taxonID	type	description	language	source
761D1C3DFFAAFFF1FF6DFCC14380FCF3.taxon	materials_examined	Type species: Papilio lea Cramer, 1777 – by present designation	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
761D1C3DFFAAFFFEFCCDF9A14022F98B.taxon	description	Male: Forewing length: 22 – 26 mm (mean: 23.8 mm; n = 13) Head: Eyes brown with lightly colored hair-like setae, white scales at base; frons dark brownish to blackish, covered with greyish and whitish scales, as well as lightly colored elongate hair-like setae; first 20 0.0326 19 0.0326 18 0.0128 0.0199 17 0.134 0.1015 16 0.1167 0.0842 15 0.1398 0.1073 14 0.118 0.0855 13 0.1214 0.0889 12 0.1291 0.0966 11 0.117 0.0845 10 0.1168 0.0843 9 0.1445 0.112 8 0.1447 0.1122 7 0.1445 0.112 6 0.1243 0.0918 5 0.1325 0.1 4 0.1298 0.0973 3 0.1384 0.1058 2 0.1492 0.1167 1 0.1385 0.1059 _ Guiana _ Guiana Cisandina French _ Continued 1708 _ 15 - _ trinitensis _ 356 Cisandina _ French trinitensis _ 3. MB- LCB Table 19 .. 20 segment of labial palpi short, covered with white scales and white hair-like scales and black hair-like scales, second segment about twice as long as eye depth and covered with white scales and white hair-like scales laterally and dorsally, in addition to brownish elongated scales, ventrally adorned with brownish hair-like scales and whitish hair-like scales longer than segment width, third segment apparently roughly one-third of second segment in length (although this can be somewhat variable), and covered with brownish scales dorsally and ventrally, with white scales laterally; antennae approximately two-fifths of forewing length, with ca. 38 – 39 antennomers (n = 4), scape rounded, about as twice as long as pedicel and wider, flagellomeres orangish, distal few flagellomeres appearing dark, covered with greyish scales with whitish scales visible on each side at base of each flagellomeres, these scales more apparent on basal flagellomeres, distal 12 – 13 flagellomeres composing club. Thorax: Brownish, dorsally with greyish scales and scales with iridescent coloration, in addition to lightly colored long hair-like scales; laterally and ventrally brownish, scattered with greyish scales and some lightly colored scales, in addition to long hair-like scales; foreleg with whitish long hair-like scales and brownish long hair-like scales, femur, tibia and tarsus similar in length, tarsus not divided into subsegments; pterothoracic legs ventrally appearing lighter compared with more greyish or darker dorsal surface, tibia with two longitudinal rows of spines on ventral surface, as well as spines present laterally, tarsus with three longitudinal rows of spines ventrally until distal end of first tarsomer, number of rows increasing to four from distal end of first tarsomer onwards, pair of tibial spurs, equal in length, present at distal end of tibia. Wing venation: Basal half of forewing subcostal vein swollen; base of cubitus swollen; forewing recurrent vein absent; discocellular vein m 1 - m 2 (i. e., dcs) curved basally, m 2 - m 3 (i. e., dcm) appearing rather straight; hindwing humeral vein present, exiting from origin of Rs and curving; origin of M 2 slightly towards M 1 than M 3 (Fig. 5). Wing shape: Forewing variably sub-triangular, apex rounded, costal margin slightly convex, outer margin somewhat variable but almost straight, inner margin almost straight; hindwing rounded, appearing slightly elongate (apparent in C. lea n. comb. and C. esmeralda n. sp. due to their rather elongated forewing shape), apex angular, costal margin slightly convex, angled inwards at base, outer margin slightly undulating, inner margin slightly curved inwards near tornus, anal lobe convex, slightly rounded (Fig. 2). Dorsal forewing: Ground color brownish, varying from being rather uniformly brownish to covered with iridescent to semi-iridescent scales (Fig. 2), submarginal and marginal bands occasionally visible as darker bands. Dorsal hindwing: Ground color similar to forewing, varying from being rather uniformly brownish to covered with iridescent to semiiridescent scales (Fig. 2); submarginal and marginal bands occasionally visible as darker bands. Ventral forewing: Ground color variable, from greyish brown to more brownish, may or may not be covered variably with iridescent scales (Fig. 2); discal band extending from radial vein, crossing discal cell, often passing just basal of origin of Cu 2, fading and terminating in cell Cu 2; concolorous postdiscal band extending from radial vein towards inner margin, terminating at 2 A if not already faded in Cu 2, more or less parallel to discal band albeit they may appear close to each other at posterior end; umbra appearing as undefined dark brownish shading extending from area near VFW ocellus (in cell M 1) to cell Cu 2, if not terminating earlier; submarginal band, appearing darker compared with discal and postdiscal band, often narrower than previous two bands, extending from apex towards tornus, rather smooth although jaggedness can be somewhat variable, appearing more or less evenly broad perhaps except for anterior and posterior end; concolorous marginal band, not jagged, narrower than submarginal band, traversing along marginal area from apex to tornus; fringe greyish; small ocellus in cell M 1, often possessing black central area and single white pupil with pale ring but may lack one or two of these elements, additional ocellus or ocelli may appear in cells R 5, M 2, M 3, and Cu 1. Ventral hindwing: Ground color range and variation similar to forewing, may or may not be covered variably with iridescent scales (Fig. 2); discal band, concolorous to that of VFW, similar in width or broader, extending from costa to inner margin, often passing origin of Rs, or just basal of origin of this vein; concolorous postdiscal band passing origin of M 2 and / or Cu 1, otherwise near origin of these veins, similar in width or broader compared with previous band and appearing more or less parallel; submarginal band, concolorous to that of VFW, similar in width or broader, more jagged, extending from apex towards tornus, posterior end occasionally fused or terminating very close to postdiscal band near inner margin; concolorous marginal band, narrower than previous band and less sinuate, traversing along marginal area from apex to tornus; fringe greyish; six submarginal ocelli in cells Rs, M 1, M 2, M 3, Cu 1, and Cu 2 (ocellus in cell Cu 2 absent in C. trinitensis n. comb.), ocelli in cells M 1 and Cu 1 similar in size (as in C. castanya n. sp., C. fida n. comb., C. sanmarcos n. comb., and C. esmeralda n. sp.) or ocellus in Cu 1 may be larger (as in C. trinitensis n. comb., C. lea n. comb., and C. philippa n. comb. & reinst. stat.), both ocelli often possessing black central area and single white pupil ringed in yellow, ocelli in cells Rs and Cu 2 smaller compared with previous two ocelli but general appearance is same, ocelli in cells M 2 and M 3 are similar in size and appearance to those in cells Rs and Cu 2 but lacking black area, ocellus in cell M 2 placed more basally compared with ocellus in cell M 3. Abdomen: Eighth tergite sclerotized in narrow anterior band and broader posterior patch; eighth sternite variable in appearance from rather narrow band to broader patch, as well as separated into two patches in some specimens (see below for further details). Genitalia: Tegumen somewhat trapezoidal in lateral view (compared with more semi-circular and / or elongated tegumen in many other euptychiines) due to convexity of dorsal margin located posteriorly and anterior portion being rather straight, this convexity of dorsal margin variable as to its degree, ventral margin convex; uncus longer than tegumen in lateral view, setae visible at base and along ventral margin towards posterior end, uncus roughly straight in lateral view, appearing somewhat broad in lateral view towards base, middle section evenly broad in dorsal view, posteriorly terminating in slightly hooked point in lateral view, rather blunt in dorsal view; brachium broad at base, longer than uncus, tapering towards apex and terminating in pointed apical edge, dorsally projecting away from uncus with posterior portion curving back in; combination of ventral arms from tegumen and dorsal arms from saccus somewhat sinuate, roughly evenly broad; appendices angulares present, but not discernible in lateral view, visible as rectangular projection in posterior view (when valva removed); saccus appearance in lateral view varying from straight to curved, but generally length not exceeding uncus; juxta (i. e., fultura inferior) present as narrow stripe; valvae, distal half setose, basal two-thirds roughly rhomboidal in lateral view, apical process approximately one-third of valva although this ratio is variable (see below for further details), varying in appearance from a narrow, curving process slightly tapering towards apex to a broader, straight process (see below for further details), y terminating in angular tip, dorsal margin just distal of costa curving interiorly forming a sharp projection visible in dorsal view in C. lea n. comb. but apparently absent in other species, costa appearing somewhat as ‘ fan-shaped’ plate, dorsal margin curving inwards and folded towards appendices angulares; phallus roughly straight, similar in length to valva in lateral in view, phallobase occupying about half of phallus, antero-dorsal opening large, sclerotized region from dorsal margin of phallobase projecting upwards and apparently forming part of manica at juncture of phallobase and aedeagus, manica covering approximately half of aedeagus, cornuti visible as roughly two sclerotized narrow patches of vesica, vesica visible at posterior opening of aedeagus (Fig. 3). Female: Forewing length: 21 – 27 mm (mean: 23.3 mm; n = 10) Similar to male except as follows: Foretarsus divided into five tarsomers; forewing somewhat rounded and broader (apparent in C. lea n. comb., C. esmeralda n. sp., C. fida n. comb., and C. sanmarcos n. comb. due to their rather elongated male forewing); overall coloration lighter; iridescent purple-ish lilac reflection on DFW variable, from absent to covering about basal half of DFW; iridescent purple-ish lilac reflection on DHW similarly variable, from absent to covering entire surface of DFW; iridescent scales around VHW tornus often less intense. Female genitalia and abdomen: Eighth tergite fully developed (as seventh tergite); papilla analis lacking posterior apothysis; lamella antevaginalis sclerotized, forming rounded to rectangular plate with wrinkled margin; sclerotized plate present at lateral side of eighth abdominal segment, anterior margin fused with lamella antevaginalis (lateral margin fused as well in C. fida n. comb. and C. sanmarcos n. comb.), dorsal margin of this lateral sclerotized plate indented around spiracle; inter-segmental membrane of seventh and eighth abdominal segment pleated and expandable, elongated weakly sclerotized region present; ductus bursae membraneous, ductus seminalis exits close to ostium bursae, ductus bursae somewhat inflated between origin of ductus seminalis and ostium bursae, corpus bursae ellipse in dorsal view, with two relatively short signa parallel to each other and apparently located laterally, bursa extending to juncture of third and fourth abdominal segment (Fig. 4). Etymology: This generic name is based on the Spanish word ‘ cisandina’, itself a compound word formed of the Latin preposition ‘ cis’, meaning ‘ on this side of’, and the Spanish adjective ‘ andina’, meaning ‘ Andean’, which is widely used to refer to the Neotropical region east of the Andes. The name refers to the fact that the species currently contained within this genus are found exclusively east of the Andean continental divide. The generic name should be regarded as a feminine noun in the nominative singular. Distribution and natural history: Cisandina n. gen. is an entirely South American genus known exclusively from east of the Andes (Fig. 8). Specifically, members are known from the cloud forests of the eastern Andes, the lowland rainforest of the Amazon basin, the Guianas, Trinidad, and the Atlantic coastal forest of Brazil and northeastern Argentina. Despite this broad range, no more than two species ever occur in sympatry. All species seem to be uncommon to very rare and are typically found in undisturbed forest. Singer and Ehrlich (1993) reported the host plant of Cisandina lea n. comb. (under the name ‘ Cissia junia’) as being an unidentified Gramineae (Poaceae) species according to their study in Trinidad, although this record is based on a plant accepted by the female in captivity in their oviposition trials. We here provide data on the immature stages of C. philippa n. comb. & reinst. stat. and C. castanya n. sp., including its natural host plant (Figs. 6 and 7), which constitute to date the only reliable immature stage data available for this genus.	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
761D1C3DFFA5FFE7FF6DF9E240AEF83D.taxon	description	(Figs. 1, 2 a – d, 3 a – d, 4 a and b, 5, 8)	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
761D1C3DFFA5FFE7FF6DF9E240AEF83D.taxon	diagnosis	Systematic placement and diagnosis: Our phylogenetic hypothesis places Cisandina lea n. comb. as sister to C. esmeralda n. sp., with strong support (Fig. 1; SH-aLRT / UFBoot = 96.5 / 96). The infra-specific genetic divergence based on DNA barcodes between the two sequenced individuals of C. lea n. comb. from two localities (LEP- 34359 from French Guiana; BC-DZ- 138 from Amazonas, Brazil) was 0.84 %, whereas interspecific divergences between C. lea n. comb. and three sampled specimens of C. esmeralda n. sp. range from 5.01 to 5.3 %. These three individuals of C. esmeralda n. sp. are from two sites, including BC-DZ- 137 (holotype) from Paraná, Brazil, and the other two specimens from Misiones, Argentina. The infraspecific genetic divergences among these three sequenced individuals of C. esmeralda n. sp. were 0.02 %. See Table 3 for further information regarding genetic divergence among Cisandina n. gen. taxa. The male of C. lea n. comb. is readily distinguished from C. esmeralda n. sp. by its iridescent to semi-iridescent bluish flush covering the DFW and DHW, whereas the DFW and DHW discal cell and adjacent area is greenish in male C. esmeralda n. sp. The broader ventral bands and larger ventral submarginal ocelli of C. lea n. sp. can be used to distinguish both sexes of these two closely related species. The female of C. lea n. comb. is also distinguished from the female of C. esmeralda n. sp. by the presence of iridescent bluish lilac reflection on the DHW, whereas this reflection appears more purplish in the female of C. esmeralda n. sp., and the ventral bands and ocelli overall appear larger in C. lea n. sp. compared with its sister species. Taxonomy: Papilio lea was described by Pieter Cramer, a Dutch merchant, in his De Uitlandsche Kapellen series (work completed by Caspar Stoll) based on an unspecified number of specimen (s) from ‘ Berbices’, a then Dutch colony (i. e. Dutch Guiana) along Rio Berbice, an area now part of the Republic of Guyana. The original description only states ‘ the blue color on both sides of the wings is shiny’. Nevertheless, Cramer provided illustrations of both wing surfaces of this taxon (pl. 151, figs. C, D), which can be used to narrow down the identity of P. lea. His drawings of this species clearly show the basal half of its DFW and all of the DHW being blue, although the anterior side of the DHW is not visible due to being covered by the forewing. The hindwing illustration shows six individual submarginal ocelli on the VHW in cells Rs, M 1, M 2, M 3, Cu 1, and Cu 2, with the ocellus in cell M 2 placed more basally compared with that in cell M 3, as well as a single ocellus present on the VFW. There is some bluish coloration visible on the DFW and DHW, especially along the ventral bands and around the VHW submarginal ocelli. The presence of six VHW ocelli excludes all Neotropical satyrine species with blue DFW and / or DHW, in genera such as Caeruleuptychia Forster, 1964, Magneuptychia, Chloreuptychia and Amiga Nakahara, Willmott, & Espeland 2019, as candidates for the species under description by Cramer, because all the species in those genera which possess bluish coloration only have five VHW ocelli, with the exception of a few species in Caeruleuptychia. Caeruleuptychia twalela Brévignon, 2005, Cae. pilata (Butler, 1867), and Cae. scripta Nakahara, Zacca, & Huertas, 2017 are the only three Caeruleuptychia species with bluish scales and six VHW ocelli, but the dorsal surface of Cae. twalela is entirely brown; the orangish rings of the VHW ocelli in cells M 1 and Cu 1 are skewed (i. e., not evenly broad) in Cae. pilata and Cae. scripta, whereas the rings of these ocelli are evenly broad in Cramer’s illustration. Perhaps one of the most notable features in Cramer’s drawing is the VHW ocellus in cell M 2 being placed more basally compared with that in cell M 3, a distinctive feature of Cisandina n. gen. This character excludes the possibility of Cramer referring to virtually any other euptychiine species apart from those discussed herein. Those few exceptions include some species in Moneuptychia Forster, 1964 such as M. vitellina Freitas & Barbosa, 2015, which can easily be distinguished by the lack of bluish coloration on the wing surface. Thus, Cramer’s P. lea represents either the female of P. lea as conceived herein, possibly C. philippa n. comb. & reinst. stat. (in which some females have bluish dorsal coloration), or one of the new species described and named in this study. However, it is very unlikely that Cramer’s illustration represents C. castanya n. sp., C. esmeralda n. sp., or C. philippa n. comb. & reinst. stat., since none of these species are known from the Guianas. Our attempt to locate syntype (s) of P. lea resulted in finding three candidate specimens in RMNH, two females each with a label indicating ‘ [Johan] Calkoen’ with the locality ‘ Surinam [e] ’ (RMNH. INS 967256 and 967257), and a male specimen with a round label indicating ‘ Verloren [van Themaat] Brasil’ (RMNH. INS 967280). Two collections now housed at RMNH, namely Joan Raye Heer van Breukelerwaard’s and Johan Calkoen’s collections, are both known to include Cramer types (de Jong 1982, Smit et al. 1986, Gernaat et al. 2012), and the former pair from ‘ Surinam [e] ’ are evidently from the latter collection. The latter male from the Verloren van Themaat collection can also be regarded as a Cramer type based on information from some existing literature on his collection (e. g., Roepke 1941). Verloren van Themaat purchased van Eyndhoven’s collection that was believed to have been sold on 14 October 1861 (see Chainey 2005; Smit et al. 1986) which is another collection believed to have included Cramer specimens (Horn and Kahle 1935, Roepke 1941). Despite the fact that Verloren van Themaat’s collection is supposed to have been destroyed due to an unfortunate battle when the collection was kept near Utrecht (Chainey 2005), we cannot exclude the possibility of some specimens ending up at Leiden where the RMNH is situated. Although lacking the abdomen, the female specimen (RMNH. INS 967257) does overall match Cramer’s illustration of P. lea, with minor differences, such as bluish scales on the DFW being somewhat restricted to cells Cu 2 and 2 A (not covering the basal half as in fig. C), but this is likely due to the color being faded over time. Nevertheless, its locality ‘ Surinam [e] ’ historically never included ‘ Berbice’ (the type locality of P. lea), which was then referred to as Dutch Guiana, and thus, it is debatable whether this female represents a syntype of P. lea or not. Confusingly, Butler (1877: 121) listed ‘ Surinam [e] ’ as the type locality for Euptychia lea (= P. lea), without any apparent evidence as to whether this was a misinterpretation or not. The only clue as without abdomen illustration; (j) illustration indicating location of signa on corpus bursae to left; (k) ventral view of lamella antevaginalis with intersegmental membrane of seventh and eighth abdominal segments expanded ([j and k] based on dissection SN- 16 - 17); C. trinitensis n. comb.: (l) dorsal view of genitalia with intersegmental membrane of seventh and eighth abdominal segments folded; illustration showing arrangement of signa to right (not to scale); (m) ventral view of lamella antevaginalis with intersegmental membrane of 7 th and 8 th abdominal segments expanded ([l and m] based on dissection SN- 20 - 85). Scale bar = 1 mm. Drawings of C. sanmarcos n. comb. are reproduced from Nakahara et al. (2018 a). to the whereabouts of the syntypes is contained in the last part of Cramer’s description ‘ [The specimen (s)] now rests in the previous collection’, referring to the last part of the description of the immediately preceding taxon, ‘ Merope’, described on the same page, ‘ She rests in the Cabinet (= collection) of the honorable gentleman A. Gevers’. As mentioned by Chainey (2005), this ‘ A. Gevers’ most likely refers to Abraham Paulusz Gevers (1712 – 1780), then the Mayor of Rotterdam, who had a natural history collection which was auctioned in 1787. Despite having no evidence of acquisition, specimens from his collection were perhaps bought by Calkoen and ended up in the RMNH, thus there exist a possibility that the provenance of the aforementioned specimen from ‘ Surinam [e] ’ is the Gevers collection, but with a changed locality. Three years later, Cramer described Papilio junia, a taxon now regarded as a junior subjective synonym of Papilio lea by Lamas (2004), again in his De Uitlandsche Kapellen. Like his description of Papilio lea, the number and sex of the examined specimen (s) used to describe P. junia cannot be unambiguously determined from the original description, except for the type locality being ‘ Surinam [e] ’. Cramer’s description is again terse, and his Dutch and French texts are accompanied with illustrations of both dorsal and ventral surfaces (pl. 292, figs. D, E). As stated in the text, the DFW and DHW are entirely bluish (described as ‘ cerulean satinlike shine’ by Cramer), with his paintings reflecting a lighter blue color relative to the darker blue coloration of the dorsal wing shown in his illustration of P. lea. The general wing pattern of the ventral surface is similar to that of his illustration of P. lea, with some minor differences such as a paler ground color, more elongated VHW submarginal ocelli, and bluish coloration on the VHW restricted to the inner margin. It must be noted that a number of painters were involved in producing copies of Cramer’s work, and there is some evidence that inevitably some variability in markings and coloration resulted, in comparison with his original drawings (Chainey 2005). Thus, these subtle dissimilarities mentioned above may not actually reflect the differences observed by Cramer himself. However, it is worth noting the small incomplete ocelli in VFW cells M 2 and M 3 visible in the illustration of P. junia, which are especially apparent on the right wing, and which appear to be absent in his illustration of P. lea. The male specimen from the Verloren collection mentioned above (RMNH. INS 967280) does exhibit ocelli on the right VFW in cells M 2 and M 3, in addition to having an extra ocellus above M 1. The male specimen from the Calkoen collection also possesses a tiny smudge-like ocellus in the right VFW cell M 2, although apparently the ocellus is absent in cell M 3. Since its description, the taxonomic viewpoint of subsequent authors concerning P. junia has been variable, ranging between some regarding it as a valid species (e. g., Butler 1877, Forster 1964) and some considering it to be conspecific with P. lea (e. g., Weymer 1911, Lamas 2004). Despite this unstable taxonomic status, it is noteworthy that these names have rarely been applied to other euptychiine species in museum collections and / or scientific articles, with few exceptions (e. g., D’Abrera 1988), an unusual situation for a group where misidentification is extremely common. Specifically, the male specimen of this taxon, as painted by Cramer (pl. 292, figs. D, E), is distinctive and is less likely to be misidentified compared with the female which can be confused with species in the same genus such as C. castanya n. sp. Recently, Nakahara et al. (2019 a) did not accept Calkoen specimens (also housed in the RMNH) as potential syntypes of Papilio ebusa Cramer 1780 because of the discrepancy in terms of the type locality indicated on the associated label. Unlike this previous case, the handwritten locality on the rounded label for RMNH. INS 967256 does match the type locality indicated in Cramer’s description of P. junia, both being ‘ Surinam [e] ’. Thus, there seems to be no reason to reject this male specimen as a syntype, and thus we here designate this male syntype as the lectotype for P. junia with the following labels separated by double-forward slashes (lectotype designation): // Calkoen Suriname // RMNH. INS 967256 //. Accepting this male specimen as a former syntype of P. junia reinforces the possibility of the aforementioned female specimen (RMNH. INS 967257) being a syntype of P. lea with an erroneous locality on the label. The fact that only a single female for a possible Cramer syntype was located at RMNH matches the fact that Cramer illustrated a female for P. lea and a male specimen for P. junia. Therefore, we designate this female from the Calkoen collection as the lectotype of P. lea with the following labels separated by double-forward slashes (lectotype designation): // Calkoen Suriname // RMNH. INS 967257 //. Distribution and natural history: Cisandina lea n. comb. is known from the Guianas, Trinidad, southern Venezuela, as well as the central and lower Amazon in Brazil, in addition to northeastern Brazil (Fig. 8). Specimens examined (129 ♂, 80 ♀): Brazil: Alagoas: São José de Lage, Usina Serra Grande, [8 ° 58 ′ S, 36 ° 3 ′ W], 400 – 500 m, 1 – 3 Aug 2003, 1 ♀, (ZUEC); Maceió, [9 ° 39 ′ S, 35 ° 46 ′ W], (Cardoso, A.), 30 Apr 1945, 1 ♀, (DZUP); Amazonas: Barcelos, Rio Aracá, Foz do Rio Curuduri, [0 ° 5 ′ 50 ″ S, 63 ° 17 ′ 22 ″ W], (Mielke, O. H. H., Casagrande, M. M.), 15 – 19 Jun 2010, 1 ♀, (DZ 49.955, BC-DZ 138) (DZUP); Ega (= Tefé), [3 ° 22 ′ S, 64 ° 42 ′ W], (Bates, H. W.), 1 ♀ [BMNH (E) - 1497637], (NHMUK); Manaus, [3 ° 7 ′ S, 60 ° 2 ′ W], (Hahnel), 1886, 1 ♀, (MNHU); Manicoré, [5 ° 49 ′ S, 61 ° 17 ′ W], (Le Moult, E.), 1 ♂ [FLMNH-MGCL- 265701], (FLMNH); Maués, [3 ° 22 ′ S, 57 ° 43 ′ W], (Le Moult, E.), 1 ♂ [FLMNH-MGCL- 265699], 1 ♀ [FLMNH- MGCL- 265703], (FLMNH); Rio Negro, 50 - km NW Manaus, (Schmidt, U.), 1 – 4 Nov 1993, 1 ♀ [FLMNH-MGCL- 265705], [dissection, SN- 20 - 31] (FLMNH); Bahia: [12 ° 59 ′ S, 38 ° 31 ′ W], 1 ♂ [BMNH (E) - 1497632], (NHMUK); Espírito Santo: [20 ° 20 ′ S, 40 ° 17 ′ W], 1 ♂ [BMNH (E) - 1497649], 1 ♂ [BMNH (E) - 1497650], 1 ♂ [BMNH (E) - 1497651], (NHMUK); Pará: [Rio] Tapajós, [4 ° 16 ′ 8 ″ S, 55 ° 59 ′ 10 ″ W], 25 m, (Bates, H. W.), 1 ♂ [BMNH (E) - 1497630], (NHMUK); Amazonas, (Fassl, A. H.), (ZSM); Bragança, [1 ° 3 ′ S, 46 ° 47 ′ W], (Mathan, M. de), 1 ♀ [BMNH (E) - 1497641], (NHMUK); Breves, [1 ° 40 ′ S, 50 ° 28 ′ W], 1 ♀ [BMNH (E) - 525609], (NHMUK); hwy Cuiabá-Santarém, km 1666, [3 ° 17 ′ 17 ″ S, 54 ° 56 ′ 22 ″ W], - 150, (Callaghan, C. J.), 28 Jul 1978, 1 ♂ [FLMNH-MGCL- 265697], (FLMNH); hwy Cuiabá-Santarém, km 958, [8 ° 3 ′ S, 55 ° 2 ′ W], (Callaghan, C. J.), 30 Jul 1978, 1 ♀ [FLMNH- MGCL- 265706] [dissection, SN- 20 - 17], (FLMNH); Itaituba on Rio Tapajós, [4 ° 16 ′ 8 ″ S, 55 ° 59 ′ 10 ″ W], (Le Moult, E.), 1 ♂ [FLMNH- MGCL- 265702], [dissection, SN- 20 - 16] (FLMNH); Itaituba, [4 ° 17 ’ S, 55 ° 59 ’ W], (Mich.), 1890, 1 ♀ [dissection, Lee D. Miller 9136], (MNHU), 1893, 1 ♂ [dissection, Lee D. Miller 9135], (MNHU); Itaituba, Rio Tapajós, [4 ° 16 ′ 8 ″ S, 55 ° 59 ′ 10 ″ W], 25 m, (Lathy), May 1932, 1 ♂ [BMNH (E) - 1670284], (NHMUK); Óbidos, [1 ° 54 ′ S, 55 ° 31 ′ W], (Le Moult, E.), 1 ♂ [FLMNH-MGCL- 265700], (FLMNH); Pará, [1 ° 0 ′ S, 51 ° 11 ′ W], 1 ♂ [BMNH (E) - 1497629], 1 ♀ [BMNH (E) - 1497640], (NHMUK), (Bates, H. W.), 1 ♀ [BMNH (E) - 1497638], (NHMUK), (Wallace), 1 ♀ [BMNH (E) - 1497639], (NHMUK); Rio Tapajós, [4 ° 16 ′ 8 ″ S, 55 ° 59 ′ 10 ″ W], 25 m, (Klug, G. G.), Chainey 2005 Cramer ep 1931, 1 ♀ [BMNH (E) - 1497729], (NHMUK); Rio Tapajós, Mujo, [2 ° 41 ′ S, 54 ° 38 ′ W], (Le Moult, E.), 1 ♂ [FLMNH-MGCL- 265698], (FLMNH), Sep 1922, 1 ♀ [FLMNH- MGCL- 265704], (FLMNH); Santarém, [2 ° 26 ′ S, 54 ° 43 ′ W], (Smith, H. H.), 1 ♂ [BMNH (E) - 1497631], 1 ♀ [BMNH (E) - 1497642], 1 ♀ [BMNH (E) - 1497643], (NHMUK); Roraima: Alto Alegre, Ilha de Maracá, [3 ° 21 ′ 41 ″ N, 61 ° 26 ′ 2 ″ W], (Mielke, O. H. H., Casagrande, M. M.), 24 – 31 Aug 1987, 3 ♂ 1 ♀ (DZ 49.925, DZ 49.935 - BC-DZ Willmott 191, DZ 49.945, DZ 49.965, 23 – 28 Feb 1988, 2 ♀ (DZ 5371, DZ 49975) (DZUP); Not located: ‘ Brasilia’, 1 ♂ [BMNH (E) - 1497635], (NHMUK); ‘ Brazil’, 1 ♂ [BMNH (E) - 1497633], 1 ♂ [BMNH (E) - 1497634], 1 ♂ [BMNH (E) - 1497636], 1 ♂ [BMNH (E) - 1497652], 1 ♂ [BMNH (E) - 1497653], 1 ♂ [BMNH (E) - 1497654], 1 ♀ [BMNH (E) - 1497647], 1 ♀ [BMNH (E) - 1497648], (NHMUK); not located: ‘ Amazon’, 1 ♂ [BMNH (E) - 1497714], 1 ♂ [BMNH (E) - 1497727], (NHMUK), (Bates, H. W.), 1 ♀ [BMNH (E) - 1497730], (NHMUK) [Bates only collected in Brazil]. French Guiana: Cayenne: Cayenne, [4 ° 56 ′ N, 52 ° 20 ′ W], 1 ♂ [BMNH (E) - 787669; (NHMUK); Saint-Laurent-du-Maroni: Bas Maroni, 1 ♂ [FLMNH-MGCL- 265707; dissection, KW- 14 - 018], (FLMNH); Maroni River, (Le Moult, E.), 1 ♀ [FLMNH- MGCL- 265708], (FLMNH); Maroni river, Maripasoula, [3 ° 41 ′ N, 54 ° 2 ′ W], (Brévignon, C.), 14 May 1987, 1 ♂ [MUSM- LEP- 103094], (MUSM); Saül, [3 ° 51 ′ 30 ″ N, 53 ° 18 ′ 14 ″ W], 200 – 450 m, (Nakahara, S.), 5 Aug 2014, 1 ♂ [FLMNH-MGCL- 209436], 1 ♀ [FLMNH-MGCL- 195747], (FLMNH); St. Laurent du Maroni, [5 ° 30 ′ N, 54 ° 2 ′ W], Jul-Sep 1915, 1 ♂ [BMNH (E) - 1497720], (NHMUK), (Le Moult), Nov, 1 ♂ [BMNH (E) - 1497721], (NHMUK); Not located: ‘ French Guiana’, (Bar, C.), 1 ♂ [BMNH (E) - 1497722], 1 ♂ [BMNH (E) - 1497723], (NHMUK). Guyana: Cuyuni-Mazaruni: Bartica, [6 ° 24 ′ N, 58 ° 37 ′ W], (Parish, H. S.), 1 ♂ [BMNH (E) - 787654], 1 ♀ [BMNH (E) - 787665], (NHMUK), (Parish, H. S.), 1 ♂ [BMNH-E- 787654], 1 ♀ [BMNH-E- 787665], (NHMUK); East Berbice-Corentyne: New River Triangle, Camp Jaguar, [3 ° 18 ′ 23 ″ N, 57 ° 35 ′ 21 ″ W], 152 m, (Steinhauser, S. R.), 13 Nov 1980, 1 ♀ [FLMNH-MGCL- 265710], (FLMNH); Upper Demerara-Berbice: Berbice, [5 ° 27 ′ N, 57 ° 57 ′ W], 1 ♂ [BMNH (E) - 787651], 1 ♂ [BMNH-E- 787651], (NHMUK); Upper Takutu-Upper Essequibo: E. Kanuku Mountains, Two Hat Mountain, [3 ° 8 ′ 48 ″ N, 59 ° 6 ′ 54 ″ W], 244 m, (Fratello, S., et al), 17 Sep – 2 Oct 2000, 1 ♂, (USNM); Essequibo River, (Büche, M.), Jul 1997, 1 ♂ [MUSM-LEP- 103093], (MUSM); Essequibo River, Aunai, [5 ° 22 ′ N, 58 ° 53 ′ W], (Whitely, H.), 1 ♂ [BMNH (E) - 787650], 1 ♂ [BMNH-E- 787650], 1 ♀ [BMNH (E) - 787664], 1 ♀ [BMNH-E- 787664], (NHMUK); Kanuku Mountains, [3 ° 12 ′ N, 59 ° 34 ′ W], 152 – 305 m, (Fratello, S., Hanner, R., Hendricks, S., Williams, R.), 21 Feb – 10 Mar 1999, 1 ♂, (USNM); Lethem, (Le Moult, E.), 8 Aug 1971, 1 ♀ [FLMNH-MGCL- 265709], (FLMNH); Not located: ‘ Guyana’, 1 ♂ [BMNH (E) - 787652], 1 ♂ [BMNH-E- 787652], 1 ♀ [BMNH (E) - 1497716], 1 ♀ [BMNH-E- 787669], (NHMUK), (Parish), 1 ♀ [BMNH-E- 787667], (NHMUK), (Parish, H. S.), 1 ♀ [BMNH (E) - 787667], (NHMUK), (Whitely), 1 ♀ [BMNH-E- 787666], (NHMUK), (Whitely, H.), 1 ♀ [BMNH (E) - 787666], (NHMUK); Demerara River, Akayma Fort, 1 ♀ [BMNH (E) - 1497715], (NHMUK). Suriname: Brokopondo: Berg-en-Dal, [5 ° 9 ′ N, 55 ° 4 ′ W], (Ellacombe, C. W.), Apr 1892, 1 ♂ [BMNH (E) - 787668], (NHMUK), (Ellacombe, C. W.), Apr 1892, 1 ♀ [BMNH-E- 787668], (NHMUK); Para: Bersaba, [5 ° 32 ′ N, 55 ° 3 ′ W], (Michls.), 1898 – 1899, 1 ♂ [dissection, M- 9137 Lee D. Miller], (MNHU); Not located: ‘ Surinam’, 1 ♂ [BMNH (E) - 1497718], 1 ♂ [BMNH (E) - 1497719], 1 ♂ [BMNH (E) - 787656], (NHMUK), (Fruhstorfer), May – Sep, 1 ♂ [BMNH-E- 787656], (NHMUK); ‘ Surinam’, (ZSM); ‘ interior Surinam’, (Ellacombe, C. W.), Sept 1892, 1 ♂ [BMNH (E) - 1497717], (NHMUK). Trinidad: Couva- Tarabaquite-Talparo: Caparo, [10 ° 34 ′ N, 61 ° 16 ′ W], (Birch, F.), 1 ♀ [BMNH-E- 787662], (NHMUK); Narieva, Tabaquite, [10 ° 23 ′ N, 61 ° 18 ′ W], 1 ♂ [BMNH (E) - 1497678], 1 ♂ [BMNH (E) - 1497679], 1 ♀ [BMNH (E) - 1497671], 1 ♀ [BMNH (E) - 1497672], (NHMUK); Diego Martin: Fort George, [10 ° 42 ′ N, 61 ° 32 ′ W], Sept. 1891, 1 ♂ [BMNH (E) - 1497660], 1 ♂ [BMNH (E) - 1497661], 1 ♂ [BMNH (E) - 1497662], 1 ♂ [BMNH (E) - 1497663], 1 ♂ [BMNH (E) - 1497664], 1 ♂ [BMNH (E) - 1497665], 1 ♀ [BMNH (E) - 1497666], 1 ♀ [BMNH (E) - 1497667], (NHMUK); Mt. Catherine upper trail, (Preston, J. & F.), 27 Feb 1982, 1 ♀ [FLMNH- MGCL- 265722], 1 ♀ [FLMNH-MGCL- 265723], (FLMNH); Port of Spain: Port of Spain, [10 ° 40 ′ N, 61 ° 31 ′ W], (Jenkins, D. W.), 27 Jan 1977, 1 ♂ [FLMNH-MGCL- 265715], (FLMNH), (Rosen, V.), 23 Oct, (ZSM); Rio Claro-Mayaro: ‘ Nariva’, 06 - 05 - 1993, 1 ♂ [BMNH (E) - 1670212; ‘ Swamp’], (NHMUK); San Juan-Laventille: Fondes Amandes Road, (Patterson, E. J.), 1 ♀ [FLMNH- MGCL- 265724], (FLMNH); Hololo Mt., [10 ° 41 ′ 29 ″ N, 61 ° 29 ′ 7 ″ W], (Morrall, J.), 4 Oct 2012, 1 ♂, (MZUJ); Maraval, [10 ° 43 ′ N, 61 ° 31 ′ W], Jan ‘ 92, 1 ♂ [BMNH (E) - 1497680], 1 ♂ [BMNH (E) - 1497681], 1 ♀ [BMNH (E) - 1497684], 1 ♀ [BMNH (E) - 1497685], (NHMUK), (Hall, A.), Nov 1931 – Feb 1932, 1 ♂, (BMB); St. Annes Valley, [10 ° 41 ′ N, 61 ° 30 ′ W], 1 ♂ [FLMNH-MGCL- 265718], (FLMNH), 1 ♂ [BMNH (E) - 787646], 1 ♂ [BMNH (E) - 787647], 1 ♂ [BMNH (E) - 787648], (NHMUK); St. Anns, [10 ° 41 ′ N, 61 ° 30 ′ W], 1 ♂ [BMNH-E- 787646], 1 ♂ [BMNH-E- 787647], 1 ♂ [BMNH-E- 787648], (NHMUK), (Hall, A.), Nov – Dec 1931, 1 ♀, (BMB); Tunapuna- Piarco: 6 mi. N Arima, [10 ° 42 ′ 12 ″ N, 61 ° 17 ′ 28 ″ W], 300 m, (Pliske, T. E.), 30 Jun 1962, 1 ♂ [FLMNH-MGCL- 265720], (FLMNH); Arima Valley, [10 ° 41 ′ N, 61 ° 17 ′ 30 ″ W], 305 – 457 m, (Breedlove, D. E.), 14 – 19 Dec 1970, 1 ♀, (CAS); Tunapuna, [10 ° 39 ′ 7 ″ N, 61 ° 23 ′ 17 ″ W], 50 m, 1 ♂ [BMNH (E) - 1497690], 1 ♀ [BMNH (E) - 787660], 1 ♀ [BMNH-E- 787660], (NHMUK); Not located: ‘ Trinidad’, [10 ° 26 ′ 17 ″ N, 61 ° 15 ′ 12 ″ W], 1 ♂ [BMNH (E) - 1497704], 1 ♂ [BMNH (E) - 1497705], 1 ♂ [BMNH (E) - 1497707], 1 ♂ [BMNH (E) - 1497708], 1 ♂ [BMNH (E) - 1497709], 1 ♂ [BMNH (E) - 1497710], 1 ♂ [BMNH (E) - 1497711], 1 ♂ [BMNH (E) - 1497712], 1 ♂ [BMNH (E) - 787643], 1 ♂ [BMNH (E) - 787655], 1 ♂ [BMNH-E- 787643], 1 ♂ [BMNH-E- 787644], 1 ♂ [BMNH-E- 787655], 1 ♀ [BMNH (E) - 1497696], 1 ♀ [BMNH (E) - 1497697], 1 ♀ [BMNH (E) - 1497699], 1 ♀ [BMNH (E) - 787657], 1 ♀ [BMNH (E) - 787658], 1 ♀ [BMNH-E- 787657], 1 ♀ [BMNH-E- 787658], (NHMUK), (Feather), 1 ♂ [BMNH (E) - 1497706], 1 ♀ [BMNH (E) - 1497688], (NHMUK), (Fountaine, M.), Dec. 1911, 1 ♂ [BMNH (E) - 1497693], 1 ♂ [BMNH (E) - 1497694], 1 ♂ [BMNH (E) - 1497695], 1 ♂ [BMNH (E) - 1497702], 1 ♂ [BMNH (E) - 1497703], 1 ♂ [BMNH (E) - 787653], 1 ♀ [BMNH (E) - 1497687], 1 ♀ [BMNH (E) - 1497698], 1 ♀ [BMNH (E) - 1497700], 1 ♀ [BMNH (E) - 1497701], 1 ♀ [BMNH (E) - 787661], 1 ♀ [BMNH (E) - 787663], (NHMUK), Dec 1911, 1 ♂ [BMNH-E- 787653], 1 ♀ [BMNH-E- 787661], 1 ♀ [BMNH-E- 787663], (NHMUK), (Hall, A.), Jan 1936, 1 ♂, 1 ♂ [Booth Mus: Collection 00 - 5938], (BMB), Oct-Dec 1920, 1 ♂, (BMB), (Kaye, W. J.), May 1898, 1 ♂ [FLMNH-MGCL- 265719], (FLMNH), May 1898, 1 ♂ [FLMNH-MGCL- 265716], (FLMNH), (Neuburger), (ZSM); Arima Valley, vicinity Asa Wright Nature Centre vic, [10 ° 43 ′ 3 ″ N, 61 ° 17 ′ 55 ″ W], (Austin, G. T.), Feb 1993, 1 ♂ [FLMNH-MGCL- 296554], [dissection, SN- 20 - 30] (FLMNH), (Gomes, O.), Feb 1993, 1 ♀ [FLMNH-MGCL- 298000], (FLMNH); Behind St. Benet’s Hall, (Preston, J. & F.), 2 Jan 1982, 1 ♀ [FLMNH- MGCL- 265721], (FLMNH); Caparo, (Birch, F.), 4 th Aug., 1 ♀ [BMNH (E) - 1497683], (NHMUK), 4 th Sept. ‘ 04, 1 ♀ [BMNH (E) - 787662], (NHMUK), (Klages, S. M.), Jan 1906, 1 ♂ [BMNH (E) - 1497682], (NHMUK); Northern Mts., (Hall, A.), Dec 1938 - Jan 1939, 1 ♀, (BMB); Port of Spain, (Rendall), II. 97, 1 ♂ [BMNH (E) - 1497691], 1 ♀ [BMNH (E) - 1497686], (NHMUK); ‘ Saint George’, (Ellacombe, C. W.), 1 ♂ [BMNH-E- 787645], 1 ♀ [BMNH-E- 787659], (NHMUK); Santa Margarita, (Preston, J. & F.), 13 Oct 1981, 1 ♂ [FLMNH-MGCL- 265717], (FLMNH); St. George, (Ellacombe, G. W.), 1 ♂ [BMNH (E) - 787645], 1 ♀ [BMNH (E) - 787659], (NHMUK), Dec. 1891, 1 ♀ [BMNH (E) - 1497668], 1 ♀ [BMNH (E) - 1497669], 1 ♀ [BMNH (E) - 1497760], (NHMUK), Oct. 1891, 1 ♂ [BMNH (E) - 1497673], 1 ♂ [BMNH (E) - 1497674], 1 ♂ [BMNH (E) - 1497675], 1 ♂ [BMNH (E) - 1497676], 1 ♂ [BMNH (E) - 1497677], (NHMUK); Stanway Parris River, Feb 1921, 1 ♂ [BMNH (E) - 1497692], (NHMUK); Symonds Valley, (Hall, A.), Apr 1930, 1 ♂, (BMB), Mar 1930, 1 ♀, (BMB). Venezuela: Amazonas: Samariapo, 120 m, (Lichy, R.), 7 Oct 1946, 1 ♀ [FLMNH- MGCL- 265714], (FLMNH); Bolívar: 80 km S El Dorado, [6 ° 11 ′ 8 ″ N, 61 ° 24 ′ 36 ″ W], (Nation, J. L.), 26 Jun 1984, 1 ♂ [FLMNH- MGCL- 265712], (FLMNH); Not located: ‘ Venezuela’, 1 ♂ [BMNH (E) - 1497659], 1 ♂ [BMNH (E) - 1497713], (NHMUK); Pitotan, (Nation, J. L.), 9 Jun 1937, 1 ♂ [FLMNH-MGCL- 265713], (FLMNH). Country unknown: no data, 1 ♀ [FLMNH- MGCL- 265711], (FLMNH), 1 ♂ [BMNH (E) - 1497655], 1 ♂ [BMNH (E) - 1497656], 1 ♂ [BMNH (E) - 1497724], 1 ♂ [BMNH (E) - 1497725], 1 ♂ [BMNH (E) - 1497726], 1 ♂ [BMNH (E) - 787649], 1 ♂ [BMNH-E- 787649], 1 ♀ [BMNH (E) - 1497728], (NHMUK), (Hall, A.), 18 Mar 1934, 1 ♀, (BMB). Doubtful locality: ‘ Cauca valley’, 1 ♂ [BMNH (E) - 1497657], (NHMUK). ‘ Haiti’, 1 ♂ [BMNH (E) - 1497731], 1 ♂ [BMNH (E) - 1497732], (NHMUK).	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
761D1C3DFFBCFFE7FCCDFF4C4507FE99.taxon	description	(Figs. 1, 2 e and f, 3 e – g, 8)	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
761D1C3DFFBCFFE7FCCDFEFC4468FA90.taxon	diagnosis	Systematic placement and diagnosis: As discussed under the immediately preceding taxon, C. esmeralda n. sp. is recovered as sister to C. lea n. comb. with a strong support (Fig. 1; SH-aLRT / UFBoot = 96.5 / 96). Our molecular phylogeny and DNA sequence divergence between these two species support its species-level status indicated by its distinctive male phenotype. Indeed, the iridescent greenish lilac scales possessed by the male of C. esmeralda n. sp. readily distinguish this species from the remainder of the genus and furthermore from all other known euptychiines. In addition, the more brownish ventral ground color, small ventral submarginal ocelli (smallest among all Cisandina n. gen. species), narrow and somewhat sinuate VHW postdiscal band, coupled with other narrow ventral bands (narrowest among all four similar-appearing Cisandina n. gen. species with iridescent scales) separate the male of this species from the other three Cisandina n. gen. species with iridescent scales discussed herein. The more elongate forewing is a character shared with the male of C. lea n. comb., but not with males of other taxa discussed herein. The female of C. esmeralda n. sp. is distinguished from the female specimens of C. lea n. comb. and C. castanya n. sp. by its lack of purple lilac scales on the dorsal surface thus the female of C. esmeralda n. sp. possesses a uniformly brown dorsal wing surface. Cisandina philippa n. comb. & reinst. stat. is similar in this respect because the female of C. esmeralda n. sp. Is brown dorsally. Nevertheless, the narrower ventral bands and smaller ventral submarginal ocelli of C. esmeralda n. sp., as well as its smaller adult size, can be used to separate females of these two species. Furthermore, the outward-curving VFW postdiscal band of the female C. esmeralda n. sp. is rather distinctive. See also ‘ Remarks’ below for further information regarding the identity of the female of this species. Male (Fig. 2 e): Forewing length 23 mm (n = 1). Head: See head description for Cisandina n. gen. above.	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
761D1C3DFFBDFFE0FCCDFBC34511F91D.taxon	description	(Figs. 1, 2 g and h, 3 h – j, 4 c and d, 6 a and b, 7 a and b, 8)	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
761D1C3DFFBDFFE0FCCDFBC34511F91D.taxon	diagnosis	Systematic placement and diagnosis: Our maximum likelihood approach found Cisandina philippa n. comb. & reinst. stat. as a sister taxon to C. lea n. comb. + C. esmeralda n. sp., with a weak to moderate support (Fig. 1; SH-aLRT / UFBoot = 80.8 / 71). The infraspecific genetic divergence among seven sequenced C. philippa n. comb. & reinst. stat. specimens varies from 0.04 % to 3.55 %, although this remarkably high maximum value (3.55 %) is due to DNA 99 - 022 (from Madre de Dios, Peru), which lacks data for the first approximately 250 nucleotides. The six Ecuadorian individuals, representing samples from Zamora-Chinchipe and Morona- Santiago provinces, exhibit a range of within-species COI divergence from 0.01 to 1.71 % with a mean of 0.626 %. Based on our COI data, genetic distance among these sampled C. philippa n. comb. & reinst. stat. individuals and four other closely related Cisandina n. gen. species sequenced for this study is a minimum of 3.74 %, suggesting an existence of a ‘ barcoding gap’. See Table 3 for further information regarding genetic divergence of Cisandina n. gen. taxa. The male of C. philippa n. comb. & reinst. stat. is readily distinguished from males of other species in the genus by its uniformly brown dorsal surface, whereas male specimens of other species exhibit either bluish or greenish iridescent scales on the dorsal surface. The female specimens of this species are also dorsally brown, which is also the case with C. esmeralda n. sp. but not two other species in the genus; see corresponding section of C. esmeralda n. sp. for further diagnostic characters to identify females. Taxonomy: Euptychia philippa was described by Arthur Gardiner Butler in his monograph of Euptychia (sensu lato), where he introduced 60 new euptychiine butterflies to science. Like many other species described by Butler, information regarding the number of examined specimens and sex was not provided in the original description of E. philippa. His description of E. philippa was also not accompanied by any illustration of this taxon, but the identity of this species can be guessed from the Latin text and interpretation of a few other aspects of Butler’s work. First, E. philippa was described in his ‘ Division II’ of Euptychia, which is one of his seven subdivisions of Euptychia he erected in his monograph of the group. Butler (1867: 481) characterized this ‘ Division II’ as ‘ wings variable above and below, of violet, blue and green’. Those species classified in this division, along with E. philippa, include E. picea Butler, 1867, E. lysidice (Cramer, 1777), E. glaucina Bates, 1865, E. aegrota Butler, 1867, E. pilata Butler, 1867, E. brixiola Butler, 1867, E. brixius (= Satyrus brixus Godart [1824]), E. coelestis Butler, 1867, E. urania Butler, 1867, E. lea, and E. junia. Despite some of these names not being considered valid today, Butler apparently grouped together species that possess iridescent wing coloration, judging from the phenotypes of the species listed by him. It is also worth noting that the two species immediately preceding E. philippa in Butler’s monograph are E. lea and E. junia, two names considered as applying to the same species by Lamas (2004), a proposal followed by the present study, and which species proves to be a member of Cisandina n. gen. in our molecular phylogeny (Fig. 1). Furthermore, the description of E. philippa begins by stating ‘ alae supra fuscae’ which translates to ‘ wings above dark brown’, and ends by noting its ventral similarity with E. junia, both statements that narrow down the candidates examined by Butler to female C. esmeralda n. sp. or what we regard here as E. philippa. If this assumption is correct, the possibility of Butler examining female C. esmeralda n. sp. can be excluded on the basis of the type locality of E. philippa being Ega [= Tefé] according to the original description, whereas C. esmeralda n. sp. is a taxon known from the Atlantic coastal forest of Brazil and Argentina. The syntype housed at the NHMUK is a male specimen with a uniformly brown dorsal surface as described by Butler and in accordance with characters and inferences discussed above. Like Butler, who considered the brown dorsal surface to be a character to justify E. philippa as a species-level taxon, some subsequent authors also followed this trend (e. g., D’Abrera 1988). On the other hand, other authors proposed an opposing taxonomic hypothesis, such as Weymer (1911), who regarded E. philippa to be a ‘ form’ of P. lea from Ega (= Tefé). Following Weymer’s (1911) proposal, in which the name was considered to be subspecific according to Article 45.6.4.1 of the ICZN (1999), Lamas (2004) also regarded this taxon as subspecific. Nevertheless, both genetic divergence based on COI and multi-locus maximum likelihood (see ‘ Systematic Placement and Diagnosis’, Fig. 1 and Table 2 for further information) are in favor of species-level status for this taxon with its uniformly brown dorsal surface. To reflect this taxonomic change and to settle its nomenclature as a senior subjective synonym of Euptychia batesii f. tersa, we here designate the aforementioned male syntype in the NHMUK as the lectotype of E. philippa, with the following labels separated by double-forward slashes, and reinstate its taxonomic status from subspecific to specific (lectotype designation, reinstated status): // B. M. TYPE No. Rh 3178 Euptychia philippa, ♂ Butl. // ♂ // Type of Species // Ega, U. Amazonas. H. W. Bates. // ♂ Ega Philippa Butl. Type // Type H. T. // Godman-Salvin Coll. 1904. - 1. Euptychia Philippa, Butl. //.	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
761D1C3DFFBDFFE0FCCDFBC34511F91D.taxon	distribution	Distribution and natural history: This species is known from the western Amazon, from the Andean foothills from Ecuador to southern Peru, east to the central Amazon, with a possible small area of sympatry with C. lea n. comb. in Tefé and the vicinity of Manaus (Fig. 8). It is sympatric with C. castanya sp. n. from central to southern Peru and in western Brazil (Rondônia). In Ecuador, this species occurs in lowland rainforest up to 1,400 m, where it is uncommon. Males and females were encountered at similar frequency, in a variety of sites, mostly in undisturbed terra firme forest, but also in floodplain secondary forest with abundant Guadua (Poaceae) bamboo. Individuals were encountered flying at varying heights above the ground, from 1 to 4 m, in both shady understory and in light gaps and at forest edges. A penultimate instar caterpillar was found on species of herbaceous bamboo, Taquara micrantha (Kunth) I. L. C. O liveira & R. P. O liveira (Poaceae) at Finca Las Piedras, Madre de Dios, Peru, on 20 April 2021 (voucher: 2020 - FLP-IMM- 0336), and the immature stages are described below. Specimens examined (39 ♂, 31 ♀): Brazil: Acre: Marechal Thaumaturgo, Foz do Rio Tejo, Reserva Extrativista Alto Juruá, estrada para o Rio Arara, (Brown, K. S., Freitas, A. V. L.), 16 Sep 1997, 1 ♂, (ZUEC); [Marechal Thaumaturgo, Boca do Rio Tejo, Reserva Extrativista Alto Juruá], 20 – 27 Aug 1997, 1 ♂ [‘ BTEJO- REAJ-AC’], (ZUEC); Senador Guiomard, Reserva Catuaba, (Mielke & Casagrande), 31 Jan – 5 Feb – 2009, 1 ♂ 1 ♀ [DZ 52.562, DZ 52. 564] (DZUP); Amazonas: Borba, Rio Abacaxis, Comunidade Paxiúba, [4 ° 28 ′ 48 ″ S, 58 ° 34 ″ W], (Mielke, O. H. H., Casagrande, M. M.), 2 – 4 Jun 2008, 1 ♀, [DZ 52.563 – BC-DZ Willmott – 140] (DZUP); Ega (= Tefé), [3 ° 22 ′ S, 64 ° 42 ′ W], (Bates, H. W.), 1 ♂ [‘ ♂ Type of Species. ’ // ’ Godman-Salvin Coll. 1904 - 1. Euptychia philippa, Butl. ’ // ’ Ega, U. Amazons. H. W. Bates. ’ // ’ ♂ Ega Philippa Butl. Type’ // ’ Type H. T. ’ // ’ B. M. TYPE No. Rh 3178. Euptychia philippa. ♂ Butl. ’], 1 ♂ [BMNH (E) - 1266946; HT of philippa], 1 ♂ [BMNH (E) - 1670283], 1 ♂ [BMNH (E) - 1670295], 1 ♀ [BMNH (E) - 1670287], 1 ♀ [BMNH (E) - 1670288], (NHMUK); Igarapé Massauari, [2 ° 54 ′ 17 ″ S, 57 ° 8 ′ 23 ″ W], (Hahnel, P.), 1 ♂, (MNHU); Manicoré, [5 ° 49 ′ S, 61 ° 17 ′ W], (Hahnel, P.), 1887, 1 ♂, (MNHU); São Paulo de Olivença, [3 ° 28 ′ S, 68 ° 57 ′ W], (Mathan, M. de), 1 ♂ [BMNH (E) - 1670280], 1 ♂ [BMNH (E) - 1670281], 1 ♂ [BMNH (E) - 1670282], 1 ♀ [BMNH (E) - 1670211], (NHMUK), Jun – Jul 1883, 1 ♂ [BMNH (E) - 1670275], 1 ♂ [BMNH (E) - 1670276], 1 ♂ [BMNH (E) - 1670277], 1 ♂ [BMNH (E) - 1670278], 1 ♂ [BMNH (E) - 1670279], 1 ♀ [BMNH (E) - 1670285], 1 ♀ [BMNH (E) - 1670286], (NHMUK), (Moss, A. M.), 1 ♀ [BMNH (E) - 525170], (NHMUK); Tefé, [3 ° 22 ′ S, 64 ° 44 ′ W], (Hahnel, P.), 1 ♂ [dissection, 9076; ‘ NEOTYPE ♂ Euptychia batesii f. tersa WeymerdesignatedbyLeeD. Miller 1989 // Teffe (= Tefé) Hhl. // Genitalia vial M- 9076 ♂ Lee D. Miller’], (MNHU) [unpublished neotype designation]; Rondônia: 1 km N Cacaulândia, [10 ° 31 ′ 30 ″ S, 62 ° 48 ′ W], 168 m, (Brock, J. P.), 28 Oct 1990, 1 ♂ [FLMNH-MGCL- 265727], [dissection, SN- 20 - 34] (FLMNH); 5 km S of Cacaulândia on Linha C- 10 at Rio Pardo off B- 65, [10 ° 23 ′ 15 ″ S, 62 ° 54 ′ 53 ″ W], (Gomes, O.), 13 Mar 1984, 1 ♂ [FLMNH-MGCL- 296552], (FLMNH), 29 Aug 1993, 1 ♀ [FLMNH-MGCL- 265729; Station # 15 forest], [dissection, SN- 20 - 11] (FLMNH); Cacaulândia, 7 km E B- 65, Fazenda Rancho Grande, [10 ° 17 ′ 58 ″ S, 62 ° 52 ′ 14 ″ W], (Austin, G. T.), 19 Nov 1992, 1 ♂ [FLMNH-MGCL- 265728], (FLMNH), (Bongiolo, G.), 14 Jun 1992, 1 ♂ [FLMNH-MGCL- 265726; Station # 3 forest], (FLMNH). Ecuador: Morona-Santiago: jct. Río Mayalico- Río Santiago, Isla de las Conchas, [3 ° 2 ′ 10 ″ S, 77 ° 58 ′ 29 ″ W], 250 m, (Hall, J. P. W., Willmott, K. R., J. C. R., J. I. R), 8, 10 Aug 2015, 1 ♂ [FLMNH-MGCL- 217579], (FLMNH); hwy km 20 Mendez- Santiago rd., [2 ° 47 ′ 6 ″ S, 78 ° 15 ′ 24 ″ W], 850 m, (Perceval, M. J.), 14 Oct 1997, 1 ♀, (MIPE); km 30 Méndez-Limón rd., Río Yungantza, [2 ° 52 ′ 13 ″ S, 78 ° 21 ′ 56 ″ W], 650 m, (Hall, J. P. W.), 1 – 3 Mar 2017, 1 ♀ [FLMNH-MGCL- 281450], (FLMNH); Santiago de Mendez, [3 ° 2 ′ 11 ″ S, 78 ° 2 ′ W], (Nakahara, S.), 16 Jun 2014, 1 ♀, (FLMNH); Napo: Río Napo, hwy Puerto Napo-Ahuano rd., Chichicorrumi, [1 ° 4 ′ 11 ″ S, 77 ° 37 ′ 45 ″ W], 450 m, (Willmott, K. R., Hall, J. P. W.), 2, 9 Jul 1993, 1 ♀, (FLMNH); Orellana: Laguna Zancudococha, military trail, [0 ° 35 ′ 16 ″ S, 75 ° 28 ′ 16 ″ W], 220 m, (Aldaz, R.), 9 – 13 Jul 2017, 1 ♂ [FLMNH-MGCL- 288722], (FLMNH); Río Aguarico, Zancudococha, [0 ° 34 ′ 23 ″ S, 75 ° 26 ′ 13 ″ W], 240 m, (Willmott, K. R., J. C. R, J. I. R., Aldaz, R.), 14 Jul 2017, 1 ♀, (INABIO); Río Napo, Boca del Río Añangu, [0 ° 31 ′ 43 ″ S, 76 ° 23 ′ 41 ″ W], 220 – 300 m, (Willmott, K. R.), 27 Oct 2005, 1 ♀ [FLMNH-MGCL- 111516], (FLMNH); Shiripuno Lodge, Mirador trail, [1 ° 4 ′ 50 ″ S, 76 ° 44 ′ 42 ″ W], 350 m, (Hall, J. P. W., Willmott, K. R., J. C. R., J. I. R.), 8,9, 11 Aug 2018, 1 ♂, 1 ♀, (FLMNH); Sucumbíos: Cerro Lumbaquí Norte, [0 ° 1 ′ 42 ″ N, 77 ° 19 ′ W], 800 – 950 m, (Willmott, K. R., Hall, J. P. W.), 21 – 23 Jul 1999, 1 ♂, (FLMNH); Zamora-Chinchipe: 3 km E El Panguí, Centro Shuar Cháarip, [3 ° 38 ′ 6 ″ S, 78 ° 33 ′ 29 ″ W], 800 m, (Willmott, K. R., Hall, J. P. W.), 4 Aug 2009, 1 ♀ [FLMNH- MGCL- 145674], (FLMNH); km 11.5 Los Encuentros-Zarza, La Libertad, [3 ° 47 ′ 54 ″ S, 78 ° 36 ′ 26 ″ W], 1,250 m, (Willmott, K. R., Hall, J. P. W.), 6, 8 Aug 2009, 1 ♂ [FLMNH-MGCL- 145675], 1 ♂ [FLMNH-MGCL- 145676], (FLMNH); Zamora, ridge to west, [4 ° 4 ′ 30 ″ S, 78 ° 58 ′ 7 ″ W], 1,400 – 1,450 m, (Willmott, K. R.), 20 May 2000, 1 ♂ [dissection, KW- 14 - 008], (FLMNH). Peru: Cuzco: Pilcopata, Villa Carmen, [12 ° 54 ′ S, 71 ° 24 ′ W], 540 m, (Brock, J.), 31 Jan 2013, 1 ♀ [MUSM-LEP- 103083], (MUSM); Quincemil, Quebrada Yanaorcco, [13 ° 16 ′ S, 70 ° 47 ′ W], 900 m, (Rodríguez, M.), Feb 2010, 1 ♀ [MUSM-LEP- 103084], (MUSM); Junín: La Merced, [11 ° 3 ′ S, 75 ° 19 ′ W], 790 – 762 m, (Watkins & Tomlinson), May – Jun 1903, 1 ♀ [BMNH (E) - 1670293], (NHMUK); Loreto: Castaña, [0 ° 48 ′ S, 75 ° 14 ′ W], 150 m, (Lamas, G.), 26 Oct 1993, 1 ♂ [MUSM- LEP- 103079], (MUSM), 29 Oct 1993, 1 ♂ [MUSM-LEP- 103080], (MUSM); Lower Río Ucayali, Río Pacaya, Aug-Sep 1912, 1 ♂ [BMNH (E) - 1670291], (NHMUK); Pebas, [3 ° 19 ′ S, 71 ° 51 ′ W], 120 m, (Hahnel, P.), 1 ♀ [dissection, 9077], (MNHU), (Mathan, M. de), Dec 1879 – Mar 1880, 1 ♂ [BMNH (E) - 1670290], (NHMUK), Nov 1906, 1 ♂ [BMNH (E) - 1497733], (NHMUK); Zona Reservada Allpahuayo-Mishana, [3 ° 57 ′ 30 ″ S, 73 ° 25 ′ 30 ″ W], 170 m, (Ramírez, J. J.), 8 Aug 2001, 1 ♀ [MUSM-LEP- 103092], (MUSM); Madre de Dios: 30 km SW Puerto Maldonado, [12 ° 36 ′ S, 69 ° 11 ′ W], 200 m, (Anderson, J. J.), 18 – 23 Oct 1982, 1 ♀ [MUSM-LEP- 103087], (MUSM); Boca Río La Torre, [12 ° 50 ′ S, 69 ° 17 ′ W], 300 m, (Lamas, G.), 12 Feb 1982, 1 ♂ [MUSM-LEP- 103082], (MUSM), 26 Sep 1981, 1 ♀ [MUSM-LEP- 103085], (MUSM), 27 Sep 1981, 1 ♀ [MUSM-LEP- 103086], (MUSM); Parque Nacional del Manu, Pakitza, [11 ° 55 ′ 48 ″ S, 71 ° 15 ′ 18 ″ W], 400 m, (Lamas, G.), 13 Oct 1990, 1 ♀ [MUSM-LEP- 103089], (MUSM), 18 Oct 1990, 1 ♂ [MUSM-LEP- 103081], 1 ♀ [MUSM-LEP- 103090], (MUSM), (Rowe, W.), 3 Nov 1990, 1 ♀ [MUSM-LEP- 103088], (MUSM); Reserva Tambopata, La Colpa, [13 ° 9 ′ S, 69 ° 37 ′ W], 250 m, (Aibar, P.), 19 Oct 2000, 1 ♀ [MUSM-LEP- 103091], (MUSM); Puno: Río Tambopata, [12 ° 36 ′ S, 69 ° 11 ′ W], 270 m, 15 Jul 1979, 1 ♂ [FLMNH-MGCL- 265725], [dissection, SN- 20 - 10] (FLMNH); San Martín: near Yurimaguas, ‘ Chambireyacu’ [= Río Chambirayacu], [5 ° 54 ′ S, 76 ° 14 ′ W], 100 m, (Mathan, M. de), Jun-Aug 1885, 1 ♂ [BMNH (E) - 1670292], (NHMUK); Moyobamba, [6 ° 2 ′ S, 76 ° 58 ′ W], 855 m, Jan-Jun 1887, 1 ♀ [BMNH (E) - 1670294], (NHMUK). Country unknown: Not located: ‘ Amaz [on] S’, 1 ♀ [BMNH (E) - 1497644], (NHMUK); no data, 1 ♂, (MNHU), 1 ♂, (MNHU). Other records: Ecuador: Napo: Tena-Puyo rd., El Capricho, [1 ° 11 ′ 14 ″ S, 77 ° 49 ′ 53 ″ W], 850 m, (Neild, A.), 20 Oct 2015, 1 ♂, (photograph live specimen) (Neild, A. F. E. (20 Oct 2015, sight record, by email with photo to KRW )); Peru: Madre de Dios: Madre de Dios, [12 ° 16 ′ S, 70 ° 55 ′ W] [ID based on DNA barcode], (DEMU) (Murray and Prowell (2005 )).	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
761D1C3DFFB8FFE2FF6DFD0942CFFB85.taxon	description	(Figs. 1, 2 i and j, 3 k – n, 4 e and f, 6 e – i, 7 c – e, 8)	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
761D1C3DFFB8FFE2FF6DFD0942CFFB85.taxon	diagnosis	Systematic placement and diagnosis. The phylogenetic analysis placed Cisandina castanya n. sp. as sister to (C. lea n. comb. + C. esmeralda n. sp.) + C. philippa n. comb. & reinst. stat., although the support for this placement is moderate (Fig. 1; SH-aLRT / UFBoot = 85.9 / 81). The single barcoded individual of C. castanya n. sp. (DZ 52.561 – BC-DZ- 139, from Acre, Brazil) shows high genetic divergence, greater than 3.74 % (ranging from 3.74 to 6.54 %; mean 5.0283 % (n = 12), see Table 3) compared with the three other Cisandina n. gen. species which form a clade to which it is supported as sister, reinforcing our taxonomic hypothesis. Males of C. castanya n. sp. are readily distinguishable from other species in the genus by their small forewing length, in addition to the presence of semi-iridescent bluish scales on the DFW and DHW. Females are distinguished from other species in the genus by possessing similar semi-iridescent bluish scales on the DFW and DHW, which extend from the inner margin of DFW and reach the discocellular vein, whereas the bluish scales on the DFW are more restricted to basal area in female C. lea n. comb. and absent in C. philippa n. comb. & reinst. stat. As explained under the immediately preceding taxon, C. castanya n. sp. is sympatric with C. philippa n. comb. & reinst. stat. and species-level diagnostic characters can be found in the immature stages as well, namely head scoli appearing more developed in C. castanya n. sp. (scoli / headcapsule width ratio = 0.14; scoli length c. 0.4 mm.) compared with C. philippa n. comb. & reinst. stat. (scoli / headcapsule width ratio = 0.08; scoli length c. 0.2 mm.). Male (Fig. 2 i): n = 3). Forewing length 22 – 23 mm (mean 22.3 mm; Head: See head description for Cisandina n. gen. above. Thorax: See thorax description for Cisandina n. gen. above. Abdomen: See abdomen description for Cisandina n. gen. above, eighth sternite rather narrow and appearing as two separated patches at anterior side. Wing venation: See wing venation description for Cisandina n. gen. above. Wing shape: See wing shape description for Cisandina n. gen. above, notable features include forewing appearing less elongate due to angle between forewing outer margin and inner margin being roughly right angle. Dorsal forewing: Notable features include, ground color brownish, iridescent purple-ish lilac scales covering most of DFW except for area anterior of costal vein, apex and distal side of cells M 1, M 2, M 3, Cu 1, although this area revealing ground color narrows posteriorly (Fig. 2 i); see also DFW description for Cisandina n. gen. above. Dorsal hindwing: Notable features include, ground color similar to forewing, iridescent purplish lilac scales covering most of DHW except for area anterior of Rs and area posterior of 3 A (Fig. 2 i); see also DHW description for Cisandina n. gen. above. Ventral forewing: Notable features include, ventral bands narrow, discal band and postdiscal band parallel to each other and traversing in slightly outward diagonal direction, ocellus in cell M 1 small and may appear insignificant with pupil being invisible, additional ocellus may be present in adjacent cells (Fig. 2 i); see also VFW description for Cisandina n. gen. above. Ventral hindwing: Notable features include, grey-ish blue scales covering areas in cells Cu 1, Cu 2 and 2 A, submarginal ocelli appearing small, ocelli in cells M 1 and Cu 1 not reaching or barely reaching veins defining these cells, small ocellus may be visible at posterior end of postdiscal band in some specimens (Fig. 2 i); see also VHW description for Cisandina n. gen. above. Genitalia: Notable features include, dorsal margin of tegumen rather smoothly curved as well as anterior portion of ventral margin of tegumen appearing ‘ bent’ in lateral view, rather straight saccus in lateral view, apical process of valva narrow and curving (Fig. 3 k – n); see also male genitalia description for Cisandina n. gen. above. Female (Fig. 2 j): n = 3). Forewing length 21 – 22 mm (mean 21.7 mm, Similar to male except as follows: Notable features include, area of iridescent purplish lilac scales on DFW restricted to discal cell, basal part of cell M 3, more than half of Cu 1, most of cells Cu 2 and 2 A (Fig. 2 j); see also female wing shape and pattern description for Cisandina n. gen. above. Female genitalia: Notable features include, lamella antevaginalis appearing as rectangular ‘ thumb-like’ plate with wrinkled margin; signa short, occupying basal half of corpus bursae, length of corpus bursae similar to ductus bursae (Fig. 4 e and f); see also male genitalia description for Cisandina n. gen. above. Type material: Holotype male, with the following labels separated by double-forward slashes: // PERU, MD, Albergue Pantiacolla 400 m 12 ° 39 ′ S, 71 ° 14 ′ W 22. vi. 2019 W. Dempwolf // SN-DNA 19 - 48 (MUSM). Paratypes (10 ♂, 22 ♀): Brazil: Acre: Senador Guiomard, Reserva Catuaba, [9 ° 37 ′ S, 68 ° 18 ′ W], (Mielke & Casagrande), 31 Jan – 5 Feb 2009, 2 ♀ [DZ 52.561] (DZUP); Mato Grosso: hwy Cuiabá- Santarém, km 715, [9 ° 58 ′ 54 ″ S, 54 ° 54 ′ 20 ″ W], 300 m, (Callaghan, C. J.), 14 Jul 1978, 1 ♂ [FLMNH-MGCL- 265741], (FLMNH); Rondônia: Cacaulândia, ‘ Big Rock Trail’, 3 km E of Fazenda Rancho Grande on Linha C- 20, [10 ° 17 ′ 47 ″ S, 62 ° 50 ′ 27 ″ W], (Austin, G. T.), 21 Sep 1992, 1 ♀ [FLMNH-MGCL- 265743], (FLMNH); 12.5 km S of Cacaulândia, off B- 65, Linha C- 2.5, [10 ° 27 ′ 16 ″ S, 62 ° 53 ′ 59 ″ W], (Austin, G. T.), 12 Nov 1990, 1 ♀ [FLMNH-MGCL- 265736], (FLMNH), 9 Dec 1990, 1 ♂ [FLMNH- MGCL- 265734], (FLMNH); 15 km S of Cacaulândia, off B- 65, Linha C- 0, [10 ° 28 ′ 43 ″ S, 62 ° 53 ′ 46 ″ W], (Austin, G. T.), 22 Apr 1991, 1 ♀ [FLMNH-MGCL- 265737], (FLMNH); 3 km N Cacaulândia, dry trail off B- 65, [10 ° 30 ′ 24 ″ S, 62 ° 48 ′ W], (Smith, J.), 23 Apr 1991, 1 ♀ [FLMNH-MGCL- 296555], (FLMNH); Cacaulândia, vicinity of Fazenda Rancho Grande, [10 ° 17 ′ 58 ″ S, 62 ° 52 ′ 14 ″ W], 180 m, (Austin, G. T.), 20 Oct 1989, 1 ♀ [FLMNH-MGCL- 265739], (FLMNH), 29 Oct 1989, 1 ♂ [FLMNH-MGCL- 265730] [dissection, SN- 20 - 12] (FLMNH); 5 km S of Cacaulândia on Linha C- 10 at Rio Pardo off B- 65, [10 ° 2 ′ 3 ′ 15 ″ S, 62 ° 54 ′ 53 ″ W], (Gomes, O.), 23 Apr 1993, 1 ♀ [FLMNH-MGCL- 265738] [dissection, SN- 20 - 13], (FLMNH), 23 Jun 1996, 1 ♀ [FLMNH-MGCL- 265742], (FLMNH), 7 Aug 1994, 1 ♀ [FLMNH-MGCL- 265744], (FLMNH), 8 Jun 1996, 1 ♀ [FLMNH-MGCL- 296559], (FLMNH); Cacaulândia, 7 km E B- 65, Fazenda Rancho Grande, [10 ° 17 ′ 58 ″ S, 62 ° 52 ′ 14 ″ W], (Austin, G. T.), 16 Sep 1992, 1 ♀ [FLMNH-MGCL- 265745] [dissection, SN- 20 - 33], (FLMNH), 26 Nov 1991, 1 ♀ [FLMNH-MGCL- 265735], (FLMNH), 29 Nov 1991, 1 ♀ [FLMNH-MGCL- 265740], (FLMNH), (Austin, G. T.), 17 Jul 1995, 1 ♀ [FLMNH-MGCL- 296557], (FLMNH), 22 Jul 1994, 1 ♂ [FLMNH-MGCL- 296553] [dissection, SN- 20 - 32], (FLMNH), 3 May 1995, 1 ♀ [FLMNH-MGCL- 296558], (FLMNH), 5 May 1995, 1 ♀ [FLMNH-MGCL- 296556], (FLMNH), (Brock, J. P.), 5 Nov 1989, 1 ♂ [FLMNH-MGCL- 265733], (FLMNH), (Gomes, O.), 27 Jun 1989, 1 ♂ [FLMNH-MGCL- 265731], (FLMNH); off B- 65, Linha C- 20 at Rio Pardo, [10 ° 17 ′ 58 ″ S, 62 ° 57 ′ 5 ″ W], (Austin, G. T.), 18 Sep 1992, 1 ♂ [FLMNH-MGCL- 265732], (FLMNH). Peru: Huánuco: Lower Ucayali, Río Pachitea, [8 ° 46 ′ S, 74 ° 32 ′ W], 150 m, (Tessman, G.), 1 ♀, (MNHU); Madre de Dios: 15 km E Puerto Maldonado, [12 ° 32 ′ S, 69 ° 3 ′ W], 200 m, (Medina, M.), 6 Feb 1990, 1 ♀ [MUSM-LEP- 100320] [dissection, SN- 16 - 20], (MUSM); Parque Nacional del Manu, Pakitza, [11 ° 55 ′ 48 ″ S, 71 ° 15 ′ 18 ″ W], 340 m, (Harvey, D. J.), 27 Apr 1991, 1 ♀ [MUSM-LEP- 100319], (MUSM), (Mielke, O. H. H.), 3 Oct 1991, 1 ♂ [MUSM-LEP- 100316], (MUSM), (Robbins, R. K.), 2 Oct 1991, 1 ♂ [MUSM-LEP- 100317] [dissection, SN- 16 - 36], (MUSM); Albergue Pantiacolla, [12 º 39 ′ S, 71 º 14 ′ W], 400 – 450 m, (Lamas, G.), 29 Oct 2016, 1 ♀, (MUSM), 400 m, (Gibson, L.), 30 Oct 2018, [SN-DNA 19 - 49], 1 ♂, (MUSM). Other records: Peru: Loreto: Río Paranapura, Chambira, [5 ° 54 ′ S, 76 ° 14 ′ W], 120 m, (Razzeto, O.), 1 ♂ [MUSM-LEP- 100318], (MUSM); Río Ucayali, Contamana, [7 ° 21 ′ S, 75 ° 0 ′ W], 135 m, (Büche, M.), Sep 1999, 1 ♂ [MUSM-LEP- 100315], (MUSM). Etymology: The specific epithet is derived from the Spanish word ‘ castaña’ and the Portuguese word ‘ castanha’, both meaning ‘ chestnut’ in English and, in much of Peru, Bolivia, and Brazil, more specifically ‘ Brazil nut’; Brazil nuts are large, emergent trees (Bertholletia excelsa Humb. & Bonpl. (Lecythidaceae )) that dominate the rainforest of the southwestern Amazon basin and whose distribution somewhat overlaps with that of this new butterfly species. These trees grow in abundance and characterize the landscape at Finca Las Piedras (Madre de Dios, Peru), where the immature stages of this new species were discovered. This species-group name is a feminine noun in apposition. Distribution and natural history: This species is distributed in the southwestern Amazon basin, from central and southern Peru eastwards into Brazil (Mato Grosso) (Fig. 8). A second-instar caterpillar and the last instar was found on a herbaceous bamboo, Olyra latifolia L. (Poaceae) at Finca Las Piedras, Madre de Dios, Peru, on 20 June 2020 (voucher: 2020 - FLP-IMM- 0188) and on 14 March 2021 (voucher: 2021 - FLP-IMM- 0234), respectively, and the immature stages are described below. The instar number (second) for 2020 - FLP-IMM- 0188 was determined based on comparing its size and appearance with other euptychiine larval instars. The host plant with the caterpillar was located at an edge habitat of a terra firme forest dominated by Brazil nut trees, with high incidence of light in the understory, fostering the growth of understory grasses. Remarks. The two male individuals from Loreto, Peru (MUSM- LEP- 100315 and MUSM-LEP- 100318) are classified as this taxon in MUSM. Nevertheless, these specimens possess some phenotypic characters not observed in other individuals examined for C. castanya n. sp. such as large ventral ocelli and presence of small ocellus at the VHW inner margin (at the posterior end of the VHW postdiscal band). Thus, we decided to exclude these individuals from the type series until further data becomes available to assess its identity.	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
761D1C3DFFB6FFECFF6DFCAB430EFE48.taxon	description	(Figs. 2 k and l, 8)	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
761D1C3DFFB6FFECFF6DFCAB430EFE48.taxon	materials_examined	Taxonomy: Gustav Weymer (1911) described Euptychia fida in his ‘ Saturnus group’ of Euptychia, based on an unspecified number of individuals from [Río] Songo [sic] and ‘ Corvico’ (= Coroico), both situated in La Paz, Bolivia. The original description compared E. fida with E. vesta Butler, 1867 (= Graphita griphe (C. Felder & R. Felder, 1867); see Nakahara et al. (2016) for further details regarding its synonymy and systematic placement), which is a distantly related taxon phenotypically resembling E. fida. However, as mentioned by Weymer (1911), male specimens of E. fida can easily be distinguished from G. griphe by the absence of androconial scales (termed ‘ raised scale-streak’ by Weymer) on the DFW. The illustration of E. fida associated with the description (on pl. 46, fig. f) and showing its dorsal surface clearly indicates the absence of DFW androconial scales, which are visible on the painting of E. vesta on the same plate. Another wing pattern character that is informative in terms of identifying these two taxa is the presence / absence of the DHW ocellus in cell Cu 2, which is also vaguely discussed by Weymer. As reflected in the illustration provided by him, this DHW ocellus is present in G. griphe and absent in E. fida, regardless of the sex. Despite referring to some ventral wing pattern differences between these two species, Weymer only provided a drawing of the ventral surface for E. vesta, and not for E. fida. Given this information, four syntypes were located, three specimens at MNHU and a single male at ZSM, all from Songo [sic], Bolivia. We were unable to locate any syntype (s) from Coroico. Three syntypes in the MNHU were mentioned and discussed in Nakahara et al. (2018 a) in association with the description of Euptychoides sanmarcos, although due to an unfortunate oversight, the specimen housed at ZSM was omitted. Nakahara et al. (2018 a) noted wing pattern differences between Ecuadorian and Bolivian specimens, including the possibility of these two populations representing two species. All examined Ecuadorian specimens (all from Zamora-Chinchipe; n = 5) possess a rather straight VHW postdiscal band, whereas this band is jagged in Bolivian specimens (n = 5, including three syntypes). The single known Peruvian specimen, a male from Ucayali [doubtful locality – see below], has a straight VHW postdiscal band, suggesting that this specimen represents the same taxon as those individuals from southern Ecuador. The COI sequence of this Peruvian specimen (LEP- 58115) exhibits a genetic divergence of 2.46 – 7 % compared with those two sequenced Ecuadorian specimens (LEP- 16705 and LEP- 10686). We also lack DNA data for Bolivian specimens, thus preventing further assessment based on molecular data and our judgement regarding taxonomic status of E. fida from southern Ecuador to central Peru must remain tentative. All known Bolivian specimens are phenotypically distinguishable from individuals collected in southern Ecuador and central Peru, suggesting that the latter population should be regarded as a distinct taxon, either as a species or subspecies. Apart from the lack of molecular data for Bolivian specimens, another piece of evidence needed to assess the taxonomic status for its neighboring northern population is material from other places in Peru. Without such data, it is impossible to determine whether the observed wing pattern differences are broadly clinal, or even potentially partially sympatric. We therefore follow Nakahara et al. ’ s (2019 a) approach of generating a parsimonious hypothesis for a single species, namely by considering it of subspecific rank and waiting for further evidence until we can test this null hypothesis. Additionally, in order to settle the nomenclature of this species, we here designate a lectotype for Euptychia fida based on the dissected female (male identification on the label apparently erroneous) specimen from Río Songo housed at the MNHU with the following labels separated by double-forward slashes (lectotype designation): // Songo stgr / Salona stgr. // LECTOTYPE ♂ Euptychia fida Weymer designated by: Lee D. Miller 1989 // Rio Songo (1,200 m) Bolivia (Yungas) 1895 – 1896. Garlepp // genitalia vial M- 9051 ♂ Lee D. Miller //. Despite having a lectotype label attached, L. D. Miller never published these designations as already noted by Nakahara et al. (2015). Specimens examined (5 ♂, 1 ♀): Bolivia: La Paz: Río Zongo, [16 ° 3 ′ 40 ″ S, 68 ° 1 ′ 2 ″ W], 1,200 m, (Garlepp), 1895 – 1896, 1 ♀ [dissection, 9051; ‘ Songa Stgr. / Salona Stgr. // Rio Songo (1,200 m) Bolivia (Yungas) 1895 – 1896. Garlepp // LECTOTYPE ♂ Euptychia fida Weymer designated by Lee D. Miller 1989 // genitalia vial M- 9051 ♂ Lee D. Miller’], (MNHU); Yungas, [16 ° 17 ′ 13 ″ S, 66 ° 48 ′ 33 ″ W], 1,200 m, 1 ♂ [BMNH (E) - 1267787; ‘ Salona Stgr. Bolivia // Yungas // Fruhstorfer Coll. B. M. 1937 - 285 // BMNH (E) 1267787 ’], (NHMUK); Zongo, [16 ° 5 ′ 46 ″ S, 68 ° 3 ′ 9 ″ W], (Garlepp), 2 ♂ [‘ Songo Bol. Garl. // LECTO-PARATYPE ♂ Euptychia fida Weymer, des. Lee D. Miller 1989 ’], (MNHU), 1 ♂ [BMNH (E) - 1267786; dissection, B. M. (N. H.) Rhopalocera vial No. 9531], (NHMUK), 1 ♂ [dissection, SA 19; ‘ Präparat Nr. SA 19 Zoolog. Staatssammlung München’ // ’ Songo Bol. Garl. ’ // ’ Para-Typus Euptychia fida Weym. ’ // ’ fida Weym. ’ // ’ Original! ’], (ZSM). Distribution and natural history: The nominate race of C. fida n. comb. is known to date only from Bolivia (Fig. 8)	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
761D1C3DFFB7FFECFF6DFDD1456AFE85.taxon	description	(Figs. 1, 2 m and n, 3 p and q, 4 h and i, 8) Description and Diagnosis: Male: Forewing length 23.5 – 24.5 mm (n = 2): Differs from the nominotypical subspecies in the following respect: VHW postdiscal band rather straight; this band traverses distally after passing Cu 2 and bends back half way between Cu 2 and 2 A (Fig. 2 m). Female: Forewing length 24.5 – 25.0 mm (n = 2): Differs from the nominotypical subspecies by the same ventral forewing character provided for the male (Fig. 2 n); differs from the male by having more broad and rounded wing shape; two small ocelli present in VFW cells M 2 and M 3 in one female (from Zamora, ridge to west, 13 Jan 2002), whereas the ocelli are absent in these cells in the other examined female (FLMNH-MGCL- 145742). Type material. Holotype male, with the following labels separated by double-forward slashes: // ECUADOR: Zamora-Chinchipe Zamora, ridge to west, 1,400 – 1,450 m 4 ° 4.50 ′ S, 78 ° 58.12 ′ W 18. v. 2000, K. Willmott // DNA voucher LEP- 14657 // Genitalic vial SN- 20 - 42 S. Nakahara // (FLMNH, to be deposited in INABIO). Paratypes (2 ♂, 2 ♀): Ecuador: Zamora-Chinchipe: km 10 Los Encuentros-El Panguí, ridge E San Roque, [3 ° 42 ′ 11 ″ S, 78 ° 35 ′ 36 ″ W], 1,050 m, (Willmott, K. R., Hall, J. P. W.), 4 Aug 2009, 1 ♀ [FLMNH-MGCL- 145742] [dissection, SN- 15 - 180], (FLMNH); km 14 Yacuambí-Saraguro rd., Juyapa, [3 ° 34 ′ 48 ″ S, 78 ° 57 ′ 2 ″ W], 1,740 m, (Willmott, K. R., J. I. R., J. C. R.), 21 Jun 2013, 1 ♂ [FLMNH-MGCL- 157449], (FLMNH); Zamora, ridge to west, [4 ° 4 ′ 30 ″ S, 78 ° 58 ′ 7 ″ W], 1,400 – 1,450 m, (Willmott, K. R.), 13 Jan 2002, 1 ♀, (FLMNH), 20 May 2000, 1 ♂, (FLMNH). Other records: Peru: Ucayali: Río Aguaytía, Previsto, [9 ° 3 ′ S, 75 ° 38 ′ W], 420 – 500 m, Oct 2006, 1 ♂ [FLMNH- MGCL- 281611; MGCL Accession # 2016 - 40 E. C. Knudson / Bordelon], (FLMNH) [this locality is doubtful, and the specimen probably originated further west, in Huánuco department]. Etymology. This species-group name is a feminine Latin adjective meaning ‘ straight, not curved’, in reference to the smoother VHW postdiscal band compared with the nominate race. Distribution and natural history. This subspecies is known from southern Ecuador (Zamora-Chinchipe) and central Peru (Fig. 8). In Ecuador, it occurs in cloud forest from 1,050 to 1,740 m, where it is rare and only known from ridge top localities. Males were attracted to rotting fish in canopy bait traps or to similar bait placed on leaves 1 m above the ground, and one female was found flying along the edge of a road through a secondary forest with abundant bamboo during the middle of the day.	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
761D1C3DFFB7FFEFFCCDFE8B4095FB83.taxon	description	(Figs. 1, 2 o and p, 3 r – t, 4 i – k, 8)	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
761D1C3DFFB7FFEFFCCDFE8B4095FB83.taxon	diagnosis	Systematic placement and diagnosis: As discussed in Nakahara et al. (2018 a), this species is closely related to C. fida n. comb. (Fig. 1). Cisandina sanmarcos n. comb. forms a strongly supported clade with C. fida n. comb. in the present study (Fig. 1; SH-aLRT / UFBoot = 100 / 100), which is also strongly supported as sister to four species discussed above (Fig. 1; SH-aLRT / UFBoot = 96.9 / 95). Nevertheless, the placement of C. sanmarcos n. comb. results in C. fida n. comb. being paraphyletic, as mentioned under the corresponding section of that species, and further discussion and justification as to its specific status can be found below. See corresponding section of C. fida n. comb. for diagnostic characters to distinguish these two taxa. Taxonomy: Nakahara et al. (2018 a) described Euptychoides sanmarcos based on three males and eight females, including the male holotype. As mentioned in the original description, we were aware of the fact that the generic classification regarding this species would need subsequent revision, albeit due to its distinctiveness and the goal of Nakahara et al. (2018 a) to describe ‘ miscellaneous taxa’ to further future investigation of their relationships, we went ahead and named it prior to its ultimate generic assessment. The sole reason for describing this species under Euptychoides was because of its sister relationship with ‘ Euptychoides ’ fida, although knowing that ‘ Euptychoides ’ fida was distantly related to Euptychia saturnus Butler, 1867, the type species of Euptychoides. Nevertheless, the holotype male of Cisandina sanmarcos n. comb. from Madre de Dios, Peru figured in the original description (Fig. 10), evidently possesses characters discussed in the original description that separate the species from its sister species, Cisandina fida n. comb. Note that the holotype of E. sanmarcos was deposited at MUSM in October 2019 subsequent to its description, as indicated in the original description. Additionally, the female specimen housed at the MZUJ listed below was not included in the type series as this individual was not known to the authors during the course of preparing Nakahara et al. (2018 a). Specimens examined (3 ♂, 10 ♀): Bolivia: La Paz: Río Zongo, [16 ° 3 ′ 40 ″ S, 68 ° 1 ′ 2 ″ W], 1,200 m, (Garlepp), 1895 – 1896, 1 ♂ [dissection, M- 9141 Lee D. Miller], (MNHU). Peru: Cuzco: Cosñipata Valley, Quebrada Quitacalzón, [13 ° 1 ′ 35 ″ S, 71 ° 29 ′ 57 ″ W], 1,050 m, (Harris, B.), 12 Aug 2009, 1 ♀ [MUSM-LEP- 103661; dissection, genitalic vial SN- 14 - 18 MUSM], (MUSM), (Kinyon, S.), 25 Sep 2011, 1 ♀ [MUSM-LEP- 103662], (MUSM); Cosñipata Valley, Quebrada Quitacalzón, [13 ° 1 ′ 35 ″ S, 71 ° 29 ′ 57 ″ W], 1,100 m, (Gibson, L.), 10 May 2012, 1 ♀ [MUSM-LEP- 103663], (MUSM), (Kinyon, S.), 22 Sep 2014, 1 ♀ [MUSM-LEP- 103666], (MUSM), (Lamas, G.), 22 Sep 2014, 1 ♀ [MUSM-LEP- 103664; dissection, genitalic vial SN- 16 - 17 MUSM], (MUSM), (Lamas, G.), 23 Oct 2010, 1 ♀ [MUSM-LEP- 103667], (MUSM); Gallito de las Rocas conservation area [13 ° 04.513 ′ S 71 ° 25.133 ′ W], 1,000 – 1,100 m, (G. Gallice), 9 Jul 2019, 1 ♀ [2019 - GR- 0077], (ASA); Huánuco: Cordillera del Sira, [9 ° 25 ′ S, 74 ° 45 ′ W], 1,380 m, (Exp. Universidad Viena), Sep 1987 - Aug 1988, 1 ♀ [MUSM-LEP- 103660], (MUSM); Madre de Dios: Cerro Pantiacolla, E Slope nr. summit, ca. 4 - km ENE Shintuya, [12 ° 38 ′ 19 ″ S, 71 ° 17 ′ 19 ″ W], 960 – 1,030 m, (Douglass, J. F.), 25 Jul 1980, 1 ♂ [dissection, SN- 14 - 149; HT sanmarcos], (MUSM); Puno: Tambopata – Candamo, Río Távara, [13 ° 26 ′ S, 69 ° 38 ′ W], 450 – 1,050 m, (Baynes, H.), 1 Aug 1995, 1 ♂ [MUSM-LEP- 103659], (MUSM), (Grados, J.), 9 Aug 1995, 1 ♀ [MUSM-LEP- 103665], (MUSM); San Martín: Jorge Chávez, [5 ° 41 ′ S, 77 ° 40 ′ W], 1,200 – 1,400 m, (Calderón, B.), 2003, 1 ♀, (MZUJ). Distribution and natural history: Cisandina sanmarcos n. comb. ranges from northern Peru (San Martín department) to La Paz, Bolivia (Fig. 8). All known sites are along the slopes of the eastern Andes in an altitudinal zone between 960 and 1,380 m. It is worth mentioning that an additional specimen of this species was collected subsequent to its description by GG at Gallito de las Rocas conservation area, Cosñipata Valley, Cuzco, Peru (13 ° 04.513 ′ S, 71 ° 25.133 ′ W) in July 2019, bringing the total number of known specimens of C. sanmarcos n. comb. to 11. GG sprayed the understory plants along the ridge with urine. This particular individual was patrolling, but it is unclear whether it was attracted to the urine or not. The forest along the ridge was primary, with remarkably extensive patches of bamboo (probably Guadua sp.) on the slopes that had most likely colonized landslides. Nothing otherwise notable was recorded in terms of its behavior.	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
761D1C3DFFB4FFEFFF6DFB844229FC80.taxon	description	(Figs. 1, 2 q and r, 3 u – w, 4 l and m, 8)	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
761D1C3DFFB4FFEFFF6DFB844229FC80.taxon	diagnosis	Systematic placement and diagnosis: Cisandina trinitensis n. comb. is moderately to strongly supported as sister to the remainder of Cisandina n. gen. (Fig. 1; SH-aLRT / UFBoot = 91.9 / 92), with three sequenced individuals (LCB 251, LCB 356, MB _ 1708 _ 15) forming a well-supported clade (Fig. 1; SH-aLRT / UFBoot = 100 / 100). Cisandina trinitensis n. comb. is readily distinguished from other species in the genus by the lack of an ocellus in VHW cell Cu 2, coupled with the presence of four submarginal ocelli on the VFW (in cells M 1, M 2, M 3, and Cu 1). However, it must be noted that some ocelli on the VFW may appear as faint or only a trace, and with the limited number of specimens examined this character may prove unreliable. Taxonomy: Brévignon and Benmesbah (2012) described Magneuptychia trinitensis based on two males and two females, including the male holotype, all collected in French Guiana. The holotype was examined prior to deposition in MOBE, and it was also figured in the original description (pl. 3, Figs. 1 and 2), where the aforementioned characters are visible. We also incorporated sequences from the holotype in our molecular phylogeny (Fig. 1; LCB 251), thus leaving no doubt as to its rather distinctive identity, which is also indicated as being sister to the rest of the genus according to the maximum likelihood tree (Fig. 1). Distribution and natural history: Cisandina trinitensis n. comb. is so far known from a handful of sites all situated in French Guiana (Fig. 8). Specimens examined (3 ♂, 2 ♀): French Guiana: Cayenne: Montagne de la Trinité, [4 ° 36 ′ 2 ″ N, 53 ° 24 ′ 43 ″ W], (Benmesbah, M.), 10 Nov 2010, 1 ♀ [AT trinitensis], (LBCB), (Hermier, B.), 30 Oct 2008, 1 ♀ [PT trinitensis], (LBCB); Nouragues Station, [4 ° 5 ′ N, 52 ° 41 ′ W], (Benmesbah, M.), 6 Sep 2010, 1 ♂, (MOBE); St-Laurent du Maroni: Maripasoula, Antecume-Pata, ‘ filet’, [3 ° 17 ′ 53 ″ N, 54 ° 4 ′ 16 ″ W], (Benmesbah, M.), 13 Mar 2012, 1 ♂ [HT trinitensis], (MOBE); Not located: ‘ French Guiana’, 1 ♂ [PT trinitensis], (MOBE). Other records: French Guiana: Cayenne: Cayenne, [4 ° 56 ′ N, 52 ° 20 ′ W], 1 ♂, 1 ♀, (NHMUK) (D’Abrera (188: 780, ‘ Euptychia? sp. ’ )); St-Laurent du Maroni: Maripasoula, Antecume- Pata, ‘ filet’, [3 ° 17 ′ 53 ″ N, 54 ° 4 ′ 16 ″ W], (Benmesbah, M.), 13 Mar 2012, 1 ♀ [PT trinitensis], (MOBE) (Brévignon and Benmesbah (2012 )).	en	Nakahara, Shinichi, Rodríguez-Melgarejo, Maryzender, Kleckner, Kaylin, Corahua-Espinoza, Thalia, Tejeira, Rafael, Espeland, Marianne, Casagrande, Mirna M., Barbosa, Eduardo P., See, Joseph, Gallice, Geoffrey, Lamas, Gerardo (2022): Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 6 (1): 1-30, DOI: 10.1093/isd/ixab028
