Cisandina, 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 : 5-12

Cisandina n. gen.

This description is intended to serve as a template description for taxa described herein, thus it accommodates both inter- and intraspecific characters without the need for a separate ‘variation’ section.

Male: Forewing length: 22–26 mm (mean: 23.8 mm; n = 13)

Head: Eyes brown with lightly colored hair-like setae, white scales at base; frons dark brownish to blackish, covered with greyish and whitish scales, as well as lightly colored elongate hair-like setae; first

20 0.0326

19 0.0326

18 0.0128 0.0199

17 0.134 0.1015

16 0.1167 0.0842

15 0.1398 0.1073

14 0.118 0.0855

13 0.1214 0.0889

12 0.1291 0.0966

11 0.117 0.0845

10 0.1168 0.0843

9 0.1445 0.112

8 0.1447 0.1122

7 0.1445 0.112

6 0.1243 0.0918

5 0.1325 0.1

4 0.1298 0.0973

3 0.1384 0.1058

2 0.1492 0.1167

1 0.1385 0.1059

_

Guiana

_ Guiana

Cisandina French _

Continued

1708

_ 15 - _

trinitensis _ 356

Cisandina _ French

trinitensis

_

3

.

MB- LCB

Table 19.. 20

segment of labial palpi short, covered with white scales and white hair-like scales and black hair-like scales, second segment about twice as long as eye depth and covered with white scales and white hair-like scales laterally and dorsally, in addition to brownish elongated scales, ventrally adorned with brownish hair-like scales and whitish hair-like scales longer than segment width, third segment apparently roughly one-third of second segment in length (although this can be somewhat variable), and covered with brownish scales dorsally and ventrally, with white scales laterally; antennae approximately two-fifths of forewing length, with ca. 38–39 antennomers (n = 4), scape rounded, about as twice as long as pedicel and wider, flagellomeres orangish, distal few flagellomeres appearing dark, covered with greyish scales with whitish scales visible on each side at base of each flagellomeres, these scales more apparent on basal flagellomeres, distal 12–13 flagellomeres composing club.

Thorax: Brownish, dorsally with greyish scales and scales with iridescent coloration, in addition to lightly colored long hair-like scales; laterally and ventrally brownish, scattered with greyish scales and some lightly colored scales, in addition to long hair-like scales; foreleg with whitish long hair-like scales and brownish long hair-like scales, femur, tibia and tarsus similar in length, tarsus not divided into subsegments; pterothoracic legs ventrally appearing lighter compared with more greyish or darker dorsal surface, tibia with two longitudinal rows of spines on ventral surface, as well as spines present laterally, tarsus with three longitudinal rows of spines ventrally until distal end of first tarsomer, number of rows increasing to four from distal end of first tarsomer onwards, pair of tibial spurs, equal in length, present at distal end of tibia.

Wing venation: Basal half of forewing subcostal vein swollen; base of cubitus swollen; forewing recurrent vein absent; discocellular vein m 1 -m 2 (i.e., dcs) curved basally, m 2 -m 3 (i.e., dcm) appearing rather straight; hindwing humeral vein present, exiting from origin of Rs and curving; origin of M 2 slightly towards M 1 than M 3 ( Fig. 5).

Wing shape: Forewing variably sub-triangular, apex rounded, costal margin slightly convex, outer margin somewhat variable but almost straight, inner margin almost straight; hindwing rounded, appearing slightly elongate (apparent in C. lea n. comb. and C. esmeralda n. sp. due to their rather elongated forewing shape), apex angular, costal margin slightly convex, angled inwards at base, outer margin slightly undulating, inner margin slightly curved inwards near tornus, anal lobe convex, slightly rounded ( Fig. 2 ).

Dorsal forewing: Ground color brownish, varying from being rather uniformly brownish to covered with iridescent to semi-iridescent scales ( Fig. 2 ), submarginal and marginal bands occasionally visible as darker bands.

Dorsal hindwing: Ground color similar to forewing, varying from being rather uniformly brownish to covered with iridescent to semiiridescent scales ( Fig. 2 ); submarginal and marginal bands occasionally visible as darker bands.

Ventral forewing: Ground color variable, from greyish brown to more brownish, may or may not be covered variably with iridescent scales ( Fig. 2 ); discal band extending from radial vein, crossing discal cell, often passing just basal of origin of Cu 2, fading and terminating in cell Cu 2; concolorous postdiscal band extending from radial vein towards inner margin, terminating at 2A if not already faded in Cu

2

, more or less parallel to discal band albeit they may appear close to each other at posterior end; umbra appearing as undefined dark brownish shading extending from area near VFW ocellus (in cell M 1) to cell Cu 2, if not terminating earlier; submarginal band, appearing darker compared with discal and postdiscal band, often narrower than previous two bands, extending from apex towards tornus, rather smooth although jaggedness can be somewhat variable, appearing more or less evenly broad perhaps except for anterior and posterior end; concolorous marginal band, not jagged, narrower than submarginal band, traversing along marginal area from apex to tornus; fringe greyish; small ocellus in cell M 1, often possessing black central area and single white pupil with pale ring but may lack one or two of these elements, additional ocellus or ocelli may appear in cells R 5, M 2, M 3, and Cu 1.

Ventral hindwing: Ground color range and variation similar to forewing, may or may not be covered variably with iridescent scales ( Fig. 2 ); discal band, concolorous to that of VFW, similar in width or broader, extending from costa to inner margin, often passing origin of Rs, or just basal of origin of this vein; concolorous postdiscal band passing origin of M 2 and/or Cu 1, otherwise near origin of these veins, similar in width or broader compared with previous band and appearing more or less parallel; submarginal band, concolorous to that of VFW, similar in width or broader, more jagged, extending from apex towards tornus, posterior end occasionally fused or terminating very close to postdiscal band near inner margin; concolorous marginal band, narrower than previous band and less sinuate, traversing along marginal area from apex to tornus; fringe greyish; six submarginal ocelli in cells Rs, M 1, M 2, M 3, Cu 1, and Cu 2 (ocellus in cell Cu 2 absent in C. trinitensis n. comb.), ocelli in cells M 1 and Cu 1 similar in size (as in C. castanya n. sp., C. fida n. comb., C. sanmarcos n. comb., and C. esmeralda n. sp.) or ocellus in Cu 1 may be larger (as in C. trinitensis n. comb., C. lea n. comb., and C. philippa n. comb. & reinst. stat.), both ocelli often possessing black central area and single white pupil ringed in yellow, ocelli in cells Rs and Cu 2 smaller compared with previous two ocelli but general appearance is same, ocelli in cells M 2 and M 3 are similar in size and appearance to those in cells Rs and Cu 2 but lacking black area, ocellus in cell M 2 placed more basally compared with ocellus in cell M 3.

Abdomen: Eighth tergite sclerotized in narrow anterior band and broader posterior patch; eighth sternite variable in appearance from rather narrow band to broader patch, as well as separated into two patches in some specimens (see below for further details).

Genitalia: Tegumen somewhat trapezoidal in lateral view (compared with more semi-circular and/or elongated tegumen in many other euptychiines) due to convexity of dorsal margin located posteriorly and anterior portion being rather straight, this convexity of dorsal margin variable as to its degree, ventral margin convex; uncus longer than tegumen in lateral view, setae visible at base and along ventral margin towards posterior end, uncus roughly straight in lateral view, appearing somewhat broad in lateral view towards base, middle section evenly broad in dorsal view, posteriorly terminating in slightly hooked point in lateral view, rather blunt in dorsal view; brachium broad at base, longer than uncus, tapering towards apex and terminating in pointed apical edge, dorsally projecting away from uncus with posterior portion curving back in; combination of ventral arms from tegumen and dorsal arms from saccus somewhat sinuate, roughly evenly broad; appendices angulares present, but not discernible in lateral view, visible as rectangular projection in posterior view (when valva removed); saccus appearance in lateral view varying from straight to curved, but generally length not exceeding uncus; juxta (i.e., fultura inferior) present as narrow stripe; valvae, distal half setose, basal two-thirds roughly rhomboidal in lateral view, apical process approximately one-third of valva although this ratio is variable (see below for further details), varying in appearance from a narrow, curving process slightly tapering towards apex to a broader, straight process (see below for further details), y terminating in angular tip, dorsal margin just distal of costa curving interiorly forming a sharp projection visible in dorsal view in C. lea n. comb. but apparently absent in other species, costa appearing somewhat as ‘fan-shaped’ plate, dorsal margin curving inwards and folded towards appendices angulares; phallus roughly straight, similar in length to valva in lateral in view, phallobase occupying about half of phallus, antero-dorsal opening large, sclerotized region from dorsal margin of phallobase projecting upwards and apparently forming part of manica at juncture of phallobase and aedeagus, manica covering approximately half of aedeagus, cornuti visible as roughly two sclerotized narrow patches of vesica, vesica visible at posterior opening of aedeagus ( Fig. 3 ).

Female: Forewing length: 21–27 mm (mean: 23.3 mm; n = 10)

Similar to male except as follows: Foretarsus divided into five tarsomers; forewing somewhat rounded and broader (apparent in C. lea n. comb., C. esmeralda n. sp., C. fida n. comb., and C. sanmarcos n. comb. due to their rather elongated male forewing);

overall coloration lighter; iridescent purple-ish lilac reflection on DFW variable, from absent to covering about basal half of DFW; iridescent purple-ish lilac reflection on DHW similarly variable, from absent to covering entire surface of DFW; iridescent scales around VHW tornus often less intense.

Female genitalia and abdomen: Eighth tergite fully developed (as seventh tergite); papilla analis lacking posterior apothysis; lamella antevaginalis sclerotized, forming rounded to rectangular plate with wrinkled margin; sclerotized plate present at lateral side of eighth abdominal segment, anterior margin fused with lamella antevaginalis (lateral margin fused as well in C. fida n. comb. and C. sanmarcos n. comb.), dorsal margin of this lateral sclerotized plate indented around spiracle; inter-segmental membrane of seventh and eighth abdominal segment pleated and expandable, elongated weakly sclerotized region present; ductus bursae membraneous, ductus seminalis exits close to ostium bursae, ductus bursae somewhat inflated between origin of ductus seminalis and ostium bursae, corpus bursae ellipse in dorsal view, with two relatively short signa parallel to each other and apparently located laterally, bursa extending to juncture of third and fourth abdominal segment ( Fig. 4 ).

Etymology: This generic name is based on the Spanish word ‘cisandina’, itself a compound word formed of the Latin preposition ‘cis’, meaning ‘on this side of’, and the Spanish adjective ‘andina’, meaning ‘Andean’, which is widely used to refer to the Neotropical region east of the Andes. The name refers to the fact that the species currently contained within this genus are found exclusively east of the Andean continental divide. The generic name should be regarded as a feminine noun in the nominative singular.

Distribution and natural history: Cisandina n. gen. is an entirely South American genus known exclusively from east of the Andes ( Fig. 8 ). Specifically, members are known from the cloud forests of the eastern Andes, the lowland rainforest of the Amazon basin, the Guianas, Trinidad, and the Atlantic coastal forest of Brazil and northeastern Argentina. Despite this broad range, no more than two species ever occur in sympatry. All species seem to be uncommon to very rare and are typically found in undisturbed forest. Singer and Ehrlich (1993) reported the host plant of Cisandina lea n. comb. (under the name ‘ Cissia junia’) as being an unidentified Gramineae ( Poaceae ) species according to their study in Trinidad, although this record is based on a plant accepted by the female in captivity in their oviposition trials. We here provide data on the immature stages of C. philippa n. comb. & reinst. stat. and C. castanya n. sp., including its natural host plant ( Figs. 6 and 7 ), which constitute to date the only reliable immature stage data available for this genus.

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. 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.

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.