Systematic Revision of a New Butterfly Genus, Cisandina Nakahara & Espeland, n. gen., with Descriptions of Three New Taxa (Lepidoptera: Nymphalidae: Satyrinae)

Abstract

We here establish a new genus in the nymphalid butterfly subtribe Euptychiina, Cisandina Nakahara & Espeland, n. gen. to harbor five species hitherto placed within two polyphyletic genera, namely Magneuptychia Forster, 1964 and Euptychoides Forster, 1964. We compiled data from over 350 specimens in 17 public and private collections, as well as DNA sequence data for all relevant species, to revise the species-level classification of this new genus. According to our multi-locus molecular phylogeny estimated with the maximum likelihood approach, Cisandina lean. comb., Cisandina philippan. comb. & reinst. stat., Cisandina fidan. comb., Cisandina sanmarcosn. comb., and Cisandina trinitensisn. comb. are proposed as new taxonomic combinations, since these species are distantly related to the type species of Magneuptychia and Euptychoides and cannot reasonably be accommodated in any other genus. Lectotypes are designated for Papilio lea Cramer, 1777, Papilio junia Cramer, 1780, Euptychia philippa Butler, 1867, and Eupytchia fida Weymer, 1911. Two new species of Cisandinan. gen. are named and described herein, C. esmeralda Nakahara & Barbosa, n. sp. and C. castanya Lamas & Nakahara, n. sp., increasing the described species diversity of the genus to seven. The immature stages of C. castanyan. sp. and C. philippan. comb. & reinst. stat. are documented along with their natural hostplants, representing the first two species of the genus with known life history information. We describe a new subspecies, Cisandina fida directa Nakahara & Willmott, n. ssp., based on a limited number of specimens from southern Ecuador and central Peru. We were unable to obtain genetic data for the nominate race of C. fidan. comb., and thus, this taxonomic hypothesis is currently based solely on phenotypic characters.

Resumen

Se establece un nuevo género de mariposas ninfálidas de la subtribu Euptychiina, Cisandina Nakahara & Espeland, n. gen. para albergar cinco especies previamente ubicadas dentro de dos géneros polifiléticos, Magneuptychia Forster, 1964 y Euptychoides Forster, 1964. Se recopiló datos de más de 350 especímenes de 17 colecciones públicas y privadas, así como datos de secuencias de ADN para todas las especies relevantes y así poder revisar la clasificación a nivel de especie de este nuevo género. De acuerdo con nuestra filogenia molecular multilocus, estimada con el enfoque de máxima verosimilitud, se propone como nuevas combinaciones taxonómicas a Cisandina lean. comb., Cisandina philippan. comb. & reinst. stat., Cisandina fidan. comb., Cisandina sanmarcosn. comb. y Cisandina trinitensisn. comb., ya que estas especies se relacionan lejanamente con las especies tipo de Magneuptychia y Euptychoides y no pueden acomodarse razonablemente en ningún otro género. Se designa lectotipos para Papilio lea Cramer, 1777, Papilio junia Cramer, 1780, Euptychia philippa Butler, 1867 y, Eupytchia fida Weymer, 1911. Adicionalmente se nombra y describe aquí dos nuevas especies de Cisandinan. gen., C. esmeralda Nakahara & Barbosa, n. sp. y C. castanya Lamas & Nakahara, n. sp., aumentando la riqueza de especies descritas del género a siete. Los estadíos inmaduros de C. castanya Lamas & Nakahara, n. sp. y C. philippan. comb. & reinst. stat., son documentados junto con su planta hospedante natural, aportando nueva información sobre el ciclo de vida del género. Además, describimos una nueva subespecie, Cisandina fida directa Nakahara & Willmott, n. ssp., con base en un número limitado de especímenes del sur de Ecuador y del centro de Perú. No obtuvimos datos genéticos para la raza nominada de C. fidan. comb., por lo que esta hipótesis taxonómica se establece únicamente con base en caracteres fenotípicos.

Resumo

Estabelecemos aqui um novo gênero na subtribo de borboletas ninfalídeas Euptychiina, Cisandina Nakahara & Espeland, n. gen. para abrigar cinco espécies até agora alocadas dentro de dois gêneros polifiléticos, Magneuptychia Forster 1964 e Euptychoides Forster 1964. Compilamos dados de mais de 350 espécimes em 17 coleções públicas e privadas, bem como dados de sequência de DNA para todas as espécies relevantes, para revisar a classificação a nível de espécie deste novo gênero. De acordo com nossa filogenia molecular multilocus estimada com a abordagem de máxima verossimilhança, são propostas como novas combinações taxonômicas: Cisandina lean. comb., Cisandina philippan. comb. & reinst. stat., Cisandina fidan. comb., Cisandina sanmarcosn. comb. e Cisandina trinitensisn. comb., uma vez que essas espécies estão distantemente relacionadas às espécies-tipo de Magneuptychia e Euptychoides e não podem ser razoavelmente acomodadas em qualquer outro gênero. Lectótipos são designados para Papilio lea Cramer 1777, Papilio junia Cramer 1780, Euptychia philippa Butler 1867 e Eupytchia fida Weymer 1911. Duas novas espécies de Cisandinan. gen. são nomeadas e descritas aqui, C. esmeralda Nakahara & Barbosa, n. sp. e C. castanya Lamas & Nakahara, n. sp., resultando no aumento da diversidade de espécies do gênero para sete. Os estágios imaturos de C. castanyan. sp. e C. philippan. comb. & reinst. stat., são documentados junto com sua planta hospedeira natural, representando duas espécies com informações de história de vida conhecidas para o gênero. Descrevemos uma nova subespécie, Cisandina fida directa Nakahara & Willmott, n. ssp., com base em um número limitado de espécimes do sul do Equador e centro do Peru. Não foi possível obter dados genéticos para a raça nominal de C. fidan. comb., e, portanto, esta hipótese taxonômica é atualmente baseada apenas em caracteres fenotípicos.

Euptychiina, Lectotype, taxonomy, Magneuptychia, Euptychoides

The nymphalid genus Magneuptychia Forster, 1964 was erected by Forster (1964) to accommodate 15 species previously associated with Euptychia Hübner, 1818 (sensu lato), by designating Papilio libye Cramer, 1767 as the type species. Nevertheless, as indicated by Forster (1964: 125) in his description, this placement of species in Magneuptychia is ‘bis auf weiteres’ (i.e., for the time being), and the genus was established based on differences in wing span and pattern, as well as male genitalic characters in comparison with Argyreuptychia Forster, 1964 without any supporting phylogenetic hypotheses. Forster’s (1964) tentative classification of Magneuptychia was followed by subsequent authors such as Miller (1968) and was one of the most diverse genera with 29 described species recognized by Lamas (2004) in his comprehensive checklist of the satyrine subtribe Euptychiina. Subsequent molecular (Murray and Prowell 2005; Peña et al. 2010, Espeland et al. 2019) and morphological (Marín et al. 2017) phylogenetic studies, however, have shown Magneuptychia to be thoroughly polyphyletic. Costa et al. (2016) discussed the taxonomy of Magneuptychia and Andrade et al. (2019) proposed generic classifications for nine putative species previously placed in Magneuptychia by transferring them into four new genera erected in that study, as well as one existing genus Satyrotaygetis Forster, 1964. However, taxonomic hypotheses in both of these two publications were based solely on comparative morphology without providing any supporting phylogenetic analysis. Following Nakahara et al. (2020a) and Zacca et al. (2021), a phylogenetic study is underway by SN and collaborators to classify taxa previously placed in Magneuptychia (sensuLamas 2004). Meanwhile, during attempts to revise another polyphyletic euptychiine genus, Euptychoides Forster, 1964, Nakahara et al. (2016), and Willmott et al. (2019) described new genera, supported by phylogenetic analyses, for taxa unrelated to the type species of that genus, Euptychia saturnus Butler, 1867. This ongoing effort to unravel the systematics of Euptychoides is still underway by this study’s authors and colleagues.

This article, therefore, aimed to further clarify the systematics of both Magneuptychia and Euptychoides, with emphasis on a clade of seven species with similar ventral wing patterns and genitalia, that are distributed entirely east of the Andes. We here propose a new genus for this clade and review its species-level taxonomy, including providing descriptions of three new species-group names.

Materials and Methods

We conducted field work in Brazil, Ecuador, and Peru to study euptychiines and collect specimens for morphological and molecular analysis. Additionally, specimens in the following public and private collections were examined and the following collection abbreviations are used:

• ASA:      Alliance for a Sustainable Amazon collection, Puerto Maldonado, Perú

• BMB:     Booth Museum of Natural History, Brighton, UK

• DEMU:   Debra Murray collection, USA

• DZUP:    Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Brazil

• FLMNH:  McGuire Center for Lepidoptera and Biodiversity (MGCL), Florida Museum of Natural History, University of Florida, Gainesville, USA

• INABIO:  Instituto Nacional de Biodiversidad, Quito, Ecuador (formerly MECN)

• MACN:   Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia’, Buenos Aires, Argentina

• MIPE:     Mike J. Perceval collection, Surrey, UK

• MNHU:    Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung an der Humboldt Universität, Berlin, Germany

• MOBE:   Mohamed Benmesbah collection, Toulouse, France

• MUSM:     Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Perú

• MZUJ:            Zoological Museum, Jagellonian University, Krakow, Poland

• NHMUK: Natural History Museum, London, UK (formerly BMNH)

• RMNH:     Rijksmuseum van Natuurlijke Historie (currently Netherlands Centre for Biodiversity Naturalis), Leiden, Netherlands

• SMT:     Senckenberg Museum für Tierkunde, Dresden, Germany

• USNM:    National Museum of Natural History, Smithsonian Institution, Washington, DC, USA

• ZSM:     Zoologische Staatssammlung München, Munich, Germany

• ZUEC:    Museu de Zoologia da Universidade Estadual de Campinas ‘Adão José Cardoso’, Campinas, Brazil.

Over 350 specimens examined in these collections were databased and georeferenced in accordance with Nakahara et al. (2019b). Label data for name-bearing types are written verbatim with labels separated by double-forward slashes.

We studied morphology using standard techniques, with abdomens and other appendages being soaked in hot (80°C) 10% KOH for 5–10 min. These appendages and genitalia were dissected and subsequently stored in glass tubes and/or small plastic vials filled with glycerine. The wing venation was visualized by clearing scales of the ventral surface using 70% ethanol. Morphological features were studied using a Leica MZ 16 stereomicroscope at magnifications up to 100×, and a Leica LED2500 (10–160×). Almost all the illustrations were made using a camera lucida attached to a Leica MZ 16 stereomicroscope, except that illustrations of head capsules of the immature stages were drawn in Adobe Illustrator under constant observation in the Leica LED2500 stereomicroscope. The study site and methods for the immature stages’ documentation largely follow Baine et al. (2019). The terminology associated with wings and genitalia follows Nakahara et al. (2018a, b), and the following abbreviations are used: DFW: dorsal forewing; DHW: dorsal hindwing; VFW: ventral forewing; VHW: ventral hindwing.

We follow Nakahara et al. (2020b) for methods for DNA extractions, internal primer design, PCR, and Sanger sequencing for the first half of the mitochondrial gene cytochrome oxidase I (COI), commonly known as the ‘DNA barcode’ (sensuHebert et al. 2003), as well as for three nuclear gene sequences: elongation factor 1-alpha (Ef1a), glyceraldehyde-3-phosphate dehydrogenase (GAPDH), and ribosomal protein S5 (RpS5). These four genes were amplified for selected individuals of the ‘Archeuptychia clade’, coupled with four out-group individuals, and used to produce a phylogenetic hypothesis to support our taxonomic proposal. Information regarding sequences used in this study, including new DNA sequence data generated, is provided with GenBank voucher codes (Table 1). We performed a phylogenetic analysis with maximum likelihood as the optimality criterion on an alignment of these mitochondrial and nuclear gene sequences in IQ-TREE v.2.0.5 (Minh et al. 2020). The data were partitioned into codon positions and best-fit substitution models individually derived through ModelFinder (Kalyaanamoorthy et al. 2017) were applied, with information provided in Table 2. In total, 20 independent analyses were conducted based on the above dataset, with branch support estimated through 2,000 replications of both ultrafast bootstrap (UFBoot) with the ‘-bnni’ option to reduce model violation, and Shimodaira-Hasegawa-like approximate likelihood ratio test (SH-aLRT). The run which gave the tree with the highest log-likelihood score for the above dataset was selected and the tree was rooted with Godartiana byses (Godart, [1824]). We also calculated infra- and interspecific genetic distances based on ‘DNA barcode’ data among Cisandinan. gen. taxa based on Kimura-2-parameter (Table 3).

Table 1.

GenBank Accession number information for DNA sequences used in this study

Voucher code	Genus	Species	COI	Ef1a	RpS5	GAPDH	Locality
BC-DZ Willmott-137	Cisandina	esmeralda	MW008458	MT999012	N/A	MT998979	Brazil: Paraná: Reservas Particulares do Patrimônio Natural (RPPN) Moreira Salles (−24.071, −53.033)
BC-DZ Willmott-138	Cisandina	lea	MW018438	N/A	N/A	N/A	Brazil: Amazonas: Foz do Rio Curuduri (−0.092, −63.287)
BC-DZ Willmott-139	Cisandina	castanya	MW008460	N/A	N/A	N/A	Brazil: Acre: Reserva Catuaba (−10.071, −67.619)
BC-DZ Willmott-250	Archeuptychia	cluena	MW008469	MT999007	MT998995	MT998984	Brazil: Espírito Santo: Reserva Ecológica de Sooretama (−19.054, −40.142)
IN014	Erichthodes	arius	MW008470	N/A	N/A	N/A	Peru: Junín: 2.5 km N San Ramón, Genova (−11.091, −75.351)
KW-090113-21	Cepheuptychia	cephus	MW008471	N/A	MT998996	MT998985	Ecuador: Orellana: Boca del Río Añangu (−0.529, −76.395)
KW-140708-02	Chloreuptychia	chlorimene	MW008474	MT999008	MT998997	MT998986	Ecuador: Orellana: Estación Científica Yasuní (−0.674, −76.397)
KW-15-025	Euptychoides	sanmarcos	MW008466	MT999005	MT998993	MT998982	Peru: Cuzco: Quebrada Quitacalzón (−13.022, −71.493)
KW-15-034	Pseudeuptychia	languida	MW008473	N/A	N/A	N/A	Peru: Cuzco: Quebrada Quitacalzón (−13.022, −71.493)
LCB353	Chloreuptychia	marica	MW008475	N/A	N/A	N/A	French Guiana: Cayenne: Nouragues Station (4.083, −52.683)
LCB356	Cisandina	trinitensis	MW008463	N/A	N/A	N/A	French Guiana: Not located: ‘French Guiana’
LEP-08983	Cisandina	philippa	MW008467	N/A	MT998990	N/A	Ecuador: Zamora-Chinchipe: La Libertad (−3.798, −78.607)
LEP-10517	Pseudeuptychia	languida	MW008472	N/A	N/A	N/A	Ecuador: Orellana: Reserva Biológica del Río Bigal, ‘the Hooch’ (−0.524, −77.419)
LEP-16705	Cisandina	fida	MW008465	MT999004	MT998992	MT998981	Ecuador: Zamora-Chinchipe: Juyapa (−3.58, −78.951)
LEP-16939	Cisandina	philippa	MW008468	MT999006	MT998994	MT998983	Ecuador: Morona-Santiago: Isla de las Conchas (−3.036, −77.975)
LEP-18697	Stevenaria	eriphule	MW008477	N/A	MT999001	N/A	Guyana: Upper Takutu-Upper Essequibo: Kanuku Mountains (3.2, −59.567)
LEP-19580	Chloreuptychia	herseis	MW008476	MT999009	MT998998	MT998987	Peru: Madre de Dios: Los Amigos Biological Station (−12.567, −70.093)
LEP-34359	Cisandina	lea	MW008459	N/A	N/A	N/A	French Guiana: St-Laurent du Maroni: Saül (3.855, −53.302)
LEP-58115	Cisandina	fida	MW008464	N/A	N/A	N/A	Peru: Ucayali: Previsto (−9.05, −75.633)
MB-1708-15	Cisandina	trinitensis	MW008462	N/A	N/A	N/A	French Guiana: Cayenne: Nouragues Station (4.083, −52.683)
LCB251	Cisandina	trinitensis	MW008461	MT999003	MT998991	MT998980	French Guiana: St-Laurent du Maroni: Antecume-Pata, ‘filet’ (3.292, −54.069)
MGCL-LOAN-523	Godartiana	byses	KU892533	N/A	KU892535	KU892534	Brazil: Bahía: Reserva Biológica de Una (−15.173, −39.103)
LEP-14777	Magneuptychia	libye	MG209754	MF192722	MG546320	MF192728	Ecuador: Zamora-Chinchipe: km 6 Los Encuentros-El Panguí (−3.731, −78.609)
NW149-9	Archeuptychia	cluena	GQ864736	GQ864830	GQ865392	GQ864926	Brazil: São Paulo: Salesópolis
CP02-50	Chloreuptychia	marica	GU20583	GU205887	GU206064	GU20594	Peru: Madre de Dios: Posada Amazonas (−12.801, −69.293)
DNA99-050	Chloreuptychia	rectilinea	AY508528	AY509055	N/A	N/A	Ecuador: Napo: no specific locality
LEP-14945	Chloreuptychia	herseis		MT999010	MT998999	MT998988	Suriname: Nickerie: Kabalebo Dam
KW-081111-49	Chloreuptychia	rectilinea		MT999011	MT999000	MT998989	Ecuador: Orellana: Boca del Río Añangu (−0.529, −76.395)
LEP-34283	Cisandina	philippa	MW018439	N/A	N/A	N/A	Ecuador: Morona-Santiago: Santiago de Mendez (−3.036, −78.033)
LEP-10686	Cisandina	fida	MW018440	N/A	N/A	N/A	Ecuador: Zamora-Chinchipe: ridge E San Roque (−3.703, −78.593)
LEP-08986	Cisandina	philippa	MW018441	N/A	N/A	N/A	Ecuador: Zamora-Chinchipe: Centro Shuar Cháarip (−3.635, −78.558)
LEP-08984	Cisandina	philippa	MW018442	N/A	N/A	N/A	Ecuador: Zamora-Chinchipe: La Libertad (−3.798, −78.607)
KW-081111-38	Cisandina	philippa	MW018443	N/A	N/A	N/A	Ecuador: Orellana: Boca del Río Añangu (−0.529, −76.395)

Table 2.

Best-fit substitution models by partition derived from ModelFinder and applied in this study

Codon position	Model
COI 1st	TIM2+F+G4
COI 2nd	K3Pu+F+I
COI 3rd	TIM+F+G4
EF1a 1st	FN+F
EF1a 2nd	F81+F+I
EF1a 3rd	TPM3+F+G4
GAPDH 1st	F81+F+I
GAPDH 2nd	F81+F+I
GAPDH 3rd	TPM+F+G4
RPS5 1st	TNe+I
RPS5 2nd	JC
RPS5 3rd	TIM2e+G4

Table 3.

Genetic distances among Cisandinan. gen. taxa based on DNA ‘barcodes’

	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17	18	19	20
1. LEP-10686_Cisandina_fida_Ecuador_Zamora-Chinchipe		0.0247	0.0004	0.0647	0.0674	0.0592	0.0794	0.0796	0.0794	0.0517	0.0519	0.064	0.0563	0.0529	0.0747	0.0516	0.0094	0.1257	0.1385	0.1059
2. LEP-58115_Cisandina_fida_Peru_Ucayali	0.0247		0.0246	0.0754	0.0782	0.0699	0.0902	0.0904	0.0902	0.0625	0.0626	0.0747	0.067	0.0636	0.0855	0.0623	0.0202	0.1365	0.1492	0.1167
3. LEP-16705_Cisandina_fida_Ecuador_Zamora-Chinchipe	0.0004	0.0246		0.0646	0.0673	0.0591	0.0793	0.0795	0.0793	0.0516	0.0518	0.0639	0.0562	0.0528	0.0746	0.0515	0.0093	0.1256	0.1384	0.1058
4. BC-DZ 138 Cisandina lea Brazil_Amazonas	0.0647	0.0754	0.0646		0.0084	0.0505	0.0501	0.0503	0.0501	0.0431	0.0432	0.0553	0.0476	0.0442	0.066	0.0429	0.0602	0.117	0.1298	0.0973
5. LEP-34359_Cisandina_lea_French	0.0674	0.0782	0.0673	0.0084		0.0533	0.0528	0.053	0.0528	0.0458	0.0459	0.058	0.0503	0.047	0.0688	0.0457	0.063	0.1198	0.1325	0.1
6. BC-DZ 139_Cisandina_castanya_Brazil_Acre	0.0592	0.0699	0.0591	0.0505	0.0533		0.0652	0.0654	0.0652	0.0376	0.0377	0.0498	0.0421	0.0387	0.0605	0.0374	0.0547	0.1115	0.1243	0.0918
7. MF546153LEPIG190-11_Cisandina_lea_Argentina_Misiones	0.0794	0.0902	0.0793	0.0501	0.0528	0.0652		0.0002	0	0.0578	0.0579	0.07	0.0623	0.0589	0.0807	0.0576	0.0749	0.1318	0.1445	0.112
8. BC-DZ 137_Cisandina_esmeralda_Brazil_Parana	0.0796	0.0904	0.0795	0.0503	0.053	0.0654	0.0002		0.0002	0.058	0.0581	0.0702	0.0625	0.0591	0.081	0.0579	0.0752	0.132	0.1447	0.1122
9. MF545572LEPIG013-11_Cisandina_lea_Argentina_Misiones	0.0794	0.0902	0.0793	0.0501	0.0528	0.0652	0	0.0002		0.0578	0.0579	0.07	0.0623	0.0589	0.0807	0.0576	0.0749	0.1318	0.1445	0.112
10. LEP-08986_Cisandina_philippa_Ecuador_Zamora-Chinchipe	0.0517	0.0625	0.0516	0.0431	0.0458	0.0376	0.0578	0.058	0.0578		0.0004	0.0125	0.0048	0.0014	0.0233	0.0001	0.0473	0.1041	0.1168	0.0843
11. LEP-08983_Cisandina_philippa_Ecuador_Zamora-Chinchipe	0.0519	0.0626	0.0518	0.0432	0.0459	0.0377	0.0579	0.0581	0.0579	0.0004		0.0127	0.0049	0.0016	0.0234	0.0003	0.0474	0.1042	0.117	0.0845
12. LEP-34283_Cisandina_philippa_Ecuador_Morona-Santiago	0.064	0.0747	0.0639	0.0553	0.058	0.0498	0.07	0.0702	0.07	0.0125	0.0127		0.0171	0.0137	0.0355	0.0124	0.0595	0.1163	0.1291	0.0966
13. LEP-16939_Cisandina_philippa_Ecuador_Morona-Santiago	0.0563	0.067	0.0562	0.0476	0.0503	0.0421	0.0623	0.0625	0.0623	0.0048	0.0049	0.0171		0.006	0.0278	0.0047	0.0518	0.1086	0.1214	0.0889
14. KW-081111-38_Cisandina_philippa_Ecuador_Orellana	0.0529	0.0636	0.0528	0.0442	0.047	0.0387	0.0589	0.0591	0.0589	0.0014	0.0016	0.0137	0.006		0.0244	0.0013	0.0484	0.1052	0.118	0.0855
15. DNA99-022_Cisandina_philippa_MD_Peru	0.0747	0.0855	0.0746	0.066	0.0688	0.0605	0.0807	0.081	0.0807	0.0233	0.0234	0.0355	0.0278	0.0244		0.0231	0.0702	0.1271	0.1398	0.1073
16. LEP-08984_Cisandina_philippa_Ecuador_Zamora-Chinchipe	0.0516	0.0623	0.0515	0.0429	0.0457	0.0374	0.0576	0.0579	0.0576	0.0001	0.0003	0.0124	0.0047	0.0013	0.0231		0.0471	0.1039	0.1167	0.0842
17. KW-15-025_Cisandina_sanmarcos_Peru_Cuzco	0.0094	0.0202	0.0093	0.0602	0.063	0.0547	0.0749	0.0752	0.0749	0.0473	0.0474	0.0595	0.0518	0.0484	0.0702	0.0471		0.1212	0.134	0.1015
18. LCB252_Cisandina_trinitensis_French_Guiana	0.1257	0.1365	0.1256	0.117	0.1198	0.1115	0.1318	0.132	0.1318	0.1041	0.1042	0.1163	0.1086	0.1052	0.1271	0.1039	0.1212		0.0128	0.0199
19. MB-1708-15_Cisandina_trinitensis_French_Guiana	0.1385	0.1492	0.1384	0.1298	0.1325	0.1243	0.1445	0.1447	0.1445	0.1168	0.117	0.1291	0.1214	0.118	0.1398	0.1167	0.134	0.0128		0.0326
20. LCB356_Cisandina_trinitensis_French_Guiana	0.1059	0.1167	0.1058	0.0973	0.1	0.0918	0.112	0.1122	0.112	0.0843	0.0845	0.0966	0.0889	0.0855	0.1073	0.0842	0.1015	0.0199	0.0326

Nomenclature

This article has been registered in ZooBank (www.zoobank.org), the official register of the International Commission on Zoological Nomenclature. The LSID (Life Science Identifier) number of the publication is: urn:lsid:zoobank.org:pub:ACFA7036-C10E-4370-BF43-D22288168F33

Results

Systematics

Our multi-locus maximum likelihood tree (Fig. 1; LnL = −11380.331) shows that Papilio lea Cramer, 1777 and Euptychia philippa Butler, 1867 (see our discussion below for the specific status of this taxon), two taxa hitherto placed in Magneuptychia, do not group closely with the type species of Magneuptychia, Papilio libye Linnaeus, 1767. This phylogenetic hypothesis also indicates that Euptychoides sanmarcos Nakahara & Lamas, 2018 and Euptychia fida Weymer, 1911, two species previously provisionally placed in Euptychoides, are strongly supported as sister to the clade including Papilio lea and Euptychia philippa (Fig. 1; SH-aLRT/UFBoot = 96.5/95). As discussed and demonstrated in a number of phylogenetic studies (e.g., Peña et al. 2010, Nakahara et al. 2016, Willmott et al. 2019), other species currently placed in Euptychoides are either in the so-called ‘Pareuptychia clade’ or ‘Splendeuptychia clade’, with the type species in the former clade. Finally, the recently described species Magneuptychia trinitensis Brévignon & Benmesbah, 2012 was moderately to strongly supported as sister to the clade containing the above-mentioned former Magneuptychia and Euptychoides species (Fig. 1; SH-aLRT/UFBoot = 91.9/92), which it also closely resembles in male genitalia and ventral wing pattern. Therefore, we herein describe a new genus to harbor all of these taxa, in addition to naming two new species and one new subspecies within the new genus.

Fig. 1.

Maximum likelihood tree (LnL = −11380.331) inferred in IQ-TREE v2.0.5, showing monophyly of Cisandinan. gen., as well as relationships of taxa within these genera. Numbers beside branches are SH-aLRT/UFBoot values. HT denotes holotype specimens.

Family Nymphalidae Rafinesque, 1815

 

Cisandina Nakahara & Espeland, New Genus

Type species: Papilio lea Cramer, 1777 – by present designation

Species Account:

• Cisandina Nakahara & Espeland, n. gen.

• Cisandina lea (Cramer, 1777) (Papilio), n. comb.

• = Papilio junia Cramer, 1780

• Cisandina esmeralda Nakahara & Barbosa, n. sp.

• Cisandina philippa (Butler, 1867) (Euptychia), n. comb. & reinst. stat.

• =Euptychia batesii f. tersa Weymer, 1911

• Cisandina castanya Lamas & Nakahara, n. sp.

• Cisandina fida fida (Weymer, 1911) (Euptychia), n. comb.

• Cisandina fida directa Nakahara & Willmott, n. ssp.

• Cisandina sanmarcos (Nakahara & Lamas, 2018) (Euptychoides), n. comb.

• Cisandina trinitensis (Brévignon & Benmesbah, 2012) (Magneuptychia), n. comb.

Generic Relationships and Diagnosis:

Based on hybrid enrichment data, Espeland et al. (2019) found ‘Magneuptychia lea philippa’ as a member of strongly supported (UFBoot > 95; Posterior Probability [PP] > 0.95) ‘Archeuptychia clade’, sister to Chloreuptychia herseis (Godart, [1824]), although with weak support (UFBoot < 75; PP < 0.75). The ‘Archeuptychia clade’ is weakly supported (UFBoot < 75; PP < 0.75) as sister to the ‘Splendeuptychia clade’. Analyses conducted here (and in ongoing research, Espeland et al., unpublished data) including additional members of the ‘Archeuptychia clade’ recovered Cisandinan. gen. as a monophyletic group with moderate support (Fig. 1; SH-aLRT/UFBoot = 91.9/92), although its closest relatives were not strongly resolved. It is worth mentioning that the monophyly of Cisandinan. gen. is strongly supported (UFBoot > 95) in a molecular phylogeny generated to describe a new genus for its potential sister group, as mentioned below. Our analysis found the sister group to be Yphthimoides eriphule (Butler, 1867) + Erichthodes arius (Weymer, 1911), with other closely related taxa including Chloreuptychia herseis, Chloreuptychia marica (Weymer, 1911), Pseudeuptychia Forster, 1964, and Archeuptychia Forster, 1964. Research is underway on the generic classification of these various clades.

Cisandinan. gen. is distinguished from other members of the ‘Archeuptychia clade’ by the configuration of the VHW submarginal ocelli, with each having only a single, central white pupil (except for the ocellus in cell M2), and which are largest in cells Cu1 and M1 and reduced in size in adjacent cells but otherwise similar in form, with the ocellus in M2 notably offset basally in comparison with the neighboring ocelli. Erichthodes arius and A. cluena (Drury, 1782) have VHW submarginal ocelli of similar shape and appearance, but they are roughly uniform in size throughout the wing, while remaining species have the ‘middle’ two ocelli (in cells M2 and M3) different in form (elongate, with stretched silvery or whitish pupils, or very much reduced) compared with the ocelli in the cells bounding them. All members of Cisandinan. gen., except for C. trinitensisn. comb., possess a sixth VHW ocellus in cell Cu2, whereas most other species in the ‘Archeupytchia’ clade lack this ocellus in cell Cu2. The male genitalia are relatively uniform and are distinguished from other ‘Archeuptychia clade’ members by having a valva that is broad basally then tapers sharply in the middle to form an elongate, narrow posterior projection (instead of gradually tapering throughout as in E. arius and A. cluena, or broad throughout as in Pseudeuptychia and Chloreuptychia). Furthermore, the reduced appendices angulares (more visible in posterior view and hardly visible in lateral view), rather broad base of brachia, sclerotized portion of manica extending from phallobase at its juncture with aedeagus (this character is also seen in E. arius and Yphthimoides eriphule (Butler, 1867)), and large antero-dorsal opening of phallus is apparently unique features within Euptychiina that are restricted to this genus. The genitalia, however, differ slightly among species in Cisandinan. gen. For example, the postero-dorsal process of the phallus (i.e. aedeagus) is shorter in C. lean. comb., C. philippan. comb. & reinst. stat., and the two new species (C. castanyan. sp. and C. esmeraldan. sp.), compared with C. fidan. comb. and C. sanmarcosn. comb. Furthermore, the lamella antevaginalis is more rectangular and smaller compared with the lateral plate of the eighth abdominal segment and the lateral margin of the lamella antevaginalis is not fused with this plate in C. lean. comb., C. philippan. comb. & reinst. stat., C. castanyan. sp., and C. esmeraldan. sp., whereas this structure is larger and more rounded with a concave posterior margin in ventral view in C. fidan. comb. and C. sanmarcosn. comb., and notably both sides of the lamella antevaginalis are fused with the lateral plate of the eighth abdominal segment in these two taxa. Despite the immature stages being known for only two species, the final instar larva of C. philippan. comb. & reinst. stat. and C. castanyan. sp. both possess rounded, stubby, and blunt head scoli, transforming from the ‘antler-like’ bifurcating head scoli of the penultimate instar, whereas most other known last instar larva of euptychiines exhibit developed horn-like scoli throughout the last two instars or simply lack scoli, such as species in the genus Hermeuptychia Forster, 1964 (e.g., Cong & Grishin 2014).

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 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 m1-m2 (i.e., dcs) curved basally, m2-m3 (i.e., dcm) appearing rather straight; hindwing humeral vein present, exiting from origin of Rs and curving; origin of M2 slightly towards M1 than M3 (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. lean. comb. and C. esmeraldan. 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).

Fig. 2.

Cisandinan. gen. adults, C. lean. 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. esmeraldan. sp., e) male holotype in dorsal view (left), ventral view (right); (f) female paratype in dorsal view (left), ventral view (right); C. philippan. 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. castanyan. 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 fidan. comb.: (k) male paralectotype in dorsal view (left), ventral view (right); (l) female lectotype in dorsal view (left), ventral view (right); C. fida directan. 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. sanmarcosn. 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. trinitesisn. 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.

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 semi-iridescent 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 Cu2, fading and terminating in cell Cu2; concolorous postdiscal band extending from radial vein towards inner margin, terminating at 2A if not already faded in Cu2, 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 M1) to cell Cu2, 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 M1, 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 R5, M2, M3, and Cu1.

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 M2 and/or Cu1, 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, M1, M2, M3, Cu1, and Cu2 (ocellus in cell Cu2 absent in C. trinitensisn. comb.), ocelli in cells M1 and Cu1 similar in size (as in C. castanyan. sp., C. fidan. comb.,C. sanmarcosn. comb., and C. esmeraldan. sp.) or ocellus in Cu1 may be larger (as in C. trinitensisn. comb., C. lean. comb., and C. philippan. comb. & reinst. stat.), both ocelli often possessing black central area and single white pupil ringed in yellow, ocelli in cells Rs and Cu2 smaller compared with previous two ocelli but general appearance is same, ocelli in cells M2 and M3 are similar in size and appearance to those in cells Rs and Cu2 but lacking black area, ocellus in cell M2 placed more basally compared with ocellus in cell M3.

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. lean. 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).

Fig. 3.

Cisandinan. gen. male genitalia, C. lean. 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. esmeraldan. 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. philippan. 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. castanyan. 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. fidan. 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. sanmarcosn. 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. trinitesisn. 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. sanmarcosn. comb. are reproduced from Nakahara et al. (2018a).

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. lean. comb., C. esmeraldan. sp., C. fidan. comb., and C. sanmarcosn. 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. fidan. comb. and C. sanmarcosn. 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).

Fig. 4.

Cisandinan. gen. female genitalia, C. lean. 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. philippan. 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. castanyan. 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. fidan. 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. sanmarcosn. comb.: (i) female genitalia without abdomen illustration; (j) illustration indicating location of signa on corpus bursae to left; (k) ventral view of lamella antevaginalis with intersegmental membrane of seventh and eighth abdominal segments expanded ([j and k] based on dissection SN-16-17); C. trinitensisn. comb.: (l) dorsal view of genitalia with intersegmental membrane of seventh and eighth abdominal segments folded; illustration showing arrangement of signa to right (not to scale); (m) ventral view of lamella antevaginalis with intersegmental membrane of 7th and 8th abdominal segments expanded ([l and m] based on dissection SN-20-85). Scale bar = 1 mm. Drawings of C. sanmarcosn. comb. are reproduced from Nakahara et al. (2018a).

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:

Cisandinan. 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 lean. 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. philippan. comb. & reinst. stat. and C. castanyan. sp., including its natural host plant (Figs. 6 and 7), which constitute to date the only reliable immature stage data available for this genus.

Fig. 5.

Cisandina lean. comb. wing venation, male to the left, female to the right. Scale bar = 10 mm.

Fig. 6.

Cisandinan. gen. immature stages, C. philippan. 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. philippan. comb. & reinst. stat., Taquara micrantha (Poaceae) in situ; (d) inflorescence material of (c); C. castanyan. 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. castanyan. sp., Olyra latifolia (Poaceae) in situ. Scale bar = 1 mm for a, b, e-g; 5 mm for h, i.

Fig. 7.

Cisandina philippan. 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 castanyan. 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 Cisandinan. gen. taxa discussed herein.

Cisandina lea (Cramer, 1777), New Combination
(Figs. 1, 2a–d, 3a–d, 4a and b, 5, 8)

• Papilio leaCramer, 1777: 87, pl. 151, figs. C, D. Lectotype, designated herein.

• =Papilio juniaCramer, 1780: 9, pl. 292, figs. D, E. Lectotype, designated herein.

• Satyrus lea: Godart [1824]: 464, 492.

• Satyrus junia: Godart [1824]: 492.

• Euptychia lea: Westwood 1851: 373, Butler 1868: 29, Kirby 1871: 52, Butler 1877: 121, Weymer 1911: 216, pl. 48, fig. f, Gaede 1931: 452, D’Abrera 1988: 768–769, figs.

• Euptychia junia: Westwood 1851: 373, Butler 1868: 30, Kirby 1871: 52, Butler 1877: 121, Kirby 1879: 135, Forster 1964: 128, Lamas 2004: 220.

• Magneuptychia lea: Forster 1964: 128, Lamas 2004: 220, Beccaloni et al. 2008: 335.

• Euptichia [sic] junia: Geyer 1832: 12, pl. [109], figs. 627, 628.

• Euptychia lea f. junia: Weymer 1911: 216.

• Euptychia lea var. junia: Gaede 1931: 452.

• Cissia junia: Singer and Ehrlich 1993: 251, fig. 1.

Systematic placement and diagnosis:

Our phylogenetic hypothesis places Cisandina lean. comb. as sister to C. esmeraldan. sp., with strong support (Fig. 1; SH-aLRT/UFBoot = 96.5/96). The infra-specific genetic divergence based on DNA barcodes between the two sequenced individuals of C. lean. comb. from two localities (LEP-34359 from French Guiana; BC-DZ-138 from Amazonas, Brazil) was 0.84%, whereas interspecific divergences between C. lean. comb. and three sampled specimens of C. esmeraldan. sp. range from 5.01 to 5.3%. These three individuals of C. esmeraldan. sp. are from two sites, including BC-DZ-137 (holotype) from Paraná, Brazil, and the other two specimens from Misiones, Argentina. The infraspecific genetic divergences among these three sequenced individuals of C. esmeraldan. sp. were 0.02%. See Table 3 for further information regarding genetic divergence among Cisandinan. gen. taxa. The male of C. lean. comb. is readily distinguished from C. esmeraldan. sp. by its iridescent to semi-iridescent bluish flush covering the DFW and DHW, whereas the DFW and DHW discal cell and adjacent area is greenish in male C. esmeraldan. sp. The broader ventral bands and larger ventral submarginal ocelli of C. lean. sp. can be used to distinguish both sexes of these two closely related species. The female of C. lean. comb. is also distinguished from the female of C. esmeraldan. sp. by the presence of iridescent bluish lilac reflection on the DHW, whereas this reflection appears more purplish in the female of C. esmeraldan. sp., and the ventral bands and ocelli overall appear larger in C. lean. sp. compared with its sister species.

Taxonomy:

Papilio lea was described by Pieter Cramer, a Dutch merchant, in his De Uitlandsche Kapellen series (work completed by Caspar Stoll) based on an unspecified number of specimen(s) from ‘Berbices’, a then Dutch colony (i.e. Dutch Guiana) along Rio Berbice, an area now part of the Republic of Guyana. The original description only states ‘the blue color on both sides of the wings is shiny’. Nevertheless, Cramer provided illustrations of both wing surfaces of this taxon (pl. 151, figs. C, D), which can be used to narrow down the identity of P. lea. His drawings of this species clearly show the basal half of its DFW and all of the DHW being blue, although the anterior side of the DHW is not visible due to being covered by the forewing. The hindwing illustration shows six individual submarginal ocelli on the VHW in cells Rs, M1, M2, M3, Cu1, and Cu2, with the ocellus in cell M2 placed more basally compared with that in cell M3, as well as a single ocellus present on the VFW. There is some bluish coloration visible on the DFW and DHW, especially along the ventral bands and around the VHW submarginal ocelli. The presence of six VHW ocelli excludes all Neotropical satyrine species with blue DFW and/or DHW, in genera such as Caeruleuptychia Forster, 1964, Magneuptychia, Chloreuptychia and Amiga Nakahara, Willmott, & Espeland 2019, as candidates for the species under description by Cramer, because all the species in those genera which possess bluish coloration only have five VHW ocelli, with the exception of a few species in Caeruleuptychia. Caeruleuptychia twalela Brévignon, 2005, Cae. pilata (Butler, 1867), and Cae. scripta Nakahara, Zacca, & Huertas, 2017 are the only three Caeruleuptychia species with bluish scales and six VHW ocelli, but the dorsal surface of Cae. twalela is entirely brown; the orangish rings of the VHW ocelli in cells M1 and Cu1 are skewed (i.e., not evenly broad) in Cae. pilata and Cae. scripta, whereas the rings of these ocelli are evenly broad in Cramer’s illustration. Perhaps one of the most notable features in Cramer’s drawing is the VHW ocellus in cell M2 being placed more basally compared with that in cell M3, a distinctive feature of Cisandinan. gen. This character excludes the possibility of Cramer referring to virtually any other euptychiine species apart from those discussed herein. Those few exceptions include some species in Moneuptychia Forster, 1964 such as M. vitellina Freitas & Barbosa, 2015, which can easily be distinguished by the lack of bluish coloration on the wing surface. Thus, Cramer’s P. lea represents either the female of P. lea as conceived herein, possibly C. philippa n. comb. & reinst. stat. (in which some females have bluish dorsal coloration), or one of the new species described and named in this study. However, it is very unlikely that Cramer’s illustration represents C. castanyan. sp., C. esmeraldan. sp., or C. philippan. comb. & reinst. stat., since none of these species are known from the Guianas.

Our attempt to locate syntype(s) of P. lea resulted in finding three candidate specimens in RMNH, two females each with a label indicating ‘[Johan] Calkoen’ with the locality ‘Surinam[e]’ (RMNH.INS 967256 and 967257), and a male specimen with a round label indicating ‘Verloren [van Themaat] Brasil’ (RMNH.INS 967280). Two collections now housed at RMNH, namely Joan Raye Heer van Breukelerwaard’s and Johan Calkoen’s collections, are both known to include Cramer types (de Jong 1982, Smit et al. 1986, Gernaat et al. 2012), and the former pair from ‘Surinam[e]’ are evidently from the latter collection. The latter male from the Verloren van Themaat collection can also be regarded as a Cramer type based on information from some existing literature on his collection (e.g., Roepke 1941). Verloren van Themaat purchased van Eyndhoven’s collection that was believed to have been sold on 14 October 1861 (see Chainey 2005; Smit et al. 1986) which is another collection believed to have included Cramer specimens (Horn and Kahle 1935, Roepke 1941). Despite the fact that Verloren van Themaat’s collection is supposed to have been destroyed due to an unfortunate battle when the collection was kept near Utrecht (Chainey 2005), we cannot exclude the possibility of some specimens ending up at Leiden where the RMNH is situated. Although lacking the abdomen, the female specimen (RMNH.INS 967257) does overall match Cramer’s illustration of P. lea, with minor differences, such as bluish scales on the DFW being somewhat restricted to cells Cu2 and 2A (not covering the basal half as in fig. C), but this is likely due to the color being faded over time. Nevertheless, its locality ‘Surinam[e]’ historically never included ‘Berbice’ (the type locality of P. lea), which was then referred to as Dutch Guiana, and thus, it is debatable whether this female represents a syntype of P. lea or not. Confusingly, Butler (1877: 121) listed ‘Surinam[e]’ as the type locality for Euptychia lea (=P. lea), without any apparent evidence as to whether this was a misinterpretation or not. The only clue as to the whereabouts of the syntypes is contained in the last part of Cramer’s description ‘[The specimen(s)] now rests in the previous collection’, referring to the last part of the description of the immediately preceding taxon, ‘Merope’, described on the same page, ‘She rests in the Cabinet (=collection) of the honorable gentleman A. Gevers’. As mentioned by Chainey (2005), this ‘A. Gevers’ most likely refers to Abraham Paulusz Gevers (1712–1780), then the Mayor of Rotterdam, who had a natural history collection which was auctioned in 1787. Despite having no evidence of acquisition, specimens from his collection were perhaps bought by Calkoen and ended up in the RMNH, thus there exist a possibility that the provenance of the aforementioned specimen from ‘Surinam[e]’ is the Gevers collection, but with a changed locality.

Three years later, Cramer described Papilio junia, a taxon now regarded as a junior subjective synonym of Papilio lea by Lamas (2004), again in his De Uitlandsche Kapellen. Like his description of Papilio lea, the number and sex of the examined specimen(s) used to describe P. junia cannot be unambiguously determined from the original description, except for the type locality being ‘Surinam[e]’. Cramer’s description is again terse, and his Dutch and French texts are accompanied with illustrations of both dorsal and ventral surfaces (pl. 292, figs. D, E). As stated in the text, the DFW and DHW are entirely bluish (described as ‘cerulean satin-like shine’ by Cramer), with his paintings reflecting a lighter blue color relative to the darker blue coloration of the dorsal wing shown in his illustration of P. lea. The general wing pattern of the ventral surface is similar to that of his illustration of P. lea, with some minor differences such as a paler ground color, more elongated VHW submarginal ocelli, and bluish coloration on the VHW restricted to the inner margin. It must be noted that a number of painters were involved in producing copies of Cramer’s work, and there is some evidence that inevitably some variability in markings and coloration resulted, in comparison with his original drawings (Chainey 2005). Thus, these subtle dissimilarities mentioned above may not actually reflect the differences observed by Cramer himself. However, it is worth noting the small incomplete ocelli in VFW cells M2 and M3 visible in the illustration of P. junia, which are especially apparent on the right wing, and which appear to be absent in his illustration of P. lea. The male specimen from the Verloren collection mentioned above (RMNH.INS 967280) does exhibit ocelli on the right VFW in cells M2 and M3, in addition to having an extra ocellus above M1. The male specimen from the Calkoen collection also possesses a tiny smudge-like ocellus in the right VFW cell M2, although apparently the ocellus is absent in cell M3. Since its description, the taxonomic viewpoint of subsequent authors concerning P. junia has been variable, ranging between some regarding it as a valid species (e.g., Butler 1877, Forster 1964) and some considering it to be conspecific with P. lea (e.g., Weymer 1911, Lamas 2004). Despite this unstable taxonomic status, it is noteworthy that these names have rarely been applied to other euptychiine species in museum collections and/or scientific articles, with few exceptions (e.g., D’Abrera 1988), an unusual situation for a group where misidentification is extremely common. Specifically, the male specimen of this taxon, as painted by Cramer (pl. 292, figs. D, E), is distinctive and is less likely to be misidentified compared with the female which can be confused with species in the same genus such as C. castanyan. sp. Recently, Nakahara et al. (2019a) did not accept Calkoen specimens (also housed in the RMNH) as potential syntypes of Papilio ebusa Cramer 1780 because of the discrepancy in terms of the type locality indicated on the associated label. Unlike this previous case, the handwritten locality on the rounded label for RMNH.INS 967256 does match the type locality indicated in Cramer’s description of P. junia, both being ‘Surinam[e]’. Thus, there seems to be no reason to reject this male specimen as a syntype, and thus we here designate this male syntype as the lectotype for P. junia with the following labels separated by double-forward slashes (lectotype designation): //Calkoen Suriname// RMNH.INS 967256//. Accepting this male specimen as a former syntype of P. junia reinforces the possibility of the aforementioned female specimen (RMNH.INS 967257) being a syntype of P. lea with an erroneous locality on the label. The fact that only a single female for a possible Cramer syntype was located at RMNH matches the fact that Cramer illustrated a female for P. lea and a male specimen for P. junia. Therefore, we designate this female from the Calkoen collection as the lectotype of P. lea with the following labels separated by double-forward slashes (lectotype designation): //Calkoen Suriname// RMNH.INS 967257//.

Distribution and natural history:

Cisandina lean. comb. is known from the Guianas, Trinidad, southern Venezuela, as well as the central and lower Amazon in Brazil, in addition to northeastern Brazil (Fig. 8).

Specimens examined (129 ♂, 80 ♀):

Brazil: Alagoas: São José de Lage, Usina Serra Grande, [8°58′S,36°3′W], 400–500 m, 1–3 Aug 2003, 1 ♀, (ZUEC); Maceió, [9°39′S,35°46′W], (Cardoso, A.), 30 Apr 1945, 1 ♀, (DZUP); Amazonas: Barcelos, Rio Aracá, Foz do Rio Curuduri, [0°5′50″S,63°17′22″W], (Mielke, O. H. H., Casagrande, M. M.), 15–19 Jun 2010, 1 ♀, (DZ 49.955, BC-DZ 138) (DZUP); Ega (= Tefé), [3°22′S,64°42′W], (Bates, H. W.), 1 ♀ [BMNH(E)- 1497637], (NHMUK); Manaus, [3°7′S,60°2′W], (Hahnel), 1886, 1 ♀, (MNHU); Manicoré, [5°49′S,61°17′W], (Le Moult, E.), 1 ♂ [FLMNH-MGCL-265701], (FLMNH); Maués, [3°22′S,57°43′W], (Le Moult, E.), 1 ♂ [FLMNH-MGCL-265699], 1 ♀ [FLMNH-MGCL-265703], (FLMNH); Rio Negro, 50-km NW Manaus, (Schmidt, U.), 1–4 Nov 1993, 1 ♀ [FLMNH-MGCL-265705], [dissection, SN-20-31] (FLMNH); Bahia: [12°59′S,38°31′W], 1 ♂ [BMNH(E)-1497632], (NHMUK); Espírito Santo: [20°20′S, 40°17′W], 1 ♂ [BMNH(E)-1497649], 1 ♂ [BMNH(E)-1497650], 1 ♂ [BMNH(E)-1497651], (NHMUK); Pará: [Rio] Tapajós, [4°16′8″S,55°59′10″W], 25 m, (Bates, H. W.), 1 ♂ [BMNH(E)-1497630], (NHMUK); Amazonas, (Fassl, A. H.), (ZSM); Bragança, [1°3′S,46°47′W], (Mathan, M. de), 1 ♀ [BMNH(E)- 1497641], (NHMUK); Breves, [1°40′S,50°28′W], 1 ♀ [BMNH(E)- 525609], (NHMUK); hwy Cuiabá-Santarém, km 1666, [3°17′17″S,54°56′22″W], -150, (Callaghan, C. J.), 28 Jul 1978, 1 ♂ [FLMNH-MGCL-265697], (FLMNH); hwy Cuiabá-Santarém, km 958, [8°3′S,55°2′W], (Callaghan, C. J.), 30 Jul 1978, 1 ♀ [FLMNH-MGCL-265706] [dissection, SN-20-17], (FLMNH); Itaituba on Rio Tapajós, [4°16′8″S,55°59′10″W], (Le Moult, E.), 1 ♂ [FLMNH-MGCL-265702], [dissection, SN-20-16] (FLMNH); Itaituba, [4°17’S,55°59’W], (Mich.), 1890, 1 ♀ [dissection, Lee D. Miller 9136], (MNHU), 1893, 1 ♂ [dissection, Lee D. Miller 9135], (MNHU); Itaituba, Rio Tapajós, [4°16′8″S,55°59′10″W], 25 m, (Lathy), May 1932, 1 ♂ [BMNH(E)-1670284], (NHMUK); Óbidos, [1°54′S,55°31′W], (Le Moult, E.), 1 ♂ [FLMNH-MGCL-265700], (FLMNH); Pará, [1°0′S,51°11′W], 1 ♂ [BMNH(E)-1497629], 1 ♀ [BMNH(E)-1497640], (NHMUK), (Bates, H. W.), 1 ♀ [BMNH(E)- 1497638], (NHMUK), (Wallace), 1 ♀ [BMNH(E)-1497639], (NHMUK); Rio Tapajós, [4°16′8″S,55°59′10″W], 25 m, (Klug, G. G.), Chainey 2005 Cramer ep 1931, 1 ♀ [BMNH(E)-1497729], (NHMUK); Rio Tapajós, Mujo, [2°41′S,54°38′W], (Le Moult, E.), 1 ♂ [FLMNH-MGCL-265698], (FLMNH), Sep 1922, 1 ♀ [FLMNH-MGCL-265704], (FLMNH); Santarém, [2°26′S,54°43′W], (Smith, H. H.), 1 ♂ [BMNH(E)-1497631], 1 ♀ [BMNH(E)-1497642], 1 ♀ [BMNH(E)-1497643], (NHMUK); Roraima: Alto Alegre, Ilha de Maracá, [3°21′41″N,61°26′2″W], (Mielke, O. H. H., Casagrande, M. M.), 24–31 Aug 1987, 3♂ 1 ♀(DZ 49.925, DZ 49.935 - BC-DZ Willmott 191, DZ 49.945, DZ 49.965, 23–28 Feb 1988, 2 ♀ (DZ 5371, DZ 49975) (DZUP); Not located: ‘Brasilia’, 1 ♂ [BMNH(E)-1497635], (NHMUK); ‘Brazil’, 1 ♂ [BMNH(E)-1497633], 1 ♂ [BMNH(E)-1497634], 1 ♂ [BMNH(E)-1497636], 1 ♂ [BMNH(E)-1497652], 1 ♂ [BMNH(E)-1497653], 1 ♂ [BMNH(E)- 1497654], 1 ♀ [BMNH(E)-1497647], 1 ♀ [BMNH(E)-1497648], (NHMUK); not located: ‘Amazon’, 1 ♂ [BMNH(E)-1497714], 1 ♂ [BMNH(E)-1497727], (NHMUK), (Bates, H. W.), 1 ♀ [BMNH(E)-1497730], (NHMUK) [Bates only collected in Brazil]. French Guiana: Cayenne: Cayenne, [4°56′N,52°20′W], 1 ♂ [BMNH(E)-787669; (NHMUK); Saint-Laurent-du-Maroni: Bas Maroni, 1 ♂ [FLMNH-MGCL-265707; dissection, KW-14-018], (FLMNH); Maroni River, (Le Moult, E.), 1 ♀ [FLMNH-MGCL-265708], (FLMNH); Maroni river, Maripasoula, [3°41′N,54°2′W], (Brévignon, C.), 14 May 1987, 1 ♂ [MUSM-LEP-103094], (MUSM); Saül, [3°51′30″N,53°18′14″W], 200–450 m, (Nakahara, S.), 5 Aug 2014, 1 ♂ [FLMNH-MGCL-209436], 1 ♀ [FLMNH-MGCL-195747], (FLMNH); St. Laurent du Maroni, [5°30′N,54°2′W], Jul-Sep 1915, 1 ♂ [BMNH(E)-1497720], (NHMUK), (Le Moult), Nov, 1 ♂ [BMNH(E)-1497721], (NHMUK); Not located: ‘French Guiana’, (Bar, C.), 1 ♂ [BMNH(E)-1497722], 1 ♂ [BMNH(E)-1497723], (NHMUK). Guyana: Cuyuni-Mazaruni: Bartica, [6°24′N,58°37′W], (Parish, H. S.), 1 ♂ [BMNH(E)-787654], 1 ♀ [BMNH(E)-787665], (NHMUK), (Parish, H.S.), 1 ♂ [BMNH-E-787654], 1 ♀ [BMNH-E-787665], (NHMUK); East Berbice-Corentyne: New River Triangle, Camp Jaguar, [3°18′23″ N,57°35′21″W], 152 m, (Steinhauser, S. R.), 13 Nov 1980, 1 ♀ [FLMNH-MGCL-265710], (FLMNH); Upper Demerara-Berbice: Berbice, [5°27′N,57°57′W], 1 ♂ [BMNH(E)-787651], 1 ♂ [BMNH-E-787651], (NHMUK); Upper Takutu-Upper Essequibo: E. Kanuku Mountains, Two Hat Mountain, [3°8′48″N,59°6′54″W], 244 m, (Fratello, S., et al), 17 Sep–2 Oct 2000, 1 ♂, (USNM); Essequibo River, (Büche, M.), Jul 1997, 1 ♂ [MUSM-LEP-103093], (MUSM); Essequibo River, Aunai, [5°22′N,58°53′W], (Whitely, H.), 1 ♂ [BMNH(E)-787650], 1 ♂ [BMNH-E-787650], 1 ♀ [BMNH(E)-787664], 1 ♀ [BMNH-E-787664], (NHMUK); Kanuku Mountains, [3°12′N,59°34′W], 152–305 m, (Fratello, S., Hanner, R., Hendricks, S., Williams, R.), 21 Feb–10 Mar 1999, 1 ♂, (USNM); Lethem, (Le Moult, E.), 8 Aug 1971, 1 ♀ [FLMNH-MGCL-265709], (FLMNH); Not located: ‘Guyana’, 1 ♂ [BMNH(E)-787652], 1 ♂ [BMNH-E-787652], 1 ♀ [BMNH(E)-1497716], 1 ♀ [BMNH-E-787669], (NHMUK), (Parish), 1 ♀ [BMNH-E-787667], (NHMUK), (Parish, H. S.), 1 ♀ [BMNH(E)-787667], (NHMUK), (Whitely), 1 ♀ [BMNH-E-787666], (NHMUK), (Whitely, H.), 1 ♀ [BMNH(E)-787666], (NHMUK); Demerara River, Akayma Fort, 1 ♀ [BMNH(E)-1497715], (NHMUK). Suriname: Brokopondo: Berg-en-Dal, [5°9′N,55°4′W], (Ellacombe, C. W.), Apr 1892, 1 ♂ [BMNH(E)-787668], (NHMUK), (Ellacombe, C.W.), Apr 1892, 1 ♀ [BMNH-E-787668], (NHMUK); Para: Bersaba, [5°32′N,55°3′W], (Michls.), 1898–1899, 1 ♂ [dissection, M-9137 Lee D. Miller], (MNHU); Not located: ‘Surinam’, 1 ♂ [BMNH(E)-1497718], 1 ♂ [BMNH(E)-1497719], 1 ♂ [BMNH(E)-787656], (NHMUK), (Fruhstorfer), May–Sep, 1 ♂ [BMNH-E-787656], (NHMUK); ‘Surinam’, (ZSM); ‘interior Surinam’, (Ellacombe, C. W.), Sept 1892, 1 ♂ [BMNH(E)-1497717], (NHMUK). Trinidad: Couva-Tarabaquite-Talparo: Caparo, [10°34′N,61°16′W], (Birch, F.), 1 ♀ [BMNH-E-787662], (NHMUK); Narieva, Tabaquite, [10°23′N,61°18′W], 1 ♂ [BMNH(E)-1497678], 1 ♂ [BMNH(E)- 1497679], 1 ♀ [BMNH(E)-1497671], 1 ♀ [BMNH(E)-1497672], (NHMUK); Diego Martin: Fort George, [10°42′N,61°32′W], Sept. 1891, 1 ♂ [BMNH(E)-1497660], 1 ♂ [BMNH(E)-1497661], 1 ♂ [BMNH(E)-1497662], 1 ♂ [BMNH(E)-1497663], 1 ♂ [BMNH(E)-1497664], 1 ♂ [BMNH(E)-1497665], 1 ♀ [BMNH(E)- 1497666], 1 ♀ [BMNH(E)-1497667], (NHMUK); Mt. Catherine upper trail, (Preston, J. & F.), 27 Feb 1982, 1 ♀ [FLMNH-MGCL-265722], 1 ♀ [FLMNH-MGCL-265723], (FLMNH); Port of Spain: Port of Spain, [10°40′N,61°31′W], (Jenkins, D. W.), 27 Jan 1977, 1 ♂ [FLMNH-MGCL-265715], (FLMNH), (Rosen, V.), 23 Oct, (ZSM); Rio Claro-Mayaro: ‘Nariva’, 06-05-1993, 1 ♂ [BMNH(E)-1670212; ‘Swamp’], (NHMUK); San Juan-Laventille: Fondes Amandes Road, (Patterson, E. J.), 1 ♀ [FLMNH-MGCL-265724], (FLMNH); Hololo Mt., [10°41′29″N,61°29′7″W], (Morrall, J.), 4 Oct 2012, 1 ♂, (MZUJ); Maraval, [10°43′N,61°31′W], Jan ‘92, 1 ♂ [BMNH(E)-1497680], 1 ♂ [BMNH(E)-1497681], 1 ♀ [BMNH(E)-1497684], 1 ♀ [BMNH(E)-1497685], (NHMUK), (Hall, A.), Nov 1931–Feb 1932, 1 ♂, (BMB); St. Annes Valley, [10°41′N,61°30′W], 1 ♂ [FLMNH-MGCL-265718], (FLMNH), 1 ♂ [BMNH(E)-787646], 1 ♂ [BMNH(E)-787647], 1 ♂ [BMNH(E)- 787648], (NHMUK); St. Anns, [10°41′N,61°30′W], 1 ♂ [BMNH-E- 787646], 1 ♂ [BMNH-E-787647], 1 ♂ [BMNH-E-787648], (NHMUK), (Hall, A.), Nov–Dec 1931, 1 ♀, (BMB); Tunapuna-Piarco: 6 mi. N Arima, [10°42′12″N,61°17′28″W], 300 m, (Pliske, T. E.), 30 Jun 1962, 1 ♂ [FLMNH-MGCL-265720], (FLMNH); Arima Valley, [10°41′N,61°17′30″W], 305–457 m, (Breedlove, D. E.), 14–19 Dec 1970, 1 ♀, (CAS); Tunapuna, [10°39′7″N, 61°23′17″W], 50 m, 1 ♂ [BMNH(E)-1497690], 1 ♀ [BMNH(E)- 787660], 1 ♀ [BMNH-E-787660], (NHMUK); Not located: ‘Trinidad’, [10°26′17″N,61°15′12″W], 1 ♂ [BMNH(E)-1497704], 1 ♂ [BMNH(E)-1497705], 1 ♂ [BMNH(E)-1497707], 1 ♂ [BMNH(E)-1497708], 1 ♂ [BMNH(E)-1497709], 1 ♂ [BMNH(E)- 1497710], 1 ♂ [BMNH(E)-1497711], 1 ♂ [BMNH(E)-1497712], 1 ♂ [BMNH(E)-787643], 1 ♂ [BMNH(E)-787655], 1 ♂ [BMNH-E-787643], 1 ♂ [BMNH-E-787644], 1 ♂ [BMNH-E- 787655], 1 ♀ [BMNH(E)-1497696], 1 ♀ [BMNH(E)-1497697], 1 ♀ [BMNH(E)-1497699], 1 ♀ [BMNH(E)-787657], 1 ♀ [BMNH(E)- 787658], 1 ♀ [BMNH-E-787657], 1 ♀ [BMNH-E-787658], (NHMUK), (Feather), 1 ♂ [BMNH(E)-1497706], 1 ♀ [BMNH(E)- 1497688], (NHMUK), (Fountaine, M.), Dec. 1911, 1 ♂ [BMNH(E)-1497693], 1 ♂ [BMNH(E)-1497694], 1 ♂ [BMNH(E)- 1497695], 1 ♂ [BMNH(E)-1497702], 1 ♂ [BMNH(E)-1497703], 1 ♂ [BMNH(E)-787653], 1 ♀ [BMNH(E)-1497687], 1 ♀ [BMNH(E)- 1497698], 1 ♀ [BMNH(E)-1497700], 1 ♀ [BMNH(E)-1497701], 1 ♀ [BMNH(E)-787661], 1 ♀ [BMNH(E)-787663], (NHMUK), Dec 1911, 1 ♂ [BMNH-E-787653], 1 ♀ [BMNH-E-787661], 1 ♀ [BMNH-E-787663], (NHMUK), (Hall, A.), Jan 1936, 1 ♂, 1 ♂ [Booth Mus: Collection 00-5938], (BMB), Oct-Dec 1920, 1 ♂, (BMB), (Kaye, W. J.), May 1898, 1 ♂ [FLMNH-MGCL-265719], (FLMNH), May 1898, 1 ♂ [FLMNH-MGCL-265716], (FLMNH), (Neuburger), (ZSM); Arima Valley, vicinity Asa Wright Nature Centre vic, [10°43′3″N,61°17′55″W], (Austin, G.T.), Feb 1993, 1 ♂ [FLMNH-MGCL-296554], [dissection, SN-20-30] (FLMNH), (Gomes, O.), Feb 1993, 1 ♀ [FLMNH-MGCL-298000], (FLMNH); Behind St. Benet’s Hall, (Preston, J. & F.), 2 Jan 1982, 1 ♀ [FLMNH-MGCL-265721], (FLMNH); Caparo, (Birch, F.), 4th Aug., 1 ♀ [BMNH(E)-1497683], (NHMUK), 4th Sept. ‘04, 1 ♀ [BMNH(E)- 787662], (NHMUK), (Klages, S.M.), Jan 1906, 1 ♂ [BMNH(E)-1497682], (NHMUK); Northern Mts., (Hall, A.), Dec 1938-Jan 1939, 1 ♀, (BMB); Port of Spain, (Rendall), II.97, 1 ♂ [BMNH(E)-1497691], 1 ♀ [BMNH(E)-1497686], (NHMUK); ‘Saint George’, (Ellacombe, C.W.), 1 ♂ [BMNH-E-787645], 1 ♀ [BMNH-E-787659], (NHMUK); Santa Margarita, (Preston, J. & F.), 13 Oct 1981, 1 ♂ [FLMNH-MGCL-265717], (FLMNH); St. George, (Ellacombe, G. W.), 1 ♂ [BMNH(E)-787645], 1 ♀ [BMNH(E)-787659], (NHMUK), Dec. 1891, 1 ♀ [BMNH(E)-1497668], 1 ♀ [BMNH(E)-1497669], 1 ♀ [BMNH(E)-1497760], (NHMUK), Oct. 1891, 1 ♂ [BMNH(E)-1497673], 1 ♂ [BMNH(E)-1497674], 1 ♂ [BMNH(E)-1497675], 1 ♂ [BMNH(E)-1497676], 1 ♂ [BMNH(E)-1497677], (NHMUK); Stanway Parris River, Feb 1921, 1 ♂ [BMNH(E)-1497692], (NHMUK); Symonds Valley, (Hall, A.), Apr 1930, 1 ♂, (BMB), Mar 1930, 1 ♀, (BMB). Venezuela: Amazonas: Samariapo, 120 m, (Lichy, R.), 7 Oct 1946, 1 ♀ [FLMNH-MGCL-265714], (FLMNH); Bolívar: 80 km S El Dorado, [6°11′8″N,61°24′36″W], (Nation, J. L.), 26 Jun 1984, 1 ♂ [FLMNH-MGCL-265712], (FLMNH); Not located: ‘Venezuela’, 1 ♂ [BMNH(E)-1497659], 1 ♂ [BMNH(E)-1497713], (NHMUK); Pitotan, (Nation, J. L.), 9 Jun 1937, 1 ♂ [FLMNH-MGCL-265713], (FLMNH). Country unknown: no data, 1 ♀ [FLMNH-MGCL-265711], (FLMNH), 1 ♂ [BMNH(E)-1497655], 1 ♂ [BMNH(E)-1497656], 1 ♂ [BMNH(E)-1497724], 1 ♂ [BMNH(E)- 1497725], 1 ♂ [BMNH(E)-1497726], 1 ♂ [BMNH(E)-787649], 1 ♂ [BMNH-E-787649], 1 ♀ [BMNH(E)-1497728], (NHMUK), (Hall, A.), 18 Mar 1934, 1 ♀, (BMB). Doubtful locality: ‘Cauca valley’, 1 ♂ [BMNH(E)-1497657], (NHMUK). ‘Haiti’, 1 ♂ [BMNH(E)-1497731], 1 ♂ [BMNH(E)-1497732], (NHMUK).

Cisandina esmeralda Nakahara & Barbosa, New Species
(Figs. 1, 2e and f, 3e–g, 8)

Euptychia lea [misidentification]: D’Abrera 1988: 768–769, figs.

Systematic placement and diagnosis:

As discussed under the immediately preceding taxon, C. esmeraldan. sp. is recovered as sister to C. lean. comb. with a strong support (Fig. 1; SH-aLRT/UFBoot = 96.5/96). Our molecular phylogeny and DNA sequence divergence between these two species support its species-level status indicated by its distinctive male phenotype. Indeed, the iridescent greenish lilac scales possessed by the male of C. esmeraldan. sp. readily distinguish this species from the remainder of the genus and furthermore from all other known euptychiines. In addition, the more brownish ventral ground color, small ventral submarginal ocelli (smallest among all Cisandinan. gen. species), narrow and somewhat sinuate VHW postdiscal band, coupled with other narrow ventral bands (narrowest among all four similar-appearing Cisandinan. gen. species with iridescent scales) separate the male of this species from the other three Cisandinan. gen. species with iridescent scales discussed herein. The more elongate forewing is a character shared with the male of C. lean. comb., but not with males of other taxa discussed herein. The female of C. esmeraldan. sp. is distinguished from the female specimens of C. lean. comb. and C. castanyan. sp. by its lack of purple lilac scales on the dorsal surface thus the female of C. esmeraldan. sp. possesses a uniformly brown dorsal wing surface. Cisandina philippan. comb. & reinst. stat. is similar in this respect because the female of C. esmeraldan. sp. Is brown dorsally. Nevertheless, the narrower ventral bands and smaller ventral submarginal ocelli of C. esmeraldan. sp., as well as its smaller adult size, can be used to separate females of these two species. Furthermore, the outward-curving VFW postdiscal band of the female C. esmeraldan. sp. is rather distinctive. See also ‘Remarks’ below for further information regarding the identity of the female of this species.

Male (Fig. 2e):

Forewing length 23 mm (n = 1).

Head:

See head description for Cisandinan. gen. above.

Thorax:

See thorax description for Cisandinan. gen. above.

Abdomen:

See abdomen description for Cisandinan. gen. above, eighth sternite appearing as two separate, somewhat broader patches.

Wing venation:

See wing venation description for Cisandinan. gen. above.

Wing shape:

See wing shape description for Cisandinan. gen., notable features include forewing appearing elongate due to angle between forewing outer margin and inner margin being obtuse.

Dorsal forewing:

Notable features include, ground color brownish, iridescent greenish lilac scales covering most of DFW except for area anterior of costal vein (Fig. 2e), apex and distal side of cells M1, M2, M3, Cu1, although this area revealing ground color narrowins posteriorly; see also DFW description for Cisandinan. gen. above.

Dorsal hindwing:

Notable features include, ground color similar to forewing, iridescent greenish lilac scales visible in discal cell and adjacent area just outside of this cell (Fig. 2e); see also DHW description for Cisandinan. gen. above.

Ventral forewing:

Notable features include ventral bands narrow, discal band and postdiscal band parallel to each other and traversing straight from near costa towards inner margin but terminating in cell Cu2, ocellus in cell M1 small somewhat insignificant with pupil apparently invisible (Fig. 2e); see also VFW description for Cisandinan. gen. above.

Ventral hindwing:

Notable features include, iridescent scales absent, postdiscal band slightly sinuate, submarginal ocelli small, ocelli in cells M1 and Cu1 not reaching veins defining these cells, ocelli in cells Rs and Cu2 smaller than previously mentioned two ocelli (Fig. 2e); see also VHW description for Cisandinan. gen. above.

Genitalia:

Notable features include, dorsal margin of tegumen rather smoothly curved as well as anterior portion of ventral margin of tegumen appearing only slightly convex, rather straight saccus in lateral view, apical process of valva somewhat broad and straight, occupying about one-fourth of valva in length (Fig. 3e–g); see also male genitalia description for Cisandinan. gen. above.

Female (Fig. 2f):

Forewing length 24 mm (n = 1).

Similar to male except as follows:

Notable features include, iridescent greenish lilac scales absent on dorsal surface which appears uniformly brown, posterior end of VFW postdiscal band curving outwards (Fig. 2f); see also female wing shape and pattern description for Cisandinan. gen. above.

Female genitalia and abdomen:

Not examined.

Type material:

Holotype male, with the following labels separated by double-forward slashes: //BRASIL, PARANÁ, MOREIRA SALLES, RPPN MOREIRA SALLES, 24° 04′ 25″ S, 53° 02′ 4610″ W, 7-X-2012 LABLEP LEG// BC-DZ Willmott 137// (DZUP).

Paratypes (13 ♂, 6 ♀):

Argentina: Misiones: Departamento de Iguazú, Parque Nacional Iguazú, Seccional Yacui, [25°40′47″S,54°10′12″W], 240 m, (Núñez Bustos, E., Kopuchian, C., Tubaro, P., Fortino, A.), 11 Apr 2011, 1 ♂, (MACN) (Lavinia et al. (2017)), 13 Apr 2011, 1 ♀, (MACN) (Lavinia et al. (2017)). Brazil: Bahia: [12°59′S,38°31′W], 4 ♂, (MNHU); Espírito Santo: [20°20′S,40°17′W], 1 ♂, (MNHU), 1 ♂, (MNHU); Leopoldina (= Santa Leopoldina?), [20°6′S,40°32′W], (Michaelis), 1894, 1 ♂, (MNHU); Linhares, [19°23′S,40°4′W], (Elias, P. C.), Feb 1972, 1 ♂ [FLMNH-MGCL-265692], (FLMNH); Rio de Janeiro: hwy km 27 Rio - Teresópolis, [6°51′57″S,78°4′35″W], (Callaghan, C. J.), 9 Oct 1971, 1 ♂ [FLMNH-MGCL-265693], [dissection, SN-20-14] (FLMNH). Paraná: São Pedro do Ivaí, RPPN Barbacena, 300m, (Mielke, Dolibaina, Carneiro, Maia), 8 Oct 2010, 1 ♀ [DZ 29.315 – BC DZ Willmott 192] (DZUP); Londrina, Fazenda Santa Helena, 650m, (Moure, Mielke, Wedderhoff), 7 Dez 1975, 2 ♀ [DZ 29.317, DZ 52. 557] (DZUP); Jussara: Horto CMNP, (Moure, Mielke, Wedderhoff), 12 Dez 1975, 1 ♂ [DZ 52.559] (DZUP); Foz do Iguaçu, (Mielke & Casagrande), 6 Sep 1985, 1 ♀ [DZ 52.556]; Moreira Salles, RPPN Moreira Salles, (Lablep), 7 Oct 2012, 1♂ 1 ♀ [DZ 49.895, DZ 49.915 (DZUP). Country unknown: Not located: ‘Surinam’-error, 1 ♂, (MNHU).

Other records (1♂, 7 ♀):

Brazil:Bahia: Bahía, [12°59′S,38°31′W], 2 ♀ (MNHU); Espírito Santo: Espírito Santo, [20°20′S,40°17′W], 1 ♀ [BMNH(E)-1497645], 1 ♀ [BMNH(E)-1497646], (NHMUK); Leopoldina (= Santa Leopoldina?), [20°6′S,40°32′W], 1 ♀, (MNHU), (Michaelis); Linhares, [19°23′S,40°4′W], (Elias, P. C.), Mar 1972, 1 ♀ [FLMNH-MGCL-265694] [dissection, SN-20-15], 1 ♀ [FLMNH-MGCL-265695], (FLMNH), May 1972, 1 ♀ [FLMNH-MGCL-265696], (FLMNH); Paraguay:Canindeyú: Armisticio, [−24.57487, −54.53756], 28 Aug 2008, 1♂ (sight record, photographed by Ulf Drechsel; Fig. 8).

Etymology:

This specific epithet is based on the Spanish and Portuguese noun ‘esmeralda’, meaning ‘emerald’, in reference to the male of this species possessing characteristic brilliant green scales on the DFW and part of the DHW. The name is a feminine noun in apposition.

Distribution and natural history:

This species is known from eastern and southeastern Brazil and northeastern Argentina, as well as by a single sight record from Paraguay (Fig. 8).

Remarks:

The barcoded female from Argentina (LEPIG190-11) is the only reliable female of C. esmeraldan. sp. to date. During the course of compiling specimens data for the present study, several females from southeastern Brazil (listed under ‘Other records’) were examined by the authors as potentially being conspecific with the male C. esmeraldan. sp. from the same area. Nevertheless, their phenotype is not consistent with the barcoded female mainly by virtue of them possessing a purple-ish sheen on the wing surface and the ventral ground color being greyish. We are not aware of any male specimen(s) corresponding to this female phenotype, and thus we decided to exclude these female individuals from the type series until further data becomes available to assess their identificatity.

Cisandina philippa (Butler, 1867), New Combination, Reinstated Status
(Figs. 1, 2g and h, 3h–j, 4c and d, 6a and b, 7a and b, 8)

• Euptychia philippaButler, 1867: 485. Lectotype, designated herein; Butler 1868: 30, Kirby 1871: 52, Butler 1877: 121; Weymer 1911: 216, Riley and Gabriel 1924: 46, D’Abrera 1988: 768–769, figs.

• Euptychia lea f. philippa: Weymer 1911: 216.

• Euptychia lea var. philippa: Gaede 1931: 452.

• Euptychia batesii f. tersaWeymer 1911: 214, pl. 49, fig. a, Lamas 2004: 220. Lectotype, designated herein.

• Magneuptychia lea philippa: Lamas 2004: 220.

Systematic placement and diagnosis:

Our maximum likelihood approach found Cisandina philippan. comb. & reinst. stat. as a sister taxon to C. lean. comb. + C. esmeraldan. sp., with a weak to moderate support (Fig. 1; SH-aLRT/UFBoot = 80.8/71). The infra-specific genetic divergence among seven sequenced C. philippan. comb. & reinst. stat. specimens varies from 0.04% to 3.55%, although this remarkably high maximum value (3.55%) is due to DNA99-022 (from Madre de Dios, Peru), which lacks data for the first approximately 250 nucleotides. The six Ecuadorian individuals, representing samples from Zamora-Chinchipe and Morona-Santiago provinces, exhibit a range of within-species COI divergence from 0.01 to 1.71% with a mean of 0.626%. Based on our COI data, genetic distance among these sampled C. philippan. comb. & reinst. stat. individuals and four other closely related Cisandinan. gen. species sequenced for this study is a minimum of 3.74%, suggesting an existence of a ‘barcoding gap’. See Table 3 for further information regarding genetic divergence of Cisandinan. gen. taxa. The male of C. philippan. comb. & reinst. stat. is readily distinguished from males of other species in the genus by its uniformly brown dorsal surface, whereas male specimens of other species exhibit either bluish or greenish iridescent scales on the dorsal surface. The female specimens of this species are also dorsally brown, which is also the case with C. esmeraldan. sp. but not two other species in the genus; see corresponding section of C. esmeraldan. sp. for further diagnostic characters to identify females.

Taxonomy:

Euptychia philippa was described by Arthur Gardiner Butler in his monograph of Euptychia (sensu lato), where he introduced 60 new euptychiine butterflies to science. Like many other species described by Butler, information regarding the number of examined specimens and sex was not provided in the original description of E. philippa. His description of E. philippa was also not accompanied by any illustration of this taxon, but the identity of this species can be guessed from the Latin text and interpretation of a few other aspects of Butler’s work. First, E. philippa was described in his ‘Division II’ of Euptychia, which is one of his seven subdivisions of Euptychia he erected in his monograph of the group. Butler (1867: 481) characterized this ‘Division II’ as ‘wings variable above and below, of violet, blue and green’. Those species classified in this division, along with E. philippa, include E. picea Butler, 1867, E. lysidice (Cramer, 1777), E. glaucina Bates, 1865, E. aegrota Butler, 1867, E. pilata Butler, 1867, E. brixiola Butler, 1867, E. brixius (= Satyrus brixus Godart [1824]), E. coelestis Butler, 1867, E. urania Butler, 1867, E. lea, and E. junia. Despite some of these names not being considered valid today, Butler apparently grouped together species that possess iridescent wing coloration, judging from the phenotypes of the species listed by him. It is also worth noting that the two species immediately preceding E. philippa in Butler’s monograph are E. lea and E. junia, two names considered as applying to the same species by Lamas (2004), a proposal followed by the present study, and which species proves to be a member of Cisandinan. gen. in our molecular phylogeny (Fig. 1). Furthermore, the description of E. philippa begins by stating ‘alae supra fuscae’ which translates to ‘wings above dark brown’, and ends by noting its ventral similarity with E. junia, both statements that narrow down the candidates examined by Butler to female C. esmeraldan. sp. or what we regard here as E. philippa. If this assumption is correct, the possibility of Butler examining female C. esmeraldan. sp. can be excluded on the basis of the type locality of E. philippa being Ega [=Tefé] according to the original description, whereas C. esmeraldan. sp. is a taxon known from the Atlantic coastal forest of Brazil and Argentina. The syntype housed at the NHMUK is a male specimen with a uniformly brown dorsal surface as described by Butler and in accordance with characters and inferences discussed above. Like Butler, who considered the brown dorsal surface to be a character to justify E. philippa as a species-level taxon, some subsequent authors also followed this trend (e.g., D’Abrera 1988). On the other hand, other authors proposed an opposing taxonomic hypothesis, such as Weymer (1911), who regarded E. philippa to be a ‘form’ of P. lea from Ega (=Tefé). Following Weymer’s (1911) proposal, in which the name was considered to be subspecific according to Article 45.6.4.1 of the ICZN (1999), Lamas (2004) also regarded this taxon as subspecific. Nevertheless, both genetic divergence based on COI and multi-locus maximum likelihood (see ‘Systematic Placement and Diagnosis’, Fig. 1 and Table 2 for further information) are in favor of species-level status for this taxon with its uniformly brown dorsal surface. To reflect this taxonomic change and to settle its nomenclature as a senior subjective synonym of Euptychia batesii f. tersa, we here designate the aforementioned male syntype in the NHMUK as the lectotype of E. philippa, with the following labels separated by double-forward slashes, and reinstate its taxonomic status from subspecific to specific (lectotype designation, reinstated status): //B.M. TYPE No. Rh 3178 Euptychia philippa, ♂ Butl.//♂//Type of Species//Ega, U.Amazonas. H.W. Bates.// ♂ Ega Philippa Butl. Type// Type H. T.// Godman-Salvin Coll. 1904.-1. Euptychia Philippa, Butl.//.

Euptychia batesii f. tersa was described by Gustav Weymer in Seitz’s Macrolepidoptera of the World. The original description did not specify the number of specimen(s) he examined nor the sex, in addition to not providing any information on its locality. Despite being unrelated to Euptychia batesii Butler, 1867 (currently regarded as a subspecies of Magneutpychia harpyia (C. Felder & R. Felder, 1867), according to Lamas (2004)), Weymer described this taxon as a form of Euptychia batesii and also compared it with a close relative of that species, Euptychia analis Godman, 1905. The original description noted that the VHW ocelli in cells M2 and M3 were formed of a circular ring with a central pupil, rather than being silver spots as in Euptychia batesii (= Neonympha harpyia) and Euptychia analis. Also, Weymer pointed out the presence/absence of the VHW ocellus in cell Cu2 as his justification for erecting this new ‘form’. Nevertheless, these phenotypes discussed by Weymer are often considered as informative characters at euptychiine species-level classification, and it is unclear why he considered tersa and batesii as conspecific. The illustration of the ventral surface associated with the original description (pl. 49, fig. a) does indeed show the phenotypic features described by Weymer, as well as the presence of the VFW ocellus in cell Cu1, which is apparently an unusual character for species discussed in this article. Lamas (2004) considered Euptychia batesii f. tersa as a junior subjective synonym of E. philippa. The female syntype, which is most likely the specimen on which Weymer based his illustration, given the presence of a VFW ocellus in cell Cu1, is housed at SMT and figured in Warren et al. (2017). We here designate this female specimen, with the following labels separated by double-forward slashes, as the lectotype of E. batesii f. tersa in order to settle its nomenclature, and follow Lamas (2004) in regarding this taxon as a junior subjective synonym of E. philippa (lectotype designation): //GART specimen ID: 02498 Exemplar + Etiketten dokumentiert specimen + label data documented 2003// LECTOTYPUS// LECTOTYPE ♀ Euptychia batesii f. tersa Weymer by G. Lamas ‘91// Stauding.& Bang-Haas Dresden, Ankauf 1961// Staatl. Museum für Tierkunde Dresden// tersa Weym.// Spec.// Original?//.

Distribution and natural history:

This species is known from the western Amazon, from the Andean foothills from Ecuador to southern Peru, east to the central Amazon, with a possible small area of sympatry with C. lean. comb. in Tefé and the vicinity of Manaus (Fig. 8). It is sympatric with C. castanyasp. n. from central to southern Peru and in western Brazil (Rondônia). In Ecuador, this species occurs in lowland rainforest up to 1,400 m, where it is uncommon. Males and females were encountered at similar frequency, in a variety of sites, mostly in undisturbed terra firme forest, but also in floodplain secondary forest with abundant Guadua (Poaceae) bamboo. Individuals were encountered flying at varying heights above the ground, from 1 to 4 m, in both shady understory and in light gaps and at forest edges. A penultimate instar caterpillar was found on species of herbaceous bamboo, Taquara micrantha (Kunth) I.L.C.O liveira & R.P.O liveira (Poaceae) at Finca Las Piedras, Madre de Dios, Peru, on 20 April 2021 (voucher: 2020-FLP-IMM-0336), and the immature stages are described below.

Specimens examined (39 ♂, 31 ♀):

Brazil: Acre: Marechal Thaumaturgo, Foz do Rio Tejo, Reserva Extrativista Alto Juruá, estrada para o Rio Arara, (Brown, K. S., Freitas, A. V. L.), 16 Sep 1997, 1 ♂, (ZUEC); [Marechal Thaumaturgo, Boca do Rio Tejo, Reserva Extrativista Alto Juruá], 20–27 Aug 1997, 1 ♂ [‘BTEJO-REAJ-AC’], (ZUEC); Senador Guiomard, Reserva Catuaba, (Mielke & Casagrande), 31 Jan – 5 Feb – 2009, 1♂ 1 ♀ [DZ 52.562, DZ 52. 564] (DZUP); Amazonas: Borba, Rio Abacaxis, Comunidade Paxiúba, [4°28′48″S,58°34″W], (Mielke, O. H. H., Casagrande, M. M.), 2–4 Jun 2008, 1 ♀, [DZ 52.563 – BC-DZ Willmott – 140] (DZUP); Ega (= Tefé), [3°22′S,64°42′W], (Bates, H. W.), 1 ♂ [‘♂ Type of Species.’//’Godman-Salvin Coll. 1904-1. Euptychia philippa, Butl.’//’Ega, U. Amazons. H.W. Bates.’//’♂ Ega Philippa Butl. Type’//’Type H.T.’//’B.M. TYPE No. Rh3178. Euptychia philippa. ♂ Butl.’], 1 ♂ [BMNH(E)-1266946; HT of philippa], 1 ♂ [BMNH(E)-1670283], 1 ♂ [BMNH(E)-1670295], 1 ♀ [BMNH(E)-1670287], 1 ♀ [BMNH(E)-1670288], (NHMUK); Igarapé Massauari, [2°54′17″S,57°8′23″W], (Hahnel, P.), 1 ♂, (MNHU); Manicoré, [5°49′S,61°17′W], (Hahnel, P.), 1887, 1 ♂, (MNHU); São Paulo de Olivença, [3°28′S,68°57′W], (Mathan, M. de), 1 ♂ [BMNH(E)-1670280], 1 ♂ [BMNH(E)-1670281], 1 ♂ [BMNH(E)-1670282], 1 ♀ [BMNH(E)-1670211], (NHMUK), Jun–Jul 1883, 1 ♂ [BMNH(E)-1670275], 1 ♂ [BMNH(E)-1670276], 1 ♂ [BMNH(E)-1670277], 1 ♂ [BMNH(E)-1670278], 1 ♂ [BMNH(E)-1670279], 1 ♀ [BMNH(E)-1670285], 1 ♀ [BMNH(E)-1670286], (NHMUK), (Moss, A. M.), 1 ♀ [BMNH(E)-525170], (NHMUK); Tefé, [3°22′S,64°44′W], (Hahnel, P.), 1 ♂ [dissection, 9076; ‘NEOTYPE ♂ Euptychia batesii f. tersa Weymer designated by Lee D. Miller 1989//Teffe (= Tefé) Hhl.//Genitalia vial M-9076 ♂ Lee D. Miller’], (MNHU) [unpublished neotype designation]; Rondônia: 1 km N Cacaulândia, [10°31′30″S,62°48′W], 168 m, (Brock, J. P.), 28 Oct 1990, 1 ♂ [FLMNH-MGCL-265727], [dissection, SN-20-34] (FLMNH); 5 km S of Cacaulândia on Linha C-10 at Rio Pardo off B-65, [10°23′15″S,62°54′53″W], (Gomes, O.), 13 Mar 1984, 1 ♂ [FLMNH-MGCL-296552], (FLMNH), 29 Aug 1993, 1 ♀ [FLMNH-MGCL-265729; Station #15 forest], [dissection, SN-20-11] (FLMNH); Cacaulândia, 7 km E B-65, Fazenda Rancho Grande, [10°17′58″S,62°52′14″W], (Austin, G. T.), 19 Nov 1992, 1 ♂ [FLMNH-MGCL-265728], (FLMNH), (Bongiolo, G.), 14 Jun 1992, 1 ♂ [FLMNH-MGCL-265726; Station #3 forest], (FLMNH). Ecuador: Morona-Santiago: jct. Río Mayalico-Río Santiago, Isla de las Conchas, [3°2′10″S,77°58′29″W], 250 m, (Hall, J. P. W., Willmott, K. R., J. C. R., J. I. R), 8,10 Aug 2015, 1 ♂ [FLMNH-MGCL-217579], (FLMNH); hwy km 20 Mendez-Santiago rd., [2°47′6″S,78°15′24″W], 850 m, (Perceval, M. J.), 14 Oct 1997, 1 ♀, (MIPE); km 30 Méndez-Limón rd., Río Yungantza, [2°52′13″S,78°21′56″W], 650 m, (Hall, J. P. W.), 1–3 Mar 2017, 1 ♀ [FLMNH-MGCL-281450], (FLMNH); Santiago de Mendez, [3°2′11″S,78°2′W], (Nakahara, S.), 16 Jun 2014, 1 ♀, (FLMNH); Napo: Río Napo, hwy Puerto Napo-Ahuano rd., Chichicorrumi, [1°4′11″S,77°37′45″W], 450 m, (Willmott, K. R., Hall, J. P. W.), 2,9 Jul 1993, 1 ♀, (FLMNH); Orellana: Laguna Zancudococha, military trail, [0°35′16″S,75°28′16″W], 220 m, (Aldaz, R.), 9–13 Jul 2017, 1 ♂ [FLMNH-MGCL-288722], (FLMNH); Río Aguarico, Zancudococha, [0°34′23″S,75°26′13″W], 240 m, (Willmott, K.R., J.C.R, J.I.R., Aldaz, R.), 14 Jul 2017, 1 ♀, (INABIO); Río Napo, Boca del Río Añangu, [0°31′43″S,76°23′41″W], 220–300 m, (Willmott, K. R.), 27 Oct 2005, 1 ♀ [FLMNH-MGCL-111516], (FLMNH); Shiripuno Lodge, Mirador trail, [1°4′50″S,76°44′42″W], 350 m, (Hall, J. P. W., Willmott, K. R., J. C. R., J. I. R.), 8,9,11 Aug 2018, 1 ♂, 1 ♀, (FLMNH); Sucumbíos: Cerro Lumbaquí Norte, [0°1′42″N,77°19′W], 800–950 m, (Willmott, K. R., Hall, J. P. W.), 21–23 Jul 1999, 1 ♂, (FLMNH); Zamora-Chinchipe: 3 km E El Panguí, Centro Shuar Cháarip, [3°38′6″S,78°33′29″W], 800 m, (Willmott, K. R., Hall, J. P. W.), 4 Aug 2009, 1 ♀ [FLMNH-MGCL-145674], (FLMNH); km 11.5 Los Encuentros-Zarza, La Libertad, [3°47′54″S,78°36′26″W], 1,250 m, (Willmott, K. R., Hall, J. P. W.), 6,8 Aug 2009, 1 ♂ [FLMNH-MGCL-145675], 1 ♂ [FLMNH-MGCL-145676], (FLMNH); Zamora, ridge to west, [4°4′30″S,78°58′7″W], 1,400–1,450 m, (Willmott, K. R.), 20 May 2000, 1 ♂ [dissection, KW-14-008], (FLMNH). Peru: Cuzco: Pilcopata, Villa Carmen, [12°54′S,71°24′W], 540 m, (Brock, J.), 31 Jan 2013, 1 ♀ [MUSM-LEP-103083], (MUSM); Quincemil, Quebrada Yanaorcco, [13°16′S,70°47′W], 900 m, (Rodríguez, M.), Feb 2010, 1 ♀ [MUSM-LEP-103084], (MUSM); Junín: La Merced, [11°3′S,75°19′W], 790–762 m, (Watkins & Tomlinson), May–Jun 1903, 1 ♀ [BMNH(E)-1670293], (NHMUK); Loreto: Castaña, [0°48′S,75°14′W], 150 m, (Lamas, G.), 26 Oct 1993, 1 ♂ [MUSM-LEP-103079], (MUSM), 29 Oct 1993, 1 ♂ [MUSM-LEP-103080], (MUSM); Lower Río Ucayali, Río Pacaya, Aug-Sep 1912, 1 ♂ [BMNH(E)-1670291], (NHMUK); Pebas, [3°19′S,71°51′W], 120 m, (Hahnel, P.), 1 ♀ [dissection, 9077], (MNHU), (Mathan, M. de), Dec 1879–Mar 1880, 1 ♂ [BMNH(E)-1670290], (NHMUK), Nov 1906, 1 ♂ [BMNH(E)-1497733], (NHMUK); Zona Reservada Allpahuayo-Mishana, [3°57′30″S,73°25′30″W], 170 m, (Ramírez, J. J.), 8 Aug 2001, 1 ♀ [MUSM-LEP-103092], (MUSM); Madre de Dios: 30 km SW Puerto Maldonado, [12°36′S,69°11′W], 200 m, (Anderson, J. J.), 18–23 Oct 1982, 1 ♀ [MUSM-LEP-103087], (MUSM); Boca Río La Torre, [12°50′S,69°17′W], 300 m, (Lamas, G.), 12 Feb 1982, 1 ♂ [MUSM-LEP-103082], (MUSM), 26 Sep 1981, 1 ♀ [MUSM-LEP-103085], (MUSM), 27 Sep 1981, 1 ♀ [MUSM-LEP-103086], (MUSM); Parque Nacional del Manu, Pakitza, [11°55′48″S,71°15′18″W], 400 m, (Lamas, G.), 13 Oct 1990, 1 ♀ [MUSM-LEP-103089], (MUSM), 18 Oct 1990, 1 ♂ [MUSM-LEP-103081], 1 ♀ [MUSM-LEP-103090], (MUSM), (Rowe, W.), 3 Nov 1990, 1 ♀ [MUSM-LEP-103088], (MUSM); Reserva Tambopata, La Colpa, [13°9′S,69°37′W], 250 m, (Aibar, P.), 19 Oct 2000, 1 ♀ [MUSM-LEP-103091], (MUSM); Puno: Río Tambopata, [12°36′S,69°11′W], 270 m, 15 Jul 1979, 1 ♂ [FLMNH-MGCL-265725], [dissection, SN-20-10] (FLMNH); San Martín: near Yurimaguas, ‘Chambireyacu’ [=Río Chambirayacu], [5°54′S,76°14′W], 100 m, (Mathan, M. de), Jun-Aug 1885, 1 ♂ [BMNH(E)-1670292], (NHMUK); Moyobamba, [6°2′S,76°58′W], 855 m, Jan-Jun 1887, 1 ♀ [BMNH(E)-1670294], (NHMUK). Country unknown: Not located: ‘Amaz[on] S’, 1 ♀ [BMNH(E)-1497644], (NHMUK); no data, 1 ♂, (MNHU), 1 ♂, (MNHU).

Other records:

Ecuador: Napo: Tena-Puyo rd., El Capricho, [1°11′14″S,77°49′53″W], 850 m, (Neild, A.), 20 Oct 2015, 1 ♂, (photograph live specimen) (Neild, A. F. E. (20 Oct 2015, sight record, by email with photo to KRW)); Peru: Madre de Dios: Madre de Dios, [12°16′S,70°55′W] [ID based on DNA barcode], (DEMU) (Murray and Prowell (2005)).

Immature Stages (Figs. 6a and b; 7a and b):

Penultimate instar:

Head capsule (Fig. 7a) dark olive, bearing pair of stubby but rather pointed somewhat ‘antler-like’ scoli on vertex, apex of scoli appearing darker; six stemmata present in lateral view, third stemma largest, first and sixth stemmata transparent thus somewhat insignificant, fifth semi-transparent; head capsule width c. 1.9 mm; scoli length c. 0.4 mm; body (Fig. 6a) appearing dark chartreuse, two jagged and somewhat undefined dorsal lines, with three creamy-whitish dorsolateral to lateral lines, pair of short caudal filaments; body length c. 18 mm. Duration 13 d (n = 1).

Ultimate instar:

Head capsule (Fig. 7b) dark brown, bearing pair of rounded, stubby and blunt scoli; six stemmata present in lateral view, third stemma largest, first and sixth stemmata transparent thus somewhat insignificant, fifth semi-transparent; labrum appearing reduced compared with previous instar; secondary setae increasing in number and length, head capsule overall rounder compared with immediately preceding instar; light area present in frontal view appearing as ‘M-shaped’ (not illustrated); head capsule width c. 2.5 mm; scoli length c. 0.2 mm; body (Fig. 6b) appearing flesh-coloured, two jagged and somewhat undefined dorsal lines, with three creamy-whitish dorsolateral to lateral lines, pair of short caudal filaments; body length c. 28 mm. Duration 13 d (n = 1).

Pupa:

Not described or illustrated. Duration 16 d (n = 1).

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 castanyan. sp. as sister to (C. lean. comb. + C. esmeraldan. sp.) + C. philippan. 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. castanyan. 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 Cisandinan. gen. species which form a clade to which it is supported as sister, reinforcing our taxonomic hypothesis. Males of C. castanyan. 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. lean. comb. and absent in C. philippan. comb. & reinst. stat. As explained under the immediately preceding taxon, C. castanyan. sp. is sympatric with C. philippan. 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. castanyan. sp. (scoli/headcapsule width ratio = 0.14; scoli length c. 0.4 mm.) compared with C. philippan. comb. & reinst. stat. (scoli/headcapsule width ratio=0.08; scoli length c. 0.2 mm.).

Male (Fig. 2i):

Forewing length 22–23 mm (mean 22.3 mm; n = 3).

Head:

See head description for Cisandinan. gen. above.

Thorax:

See thorax description for Cisandinan. gen. above.

Abdomen:

See abdomen description for Cisandinan. gen. above, eighth sternite rather narrow and appearing as two separated patches at anterior side.

Wing venation:

See wing venation description for Cisandinan. gen. above.

Wing shape:

See wing shape description for Cisandinan. 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 M1, M2, M3, Cu1, although this area revealing ground color narrows posteriorly (Fig. 2i); see also DFW description for Cisandinan. 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 Cisandinan. 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 M1 small and may appear insignificant with pupil being invisible, additional ocellus may be present in adjacent cells (Fig. 2i); see also VFW description for Cisandinan. gen. above.

Ventral hindwing:

Notable features include, grey-ish blue scales covering areas in cells Cu1, Cu2 and 2A, submarginal ocelli appearing small, ocelli in cells M1 and Cu1 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 Cisandinan. 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 Cisandinan. gen. above.

Female (Fig. 2j):

Forewing length 21–22 mm (mean 21.7 mm, n = 3).

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 M3, more than half of Cu1, most of cells Cu2 and 2A (Fig. 2j); see also female wing shape and pattern description for Cisandinan. 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 Cisandinan. 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. castanyan. 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.

Immature Stages (Figs. 6e–i, 7c–e):

Egg:

Not Recorded.

First instar:

Not Recorded.

Second instar:

Head capsule (Fig. 7c) light brown, bearing pair of stubby but rather pointed, somewhat ‘antler-like’ scoli on vertex, apex of scoli appearing darker; six stemmata present in lateral view, third stemma largest, first and sixth stemmata transparent thus somewhat insignificant, fifth semi-transparent; head capsule width c. 1.4 mm; scoli length c. 0.4 mm; body (Fig. 6e) pale green, with creamy-whitish dorso-lateral lines and similar but narrower lateral lines, pair of short caudal filaments; body length c. 11 mm. Molted to third instar 6 d after collecting in the field (on June 26th 2020; n = 1).

Third instar:

Head capsule (Fig. 7d) light brown, bearing pair of stubby but rather pointed, somewhat ‘antler-like’ scoli on vertex, apex of scoli appearing lighter; six stemmata present in lateral view, third stemma largest, first and sixth stemmata transparent thus somewhat insignificant, fifth semi-transparent; head capsule width c. 2.1 mm; scoli length c. 0.5 mm; body (Fig. 6f) appearing chartreuse, two jagged and somewhat undefined dorsal lines, with three creamy-whitish dorsolateral to lateral lines, pair of short caudal filaments; body length c. 14 mm. Duration 13 d (n = 1).

Fourth (ultimate) instar:

Head capsule (Fig. 7e) dark brown, bearing pair of rounded, stubby and blunt scoli on vertex, scoli appearing lighter posteriorly; six stemmata present in lateral view, third stemma largest, first and sixth stemmata transparent thus somewhat insignificant, fifth semi-transparent; labrum appearing reduced compared with previous instars; secondary setae increasing in number and posterior portion of head capsule broadens compared with immediately preceding instar; light area present in frontal view appearing as ‘M-shaped’ (not illustrated); head capsule width c. 2.8 mm; scoli length c. 0.4 mm; body flesh-colored, two jagged and somewhat indistinctive dorsal lines, with three creamy-whitish dorsolateral to lateral lines, pair of short caudal filaments; body length c. 24 mm. Duration 20 d (n = 1).

Pupa:

Short and relatively smooth, with reduced ocular caps, overall appearing somewhat mottled, ventrally, including wing case, creamy white scattered with dark brown, dorsally appearing darker brownish except for rather pinkish two longitudinal rows of protuberances on abdomen and small areas of creamy-white elsewhere, cremaster short (Fig. 6h and i). Duration 13–15 d (n = 2).

Adult:

Two males, eclosed on 13 August 2020 (2020-FLP-IMM-0188) and 5 April 2021 (2021-FLP-IMM-0234).

Cisandina fida (Weymer, 1911), New Combination
Cisandina fida fida (Weymer, 1911), New Combination
(Figs. 2k and l, 8)

• Euptychia fidaWeymer, 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 fidan. comb. is paraphyletic, although the sister relationship between Peruvian C. fidan. comb. (LEP-58115) and C. sanmarcosn. 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 fidan. comb. has a jagged VHW postdiscal band (but see also diagnostic characters for the following taxon), whereas this band is rather straight in C. sanmarcosn. comb.; C. fidan. comb. possesses whitish coloration distal to the VHW postdiscal band, whereas this whitish coloration is absent in C. sanmarcosn. 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. fidan. comb. revealed some individuals (e.g., FLMNH# 145742) in which the VFW postdiscal band terminates at 2A, as in C. sanmarcosn. 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. fidan. comb. is known to date only from Bolivia (Fig. 8)

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 Cu2 and bends back half way between Cu2 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 M2 and M3 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.

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, inNakahara 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. fidan. comb. (Fig. 1). Cisandina sanmarcosn. comb. forms a strongly supported clade with C. fidan. 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. sanmarcosn. comb. results in C. fidan. 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. fidan. 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 sanmarcosn. 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 fidan. 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 sanmarcosn. 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. sanmarcosn. 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.

Cisandina trinitensis (Brévignon & Benmesbah, 2012), New Combination
(Figs. 1, 2q and r, 3u–w, 4l and m, 8)

• Euptychia? sp.: D’Abrera 1988: 780, figs.

• Magneuptychia trinitensisBrévignon & Benmesbah, 2012: 47, pl. 3, Figs. 1–4, pl. 4, Figs. 8, 8a.

Systematic placement and diagnosis:

Cisandina trinitensisn. comb. is moderately to strongly supported as sister to the remainder of Cisandinan. gen. (Fig. 1; SH-aLRT/UFBoot=91.9/92), with three sequenced individuals (LCB251, LCB356, MB_1708_15) forming a well-supported clade (Fig. 1; SH-aLRT/UFBoot = 100/100). Cisandina trinitensisn. comb. is readily distinguished from other species in the genus by the lack of an ocellus in VHW cell Cu2, coupled with the presence of four submarginal ocelli on the VFW (in cells M1, M2, M3, and Cu1). However, it must be noted that some ocelli on the VFW may appear as faint or only a trace, and with the limited number of specimens examined this character may prove unreliable.

Taxonomy:

Brévignon and Benmesbah (2012) described Magneuptychia trinitensis based on two males and two females, including the male holotype, all collected in French Guiana. The holotype was examined prior to deposition in MOBE, and it was also figured in the original description (pl. 3, Figs. 1 and 2), where the aforementioned characters are visible. We also incorporated sequences from the holotype in our molecular phylogeny (Fig. 1; LCB 251), thus leaving no doubt as to its rather distinctive identity, which is also indicated as being sister to the rest of the genus according to the maximum likelihood tree (Fig. 1).

Distribution and natural history:

Cisandina trinitensisn. comb. is so far known from a handful of sites all situated in French Guiana (Fig. 8).

Specimens examined (3 ♂, 2 ♀):

French Guiana: Cayenne: Montagne de la Trinité, [4°36′2″N,53°24′43″W], (Benmesbah, M.), 10 Nov 2010, 1 ♀ [AT trinitensis], (LBCB), (Hermier, B.), 30 Oct 2008, 1 ♀ [PT trinitensis], (LBCB); Nouragues Station, [4°5′N,52°41′W], (Benmesbah, M.), 6 Sep 2010, 1 ♂, (MOBE); St-Laurent du Maroni: Maripasoula, Antecume-Pata, ‘filet’, [3°17′53″N,54°4′16″W], (Benmesbah, M.), 13 Mar 2012, 1 ♂ [HT trinitensis], (MOBE); Not located: ‘French Guiana’, 1 ♂ [PT trinitensis], (MOBE).

Other records:

French Guiana: Cayenne: Cayenne, [4°56′N,52°20′W], 1 ♂, 1 ♀, (NHMUK) (D’Abrera (188: 780, ‘Euptychia? sp.’)); St-Laurent du Maroni: Maripasoula, Antecume-Pata, ‘filet’, [3°17′53″N,54°4′16″W], (Benmesbah, M.), 13 Mar 2012, 1 ♀ [PT trinitensis], (MOBE) (Brévignon and Benmesbah (2012)).

Discussion

We here introduce another new euptychiine genus, Cisandinan. gen., to accommodate five existing species previously associated with Magneuptychia and Euptychoides, as well as describe three new taxa within the new genus. The highly polyphyletic natures of Magneuptychia and Euptychoides have been indicated by a number of other studies (e.g., Peña et al. 2010, Nakahara et al. 2016, Espeland et al. 2019), and our multi-locus phylogeny justifies these taxonomic changes as discussed above. Here, we considered several possible generic arrangements for these seven species, including recognizing three genera for the three phenotypically compact groups of Cisandina species, namely the C. lea subgroup, C. fida subgroup and C. trinitensis subgroup. In fact, our initial approach was to describe two genera, one for the C. lea subgroup and the other for the C. fida subgroup. Nevertheless, as DNA sequence data supported M. trinitensis as the sister to both of these genera, we decided instead to describe a single genus to harbor this morphologically relatively compact group of seven species. Indeed, the male genitalia and female genitalia can be considered as rather homogeneous among these seven species, with some characteristic features such as 1) reduced appendices angulares, 2) broad base of brachia, 3) large antero-dorsal opening of phallus, 4) sclerotized portion of manica, and 5) heavily sclerotized wrinkled lamella antevaginalis partially fused with lateral plate of eighth abdominal segment. These traits (1) to (5) are apparently not shared with other species in the ‘Archeuptychia clade’, except for the sclerotized region of manica, which is apparently also present in Erichthodes arius and Yphthimoides eriphule, and these characters (1) to (4) can be considered as rare character states even among the entire subtribe Euptychiina. Although the support is not high for its monophyly (Fig. 1; SH-aLRT/UFBoot=86.7/69), the potential sister group (Erichthodes arius + Yphthimoides eriphule) of Cisandinan. gen. also has rather distinctive genitalic characters. Apart from the manica being sclerotized at the juncture of the phallus and aedeagus, the male genitalia of these two species possess a broad, band-like juxta (in lateral and ventral view), a narrow uncus (of width similar to the brachia) which curves downwards, and the basal portion of the valva is tapering, and a study is underway by SN and EPB to describe a new genus for this clade. The distinctive genitalia of its sister group reinforces the taxonomic significance of the homogenous genitalic features of Cisandinan. gen., although we have not been able to examine the female genitalia of E. arius. Coupled with these genitalic characters, the slightly displaced VHW ocellus in cell M2 serves as a diagnostic character to separate these seven species from E. arius and Y. eriphule, and virtually all other members of the ‘Archeuptychia clade’. Despite genitalic characters suggesting that a single genus is the best taxonomic solution, there are some significant biological and distributional differences between species in the lea and fida subgroups, which are worth emphasizing. As summarized above, all four species in the lea subgroup inhabit lowland forests of the Amazon basin, Guianas, Trinidad, and Atlantic coastal forest of Brazil, being recorded from elevations up to 1,400 m. On the other hand, the two species in the fida subgroup are residents of tropical east Andean middle elevation cloud forests from southern Ecuador to Bolivia, confined to an altitudinal zone between 960 and 1,740 m. Thus, species in the lea and fida subgroups occur in allopatry with somewhat different habitat preferences, which may result in different biological traits, although the lack of natural history information for taxa recognized in the fida subgroup prevents further recognition of such differences. In fact, the immature stage morphology of two species in the lea subgroup, C. castanyan. sp. and C. philippan. comb. & reinst. stat. further supports our taxonomic proposal of establishing Cisandinan. gen. by their short, rounded and stubby head scoli in the last instar larvae, transforming from the well-developed bifurcating head scoli of the penultimate instar, which is a unique feature for euptychiines, as explained above. The overall phenotype of species in the lea subgroup is different from species in the fida subgroup in exhibiting iridescent blue or greenish wing coloration, whereas the wings of the latter subgroup are basically brownish without any lilac reflection. These phenotypic and ecological differences might merit generic recognition, but in that case a monotypic genus would be required for M. trinitensis, which seems unreasonable given the number of shared unique genitalic features with species in the lea and fida subgroups. We, therefore, decided to resolve this subjective issue by establishing a single genus for these seven species.

Nakahara et al. (2019b) made a judgement of accepting paraphyletic species, a situation presents in this article as well. We are fully aware of ‘the untenability of paraphyletic groups’ for taxonomic purposes (e.g., de Queiroz and Gauthier 1990), and the advantages of recognizing clades as taxa. Nevertheless, many ‘species’ (i.e., specific epithets) are established without testing their monophyly and it is not uncommon that they do not represent monophyletic entities due to the mode of speciation (e.g., Nosil et al. 2002), a situation also discussed in Nakahara et al. (2019a). Furthermore, multiple molecular markers should ideally be used to provide reliable evidence of monophyly or paraphyly of a species. Cisandina fidan. comb. and C. sanmarcosn. comb. are phenotypically readily distinguishable and their broad sympatry in central Peru and Bolivia refutes the possibility of subspecific status of the latter taxon. We were unable to obtain genetic data for the nominate race of C. fidan. comb., so this lack of data also might have contributed towards the unexpected result of recovering C. fidan. comb. as paraphyletic.

In summary, we contributed to resolving the systematics of two polyphyletic genera, Magneuptychia and Euptychoides, by removing five species in total and erecting a new genus to accommodate them. We further described two new species and one subspecies related to these taxa, and provided a phylogenetic hypothesis for all the species discussed in this study. Throughout this article and in other studies on taxa associated with Magneuptychia (Nakahara et al. 2020a), we have highlighted the importance of a biological classification supported by a phylogenetic hypothesis. We continue to revise the systematics of Euptychiina by following this trend, including a forthcoming study on a clade collapsed in Fig. 1.

Acknowledgments

We would like to express our gratitude to all curators and collectors who allowed access to or provided information about specimens under their care, including Gerald Legg (BMB), Wolfram Mey (MNHU), Mike J. Perceval (MIPE), Mohamed Benmesbah (MOBE), Tomasz Pyrcz (MZUJ), Blanca Huertas (NHMUK), Robert Robbins and Brian Harris (USNM), Axel Hausmann (ZSM), and André V. L. Freitas (ZUEC); Hajo Gernaat for translating relevant parts of the original description; Ulf Drechsel for allowing us to use his photograph of C. esmeraldan. sp.; Ezequiel Núñez for photographing relevant specimens; Lynn Clark and Fabrício Ferreira for generously identifying the host plants; all those who contributed to digitizing specimen data, especially Robyn Crowther and Evelyn Gamboa; three anonymous reviewers for conducting post-submission reviews and providing numerous helpful comments and suggestions. SN acknowledges University of Florida’s Entomology Department and Alliance for Sustainable Amazon for support. MRM thanks Diana Silva for their continuous support, and Consejo Nacional de Investigaciones Científicas y Técnicas – CONICET for doctoral fellowship. EPB (2016/15873-8, 2018/21432-0) thanks Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) (162673/2020-5). MMC (CNPq302084/2017-7) thanks Conselho Nacional de Desenvolvimento Científico e Tecnológico. KRW acknowledges the US National Science Foundation (DEB-1256742) for support. Rearing relevant to the present study at Finca Las Piedras was conducted under permit no. 187-2017-SERFOR/DGGSPFFS; Brazilian butterfly species are registered in the Brazilian Sistema Nacional de Gestão do Patrimônio Genético e do Conhecimento Tradicional Associado (National System for the Management of Genetic Heritage and Associated Traditional Knowledge) SISGEN (A37D0EF).

Author Contributions

SN wrote the paper and performed morphological work; SN, KK, ME, and KRW performed relevant molecular lab work and analysis; MRM, TCE, RT, JS and GG contributed towards obtaining life history information and incorporating into the paper; EPB and MMC assisted with specimens under their care; SN, MRM, KK, TCE, ME, MMC, EPB, GG, GL, and KRW edited and revised the paper.

Version of Record, first published online January 6, 2022 with fixed content and layout in compliance with Art. 8.1.3.2 ICZN