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 Author Kir’Yanov, Alexander V. text Zootaxa 2019 2019-12-10 4706 3 408 426 journal article 24673 10.11646/zootaxa.4706.3.2 5bf16196-176c-4d6c-af82-7121d4bc968e 1175-5326 3568825 B42DC76D-0C0F-4EFD-B573-D14C383961DF 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 ” originates from the Quechua language, “ ccoto ” meaning “ stars ” (another translation — “ home ”) and “ huasi ” — “ meeting ”. 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 “ limited 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 forewings 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 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 hindwings 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. 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. 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. FIGURE 14. Type specimens, of C. misti ccota , males. (a), ♂ holotype; (b–d), ♂ paratypes; on left—dorsal view, on rightventral view. Cotahuasi canyon, Central Peru, 2800–3100 m. 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. 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 region. 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 9 th 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 convex 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 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 narrower 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 distally, 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 (stronger 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, 9 th and 10 th segments are fused, 9 th segment broader than 10 th 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. 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.