Viloria & Costa

ANARTIA

Publicación del Museo de Biología de la Universidad del Zulia

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

https://doi.org/10.5281/zenodo.18111812 / Anartia, 41 (diciembre 2025): 32-39

http://zoobank.org/urn:lsid:zoobank.org:pub:C8B9042B-D666-4A48-8219-259B5B0018BA

Fernanmorana, new genus of butterflies from the Amazon

region (Lepidoptera: Nymphalidae, Satyrinae)

Fernanmorana, nuevo género de mariposas de la región amazónica

(Lepidoptera: Nymphalidae, Satyridae)

Ángel L. Viloria1,2 & Mauro Costa3

1 Centro de Ecología, Instituto Venezolano de Investigaciones Científicas (IVIC), km 11 carretera Panamericana, Altos de Pipe,

estado Miranda, Venezuela. https://orcid.org/0000-0002-5747-4747

2 Coordinación de Ciencia, Tecnología y Educación, Organización del Tratado de Cooperación Amazónica (OTCA), SEPN 510,

Bloco A, 3° Andar, Asa Norte, CEP 70750-521, Brasilia, D. F., Brasil

3 Museo del Instituto de Zoología Agrícola, Facultad de Agronomía, Universidad Central de Venezuela,

Maracay, estado Aragua, Venezuela. https://orcid.org/0009-0000- 7771-3904

Correspondence: sebastianviloriacarrizo@gmail.com

(Received: 10-09-2025 / Accepted: 26-12-2025 / On line: 31-12-2025)

ABSTRACT

Based on comparative morphological evidence and previously published phylogenetic hypotheses, a new genus of satyrine

nymphalid butteries, Fernanmorana Viloria & Costa, gen. nov., is diagnosed and described. It is native to the lowlands of

the Amazon region and so far known from Brazil and Venezuela. e taxonomy of the new genus is briey discussed, and

some bionomic data of its type species, Fernanmorana insignis (Butler, 1867), comb. nov., are provided.

Keywords: Brazil, Euptychia ordinata, Maroa, Raudal del Danto, Serranía del Cuao, Tocantins, Venezuela, Yavita.

RESUMEN

Con base en evidencia morfológica comparativa e hipótesis logenéticas previamente divulgadas, se diagnostica y describe

un nuevo género de mariposas ninfálidas satirinas, Fernanmorana Viloria & Costa, gen. nov. El mismo es originario de las

tierras bajas de la región amazónica y hasta la fecha se conoce en Brasil y Venezuela. Se discute brevemente la taxonomía

del nuevo género y se proporcionan algunos datos bionómicos de su especie tipo, Fernanmorana insignis (Butler, 1867),

comb. nov.

Palabras clave: Brasil, Euptychia ordinata, Maroa, Raudal del Danto, Serranía del Cuao, Tocantins, Venezuela, Yavita.

in the Amazonas state of Venezuela. e main objective of

this initiative, in addition to traveling by kayak and row-

boat along the aforementioned waterways from San Car-

los de Río Negro to Puerto Ayacucho, was to test, for the

rst time, internet communication from selected remote

locations of the planet using the rst commercially avail-

able laptops equipped with satellite antennas.

is initiative was called the Amazon Netspedition

and took place between July and October 1996, under the

INTRODUCTION

In 1996, while one of the authors (ÁLV) was pursu-

ing doctoral studies at the Natural History Museum in

London, he presented to the Venezuelan diplomatic au-

thorities in the United Kingdom an initiative by a group

of postgraduate students (engineers) from the Imperial

College London to undertake a recreational and scientic

expedition to the rivers Negro, Casiquiare, and Orinoco

New genus of butterflies from the Amazon region

leadership of Anthony Heenan (Anonymous 1996). e

group included the Venezuelan agronomist Jesús Cama-

cho, professor of entomology and curator of the Arthro-

pod Museum of the University of Zulia (MALUZ, Mara-

caibo, Venezuela), who made extensive insect collections

during the expedition, particularly Lepidoptera and Cole-

optera. Some of the information posted on the Netspedi-

tion website (http://sunsite.doc.ic.ac.uk/netspedition)

referred to the discovery of unusual buttery species, some

of which were discussed in the rst part of the work of A.

F. E. Neild, published at the end of that year (Neild 1996).

Due to bureaucratic delays in obtaining permits, the en-

tomological material was conscated by the authorities of

the Fauna Division of the Ministry of the Environment

and Natural Resources in Caracas, and unfortunately was

not available for study until years later, when some of it

was returned to MALUZ. In 2002, it became possible

to display, mount, preserve, and examine these samples,

which also allowed for the preliminary identication of

most buttery species (Lepidoptera, Papilionoidea). e

nymphalid species of the subfamily Satyrinae were studied

by Viloria & Camacho (2002), with several cases of provi-

sional identications, mainly based on the work of Butler

(1867), Forster (1964), and D’Abrera (1988).

In the collection of Amazonian Satyrinae of the

Netspedition 1996, a male specimen of ‘Euptychia’ in-

signis Butler was identied, the rst one known from the

Venezuelan territory, which, due to its morphological pe-

culiar aspect, especially wing color pattern, could not be

placed within any of the genera of Euptychiina described

by Forster (1964).

In more recent dates Lamas (2004) detected the impos-

sibility of placing this buttery species in any of the gen-

era of Nymphalidae Satyrinae described for the American

continent. Freitas et al. (2018), Marín Uribe et al. (2019),

Nakahara et al. (2019) and Espeland et al. (2023) followed

this opinion, adding important genetic evidence that re-

inforces the notion that this taxon should be placed in its

own genus. Herein we proceed to diagnose and describe it.

MATERIALS AND METHODS

A comparative study has been carried out on the mor-

phology of the venation and the design pattern and color-

ation of the wings, particularly the band system and ocelli

formula, as well as the male genital chitinous structures

of the taxon mentioned in the introduction. e unique

combinations of characters were used as criteria for the

denition of the new genus.

Description 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.

Dry-preserved, pinned, and displayed specimens were

examined. e observations, photographs and drawings

were made with and without magnication (in the rst

case with manual magnifying glasses and stereoscopic

microscopes of dierent 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 diaphanizations were performed using diluted com-

mercial chlorine and immediate washing with distilled

water, dehydration with ethanol and preservation by im-

mersion in euparal between 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 butteries was carried out by controlled

immersion in caustic solution and subsequent washing in

water. Once these preparations were examined, they were

stored in a solution of ethanol and glycerin. ese techni-

cal procedures have been described in more detail by Vilo-

ria & Costa (2022) and Viloria (2022).

Acronyms: IVIC: Centro de Ecología, Instituto Vene-

zolano de Investigaciones Cientícas, Altos de Pipe, Ven-

ezuela; MALUZ: Museo de Artrópodos de la Universidad

del Zulia, Facultad de Agronomía, Maracaibo, Venezuela;

MC: Mauro Costa collection, 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.

Material examined: BRAZIL: 1♀ Tonantins [sic],

Amazons [white rectangular label, printed], Godman &

Salvin Coll. 1904-1, Euptychia insignis Btl. [white rectan-

gular labe, printedl] Type of species [white rectangular la-

bel, printed], B.M. Type Rh 3867, Euptychia insignis Butl.

♀ [white square label, part printed, part handwritten],

Syn-type [small round label circled in light blue, printed],

Type HT [small round label circled in red, printed], Eup-

tychia insignis Butler Monog. [larger light blue rectangu-

lar label, handwritten]. is specimen corresponding well

with the illustration of the original description (Butler

1867: pl. 40, g. 12), herein designated LECTOTYPE of

Euptychia insignis Butler, 1867 [NHMUK]; VENEZU-

ELA: 1♂, Amazonas, Mcpio. Guainía, camino Yavita-Ma-

roa, 2° 55’ 16” N, 67° 26’ 25” W, 300 m, 29-31.viii.1996,

cols. J. Camacho, A. Heenan [wing prep. ALV062-19;

genit. prep. ALV599-14] [MALUZ]; 1♀, Amazonas,

Raudal del Danto [5° 2’ 30” N; 67° 33’ 30” W], 150 m,

13.i.2024, H. Camico [IVIC], 5 ♂♂ Amazonas, Raudal

del Danto [5° 2’ 30” N; 67° 33’ 30” W], 150 m, 25.x.2024,

Viloria & Costa

H. Camico [3 IVIC, 1 MIZA, 1 MC]; 2♂♂ Amazonas,

Raudal del Danto [5° 2’ 30” N; 67° 33’ 30” W], 150 m,

30.x.2024, H. Camico [1 IVIC, 1 MALUZ].

RESULTS

Fernanmorana Viloria & Costa, gen. nov.

(Figs. 1 [type species, male wing venation],

2 [type species, male genitalia], 3 [type species, habitus,

male and female])

https://zoobank.org/urn:lsid:zoobank.org:act:AE53C6EF-9EF1-44C3-857A-

8E207FFAD9AD

Type species: Euptychia insignis Butler, 1867: 501, pl.

40, g. 12. Herein designated.

Diagnosis. Wing pattern (Fig. 3): dorsal: uniform dark

brown, without ornamentation; ventral forewing without

ocelli and lacking discal band or line; hindwing with a

distinct, straight dark discal stripe or line, running from

anterior margin, almost reaching anal margin; postdiscal

line running parallel but only approximately to half wing

width; hindwing distinctly patterned with a white postdi-

cal band NOT parallel to the line-stripe system, but in-

stead crossing the wing approximately from the ápex to

the middle of the anal margin. is feature is unlike that

of most other white banded satyrid from the lowlands of

Tropical America (e. g., Argentaria Huertas & Wilmott,

2023, Cristalinaia Freitas, Barbosa & Zacca, 2019, Sau-

rona Huertas & Willmott, 2023) and only comparable

to that of Forsterinaria pronophila (Butler, 1867) or the

species of Splendeuptychia Forster, 1964, from which it

diverges in their dierent system of postdiscal ocelli. ey

even belong to dierent clades (Espeland et al. 2023).

Postdiscal ocellar elements well developed, inmerse in a

diuse cloud of yellow color, possibly resulting from the

expansion and fusion of the “yellow rings” of the ocelli,

which are usually 3 to 4, from M2 to Cu2 (notably larger

and prominent in Cu1, oen vestigial or absent in M2),

black, with white pupils, triangular or v-shaped in M2

Description. Eyes glabrous. Antennae dark, with some

white scales on the joints of their segments, threadlike

(liform), thin, barely reaching a third of the length of

the wing costa; club faintly formed, barely perceptible.

Forewings subtriangular, apex subtruncate, smooth, outer

margin gently convex. Hindwings suboval, apex and anal

angle rounded, anterior margin slightly angled at the end

of Sc +R1, outer margin arched, with very slight crenula-

tion, anal margin straight. Venation (Fig. 1, male). Fore-

wing: Sc notably inated in its basal two-hs, ending at

the anterior two-hs of the costa length; independent

R1 emerges midway through the discal cell and terminates

approximately three-hs of the costa length; R2 and R3-

R5 emerge together from the anterior apex of the discal

cell, R2 terminating at one-quarter of the costa length, R3

emerges one-third of the length of R4-R5 and terminates

before the wing apex, R4 and R5 diverge approximately

halfway along the radial axis of R3-R5, R4 terminating at

the wing apex itself; free end of R5 short, terminating at the

posterior limit of the wing apex; M1 long and completely

independent, emerges posteriorly and distinctly separated

from the origin of R2-R5 (between them a small, straight

vein r5-m1, forming a very dierentiated anterior exten-

sion of the discal cell); M2 runs parallel to M1 and is of the

same length; m1-m2 runs between them at the end of the

cell, strongly V-shaped, with its angle pointing towards the

base of the wing, and in it a vestige of a recurrent veinlet

within the cell; M3 divides equidistantly the anterior and

posterior portions of the wing, runs approximately paral-

lel to M2, between them m2-m3 is straight and approxi-

Figure 1. Wing venation of a male individual of Fernanmorana

insignis (Butler, 1867) comb. nov., type species of its genus (av-

erage forewing length, from base to apex: 23.8 mm, n=7). Ven-

ezuela, Amazonas, Mcpio. Guainía, camino Yavita-Maroa (wing

prep. ALV062-19).

New genus of butterflies from the Amazon region

Figure 2. Genitalia of a male individual of Fernanmorana insignis (Butler, 1867) comb. nov., type species of its genus. Venezuela,

Amazonas, Mcpio. Guainía, camino Yavita-Maroa (wing prep. ALV599-14). Upper, dorsoventral view of genital armature (without

aedeagus); lower, lateral view of genital armature (without aedeagus)n; below dorsal (upper) and lateral (lower) wiews of aedeagus.

Illustrations produced at 40x magnication, horizontal line below drawings represents 1 mm.

Viloria & Costa

mately four times longer than r5-m1; Cu1, almost straight,

parallel to M3, emerges in the distal quarter of the discal

cell; Cu2, straight, parallel to Cu1, emerges approximately

two-hs of the cell length, measured from its base, as

long as two-thirds of the distance from the base to the

wing apex; A2, independent, slightly thickened at its basal

sixth, runs parallel to the anal margin of the wing. e

origin of the cubital veins is greatly inated, between the

base of the wing and halfway between this and the origin

of Cu2 (but only half the thickness of the inated base of

the Sc). Hindwing: Hu present and simple, without appar-

ent bifurcations, well developed on the anterior basal lobe

of the wing base, curved distally; Sc+R1 well developed

from the basal quarter of the anterior limit of discal cell,

curved, ending on the anterior margin at a point equidis-

tant between the wing base and the distal end of Rs; discal

cell subtriangular; Rs emerging from the anterior angle of

the cell, curved and distinct, ending at the rounded point

of the wing apex; M1 emerging posterior to the anterior

angle of the cell, not far from the origin of Rs, runs almost

straight to the outer margin of the wing; two slightly con-

vex transverse veins close the discal cell distally: m1-m2

and m2-m3, both approximately the same length; between

them emerges M2, which measures approximately half the

length between the base and the anal angle of the wing;

independent M3, shorter and more curved than M2, arises

at the most distal point of the cell; a little further towards

the base emerges Cu1, straight; m2-m3 and m3-cu1 form

a 90° angle; independent Cu2, a little longer than half the

length of the wing, arises in the nal third of the posterior

vein of the cell; A2 runs completely independently from

the base of the wing to the anal angle; independent A3,

Figure 3. Habitus of male (a dorsal, b ventral) and female (c dorsal, d ventral) individuals of Fernanmorana insignis (Butler, 1867)

comb. nov., type species of its genus. Venezuela, Amazonas, Raudal del Danto, 150 m.

New genus of butterflies from the Amazon region

from the base of the wing to its anal margin, ending at a

basal third of its length.

Male genitalia (Fig. 2). Armature well developed and

stylized; tegumen small and distinctly dome-like, very

well distinguished from the uncus, which is one and a half

times wider than the tegumen at its base, but also much

depressed dorsoventrally at that level, then immediately

compressed laterally, with a prominent crest (“inated at

its base in lateral view” sensu Espeland et al. 2023). Uncus

is also twice as long as tegumen, lanceolate, and ended in

a tip curved downwards; subunci with long, stylized bra-

chia, reaching beyond half the length of the uncus; saccus

thin, attened, as long as the brachia of subunci; valves

sub-rectangular, very long, longer than length of tegumen

+ uncus; aedeagus attened, wide, especially at distal ex-

treme, basal extreme curved downwards.

Etymology: e name Fernanmorana, feminine, comes

from the shortening, modication and arbitrary combina-

Figure 5. View of a trail used by the Piaroa indigenous people of the Cuao River region in Venezuela, inhabited by Fernanmorana

insignis (Butler, 1867) comb. nov.

Figure 4. Habitat of Fernanmorana insignis (Butler, 1867) comb. nov., type species of its genus. Venezuela, Amazonas, Raudal del

Danto, 150 m.

Viloria & Costa

tion of the Spanish surnames Fernández and Morán, and

is proposed here as a tribute to the distinguished Ven-

ezuelan physician, scientic researcher, and technologist,

Humberto Avelino Fernández-Morán (Maracaibo, 1924 –

Stockholm, 1999). His work and legacy had a global reach

and have been of paramount importance to the develop-

ment of modern science in Venezuela (Matos Romero

1986, Jiménez Maggiolo 1998, Rivas Cols 2005, Hernán-

dez Fonseca & Valbuena 2008, Requena 2011, Esparza &

Padrón 2020, Carvalho Kassar 2025, Molina Vílchez &

García Tamayo, in press). In his late years, H. Fernández-

Morán developed a keen interest in the preservation of the

Amazon river basin and its natural resources.

Distribution: Locally known in the lowlands of the

Amazon region, from the Tocantins river area (type speci-

men) to the Amazonas state in southern Venezuela (ma-

terial herein examined from Maroa-Yavita, Raudal del

Danto, Cuao river, but also Siapa region [R. Mattei, pers.

comm.]). It is probably widely distributed across the Ama-

zon basin.

Bionomic data: e type species of Fernanmorana

gen. nov. Fermanmorana insignis (Butler) new. comb.,

is a natural dweller of the tropical forests of the Amazon

region lowlands. It has been observed to be at least sea-

sonally abundant in some Venezuelan localities, such as

Raudal del Danto, in the río Cuao region (H. Camico,

pers. comm.) and Siapa, río Negro region (R. Mattei, pers.

comm.), mostly associated to thick undergrowth second-

ary vegetation (Figs. 4 &5) and ying together with other

satyrines among thick intrincate bamboo aggregations

(R.Mattei observations).

DISCUSSION AND CONCLUSION

Lamas (2004), Marín Uribe et al. (2019), Nakahara et

al. (2019) and Espeland et al. (2023) considered classifying

Euptychia insignis Butler and Euptychia ordinata Weymer,

1911, together within the same genus. We have had no ac-

cess to specimens of E. ordinata, originally described from

Bolivia, to test this hypothesis. Although both taxa look

supercially similar, we noticed in the photograph of the

type specimen of E. ordinata, once available publicly in the

Butteries of America website, that its ocelli bear a very

excentric silvery pupilla. In our experience this character

trend is not irrelevant for the generic classication of saty-

rine butteries, therefore, we declined to classify E. ordi-

nata under Fernanmorana gen. nov. Recently, two super-

cially identical Amazonian satyrine species, Stephenympha

pauliana Viloria (2022) and Xikrin ueharapradoi Freitas

& Barbosa (in Barbosa et al. 2023) were described in two

dierent and very distinct genera.

Espeland et al. (2023) demonstrated and asserted that

E. ordinata belongs in their Amphidecta-Clade, estab-

lishing not only some morphological likeness with some

members of Amphidecta Butler, 1867, but also their ge-

netic closeness.

Species belonging to Fernanmorana, gen. nov.:

Fernanmorana insignis (Butler, 1867), comb. nov.

Euptychia insignis Butler, 1867: 501, pl. 40, g. 12.

Euptychia insignis Butler; Butler, 1868: 38; Kirby,

1871: 56; Weymer, 1911: 213; Gaede, 1931: 451;

D’Abrera, 1988: 785 [gs. row 4, male habitus

dorsal, ventral]; Lamas, 2004: 223; Freitas et al.,

2018: 156; Marín Uribe et al., 2019: 97 [as incer-

tae sedis, Barbosa et al., in prep.]; Nakahara et al.,

2019: 14, 18; Espeland et al., 2023: 23, 59 [as a

new genus, Nakahara et al., in prep.]

[incertae sedis] insignis Butler; Lamas, 2004: 223.

‘Euptychia’ insignis Butler; Espeland et al., 2023:

13,14, 25.

DISCLAIMER

e authors declare no conicts of interest among

themselves. ALV was responsible for the conceptualiza-

tion, methodology, writing, and line drawings of this arti-

cle. MC and ALV jointly provided the resources, and MC

is the author of the photographic illustrations.

ACKNOWLEDGEMENTS

We thank Jesús Camacho (†), Anthony Heenan and

Hernando Camico for collecting the entomological ma-

terial we were able to examine for this work. Roberto

Mattei provided important bionomic information about

F. insignis and its occurrence in Siapa, Amazonas, Vene-

zuela. Ionesco Troconis, Gilson Rivas and María Eugenia

Andara assisted in processing images, reading and editing

this article.

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