Cisandina castanya Lamas & Nakahara, 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 : 23-24

Cisandina castanya Lamas & Nakahara , New Species

( Figs. 1 , 2i and j , 3k–n , 4e and f , 6e–i , 7c–e , 8 )

Caeruleuptychia sp. n. 2: Robbins, Lamas, Mielke, Harvey & Casagrande, 1996: 231.

Caeruleuptychia n. sp. Lamas, MS: Lamas 2004: 218.

Systematic placement and diagnosis. The phylogenetic analysis placed Cisandina castanya n. sp. as sister to ( C. lea n. comb. + C. esmeralda n. sp.) + C. philippa n. comb. & reinst. stat., although the support for this placement is moderate ( Fig. 1 ; SH-aLRT/UFBoot = 85.9/81). The single barcoded individual of C. castanya n. sp. (DZ 52.561–BC-DZ-139, from Acre, Brazil) shows high genetic divergence, greater than 3.74% (ranging from 3.74 to 6.54%; mean 5.0283% (n = 12), see Table 3) compared with the three other Cisandina n. gen. species which form a clade to which it is supported as sister, reinforcing our taxonomic hypothesis. Males of C. castanya n. sp. are readily distinguishable from other species in the genus by their small forewing length, in addition to the presence of semi-iridescent bluish scales on the DFW and DHW. Females are distinguished from other species in the genus by possessing similar semi-iridescent bluish scales on the DFW and DHW, which extend from the inner margin of DFW and reach the discocellular vein, whereas the bluish scales on the DFW are more restricted to basal area in female C. lea n. comb. and absent in C. philippa n. comb. & reinst. stat. As explained under the immediately preceding taxon, C. castanya n. sp. is sympatric with C. philippa n. comb. & reinst. stat. and species-level diagnostic characters can be found in the immature stages as well, namely head scoli appearing more developed in C. castanya n. sp. (scoli/ headcapsule width ratio = 0.14; scoli length c. 0.4 mm.) compared with C. philippa n. comb. & reinst. stat. (scoli/headcapsule width ratio=0.08; scoli length c. 0.2 mm.).

Male ( Fig. 2i ): n = 3).

Forewing length 22–23 mm (mean 22.3 mm;

Head: See head description for Cisandina n. gen. above.

Thorax: See thorax description for Cisandina n. gen. above. Abdomen: See abdomen description for Cisandina n. gen. above, eighth sternite rather narrow and appearing as two separated patches at anterior side.

Wing venation: See wing venation description for Cisandina n. gen. above.

Wing shape: See wing shape description for Cisandina n. gen. above, notable features include forewing appearing less elongate due to angle between forewing outer margin and inner margin being roughly right angle.

Dorsal forewing: Notable features include, ground color brownish, iridescent purple-ish lilac scales covering most of DFW except for area anterior of costal vein, apex and distal side of cells M 1, M 2, M 3, Cu 1, although this area revealing ground color narrows posteriorly ( Fig. 2i ); see also DFW description for Cisandina n. gen. above.

Dorsal hindwing: Notable features include, ground color similar to forewing, iridescent purplish lilac scales covering most of DHW except for area anterior of Rs and area posterior of 3A ( Fig. 2i ); see also DHW description for Cisandina n. gen. above.

Ventral forewing: Notable features include, ventral bands narrow, discal band and postdiscal band parallel to each other and traversing in slightly outward diagonal direction, ocellus in cell M 1 small and may appear insignificant with pupil being invisible, additional ocellus may be present in adjacent cells ( Fig. 2i ); see also VFW description for Cisandina n. gen. above.

Ventral hindwing: Notable features include, grey-ish blue scales covering areas in cells Cu 1, Cu 2 and 2A, submarginal ocelli appearing small, ocelli in cells M 1 and Cu 1 not reaching or barely reaching veins defining these cells, small ocellus may be visible at posterior end of postdiscal band in some specimens ( Fig. 2i ); see also VHW description for Cisandina n. gen. above.

Genitalia: Notable features include, dorsal margin of tegumen rather smoothly curved as well as anterior portion of ventral margin of tegumen appearing ‘bent’ in lateral view, rather straight saccus in lateral view, apical process of valva narrow and curving ( Fig. 3k–n ); see also male genitalia description for Cisandina n. gen. above.

Female ( Fig. 2j ): n = 3).

Forewing length 21–22 mm (mean 21.7 mm,

Similar to male except as follows: Notable features include, area of iridescent purplish lilac scales on DFW restricted to discal cell, basal part of cell M 3, more than half of Cu 1, most of cells Cu 2 and 2A ( Fig. 2j ); see also female wing shape and pattern description for Cisandina n. gen. above.

Female genitalia: Notable features include, lamella antevaginalis appearing as rectangular ‘thumb-like’ plate with wrinkled margin; signa short, occupying basal half of corpus bursae, length of corpus bursae similar to ductus bursae ( Fig. 4e and f ); see also male genitalia description for Cisandina n. gen. above.

Type material: Holotype male, with the following labels separated by double-forward slashes: // PERU, MD, Albergue Pantiacolla 400

m 12° 39 ′ S, 71° 14 ′ W 22.vi.2019 W. Dempwolf// SN-DNA19-48 (MUSM).

Paratypes (10 ♂, 22 ♀): Brazil: Acre: Senador Guiomard, Reserva Catuaba , [9°37 ′ S, 68°18 ′ W], (Mielke & Casagrande), 31 Jan–5 Feb 2009, 2 ♀ [DZ 52.561] ( DZUP) ; Mato Grosso: hwy Cuiabá- Santarém , km 715, [9°58 ′ 54 ″ S, 54°54 ′ 20 ″ W], 300 m, (Callaghan, C. J.), 14 Jul 1978, 1 ♂ [FLMNH-MGCL-265741], ( FLMNH) ; Rondônia: Cacaulândia, ‘ Big Rock Trail’ , 3 km E of Fazenda Rancho Grande on Linha C-20, [10°17 ′ 47 ″ S, 62°50 ′ 27 ″ W], (Austin, G. T.), 21 Sep 1992, 1 ♀ [FLMNH-MGCL-265743], ( FLMNH) ; 12.5 km S of Cacaulândia, off B-65, Linha C-2.5, [10°27 ′ 16 ″ S, 62°53 ′ 59 ″ W], (Austin, G. T.), 12 Nov 1990, 1 ♀ [FLMNH-MGCL-265736], ( FLMNH), 9 Dec 1990, 1 ♂ [ FLMNH- MGCL-265734], ( FLMNH) ; 15 km S of Cacaulândia, off B-65, Linha C-0, [10°28 ′ 43 ″ S, 62°53 ′ 46 ″ W], (Austin, G. T.), 22 Apr 1991, 1 ♀ [FLMNH-MGCL-265737], ( FLMNH) ; 3 km N Cacaulândia , dry trail off B-65, [10°30 ′ 24 ″ S, 62°48 ′ W], (Smith, J.), 23 Apr 1991, 1 ♀ [FLMNH-MGCL-296555], ( FLMNH) ; Cacaulândia, vicinity of Fazenda Rancho Grande , [10°17 ′ 58 ″ S, 62°52 ′ 14 ″ W], 180 m, (Austin, G. T.), 20 Oct 1989, 1 ♀ [FLMNH-MGCL-265739], ( FLMNH), 29 Oct 1989, 1 ♂ [FLMNH-MGCL-265730] [dissection, SN-20-12] ( FLMNH) ; 5 km S of Cacaulândia on Linha C-10 at Rio Pardo off B-65, [10°2 ′ 3 ′ 15 ″ S, 62°54 ′ 53 ″ W], (Gomes, O.), 23 Apr 1993, 1 ♀ [FLMNH-MGCL-265738] [dissection, SN-20-13], ( FLMNH), 23 Jun 1996, 1 ♀ [FLMNH-MGCL-265742], ( FLMNH), 7 Aug 1994, 1 ♀ [FLMNH-MGCL-265744], ( FLMNH), 8 Jun 1996, 1 ♀ [FLMNH-MGCL-296559], ( FLMNH) ; Cacaulândia, 7 km E B-65, Fazenda Rancho Grande , [10°17 ′ 58 ″ S, 62°52 ′ 14 ″ W], (Austin, G. T.), 16 Sep 1992, 1 ♀ [FLMNH-MGCL-265745] [dissection, SN-20-33], ( FLMNH), 26 Nov 1991, 1 ♀ [FLMNH-MGCL-265735], ( FLMNH), 29 Nov 1991, 1 ♀ [FLMNH-MGCL-265740], ( FLMNH), (Austin, G.T.), 17 Jul 1995, 1 ♀ [FLMNH-MGCL-296557], ( FLMNH), 22 Jul 1994, 1 ♂ [FLMNH-MGCL-296553] [dissection, SN-20-32], ( FLMNH), 3 May 1995, 1 ♀ [FLMNH-MGCL-296558], ( FLMNH), 5 May 1995, 1 ♀ [FLMNH-MGCL-296556], ( FLMNH), (Brock, J. P.), 5 Nov 1989, 1 ♂ [FLMNH-MGCL-265733], ( FLMNH), (Gomes, O.), 27 Jun 1989, 1 ♂ [FLMNH-MGCL-265731], ( FLMNH) ; off B-65, Linha C-20 at Rio Pardo , [10°17 ′ 58 ″ S, 62°57 ′ 5 ″ W], (Austin, G. T.), 18 Sep 1992, 1 ♂ [FLMNH-MGCL-265732], ( FLMNH) . Peru: Huánuco: Lower Ucayali, Río Pachitea , [8°46 ′ S, 74°32 ′ W], 150 m, (Tessman, G.), 1 ♀, ( MNHU) ; Madre de Dios: 15 km E Puerto Maldonado , [12°32 ′ S, 69°3 ′ W], 200 m, (Medina, M.), 6 Feb 1990, 1 ♀ [MUSM-LEP-100320] [dissection, SN-16-20], ( MUSM) ; Parque Nacional del Manu, Pakitza , [11°55 ′ 48 ″ S, 71°15 ′ 18 ″ W], 340 m, (Harvey, D. J.), 27 Apr 1991, 1 ♀ [MUSM-LEP-100319], ( MUSM), (Mielke, O. H. H.), 3 Oct 1991, 1 ♂ [MUSM-LEP-100316], ( MUSM), (Robbins, R. K.), 2 Oct 1991, 1 ♂ [MUSM-LEP-100317] [dissection, SN-16-36], ( MUSM) ; Albergue Pantiacolla , [12º39 ′ S, 71º14 ′ W], 400–450m, (Lamas, G.), 29 Oct 2016, 1 ♀, ( MUSM), 400m, (Gibson, L.), 30 Oct 2018, [SN-DNA19-49], 1 ♂, ( MUSM) .

Other records: Peru: Loreto: Río Paranapura, Chambira , [5°54 ′ S, 76°14 ′ W], 120 m, (Razzeto, O.), 1 ♂ [MUSM-LEP-100318], ( MUSM) ; Río Ucayali, Contamana , [7°21 ′ S, 75°0 ′ W], 135 m, (Büche, M.), Sep 1999, 1 ♂ [MUSM-LEP-100315], ( MUSM) .

Etymology: The specific epithet is derived from the Spanish word ‘castaña’ and the Portuguese word ‘castanha’, both meaning ‘chestnut’ in English and, in much of Peru, Bolivia, and Brazil, more specifically ‘ Brazil nut’; Brazil nuts are large, emergent trees ( Bertholletia excelsa Humb. & Bonpl. ( Lecythidaceae )) that dominate the rainforest of the southwestern Amazon basin and whose distribution somewhat overlaps with that of this new butterfly species. These trees grow in abundance and characterize the landscape at Finca Las Piedras (Madre de Dios, Peru), where the immature stages of this new species were discovered. This species-group name is a feminine noun in apposition.

Distribution and natural history: This species is distributed in the southwestern Amazon basin, from central and southern Peru eastwards into Brazil (Mato Grosso) ( Fig. 8 ). A second-instar caterpillar and the last instar was found on a herbaceous bamboo, Olyra latifolia L. ( Poaceae ) at Finca Las Piedras, Madre de Dios, Peru, on 20 June 2020 (voucher: 2020-FLP-IMM-0188) and on 14 March 2021 (voucher: 2021-FLP-IMM-0234), respectively, and the immature stages are described below. The instar number (second) for 2020-FLP-IMM-0188 was determined based on comparing its size and appearance with other euptychiine larval instars. The host plant with the caterpillar was located at an edge habitat of a terra firme forest dominated by Brazil nut trees, with high incidence of light in the understory, fostering the growth of understory grasses.

Remarks. The two male individuals from Loreto, Peru (MUSM- LEP-100315 and MUSM-LEP-100318) are classified as this taxon in MUSM. Nevertheless, these specimens possess some phenotypic characters not observed in other individuals examined for C. castanya n. sp. such as large ventral ocelli and presence of small ocellus at the VHW inner margin (at the posterior end of the VHW postdiscal band). Thus, we decided to exclude these individuals from the type series until further data becomes available to assess its identity.

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.