Cisandina philippa ( Butler, 1867 ) 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 : 20-22

Cisandina philippa ( Butler, 1867) , New Combination, Reinstated Status

( Figs. 1 , 2g and h , 3h–j , 4c and d , 6a and b , 7a and

b, 8 )

Euptychia philippa Butler, 1867: 485 . Lectotype, designated herein; Butler 1868: 30, Kirby 1871: 52, Butler 1877: 121; Weymer 1911: 216, Riley and Gabriel 1924: 46, D’Abrera 1988: 768–769, figs.

Euptychia lea f. philippa: Weymer 1911: 216 .

Euptychia lea var. philippa: Gaede 1931: 452 .

Euptychia batesii f. tersa Weymer 1911: 214, pl. 49, fig. a, Lamas 2004: 220. Lectotype, designated herein.

Magneuptychia lea philippa: Lamas 2004: 220 .

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 DNA99-022 (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. //.

Euptychia batesii f. tersa was described by Gustav Weymer in Seitz’s Macrolepidoptera of the World. The original description did not specify the number of specimen(s) he examined nor the sex, in addition to not providing any information on its locality. Despite being unrelated to Euptychia batesii Butler, 1867 (currently regarded as a subspecies of Magneutpychia harpyia (C. Felder & R. Felder, 1867) , according to Lamas (2004)), Weymer described this taxon as a form of Euptychia batesii and also compared it with a close relative of that species, Euptychia analis Godman, 1905 . The original description noted that the VHW ocelli in cells M 2 and M 3 were formed of a circular ring with a central pupil, rather than being silver spots as in Euptychia batesii (= Neonympha harpyia ) and Euptychia analis . Also, Weymer pointed out the presence/absence of the VHW ocellus in cell Cu 2 as his justification for erecting this new ‘form’.Nevertheless, these phenotypes discussed by Weymer are often considered as informative characters at euptychiine species-level classification, and it is unclear why he considered tersa and batesii as conspecific. The illustration of the ventral surface associated with the original description (pl. 49, fig. a) does indeed show the phenotypic features described by Weymer, as well as the presence of the VFW ocellus in cell Cu 1, which is apparently an unusual character for species discussed in this article. Lamas (2004) considered Euptychia batesii f. tersa as a junior subjective synonym of E. philippa . The female syntype, which is most likely the specimen on which Weymer based his illustration, given the presence of a VFW ocellus in cell Cu 1, is housed at SMT and figured in Warren et al. (2017). We here designate this female specimen, with the following labels separated by double-forward slashes, as the lectotype of E. batesii f. tersa in order to settle its nomenclature, and follow Lamas (2004) in regarding this taxon as a junior subjective synonym of E. philippa (lectotype designation): //GART specimen ID: 02498 Exemplar + Etiketten dokumentiert specimen + label data documented 2003// LECTOTYPUS // LECTOTYPE ♀ Euptychia batesii f. tersa Weymer by G. Lamas ‘91// Stauding.& Bang-Haas Dresden, Ankauf 1961// Staatl. Museum für Tierkunde Dresden// tersa Weym.// Spec.// Original?//.

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. Rh3178. 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.Miller1989//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)) .

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. 6. Cisandina n. gen. immature stages, C. philippa n. comb. & reinst. stat. based on 2020-FLP-IMM-0336:(a) penultimate instar in dorso-lateral view; (b) last instar in dorso-lateral view;(c) host plant of C. philippa n. comb. & reinst. stat., Taquara micrantha (Poaceae) in situ; (d) inflorescence material of (c); C. castanya n. sp. based on 2020-FLP-IMM-0188:(e) second instar in lateral view; (f) third instar in lateral view; (g) fourth instar in lateral view; (h) pupa in lateral view;(i) pupa in ventral view; (j) host plant of C. castanya n. sp., Olyra latifolia (Poaceae) in situ. Scale bar = 1 mm for a, b, e-g; 5 mm for h, i.

Fig. 7. Cisandina philippa n. comb. & reinst. stat. head capsule illustration based on 2020-FLP-IMM-0336: (a) penultimate instar in frontal view; (b) last instar in frontal view. Cisandina castanya n. sp. head capsule illustrations based on 2020-FLP-IMM-0188: c) second instar in frontal view; (d) third instar in frontal view; (e) fourth (last) instar in frontal view. Scale bar = 1 mm.

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