Á. L. Viloria

ANARTIA

Publicación del Museo de Biología de la Universidad del Zulia

ISSN 1315-642X (impresa) / ISSN 2665-0347 (digital)

Anartia, 34 (junio 2022): 28-42

http://zoobank.org/urn:lsid:zoobank.org:pub:0D754F2B-056C-4EE3-82B5-4F5AF752E4E4

Two new satyrine butteries from Venezuela

(Insecta: Lepidoptera: Nymphalidae)

Dos nuevas mariposas satíridas de Venezuela

(Insecta: Lepidoptera: Nymphalidae)

Ángel L. Viloria

Centro de Ecología, Instituto Venezolano de Inestigaciones Cientícas (IVIC), km 11 carretera Panamericana, Altos de Pipe,

estado Miranda 1204, Venezuela.

Correspondence: sebastianiloriacarrizo@gmail.com; aviloria@ivic.gob.ve

(Received: 02-11-2021 / Accepted: 15-05-2022 / On line: 30-09-2022)

ABSTRACT

Two new satyrine butteries known so far only from the Venezuelan territory are diagnosed, described, illustrated and

compared: Rudyphthimoides iseai gen. nov. et sp. nov. (Cordillera de la Costa, northern Venezuela) and Stephenympha

pauliana gen. nov. et sp. nov. (Serranía del Cuao, Amazonas, southern Venezuela). Seven new combinations, two new

synonymies are established, and the specic status of two taxa are revised: Rudyphthimoides anis (Butler, 1867) comb.

nov., R. erigone (Butler, 1867) sp. restit., stat. rev. et comb. nov., R. guenzeli (Anken, 1994) stat. rev. et comb. nov., R.

maepius (Godart, [1824]) comb. nov. (= Euptychia argyrospila Butler, 1867 syn. nov., = Euptychia perfuscata Butler, 1867

syn. nov.), R. mythra (Weymer, 1911) comb. nov., Stephenympha arius (Weymer, 1911) comb. nov. and S. eriphule (But-

ler, 1867) comb. nov.

Keywords: Amazon, Aragua, Coastal range, Cuao Mountains, nomenclature, new combinations, Satyrini, synonymy.

RESUMEN

Se diagnostican, describen, ilustran y comparan dos nuevas mariposas satíridas conocidas hasta ahora únicamente del terri-

torio venezolano: Rudyphthimoides iseai gen. nov. et sp. nov. (Cordillera de la Costa, norte de Venezuela) y Stephenympha

pauliana gen. nov. et sp. nov. (Serranía del Cuao, Amazonas, sur de Venezuela). Se establecen siete nuevas combinaciones,

dos nuevas sinonimias y se revisa el status de dos especies: Rudyphthimoides anis (Butler, 1867) comb. nov., R. erigone

(Butler, 1867) sp. restit., stat. rev. et comb. nov., R. guenzeli (Anken, 1994) stat. rev. et comb. nov., R. maepius (Godart,

[1824]) comb. nov. (= Euptychia argyrospila Butler, 1867 syn. nov., = Euptychia perfuscata Butler, 1867 syn. nov.), R.

mythra (Weymer, 1911) comb. nov., Stephenympha arius (Weymer, 1911) comb. nov. y S. eriphule (Butler, 1867) comb.

nov.

Palabras clave: Amazonas, Aragua, Cordillera de la Costa, nomenclatura, nuevas combinaciones, Satyrini, Serranía del

Cuao, sinonimia.

INTRODUCTION

e objective of this work is to describe two new spe-

cies of butteries of the family Nymphalidae, subfamily

Satyrinae, coming respectively from two distant localities

of the Venezuelan territory. e rst case corresponds to a

series of individuals of a previously unnoticed taxon, cap-

tured 70 years ago by Francisco Fernández-Yépez in the

surroundings of the town of El Limón, currently part of

the city of Maracay, in the foothills of the Cordillera de La

New Satyrinae from Venezuela

Costa (450–500 m a.s.l.), Aragua state. is series of eight

specimens, preserved in the collection of the Museo del

Instituto de Zoología Agrícola of the Universidad Central

de Venezuela, was rediscovered in 2014 and studied by the

present author while making taxonomic identications to

update the inventory of satyrine butteries in Venezuela.

e second case concerns the unexpected nding made in

2016 by Mauro Costa on his rst expedition to the Ser-

ranía del Cuao, Amazonas state (Costa et al. 2019) of a

hitherto undescribed species of Satyrinae, related to Eup-

tychia eriphule Butler, 1867. As a result of this expedition

we had a single specimen from the slopes of Cerro Woroi

at 500 m above sea level. In 2020, two additional individu-

als were obtained at the base of Cerro Paraka (600 m), not

far from the type locality, whose study conrmed the pe-

culiar characteristics and identity of this taxon.

It was not possible to classify the new species within

the known genera of Neotropical Satyrinae. e peculiar

combinations of their morphological characteristics mer-

ited the proposal and diagnoses of two new genera for the

respective allocation of each of these taxa. e generic

traits diagnosed in each case are shared with a few butter-

y species formerly assigned to genera such as Erichthodes

Forster, 1964 or Yphthimoides Forster, 1964. ese species

have been consequently transferred to the new genera.

MATERIAL AND METHODS

A comparative study has been carried out on the mor-

phology of the venation and the design pattern and col-

oration of the wings, particularly the ocellar formula, as

well as the male genital chitinous structures of the genera

and species of papilionoid Lepidoptera (Nymphalidae:

Satyrinae) described here and some of the taxa that are

apparently related to them. Once the peculiarities of each

taxon were detected, the unique combinations of charac-

ters were used as criteria for the denition of the new gen-

era and new species. For the establishment of taxonomic

hierarchies, the congruence of the localities of origin or

geographic distribution areas of the dierent species was

also considered.

Descriptions follow the nomenclature of wing vena-

tion (and cells) of the Comstock-Needham system (Miller

1970), and the modied terminology of Klots (1970)

was used for the description of male genital structures.

A system of homological equivalences of the ocelli in the

wings was used based on the basic concepts of the ground

plan developed by Schwanwitsch (1924). Dry-preserved,

pinned, and displayed specimens were examined. e ob-

servations, photographs and drawings were made with

and without magnication (in the rst case with manual

magnifying glasses and stereoscopic microscopes of dier-

ent models and brands, natural and articial lighting and

accessories such as the camera lucida and photographic

camera), the lengths were taken with a drawing compass

and ruler, manual and ocular microscales. Wing diaphani-

zations were performed using diluted commercial chlorine

and immediate washing with distilled water, dehydration

with ethanol and preservation by immersion in euparal be-

tween glass slides and covers. e soening and digestion

of eshy tissues and abdominal fat for the microdissection

of the chitinous structures of the male genitalia of but-

teries was carried out by controlled immersion in caus-

tic solution and subsequent washing in water. Once these

preparations were examined, they were stored in a solution

of ethanol and glycerin. ese technical procedures have

been described in more detail by Viloria & Costa (2022).

Abbreviations and acronyms: FW: Forewing; FWL:

Forewing length; HW: Hindwing; IVIC: Centro de

Ecología, Instituto Venezolano de Investigaciones Cientí-

cas, Altos de Pipe, Venezuela; MC: Mauro Costa collec-

tion, Caracas, Venezuela; MIZA: Museo del Instituto de

Zoología Agrícola, Facultad de Agronomía, Universidad

Central de Venezuela, Maracay, Venezuela; NHMUK:

e Natural History Museum, London, United Kingdom;

SNSB: Zoologische Staatssammlung München, Germany.

RESULTS

Rudyphthimoides Viloria, gen. nov.

(Figs. 1a [type species, wing venation], b [type species,

male genitalia], 2a [type species male, dorsal], b [type

species, male, ventral], c [type species, female, dorsal,

d [type species, female, ventral], 3a–f [male genitalia of

dierent species])

http://zoobank.org/urn:lsid:zoobank.org:act:775EEC39-D43A-4065-

9DD4-39B5BB20595F

Type species: Rudyphthimoides iseai Viloria, sp. nov.,

herein designated.

Diagnosis: Similar in size and habitus to Emeryus

Zacca, Casagrande & Mielke, 2020, especially on ventral

view, sharing the possession of all postdiscal ocelli with

double pupils. However, it diers from Emeryus in hav-

ing distinctive male genitalia; Emeryus is characterized

by elongate, rectangular and spatulate valvae, while spe-

cies of Rudyphthimoides gen. nov., have a dierent shape

in their valvae, which are triangular, broad at base, with

a distal projection in the shape of a hook, more similar to

those of the species of Malaveria Viloria & Benmesbah,

[2021]; however they dier from the latter, apart from

other characters mentioned below, in a more developed

Á. L. Viloria

inner ampullar process at the base of the hooked extrem-

ity. In Rudyphthimoides gen. nov., FW veins R2-R5 and

M1 originate together from anterior angle of the distal

extremity of discal cell (R2-R5 separated from M1 in Ma-

laveria); in Rudyphthimoides gen. nov., the recurrent vein

in FW discal cell emerges as a prolongation of M2, thus

dividing m1-m2 from m2-m3 (emerging approximately

in the middle of m1-m2 in Malaveria); Cissia Doubleday,

1848, Magneuptychia Forster, 1964, Paryphthimoides For-

ster, 1964 and Emeryus, all lack a recurrent vein in FW

discal cell and have well developed humeral vein in HW

(apparently absent or not developed in Rudyphthimoides

gen. nov.). Double pupils are present in all ventral ocelli

(even reduced or vestigial) of all species of Rudyphthimoi-

des (unlike Modestia Viloria & Benmesbah, [2021] and

Malaveria which have monopupilled ventral FW ocelli,

and usually – but not always– single pupils in HW R5 and

M1). Wings of the species of Modestia, Malaveria, Mag-

neuptychia, and Stephenympha gen. nov., have no dorsal

ocelli, the same occurs in Emeryus, in which sometimes

there appears a diuse ocellar dark mark on HW Cu1; in

contrast, members of Rudyphthimoides gen. nov., always

bear at least one HW dorsal ocellus with single pupil in

Cu1 (another one smaller in Cu2 may also appear in some

species or in females, plus dark dots on R5, M1 and M3).

Etymology: Rudyphthimoides is an arbitrary combi-

nation of the existing generic name of satyrine butteries

Yphthimoides, and the name of the outstanding Venezue-

lan singer, composer and musician, Rudy Márquez (b. Ro-

dolfo Márquez van Stenis, Caracas, December 7, 1942).

e author of the present discovery and description wants

to honor Master Rudy Márquez for his remarkable and

long professional career, and for his notable contribution

to the cultural development of rock and pop music in Ven-

ezuela during the last six decades.

Distribution: Tropical lowlands of South America, east

of the Andes.

Taxonomic comments: During the process of the pres-

ent publication, Barbosa et al. (2022, March) produced

a relevant and extensive article on the phylogenetic sys-

tematics of the genus Yphthimoides. ey established

the following new combinations: Malaveria anis (But-

ler), Malaveria maepius (Godart) and Malaveria mythra

(Weymer), while considering Yphthimoides argyrospila

(Butler) a valid taxon in uncertain generic position, and

Euptychia eriphule Butler a representative of a new genus.

e latter hypothesis is herein independently conrmed

(see below), but the author of the present contribution

disagrees with the assignations of the other taxa to Mala-

veria. Several morphological characters pointed out in its

diagnosis discriminate Rudyphthimoides gen. nov., from

Figure 1. Morphological features of Rudyphthimoides iseai gen.

et sp. nov.; a. Wing venation of a male individual; b. male geni-

talia: above, dorsal and lateral view of aedeagus; center, lateral

view of genital armature, le valva detached; below, dorsal view

of tegumen, uncus, subunci, vinculum and saccus.

New Satyrinae from Venezuela

Malaveria. ese characters are unequivocally shared by

the following taxa.

Species belonging to Rudyphthimoides:

Rudyphthimoides anis (Butler, 1867), comb. nov.

Euptychia anis Butler, 1867a: 469, pl. 39, g. 9.

[♀, Rio de Janeiro, Brazil]

Euptychia anis Butler; Butler, 1868: 20; 1877a:

118; Kirby, 1871: 49; 1877: 132;

Weymer, 1911: 204; Riley & Gabriel, 1924: 5; Gae-

de, 1931: 437; D’Abrera, 1988: 789, 873; Lamas,

2006: 122-123, g. 5 (female lectotype).

Neonympha thobiei Capronnier, 1881: 102. [♂,

Campos, Brazil]

Euptychia thobiei (Capronnier); Weymer, 1911:

199; Gaede, 1931: 465; D’Abrera, 1988: 762

[row 6] male dorsal and ventral (misidentica-

tions), 877.

Euptychoides anis (Butler); Forster, 1964: 98;

Brown & Mielke, 1967: 91; Brown, 1987: 157.

[Euptychia argyrospila Butler; D’Abrera, 1988: 772-

773 row 1, gs. male dorsal & ventral (misidenti-

cation)]

“Euptychia” nr. thobiei (?) (Capronnier); Emmel &

Austin, 1990: 10.

Yphthimoides wuerttembergiae Anken, 1999: 269,

275-278, gs. 9 (male dorsal), 10 (male ventral),

11 (male genitalia) [male, Poté, Minas Gerais,

Brazil]

Figure 2. Habitus of Rudyphthimoides iseai gen. et sp. nov. (both sexes); a. male Holotype, dorsal view; b. same, ventral view; c. female

Allotype, dorsal view; d. same, ventral view.

Á. L. Viloria

Cissia thobiei (Capronnier); Canals, 2003 (misi-

dentication of Paryphthimoides terrestris grevei

Zacca, Casagrande & Mielke, 2021; Zacca et al.,

2021: 86 (as a misidentication).

Yphthimoides anis (Butler); Freitas, 2004: 10, g.

3 row b [2]; Lamas, 2004: 223; 2006: 123; Petti-

rossi, 2010: 19, 21; Marín, 2011: 50, g. 79 (male

habitus, dorsal and ventral), 56, g. 6 (phyloge-

netic tree), 60, g. 7 (phylogenetic tree), 67, 69,

71, 73, 75, 108; Silva et al., 2015: 6; Marín et al.,

2019: 96; Zacca et al., 2021: 86; Barbosa et al.,

2022: 1, 4, 9, 11, 16.

Magneuptychia thobiei (Capronnier); Piñas-Rubio,

2004: 33, 81, 86, gs. 255-258 (misidentication

of an undescribed taxon).

Yphthimoides wuerttembergiae Anken; Lamas, 2004:

223 (synonymy established); 2006: 123 (as a sy-

nonym).

Neonympha thobiei Capronnier; Lamas, 2006: 122-

123, g. 6 [male lectotype] (synonymy establis-

hed); Zacca et al., 2021: 86 (as a synonym).

Malaveria anis (Butler); Barbosa et al., 2022: 9, 16.

Rudyphthimoides erigone (Butler, 1867), sp. restit.,

stat. rev., comb. nov.

Euptychia erigone Butler, 1867a: 466, pl. 39, g. 5

[♂, São Paulo [de Olivença], Brazil]

Euptychia erigone Butler; Butler, 1868: 18; 1877a:

118; 1877b: 112; Kirby, 1871: 48; 1877: 132;

Druce, 1876: 213; Staudinger, 1887: 226, pl.

81; Weymer, 1911: 203, pl. 47 [row d]; Riley &

Figure 3. Lateral view of the male genitalia of ve dierent species of Rudyphthimoides gen. nov.; a. R. anis (Butler) comb. nov. (=

Yphthimoides wuerttembergiae Anken, reproduced from Anken 1999); b. R. erigone (Butler) sp. restit., stat. rev., comb. nov. (repro-

duced from Forster 1964); c. R. guenzeli (Anken) stat. rev., comb. nov. (reproduced from Anken 1994); d. R. maepius (Godart) comb.

nov. (reproduced from Hayward, [1964]); e. R. maepius (Godart) comb. nov. (= Euptychia argyrospila Butler, syn. nov., reproduced

from Forster 1964); f. R. mythra (Weymer) comb. nov. (reproduced from Forster 1964).

New Satyrinae from Venezuela

Gabriel, 1924: 20; Gaede, 1931: 445; D’Abrera,

1988: 789, 874; Lamas, 2004: 223 [erroneously

as a synonym of Yphthimoides maepius (Godart)].

Yphthimoides erigone (Butler); Forster, 1964: 101,

g.97 (male genitalia), 103; Anken, 1994: 331;

DeVries et al., 1997: 355, 359; Brown & Freitas,

2000: 104; Murray & Prowell, 2004: 70, 72, g.

1, 73, g. 2; 75, g. 3; 76, g. 4 (all gures phylo-

genetic trees).

Yphthimoides maepius erigone (Butler); T. Racheli &

L. Racheli, 2001: 326.

Yphthimoides erigone ? (Butler): Motta, 2002: 158.

[Cissia maepius (Godart); Piñas-Rubio, 2004: 31,

76, 80, gs. 227 (female, habitus, dorsal), 228

(female, habitus, ventral), 229 (male, habitus,

dorsal), 230 (male, habitus, ventral), 231 (male,

habitus, dorsal), 232 (male, habitus, dorsal) (mi-

sidentications)].

Rudyphthimoides guenzeli Anken, 1994, stat. rev.,

comb. nov.

Yphthimoides guenzeli Anken, 1994: 327, 328-331,

gs. 1a (male, habitus, dorsal), 1b (male, habitus,

ventral), g. 2 (male genitalia). [♂, Navirai, Mato

Grosso do Sul, Brazil]

Yphthimoides guenzeli Anken; Anken 1999: 269,

277, 278.

Yphthimoides maepius guenzeli Anken; Lamas, 2004:

223; Marín et al., 2019: 97.

Rudyphthimoides iseai Viloria, sp. nov. [♂ & ♀, El Li-

món, Aragua, Venezuela]

Rudyphthimoides maepius (Godart, [1824]), comb.

nov.

Satyrus maepius Godart, [1824]: 490. [♀, Guyane]

Euptychia maepius (Godart); Westwood, 1851: 373;

Butler, 1867a: 465; 1868: 28; 1877a: 122; Kir-

by, 1871: 51; Weymer, 1911: 202, pl. 47 [row] d;

Gaede, 1931: 453-454; Hayward, [1964]: 512,

513, g. 97 (male genitalia).

Neonympha maepius (Godart); Herrich-Schäer,

1865: 70.

Euptychia argyrospila Butler, 1867a: 467, pl. 11, g.

12, syn. nov. [♀, Ega, Amazonas, Brazil]

Euptychia argyrospila Butler; Butler, 1868: 18;

1877a: 118; Kirby, 1871: 48; Weymer, 1911:

203; Möschler, 1877: 323; Gaede, 1931: 439.

Euptychia perfuscata Butler, 1868: 18, syn. nov. [♂,

Pará, Brazil]

Euptychia argyrospila Butler; Butler, 1868: 18; Kir-

by, 1877: 132; D’Abrera, 1988: 772-773 [row 1,

gs. male dorsal & ventral (misidentication of

R. anis (Butler))].

Euptychia perfuscata Butler; Butler, [1870]: 13; Kir-

by, 1871: 48; Weymer, 1911: 209, pl. 48 [row] d;

Gaede, 1931: 460.

Euptychia nausiaca Möschler, 1883: 320, pl. 17, g.

14 [♀, Surinam] (synonymy established by La-

mas, 2004c: 223)

Euptychia sp.; Sanders, 1904: 364, 371, pl. 6, g. 8

(male habitus, dorsal).

Euptychia nausiaca Möschler; Weymer, 1911: 203.

Euptychia argyrospila Butler var. nausicaa [sic] Mös-

chler; Gaede, 1931: 439.

Yphthimoides argyrospila (Butler); Forster, 1964:

103, 104, g. 102 (male genitalia), 105; Mielke

& Casagrande, 1992: 180; Anken, 1994: 327,

330, 331; 1999: 269, 277, 278; Kochalka et al.,

1996:210; Lamas, 2004: 223; Emery et al., 2006:

90; Gareca et al., 2006: 52; Gareca & Reichle,

2007: XII; Brévignon, 2008: 81; Pulido & An-

drade, 2010: 541, 556; Marín et al., 2019: 97;

Barbosa et al., 2022: 2, 9, 11, 15.

Yphthimoides nausicaa [sic] (Möschler); Forster,

1964: 103

[Yphthimoides maepius (Godart); Lewis, 1973: 58,

g. 10 (female habitus, ventral) (misidentica-

tion of Cissia penelope (Fabricius))]

Euptychia moepius [sic] (Godart); D’Abrera, 1988:

772 [rows 6 & 7, gs. (3 males)].

“Euptychia” perfuscata Butler; Emmel & Austin,

1990: 10.

Yphthimoides maepius (Godart); Kochalka et al.,

1996: 210; Lamas et al., [1997]: 65; T. Racheli

& L. Racheli, 2001: 326; Murray, 2001: 115,

128,129, 130, 220, g. 3.29, 236, 237, 268, 282,

g. 4.2, 290, g. 4.6; 297, g. 4.9, 301, g. 4.11,

303, g. 4.12, 308, g. 4.4 (all gures phylogene-

tic trees); Beccaloni et al., 2008: 345 (hostplant);

Marín et al., 2019: 97; Espeland et al., 2019: 118,

g. 1 (phylogenetic tree); Garwood et al., 2021:

148; Ramos-Artunduaga et al., 2021: 62; Barbosa

et al., 2022: 1, 4, 9,11, 16.

Yphthimoides moepius [sic] (Godart); T. Racheli &

L. Racheli, 2001: 326 (as a misspelling).

Maressia maepius (Godart) nom. nud.; Murray,

2001: 238.

Marissia maepius (Godart) nom. nud.; Murray,

2001: 331, 335, g. 5.1, 339, g. 5.3, 343, g. 5.4

(all gures phylogenetic trees).

Yphthimoides maepius maepius (Godart); Lamas,

2004: 223; Emery et al., 2006: 90; Brévignon,

2008: 81; Marín et al., 2019: 97.

Á. L. Viloria

Yphthimoides maepius perfuscata (Butler); Lamas,

2004: 223; Marín et al., 2019: 97.

Cissia maepius (Godart); Piñas-Rubio, 2004: 31, 76,

80, gs. 227 (female, habitus, dorsal), 228 (fema-

le, habitus, ventral), 229 (male, habitus, dorsal),

230 (male, habitus, ventral), 231 (male, habitus,

dorsal), 232 (male, habitus, dorsal) (misidenti-

cations of Rudyphthimoides erigone (Butler),

comb. nov.)

[? ] argyrospila (Butler); Barbosa et al., 2022: 9.

Malaveria maepius (Godart); Barbosa et al., 2022:

9, 16.

Rudyphthimoides mythra (Weymer, 1911), comb. nov.

Euptychia mythra Staudinger, in litt.; Weymer, 1911:

205, pl. 47 [row e]. [♂, Bolivia]

Euptychia mythra Weymer; Gaede, 1931: 456;

D’Abrera, 1988: 774, 775 [row 1, gs. Male dor-

sal & ventral]

Yphthimoides mythra (Weymer); Forster, 1964: 103,

104, g. 104 (male genitalia), 106; Brown & Mie-

lke, 1967: 91; Anken, 1994: 331; 1999: 273, 278;

Lamas, 2004: 223; Emery et al., 2006: 90; Gareca

et al., 2006: 52; Marín et al., 2019: 97; Barbosa et

al., 2022: 1, 2, 9, 11, 16.

Yphthimoides mythe Staudinger, in litt., Forster,

1964: 106, nomen nudum (synonymy established

by Lamas, 2004: 223).

Yphthimoides cf. mythra (Weymer); Kochalka et al.,

1996: 210.

Yphthimoides mythra ? (Weymer); Motta, 2002:

158.

Malaveria mythra (Weymer); Barbosa et al., 2022:

9, 16.

Rudyphthimoides iseai Viloria, sp. nov.

(Figs. 1a, b; 2a, b, c, d)

http://zoobank.org/urn:lsid:zoobank.org:act:885AF61E-5E3C-401B-93AE-

6936017895A8

Type material: H ♂, Venezuela, AR[agua],

El Limón, 450 m, 30-I-[19]53, F. Fernández Y. col.; A-

 ♀, Venezuela, edo. Aragua, Maracay, El Limón,

5-I-[19]53, F. Fernández Y. col.; P: (3 ♂ ♂, 3

♀♀) 1 ♂, Venezuela; AR[agua], El Limón cr. Maracay,

8-IV-[19]52, F. Fernández Y. col.; 1 ♀, same data, 17-IV-

[19]52; 1 ♂, Venezuela, E. Aragua, Maracay, El Limón,

500 m, 11-XII-[19]52, F. Fernández Y. col. (genit. prep.

ALV518-14); 1 ♀, same data, 12-XII-[19]52; 1 ♀, same

data as holotype, 23-I-[19]53; 1 ♂, same data as holotype,

24-II-[19]53 (wing prep. ALV059-19) [MIZA].

Description: Male (Figs. 1a, b, 2a, b). FWL: 20-22 mm

(mean: 20.87; n = 4). Body brown, ventrally lighter. Palpi

twice as long as head, creamy ochraceous with brown elon-

gated scales below. Antenna thin, dark brown, club formed

gradually, reaching two thirds of the length of the costa.

FW triangular, costal margin slightly arched, anal margin

almost straight, outer margin regular and smooth; HW

rounded, suboval, costal and anal margins slightly curved,

outer margin slightly wavy. Dorsal ground color brown,

slightly darker towards distal quarter; one wavy submar-

ginal dark line, faded on FW; two marginal dark lines

running parallel to each other, lighter brown in between;

dense marginal scaling forming fringes along outer mar-

gins. FW with darker color over the veins distally closing

discal cell; one dark (almost black) subapical ocellar spot

in cell M1, with orange halo. HW upperside with irregu-

lar postdiscal darker line, and a darker mark over vein m1-

m2; darker postdiscal circular spots, very diuse on R5,

M1 and M3; prominent postdiscal black ocellus on Cu1,

with centered white pupil, ringed with yellow-orange,

nely circled with dark brown; another similar but smaller

(a third of Cu1’s diameter) on Cu2. Ventral ground color

very light, creamy brown; basal half of wings densely dust-

ed with brown; irregular postbasal darker lines on both

wings, almost straight; another one similar, limiting dis-

cal and postdiscal areas, more regular and straight; diuse

umbra developed on both wings in the middle of discal

area, above which a series of double-pupilled ocelli, as fol-

lows: FW, three to four ocelli in cells M1(black), M2, M3

and (sometimes) Cu1, last three not black but brown and

less well dened, all three or four circled with yellow; HW,

ve to six double-pupilled ocelli occupying cells R5, M1,

M2, M3, Cu1 and Cu2 (vestigial), those on R5, M1, and

Cu1 black, others brown, all circled with yellow, pupils of

R5, M2 and M3 silver, the others white; both wings with

submarginal dark brown line, wavy or zig-zagging; two

parallel ne lines running along outer margins.

Genitalia (Fig. 1b). Tegumen well developed, semi-

globular; uncus almost twice as long as tegumen’s dome,

laceolate; subunci stylized, two thirds length of uncus,

emerging laterally from below base of uncus and directed

upwards; vinculum thin but strong; saccus as long as sub-

unci, semitubular, attened at base, moderately inated

at anterior extremity; valvae subtriangular, very broad at

base, with an inner aileron developed from ampulla as a

process at the base of the digitiform extremity, which is

curved inwards as a hook; aedeagus straight, depressed,

with broad basal opening (dorsal).

Female (Figs. 2c, d). FWL: 19-23 mm (mean: 21.25;

n = 4). Similar to male, but larger on average, slightly

lighter in color. Dorsal FW ocellar spot reduced, as well as

New Satyrinae from Venezuela

dorsal HW Cu2 ocellus. HW ventral lines on both sides

of discal area more irregular and zig-zagging.

Etymology: is buttery is named in memoriam of

the Venezuelan composer, arranger and virtuoso musician,

Professor Dr. Douglas Isea (b. Douglas Alcíades Isea Re-

villa, Maracaibo, September 27, 1951; d. April, 24, 2020).

Isea’s exceptional abilities and own style in the execution

of the Venezuelan cuatro was inuential in the develop-

ment of modern trends of local folk and popular music, as

well as pioneering the adaptation of cuatro performing to

accompany classic orchestra. He was responsible for intro-

ducing cuatro in worldwide recognized examples of rock

and pop music.

Distribution, habitat and behaviour: Nothing is

known about the habitat preferences of Rudyphthimoides

iseai gen. et sp. nov. is species is probably locally com-

mon in northern Venezuela, but only known so far at low

elevations on the foothills of the Cordillera de La Costa.

ere is no mention of it, or any other taxon with similar

aspect that could have been confused with it, in previous

records or species lists for the region (Martorell 1939, For-

ster 1949, Beebe 1951, Osuna 2000, Sandoval-Cabrera

et al. 2008, Ríos-Málaver et al. [2021]). Two records of

Emeryus argulus magnum Zacca, Casagrande & Mielke

(Zacca et al. 2020) in the Parque Nacional Henri Pittier

(Rancho Grande, 1,125 m) might be either mislabelled

specimens or misidentications of R. iseai gen. et sp. nov.

e present author does not know of any reliable record

of butteries of the genus Emeryus, north of the Orinoco

River, in Venezuela.

Stephenympha Viloria, gen. nov.

(Figs. 4a [type species, wing venation], b [wing venation

S. eriphule (Butler) comb. nov.]; 5a [type species, male

habitus, dorsal,], b [type species, male habitus, ventral,],

c [male habitus S. eriphule (Butler) comb. nov., dorsal],

d [male habitus, S. eriphule (Butler) comb. nov., ventral];

6a [type species, male genitalia], b [male genitalia,

S. eriphule (Butler) comb. nov.], c [male genitalia,

S. arius (Weymer) comb. nov.])

http://zoobank.org/urn:lsid:zoobank.org:act:91261755-8F4D-4944-8B10-

3FCBC0E9CD0D

Type species: Stephenympha pauliana Viloria, sp. nov.,

herein designated.

Diagnosis: e members of Stephenympha gen. nov.,

may look supercially similar to many related medium-

sized neotropical lowland satyrine buttery (e. g., species

of Cissia and Vareuptychia Forster, 1964), but they are un-

like any other in several aspects: 1. ey lack any visible

dorsal marking or ocelli (several other genera do, like Mag-

neuptychia, Emeryus, Modestia, Malaveria and Hermeup-

tychia Forster, 1964); 2. ey have ve postdiscal, mono-

pupilled ocelli on ventral HW (R5 to M3), sometimes a

sixth, supranumerary, minute ocelli, also develops on the

anal margin (cell A2), and a well-developed to vestigial

ocellus may appear on ventral FW M3, which is peculiar;

3. In the male genitalia the uncus is unique, as it is heavily

bended downwards, either at its base or half way its length;

the subunci emerging laterally, usually elevated above the

base of the uncus. e character of its uncus is unique and

might represent a synapomorphy. As for its wing venation,

Stephenympha gen. nov., is similar to several other genera

(e. g., Cissia, Magneuptychia, Paryphthimoides, Emeryus

and Malaveria) in that their FW veins R2, R3-R5 and M1

all emerge together from the anterior angle of the distal

extremity of discal cell, and in having no recurrent vein in

FW discal cell, and a similar disposition of crossveins m1-

m2 and m2-m3.

Etymology: is genus of nymphalid buttery is

named in remembrance of the internationally renowned

Venezuelan musician, actor, composer and extraordinary

singer, Henry Stephen (b. Henry Augustus Stephen Pierre,

Cabimas, July 15, 1941; d. Caracas, April 5, 2021).

Distribution: West of the Andes, Amazonian lowlands

and Pantepui.

Taxonomic comments: De Lesse (1967: 36-38, g.

2) dissected, examined, and illustrated the male genitalia

of a buttery from Coroico, Bolivia, in all its appearance

consistent with the species Euptychia arius, originally de-

scribed and illustrated by Weymer (1911: 198, pl. 47 [row

a]). In this way, he detected the disparity between the im-

age of the male genitalia illustrated by Forster (1964: 100

g. 93) for Yphthimoides arius (Weymer) and the structure

dissected and illustrated by him. Judging by the presence

of the intact abdomen in the cotype from Mapiri [Bolivia]

preserved at the Zoologische Staatssammlung München,

and despite the fact that it bears the label “Präparat Nr

SA25” placed by Forster, it is evident that it was not dis-

sected and therefore consequently never illustrated, so it

follows that Forster’s Figure 93 necessarily corresponds to

the genitalia of an indeterminate species that is not arius.

us, relying on De Lesse’s excellent illustration (repro-

duced here in Figure 6c) it is possible to conrm the ex-

traordinary orientation of the uncus in this species, point-

ing directly downwards, due to its emergence at right angle

in relation to the natural axis of the tegumen. Such charac-

ter led De Lesse to suggest the creation of a new genus for

Euptychia arius.

We have discovered in our comparative studies the

existence of at least two other species of Amazonian dis-

tribution (both found in Venezuela, one described here)

Á. L. Viloria

that share a similar trait, although less extreme than that

of arius. ese species, mentioned below, also share with

arius the basic ocellar pattern on the ventral side of their

wings. ese homologies allow for grouping these three

taxa within Stephenympha gen. nov.

Species belonging to Stephenympha:

Stephenympha arius (Weymer, 1911), comb. nov.

Euptychia arius Weymer, 1911: 198, pl. 47 [row a].

[♂, Mapiri, Bolivia]

Euptychia arius Weymer; Gaede, 1931: 439;

D’Abrera, 1988: 789, 873.

Yphthimoides arius (Weymer); Forster, 1964: 100,

g. 93 (male genitalia, erroneous), 102, 103;

Anken, 1994: 331.

Euptychia (s. l.) arius Weymer; De Lesse, 1967: 36,

g. 2 (male genitalia), 37-38.

Erichthodes arius (Weymer); Lamas, 2004: 219; Ga-

reca et al., 2006: 50.

Erichthodes s. l. arius (Weymer); Brown et al., 2007:

473.

Stephenympha eriphule (Butler, 1867), comb. nov.

Euptychia eriphule Butler, 1867a: 468, pl. 39, g. 6.

[♂, Pernambuco, Brazil]

Euptychia eriphule Butler; Butler, 1868: 19; 1877a:

118; Kirby, 1871: 49; Weymer, 1911: 203; Riley

& Gabriel, 1924: 20; Gaede, 1931: 446; D’Abrera,

1988: 773 [row 5, g. female ventral].

Yphthimoides eriphule (Butler); Lamas, 2004: 223;

Brévignon, 2008: 80-81, 82, 84, 90 gs. 67a, b

(male genitalia), 67c (male habitus, dorsal le,

ventral right), 94; Brévignon & Benmesbah,

2012: 52; Marín et al., 2019: 97; Barbosa et al.,

2022: 1, 3, 9, 11, 16.

[New genus 1] eriphule (Butler); Barbosa et al.,

2022: 9.

Stephenympha pauliana Viloria, sp. nov. [♂, Cerro

Cuao, Amazonas, Venezuela]

Stephenympha pauliana Viloria, sp. nov.

(Figs. 4a; 5a, b; 6a)

http://zoobank.org/urn:lsid:zoobank.org:act:713D5AA1-B5CC-4E8A-

9ED0-AE5749C562DB

Type material: H ♂, Venezuela, Amazonas,

Serranía del Cuao, subida al Cerro Woroi, 500 m, 20-

III-2016, M. Costa (genit. prep. ALV361-17) [MIZA];

P, 2 ♂♂, Venezuela, Amazonas, Serranía del

Cuao, Cerro Paraka, 600 m, 14/21-II-2020, M. Costa [1

IVIC (wing prep. ALV060-21), 1 MC].

Figure 4. Wing venation of male individuals of Stephenympha gen. nov.; a. S. pauliana sp. nov.; S. eriphule (Butler) comb. nov.

New Satyrinae from Venezuela

Description: Male (Figs. 4a; 5a, b; 6a). FWL: 18–21.5

mm (mean: 19.33, n = 3). Body dorsally dark chocolate

brown, ventrally lighter; antennae, dark brown, thin, club

formed gradually, reaching two hs of costa; palpi almost

twice as long as head, dark brown, with sparse elongated

light brown scales, laterally very light brown. FW subtri-

angular, costa moderately curved, outer margin smooth,

anal margin straight, apex and tornus rounded; HW

suboval, costa moderately curved, as well as anal margin,

outer margin barely scalloped, apex and tornus rounded.

Upperside ground color dark chocolate brown, slightly

darker on anterior tip of FW apex and especially on dis-

cal cell and adjacent costal area, where there appears to

develop an androconial patch (Fig. 5a); both wings with

three marginal lines running parallel, the inner one wavy,

the middle one more regular, and the outer one running

through margin and forming the fringes, all three diuse

on FW, distinct on HW. Underside ground color light

chocolate brown, tending to olive green towards center

of wings; submarginal and marginal areas lighter, darker

postdiscal umbra on FW; two dark brown stripes limiting

both sides of discal area, the postbasal one slightly curved

distally, as well as the distal one on HW; the latter straight

on FW; small dark maculae at distal extremity of discal

cells; submarginal dark line wavy, more markedly on HW;

two neat, dark marginal lines running parallel along outer

Figure 5. Habitus of male individuals of two species of Stephenympha gen. nov.; a. S. pauliana sp. nov., Holotype, dorsal view; b. Same,

ventral view; d. S. eriphule (Butler) comb. nov., Venezuela, edo. Bolívar, Auyántepui, Guayaraca, 1,000 m, dorsal view; e. Same, ventral

view.

Á. L. Viloria

margins; a series of ve postdiscal ocelli on HW underside,

occupying cells R5 to Cu1, respectively, nely ringed with

ochraceous yellow, the rst one smaller than the others;

those on R5, M1 and Cu1 black, with white central pupils,

but with silver scales, either scattered (R5 more than M1)

or concentrated at basal border of ocellus (Cu1) forming a

silver lunula; two ocelli, M2 and M3, silver, without a vis-

ible white pupil, but with some dusting of black and white

scales. is ocellar pattern is very similar to that of S. er-

iphule (Butler) comb. nov.

Genitalia (Fig. 6a). Tegumen semiglobular, somewhat

depressed, basally 1.5 times wider than distally; uncus thin

and stylized, as long as tegumen, rising at the lowest level

of distal extremity of the latter, emerging inclined down-

wards approximately 45° in relation to it, halfway its length

it bends further 30–40° downwards; subunci as long as un-

cus, but more stylized and regularly curved, emerging lat-

erally at distal extremity of tegumen, inclined upwards and

inwards at base, developing at about same position of max-

imum elevation of tegumen, their tips touching each other

in middle axis; vinculum stylized, more or less straight

both in lateral and dorsal views; saccus slightly shorter

than tegumen, tubular and slightly depressed at base; valva

subtriangular, large and broad, ending in a digitiform dis-

tal process, nearly a h of its maximum length; a subter-

minal, shorter, secondary inner process; aedeagus regular,

depressed; both extremities slightly curved upwards.

Female. Unknown.

Etymology: is new species is named aer Paul Mc-

Cartney (b. James Paul McCartney, Liverpool, UK, June

18, 1942), an animal rights advocate, but best known and

celebrated as a highly creative and original songwriter,

singer and musician. His remarkable artistic work has

achieved an unprecedented cultural impact worldwide

over the last sixty years.

Distribution, habitat and behaviour: Stephenympha

pauliana gen. et sp. nov., might be a widespread Amazo-

nian species. However, it is known only from the foothills

of the Serranía del Cuao (500–600 m), a Tepuyan moun-

tainous region in northwestern Amazonas state in Venezu-

ela, which belongs in the Guiana Shield. e habitat of S.

pauliana gen. et sp. nov., is somewhat particular. It is with-

in an area in which there are many rocks and stones and

little soil lling the space between them, with small and

thin trees, which makes it possible to walk freely among

them. e oor is always covered by dry leaves where this

buttery rests from time to time. It has neither been found

anywhere else nor at dierent elevations. e fact that the

only individuals known where found exactly in the same

kind of places indicates that this is the habitat preferred

by this species (M. Costa, pers. comm. 2022). More data

on the geographical position, the environment and some

features of the vegetation of this sector of the Serranía del

Cuao have been described by Costa et al. (2019, 2020,

[2021]), although mainly what corresponds to the pan-

tepuyan life zone, which develops at higher altitudes.

Additional material examined: Stephenympha arius

(Weymer) comb. nov.: 1 male, cotypus Euptychia arius

Weymer, [Bolivia], Mapiri, collection v. Rosen, Präparat

Nr SA25 Zoolog. Staatssammlung München [SNSB];

Figure 6. Comparison of the male genitalia of three species of Stephenympha gen. nov.; a. S. pauliana sp. nov., Holotype (genit. prep.

ALV361-17): above, dorsal and lateral view of aedeagus; center, lateral view of genital armature, le valva detached; below, dorsal view

of male genital armature (aedeagus removed); b. S. eriphule (Butler) comb. nov., lateral view of entire genital capsule, aedeagus in

natural position (genit. prep. ALV516-14); c. S. arius (Weymer) comb. nov., lateral view of entire genital capsule, aedeagus in natural

position (reproduced from De Lesse 1967).

New Satyrinae from Venezuela

Stephenympha eriphule (Butler) comb. nov.: 1 male, syn-

type, Euptychia eriphule Butler Monog., [Brazil], Per-

nambuco, pres. By M. S. Smith 45-70, B. M. Type No.

Rh.3204 [NHMUK]; 1 female, (ST-068-1), [Venezuela],

edo. Bolívar, Qda. El Jaspe, 24-XII-1983, [M. Costa] (ge-

nit. prep. ALV516-14); 1 male, Venezuela, edo. Bolívar,

Auyántepui, Guayaraca, 1,000 m, 28.III.2013, [M. Costa]

(genit. prep. ALV517-14); 1 male, (12z), same data, 28-

III-2015; 2 males (10z, 11z), same data, 5-IV-2015 (1

wing prep. ALV061-19); 2 males (28b), Venezuela, edo.

Bolívar, Akopán Tepui, entre Yunek y Piaima Parú, 900

m, 31-XII-2013, M. Costa; 1 male (4a), Venezuela, edo.

Bolívar, Ptarí Tepui, Salto Iworé, 1,400 m, 19-XII-2014,

M. Costa [MC].

ACKNOWLEDGEMENTS

I thank Mauro Costa for selessly providing part of

the biological samples and information used in this study,

and for sharing his inspiring enthusiasm about exploring

the remotest areas of Venezuela in search for new butter-

ies. José A. Clavijo, Jürg De Marmels, Marco A. Gaiani,

uintín Arias and Luis J. Joly are warmly acknowledged

for granting me open access to the collections of MIZA

and for their hospitality in Maracay. Héctor Suárez, Mau-

ro Costa, Indiana Cristóbal Ríos-Málaver, and Rafael Car-

reño assisted me in several ways to produce the illustra-

tions for this article. Tito R. Barros, Gilson A. Rivas, I. B.

Petit, Neva Mora and María Eugenia Andara read partial

dras of this article and helped editing its content.

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