Á. L. Viloria & M. Costa
18
ANARTIA
Publicación del Museo de Biología de la Universidad del Zulia
ISSN 1315-642X (impresa) / ISSN 2665-0347 (digital)
Anartia, 34 (junio 2022): 18-27
http://zoobank.org/urn:lsid:zoobank.org:pub:5902C23E-7DFB-4826-BDB8-953DAF32033B
A new subspecies of Oxeoschistus puerta (Westwood, 1851),
from the Sierra de Perijá, western Venezuela
(Lepidoptera: Nymphalidae: Satyrinae)
Una nueva subespecie de Oxeoschistus puerta (Westwood, 1851), de la
Sierra de Perijá, Venezuela occidental (Lepidoptera: Nymphalidae: Satyrinae)
Ángel L. Viloria1 & Mauro Costa2
1Centro de Ecología, Instituto Venezolano de Inestigaciones Cientícas (IVIC), km 11 carretera Panamericana, Altos de Pipe,
estado Miranda 1204, Venezuela.
2Residencias Las Cumbres, avenida Las Acacias, La Florida, Caracas 1050, Distrito Capital, Venezuela.
Correspondence: sebastianiloriacarrizo@gmail.com; aviloria@ivic.gob.ve
(Received: 02-11-2021 / Accepted: 15-02-2022 / On line: 30-09-2022)
ABSTRACT
Oxeoschistus puerta sabinoi ssp. nov., a pronophiline buttery (Lepidoptera: Nymphalidae: Satyrinae) from the montane
and upper montane forests (1,100-1,900 m a.s.l.) of the Venezuelan slope of the Sierra de Perijá, Zulia state, Venezuela, is
described, illustrated and compared with its close congeners. e taxon O. puerta magnus Pyrcz & Viloria, 2007, from the
Serranía del Tamá, is revised and newly identied as Oxeoschistus simplex magnus comb. nov.
Keywords: Northern Andes, Lake Maracaibo basin, Pronophilina, Satyrini.
RESUMEN
Se describe, ilustra y compara Oxeoschistus puerta sabinoi ssp. nov., una mariposa pronolina (Lepidoptera: Nymphalidae:
Satyrinae) procedente de los bosques montanos y altimontanos (1.100-1.900 m s.n.m.) de la vertiente venezolana de la Sie-
rra de Perijá, estado Zulia, Venezuela. Se revisa el taxón O. puerta magnus Pyrcz & Viloria, 2007, de la Serranía del Tamá, y
se reidentica como Oxeoschistus simplex magnus comb. nov.
Palabras clave: Andes septentrionales, cuenca del Lago de Maracaibo, Pronophilina, Satyrini,
acteristic to the Paramo Province of the South American
transition zone.
e study of the buttery fauna of the Perijá Moun-
tains began in the late 19th Century (Godman & Salvin
1880). Viloria (1990b, 1997) compiled a number of bib-
liographic sources with references to a few entomological
expeditions to this range and the adjacent lowlands. ese
works also included the citation of systematic revisions of
some Neotropical buttery genera with mention to species
and subspecies from Perijá and papers with descriptions
of several of its endemic taxa. Starting in the late 1970s,
INTRODUCTION
e Sierra de Perijá is a mountain range at the north-
ernmost tip of the Andes. Its main ridge (maximum eleva-
tion ca. 3,670 m a.s.l.) divides the Magdalena River (Co-
lombia) and the Lake Maracaibo (Venezuela) basins. So,
it harbors slightly dierent biological elements on each
slope, which according to Morrone et al. (2022), belong to
the adjacent biogeographical Provinces of Magdalena and
Guajira (Pacic Dominion), as well as higher elevation
Andean elements, part of which are endemics and char-
New subspecies of Oxeoschistus puerta
19
interest in the butteries of this region increased remark-
ably, and a number of taxonomic and faunistic studies
have been accomplished to improve the knowledge of the
regional Papilionoidea, most notably the Nymphalidae
(Adams 1977, 1984, 1985, Adams & Bernard 1979 [Co-
lombian slope], Viloria 1989, 1990a, b, 1991a, b, c, 1992,
1994, 1995, 2000, 2005, [2006], Pineda 1991, Viloria et
al. 1992, 2008, Neild 1996, 2008 [Venezuelan slope], Pu-
lido, 2007, Pulido & Andrade 2007, 2008, 2010, Andrade
2008, Pulido et al. 2010, 2011 [Colombian slope]).
Among the nymphalid butteries of the Sierra de Perijá,
the members of the Satyrinae have been particularly well
studied, especially the montane and high Andean members
of the subtribe Pronophilina, whose inventory was rst
achieved by Adams & Bernard (1979), followed by Viloria
(1990b) and Pulido & Andrade (2007, 2010). Only Vilo-
ria (op. cit.) detected and reported the presence of one spe-
cies of Oxeoschistus Butler, 1867 in this mountain range.
e genus Oxeoschistus was erected by Butler (1867).
He dened it by having forewings elongated and subfal-
cate with outer margin moderately wavy (minimally sinu-
ous) and notably wavy in hindwings; labial palpi long and
wavy. One diagnostic character was also the “forewing
cell bifurcated at the apex”, that is to say, divided by the
presence of a prominent recurrent vein entering the discal
cell, being the “lower fork” greater and arched, with three
angles, and the “upper fork” pointed and originating the
ve radial veins and M1 (Butler 1867: 267, [g.] 2, but
see Figure 3 of the present work). e type species of the
genus, by original designation, is Pronophila puerta West-
wood, 1851, a taxon endemic to the middle elevations of
the Cordillera de la Costa in northern Venezuela.
Forty years later, the genus was revised by ieme
(1907) and more recently by Pyrcz et al. (2020) (in part).
Judging from its morphological anities – palpi, legs, wing
pattern and venation, genitalic structure – (Reuter 1896,
Miller 1968, Adams 1986, Viloria 2007, Pyrcz 2010), and
genetic (molecular) relationships (Matz & Brower 2016),
Oxeoschistus belongs in the tribe Satyrini Boisduval, 1833,
subtribe Pronophilina Reuter, 1896, and should be placed
close to its type genus, Pronophila Doubleday, 1849.
Modern taxonomy of the species of Oxeoschistus has
been the subject of two arrangements (Lamas et al. 2004,
Pyrcz 2000). However, its species have been catalogued
or listed with synonymy by several authors since the 19th
Century (Kirby 1871, ieme 1907, Weymer 1911, Gaede
1931, D’Abrera 1988, Lamas et al. 2004, Pyrcz 2010, Pyrcz
et al. 2020).
e most recently described taxa are from Peru (O.
puerta garleppi Lamas 2003, O. iphigenia Pyrcz 2004),
Venezuela (O. romeo Pyrcz & Fratello 2005, O. puerta
magnus Pyrcz & Viloria 2007), Colombia (O. simplex tri-
plex Pyrcz & Salazar in Pyrcz & Rodríguez 2007), Costa
Rica and Honduras (respectively, O. tauropolis mitsuko
Pyrcz & Nakamura and O. hilara lempira Pyrcz in Pyrcz
et al. 2020).
e species of Oxeoschistus known to Venezuela have
been described, illustrated, reported or cited by West-
wood (1851), Raymond (1982), Viloria (1990b), Fratello
(2004), Pyrcz & Fratello (2005), Pyrcz & Viloria (2007),
Viloria et al. (2010), Viloria & Costa (2019) and Orellana
et al. (2019) (for a conclusive identication of the taxa
present in Venezuela, see discussion). In this paper we de-
scribe a new subspecies of Oxeoschistus puerta, which cor-
responds with the taxon referred to by Viloria (1990b) as
Oxeoschistus sp., and Oxeoschistus puerta ssp. nov. (Viloria
2000, Lamas et al. 2004).
MATERIAL AND METHODS
e butteries studied for this report were collected
in the Sierra de Perijá, western Venezuela in three dier-
ent occasions: December 1950, by Francisco Fernández
Yépez, October 1989, by Ángel L. Viloria and April 2011,
by Mauro Costa. It was rst detected as a new taxon dur-
ing the studies conducted by Viloria in 1988-1990 for his
rst degree thesis (Viloria 1990b). us, the specimens of
the type series have been examined several times ever since.
We have used standard techniques of observation and cap-
turing the adult insects in the eld (hand nets), ordinary
methods of setting the specimens, pinned for dry preser-
vation in entomological cabinets. e right wings of one
male specimen were removed, diaphanized by immersion
in diluted commercial bleach 1% for approximately 10
minutes, then profusely washed in distilled water, trans-
ferred subsequently to ethanol 70% for 15 minutes, and
then to absolute ethanol. ey were immediately xed and
permanently preserved in euparal between a large glass mi-
croscope slide (76 × 51 mm) and its cover (64 × 50 mm).
One male abdomen was removed and soaked overnight
in KOH 10% solution to digest and remove esh and fat
from the chitinous structures. Genitalic structures were
mechanically removed, transferred to distilled water for
8 hh and cleaned with microdissecting tools under a ste-
reoscopic microscope. Aer study, these microstructures
were stored in plastic vials with a solution of 70% ethanol
and 30% glycerine. Slides and vials have been externally
labelled and catalogued. Both wing veins and genitalic
structures were drawn with the aid of a camera lucida. Ob-
serving the specimens of the type series with and without
magnication, and their dissected organs and structures,
allowed for comparisons with other taxa (actual specimens
Á. L. Viloria & M. Costa
20
preserved, photographs and drawings), which eventually
led to the diagnosis and description of the new taxon.
Our criteria to establish the classication of the new
taxon were the stable dierences of some features of its
wing color and design patterns, as well as the lack of sig-
nicant variation of its wing venation and male genita-
lia, when compared to typical Oxeoschistus puerta puerta
(Westwood, 1851). However, we also took into account
the geographic distribution of the several species and sub-
species of Oxeoschistus to evaluate spatial patterns, both
horizontal (latitudinal) and perpendicular (altitudinal)
and consider the more plausible hypothesis to justify the
new geographic race of Oxeoschistus puerta as a subspecies.
e following acronyms and abbreviations are used in
this text: FW: Forewing; FWL: Forewing length; HW:
Hindwing; MBLUZ: Museo de Biología de la Universi-
dad del Zulia, Facultad Experimental de Ciencias, Mara-
caibo, Venezuela; MC: Mauro Costa collection, Cara-
cas, Venezuela; MIZA: Museo del Instituto de Zoología
Agrícola, Facultad de Agronomía, Universidad Central de
Venezuela, Maracay.
RESULTS
Oxeoschistus puerta sabinoi Viloria & Costa, ssp. nov.
(Figs. 1a, b, c, d, 3, 4)
http://zoobank.org/urn:lsid:zoobank.org:act:4E01D3CF-5820-4F99-AE67-
0B775AF9AAEB
Oxeoschistus sp.; Viloria, 1990b: 244–246, 269, 270,
271, gs. 157 (male habitus, dorsal), 158 (male habi-
tus, ventral), 159 (male genitalia)(MBLUZ-0557).
Oxeoschistus puerta ssp. nov. Viloria, MS; Viloria, 2000:
268.
Oxeoschistus puerta [n. ssp.] Viloria, MS; Lamas et al.,
2004: 211.
Figure 1. Habitus of Oxeoschistus puerta sabinoi ssp. nov. a. Holotype male, dorsal view; b. Same, ventral view; c. Allotype female,
dorsal view; d. Same, ventral view.
New subspecies of Oxeoschistus puerta
21
Figure 2. Habitus of two species of Oxeoschistus Butler from Venezuela. a. Male, dorsal view O. puerta puerta (Westwood); b. Same,
ventral view; c. Female, dorsal view O. puerta puerta; d. Same, ventral view (individuals from Colonia Tovar, 1,650m, Aragua State,
Venezuela); e. Male, dorsal view O. simplex magnus Pyrcz & Viloria, comb. nov.; f. Same, ventral view (individual from Matamula,
1,800m, Táchira State, Venezuela).
Á. L. Viloria & M. Costa
22
Type material: H ♂, Ven.[ezuela], [Sierra de]
Perijá, 10° 19’ 35” N, 72° 35’ 27” W, 1,900 m, 06-IV-2011,
[M. Costa] [MIZA]; A ♀, same data as holo-
type; P: 1 ♂, same data as holotype (wing prep.
ALV062-19)[MC]; 1 ♂, [Venezuela, Sierra de Perijá], SM,
Kunana, [10° 03’ 08” N, 72° 47’48” W, 1,127 m], 30-XII-
[19]50, Expedición La Salle [MIZA]; 1 ♂, Venezuela, Si-
erra de Perijá, SM, Kunana, [10° 03’ 08” N, 72° 47’48” W,
1,127 m], 24-X-1989, Á. L. Viloria leg. (MBLUZ-0557,
genit. prep. ALV-s/n) [MBLUZ].
Diagnosis: Oxeoschistus puerta sabinoi ssp. nov., from
the Sierra de Perijá, diers from the typical subspecies O.
puerta puerta (Westwood, 1851), from the Cordillera de
la Costa of Venezuela, in the following characters: Male.
FW upperside postdiscal dark brown dot on M3 absent;
underside postdiscal brick-orange patch on M1 darker;
postdiscal patch on M2 slightly wider on both sides, as
well as that of M1; postdiscal patch on R5 basally longer;
postdiscal patch on R3 more developed; FW underside
ground color with more distinct borders, darker on the
basal two thirds of the wing, as well as in submarginal and
marginal areas. HW upperside postdiscal dark brown dots
less distinct, especially on M3; a pair of minute dots on
Cu2. HW underside ocelli in R5 and M1 brown instead
of black, that on M3 black but reduced. Female. HW up-
perside lacking postdiscal brown dots on M1 and M2, that
on M3 and a pair on Cu2 notably reduced; dot on Cu1 re-
duced, with a white pupil. HW underside postdiscal ocelli
in R5, M1, M2 and M3 lacking black.
Description: Male (Figs. 1a, b, 3, 4). FWL: 34-35
mm (mean 34.5 mm). Body dorsally dark coee, almost
black, ventrally light brown, legs light brown. Antennae
dark coee, reaching two hs of costa, club gradually
formed. Palpi twice as long as head, dorsally and later-
ally cream-white, ventrally densely covered with long
black scales. FW subtriangular, apex and tornus rounded,
outer margin moderately scalloped, pronounced at dis-
tal end of vein M1. HW suboval, costal margin basally
straight to two thirds of its length, outer margin mark-
edly scalloped. FW upperside ground color coee brown,
postdiscal area occupied by a brick-orange band formed
by contiguous patches (divided by crossing coee brown
veins); three postdiscal circular dark brown spots on M1,
Cu1 and Cu2, respectively, Cu1 twice the diameter of
the others. HW upperside ground color coee brown; a
wide brick-orange postdiscal-submarginal band; a suu-
sion of the same color on marginal area from vein M3 to
tornus, nely bordered with dark coee along outer mar-
gin; over the band one vestigial, very diused dark spot
on M1, another one more visible on M3, another one
more distinct on Cu1, with tiny white pupil; two small
Figure 3. Wing venation of Oxeoschistus puerta sabinoi ssp. nov.,
male paratype (wing prep. ALV062-19, scale bar: 1 cm).
Figure 4. Male genitalia of Oxeoschistus puerta sabinoi ssp.
nov. (Paratype, MBLUZ-0557, genit. prep. ALV-s/n, scale bar:
1mm). Re-drawn from Viloria (1990b).
New subspecies of Oxeoschistus puerta
23
anal dark spots on cell Cu2. Basal two thirds of FW un-
derside ground color hazelnut, chocolate brown towards
anal margin, coee brown towards anterior half of discal
area, markedly over recurrent vein inside discal cell; costal
line light ochraceous, more visible along two basal thirds;
veins dark brown; brick-orange postdiscal band running
only from vein M2 to tornus, progressively widening in
that direction, twice as broad in tornus as in cell M2; two
ocellar postdiscal dark brown dots on M1(with tiny pu-
pil) and Cu1, respectively; subapical white dot (2 mm
diameter) on R5, contiguous to another one smaller in
R4, just in branching of veins R4 and R5; distal to these
dots an area of lighter pale brown that reaches the costa; a
cream-white postdiscal wedge from costa to anterior base
of cell M1. HW underside ground color hazelnut dusted
with ochraceous brown on basal, anal and distal half of
marginal areas; cream-white stripe, running from costa on
limit of discal and postdiscal area, across the middle of
discal cell, more or less straight to vein A2; on the costa
it reaches the anterior extremity of the wide cream-white
postdiscal band, which runs from apex to tornus; its in-
ner margin more or less regular, slightly sigmoidal, con-
cave from costal margin to vein M3 and slightly convex
from there to tornus; its distal margin dentate, pointing
distad on apex (vein Rs), veins M1 and M3; from M3
to Cu1 and Cu2 the shape of that margin is more or less
scalloped but not regular; a series of seven round ocelli in
the middle of the postdiscal band, four ocelli of similar
size and aspect in cells R5, M1, M2 and M3, respectively,
with dark hazelnut centers and white pupils (R5 and M3
with some black around pupils), widely surrounded by
yellow-orange and nely circled with hazelnut; one larger
ocellus with black center and larger pupil on Cu1; two
small, twin, black ocelli on Cu2 (in tornus), the last three
ocelli surrounded with a diuse mixture of hazelnut and
yellow-orange.
Genitalia. Tegumen attened; uncus conical and
slightly curved downwards, as long as tegumen, emerging
at same level of the latter; subunci well developed, as long
as uncus, but emerging laterally and just behind its base;
vinculum strong but thin; saccus tubular, slightly shorter
than subuncus; Valvae subrectangular in lateral view, 1.3
times the length of tegumen + uncus, distal extremity al-
most as deep as base, with some rugosity around its distal
extremity, more prominent on ampulla. Aedeagus straight
and robust, slightly thicker than saccus, and slightly longer
than valvae.
In general the male genitalia of this subspecies does not
dier signicantly from that of O. puerta puerta.
Female (Figs. 1c, d). FWL: 36 mm. Similar to male
but slightly larger and paler in coloration. FW upperside
with prominent postdiscal-subapical wedge on costal
area (as on underside); postdiscal dark dots larger, one
in cell M3.
Etymology: e name of this new taxon is dedicated to
the memory of Sabino Romero Izarra (b. Sierra de Perijá,
Venezuela, February 9, 1965), uatpú (chief ) of the indig-
enous community of Chaktapa, in the Yasa river basin,
southern Venezuelan slope of the Sierra de Perijá. In the
present century, Sabino was so far the most outstanding
defender in the struggle of the Yukpa ethnic group for the
recovery of their original lands. He was murdered in Los
Angeles del Tukuko on March 3, 2013.
Habitat and behaviour: e dierent species and sub-
species of Oxeoschistus known in Venezuela (see discus-
sion) have similar habits. ey are insects associated with
montane and upper montane forests, recorded between
1,000 and 2,000 m. However, only the rst two known in-
dividuals of O. puerta sabinoi ssp. nov., come from a bam-
boo forest at around 1,100 m on the trail that connects the
Yukpa populations of Ayajpaina and Kunana in the central
sector of the Sierra de Perijá, on its Venezuelan slope. At
this altitude the presence of the species is so rare that it
could be considered occasional. However, we have noticed
that although more frequently found above 1,500 m, they
are always quite localized in dense forested sectors, with
the presence of bamboo (Guadua spp. and Chusquea spp.),
presumably their host plants. ey seldom y in clearings
or open spaces. ey do not y if there is not enough sun-
light entering the forest; their ight is erratic and can be
remarkably fast when the animals are startled. at is why
they are dicult to catch on the wing. ey prefer to move
between 1 and 3 meters from the ground.
Females and males of Oxeoschistus in the Tamá, Perijá,
Cordillera de la Costa and Pantepui are attracted by fer-
mented plantain baits, whose sugary uids are among their
favorite foods. In this way, once their presence is detected,
it is easier to observe and capture them when feeding on
these fruits (gs. 5–7).
Pyrcz et al. (2020) commented on behavioral trends,
habitat and distribution of the species of Oxeoschistus (s. l.)
in Central America.
DISCUSSION
Adams (1986) pointed out the presence of distinctive
yellow-orange rings (iris) in the ocelli of the hindwing
underside of three north Andean taxa of this genus (sensu
stricto). He also oered morphological criteria and biogeo-
graphic reasons to consider each of them a separate species:
O. puerta (Westwood), O. protogenia (Hewitson, 1862)
and O. pervius ieme, 1907. We agree with this point of
Á. L. Viloria & M. Costa
24
Figure 5. Adult female Oxeoschistus puerta puerta (Westwood), resting on a palm leave. Near Colonia Tovar, 1,650 m, Aragua State,
Cordillera de La Costa, northern Venezuela (photo M. Costa).
Figure 6. Adult female Oxeoschistus puerta puerta (Westwood) baited with rotten fruit (plantain). Neighbouring forest of the Colonia
Tovar, 1,650 m, Aragua State, Cordillera de La Costa, northern Venezuela (photo M. Costa).
New subspecies of Oxeoschistus puerta
25
view. Our new taxon bears those yellow-orange rings and
is morphologically (wing pattern, male genitalia) and bio-
geographically closer to typical O. puerta from the Cor-
dillera de La Costa, therefore we comparatively consider
sabinoi ssp. nov., a subspecies of puerta. Both Oxeoschistus
protogenia and O. simplex Butler, 1868 y parapatrically,
the rst below (800-1,000 m) the second one (1,800-
2,350 m), in the Cordillera Oriental of Colombia. Oxeos-
chistus simplex belongs to a group of species that lacks the
presence of the above mentioned yellow-orange rings. e
taxon magnus, described by Pyrcz & Viloria (2007), clear-
ly lacking the yellow-ringed ocelli, being but slightly dier-
ent from typical O. simplex, and distributed in the barely
disjunct northernmost extreme of the Cordillera Oriental
(El Tamá), should be regarded as a subspecies of the latter
taxon. We herein establish this new combination.
us, there are three dierent species of Oxeoschistus
recorded in Venezuela, one with two subspecies:
Oxeoschistus puerta puerta (Westwood, 1851) (Figs. 4a, b,
c, d, 5, 6). Type locality: Venezuela [Cordillera de La
Costa].
Oxeoschistus puerta sabinoi Viloria & Costa, ssp. nov.
(Figs. 1a, b, c, d, 2, 3). Type locality: Venezuela, Sierra
de Perijá, 10° 19’ 35” N, 72° 35’ 27” W, 1,900 m.
Oxeoschistus simplex magnus Pyrcz & Viloria, 2007, comb.
nov. (Figs. 4e, f ). Type locality: Venezuela, Táchira,
Delicias, 1,000 m.
Oxeoschistus romeo Pyrcz & Fratello, 2005 (Fig. 7). Type
locality: Guyana, N. slope of Mt. Roraima, 2nd Camp,
5° 16’ N, 60° 44’ W, 1,300 m.
ACKNOWLEDGEMENTS
We thank Tito R. Barros, Nelson Delgado (†), Leonel
Lanier, Jesús Camacho, Tomasz W. Pyrcz, Florencio Ber-
nal, Guillermo Morales, Rafael Carreño and Lusbi Por-
tillo for their companionship and assistance during eld
activities associated with the discovery of the new subspe-
cies herein described (1989 and 2011). Professors Francis
Geraud, Magally uiróz (†), Ramón Acosta (†) and José
Moscó (†) (La Universidad del Zulia), helped the rst au-
thor in many ways during the early days of surveying the
satyrine butteries of the Sierra de Perijá (1984-1993).
Jürg De Marmels, Luis D. Otero, José A. Clavijo and Car-
los Bordón (†) are warmly thanked for their hospitality
and assistance at MIZA. Héctor Suárez (IVIC) took the
photographs of the type material. e rst dras of this
article were critically read by I. B. Petit, M. uiróz, F. Ger-
aud and Gilson Rivas. e latter, together with Neva Mora
and María Eugenia Andara were responsible for the nal
editing of text and gures.
REFERENCES
Adams, M. J. 1977. Trapped in a Colombian Sierra. Geographi-
cal Magazine 49(4): 250–254.
Adams, M. J. 1984. Northern Andean butteries - Search and
research. e Alpine Journal 1984: 90–96.
Adams, M. J. 1985. Speciation in the pronophiline butteries
(Satyridae) of the northern Andes. Journal of Research on the
Lepidoptera Supplement 1: 33–49.
Adams, M. J. 1986. Pronophiline butteries (Satyridae) of the
three Andean Cordilleras of Colombia. Zoological Journal of
the Linnean Society 87(3): 235–320.
Adams, M. J. & G. I Bernard. 1979. Pronophiline butteries (Sa-
tyridae) of the Serranía de Valledupar, Colombia-Venezuela
border. Systematic Entomology 4(2): 95–118.
Andrade, M. G. 2008. Descubierta nueva especie de mariposa.
El Tiempo (Bogotá), 9 marzo 2008, pp. 14.
Butler, A. G. 1867. Revision of the group of lepidopterous
insects hitherto included in the genus Pronophila of West-
wood. Annals and Magazine of Natural History (3)20(118):
266–268.
Figure 7. ree adult individuals of Oxeoschistus romeo Pyrcz &
Fratello, an endemic to the Pantepui, feeding on rotting plan-
tain. Sierra de Lema, 1,400 m, Bolívar State, southeastern Vene-
zuela (photo M. Costa).
Á. L. Viloria & M. Costa
26
D’Abrera, B. 1988. Butteries of the Neotropical Region. Part V.
Nymphalidae (Conc.) & Satyridae. Victoria, Black Rock:
Hill House, [viii] + pp. 679–877.
Fratello, S. A. 2004. New species from Mt. Roraima. News of the
Lepidopterists’ Society 46(4): 122–126.
Gaede, M. 1931. Familia Satyridae. In: Strand, E. (ed.). Lepidop-
terorum Catalogus 43: 1−320; 46: 321–544; 48: 545−759.
Godman, F. D. C. & O. Salvin. 1880. A list of diurnal Lepidop-
tera collected in the Sierra Nevada of Santa Marta, Colom-
bia, and the vicinity. Transactions of the Entomological Society
of London 1880(3): 119–132, pls. 3–4.
Kirby, W. F. 1871. A synonymic catalogue of diurnal Lepidoptera.
London: John van Voorst, vii + 690 pp.
Lamas, G. 2003. Las Mariposas de Machu Picchu. Guía ilustrada
de las mariposas del Santuario Histórico Machu Picchu, Cuzco,
Perú. Lima: PROFONANPE, [vi] + 221 pp., 34 pls.
Lamas, G., Á. L. Viloria & T. W. Pyrcz. 2004. Nymphalidae.
Satyrinae. Tribe Satyrini. Subtribe Pronophilina. pp. 206–
215. In: Lamas, G. (ed.): Checklist: Part 4A. Hesperioidea
– Papilionoidea. In: Heppner, J. B. (ed.): Atlas of Neotropical
Lepidoptera. Volume 5A. Gainesville: Association for Tropi-
cal Lepidoptera / Scientic Publishers.
Matz, J. & A. V. Z. Brower. 2016. e south temperate Prono-
philina (Lepidoptera: Nymphalidae: Satyrinae): a phyloge-
netic hypothesis, redescriptions and revisionary notes. Zoo-
taxa 4125(1):1–108.
Miller, L. D. 1968. e higher classication, phylogeny and
zoogeography of the Satyridae (Lepidoptera). Memoirs of the
American Entomological Society 24: [6] + iii + 174 pp.
Neild, A. F. E. 1996. e Butteries of Venezuela. Part 1: Nym-
phalidae I (Limenitidinae, Apaturinae, Charaxinae). A com-
prehensive guide to the identication of adult Nymphalidae,
Papilionidae, and Pieridae. Greenwich, London: Meridian
Publications, 144 pp., 32 pls.
Neild, A. F. E. 2008. e Butteries of Venezuela. Part 2:
Nymphalidae II (Acraeinae, Libytheinae, Nymphalinae,
Ithomiinae, Morphinae). A comprehensive guide to the iden-
tication of adult Nymphalidae, Papilionidae, and Pieridae.
Greenwich, London: Meridian Publications, 275 pp., 84 pls.
Orellana, A. M., J. M. González & Á. L. Viloria. 2019. Adicio-
nes, recticaciones y actualizaciones a “Mariposas de Vene-
zuela” por éophile Raymond. VI Satyrinae (Insecta: Lepi-
doptera: Nymphalidae). Anartia 29: 54–67.
Pineda, A. 1991. En la Sierra de Perijá la vida viaja en alas de
mariposas. Ciencias (Maracaibo) 7(49): 15.
Pulido-B., H. W. 2007. High altitudes [sic] butteries (Lepi-
doptera: Rhopalocera) of the Serranía de Perijá, Colombia.
II ELEN, Encuentro de Lepidopterología Neotropical. Ciudad
de Panamá, Panamá, [29 April – 3 May 2007], pp. [14–15]
[abstract].
Pulido-B., H. W. & M. G. Andrade-C. 2007. Mariposas de las
partes altas de la Serranía de Perijá. pp. 235–248. In: Rangel-
Ch., J. O. (ed.): Colombia, diversidad biótica V. La alta mon-
taña de la Serranía de Perijá. Bogotá: Universidad Nacional
de Colombia, Instituto de Ciencias Naturales.
Pulido-B., H. W. & M. G. Andrade-C. 2008. A new species of
Forsterinaria Gray, 1973 (Lepidoptera: Nymphalidae: Sa-
tyrinae) from the Serranía del Perijá, Cesar, Colombia. Cal-
dasia 30(1): 189–195.
Pulido-B., H. W. & M. G. Andrade-C. 2010. Las mariposas de
la Serranía de Perijá. pp. 509–559. In: Rangel-Ch., J. O. (ed.):
Colombia, diversidad biótica VIII. Media y baja montaña de
la Serranía de Perijá. Bogotá: Universidad Nacional de Co-
lombia, Instituto de Ciencias Naturales.
Pulido-B., H. W., M. G. Andrade-C. & J. Llorente-Bousquets.
2010. Taxonomía y distribución de Nathalis (Lepidoptera:
Pieridae) en Colombia. Revista de Biología Tropical (Interna-
tional Journal of Tropical Biology) 58(1): 273–285.
Pulido-B., H. W., M. G. Andrade-C., C. Peña & G. Lamas.
2011. Two new taxa of Euptychia Hübner, 1818 (Lepidop-
tera: Nymphalidae: Satyridae) from the Andes of Colombia
and Peru. Zootaxa 2906: 43–51.
Pyrcz, T. W. 2004. Pronophiline butteries of the highlands of
Chachapoyas in northern Peru: faunal survey, diversity and
distribution patterns (Lepidoptera, Nymphalidae, Satyri-
nae). Genus 15(4): 455–622.
Pyrcz, T. W. 2010. Wybrane zagadnienia z taksonomii, zoogeo-
grai I ewolucji faun górskich na przykładzie grupy modelowej
motyli z plemienia Pronophilini (Nymphalidae). Olsztyn:
Wydawnictwo Mantis, 245 pp. + [iii].
Pyrcz, T. W. & S. A. Fratello. 2005. Cloud forest buttery fauna
of the Pantepui - Poor or poorly known? Description of new
species and records of new genera of Pronophilina: Eretris
agata and Oxeoschistus romeo (Nymphalidae: Satyrinae).
Journal of the Lepidopterists’ Society 59(4): 200–211.
Pyrcz, T. W. & G. Rodríguez. 2007. Mariposas de la tribu
Pronophilini en la Cordillera Occidental de los Andes de
Colombia (Lepidoptera: Nymphalidae, Satyrinae). Shilap
35(140): 455–489.
Pyrcz, T. W. & Á. L. Viloria. 2007. Erebiine and pronophiline
butteries of the Serranía del Tamá, Venezuela-Colombia
border (Lepidoptera: Nymphalidae: Satyrinae). Tropical
Lepidoptera 15(1/2): 18–52.
Pyrcz, T. W., A. Zubek, P. Boyer, I. Nakamura, B. Wacławik &
K. Florczyk. 2020. Revisional notes on the cloud forest but-
tery genus Oxeoschistus Butler in Central America (Lepi-
doptera: Nymphalidae: Satyrinae). Neotropical Entomology
49(3): 392–411.
Raymond, T. 1982. Mariposas de Venezuela. [1st ed.]. Caracas:
Ediciones Corpoven / Armitano Editor, 277 pp. + [iii].
Reuter, E. 1896. Über die Palpen der Rhopalocera. Ein Beitrag
zur Erkenntnis der verwandtschalichen Beziehungen unter
den Tagfaltern. Acta Societatis Scientiarum Fennicae 22(1): i-
xvi + 577 pp. + [i], 6 pls.
ieme, O. 1907. Monographische bearbeitung der Gattungen
Lasiophila Felder, Daedalma Hew., Catargynnis Röber, Ox-
eoschistus Butler, Pronophila Westwood, Corades Doubl. Hew.
Berliner Entomologische Zeitschri 51(2/3): 99–234, pls. 3–5.
Viloria, Á. L. 1989. Estudio preliminar de la fauna de lepidóp-
teros de Perijá, Estado Zulia. Parte I. Nymphalidae: Itho-
New subspecies of Oxeoschistus puerta
27
miinae. XI Congreso Venezolano de Entomología. Maracaibo
(Venezuela), July 5-8, 1989, pp. 54 [abstract].
Viloria, Á. L. 1990a. Consideraciones sobre los niveles de ende-
mismo en mariposas (Lepidoptera: Rhopalocera) de Perijá,
frontera colombo-venezolana. XI Congreso Latinoamericano
de Zoología. Cartagena (Colombia), April 1990, pp. 68 [abs-
tract].
Viloria, Á. L. 1990b. Taxonomía y distribución de los Satyridae
(Lepidoptera: Rhopalocera) en la Sierra de Perijá, ontera
colombo-venezolana. Maracaibo: Facultad Experimental de
Ciencias, Universidad del Zulia, [ii] + xxxviii + 296 pp. +
[ii] [thesis].
Viloria, Á. L. 1991a. Catatumbo un centro de evolución, evi-
dencia de una mayor amplitud en base a la distribución geo-
gráca de algunos lepidópteros aposemáticos del oeste de
Venezuela. p. 37. V Jornadas Cientícas de la Facultad Expe-
rimental de Ciencias de la Universidad del Zulia. Maracaibo
(Venezuela), June 26-28, 1991 [abstract].
Viloria, Á. L. 1991b. Los páramos de Perijá. Natura (Caracas)
93: 25–29.
Viloria, Á. L. 1991c. Estudio preliminar de la fauna de lepi-
dópteros de Perijá, Colombia-Venezuela. Parte II. Satyridae.
p.215. XII Congreso Venezolano de Entomología. Mérida (Ve-
nezuela), July 1-4, 1991 [abstract].
Viloria, Á. L. 1992. Taxonomía y distribución de los Satyridae
(Lepidoptera: Rhopalocera) en la Sierra de Perijá, frontera
colombo-venezolana. pp. 199–200. In: Vicerrectorado Aca-
démico (ed.). Producción intelectual de LUZ, Facultad Expe-
rimental de Ciencias, años 1983-1991. Maracaibo: Univer-
sidad del Zulia, Biblioteca Central “Gral. Rafael Urdaneta”,
Impresos Chirinos.
Viloria, Á. L. 1994. Dos expediciones a los páramos de la Sierra
de Perijá, frontera colombo-venezolana, 1989. El Guácharo,
Boletín de Divulgación Espeleológica (Caracas) 35: 16–21.
Viloria, Á. L. 1995. Informe general de la expedición a Cerro
Pintao (Cerro Viruela), frontera colombo-venezolana. Mar-
zo 1993. El Guácharo, Boletín de Divulgación Espeleológica
(Caracas) 36: 61–64.
Viloria, Á. L. 1997. Fuentes para el estudio de la Sierra de Perijá.
Maracaibo: Universidad del Zulia, 176 pp.
Viloria, Á. L. 2000. Estado actual del conocimiento taxonómico
de las mariposas (Lepidoptera: Rhopalocera) de Venezuela.
pp. 261–274. In: Martín-Piera, F., J. J. Morrone & A. Melic
(eds.). Hacia un Proyecto CYTED para el inentario y esti-
mación de la diversidad entomológica en Ibero-américa: PrI-
BES-2000. m3m-Monografías Tercer Milenio, vol. 1. Zara-
goza: Sociedad Entomológica Aragonesa.
Viloria, Á. L. 2005. Las mariposas (Lepidoptera: Papilionoi-
dea) y la regionalización biogeográca de Venezuela. pp.
441–459. In: Llorente Bousquets, J. E. & J. J. Morrone (eds.).
Regionalización biogeográca en Iberoamérica y tópicos anes.
Primeras Jornadas Biogeográcas de la Red Iberoamericana
de Biogeografía y Entomología Sistemática (RIBES XII.I-
CYTED). México, D. F.: Las Prensas de Ciencias, Facultad
de Ciencias, UNAM.
Viloria, Á. L. [2006]. Cuatro noticias zoológicas de interés
bioespeleológico. Boletín de la Sociedad Venezolana de Espe-
leología 39: 83–87.
Viloria, Á. L. 2007. e Pronophilina: synopsis of their biol-
ogy and systematics (Lepidoptera: Nymphalidae: Satyrinae).
Tropical Lepidoptera 15(1-2): 1–17.
Viloria, Á. L. & M. Costa. 2019. Endemic butteries. pp. 193–
222. In: Rull, V., T. Vegas-Vilarrúbia, O. Huber & C. Señaris
(eds.). Biodiversity of Pantepui: e pristine “Lost World” of
the Neotropics. Cambridge, Massachussets: Elsevier.
Viloria, Á. L., F. Herrera & C. Galán. 1992. Resultados prelimi-
nares del estudio del material biológico colectado en Mesa
Turik y cuenca del Río Socuy. Boletín de la Sociedad Venezo-
lana de Espeleología 26: 7–9.
Viloria, Á. L., T. W. Pyrcz & A. Orellana. 2010. A survey of the
Neotropical montane butteries of the subtribe Pronophi-
lina (Lepidoptera, Nymphalidae) in the Venezuelan Cordil-
lera de la Costa. Zootaxa 2622: 1–41.
Viloria, Á. L., A. D. Warren & G. T. Austin. 2008. A spectacular
new Dalla Mabille, 1904 from Venezuela-Colombia (Hesperi-
idae: Heteropterinae). Bulletin of the Allyn Museum 156: 1–12.
Westwood, J. O. 1851. pp. 327–466, pls. 59, 67–77. In: Dou-
bleday, E.: e genera of diurnal Lepidoptera: comprising their
generic characters, a notice of their habits and transformations,
and a catalogue of the species of each genus, ol. 1. London:
Longman, Brown, Green & Longmans.
Weymer, G. 1911. 4. Familie: Satyridae. pp. 193–240, pls. 42–
50. In: Seitz, A. (ed.). Die Gross-Schmetterlinge der Erde, 5.
Stuttgart: A. Kernen.
