A new species of butterfly (Lepidoptera: Nymphalidae) from the highlands of Southeastern Brazil Author Barbosa, Eduardo P. Departamento de Biologia Animal, Instituto de Biologia, Universidade Estadual de Campinas, Campinas, São Paulo. & Museu de Zoologia “ Adão José Cardoso ”, Universidade Estadual de Campinas, Campinas, São Paulo. & edu 1482 @ gmail. com; https: // orcid. org / 0000 - 0002 - 8885 - 454 X Author Aguiar, Tamara M. C. Departamento de Biologia Animal, Instituto de Biologia, Universidade Estadual de Campinas, Campinas, São Paulo. & Museu de Zoologia “ Adão José Cardoso ”, Universidade Estadual de Campinas, Campinas, São Paulo. & Programa de Pós-Graduação em Biologia Animal, Instituto de Biologia, Universidade Estadual de Campinas & tamara. aguiarcm @ gmail. com; https: // orcid. org / 0000 - 0002 - 0711 - 8226 Author Rosa, Augusto H. B. Departamento de Biologia Animal, Instituto de Biologia, Universidade Estadual de Campinas, Campinas, São Paulo. & Museu de Zoologia “ Adão José Cardoso ”, Universidade Estadual de Campinas, Campinas, São Paulo. & Programa de Pós-Graduação em Biologia Animal, Instituto de Biologia, Universidade Estadual de Campinas & augustohbrosa @ hotmail. com; https: // orcid. org / 0000 - 0002 - 0151 - 114 X Author Zacca, Thamara Departamento de Biologia Animal, Instituto de Biologia, Universidade Estadual de Campinas, Campinas, São Paulo. & Museu de Zoologia “ Adão José Cardoso ”, Universidade Estadual de Campinas, Campinas, São Paulo. & zacca. butterfly @ gmail. com; https: // orcid. org / 0000 - 0002 - 1398 - 0381 Author Freitas, André V. L. Departamento de Biologia Animal, Instituto de Biologia, Universidade Estadual de Campinas, Campinas, São Paulo. & Museu de Zoologia “ Adão José Cardoso ”, Universidade Estadual de Campinas, Campinas, São Paulo. & baku @ unicamp. br; https: // orcid. org / 0000 - 0002 - 5763 - 4990 text Zootaxa 2020 2020-07-22 4819 1 128 142 journal article 10.11646/zootaxa.4819.1.6 1175-5326 3955941 EFA18EF4-B927-4A7F-AE77-A128EEF0923F Carminda surpresa Barbosa, Aguiar, Rosa, Zacca & Freitas , sp. nov. Figures 1–4 Type material. Holotype : Deposited in the Museu de Zoologia da Universidade Estadual de Campinas ( ZUEC ), Campinas , São Paulo , Brazil , with the following four labels separated by transverse bars : / HOLOTYPUS / Brazil , São Paulo , Campos do Jordão , Brejo Caminho para Pico do Itapeva , 22°45’S 45°33’W , 1735 m , 11.II.2018 , A. H. B. Rosa leg . / ZUEC LEP 11052 / Holotypus Carminda surpresa Barbosa, Aguiar, Rosa, Zacca & Freitas , det . 2019 /. Allotype : Deposited in the Museu de Zoologia da Universidade Estadual de Campinas ( ZUEC ), Campinas , São Paulo , Brazil . With the following four labels separated by transverse bars : / ALLOTYPUS / Brazil , São Paulo , Campos do Jordão , Brejo Caminho para Pico do Itapeva , 22°45’S 45°33’W , 1735 m , 28.I.2018 , A. H. B. Rosa & E. K. S. Ramos leg . / ZUEC LEP 11049 / Allotypus Carminda surpresa Barbosa, Aguiar, Rosa, Zacca & Freitas , det . 2019 /. Paratypes (all from Campos do Jordão , São Paulo , Brazil ). No specific locality , 1 male , 27.II.1983 , K. S. Brown Jr. leg ., ( ZUEC LEP 11059), ( ZUEC ) ; Brejo Caminho para Pico do Itapeva , 22°45’S 45°33’W , 1735 m , 2 males , 28.I.2018 , A. H. B. Rosa & E. K. S. Ramos leg ., DNA Voucher YPH–0781, ( ZUEC LEP 11047, 11048 ) ( ZUEC ) , 1 male , II.2018 , A. H. B. Rosa leg ., ex. Larva, ST5 ( ZUEC LEP 11050), ( ZUEC ) , 1 male , 02.II.2018 , E. P. Barbosa leg ., ( ZUEC LEP 11051), ( ZUEC ) , 2 males , 11.II.2018 , A. H. B. Rosa leg ., ( ZUEC LEP 11053, 11054 ), ( ZUEC ) , 1 male , 21.II.2018 , A. H. B. Rosa leg ., DNA Voucher YPH–0793, ( ZUEC LEP 11055) ( ZUEC ) , 1 male , 05.III.2018 , A. H. B. Rosa leg ., DNA Voucher YPH–0689, ( ZUEC LEP 11056) ( ZUEC ) ; 1 male , 1 female , III–V.2018, A- 358, criação, ex. Larva ( ZUEC-AVLF ) ; Novo Capivari , Mirante Ferradura , 22°43’S 45°31’W , 1745 m , 1 female , 08.II.2017 , A. H. B. Rosa leg ., (DNA Voucher YPH–0689), ( ZUEC LEP 11057), ( ZUEC ) , Novo Capivari , 22°44’S 45°32’W , 1735 m , 1 male , 14.III.2018 , A. H. B. Rosa leg ., ( ZUEC LEP 11058), ( ZUEC ) . Diagnosis. Carminda surpresa sp. nov. is superficially similar to Yphthimoides ochracea (Butler) and Yphthimoides straminea (Butler) , both sympatric and syntopic with the new taxon here described. It can be distinguished from both species of Yphthimoides by the following characters: 1) the absence of a well-defined thin stripe crossing both wings at one third distance from wing base (present in both species of Yphthimoides ); 2) the presence of a faint yellowish stripe crossing the VHW (absent in Y. straminea and more conspicuous in Y. ochracea ). Carminda surpresa sp. nov. can be easily distinguished from other species of Carminda by the following characters: 1) the ventral wings are mostly homogeneous, lacking the dark and white mottled pattern present in other species of Carminda ; 2) the ocelli on the VHW are small and incomplete, usually only three are visible (rarely four) and the pupils are occasionally absent, whereas in the other species of Carminda usually four large ocelli with double white pupils are present; 3) a faint yellowish stripe crossing the VHW in the medial area is present in some individuals. TABLE 1. Data of evaluated specimens of Carminda and outgroups with code, locality and GenBank accession numbers.

Species	DNA voucher	Locality	COI	GAPDH	RPS5
Atlanteuptychia ernestina	YPH-0191	Brazil, São Paulo, Jundiaí, Serra do Japi	KP994863	KU340904	KU340937
Paleonympha opalina	EW25-21	Taiwan, Pingtung County, Hsiaokuehu, Wutai Shiang	DQ338880	GQ865010	GQ865479
Megisto cymela	CP21-04	USA, Rhode Island, Valley Falls	GQ357208	GQ357439	GQ357569
Cissia rubricata	LEP-19315	USA, Arizona, Southwestern Research Station, Portal	MG209757	MG546314	MG546327
Cissia proba	CP01-30	Peru, Madre de Dios, Tambopata Research Center	GQ864751	GQ864945	GQ865416
Cissia pompilia	07-SRNP-100013	Costa Rica, Area de Conservacion Guanacaste, Sector San Cristobal, Estacion San Gerardo	JQ573901	—	—
Cissia pompilia	MAL-02577	Mexico, Yucatan, Valladolid, carretera a Xocen	GU659737	—	—
Cissia penelope	DNA99-028	Ecuador, Napo	AY508530	—	—
Cissia penelope	CP07-58	Peru, Junin, La Solitaria-Quebrada Siete Jeringas	GU205833	GU205945	GU206005
Cissia phronius	NW126-7	Brazil, São Paulo, Atibaia	DQ338807	GU205973	GU206034
Cissia phronius	BLU378	Brazil, São Paulo, Ribeirão Grande, Trilha do Capinzal	KU340890	—	—
Cissia eous	YPH-0422	Brazil, Rio Grande do Sul, Alegrete	KU340872	KU340920	KU340955
Cissia eous	BLU619	Brazil, Paraná, Foz do Iguaçu	KU340894	—	—
Cissia eous	YPH-0104	Brazil, São Paulo, Campinas, Unicamp, Mata da Zoologia	KU340856	KU340901	KU340934
Stegosatyrus ocelloides	YPH-0527	Brazil, Minas Gerais, Serra do Cipó	KU340879	—	KU340966
Stegosatyrus ocelloides	YPH-0530	Brazil, Minas Gerais, Serra do Cipó	KU340880	KU340925	KU340967
Stegosatyrus ocelloides	YPH-0543	Brazil, Minas Gerais, Serra do Cipó	KU340883	KU340928	KU340970
Graphita griphe	YPH-0361	Colombia, Cali, San Antonio	KU340866	KU340915	KU340948
Graphita griphe	YPH-0362	Colombia, Cali, San Antonio	KU340867	—	KU340949
Pharneuptychia phares	YPH-0465	Argentina, Santiago del Estero, Villa Punta	MN264790	MN264901	MN265039
Moneuptychia giffordi	YPH-0480	Brazil, Minas Gerais, Belo Horizonte, Pq. Serra Rola Moça	KP890215	KU340924	KU340959
Moneuptychia itapeva	YPH-0316	Brazil, Minas Gerais, Nova Lima	KP890216	MN264907	MN265046
Moneuptychia soter	YPH-0312	Brazil, São Paulo, Jundiaí, Serra do Japi	KP890212	KU340909	KU340942
Yphthimoides yphthima	YPH-0003	Brazil, Minas Gerais, Tiradentes	KT880198	KT880208	KT880213
Yphthimoides pacta	YPH-0302	Brazil, Distrito Federal, Brasília	KT736112	MN264954	MN265094
Yphthimoides straminea	YPH-0015	Brazil, Minas Gerais, Serra do Caraça	KU525703	MN264963	MN311532
Yphthimoides patricia	YPH-0282	Brazil, Minas Gerais, Santana do Riacho, Serra do Cipó	KT736117	MT036326	MT036337
Carminda griseldis	YPH-0394	Brazil, São Paulo, Alto do Capivari, Campos do JordaÞo	KU340869	KU340917	KU340952

......continued on the next page TABLE 1. (Continued)

Species	DNA voucher	Locality	COI	GAPDH	RPS5
Carminda griseldis	NW127-17	Brazil, Mato Grosso, Extrema	GU205853	GU205965	GU206025
Carminda griseldis	YPH-0462	Brazil, Espírito Santo, Caparaó	KU340875	KU340923	KU340958
Carminda griseldis	YPH-0785	Brazil, São Paulo, Pindamonhangaba	MT036323	MT036335	—
Carminda griseldis	YPH-0814	Brazil, São Paulo, Campos do Jordão	MT036324	MT036336	MT036343
Carminda griseldis	YPH-0857	Brazil, São Paulo, Campos do Jordão	MT036325	—	—
Carminda paeon	B-17-41	Brazil, São Paulo	DQ338801	—	—
Carminda paeon	NW126-11	Brazil, São Paulo, Ribeirão das Pedras	GQ864792	GQ865004	GQ865473
Carminda paeon	BLU324	Brazil, São Paulo, Jundiaí, Serra do Japi	MG209740	—	—
Carminda paeon	BLU325	Brazil, São Paulo, Jundiaí, Serra do Japi	MG209741	—	—
Carminda paeon	BLU326	Brazil, São Paulo, Jundiaí, Serra do Japi	MG209742	—	—
Carminda paeon	YPH-0357	Brazil, Paraná, Foz do Iguaçu, Trilha Poço Preto	MG209761	MG546318	MG546323
Carminda paeon	YPH-0778	Brazil, São Paulo, Pindamonhangaba	MT036322	MT036334	—
Carminda umuarama	YPH-0437	Brazil, Minas Gerais, Pq. Itatiaia, Itamonte	KU340873	KU340921	KU340956
Carminda umuarama	YPH-0459	Brazil, Espírito Santo, Caparaó	KU340874	KU340922	KU340957
Carminda umuarama	YPH-0782	Brazil, São Paulo, Pindamonhangaba	MT036318	MT036331	—
Carminda umuarama	YPH-0833	Brazil, São Paulo, Pindamonhangaba	MT036319	MT036332	MT036341
Carminda umuarama	YPH-0834	Brazil, São Paulo, Pindamonhangaba	MT036320	MT036333	MT036342
Carminda umuarama	MGCL-LOAN-297	Brazil, São Paulo, Pindamonhangaba	MT036321	—	—
Carminda surpresa	YPH-0780	Brazil, São Paulo, Campos do Jordão	MT036315	MT036328	MT036339
Carminda surpresa	YPH-0781	Brazil, São Paulo, Campos do Jordão	MT036316	MT036329	—
Carminda surpresa	YPH-0793	Brazil, São Paulo, Campos do Jordão	MT036317	MT036330	MT036340
Carminda surpresa	YPH-0689	Brazil, São Paulo, Campos do Jordão	MT036314	MT036327	MT036338

TABLE 2. Genetic distances among four species of Carminda genus.

1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17	18	19	20	21
1. YPH-0394 C. griseldis
2. NW127-17 C. griseldis	0.4
3. YPH-0462 C. griseldis	4.1	4.5
4. YPH-0785 C. griseldis	0.0	0.4	4.1
5. YPH-0814 C. griseldis	0.0	0.4	4.1	0.0
6. B-17-41 C. paeon	8.2	8.7	7.6	8.2	8.2
7. NW126-11 C. paeon	8.2	8.7	7.6	8.2	8.2	0.0
8. BLU324 C. paeon	8.2	8.7	7.6	8.2	8.2	0.0	0.0
9. BLU325 C. paeon	8.2	8.7	7.8	8.2	8.2	0.2	0.2	0.2
10. BLU326 C. paeon	8.2	8.7	7.8	8.2	8.2	0.2	0.2	0.2	0.0
11. YPH-0357 C. paeon	7.8	8.2	7.8	7.8	7.8	2.5	2.5	2.5	2.3	2.3
12. YPH-0778 C. paeon	8.5	8.9	8.5	8.5	8.5	2.3	2.3	2.3	2.1	2.1	0.6
13. YPH-0437 C. umuarama	6.9	7.4	7.4	6.9	6.9	4.3	4.3	4.3	4.5	4.5	4.1	4.3
14. YPH-0459 C. umuarama	7.1	7.6	8.0	7.1	7.1	4.5	4.5	4.5	4.7	4.7	4.3	4.5	0.9
15. YPH-782 C. umuarama	6.9	7.4	7.4	6.9	6.9	4.3	4.3	4.3	4.5	4.5	4.1	4.3	0.0	0.9
16. YPH-0833 C. umuarama	6.9	7.4	7.4	6.9	6.9	4.3	4.3	4.3	4.5	4.5	4.1	4.3	0.0	0.9	0.0
17. YPH-0834 C. umuarama	6.7	7.1	7.1	6.7	6.7	4.1	4.1	4.1	4.3	4.3	3.9	4.1	0.2	1.1	0.2	0.2
18. MGCL-LOAN-297 C. umuarama	6.5	6.9	7.4	6.5	6.5	3.9	3.9	3.9	4.1	4.1	3.7	3.9	0.4	0.9	0.4	0.4	0.6
19. YPH-0780 C. surpresa	3.3	3.7	3.7	3.3	3.3	7.0	7.0	7.0	7.0	7.0	6.5	7.2	6.9	7.2	6.9	6.9	6.7	6.5
20. YPH-0781 C. surpresa	3.3	3.7	3.7	3.3	3.3	7.0	7.0	7.0	7.0	7.0	6.5	7.2	6.9	7.2	6.9	6.9	6.7	6.5	0.0
21. YPH-0793 C. surpresa	3.3	3.7	3.7	3.3	3.3	7.0	7.0	7.0	7.0	7.0	6.5	7.2	6.9	7.2	6.9	6.9	6.7	6.5	0.0	0.0
22. YPH-0689 C. surpresa	3.3	3.7	3.7	3.3	3.3	7.0	7.0	7.0	7.0	7.0	6.5	7.2	6.9	7.2	6.9	6.9	6.7	6.5	0.0	0.0	0.0

Description of adults . Male ( Fig. 1 A–B). Forewing length 19–22 mm (mean = 20.4 mm , SD = 0.81, n = 11); hind wing length 17–19 mm (mean = 17.6 mm , SD = 0.67, n = 11). Eyes dark brown, covered by sparse brown setae. Palpus length 1.5 times head height, beige, with long dark brown and white hairs. Antenna 9–10 mm in length, with 42–43 antennomeres extending to mid-costa; shaft rust brown, dorsally covered by dark brown scales, club with 14 antennomeres, not conspicuously developed. Male foreleg covered by long dark brown hairs. DFW covered with brown scales with no marks except for a faint thick submarginal line extending from R 4 to Cu 2 . VFW dark brown with no marks except for a speckled pattern of dark brown dots at apex and subapical area. DHW covered with brown scales and presenting a thin marginal line a thick submarginal line extending from Rs to 1A+2A; presence of an oval patch of androconial scales between 1A+2A and 3A, near anal margin at proximity of tornus region. VHW entirely dark brown with a very thin marginal line extending from M 1 to 1A+2A and a fainted yellowish stripe crossing VHW; four small ocelli composed of a thin ring of yellowish-ochre scales surrounding a patch of black scales; four ocelli are located at Rs–M 1 (ocellus 1), M 1 –M 2 (ocellus 2), Cu 1 –Cu 2 (ocellus 3) and Cu 2 –1A+2A (ocellus 4); ocelli 2 and 3 are larger and ocellus 1 is absent in some individuals. FIGURE 1. A, B. Holotype male (dorsal, ventral) and C, D. Allotype female (dorsal, ventral) of Carminda surpresa sp. nov. Scale bar = 1 cm. Male genitalia ( Figs. 2 A–F). Tegumen suboval in lateral view with anterior region projected; appendix angularis short; ventral projections of tegumen and dorsal projections of saccus fused and angulated at mid-region; saccus very short when compared to valva length; anterior part of uncus robust and posterior part tapering towards tip in lateral view, where it folds down ventrally; uncus spatulate, constricted near junction with tegumen; gnathos almost half size of valva, sinuous, curved upwards in lateral view and inward near tip in dorsal view; valva trapezoid and robust in lateral view, slightly tapering towards posterior tip, which presents a small group of mid-sized “teeth”; in dorsal view valva bends inward towards mid-line; costa developed and bearing a bunch of small “teeth”; fultura superior absent; fultura inferior present as a sclerotized V-shape stripe; phallus 2/3 length of valva, sinuous in lateral view, anterior region digitiform, posterior region about 2 times longer than anterior region, covered with small knobs randomly distributed on surface ( Fig. 2E, F ) and apex bipartite in dorsal view; vesica presenting very small cornuti ( Fig. 2F ). FIGURE 2. Male genitalia of Carminda surpresa sp. nov. A . Lateral view; B . Dorsal view; C . Ventral view; D. Detailed ventral view of valvae tip and the small “teeth” at costa region; E. Aedeagus (lateral above, dorsal below); F . Magnified view of anterior portion of aedeagus showing details of randomly distributed knobs and the small cornuti; G, H. Female genitalia (lateral left, ventral right). Female ( Figs. 1C, D ). Forewing length 23–24 mm (n = 2); hind wing length 19–21 mm (n = 2); antenna 10 mm (n = 2). General color and pattern very similar to that of males, with wings more rounded. FIGURE 3. Immature stages of Carminda surpresa sp. nov. A . Freshly laid egg in lateral view; B . Egg in lateral view showing the beginning of larva development; C, D . First instar (dorsal, lateral); E, F . Second instar near molting (dorsal, ventral); G, H. Third instar (dorsal, lateral); I, J . Fourth (last) instar (dorsal, lateral); K, L, M . Pupa (ventral, lateral, dorsal). Female genitalia ( Fig 2G, H ). Eighth tergite weakly sclerotized; papilla analis ovoid, sclerotized at base, covered by long hairy-like setae at distal region, apophysis posterior absent; lamella antevaginalis as a semicircle; lateral expansions of lamella antevaginalis with one spiracle located at upper margin; ductus bursae membranous; corpus bursae almost 1.5 times longer than ductus bursae; signa dorsally paired. Variation. Variation in external morphology seems limited, and is restricted to some wing pattern elements. A yellowish irregular thick medial stripe crossing from costa to 1A+2A on the VHW is present in some individuals. The number of ocelli on the VHW can vary: while most individuals have three ocelli, a fourth tiny ocellus is present in some individuals. In addition, in several individuals the larger ocelli (ocelli 2 and 4) can have tiny double pupils. Etymology. The specific epithet, the Portuguese word for “surprise”, alludes to the fact that it was a surprise to discover a new species in a well sampled locality, and belonging to a small genus endemic to the Atlantic Forest. It is a name in apposition. Immature stages. Egg ( Figs. 3 A–B). White and round, chorion sculptured with several hexagonal cells. Height 0.96–1.01 mm (n = 4); diameter 0.92–0.94 mm (n = 4); duration 10–13 days (n = 6). First instar ( Figs. 3 C–D). Head capsule width 0.68–0.77 mm ; head scoli 0.08 mm (n = 10). Head black, bearing a pair of short scoli on vertex, each with two long narrow black setae. Third stemma larger than other stemmata. Body light green, with white longitudinal red stripes; caudal filaments very short. Legs and prolegs light green. Setae light green, all dorsal and subdorsal setae clubbed at tip. Maximum length 6 mm . Duration 7–8 days (n = 5). Second instar ( Figs. 3 E–F). Head capsule width 0.94–1.16 mm ; head scoli 0.19–0.22 mm (n = 8). Head light brown, with two diverging short scoli on vertex. Body brown, with dark longitudinal stripes; caudal filaments short. Legs and prolegs light brown. Maximum length 10 mm . Duration 5–6 days (n = 5). Third instar ( Figs. 3 G–H). Head capsule width 1.50–1.84 mm ; head scoli 0.31–0.37 mm (n = 7). Similar to second instar in color and general shape; body with several longitudinal zigzag stripes. Maximum length 18 mm . Duration 5–9 days (n = 4). FIGURE 4. Natural habitat and adults of Carminda surpresa sp. nov. A, B . General view of the wet grasslands where the species can be found; C . Adult male perched with wings closed on an inflorescence of Baccharis sp. ( Asteraceae ); D. Adult female perched on a grass leaf with wings opened. Fourth (last) instar ( Figs. 3 I–J). Head capsule width 2.66–3.46 mm ; head scoli 0.52–0.64 mm (n = 3). Head light brown, with two diverging short scoli on vertex. Body light brown with several zigzag longitudinal stripes; legs and prolegs light brown; caudal filaments short. Maximum length 30 mm . Duration 14–15 days (n = 3). A single larva presented a total of five instars, and in this case, the fourth instar was very similar to the third instar, just slightly bigger, and the last (fifth) instar was identical to that described above. Pupa ( Figs. 3 K–M). Short and smooth; mostly dark brown, with short ocular caps; cremaster broad and dark in ventral portion; dorsal abdomen with a paired series of short subdorsal light brown protuberances. Total length 9–10 mm . Duration 15–18 days (n = 4). FIGURE 5. Maximum likelihood tree obtained with RAxML showing the phylogenetic relationship of Carminda surpresa sp. nov. with other species of Carminda and with other sampled genera in Euptychiina subtribe. Number below branches are bootstrap values (values below 75 are not shown). Distribution, habitat and Natural history . Carminda surpresa sp. nov. is only known from three nearby sites above 1700 m altitude in Campos do Jordão, São Paulo , southeastern Brazil . The three sites are part of the system of natural grasslands that occur above 1600 m in the Serra da Mantiqueira mountain range. The species is associated with wet natural grasslands where the soil is permanently flooded ( Figs. 4 A–B), and are apparently restricted to this specific habitat. A single observation of an individual flying fast through a dry grassland adjacent to a wet grassland was recorded. Adults were observed flying only during the summer months, from January to March. Although no formal population studies have been done, daily counts suggest relatively small populations, with less than 10 individuals observed in a single day. Adults spent most of their time flying and landing within grass tufts of Cortaderia sp. ( Poaceae ) (a common grass species in the wet grasslands) ( Figs. 4 C–D). Courtship behavior and mating were not observed. Oviposition behavior was not observed in the field, and the host plant in nature is unknown. However, in the laboratory, larvae easily accepted Bambusa gracilis (Poaceae) . Phylogenetic relationships ( Fig. 5 ). Based on the Maximum Likelihood analysis using three genes (COI, GAPDH and RpS5), the genus Carminda is monophyletic and well-supported (bootstrap support = 94) and Carminda surpresa sp. nov. is sister to C. griseldis , forming a clade sister to C. umuarama + C. paeon , in the “ Megisto clade”, and both clades are well-supported (bootstrap support = 100, each).