ecuador,
Napo, Rio Napo 22–24 August 1978 (LACM); 1M Ecuador, Oriente Rio Finoly (sic), 22 July 1980 (LACM);
Ecuador, Sucumbios, Garzacocha, 16–22 September 1994, dissection CMP 13-23 (UFL); 1M Ecuador, Lake
Taracoa on Rio Napo, 25 June 1980 (UFL); 6M Ecuador, Pastaza, Shiripuno 31 May 2008 dissection CMP 13-26,
31 May 2008, 5 August 2008, 3 September 2008, 3 November 2008, 5 November 2008 (PJD); 2M Ecuador,
Concepción, September 1929, one dissection CMP 14-09 (LACM); 2M Peru, Iquitos, Explorama Lodge 22–28
October 1989, dissection CMP 13-15 (UFL); 3M Peru, Loreto, Rio Amazonas, Iquitos, 9–12 September 1990
(LACM); 1M Peru, Iquitos, June 1919 dissection CMP 14-16 (CMNH); 1M Peru, Yurimaguas, 12 April 1920,
dissection CMP 14-19 (CMNH); 1M Peru, Yurimaguas, 26 April 1920 (CMNH); FEMALE: 1F Colombia, Rio
Orteguaza, Rastrojo, August–September 1947, studied by M. Clifton as part of a paratype-to-be series, labeled as
C. ereba medea (CMNH); 1F Colombia, no date, studied by M. Clifton as part of a paratype-to-be series, labeled as
C. ereba (CMNH); 1F Colombia, Putumayo, Orito Rd. 400 m, 3 July 1981, dissection CMP 13-37 (UFL); 2F
Colombia, Amazonas, Rio Loreto Yacu 8 July–August 16 1981, one dissection CMP 14-06 (LACM); 2F Ecuador,
Yasuni National Park, 8 September 2002, dissection CMP 13-29 and 13-35 (UFL); 1F Ecuador, Napo, Rio
Misahualli, June 1988 (UFL); 1F Ecuador, Napo, Rio Napo, Misahualli 1700 ft. 9 August 1998 (UFL); 1F
Ecuador, Napo, Laguna Taracoa, Rio Napo 800 m, no date (UFL); 1F Ecuador, Napo, Limoncocha, 6 June 1977
dissection CMP 13-39 (UFL); 2F Ecuador, Oriente, Rio Arajuno 700–1000 m, 29 April 1941 and 20 March 1968
(UFL); 6F Ecuador, Pastaza, Shiripuno 3 January 2009 dissection CMP 14-01, and 5 May 2008, 2 August 2008, 5
August 2008, 4 February 2009, 3 May 2009 (PJD); 1F Peru, Loreto, Iquitos, October 2000 (UFL); 1F Peru, Iquitos,
Explorama Inn, 1980’s, dissection CMP 14-15 (LACM).
Cithaerias aurora (C. Felder & R. Felder) STAT. REV.
(Figs 4, 5a–e, 6c–e, 8d, 9)
Callitaera phantoma Fassl NEW SYN.
Callitaera pireta aura Langer NEW SYN.
Cithaerias juruaënsis [sic] D’Almeida
Type material: The type specimen is in the Natural History Museum, UK http://www.butterfliesofamerica.com/L/
ih/n_cithaerias0025_i.htm (last accessed 22 June 2014).
Distribution. Brazil, Colombia and Peru, east of the Andes. Available collection records suggest that this
species is not sympatric with C. cliftoni. See map (Fig. 9) for examined specimens.
Diagnosis. We define this species based on the following combination of characters (numbered in Fig. 4): (1)
male HW marginal band very thin; (2) male HW submarginal band usually thin, clearly separated from marginal
band and slightly staggered; (3) male HW postmedial band that outlines the ocellus usually complete, reaching vein
M3; (4) male HW postmedial band usually thin, forming a staggered pattern, complete across cells M3 through
Cu2 or reduced/absent from cell M3 depending on the locality; (5) distance between male HW submarginal and
postmedial bands usually larger than the width of the cells, variable between cells; (6) male HW rose scaling
diffuse but denser than in C. cliftoni, usually not entering discal cell, or only entering the distal portion of discal
cell; (7) female with much wider HW brown bands than male, forming arches in each cell; (8) female HW scaling
usually limited to postmedial but sometimes reaching the medial area, vivid rose (more so than C. cliftoni). In both
sexes the FW brown bands vary from incomplete (below discal cell only) to nearly absent. Male genitalia (Fig.
6c–e): in lateral view the valva is slightly shorter than that of C. cliftoni, which also lacks inner projection as seen
in ventro-lateral view; in dorsal view the lateral edges of uncus plus tegumen are rounded; and in ventral view the
rounded shape of the sclerotized subscaphium bears large spines—a key difference between C. aurora and C.
cliftoni. Female genitalia (Fig. 8d): note the rectangular sterigma (slightly variable in shape and width between
specimens), and the heavily sclerotized post-sterigma indentation.
Variation in wing pattern (Figs 4, 5ae). In both males and females HW postmedial and marginal bands
shows some variation in width (more pronounced in females), and also the distance between bands. Ocellus below
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M1 varies in size. Male HW rose scaling varies from being limited to the postmedial area to barely entering the
discal cell. We found much geographical variation in the density of scaling and color of the male HW (from pale to
vivid rose), which was the basis for the description of subspecies tambopata (denser scaling, vivid rose). Note that
specimens from Rondônia, Brazil also have a vivid rose coloration (compare Fig. 5a and d).
Subspecies: Cithaerias aurora tambopata Lamas, NEW COMB. Holotype is in the Museo Nacional de
Historia Natural, Universidad Mayor de San Marcos, Lima, Peru http://www.butterfliesofamerica.com/L/ih/
n_cithaerias0033_i.htm (last accessed 22 June 2014). This proposal is justified based on two main sources of
evidence: (1) the close similarity of male valva and subscaphium morphology and sclerotization (compare
specimens from Brazil in Fig. 6c–d to one from Peru in Fig. 6e), and (2) a regional trend in the broad spacing
between HW submarginal and postmedial bands (character (5) in Fig. 4g, see also Fig. 5a).
FIGURE 4. C. aurora; dorsal view on the left, ventral on the right. Collecting localities and genitalia dissection codes appear below
each specimen.
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Remarks. The type specimens of aurora and aura are both faded and nearly identical, and the type material of
phantoma appears to have been lost (see Lamas 1998). Most specimens recognized here as C. aurora have a wider
distance between male HW submarginal and postmedial bands than the aforementioned types, but a few resemble
the pattern of those types. See Historical Information for further details.
Material examined: Cithaerias aurora aurora MALE: 2M Brazil, Rondônia, Fazenda Rancho Grande, 62
km S Ariquemes, 17 August 1993, dissection CMP 13-19, 10 October 1993 (UFL); 1M Brazil, Rondônia, Fazenda
Rancho Grande, 62 km S Ariquemes, 29 September–10 October 1992 (USNM); 1M Brazil, Rondônia, Jaru,
August 1976 (UFL); 1F Brazil, Amazonas, São Paulo de Olivença, 5 November 1971, dissection CMP 13-16
(UFL); 11M Brazil, Amazonas, São Paulo de Olivença, January–April 1923, seven dissected by M. Clifton, slides
102, 164, 169, 170, 171, 174, 176 (CMNH); 1M Brazil, Amazonas, São Paulo de Olivença October 1973 (USNM);
1M Brazil, Amazonas, Tefé, 20 February 1920 and 22 December 1926 dissected by M. Clifton, slides 165 and 166
(CMNH); 5M Brazil, Amazonas, Arimã, November–December 1922, dissected by M. Clifton, slides 139, 140,
141, 142, 144 (CMNH); 25M Brazil, Amazonas, Nova Olinda, Rio Purus, May–July 1922, dissected by M. Clifton,
slides 101, 103, 104, 105, 136, 147, 148, 149, 150, 151, 152, 153, 154, 155, 156, 157, 158, 159, 160, 161, 166, 167,
168, 175 (CMNH); 1M Colombia, Amazonas, Rio Loreto Yacu 8 July–August 16 1981, dissection CMP 14-07
(LACM); 1M Peru, Tingo Maria, March 1981, dissection CMP 13-14 (UFL); 1M Peru, Huanuco, Tingo Maria,
Leonces Prado, 23 June 1982, dissection CMP 13-24 (UFL); 1M Peru, Tingo Maria, 28 January 1972 (USNM);
6M Peru, Junin, Satipo, October 1947, dissection CMP 2M Peru, Junin, Satipo, May and October 1983 (USNM);
14–17 and M. Clifton slides 112, 113 (CMNH); 3M Peru, Sani Benj (sic) 840 m 25 August 1935, 6 September
1935, 31 August 1935 dissected by M. Clifton, slide 221 (CMNH); FEMALE: 2F Brazil, Rondônia, Fazenda
Rancho Grande, 62 km S Ariquemes, 6 August 1992, dissection CMP 13–33, 17 August 1993 (UFL); 1M Brazil,
Rondônia, vic. Caucalandia, 14 October 1991 (USNM); 1F Brazil, Rondônia, Jaru, 6 August 1976 (UFL); 1F
Brazil, Amazonas, São Paulo de Olivença, 5 June 1972 (UFL); 4F Brazil, Amazonas, São Paulo de Olivença,
January–March 1923 (CMNH); 1F Brazil, Amazonas, São Paulo de Olivença June 1964 (USNM); 1F Brazil,
Amazonas, Arimã, November 1922 (CMNH); 7F Brazil, Amazonas, Nova Olinda, Rio Purus, May–July 1922
(CMNH); 1F Brazil, Amazonas, Manacapuru July 1923 (CMNH); 1F Colombia, Amazonas 13 April 1986,
dissection CMP 14-08 (LACM); 1F Ecuador, Mera 15 December 1956 (likely mislabeled) (LACM); 1F Peru,
Pucallpa, 19 May 1960, dissection CMP 14–21 (USNM); 2F Peru, Junin, Satipo no date, dissection CMP 14–18
and M. Clifton slide 214 (CMNH); 2F Peru, Junin, Satipo, October 1983, and no date (USNM); 1F Peru, Huanuco,
Tingo Maria, Leonces Prado, 23 June 1982 (UFL); 1F Peru, Huanuco, Tingo Maria, 7 August 1979, dissection
CMP 13–40 (UFL). Cithaerias aurora tambopata MALE: 1M Peru, Cuzco, Kosnipata, 5 km N Picopata 1000 m,
12 November 1989, dissection CMP 13–18 (UFL); 1M Peru, Madre de Dios, Cerro Pantiacolla, Shintuya 460 m,
15 July 1980, dissection CMP 13–25 (UFL); 1M Peru, Tambopata 18 June 1985 (UFL); 7M Peru, Puerto
Maldonado, various dates in 2004 and 2005 plus 25 April 2005, dissection CMP 13–20 and 13 October 2004,
dissection CMP 13–27 (PJD); 3M Peru, Madre de Dios, Manu, Pakitza, 22, 26, 28 April 1991 (USNM); 1M Peru,
Rio Madre de Dios 10 May 1975 (USNM);. FEMALE: 1F Peru, Madre de Dios, Cerro Pantiacolla, Shintuya 460
m, 25 June 1980, dissection CMP 13–34 (UFL); 1F Peru, Puerto Maldonado, Tambopata Reserve, 22 October 1983
(UFL); 4F Peru, Puerto Maldonado, various dates in 2004 and 2005 plus 13 August 2005 dissection CMP 14-02
(PJD). 1F Peru, Cuzco, Pompawaxo (sic) 1000 m, April 1997 (UFL); 1F Peru, Madre de Dios, Cerro Pantiacolla,
Shintuya 460 m, 20 July 1980 (UFL); 1F Peru, Madre de Dios, Manu, Pakitza, 4 May 1991 (USNM); 1F Peru, Rio
Madre de Dios 16 May 1975 (USNM).
Cithaerias aurorina (Weymer) STAT. REV.
(Figs 5f, 7a, 9)
Callitaera merolina Zikan
Type material: Lectotype is in the Zoologisches Museum, Berlin, Germany http://www.butterfliesofamerica.com/
L/ih/n_cithaerias0027_i.htm (last accessed 22 June 2014).
Distribution. Colombia, east of the Andes, and possibly the neighboring area in Brazil. This species might be
sympatric with both C. cliftoni and C. aurora. See map (Fig. 9) for examined specimen.
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FIGURE 5. ae, C. aurora; f, C. aurorina; gi, C. pyritosa; dorsal view on the left, ventral on the right. Collecting localities
and genitalia dissection codes appear below each specimen.
Diagnosis. A single male C. aurorina was examined, which precludes a sound diagnosis. Nonetheless, it
matches the photograph in Constantino (1995:73 Fig. 13) in which (1) the postmedial and submarginal bands are
slightly arched within the cells, and (2) HW rose scaling is quite diffuse and extended through the discal cell
(numbered in Fig. 5f), a character that is also present in C. cliftoni and C. aurora specimens (variable in both).
Male genitalia (Fig. 7a): note the large genitalia capsule; in lateral and ventral views, the valva apex is narrower
and more elongate than other rose colored species; in dorsal view the tegument plus uncus narrows posteriorly; and
in ventral view the rounded shape of the weakly sclerotized subscaphium bears small spines. Females were not
available for examination.
Subspecies. None.
Remarks. The specimen in Fig. 5f, and that illustrated by Constantino (1995:73 Fig. 13) resemble the
lectotype of C. merolina more closely than the lectotype of C. aurorina (which has straight HW postmedial and
submarginal lines). Male genitalia (Fig. 7a) match the illustrations of C. merolina by Zikán (1942:275, Fig. 29) and
C. pireta aurorina by Constantino (1995:75, Fig. 44). Although the identification of this taxon is consistent
between Constantino (1995) and this study, non-destructive examination of the C. aurorina lectotype genitalia may
provide conclusive identification (its abdomen seems intact). It might also provide evidence that C. merolina is a
synonym of C. aurorina. See Historical Information, Constantino (1995) and Lamas (1998) for additional details.
Material examined: MALE: 1M Colombia, Amazonas, Rio Loreto Yacu, 11 April 1983, dissection CMP 13-
13 (UFL).
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FIGURE 6. Male genitalia in lateral and ventral views, tegument and uncus in dorsal view, and subscaphium in ventral view.
Valva setae shown on one side only. When not clearly visible in the lateral view illustration, the phallus was shown separately.
The position of the phallus varies between illustrations, which does not reflect true variation in length. Given that the two parts
of the subscaphium (subanal plate and adjacent sclerotized area) rest on the membranous diaphragma, the variation in the shape
of the subanal plate includes preparation artifacts (compare ce which belong to the same species). Scale bar 1 mm for all
images. a, C. pireta dissection CMP 13-09; b, C. cliftoni dissection LGA 12-05; ce, C. aurora dissections CMP 13-16, CMP
13-19 and CMP 13-18.
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FIGURE 7. Male genitalia in lateral and ventral views, tegument and uncus in dorsal view, and subscaphium in ventral view.
Valva setae shown on one side only. When not clearly visible in the lateral view illustration, the phallus was shown separately.
Given that the two parts of the subscaphium (subanal plate and adjacent sclerotized area) rest on the membranous diaphragma,
the variation in the shape of the subanal plate includes preparation artifacts. Scale bar Scale bar 1 mm for all images. a, C.
aurorina dissection CMP 13-13; b, C. pyritosa dissection CMP 14-04.
Cithaerias pyritosa (Zikán)
(Figs 5g–i, 7b, 8e, 9)
Cithaerias similigena D’Almeida
Callitaera pireta amaryllis Bryk
Type material: Lectotype is in the Instituto Oswaldo Cruz, Rio de Janeiro, Brazil http://
www.butterfliesofamerica.com/L/ih/n_cithaerias0037_i.htm (last accessed 22 June 2014).
Distribution. Colombia east of the Andes, and Brazil, and might be sympatric with C. aurorina (see
Constantino 1995) and C. aurora. See map (Fig. 9) for examined specimens.
Diagnosis. We define this species based on the following combination of characters (numbered in Fig. 5g): (1)
male HW submarginal band well separated from marginal band, slightly staggered, usually merging with marginal
band in cell M2 to form a brown triangle (arrow in Fig. 5g); (2) male HW postmedial band that outlines the ocellus
complete across cells Rs through M2; (3) male HW postmedial band somewhat broad, slightly arched in each cell;
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(4) male HW scaling dense, deep rose with overtones of iridescent purple in some specimens, and usually localized
in cells M3, Cu1 and Cu2 (absent or barely entering cell M2); (5) female similar to male but with wider brown
bands and lighter color scaling (lacking the purple sheen), wing pattern of the single specimen available to us
seems indistinguishable from C. aurora. Male genitalia (Fig. 7b): note the extremely long phallus and
saccus—unique to this species, plus the narrow rounded subscaphium that bears well-sclerotized small spines.
Female genitalia (Fig. 8e): the rectangular sterigma is smaller and more deeply concave than in C. aurora, and it
bears an extremely large ostium bursae. Note also the enlarged antrum plus ductus bursae.
FIGURE 8. Female genitalia in ventral view, including schematic representation of papilla anales and abdomen outline. Scale
bar Scale bar 1 mm for line drawings only. a, C. pireta dissection CMP 13–28, the arrow indicates lateral portion of the
sterigma that is much reduced (and not illustrated) in other species; bc, C. cliftoni dissections CMP 13–35 and 13–29; d, C.
aurora dissection CMP 13–33; e, C. pyritosa dissection CMP 14–20. f, characteristic aspect of a mated female; note that the
scaling anterior to the sterigma have been lifted by the mating plug.
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FIGURE 9. Map showing the collecting localities of examined specimens (excluding dubious localities).
Variation in wing pattern. Although few specimens were available to us, some variation was observed. In
males the coloration can be pure rose (Brazil) or have overtones of purple (Colombia), and the brown scale-less
triangle below M2 may be solid (brown bands amalgamated; Fig. 5g) or open (brown bands not amalgamated; Fig
5h).
Subspecies. None.
Remarks. Although it is not possible to evaluate the length of the phallus and saccus from the original
description of C. pyritosa, based on type specimen photographs and localities, we concur with Lamas (1998) that
C. similigena and C. pireta amaryllis are synonyms of C. pyritosa. The male genitalia photograph of C. similigena
in D’Almeida (1951) is quite similar to C. pyritosa in Fig. 7b, but the phallus is proportionately longer. It is also
important to note that the proportions of the male genitalia parts in lateral view illustrated by Constantino (1995) do
not match those in D’Almeida (1951; as C. similigena) nor this study (Fig. 7b). This variation warrants further
examination.
Material examined: MALE: 1M Brazil, Amazonas, Tonantins, August 1923 M. Clifton dissection slide 239
(CMNH); 1M Colombia, Amazonas, Rio Tacana, 1–13 November 1946 (CMNH); 1M Colombia, Amazonas, Rio
Loreto Yacu, 8 July–16 August 1981, dissection CMP 14-04 (LACM); 2M Colombia, Leticia, 19 November 1975
(USNM). FEMALE: 1F Colombia, Amazonas, Rio Tacana, 1–13 November 1946, dissection CMP 14-20
(CMNH); 1F Brazil, 30 June 1919 (specimen has an unusual number of eyespots, identification is tentative)
(CMNH).
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Discussion
A review of the taxonomic history of the rose colored Cithaerias revealed longstanding issues with respect to
species definitions and nomenclature that had never been resolved. In part this was due to historical reliance on
wing pattern variation rather than on the comparative genitalic differences of both sexes. Based on original
descriptions and illustrations this study provides a discussion and clarification of the nomenclature pertinent to
understanding the rose colored Cithaerias. We also documented comparative patterns of morphological divergence
of genitalia and wing patterns for both sexes. This allowed a revision of species definitions and a better
documentation of intraspecific variation in wing color in these species.
This study uncovered several intriguing aspects of wing pattern coloration. For instance, some pattern
elements of the nymphalid groundplan are expressed as melanized stripes directly on the wing membrane. To our
knowledge this has not been noted explicitly for any other nymphalid (see Nijhout 1991). The presence of rose
scaling only on the dorsal hind wing seems to suggest that the intensity of color on the dorsal surface of the wing is
more important than that on the ventral surface. Field observations in Central and South America by PJD indicate
Cithaerias individuals raise and lower their semi-opened hind wings while on the forest floor, oriented in such a
way as to direct the dorsal surfaces toward dawn or dusk sunlight. This suggests that the dorsal color of Cithaerias
may be important for sexual signaling, something that has been shown in Bicyclus anynana (Butler) (Oliver et al.
2009). Of interest is that the hindwing ocellus below M1 is expressed on both wing surfaces, suggesting that border
ocelli are more conserved than other pattern elements. Indeed, this ocellus is highly conspicuous in all species of
Cithaerias, and usually the only complex ocellus present on this wing (there are exceptions, not illustrated).
Given that Cithaerias wings are mostly transparent, it is not surprising that male and female genitalia vary
more between species than does wing color pattern. We observed that species that broadly overlap in distribution
with others (C. aurorina and C. pyritosa) diverged more strongly in genitalia morphology than those that are
allopatric (C. cliftoni and C. aurora), suggesting character displacement as a mechanism of reproductive isolation
(see Sota & Kubota 1998 for an example of character displacement in beetle genitalia). Cithaerias aurorina, C.
pyritosa and C. aurora are sympatric (Fig. 9), and these taxa show allometric differences in genitalia morphology.
While the entire male genitalic capsule of C. aurorina is larger in size (female not available), only the phallus and
saccus of C. pyritosa are disproportionately long, corresponding to a long female antrum and ductus bursa. A
separate study will investigate the morphological divergence in Cithaerias genitalia (Penz in prep.).
Although we view species as evolving biological entities that vary in space and time, taxonomic nomenclature
and type specimens are static, and yet may not represent the most common phenotype within a given taxon. For
example, the type specimen of C. cliftoni (see Historical Information item (12)) has a relatively uncommon color
pattern within the range of variation of this species (Fig. 3), therefore poorly illustrating the biological entity as a
whole. The same could be said for the type of C. aurora (see variation in Figs 4 and 5a–e). The fact that type
specimens are preferably kept intact is also problematical for Cithaerias taxonomy, given the importance of
genitalia morphology for species definitions. In fact, some females of C. cliftoni and C. aurora have identical wing
color and can only be distinguished by the shape of their sterigma. The lack of detailed collecting locality also led
to much confusion in the taxonomy of the rose colored Cithaerias, and this is unfortunately a common problem of
most early species descriptions. Finally, preliminary examination of the blue and purple/rose species uncovered
morphological variation within species that parallels what has been described here for rose colored taxa (Penz,
work in progress). Lowland rain forest Cithaerias butterflies clearly display morphological attributes that are tied
to interesting behaviors, such as elongate forewings adapted to gliding flight (Cespedes, Penz & DeVries, work in
progress), and pre-mating reproductive isolation through genitalia divergence. The taxonomic issues resolved here
can be considered the kernel of what is to come next.
Acknowledgments
For specimen loans, we thank Andrew Warren (McGuire Center for Lepidoptera and Biodiversity, University of
Florida), John Rawlins (Carnegie Museum of Natural History), Brian Harris (United States National Museum,
Smithsonian Institution), and Brian Brown (Natural History Museum of Los Angeles County). For comments on
the manuscript, we thank Mirna Casagrande (Universidade Federal do Paraná), Olaf Mielke (Universidade Federal
Zootaxa 3873 (5) © 2014 Magnolia Press
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559
CITHAERIAS BUTTERFLIES
do Paraná), and Javier Munera. This study is dedicated to our friend and colleague Isidro Chacón for his love of
natural history and many contributions to the study of butterflies.
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... As a group, the eleven species included in the genus range from Central America to Amazonia, but do not occur in the Brazilian Atlantic Forest. While some Cithaerias species are remarkably similar and a positive identification requires examination of genitalia (see Penz et al. 2014), others can be confidently separated based on hind wing colors. The study by Penz et al. (2014) divided Cithaerias into three groups based on hind wing color similarity (blue, purple-rose, and rose-colored species). ...
... While some Cithaerias species are remarkably similar and a positive identification requires examination of genitalia (see Penz et al. 2014), others can be confidently separated based on hind wing colors. The study by Penz et al. (2014) divided Cithaerias into three groups based on hind wing color similarity (blue, purple-rose, and rose-colored species). Although this approach was convenient for comparing similar rose-colored species that have been frequently confused or misidentified, a DNA-based analysis showed that rose-colored taxa do not form a monophyletic group (Matos-Maraví et al. 2019)-C. ...
... Methodology for specimen examination and format of species diagnoses follow Penz et al. (2014). To study genitalia, abdomens were soaked in warm 5% KOH solution until soft. ...
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... Taxonomic work on this charismatic group involves numerous researchers over the last 150 years (e.g. Herrich-Schäffer, 1864;Weymer, 1910;Miller, 1968;Constantino, 1995;Lamas, 1997;Penz et al., 2014;Paluch et al., 2015;Willmott, 2015;Zacca et al., 2016). Haeterini is a monophyletic tribe (Wahlberg et al., 2009;Chazot et al., 2019), and current taxonomic understanding (Lamas, 2004;Penz et al., 2014;Willmott, 2015;Zacca et al., 2016) is that Haeterini consists of 29 described species and 39 subspecies. ...
... Herrich-Schäffer, 1864;Weymer, 1910;Miller, 1968;Constantino, 1995;Lamas, 1997;Penz et al., 2014;Paluch et al., 2015;Willmott, 2015;Zacca et al., 2016). Haeterini is a monophyletic tribe (Wahlberg et al., 2009;Chazot et al., 2019), and current taxonomic understanding (Lamas, 2004;Penz et al., 2014;Willmott, 2015;Zacca et al., 2016) is that Haeterini consists of 29 described species and 39 subspecies. However, these estimates remain contentious because published taxonomic studies have focused mostly on smaller groups within Haeterini, have relied on different morphological character systems (wing coloration, genitalia shape or male androconial organs), and have not taken into account variation in DNA sequence data. ...
... Haeterini butterflies were collected by the authors and collaborators throughout most of the geographical range of the tribe, including localities from northern Costa Rica to southeastern Brazil. Specimens were identified to the species and subspecies level following published taxonomic revisions (Constantino, 1995;Lamas, 1997;Penz et al., 2014;Paluch et al., 2015;Willmott, 2015;Zacca et al., 2016) and comparing wing morphology with type specimen photographs at https://www .butterfliesofamerica.com (accessed August 2018). We sampled all five Haeterini genera, including 18 out of 29 currently valid species, and 18 out of 39 subspecies (Lamas, 2004;Penz et al., 2014;Paluch et al., 2015;Willmott, 2015;Zacca et al., 2016). ...
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... brown bands than male, forming arches in each cell; female HW scaling usually limited to postmedial area but sometimes reaching the 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.. (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 diff ...
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... www.neotropicalbutterflies.com) and the taxonomic literature [51][52][53][54][55]. We used taxonomic references to confirm the current taxonomy of cryptic butterflies [56][57][58][59][60][61][62][63] and all butterfly identifications were verified by an expert Figure 1. Location of the study and sampling points in the Uauaçu Lake region, state of Amazonas, Brazil. ...
... www.neotropicalbutterflies.com) and the taxonomic literature [51][52][53][54][55]. We used taxonomic references to confirm the current taxonomy of cryptic butterflies [56][57][58][59][60][61][62][63] and all butterfly identifications were verified by an expert taxonomist. All collected butterflies were deposited in the Entomological Collection of the National Institute of Amazonian Research (INPA) and some individuals of the most abundant species were also deposited in the Zoological Collection of the Federal University of Amazonas (UFAM) and at the National Museum (UFRJ). ...
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While the references in the following chapters of these volumes by necessity will be focussed on twentieth-century works, a brief outline is given below of principal pre-1900 contributions to the foundation of contemporary lepidopterology (as covered in the present Handbook). © 1998 by Walter de Gruyter GmbH & Co., Berlin. All right reserved.
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We demonstrate experimentally that differences in genital characters impose a direct cost of interspecific copulation on two closely related carabid species, Carabus (Ohomopterus) maiyasanus and C. (O.) iwawakianus, that share a narrow hybrid zone. Males of both species attempted copulation indiscriminately between conspecific and heterospecific females. Females experiencing heterospecific mating often suffered mortality due to rupture of their vaginal membranes. Those without fatal injury laid eggs which developed into Fl adults, but the fertilization rate was much lower than for intraspecific pairs. Males of C. maiyasanus, but not C. iwawakianus, often had broken genital parts (copulatory pieces) following interspecific copulations, which may prevent normal copulation in subsequent matings. Because of female mortality and low fertilization rate, the estimated fitness cost of interspecific mating was very large in terms of the reduction in the number of offspring thatching larvae) for both sexes and both species. Thus, genital lock-and-key appears to exert significant selection against hybridization in the hybrid zone of these carabid beetles.
  • Zelaya Nicaragua
  • Dept
Nicaragua, Zelaya Dept. Nueva Guinea, 27 October 1976 (UFL);
  • Madden Panama
  • Dam
2M Panama, Madden Dam, 19 and 22 April 1966 (UFL);