Cisandina fida directa Nakahara & Willmott, 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

Cisandina fida directa Nakahara & Willmott , New Subspecies

( Figs. 1 , 2m and n , 3p and q , 4h 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 2A ( Fig. 2m ).

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. 2n ); 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.

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.