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https://doi.org/10.11646/zootaxa.4706.3.2
http://zoobank.org/urn:lsid:zoobank.org:pub:B42DC76D-0C0F-4EFD-B573-D14C383961DF
408 Accepted by J. De Prins: 29 Oct. 2019; published: 10 Dec. 2019
Article ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Zootaxa 4706 (3): 408–426
https://www.mapress.com/j/zt/
Copyright © 2019 Magnolia Press
Species status of Colias misti (Pieridae: Coliadinae) from the Arequipa region,
South Peru, and a description of its new subspecies, from Cotahuasi canyon,
South-Central Peru
ALEXANDER V. KIR’YANOV
Centro de Investigaciones en Optica,Loma del Bosque 115, Col. Lomas del Campestre, Leon 37150, GTO., Mexico
E-mails: alejandrokir@gmail.com; kiryanov@cio.mx
Abstract
A new species of the genus Colias Fabricius from the Arequipa region of Peru, C. misti Kir’yanov sp. nov., stat. nov., is
established by raising it from the subspecies rank, C. lesbia misti Kir’yanov 2017. This taxonomic act is justified after
detailed comparison of C. misti with representatives of phenotypically similar other South American Colias, including
the subspecies of C. lesbia. The newly presented taxa are diagnosed by adult morphology, bionomics, and male genitalia.
The diagnosis of a new subspecies Colias misti ccota Kir’yanov ssp. nov., discovered in the World’s deepest canyon
Cotahuasi (Peru), is based on different and stable characters in phenotypes, genitalia, and bionomics of C. misti ccota and
C. misti misti. Both subspecies of C. misti are endemic to the Western slopes of the Andes. It is also demonstrated that the
shape of the male aedeagus is diagnostic for a reliable identification of the South American Colias.
Key words: bionomics, diagnosis, genitalia, morphology, taxonomy
Resumen
Colias misti Kir’yanov sp. nov., stat. nov. del género Colias Fabricius de la región de Arequipa, Perú, es establecida
mediante el ascenso de categoría taxonómica de la subespecie C. lesbia misti Kir’yanov 2017. Este acto taxonómico
se justifica luego de una detallada comparación entre C. misti con otros representantes fenotipicamente similares de
Colias de Sudamérica, incluyendo las demás subespecies de C. lesbia. La diagnosis del nuevo taxón es presentada, en
términos de morfología del adulto, bionomía y la genitalia masculina. Además, se describe una nueva subespecie de C.
misti proveniente del cañón de Cotahuasi (Peru), el más profundo del Mundo, Colias misti ccota Kir’yanov ssp. nov.;
esta subespecie se basa en rasgos notablemente diferentes y estables de C. misti ccota y C. misti misti y la similitud de los
genitales y las preferencias ecológicas, inherentes a estas dos entidades. Ambas subespecies de C. misti son endémicas
de las laderas Occidentales de los Andes. También se demuestra que la forma del edeago del macho es un carácter
diagnóstico para realizar una identificación confiable de las Colias de Sudamérica.
Palabras clave: bionomía, diagnosis, genitalia, morfología, taxonomía
Introduction
In South America, the genus Colias contains only a few valid species of the C. flaveola Blanchard, 1852 group
with the ground coloration of wings in both sexes white to black and another group of species that can be treated
together due to the orange ground color of wings in males: nominotypical C. dimera Doubleday, 1847, C. lesbia
(Fabricius, 1775) (five or six subspecies), C. vauthierii Guérin-Méneville, [1830] (two subspecies), C. euxanthe
Felder & Felder 1865 (four or five subspecies) (Berger 1986; Shapiro 1991; Verhulst 2000a, 2000b, 2013; Lamas
2004; Grieshuber et al. 2012; Hammond & McCorkle 2018) and nominotypical C. alticola Godman & Salvin 1891
which is recently restored as a bona species based on molecular data (Kir’yanov 2018). The C. flaveola group is
split by some authors (Berger 1986; Shapiro 1985, 1991; Verhulst 2000a, 2000b, 2013, 2016; Benyamini et al. 2014;
VERIFICATION OF SPECIES STATUS OF COLIAS MISTI FROM PERU Zootaxa 4706 (3) © 2019 Magnolia Press · 409
Kir’yanov 2017b, Kir’yanov & Valencia 2018) into the following taxa: flaveola Blanchard, 1852, mossi Rothschild,
1913, C. weberbaueri Strand, 1912, nigerrima Fassl, 1915, blameyi Jörgensen, 1916, mendozina Breyer, 1939,
erika Lamas, 1981, benyaminii Verhulst, 2016, cora Kir’yanov, 2017, laura Kir’yanov, 2017, and harai Kir’yanov
& Valencia, 2018. Most South American Colias species are distributed along the Andes (with C. lesbia also occur-
ring in the plains of the Central and Eastern parts of the continent) whilst C. vauthierii is spread into the lowlands
of Patagonia and Tierra del Fuego.
The present study is focused on the Colias species, phenotypically belonging to C. lesbia complex, of which
wings of males are always orange whereas wings of females are either orange or whitish (alba form). Among
the valid taxa, currently assigned to C. lesbia complex (Berger 1986; Verhulst 2000a, 2000b, 2013; Lamas 2004;
Worthy 2005; Grieshuber et al. 2012), the following subspecies are recognized: pyrrhothea Hübner, 1823, dinora
Kirby, 1881, andina Staudinger, 1894, mineira Zikán, 1940, verhulsti Berger, 1983, roberti Salazar, 2002 and misti
Kir’yanov, 2017 (Figs. 1, 2).
The purpose of this article is to raise misti Kir’yanov, 2017 to species rank and to describe a new subspecies
within the C. misti.
Material and Methods
Three closely related species of Colias were collected during the same season (March–April: 2013–2017) in the
Arequipa region (Fig. 3): a very rare C. lesbia andina of which only two figured specimens were caught during five
years of fieldwork, locally common C. euxanthe hermina and C. misti. The biotope of both C. lesbia andina and
C. flaveola erika is shown in Fig. 4 and the biotope of Colias misti is shown in Fig. 5, where it may be locally and
seasonally abundant. There is a gap of less than 30 km in a direct line between the type locality of C. misti and the
area where C. lesbia and C. euxanthe occur.
During the recent years, the author has created a set of genitalia vouchers for all of the South American Colias
species. They were prepared using a technique developed at the Shovkun Dissection Laboratories (ShDL, Russian
Federation) by a standardized procedure as follows: the genitalia were dissected after submerging abdomens in 10%
KOH for approximately 48 hours. Particularly, for male specimens, the aedeagus was removed from the capsule and
photographed prior to everting the vesica with a syringe of 99% isopropyl alcohol. All parts of the aedeagus were
stained using eosin and slide-mounted with Euparal on glass slides. The genitalia vouchers fixed on the slides were
photographed using a Bresser MikroCam 10.0MP camera and processed with Adobe Photoshop Express (a free ap-
plication for Androids). For the vouchers shown, frontal view was adopted to illustrate complete female genitalia
and, also, valva and pseudouncus of male genitalia, whereas lateral view was adopted to illustrate aedeagus (Figs.
6, 7, and 9).
TABLE 1. Taxa, male genitalia of which are presented in this study along with localities.
C. dimera
(Central Ecuador, Cotopaxi)
C. vauthierii cunninghamii
(South Chile, Torres del Paine)
C. euxanthe euxanthe
(North-East Peru, Amazonas)
C. lesbia dinora ( =meieri =roberti)
(Central Ecuador, Cotopaxi)
C. vauthierii vauthierii
(Central Chile, Temuco)
C. euxanthe stuebeli
(North Peru, Cajamarca)
C. lesbia lesbia ( =pyrrhothea)
(Central Argentina, Barreal)
C. vauthierii minuscula ( =vauthierii)
(North Chile, Portillo)
C. euxanthe coeneni ( =stuebeli)
(North-West Peru, Udima)
C. lesbia andina
(South Peru, Puno)
C. misti misti
(South Peru, Arequipa)
C. euxanthe hermina ( =puna)
(South Peru, Puno)
C. lesbia verhulsti
(South Peru, Cusco)
C. misti ccota
(South-Central Peru, Cotahuasi)
C. alticola
(Central Ecuador, Cotopaxi)
The specimens from which the male genitalia were dissected and the typical adults of the nominotypical spe-
cies of the South American Colias are illustrated in Figs. 7, 8; see also Table 1. The adults from which the genitalia
vouchers are prepared were caught as close as possible to their type localities. At least two to three specimens of
each subspecies of C. lesbia complex were examined.
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FIGURE 1. Adults of the subspecies of the Colias lesbia / Colias misti complex from the type or nearby localities: (a), C.
lesbia lesbia ♂, Barreal, Argentina; (b), C. lesbia lesbia ♀, Barreal, Argentina; (c), C. lesbia dinora ♂, Chimborazo, Ecuador;
(d), C. lesbia dinora ♀, Chimborazo, Ecuador; (e), C. lesbia andina ♂, Puno, Peru; (f), C. lesbia andina ♀, Puno, Peru; (g), C.
lesbia verhulsti ♂, Cuzco, Peru; (h), C. lesbia verhulsti ♀, Cuzco, Peru; (i), C. misti ♂, holotype, Arequipa, Peru; (j), C. misti
♀, paratype, Arequipa, Peru.
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FIGURE 2. Distribution map of the Colias lesbia / Colias misti complex. The circles mark the type localities and the dashed
ellipses the approximate occurrence areas. 1, gray for C. lesbia lesbia; 2, purple for C. lesbia andina; 3, yellow for C. lesbia
verhulsti; 4, blue for C. lesbia dinora; 5, green for C. lesbia mineira; a, red for C. misti misti; b, orange for C. misti ccota.
The genitalia vouchers are preserved at the author’s private Colias collection in Moscow, Russian Federation.
The geographical distribution of C. lesbia and C. misti is mapped in Fig. 2.
Taxonomic account
Summary of phenotypical and ecological characters, diagnosing C. misti from the members of C. lesbia
complex
(Figs. 1, 3 and 8)
—dirty-yellow or ochre coloration of the wings in males of C. misti, in all other subspecies of C. lesbia complex
the wings in males are bright orange;
—apex of forewings rounded in both sexes of C. misti, in all other subspecies of C. lesbia complex apex of the
forewing is pointed;
—costal and dorsal margins of forewings are convex in both sexes of C. misti, in all other subspecies of C.
lesbia costal and dorsal margins of forewings are straight;
—pale-yellow spots, hardly perceptible, are present in the discoidal area of the male hindwings in C. misti, in
all other subspecies of C. lesbia complex, particularly in geographically adjacent to C. misti subspecies andina and
verhulsti, these spots are brightly orange and highly perceptible on the ground coloration of the male hindwings;
—cilia in C. misti are always whitish or pale-yellowish, similar in color with the ground coloration of the
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wings, in all other subspecies of C. lesbia complex, cilia are more contrasting in tint to the ground coloration of the
wings;
—C. misti adults are on average smaller (wingspan 27–37 mm) than those belonging to the other subspecies of
C. lesbia (wingspan 33–53 mm), except verhulsti that has wingspan similar to C. misti;
—C. misti occurs at altitudes from 1800 to 3000 m, occupying habitats at lower altitudes than the Andean sub-
species of C. lesbia such as andina or verhulsti which occur at altitudes from 3400 and to 4400 m.
On the basis of morphological and ecological characters the taxon C. misti is considered as a bona species. Addition-
ally, differential diagnostic genitalia characters are provided to confirm the raise of C. misti to species rank.
FIGURE 3. Adults. (a), C. lesbia andina ♂; (b), C. lesbia andina ♀; (c), C. euxanthe hermina ♂; (d), C. euxanthe hermina ♀;
(e), C. misti misti ♂ holotype; (f) C. misti misti ♀, paratype. Collecting locality—vicinities of Arequipa city, Dept. Arequipa,
South Peru.
Summary of genital morphological characters, diagnosing C. misti from the members of C. lesbia com-
plex and other related Colias species
(Figs. 6, 7 and 9)
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No irregularities or deviations were found in genitalia structures of the taxa studied. It was concluded that the mor-
phological characters of aedeagus are diagnostic. The morphological parts of male genitalia in Colias species are of
different diagnostic significance: valva and pseudouncus in Colias are usually of minor value for species identifica-
tion, though the shape of the pseudouncus in most of the cases can serve as a reference point to group species; on
the contrary, aedeagus shows pronounced and stable diagnostic differences (Figs. 6e–h, 9):
FIGURE 4. Biotope of C. euxanthe hermina, C. lesbia andina and C. flaveola erika: Pampa Cañahuas, Dept. Arequipa, Peru,
4000–4100 m with Astragalus sp. plants.
FIGURE 5. Biotope of C. misti misti ~10 km South from El Misti volcano, Dept. Arequipa, Peru, 2700–2800 m with Medicago
sativa plants, abundant in the area (foreground).
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FIGURE 6. Female genitalia a–d (frontal view): (a), C. lesbia andina; (b), C. lesbia verhulsti; (c), C. misti misti; (d), C. misti
ccota. Male genitalia e–h: (e), C. lesbia andina; (f), C. lesbia verhulsti; (g), C. misti misti; (h), C. misti ccota. 1st line, valva,
frontal view; 2nd line, pseudouncus, frontal view; 3rd line, aedeagi, lateral view; 4th line, aedeagi tips (red-framed).
—pseudouncus is trapezoidal in C. misti, it is more triangular in C. lesbia;
—cornuti on both sides of the tip of the aedeagus (highlighted in red color in Figs. 6e–h, 9) in all examined
specimens of C. lesbia are present but they are absent in the other related Colias species;
—the tip of the aedeagus in C. misti as well as in C. vauthierii is notably curved downwards, in all subspecies
of C. lesbia is slightly curved upwards;
—cornutus on the tip of the aedeagus is obscure or absent in C. misti, a single small cornutus on the tip of the
aedeagus is present in C. vauthierii;
—the tip of the aedeagus in C. vauthierii and C. misti is with rough edges, in all subspecies of C. euxanthe it is
always smooth;
—the girth of the aedeagus in C. misti is similar to that in C. lesbia, in C. euxanthe the aedeagus is very thin and
never bears cornuti along its surface.
The female genitalia characters in Colias vary slightly and hence serve poorly for the identification purposes;
nonetheless we briefly diagnose some of their details.
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FIGURE 7. Aedeagi, lateral view. (a), C. dimera; (b), C. lesbia dinora; (c), C. lesbia lesbia; (d), C. lesbia andina; (e), C. lesbia
verhulsti; (f), C. vauthierii cunninghamii; (g), C. vauthierii vauthierii; (h), C. vauthierii minuscula; (i) C. misti misti, paratype;
(j) C. misti ccota, paratype; (k) C. euxanthe euxanthe; (l), C. euxanthe stuebeli; (m), C. euxanthe coeneni; (n), C. euxanthe her-
mina and (o), C. alticola.
The female genitalia of C. misti misti and C. misti ccota (see the description below) are similar to each other
(Fig. 6a–d) whilst differ but slightly from those of C. lesbia andina and C. lesbia verhulsti. The differences found in
the terminal abdominal segments which may serve for the diagnostic purposes are as follows:
—in both subspecies of C. misti the papillae anales almost merge, in C. lesbia andina and C. lesbia verhulsti the
papillae anales are split;
—in both subspecies of C. misti the posterior apophyses are with straight margins, in C. lesbia andina and C.
lesbia verhulsti the posterior apophyses are curved (especially in C. lesbia verhulsti).
Thus, the pattern of the genitalia examined, especially the morphology of the aedeagus in C. misti, compared
with the aedeagi of all related orange South American Colias species (Fig. 7), confirms the raise of C. misti to
species status. The details of the aedeagus terminal part, allowing to delimit C. misti as a bona species from the
geographically adjacent species C. lesbia, C. euxanthe and C. vauthierii, are presented in Fig. 9. The knowledge
about the genital morphology of South American Colias species was poor so far (Berger 1986; Petersen 1963), so
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the diagnostic characters, presented above for the first time, may be useful for further studies on this peculiar species
group of the genus.
FIGURE 8. Male adults of South American orange Colias species (nominative subspecies).
FIGURE 9. Diagnostic features (red-framed) of the aedeagi tips, for Colias lesbia, C. vauthierii, C. misti and C. euxanthe.
VERIFICATION OF SPECIES STATUS OF COLIAS MISTI FROM PERU Zootaxa 4706 (3) © 2019 Magnolia Press · 417
Remark. According to ICZN (Art. 46.2), when a nominal taxon is raised in rank in the species group its
name-bearing type remains the same. Thus, the type specimens of C. misti stay the same as they were indicated in
(Kir’yanov 2017a).
Description of a new subspecies of Colias misti
When visiting the World’s deepest canyon, Cotahuasi (Central Peru) (Fig. 10), in April 2016 and April 2017, it
was found and explored in more detail a place where the specimens phenotypically similar to Colias misti fly. The
observations led to the description of a new subspecies presented below.
It was established that the butterflies belonging to the new subspecies inhabit small patches inside the canyon
at altitudes of 2800–3100 m with abundant vegetation (Fig. 11). It is the sole Colias taxon there, probably being
endemic, as outside the canyon, at higher altitudes (3900–4500 m) with landscapes of the puna type (Fig. 12) and
very different weather conditions, this newly described Colias was never encountered. Instead, three other clouded
yellows were recorded in the puna biotope: the orange C. lesbia andina and C. euxanthe hermina (Fig. 13) along
with the gray and white subspecies of C. flaveola (Kir’yanov 2017b).
The econiche of the new subspecies of C. misti is separated by parapatry from the other Colias species, occur-
ring in this region. The distribution area of the new subspecies is in lower altitude than puna, the biotope of phe-
notypically related Colias species. Therefore, together with differences in morphological characters, the different
bionomics supports the need of the description of a new subspecies within C. misti.
C. misti ccota Kir’yanov ssp. nov.
(Fig. 14, Fig. 15)
Type material. Holotype (HT) ♂, (~5–10 km from Cotahuasi village, Cotahuasi canyon, ~30–40 km North-West
from Coropuna volcano, Dept. Arequipa, Peru, 2800–3100 m), 14–16/IV/2016, A. Kir‘yanov leg. &, coll. (The
State Darwin Museum, Moscow, Russia). Paratypes (PT) 11 ♂♂, 7 ♀♀, 14–16/IV/2016, 12–16/IV/2017, the same
locality and collector, (the collection of A. Kir’yanov, Moscow, Russia; El Museo de Historia Natural, Universidad
de San Marcos, Lima, Peru).
Remark. For a rectification of the current status of the two specimens (male and female), tentatively attributed
as “Colias lesbia verhulsti(?)” in the earlier published literature sources, see Table 2.
TABLE 2. Verified type specimens and their illustrations.
Taxon name Reference and illustration Identification and illustration in present article
“Colias lesbia verhulsti(?)” Kir’yanov 2017a: Fig. 5a(2
—upper specimen) ♀
Colias misti ccota: Fig. 15(b) PT♀
“Colias lesbia verhulsti(?)” Kir’yanov 2017a: Fig. 5a(2
—lower specimen) ♂
Colias misti ccota: Fig. 16(left specimen) PT♂
“Colias lesbia verhulsti(?)” Kir’yanov 2017b: Fig. 17b ♀ Colias misti ccota: Fig. 15(b) PT♀
Etymology. The name stems from the name of its type locality, Cotahuasi canyon. The word “Cotahuasi” origi-
nates from the Quechua language, “ccoto” meaning “stars” (another translation — “home”) and “huasi” — “meet-
ing”. Given the pronounced sheen that males of C. misti ccota demonstrate (Fig. 14), they may be associated with
shining stars; another association is that this Colias taxon is endemic to Cotahuasi canyon. For Spanish speaking
people, “ccota” may be associated with the word “cota” that may, in turn, be referred to as “acotado” meaning “lim-
ited territory”. So, the name “ccota” also points to the endemism of the subspecies C. misti ccota, which is strictly
limited to Cotahuasi canyon, Peru.
Differential diagnosis. C. misti ccota is a small (on average smaller than C. misti misti) butterfly with the fore-
wings more elongated at the apex than those of C. misti misti; this morphological character is more evident in ccota
females than in males. Although the forewings in both subspecies, in both males and females, are convex in termen,
in C. misti ccota the convex shape more evident at the apex whilst in C. misti misti the convex shape is more evident
at the anal angle (Figs. 14, 15).
Whereas in males of C. misti ccota both the fore- and hindwings are deeply orange, with a slight yellowish
tinge, the fore- and hindwings in males of C. misti misti are dirty-yellow or ochre in coloration and matt. The discal
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FIGURE 10. Map of Cotahuasi canyon and surroundings, including the highest peaks: volcanos Coropuna (6425 m), Solimana
(6093 m) and Firura (5498 m). The type locality of C. misti ccota is shown by the orange square.
FIGURE 11. Biotope of C. misti ccota (Cotahuasi canyon, ~30 km North-West from Coropuna volcano, Dept. Arequipa, Peru,
2800-3100 m), with abundant clovers and alfalfa plants.
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FIGURE 12. Biotope of C. euxanthe hermina, C. lesbia andina and C. flaveola laura (Lauripampa near glacier Firura, northern
part of Cotahuasi canyon, Dept. Arequipa, Peru, 3900–4200 m).
FIGURE 13. Adults. (a), C. lesbia andina ♂; (b), C. euxanthe hermina ♂; on left—dorsal view, on right—ventral view. Vicinity
of glacier Firura, northern part of Cotahuasi canyon, Central Peru, 4000–4200 m.
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FIGURE 14. Type specimens, of C. misti ccota, males. (a), ♂ holotype; (b–d), ♂ paratypes; on left—dorsal view, on right—
ventral view. Cotahuasi canyon, Central Peru, 2800–3100 m.
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FIGURE 15. Type specimens, of C. misti ccota, females. (a–d), ♀ paratypes; on left—dorsal view, on right—ventral view.
Cotahuasi canyon, Central Peru, 2800–3100 m.
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spots on the forewings are more developed in C. misti ccota than in C. misti misti. The discal spots in C. misti ccota
on the hindwings are well perceptible against the ground color, in C. misti misti—barely perceptible. The ventral
surface of both the fore- and hindwings is bright yellow in C. misti ccota, it is much brighter than in C. misti misti.
The row of submarginal spots is more evident in C. misti ccota than in C. misti misti. The discal spots on the hind-
wings are superimposed with bars, rather short and heavily dusted with dark scales in C. misti ccota, in C. misti misti
the bars of discal spots on the hindwings are longer but less developed. Cilia are yellow in C. misti ccota, in C. misti
misti cilia are white or yellowish.
In female of C. misti ccota the wings are bicolor, in C. misti misti the coloration of the fore- and hindwings is
unicolored. Females of C. misti ccota are always of alba form, females of C. misti misti can be of yellow-orange
form though they are extremely rare. In C. misti ccota the costal area of the forewings is strongly suffused with
black scales, being in some specimens completely black, in C. misti misti, this area is lighter. The discal spots on the
forewings are more developed in C. misti ccota than in C. misti misti. The border of the forewings is broader and
more strongly suffused with black scales in C. misti ccota than in C. misti misti, bearing a series of slightly elliptical
white or yellowish spots. On the hindwings, the border is less developed in C. misti ccota. The row of submarginal
spots is less expressed than in males. The discal bars on the hindwings are less developed in C. misti ccota than in
C. misti misti. Cilia are pinkish or pink-yellowish in C. misti ccota, in C. misti misti cilia are whitish.
Male genitalia of C. misti ccota are close to those of C. misti misti (Fig. 6g, h); furthermore, in aedeagi, both
subspecies superficially resemble C. vauthierii but significantly differ from the other pale orange Colias of the re-
gion. The differences between C. misti ccota and C. misti misti is the size of aedeagus and the shape of appendix of
coecum: the aedeagus in C. misti ccota is bigger, more robust than in C. misti misti; the appendix of coecum in C.
misti ccota is straight, not forked through the length whereas in C. misti misti it is forked at the end.
Female genitalia of C. misti ccota are almost undistinguishable from those of C. misti misti (Fig. 6c, d) except
the differences in the shape of segment 9. The 9th segment is broader in C. misti ccota than in C. misti misti.
Compared with the other orange Colias of the region, C. euxanthe hermina (Fig. 13b), C. misti ccota clearly
differs from the latter by bright-yellow underside of the wings in males (Fig. 14), in males of C. euxanthe hermina
the underside of the wings is dirty-yellow or ochre. In males of C. euxanthe hermina the subdorsal area of hindwings
and the whole surface of underside of hindwings have a suffusion with dark scales, this suffusion is very weak in
males of C. misti ccota. Both the shape of wings (more elongated) and the type of coloration (bicolored) in females
of C. misti ccota (Fig. 15) demonstrate the diagnostic differences from the females of C. euxanthe hermina. The
shape of wings in C. euxanthe hermina is more or less rectangular and wings are never bicolored. Concerning the
diagnosis of males of C. misti ccota and the males of parapatric C. lesbia andina, both species differ unambiguously
(Figs. 13a, 16).
Females of C. misti ccota bear a strong suffusion with black scales of the costal margin of the forewings, this
suffusion was never observed in females of C. euxanthe hermina, as well as in females of other subspecies of C.
euxanthe (Fig. 3d).
The differential diagnosis of the genital morphological characters, delimiting C. misti from the other orange
South American Colias, is fully applicable to the subspecies C. misti ccota (Figs. 6, 7, 9).
The new subspecies occurs in the midlands on slopes of Cotahuasi canyon at altitudes varying from 2800 to
3100 m (Fig. 11), while the biotope of C. misti misti is restricted to the natural depressions at altitudes from 2000 to
3000 m, surrounded by high (3800–4300 m) mountains, not so far from Arequipa city (Fig. 5). The localities where
C. misti misti occurs are more affected by the anthropogenic factors, one of which is alfalfa planting, than the type
locality of C. misti ccota. Alfalfa is a hostplant for C. misti misti, while the host plant of C. misti ccota is one of the
Trifolium species. Preimaginal stages of C. misti ccota remain unknown.
Description. Male (Fig. 14). Wingspan 29–39 mm.
Head brown, eyes brownish-black. Antennas brown with slightly yellowish tips of clubs. Labial palpi short
(slightly longer than eye diameter), yellowish-pinkish-brown, straight.
Thorax black. Abdomen black, dorsally slightly dusted with yellow scales and barely covered with grayish
hairs; ventrally grayish-yellow. Legs pinkish. The forewings are slightly convex in costal margin and notably con-
vex in termen; dorsum straight. The hindwings are rounded, with anal angle slightly protruded. The dorsal surface
of both the fore- and hindwings is deeply orange, with a slight yellowish tinge in the hindwings (especially, in the
costal area and basal margin towards thorax). The basal area of both the fore- and hindwings is moderately suffused
with dark scales. The costal margin of the forewings is strongly suffused with black scales. The marginal area of the
VERIFICATION OF SPECIES STATUS OF COLIAS MISTI FROM PERU Zootaxa 4706 (3) © 2019 Magnolia Press · 423
forewings is also black; it is broad in the apex and steadily narrows towards the anal angle. The discal black spots
on the forewings are well developed, comma-shaped. The marginal area of the hindwings is black; it is much nar-
rower than in the forewings, fading towards the anal angle. On the hindwings, the discal spots are bright red, well
perceptible against the ground color; the spots are core-empty; they are superimposed with bars of the same color.
The ventral surface of both the fore- and hindwings is bright yellow. On the ventral side, the discal white spots on
the hindwings are superimposed with bars, rather short and heavily dusted with darkly brown scales. The row of
darkly brown submarginal spots is evident on the underside of both the fore- and hindwings; these are mirrored on
the upperside as more darkened—against the ground coloration of the wings—spots. Cilia are yellow. Androconial
patches are always present near the thorax in the basal-costal area of the hindwings.
Male genitalia (Figs. 6h, 7j). Tegumen broad. Uncus large, at terminal point hooked downwards; pseudouncus
elongated, slightly trapezoidal. Saccus barely elliptic, almost rounded. Valva strongly convex, thick, tapering dis-
tally, slightly separating on the distal and proximal parts, bears a small dentate appendix at the end. Aedeagus is long
and broad, triangular at the tip, bears tiny cornutus; appendix of coecum is parallel straight through the length, not
forking to the end.
Female. (Fig. 15). Wingspan 28–35 mm.
Head, eyes and labial palpi as in males. Antennas homogeneously brown with slight wine tinge.
Thorax and abdomen black, the latter slightly covered with grayish hairs on the dorsal side; ventrally grayish.
Legs pinkish but darker than in males. Both the fore- and hindwings are shaped similarly to those of males but are
more elongated towards apical areas. The dorsal surface of both the fore- and hindwings is light: the forewings are
always white while the hindwings—whitish but always with a yellowish or yellow-greenish tinge (only females of
the form alba have been found so far). The costal area of the forewings is strongly suffused with black scales (stron-
ger than in males), being in some specimens completely black. The discal spots on the forewings are well developed,
more rounded than in males. The termen area of the forewings is broad, strongly suffused with black scales, bearing
in submarginal area a series of big slightly elliptical white or yellowish spots. On the hindwings, the suffusion in
termen area is less developed; the discal spots on the hindwings are white, barely perceptible against the ground
color. The ventral surface of the wings is bicolor: the forewings are white or yellowish, the hindwings are heavily
suffused with yellow-greyish or green-greyish scales. The row of submarginal spots is oblique. The discal bars on
the hindwings are weakly developed. Cilia are pinkish or pink-yellowish.
FIGURE 16. Paratype specimen of C. misti ccota ♂ (on left); a specimen of C. lesbia andina ♂ (on right). Cotahuasi, Dept.
Arequipa, Peru.
Female genitalia (Fig. 6d). Papillae anales united, 9th and 10th segments are fused, 9th segment broader than 10th
one. Posterior apophyses straight. Anterior apophyses reduced to spots. Ductus bursae conic, membranous, antrum
as long as one third of ductus bursae; signum rectangular, with surface densely covered with small teeth. Corpus
bursae rounded, twice in volume compared to appendix bursae, which is also rounded.
Distribution. Currently known only from the type locality, Cotahuasi canyon, Dept. Arequipa, Central Peru
(Fig. 10). Most probably endemic.
KIR’YANOV
424 · Zootaxa 4706 (3) © 2019 Magnolia Press
Hostplant. One of the clovers (Trifolium sp.), abundant in the type locality (Fig. 11).
Bionomics. C. misti ccota occurs in the midlands on slopes of Cotahuasi canyon at altitudes varying from 2800
to 3100 m; it is observed in the meadows with abundant vegetation (Fig. 11). Till now, adults of C. misti ccota were
recorded only in April, the season when its type locality is in full blossom.
Discussion
The habitats and the flight period of three orange Colias species, C. euxanthe hermina, C. lesbia andina and C. misti
misti, merge. However, C. misti misti is found at medium altitudes, where its hostplant (Medicago sativa L.) grows,
while the other closely related species fly in significantly higher situated biotopes of puna type where at least three
species of Astragalus, their probable hostplant(s), are found. Adults of C. misti misti were never encountered in
puno (>4000 m) (Fig. 4), where C. euxanthe hermina, C. lesbia andina, and C. flaveola erika occur, and, vice versa,
there are no records of the C. euxanthe hermina, C. lesbia andina, and C. flaveola erika taxa within the distribution
area of C. misti misti, restricted to <3000 m (Fig. 5).
Thus, C. misti misti cooccurs with the other orange local Colias species, C. lesbia andina and C. euxanthe
hermina (Fig. 3), in parapatry. The other subspecies, Colias misti ccota, cohabits in parapatry with the local orange
Colias, C. lesbia andina and C. euxanthe hermina (Figs. 13, 16), the latter inhabiting the biotopes at mid and high
altitudes within Cotahuasi canyon area and the Coropuna volcano (Figs. 10, 11, 12).
Both subspecies of C. misti are endemic to the Western slopes of the Andes (Fig. 2), which are probably rem-
nants of a common ancestor with a broader distribution in the past; both the taxa inhabit the lowland “refugia” with
locally abundant vegetation, surrounded by high mountains that form natural barriers for gene flow from and to
outside the “refugia”. Furthermore, the similarity of the aedeagal morphology in males of C. misti misti and C. misti
ccota as well as of C. vauthierii (Figs. 6, 7, 9), points to their close taxonomic position. In this regard, it is interesting
to add that C. misti and C. vauthierii occur in the regions placed not far from each other (Fig. 2), with their distribu-
tion areas being separated by the lifeless Atacama desert.
The Andean Colias is a concise group of taxa which seem to be in a rapid speciation process with numerous
possible hybridization events involved, which frequently leads to difficulties in delimiting species. The cases of
sympatry of Colias species without hybridization that would justify the specific distinctiveness are rarely encoun-
tered within this group, with the exceptions of two pairs of species being C. dimera—C. lesbia in northern South
America and C. vauthierii—C. lesbia in few localities of southern South America. It is presumed that hybridization
between such Andean Colias species as C. flaveola, C. euxanthe, C. lesbia, and C. misti is a common phenomenon,
which complicates species differentiation by the molecular tools (Kir’yanov 2018). Obtaining full genome patterns
of questionable Colias species and subspecies may resolve the existing problems of their taxonomic status and sys-
tematic position.
Conclusion
Summarizing, I present here the following characteristics on new species C. misti and its new subspecies C. misti
ccota: (i) they occur in small “refugia” inside canyons or lowlands at altitudes of 2000–3000 m, feeding on Tri-
folium and Medicago sativa; (ii) the morphology of aedeagi of both taxa, C. misti misti and C. misti ccota, is a
diagnostic character reliably separating them from the rest of phenotypically similar orange Colias occurring in the
same region.
Acknowledgements
The author gratefully acknowledges the staff of ShDL (Shovkun Dissection Laboratory), and especially Kseniya
Proskuryakova (Moscow, Russia), for their efforts to prepare the genitalia slides, Jose Cerdeña (Arequipa, Peru)
for his guidance in the fieldwork and Maxim Markhasiov for fruitful discussions. The author greatly values the
comments and the linguistic suggestions of Robert Worthy (Caterham, UK) and Arthur Shapiro (Davis, CA, USA)
which enhanced the quality of the this article.
VERIFICATION OF SPECIES STATUS OF COLIAS MISTI FROM PERU Zootaxa 4706 (3) © 2019 Magnolia Press · 425
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