Cisandina lea ( Cramer, 1777 ) Nakahara & Rodríguez-Melgarejo & Kleckner & Corahua-Espinoza & Tejeira & Espeland & Casagrande & Barbosa & See & Gallice & Lamas, 2022

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), pp. 1-30 : 12-18

Cisandina lea ( Cramer, 1777) , New Combination

( Figs. 1 , 2a–d , 3a–d , 4a and b , 5, 8 )

Papilio lea Cramer, 1777: 87 , pl. 151, figs. C, D. Lectotype, designated herein.

= Papilio junia Cramer, 1780: 9 , pl. 292, figs. D, E. Lectotype, designated herein.

Satyrus lea: Godart [1824] : 464, 492.

Satyrus junia: Godart [1824] : 492.

Euptychia lea: Westwood 1851: 373 , Butler 1868: 29, Kirby 1871: 52, Butler 1877: 121, Weymer 1911: 216, pl. 48, fig. f, Gaede 1931: 452, D’Abrera 1988: 768–769, figs.

Euptychia junia: Westwood 1851: 373 , Butler 1868: 30, Kirby 1871: 52, Butler 1877: 121, Kirby 1879: 135, Forster 1964: 128, Lamas 2004: 220.

Magneuptychia lea: Forster 1964: 128 , Lamas 2004: 220, Beccaloni et al. 2008: 335.

Euptichia [sic] junia: Geyer 1832: 12 , pl. [109], figs. 627, 628.

Euptychia lea f. junia: Weymer 1911: 216 .

Euptychia lea var. junia: Gaede 1931: 452 .

Cissia junia: Singer and Ehrlich 1993: 251 , fig. 1.

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 2A (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 7th and 8th 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. (2018a).

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. (2019a) 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.), 4th Aug., 1 ♀ [BMNH(E)-1497683], (NHMUK), 4th 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).

Figures

Fig. 1. Maximum likelihood tree (LnL = −11380.331) inferred in IQ-TREE v2.0.5, showing monophyly of Cisandina n. gen., as well as relationships of taxa within these genera. Numbers beside branches are SH-aLRT/UFBoot values. HT denotes holotype specimens.

Fig. 2. Cisandina n. gen. adults,C. lea n. comb.: (a) male lectotype of Papilio lea in dorsal view;(b) male lectotype of Papilio lea in ventral view (associated labels to the right); (c) female lectotype of Papilo junia in dorsal view;(d) female lectotype of Papilo junia in ventral view (associated labels to the right);C. esmeralda n. sp., e) male holotype in dorsal view (left), ventral view (right); (f) female paratype in dorsal view (left),ventral view (right); C. philippa n. comb. & reinst. stat.: (g) male indivdual in dorsal view (left), ventral view (right) (MUSM-LEP-103081); (h) female individual in dorsal view (left), ventral view (right) (MUSM-LEP-103092); C. castanya n. sp.: (i) male holotype in dorsal view (left), ventral view (right); (j) female paratype in dorsal view (left), ventral view (right) (BC-DZ-139); C. fida fida n. comb.: (k) male paralectotype in dorsal view (left), ventral view (right); (l) female lectotype in dorsal view (left), ventral view (right); C. fida directa n. ssp.: (m) male holotype in dorsal view (left), ventral view (right); (n) female paratype in dorsal view (left), ventral view (right) (DNA voucher LEP-14650, but sequence not obtained); C. sanmarcos n. comb.: (o) male holotype in dorsal view (left), ventral view (right); (p) female paratype in dorsal view (left), ventral view (right) (MUSM-LEP 103661);C. trinitesis n. comb.: (q) male holotype in dorsal view (left), ventral view (right);(r) female allotype in dorsal view (left),ventral view (right). Scale bar = 10 mm.

Fig. 3. Cisandina n. gen. male genitalia, C. lea n. comb.: (a) lateral view of genitalic capsule without phallus;(b) posterior view of juxta;(c) lateral view of phallus; (d) posterior view of genitalic capsule indicating the reduced appendices angulares ([a to c] based on dissection KW-14-18, [d] based on dissection SN-20-16); C. esmeralda n. sp.: (e) lateral view of genitalic capsule without phallus;(f) posterior view of juxta; (g) lateral view of phallus (e to g) based on dissection SN-20- 14); C. philippa n. comb. & reinst. stat.: (h) lateral view of genitalic capsule with phallus; (i) posterior view of juxta; (j) lateral view of phallus ([h to j] based on dissection SN-20-34); C. castanya n. sp.: (k) lateral view of genitalic capsule without phallus; (l) posterior view of juxta; (m) lateral view of phallus; (n) vesica everted to visualize cornuti ([k to m] based on dissection SN-20-12, [n] based on SN-20-32); C. fida n. comb.: (o) lateral view of genitalic capsule without phallus; (p) posterior view of juxta; (q) lateral view of phallus ([o to q] based on dissection SN-20-40); C. sanmarcos n. comb.: (r) lateral view of genitalic capsule with phallus; (s) posterior view of juxta; (t) lateral view of phallus ([r to t] based on dissection SN-14-149); C. trinitesis n. comb.: (u) lateral view of genitalic capsule without phallus; (v) posterior view of juxta; (w) lateral view of phallus ([u to w] based on dissection SN-20-84). Scale bar = 1 mm. Drawings of C. sanmarcos n. comb. are reproduced from Nakahara et al. (2018a).

Fig. 4. Cisandina n. gen. female genitalia, C. lea n. comb.: (a) dorsal view of genitalia with intersegmental membrane of seventh and eighth abdominal segments folded; illustration showing arrangement of signa below (not to scale); (b) ventral view of lamella antevaginalis with intersegmental membrane of 7th and 8th abdominal segments expanded ([a and b] based on dissection SN-20-53); C. philippa n. comb. & reinst. stat.: (c) dorsal view of genitalia with intersegmental membrane of seventh and eighth abdominal segments expanded; illustration showing arrangement of signa to right (not to scale); (d) ventral view of lamella antevaginalis with intersegmental membrane of seventh and eighth abdominal segments expanded ([c and d] based on dissection SN-20-11); C. castanya n. sp.: (e) dorsal view of genitalia with intersegmental membrane of seventh and eighth abdominal segments folded; (f) ventral view of lamella antevaginalis with intersegmental membrane of seventh and eighth abdominal segments expanded ([e and f] based on dissection SN-20-13); C. fida n. comb.: (g) dorsal view of genitalia with intersegmental membrane of seventh and eighth abdominal segments folded; (h) ventral view of lamella antevaginalis with intersegmental membrane of 7th and 8th abdominal segments expanded ([g and h] based on dissection SN-15-180);C. sanmarcos n. comb.: (i) female genitalia

Fig. 8. Distribution map for Cisandina n. gen. taxa discussed herein.