Cisandina sanmarcos ( Nakahara & Lamas, 2018) 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 : 26-27

Cisandina sanmarcos ( Nakahara & Lamas, 2018) , New Combination

( Figs. 1 , 2o and p , 3r–t , 4i–k , 8 )

Magneuptychia sp. n.: Lamas & Grados [1997]: 58 Magneuptychia n. sp. Lamas, MS: Lamas 2004: 220 Euptychoides sanmarcos Nakahara & Lamas , in Nakahara et al. 2018a: 12–14, figs. 10–11, 16.

Systematic placement and diagnosis: As discussed in Nakahara et al. (2018a), 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. (2018a) 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. (2018a) 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. (2018a).

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.

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.