Cisandina fida fida ( Weymer, 1911 ) 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 : 25-26

Cisandina fida fida ( Weymer, 1911) , New Combination

( Figs. 2k and l , 8 )

Euptychia fida Weymer, 1911: 196 , pl. 46, fig. f. Lectotype, designated herein.

Euptychia fida: Gaede 1931: 446 .

Euptychoides fida: Forster 1964: 98 , fig. 87; Lamas 2004: 219.

Systematic placement and diagnosis: According to our molecular data, Cisandina fida n. comb. is paraphyletic, although the sister relationship between Peruvian C. fida n. comb. (LEP-58115) and C. sanmarcos n. comb. (KW-15-025) is poorly supported ( Fig. 1 ; SH-aLRT/ UFBoot=54.4/64). Discussion of this paraphyly is further developed in the ‘discussion’ section below. As mentioned in Nakahara et al. (2018a), these two species can be easily distinguished based on the following phenotypic characters: the nominate phenotype of Cisandina fida n. comb. has a jagged VHW postdiscal band (but see also diagnostic characters for the following taxon), whereas this band is rather straight in C. sanmarcos n. comb.; C. fida n. comb. possesses whitish coloration distal to the VHW postdiscal band, whereas this whitish coloration is absent in C. sanmarcos n. comb. and all other species discussed herein. Nakahara et al. (2018a) mentioned the length of VFW postdiscal band as a diagnostic character to distinguish these two species, but examination of additional Ecuadorian C. fida n. comb. revealed some individuals (e.g., FLMNH# 145742) in which the VFW postdiscal band terminates at 2A, as in C. sanmarcos n. comb.

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 Cu2, 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. (2018a) in association with the description of Euptychoides sanmarcos , although due to an unfortunate oversight, the specimen housed at ZSM was omitted. Nakahara et al. (2018a) 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 (2019a) 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, SA19; ‘ 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 )

Figures

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

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.