Taxonomic notes and descriptions of new satyrine butterflies from tropical America
7
ANARTIA
Publicación del Museo de Biología de la Universidad del Zulia
ISSN 1315-642X (impresa) / ISSN 2665-0347 (digital)
Anartia, 31 (diciembre 2020): 7-62
http://zoobank.org/ urn:lsid:zoobank.org:pub:A422293D-B23C-4C6F-A218-AF5969D39A8C
Taxonomic notes on Euptychia modesta Butler, 1867,
Neonympha alcinoe C. Felder & R. Felder, 1867
and Euptychia pamela Hayward, 1957
(Lepidoptera: Nymphalidae: Satyrinae), with descriptions
of three new genera, five new species and two new subspecies
from Central and South America
Notas taxonómicas sobre Euptychia modesta Butler, 1867,
Neonympha alcinoe C. Felder & R. Felder, 1867 y Euptychia pamela Hayward, 1957
(Lepidoptera: Nymphalidae: Satyrinae), con descripciones de tres géneros nuevos,
cinco especies y dos subespecies nuevas de Centroamérica y Suramérica
Mohamed Benmesbah
1
, Ángel L. Viloria
2,3
& Jérome Murienne
4
1
28T avenue des Pyrénées 31880 La Salvetat-Saint-Gilles, Francia.
2
Centro de Ecología, Instituto Venezolano de Investigaciones Científicas (IVIC), Apartado Postal 20632, Caracas 1020-A, Venezuela.
3
Simon Bolivar Professor, Centre of Latin American Studies, University of Cambridge, Alison Richard Building, 7 West Road,
Cambridge CB3 9DT, Reino Unido.
4
Laboratoire Évolution & Diversité Biologique (UMR5174) - Université de Toulouse Midi-Pyrénées, CNRS, IRD, UPS - 118,
route de Narbonne, Bât. 4R1, F-31062 Toulouse Cedex 9, Francia.
Correspondence: M. Benmesbah: modjojojo66@hotmail.com
(Received: 26-09-2020 / Accepted: 04-12-2020 / On line: 26-02-2021)
ABSTRACT
e identity, status and taxonomic position of seven species of Neotropical satyrine butteries are discussed and dened:
Euptychia modesta Butler, 1867, Euptychia nebulosa Butler, 1867, Neonympha alcinoe C. Felder & R. Felder, 1867, Eup-
tychia mimas Godman, 1905, Euptychia mimula Hayward, 1954, Euptychia pamela Hayward, 1957 and Cissia gomezi
Singer, DeVries & Ehrlich, 1983. ese taxa lately remained in genera such as Cissia Doubleday, 1848, Magneuptychia
Forster, 1964 and Yphthimoides Forster, 1964. Based on the particular combination of selected morphological characters
and comparative DNA analysis of these and other evidently related species, three new genera (with seven new combina-
tions), ve new species and two new subspecies are proposed, diagnosed and described: Modestia Viloria & Benmesbah,
gen. nov., with four taxa, M. modesta (Butler, 1867), comb. nov., M. gomezi (Singer, DeVries & Ehrlich, 1983), comb.
nov., M. remypignoux Benmesbah & Viloria, sp. nov. and M. remypignoux shueyi Benmesbah & Viloria, ssp. nov.; Ma-
laveria Viloria & Benmesbah, gen. nov., with nine taxa, M. alcinoe (C. Felder & R. Felder, 1867), comb. nov., M. nebu-
losa (Butler, 1867), comb. nov., M. mimas (Godman, 1905), comb. nov., M. ballo Benmesbah & Viloria, sp. nov., M.
rodriguezi Benmesbah & Viloria, sp. nov., M. rodriguezi risaralda Benmesbah & Viloria, ssp. nov.
, M. bottoi Benmesbah
& Viloria, sp. nov., M. duponti Benmesbah & Murienne, sp. nov. and M. mimula (Hayward, 1954), comb. nov.; and
Koutalina Viloria & Murienne, gen. nov., for a single known species, K. pamela (Hayward, 1957), comb. nov. Lectotypes
of Euptychia modesta, Neonympha alcinoe and Neonympha sylvina C. Felder & R. Felder, 1867, are designated. Variability
M. Benmesbah, Á. L. Viloria & J. Murienne
8
in morphology and aspect of the male genitalia is discussed, as well as number of pupils in ocelli and the occasional pres-
ence of supernumerary anal ocellus on ventral hindwing.
Key words: Argentina, Belize, Bolivia, Cissia, Colombia, Costa Rica, DNA, Ecuador, Euptychiina, French Guiana, Mag-
neuptychia, Panama, Paraguay, Peru, Venezuela, Yphthimoides.
RESUMEN
Se discuten y denen la identidad, el estatus y la posición taxonómica de siete especies de mariposas satirinas neotropicales:
Euptychia modesta Butler, 1867, Euptychia nebulosa Butler, 1867, Neonympha alcinoe C. Felder & R. Felder, 1867, Eup-
tychia mimas Godman, 1905, Euptychia mimula Hayward, 1954, Euptychia pamela Hayward, 1957 y Cissia gomezi Singer,
DeVries & Ehrlich, 1983. Estos taxones permanecían últimamente en géneros tales como Cissia Doubleday, 1848, Mag-
neuptychia Forster, 1964 e Yphthimoides Forster, 1964. Con base en la combinación particular de caracteres morfológicos
selectos y el análisis comparativo de ADN de éstas y otras especies evidentemente relacionadas, se proponen, diagnostican
y describen tres nuevos géneros (con siete combinaciones nuevas), cinco nuevas especies y dos nuevas subespecies: Modestia
Viloria & Benmesbah, gen. nov., con cuatro taxones, M. modesta (Butler, 1867), comb. nov., M. gomezi (Singer, DeVries
& Ehrlich, 1983), comb. nov., M. remypignoux Benmesbah & Viloria, sp. nov. y M. remypignoux shueyi, Benmesbah &
Viloria, ssp. nov.; Malaveria Viloria & Benmesbah, gen. nov., con nueve taxones, M. alcinoe (C. Felder & R. Felder, 1867),
comb. nov., M. nebulosa (Butler, 1867), comb. nov., M. mimas (Godman, 1905), comb. nov., M. ballo Benmesbah &
Viloria, sp. nov., M. rodriguezi Benmesbah & Viloria, sp. nov., M. rodriguezi risaralda Benmesbah & Viloria, ssp. nov., M.
bottoi Benmesbah & Viloria, sp. nov., M. duponti Benmesbah & Murienne, sp. nov. y M. mimula (Hayward, 1954), comb.
nov.; y Koutalina Viloria & Murienne, gen. nov., para una sola especie conocida, K. pamela (Hayward, 1957), comb. nov.
Se designan los lectotipos de Euptychia modesta, Neonympha alcinoe y Neonympha sylvina C. Felder & R. Felder, 1867. Se
discute la variabilidad en la morfología y el aspecto del genital masculino, así como el número de pupilas en los ocelos y la
presencia ocasional de un ocelo anal supernumerario en la cara ventral del ala posterior.
Palabras clave: ADN, Argentina, Belize, Bolivia, Cissia, Colombia, Costa Rica, Ecuador, Euptychiina, Guyana Francesa,
Magneuptychia, Panamá, Paraguay, Perú, Venezuela, Yphthimoides.
on, a number of studies in comparative morphology, but
also others combining traditional approaches with mo-
lecular cladistics at dierent degrees, have resulted in the
progressive resolution of the evident polyphyly of Mag-
neuptychia sensu lato (Benmesbah et al. 2018, Andrade et
al. 2019, Nakahara et al. 2020). e taxonomy of several
of the remaining species within this genus (as presented
by Forster 1964 and later Lamas 2004) is still in need of
redenition. Several eorts are being currently addressed
to this worthy task.
e aim of this paper is to provide additional infor-
mation and propose a new taxonomical arrangement to
contribute to a more coherent taxonomic reorganization
of some buttery taxa formerly allocated in this complex
genus.
e discovery in French Guiana of a new species, pre-
viously misidentied as Magneuptychia modesta (Butler,
1867) (Brévignon 2008, Brévignon & Benmesbah 2012)
prompted us to examine similar looking taxa classied for
over 50 years within Magneuptychia sensu lato. Our com-
parative studies, based on morphological character and
molecular analyses, revealed the compared set of species to
belong to two distinct entities, which are herein described
INTRODUCTION
Neotropical nymphalid butteries of the subfamily Sa-
tyrinae have been the subject of fast growing interest dur-
ing the past two decades. A look over published work on
their systematics during the last three years accounts for an
exceptional example of fruitful developments: more than
30 papers, of which about half are revisionary (Barbosa et
al. 2018, Benmesbah et al. 2018, Henao-Bañol & Meneses
[2018], Matos-Maraví et al. 2019, Nakahara et al. 2018a,
2018b, 2019a, 2019b, 2019c, 2020, Pyrcz & Nuñez 2018,
Pyrcz et al. 2018a, 2018b, 2018c, 2019a, 2019b, 2020,
Willmott et al. 2018, 2019, Zacca et al. 2018, 2020a,
2020b, 2020c, Andrade et al. 2019, Brévignon et al. 2019,
Costa et al. 2019, 2020, Espeland et al. 2019, Freitas et al.
2018, 2019, Henao-Bañol 2019, Nakahara & Gallardo
2019, Orellana et al. 2019, Ríos Málaver 2019, Viloria &
Luis Martínez 2019).
Based on morphological characters, Costa et al. (2016)
re-dened the genus Magneuptychia Forster, 1964 and re-
stricted it to three species: Magneuptychia libye (Linnaeus,
1767), type of the genus by original designation, M. liby-
oidea (Butler, 1866) and M. lethra (Möschler, 1883). Later
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
9
as new genera: Modestia Viloria & Benmesbah, gen. nov.
and Malaveria Viloria & Benmesbah, gen. nov. ese two
genera appear to be close to each other and exhibit some
external similarities, which has led to confusion in the past
identications of the species involved.
We discuss the occasional presence of an additional anal
ocellus on VHW, the relative importance of the number of
pupils on VHW ocelli, the intra- and inter-specic vari-
ability in male genitalia and the limits of the DNA analy-
sis. In addition to the new taxa described here, we found
that several cryptic species have still to be described. Due
to very close habitus and genitalic morphology, scarcity of
comparative material as well as accurate collecting data, we
were able to describe with sucient condence only the
taxa for which we had the appropriate information.
We designate the Lectotypes for Euptychia modesta
Butler, 1867 (Fig. 12a), Neonympha sylvina C. Felder & R.
Felder, 1867 (Fig. 12b) and Neonympha alcinoe C. Felder &
R. Felder, 1867 (Fig. 12c) in order to x their identity and
provide the stability necessary for our nomenclatural acts.
Applying similar criteria and using the same methods,
we also investigated the identity of Euptychia pamela Hay-
ward, 1957. e resulting evidence allowed us to recog-
nize and restore its full specic status, and to justify its
separation from any known genera of the Satyrinae. We
therefore describe the new genus Koutalina Viloria & Mu-
rienne, gen. nov., to include this species.
MATERIAL AND METHODS
Specimens from the following public and private col-
lections (acronyms used throughout the text) were exam-
ined for comparative morphological study and to compile
distribution data:
- AN: Andrew Neild collection, Saint Albans, United
Kingdom.
- DT: David Trembath collection, Dorking, United
Kingdom.
- DZUP: Entomological Collection Padre Jesus Santia-
go Moure, Curitiba, Paraná, Brazil.
- ICRM: Indiana Cristóbal Ríos Málaver collection, Ma-
nizales, Colombia.
- IVIC: Instituto Venezolano de Investigaciones Cien-
tícas, Centro de Ecología, reference collection, Vene-
zuela.
- JMD: John MacDonald collection, Starkville, Missis-
sippi, United Sates of America.
- JS: John Shuey collection, Indiana, United Sates of
America.
- LCB: Lalita and Christian Brévignon collection, Ma-
toury, French Guiana.
- MB: Mohamed Benmesbah collection, La Salvetat-
Saint-Gilles, France.
- MBLUZ: Museo de Biología de La Universidad del
Zulia, Maracaibo, Venezuela.
- MIZA: Museo del Instituto de Zoología Agrícola, Uni-
versidad Central de Venezuela, Maracay, Venezuela.
- MNHN: Muséum National d’Histoire Naturelle, Paris,
France.
- NHMUK: Natural History Museum, London, United
Kingdom.
- ZUEC: Museu de Zoologia da Universidade Estadual
de Campinas, São Paulo, Brazil.
e following abbreviations are used throughout the
text: DFW–dorsal forewing; DHW–dorsal hindwing;
FW–Forewing, FWL–Forewing length, HW–Hindwing,
TL–Type locality, VFW–ventral forewing; VHW–ven-
tral hindwing. “*” indicates specimens from which genita-
lia have been dissected for study.
A list of the specimens selected for DNA analysis is
given.
For morphological examination, the specimens stud-
ied had their abdomen removed and soaked in a test tube
containing a heated 10% potassium hydroxide (KOH) or
caustic soda (NaOH) solution before dissection of their
genitalia. Male and female genitalia preserved in glycerol
were contained in microvials and each one attached to
their respective specimen with corresponding label data.
Genitalia were dissected and examined when material was
available but in some cases (lack of material, type speci-
mens) this was not possible. Wing venation preparations
were performed by dislocating forewing and hindwing
from the body; to remove scales, wings were soaked in a
90° alcohol solution for 30 seconds, rinsed with water and
then soaked in a caustic soda solution (NaOH, 10%) for
one minute; aer being rinsed again and dried, wings were
mounted on glass slides; references to specimens were at-
tached to each corresponding slide. In most cases only a
single specimen per species was prepared and analyzed.
Wing venation preparation was not possible for two spe-
cies (M. bottoi sp. nov. and M. gomezi comb. nov.) be-
cause only unique specimen were available. Venation was
compared between the genera based on their respective
type species: Magneuptychia: type species Papilio libye
Linnaeus, 1767; Emeryus Zacca, Casagrande & Mielke in
Zacca et al. (2020a): type species Satyrus argulus Godart,
[1824]; Paryphthimoides Forster, 1964: type species Neo-
nympha poltys Prittwitz, 1865; Cissia Doubleday, 1848:
type species Papilio clarissa Cramer, 1775 = Papilio pe-
nelope Fabricius, 1775. Imago habitus, genitalia and wing
venation photos were taken with a Canon Powershot G7X
reex digital camera. Illustrations were edited using the
M. Benmesbah, Á. L. Viloria & J. Murienne
10
soware GIMP v.2.8 (GIMP team 2016). Specimens were
examined by at least one of the authors directly or via pho-
tographs, with the agreement of the photographer and the
institution authorities where the specimens are housed.
Due to the complexity found in the morphological
variation of the buttery taxa studied, the type series,
when possible, has been restricted to a small and coher-
ent biogeographical area, although other specimens were
available. For the same reasons, only data from specimens
which were identied with good condence were included
in distribution maps and additional data, undetermined
specimens were listed under “sp”. We attempted to review
all major publications where relevant taxonomic names
were cited since the year of their descriptions, with the
caveat that some inevitably have been unintentionally
missed. e original descriptions of Papilio argante Cra-
mer, 1779, Satyrus argulus Godart, [1824], Satyrus gri-
mon Godart, [1824], Neonympha alcinoe C. Felder & R.
Felder, 1867, Neonympha numeria C. Felder & R. Felder,
1867, Neonympha numilia C. Felder & R. Felder, 1867,
Neonympha sylvina C. Felder & R. Felder 1867, Euptychia
huebneri Butler, 1867, Euptychia modesta Butler, 1867,
Euptychia nebulosa Butler, 1867, Neonympha melobosis
Capronnier, 1874, Euptychia ambigua Butler, 1876, Eup-
tychia benedicta Butler, 1877, Euptychia mimas Godman,
1905, Euptychia mimula Hayward, 1954, Euptychia pa-
mela Hayward, 1957 and Cissia gomezi Singer, DeVries &
Ehrlich, 1983, were consulted and translated into French
and English. Images of the type specimens of all the above
mentioned species were examined except for E. pamela, P.
argante, N. melobosis and S. grimon, whose types have not
been located yet or are supposed to be lost. e taxonomic
classication follows Lamas (2004) and the descriptive
terminology of the wings and genitalia follows Klots
(1970), Neild (1996), Kristensen (2003), Peña & Lamas
(2005) and Benmesbah et al. (2018) with a few modica-
tions (Figs. 1, 2, 4a).
Because of the very similar habitus exhibited by each
species treated in this work, we have chosen to describe
here their general aspect, to avoid unnecessarily long de-
scriptions and repetition of common elements. Only some
particular relevant characters are given under the account
of each species.
Male (Figs. 00, 000): Head: brown; front and vertex
white mixed with brown; eyes hairy and dark brown; an-
tennae light brown with white scales at base on each ag-
ellomere; antennal club dark with tip light brown; labial
palpi with elongated white scales medially, and mixed dark
brown and white scales laterally. In one species Koutalina
pamela (Butler) comb. nov. front and vertex brown; eyes
glabrous, dark brown; antennal club dark; labial palpi with
elongated white scales medially, and short white scales lat-
erally on either sides of a central row of longer dark brown
scales. orax: dark covered with blond setae with green-
ish-blue reection; tegulae light brown mixed with red-
dish and greenish iridescent scales; prothoracic legs whit-
ish; meso- and metathoracic legs light brown, coxae and
femora setose. Abdomen: dark covered by blond to brown
setae with a greenish-ginger reection. FWL: 0-00 mm,
average 0.00 mm (x specimens); FW subtriangular, costal
margin convex, apex rounded, outer margin straight, inner
margin slightly concave, cubital and radial veins swollen
at their base. DFW: ground color brown or greyish brown
almost uniform; VFW: ground color brown or greyish
brown; submedian and median lines more or less parallel,
straight, perpendicular to inner margin running from radi-
us to 2A, median line passing outside the cell and subme-
dian through the middle of the cell, crossing Cu2 origin;
submarginal band dark brown, little marked, barely visible,
straight from costa to 2A, parallel to precedent lines; one
small black ocellus circled by a yellowish /brownish ring in
M1-M2 with one or two small white pupils, two others re-
duced and much less dened, if not vestigial or invisible, in
M2-M3 and M3-Cu2; submarginal line thicker, scalloped,
running from radius to inner margin, pointing distally in
R5 to 2A; marginal line straight, thin, parallel and close to
outer margin. DHW: ground color same as DFW; outer
margin slightly scalloped; marginal and submarginal lines
observable through translucent surface, thin marginal area
outside marginal line lighter at tornus; one species K. pa-
mela comb. nov., with black ocelli surrounded by yellow
ring with 2 tiny white pupils, more or less well dened,
in Cu1-Cu2
and Cu2-2A. VHW: ground color same as
VFW; one small basal line very close to base, almost non
visible; submedian and median lines perpendicular to cos-
ta, median line passing through the distal extremity of the
cell, and submedian line through the middle of the cell,
both lines running from costa to inner margin; a thin dark
line almost non visible, very close and parallel to inner
margin, joins submedian, median, submarginal and mar-
ginal lines; submarginal line thicker, from costa to inner
margin, scalloped, composed by more or less straight or
curved adjacent segments, basally convex from M1-M2 to
Cu2-2A; marginal dark line thin parallel to outer margin
from costa to tornus, one species K. pamela comb. nov.,
with marginal line clearly wider between anal part of Cu2-
2A and tornus; submarginal area with ve ocelli, one in
each space between Rs and Cu2, of variable size, usually
black circled with yellowish/brownish ring and centered
by one or two white pupils, ocellus in M1-M2 well dened
with one or two pupils, ocellus in
Cu1-Cu2
well dened
with two white pupils, ocelli in M2-M3 and M3-Cu1
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
11
more or less well dened, ocellus in M2-M3 usually ovoid
slightly displaced basally compared to the others, ocellus
in Rs-M1
usually much smaller than precedents, usually
well dened, centered by one or two white pupils, all these
ocelli surrounded by a thin submarginal dark area; one
species K. pamela comb. nov., with a sixth ocellus with
two pupils in Cu2-2A.
Female (Figs. 00, 000) – FWL: 0-00 mm, average
0.00 mm (x specimens). Usually similar to male, but dif-
fers from that by upper side darker or lighter, and global
shape of FW more rounded and less elongated with outer
margin shorter, giving a more crenulated aspect to HW in
some species.
Male genitalia were positioned in a standardized way
to best allow comparison. e angle of view in which they
are studied and illustrated has an important inuence over
their perceived aspect (see discussion). For the entire male
genitalic structure, it is shown the upper and lateral view
(one valva removed); external, inner and upper view for
valvae; upper and lateral view for aedeagi. For each species,
when possible, several specimens where dissected to evalu-
ate intraspecic morphological stability and variability;
the number of specimens dissected are indicated in the ex-
amined material section and marked by an asterisk “*”. We
focused mainly on the shape of valvae and aedeagi, which
seem, from our experience, to reveal the most informative
and discriminating characters. In case of infraspecic vari-
ation, we choose to illustrate a specimen that was represen-
tative of the majority of the specimens studied. Comments
on variability of the male genitalia are also discussed (see
discussion).
Female genitalia are also illustrated, but due to few
available material at our disposition, not all species could
be studied and few specimens where dissected.
DNA extraction was performed using a Chelex proto-
col (Casquet et al. 2012) as in Benmesbah et al. (2019). We
chose to amplify a short fragment of the 16S mitochon-
drial rRNA using the Ins16S_1 primer pair (Clarke et al.
Figure 1. Terminology of wing pattern elements.
Figure 2. Terminology of male genitalia elements: lateral view
and aedeagus: aa: appendix angularis; ar: anterior region of ae-
deagus; c: costa; ct: cornuti; Sb: subuncus; pr: posterior region
of aedeagus; s: saccus; t: tegumen; u: uncus; v: valva.
M. Benmesbah, Á. L. Viloria & J. Murienne
12
2014): Ins16S_1-F: TRRGACGAGAAGACCCTATA;
Ins16S_1-R: TCTTAATCCAACATCGAGGTC. De-
spite being originally developed for metabarcoding stud-
ies, this fragment has been shown to provide good resolu-
tion at the species level in several insect groups (Kocher et
al. 2017a, 2017b, Talaga et al. 2017). Because of its small
size (ca. 220 bp), it allows to provide good amplication
results even from degraded material. Amplication and
sequencing was performed on 101 specimens following
Kocher et al. 2017a, 2017b, of which 84 specimens were
represented in Table 1 to avoid unnecessary and non-
signicant redundancy of the same sequence. Sequences
were aligned using Muscle (Edgar 2004) and a Maximum
Likelihood analysis was performed in Raxml-ng with a JC
substitution model (Jukes & Cantor 1969) as inferred by
Modeltest-ng (Darriba et al. 2020) and a midpoint root-
ing.
RESULTS
Modestia Viloria & Benmesbah, gen. nov.
(Figs. 3a-3d)
http://zoobank.org/ urn:lsid:zoobank.org:act:3D9F6817-3BF0-48F4-9E65-
1695D697AB44
Type species: Euptychia modesta Butler, 1867a;
herein designated.
Diagnosis (Figs. 3, 4, 5)
Supercially similar to members of the genera Emeryus,
Malaveria Viloria & Benmesbah, gen. nov., Koutalina
Viloria & Murienne, gen. nov. and Paryphthimoides (sensu
Zacca et al. 2020c). It diers from the latter in its larger
size and in the number of VHW ocelli: ve in Modestia
Viloria & Benmesbah, gen. nov., six in Paryphthimoides
as in Koutalina gen. nov. e ocellar pattern of Modestia
gen. nov., is characterized by a subapical VFW ocellus in
M1-M2 invariably monopupillate (two pupils in Malave-
ria gen. nov. and Koutalina gen. nov., two vestigial in Em-
eryus). Ventral HW with ve ocelli, the ones in Rs-M1 and
M1-M2 with single white pupil (ve ocelli in Emeryus and
Malaveria gen. nov., six in Koutalina gen. nov., but in all
these cases with double pupils, see discussion). e upper
surface of both wings is brown, uniform, devoid of ocelli
(similar in Magneuptychia and Malaveria gen. nov., but
dierent in Yphthimoides Forster, 1964, Paryphthimoides,
Emeryus and Koutalina gen. nov., in which there is always
at least one or two anal HW ocelli). Forewing venation
is most similar to Emeryus, especially regarding the ori-
gin of R2, proximal and close to the separation of R and
M1, but Modestia gen. nov., diers by m1-m2 less curved
and m2-m3 more curved than in Emeryus. Modestia gen.
nov., diers from Magneuptychia, Paryphthimoides, Cissia
and Koutalina gen. nov., also by the more curved m2-m3
(straight in all these genera). Hindwing venation, although
supercially similar with all the following genera, diers
in the signicantly higher ratio between discal length/M3
length in Modestia gen. nov., when compared to Magneup-
tychia, Emeryus, Paryphthimoides, Cissia, Malaveria gen.
nov. and Koutalina gen. nov. (Figs. 3, 4, 5). Additional
dierences with Malaveria gen. nov., are discussed under
the latter. Male genitalia in the species of Modestia gen.
nov., bear simple, not ornamented, moderately elongated
valvae, with an internal bulge at its distal part, less robust
at distal half than those of Paryphthimoides, not digitiform
like in Malaveria gen. nov., or not distally broadened like
in Emeryus or Koutalina gen. nov.
DNA analysis shows a dierence ratio between the re-
spective type species of the following genera in comparison
to Modestia gen. nov., of: 5.9% for Magneuptychia, 14.4%
for Cissia (sensu stricto), 10% for Malaveria gen. nov., and
10.9% for Koutalina gen. nov., which are coherent with
generic distances (Tables 1 and 2).
Description (Figs. 3a-3d)
Butteries of medium size (FWL: 19-23mm), with-
out sexual dimorphism. Wings dorsally brown, devoid
of ocelli. Ventrally ground color brown with two darker
stripes or bands, crossing both wings more or less in paral-
lel, one through the middle of the discal cell (submedi-
an), the other one median; two dark brown lines, one in-
dented, submarginal, the other marginal, smooth, parallel
to outer margin and not wider at tornus. Ventral ocelli,
when not vestigial, all black, with white pupils, iris yellow,
circled with brown. FW with a subapical M1-M2 ocel-
lus with one pupil, depending on the species there may
appear other similar ocelli in M2-M3 and M3-Cu1. Fe-
males tend to develop more ventral FW ocelli. HW with
ve ocelli, Rs-M1 and M1-M2 with single pupil, M2-M3,
M3-Cu1 and Cu1-Cu2 with double pupils (but some-
times pupils in M2-M3 and M3-Cu1 either fused, vesti-
gial or not developed). FW venation: Sc markedly swollen
at basal third, ending at three hs of costa, R1 emerging
at fourth h of discal cell, R2 and R3-R5 emerging close
to each other, but clearly separated, R3 emerging half the
length of distance between the root of R3-R5 and the bi-
furcation of R4 and R5; rs-m1 very short and straight, a
sixth of the length of m1-m2 (which is convex towards
base), m2-m3 slightly curved, twice the length of m1-m2,
about same length of m3-cu1; cu1-cu2 straight, twice as
long as m3-cu1, Cu2-A2 notably swollen at basal half, A2
independent; HW venation: Humeral present; Sc + R1
ending at rst third of costa, Rs emerging nearly at half
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
13
Table 1. Phylogenetic tree representing genetic distances and relationships among buttery species of the satyrine genera
Modestia gen. nov., Malaveria gen. nov., and Koutalina gen. nov. together with representatives of four additional genera
of the Neotropical Euptychiina.
M. Benmesbah, Á. L. Viloria & J. Murienne
14
Table 2. Representation of the percentage of genetic similarity between the dierent satyrine taxa sampled for this study. ese results were obtained from the
analysis of a short fragment of 218-220 base pairs (16S marker) of mitochondrial rDNA.
length of discal cell and ending at apex; m1-m2 slightly
convex towards base, m2-m3 three times the length of
rs-m1, and twice the length of m1-m2. Male genitalia
characterized by a short globular tegumen, long lanceo-
late uncus, twice the length of the tegumen, subunci thin,
two thirds the length of uncus, vinculum dorsally strong,
ventrally thin, attached to a stylized, long saccus (as long
as the brachia of the subunci), valvae elongated, gradually
thinner towards distal extremity, with a non-conspicuous
ampullar protuberance, with tiny ornamentations, as long
as tegumen plus uncus and presenting an internal bulge at
its distal extremity; aedeagus thin and at least as long as
valva (Figs. 13j-k, 14i-j, 15j-k, 16h-i), straight or slightly
curved upwards.
Derivatio nominis
Modestia is a feminine noun that means modesty,
which is the quality of being modest or humble. e name
is also intended to produce a euphonic combination with
the name of the type species of the genus, Euptychia mod-
esta Butler, 1867a.
Modestia gen. nov., is basically a genus embracing spe-
cies of lowland Neotropical forests.
Species belonging to Modestia Viloria & Benmesbah,
gen. nov.:
Modestia gomezi (Singer, DeVries & Ehrlich, 1983), comb.
nov.
Cissia gomezi Singer, DeVries & Ehrlich, 1983: 114,
gs. 2C, 3C [TL: Costa Rica]; DeVries, 1987:
274, pl. 48 gs. 32, 33, 35c; Matamoros & Seal,
1994: 9, 17, 44, 70; Maes, 1995: 26, 27; Pelz,
1997: 41; Núñez Miño 2004: 29, Fratello, 2007:
1-5; Wiemers & Fiedler, 2008: 290; Albert et al.,
2010: 68; Costa et al., 2016: 199; See et al., 2018:
52; Zacca et al., 2018: 356, 357.
Euptychia gomezi (Singer, DeVries & Ehrlich, 1983);
D’Abrera, 1988: 789.
Magneuptychia gomezi (Singer, DeVries & Ehrlich,
1983); Lamas, 2004: 220; Wiemers & Fiedler,
2008: 290; Marín & Uribe, 2009: 28, 29; Albert
et al., 2010: 68; Garwood & Lehman, 2011: 273
[row 3, gs.]; See et al., 2018: 52.
“M.” gomezi (Singer, DeVries & Ehrlich, 1983); Cos-
ta et al., 2016: 204.
Modestia modesta (Butler, 1867a), comb. nov.
Euptychia modesta Butler, 1867a: 473 [TL: Bra-
zil (Pará)]; Butler, 1867b: pl. 12, g. 12; Butler,
1868: 21; Kirby, 1871: 49; Weymer, 1911: 206;
Hall, 1939: 33; D’Abrera, 1988: 776 [row 2, gs.].
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
15
[Euptychia gracilis Weymer 1911: D’Abrera, 1988:
780 [row 6, g. male] (misidentication)].
Euptychia ambigua Butler var. modesta Butler; Gae-
de, 1931: 438.
Yphthimoides modesta (Butler, 1867a); Forster,
1964: 102, g. 99 (male genitalia).
[Cissia alcinoe (C. Felder & R. Felder, 1867); De-
Vries, 1987: 276,pl. 41 g. 1; Singer & Ehrlich,
1993: 51 (misidentications)]
Magneuptychia modesta (Butler, 1867a); Lamas,
2004: 220; Cock, 2014: 11; Nakahara et al.,
2015: 143; Sambhu & Nankishore, 2018: 119.
[Magneuptychia alcinoe (C. Felder & R. Felder,
1867); Chacón & Montero 2007: pl. 174;
Garwood & Lehman, 2011: 274 [row 2, g.] (mi-
sidentications)]
[Megeuptychia antonoe (Cramer, 1775); Garwood
& Lehman, 2011: 275 [row 4, g.] (misidenti-
cation)]
Modestia remypignoux remypignoux Benmesbah & Vilo-
ria, sp. nov. [TL: French Guiana]
[Euptychia modesta Butler, 1867a var.]; Butler,
1867a: 473; 1868: 21.
[Euptychia nebulosa Butler, 1867a; Godman & Sal-
vin, 1880: 86, pl. 8, g. 10 (misidentication)]
Modestia remypignoux shueyi Benmesbah & Viloria ssp.
nov. [TL: Belize]
Malaveria Viloria & Benmesbah, gen. nov.
(Figs. 3e-3h)
http://zoobank.org/urn:lsid:zoobank.org:act:40D0F5BB-5728-42EB-B7BF-
2704D0BEC845
Type species: Euptychia nebulosa Butler, 1867a;
herein designated.
Diagnosis (Figs. 3, 4, 5)
Malaveria gen. nov., is most similar in size and appear-
ance to Modestia gen. nov., which has been diagnosed
above; their dierences can be summarized as follows:
1.FW venation in Malaveria gen. nov. is characterized
by a tiny basally-pointing recurrent vein on discocellular
m1-m2, present in all the species treated here as belong-
ing to Malaveria gen. nov. (absent in Modestia gen. nov.);
2.VHW ocelli in M1-M2 and Rs-M1 are bipupillate in all
known species of Malaveria gen. nov., instead of mono-
pupillate in Modestia gen. nov. (see discussion); 3. dorsal
edge of the valvae in male genitalia presenting a marked
bulge in Malaveria gen. nov. (straight in Modestia gen.
nov.); 4. distal extremity of the valvae digitiform and
hooked in Malaveria gen. nov., instead of relatively wider
and gradually pointed in Modestia gen. nov.; 5. presence
of a small and well dierentiated internal horn emerg-
ing more basally in the valvae of the species of Malaveria
gen. nov., instead of a bulge at the distal part of the valvae
(Modestia gen. nov.); 6. aedeagus in Malaveria gen. nov.,
equally long (as much as valvae) but relatively thicker and
more straight than in Modestia,
DNA analysis shows dierent percentages of diver-
gence between the respective type species of the follow-
ing genera in comparison to Malaveria gen. nov.: 9.1%
for Magneuptychia, 13.6% for Cissia (sensu stricto), 10%
for Modestia gen. nov., and 10.9% for Koutalina gen.
nov., which are coherent with generic distances (Tables1
and 2).
Description
Butteries of medium size (FWL: 19-26mm). Wings
dorsally brown except for two darker marginal lines,
more visible on HW. Ventrally ground color brown with
two darker reddish lines, crossing both wings more or less
in parallel, one submedian, the other one discal, they can
be straight, moderately or strongly undulated, depend-
ing on the species; two dark brown lines, one thicker,
indented, submarginal, meeting the discal one near the
anal margin of HW, the other thinner, marginal, smooth,
parallel to outer margin. Ventral ocelli, when not vesti-
gial, all black, with white pupils, iris yellow, circled with
brown. FW with a subapical M1-M2 ocellus with two
tiny pupils. HW with ve ocelli or ocellar vestigia, in
Rs to Cu2, all when apparent, with double pupils (but
sometimes pupils in Rs fused in one). FW venation: Sc
broadly swollen at basal third, thinning abruptly, end-
ing at half costa; R1 emerging at distal h of discal cell,
R2-R5 emerging at distal tip of discal cell, R2 and R3-R5
branching about 2millimeters further towards the apex,
R3 and R4 separating equidistantly, at rst and second
third of their root, respectively, R5 ending at apex; r2-
5-m1 very short and straight, a sixth of the length of m1-
m2, which is convex towards base with a tiny recurrent
vein at angle, inside discal cell; m2-m3 slightly concave
towards base, twice the length of m1-m2, about same
length of m3-cu1; cu1-cu2 one and a half times as long as
m3-cu1; Cu2-A2 moderately inated at basal two hs;
A2 independent. HW venation: Humeral present, but
short and simple; Sc thin, ending at basal third of costa;
Rs and M1 branching separately but close to each other
at half length of discal cell; m1-m2 slightly curved to-
wards the cell; M2 emerging equidistantly from roots of
M1 and M3; cu1-cu2 2 times the length of m3-cu1; A2
and A3 independent. Male genitalia: tegumen globular,
long, lanceolate uncus, almost twice the length of the
tegumen, subunci thin, as long as uncus, usually pointing
M. Benmesbah, Á. L. Viloria & J. Murienne
16
Figure 3. New genera, male wing venation, habitus and genitalia. a. Modestia gen. nov., venation; b. Habitus ventral; c. Habitus dorsal;
d.Male genitalia lateral view; e. Malaveria gen. nov., venation; f. Habitus ventral; g. Habitus dorsal; h. Male genitalia lateral view; i.Koutali-
na gen. nov., venation; j. Habitus ventral; k. Habitus dorsal; l. Male genitalia lateral view. (Scale habitus: 1 cm; scale genitalia: 1mm).
upwards; vinculum strong and relatively thick, attached
to a relatively long, tubular saccus (slightly shorter than
uncus), valvae robust at basal half, laterally quadran-
gular, with a very broad base (half the lateral length of
vinculum), abruptly thinner towards distal half, which
becomes a digitiform extremity, at whose base emerges
internally a distinctive horn-like protuberance; aedeagus
regular and at least as long as valva, broader at base,
straight or just slightly sinuous.
Derivatio nominis
Malaveria is a feminine Latinization of the Spanish
family name Málaver. It is dedicated to our friend and col-
league Indiana Cristóbal Ríos Málaver, a talented young
scientist in Neotropical lepidopterology and an expert in
the butteries of Colombia, his native country. Cristo – as
he is known among lepidopterists – is certainly the person
who best knows the biology of the type species of this ge-
nus, Euptychia nebulosa Butler, 1867a, an endemic of the
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
17
Cordillera de la Costa in Venezuela, where Cristo devel-
oped buttery studies for his MSc thesis.
All species of Malaveria gen. nov., are basically denizens
of middle elevation montane forests in Tropical America.
Species belonging to Malaveria Viloria & Benmesbah, gen.
nov.:
Malaveria alcinoe (C. Felder & R. Felder, 1867), comb.
nov.
Neonympha alcinoë C. Felder & R. Felder, 1867: 477
[TL: Colombia (Bogotá), herein restricted].
Euptychia alcinoë (C. Felder & R. Felder, 1867);
Butler, 1868: 25; Kirby, 1871: 51; Weymer, 1911:
209, pl. 48 row c.
Euptychia benedicta Butler, 1877: 124, pl. 12, g. 14
[TL: Ecuador]; Kirby, 1877: 843; Weymer, 1911:
206-207; D’Abrera, 1988: 778 [row 1, gs.]; La-
mas, 2004: 220 (as a synonym of Magneuptychia
alcinoe (C. Felder & R. Felder, 1867))
Euptychia ambigua (Butler, 1867a) var. benedicta
Butler, 1877; Gaede, 1931: 437-438.
Euptychia alcinoe (C. Felder & R. Felder, 1867);
Gaede, 1931: 437; D’Abrera, 1988: 777 [row 8,
gs.].
Yphthimoides benedicta (Butler, 1877); Forster,
1964: 101 g. 94 (male genitalia, probably erro-
neous), 104.
Cissia alcinoe (C. Felder & R. Felder, 1867); DeVries,
1987:276,pl. 41 g. 1 (probably misidentica-
tion of Modestia remypignoux shueyi ssp. nov.);
Singer & Ehrlich, 1993: 251 (misidentication
of Modestia modesta (Butler, 1867a), comb. nov.,
from Costa Rica and Trinidad, respectively)
Magneuptychia alcinoe (C. Felder & R. Felder,
1867); Lamas, 2004: 220; Chacón & Montero
2007: pl. 174; Garwood & Lehman, 2011: 274
[row 2, g.] (last two misidentications of Mo-
destia modesta (Butler, 1867a), comb. nov., from
Costa Rica); Beccaloni et al., 2008: 335; Nakaha-
ra et al., 2015: 136, 137-139 g. 1d (male synty-
pe), 142, 144, 145.
Malaveria ballo Benmesbah & Viloria, sp. nov. [TL:
Ecuador (Pichincha)]
Malaveria bottoi Benmesbah & Viloria, sp. nov. [TL: Co-
lombia (Cundinamarca)]
Malaveria duponti Benmesbah & Murienne, sp. nov. [TL:
Bolivia (La Paz)]
[Yphthimoides nebulosa (Butler, 1867a); Forster,
1964: 104 g. 105 (male genitalia), 106 (misi-
dentication)]
Malaveria mimas (Godman, 1905), comb. nov.
Euptychia mimas Godman, 1905: 187, pl. 10, g. 6
(male) [TL: Bolivia (Coroico)]; Weymer, 1911:
2010, t. 48 d; Gaede 1931: 454; D’Abrera, 1988:
777 [row 7, gs.]
Yphthimoides mimas (Godman, 1905); Forster,
1964: 101 g. 96 (male genitalia), 102, 105.
Magneuptychia mimas (Godman, 1905); Lamas,
2004: 220; Nakahara et al., 2015: 136, 137, 139
g. 1e (male syntype), 142.
Malaveria mimula (Hayward, 1954), comb. nov.
Euptychia mimula Hayward, 1954: 16-17, g. 1
(male genitalia) [TL: Bolivia (Chulumani)];
D’Abrera, 1988: 789.
Yphthimoides mimula (Hayward); Forster, 1964:
102, 105; Lamas, 2004: 223.
Malaveria nebulosa (Butler, 1867a), comb. nov.
Euptychia nebulosa Butler, 1867a: 474 [TL: Vene-
zuela (Cordillera de la Costa)]; 1867b: pl. 12,
g. 2; 1868: 26; Kirby, 1871: 50; Godman & Sal-
vin, 1880: 86, pl. 8, g. 10 (misidentication of
Modestia remypignoux sp. nov., from Panama);
Hall, 1939: 236 (misidentication of Modestia
remypignoux sp. nov., from Guyana); D’Abre-
ra, 1988: 776 [row 3, gs. male] (misidentica-
tionof Malaveria mimula comb. nov.).
Yphthimoides nebulosa (Butler, 1867a); Forster,
1964: 104 g. 105 (male genitalia), 106 (misi-
dentication of Malaveria duponti sp. nov., from
Bolivia); Brown & Mielke, 1967: 91 (misiden-
tication of ‘Paryphthimoides’ grimon (Godart,
[1824]) from Brazil).
Magneuptychia nebulosa (Butler, 1867a); Lamas,
2004: 220; Nakahara et al. 2015: 135-147 gs. 2a
(head), 2b, c (male genitalia), 2d, e, f (female ge-
nitalia), 3 (distribution map), 4 (habitus).
Malaveria rodriguezi rodriguezi Benmesbah & Viloria, sp.
nov. [TL: Colombia (Antioquia)]
Malaveria rodriguezi risaralda Benmesbah & Viloria, ssp.
nov. [TL: Colombia (Risaralda)]
Koutalina Viloria & Murienne, gen. nov.
(Figs. 3i-3l)
http://zoobank.org/ urn:lsid:zoobank.org:act:610B37EC-1C54-4073-81DB-
80DF1CAC95EF
Type species: Euptychia pamela Hayward, 1957;
herein designated.
Diagnosis (Figs. 3, 4, 5, 29)
Koutalina gen. nov., diers from all other genera of the
Euptychiina by the presence in males of a large dark patch
M. Benmesbah, Á. L. Viloria & J. Murienne
18
Figure 4. Species of Malaveria gen. nov., male wing venation with focus on forewing cell (x3). a. Venation terminology; b. M. nebu-
losa comb. nov.; c. M. mimas comb. nov.; d. M. alcinoe comb. nov.; e. M. ballo sp. nov.; f. M. rodriguezi sp. nov.; g. M. mimula;
h. M. duponti sp. nov. (Scale: 1 cm).
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
19
of modied scales on DFW probably representing an-
droconia (see under Koutalina pamela comb. nov.). Fur-
thermore Koutalina gen. nov., diers from Modestia gen.
nov., and Malaveria gen. nov., by the following major fea-
tures: 1. Koutalina, gen. nov., presents a sixth supplemen-
tary ocellus on VHW, near the tornus, which is absent in
Modestia gen. nov. and Malaveria gen. nov.; 2. on DHW
Koutalina gen. nov. has a bipupillate ocellus in Cu1-Cu2,
which is missing in Modestia gen. nov. and Malaveria
gen. nov.; 3. VHW marginal line broadens at tornus in
Koutalina gen. nov., which does not occur in Modestia
gen. nov. nor Malaveria gen. nov.; 4. singular spatulate
aspect of the distal extremity of the valvae in Koutalina,
gen. nov., very distinctive and dierent from those of Pa-
ryphthimoides and Emeryus. Even if Koutalina gen. nov.,
bears six VHW ocelli, it diers from Magneuptychia in the
position of the sixth ocellus (Cu2-2A in Koutalina gen.
nov., 1A-2A-3A in Magneuptychia) and also in wing vena-
tion and male genitalic morphology. Koutalina gen. nov.,
phenotypically diers from Euptychoides Forster, 1964 in
ocellus Cu1-Cu2 on VHW, which bears a single pupil in
the latter (with the notable exception of E. pseudosaturnus
Forster, 1964). Furthermore, typical Euptychoides species
have only ve VHW ocelli and have a serrate process in
the valvae of male genitalia; these characters are not found
in Koutalina gen. nov. Species of the genus Optimandes
Marín, Nakahara & Willmott, 2019, which might show
a supercial similarity to Koutalina pamela (Hayward,
1957), comb. nov., have only ve ocelli on VHW, and
those are all monopupillate. e same kind of singled-pu-
pil ocelli are found in all species of Hermeuptychia Forster,
1964, whose largest species, such as H. harmonia (Butler,
1867a), resemble the habitus of Koutalina gen. nov.
DNA analysis shows dierent divergence ratios be-
tween the respective type species of the following genera
in comparison to Koutalina gen. nov., of: 9% for Mag-
neuptychia, 7.2% for Cissia (sensu stricto), 10.9% for Ma-
laveria gen. nov. and 12.7% for Euptychoides, which are
coherent with generic distances (Tables 1 and 2).
Description (Figs. 3i-3l)
Butteries of medium size (FWL: 19-23mm). Wings
dorsally brown with a large dark patch of modied scales
on FW, presenting a bipupillate ocellus in Cu1-Cu2 and
Figure 5. Comparative venation and habitus of male imagos of the type species of similar genera: a. Magneuptychia venation (type species
M. libye); b.Male ventral; c. Dorsal; d. Emeryus venation (type species E. argulus); e. Male ventral; f. Dorsal; g. Paryphthimoides venation
(type species P. poltys); h. Female ventral; i. Dorsal; j. Cissia venation (type species C. penelope); k. Male ventral; l. Dorsal. (Scale: 1 cm).
M. Benmesbah, Á. L. Viloria & J. Murienne
20
occasionally an additional one much smaller in Cu2-
1A+2A; two darker marginal lines more visible on HW.
Ventrally ground color brown (lighter) with two darker
grossly straight lines crossing both wings more or less
in parallel, one submedian, the other median; twodark
brown lines, one irregularly undulated, submarginal,
meeting the median line near the anal margin of HW, the
other thinner, marginal, parallel to outer margin, notably
broadening in tornus. Ventral ocelli, when not vestigial,
all black, with double white pupils, iris ochreous yellow,
nely circled with brown. FW with a subapical ocellus in
M1-M2 with two tiny pupils. HW with six ocelli from
Rs to 1A+2A, all with double pupils (but sometimes
pupils barely visible in M2 and M3). FW venation: Sc
broadly swollen at basal third of the cell, ending approxi-
mately at half costa; R1 emerging at distal sixth of discal
cell, R2 and R3-R5 branching at distal tip of discal cell,
R3 emerging half way between the end of discal cell and
the nal branching of R4 and R5; r2-5-m1 very short
and straight, m1-m2 and m2-m3 notably straight; m2-
m3 four times as long as m1-m2, and m1-m2 twice as
long as r2-5-m1; m3-cu1 and cu1-cu2 straight, all giving
the cell a quadrangular aspect; cu1-cu2 twice as long as
m3-cu1, and about same length as m2-m3; Cu2-A2 only
slightly swollen at base; 1A+2A independent. HW ve-
nation: Humeral present, thick and well developed but
simple; Sc thin, ending at basal third of costa; Rs and M1
branching separately; m1-m2 curved towards the cell;
M2 emerging closer to M1 than to M3; cu1-cu2 twice
the length of m3-cu1; 1A+2A and A3 independent
but running together at base. Male genitalia: tegumen
globular, uncus lanceolate, broad, about same length of
tegumen, subunci thin, not reaching the length of uncus;
vinculum stylized and sigmoidal in lateral view, saccus
tubular, thick, as long as tegumen, valvae strong but elon-
gated, equally thick at base and basal half, slightly thin-
ner towards distal half, which ends in a spatulate extrem-
ity, reminiscent of the shape of a wooden kitchen spoon;
aedeagus thin, as long as valva, slightly curved upwards.
Derivatio nominis
Koutalina is a Latinized feminine diminutive for kou-
táli (κουτάλι), the Greek word for spoon. is name is
proposed for this new genus of butteries because male in-
dividuals of its only known species, K. pamela (Hayward,
1957), comb. nov., bear a distinctive genitalia, whose val-
vae have a spoon-shaped tip.
Koutalina gen. nov., is so far represented by a single
known species that inhabits medium elevations in the An-
des of Bolivia and Peru.
Species belonging to Koutalina Viloria & Murienne, gen.
nov.:
Koutalina pamela (Hayward, 1957), comb. nov.
Euptychia pamela Hayward, 1957: 113-115, 119
g.3 (male genitalia).
Euptychia pamela Hayward; Forster, 1964: 105
(erroneously as a synonym of Yphthimoides be-
nedicta (Butler, 1877)); Lamas, 2004: 220 (erro-
neously as a synonym of Magneuptychia alcinoe
(C. Felder & R. Felder, 1867)).
Euptychia? sp.; D’Abrera, 1988: 777 [row 6, g.]
(misidentication).
DISCUSSION
Extended comparison between Modestia gen. nov. and
Malaveria gen. nov.: Ocellar pattern and male genitalia
Number of pupils in VHW ocelli (Figs. 6-9)
Ocellus M1-M2
On VHW the number of pupils in ocelli M1-M2
and Rs-M1 can be subject to some apparent variation.
In Modestia gen. nov., pupils are well dened, formed
by several white/silver scales, placed in the center of the
ocellus (Fig.6a). In Malaveria gen. nov., the two pupils
in M1-M2 are formed typically by fewer silver scales,
both disposed on a line passing through the center of
the ocellus and parallel to the median line. e more
anterior pupil is usually larger than the posterior one
(Figs. 6b-6d), which can be represented by at least one
or two scales. Ocellus in Rs-M1 seems to be too small
to always allow expression of the pupils, it is probably a
bipupillate ocellus but the silver scales can either repre-
sent clearly two dierent small pupils, be fused in one big
pupil or simply not be present. It is more or less the same
for ocelli M2-M3 and M3-Cu1, which are of almost the
same small size as the latter in both genera, and its pupils
are only well and constantly expressed in M. r. remypig-
noux sp. nov. e examination of specimens belonging
to Malaveria gen. nov. and Modestia gen. nov., shows
that, when silver scales are present, these two ocelli are
usually bipupillate. In Modestia gen. nov. (19 Modestia
modesta comb. nov. and 19 M. r. remypignoux sp. nov.,
examined), ocelli M1-M2 and Rs-M1 are always well
dened and monopupillate. None of the specimens ex-
amined showed two pupils. In Malaveria gen. nov. (15
Malaveria ballo sp. nov., 1 Malaveria bottoi sp. nov., 5
Malaveria mimas comb. nov. and 7 Malaveria nebulosa
comb. nov. examined), ocellus M1-M2 shows generally
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
21
two silver pupils but only one in some specimens. e
study with stereoscopic microscope has shown that the
apparent single pupil is in fact not a variation but an ar-
tefact due to several factors:
- Fusion of the two pupils, which appear larger than a
central unique average pupil. For example, in M. mimas
comb. nov.: 1. Figs. 7a-7b. normal bipupillate ocelli;
2.Figs. 7c-7d. in the same specimen a transitional varia-
tion with a bipupillate ocellus on one wing and a fusion
of the two pupils on the other; 3. Figs. 7e-7f. variation
where the two pupils are almost completely fused on
both wings. Another example in M. ballo sp. nov.,
where the two pupils have fused together in a large cen-
tral unique pupil (Fig. 6e).
Figure 6. Detail of ocellus M1-M2 on VHW: a. Modestia modesta comb. nov. (MB-0023); b. Malaveria ballo sp. nov. (MB-0070);
c. M. ballo sp. nov. (MB-0076); d. M. ballo sp. nov. (MB-0074); e. M. ballo sp. nov. (MB-0257); f. M. ballo sp. nov. (MB-0016).
Figure 7. Detail of ocellus M1-M2 on VHW, fusion of the two pupils: a-b. Malaveria mimas comb. nov. (MB-0014) le and right
HW; c-d. M. mimas comb. nov. (MB-0012) le and right HW; e-f. M. mimas comb. nov. (MB-0010) le and right HW.
Figure 8. Detail of ocellus M1-M2 on VHW, no expression or reduction of one of the two pupils: a-b. Malaveria nebulosa comb. nov.
(MB-0004) le and right HW; c-d. M. nebulosa comb. nov. (MB-0007) le and right HW; e-f. M. nebulosa comb. nov. (MB-0003)
le and right HW.
Figure 9. Detail of ocellus Cu1-Cu2 on VHW, fusion of the two pupils: a-b. Modestia modesta comb. nov. (MB-0022) le and right
HW; c-d. M. modesta comb. nov. (MB-0023) le and right HW; e-f. M. modesta comb. nov. (MB-0030) le and right HW.
M. Benmesbah, Á. L. Viloria & J. Murienne
22
- No expression, or loss of scales of one of the pupils (usua-
lly the smallest one), the apparent pupil is then not in the
center (usually in the more anterior position). For exam-
ple, in M. nebulosa comb. nov.: Figs. 8a-8b: normal bipu-
pillate ocelli; Figs. 8c-8d: transitional variation with one
of the two pupils only represented by a single scale (note,
in this particular specimen a few scales aberrantly loca-
ted, look like forming a third pupil); Figs. 8e-8f: in the
same specimen one ocellus still has two pupils composed
by a few scales, while the other ocellus presents a single,
eccentric, tiny pupil, due to the non-expression of the se-
cond pupil; Fig. 6f: apparent loss of the smallest pupil
due to the worn condition of the specimen.
Ocellus Cu1-Cu2
Contrastingly in both genera, the number of pupils
(two) in ocellus Cu1-Cu2 is very stable. Only one speci-
men (over 66 examined) showed an apparent unique pu-
pil (Figs. 9a-9b.). e mechanism involved is probably the
same as explained above: fusion of the two pupils like in
Figs. 9c-9f. It is worth noting that the only syntype of Neo-
nympha sylvina C. Felder & R. Felder, 1867, we are aware
of (Fig. 12b.) presents a single pupil in ocellus Cu1-Cu2
on both VHWs (see discussion in description of M. r. re-
mypignoux sp. nov.).
It appears from our analysis that the number of pupils
in ocelli M1-M2 and Cu1-Cu2 is a primary character
that easily dierentiates phenotypically these two genera.
e “apparent” single pupil is an artifact of the mecha-
nisms above mentioned. In most cases, it can be unmasked
through careful examination. Based on our experience, we
recognize the number of ocelli, their composition and the
number of pupils as good elements to establish morpho-
logical homologies to support taxonomic arrangements at
generic level in the Satyrinae. is hypothesis should be
tested with other buttery groups and compared with male
genitalia morphology and DNA analysis to explore if they
are consistent to support new taxonomic arrangements.
Variation in distal extremity of the valvae (Figs. 10)
Generally, most satyrine species are distinguishable
from their congeners by the morphology of their male
genitalia (Nakahara et al. 2015). Other groups of but-
teries have shown male genitalia morphology not to be
discriminating between closely related species (e. g., the
genus Adelpha Hübner, 1819; see Willmott 2003). A pre-
liminary study of Nakahara et al. (2015) pointed out that
male genitalia in M. nebulosa comb. nov. and M. alcinoe
comb. nov., were very close and subject to some variation.
In this study the examination of more than 60 male geni-
talia belonging to Modestia gen. nov. and Malaveria gen.
nov., showed that intraspecic variation is much more
important than previously estimated. e range of varia-
tion overlaps within species respectively belonging to each
genus and causes some confusion. e distal extremity of
the valvae seems to be the potentially most diagnostic part
of the genitalia at the specic level. Unfortunately, it is
subject to a wide range of variation regarding the presence,
location and development of the inner spiny process and
the membranous part of the valva, some specimens exhib-
iting dierent distal shapes for each valvae (Figs. 10a-10l.).
Nevertheless, although for some species we did not man-
age to detect specic genitalia characters that dierentiate
them from others, we have been able to discern the basal
genitalia pattern for each species. Our conclusions were
limited by the number of specimens available for dissec-
tion for each taxon. Genitalia morphology variation and
basal genitalia pattern will be discussed under each taxon
Another crucial point is the use of a standardized meth-
od to reliably compare male genitalia morphology. e an-
gle of view is of foremost importance, as even a little varia-
tion of a few degrees can result in a completely dierent
appreciation of the valva morphology. A tri-dimensional
comparison is essential to appreciate subtle morphological
dierences. Illustrations of two-dimensional photographs
to represent genitalia are seriously limited as a source of
information. We do insist on the fact that, when studying
closely related species, dierences in male genitalia be-
tween two or more species must be conrmed by the dis-
section of several specimens, and if possible, examination
and comparison by the same technician.
Inner margin supernumerary ocellus on VHW (Figs. 11)
e inner margin ocellus on VHW is a diagnostic
character for the genus Magneuptychia (sensu Costa et al.,
2016), which comprises at least 3 species: M. libye (Lin-
naeus, 1767), M. libyoidea (Butler, 1866) and M. lethra
(Möschler, 1883). During our analyses, we found some in-
dividuals expressing this ocellus in species that normally do
not exhibit it. Here we illustrate some examples from dif-
ferent genera: ‘Magneuptychia’ tricolor (Hewitson, 1850),
Megisto cymela (Cramer, 1777), Modestia remypignoux sp.
nov. and Euptychia marceli Brévignon, 2005 (Figs.11a-
11d, 11g-11h). By way of contrast, we have also illustrated
a male specimen of Magneuptychia libye from Tobago, in
which this ocellus is absent when it should normally be
present (Figs. 11e-11f). e taxon Euptychia benedicta
described by Butler (1877) is apparently based on a single
female specimen from Ecuador, which was appropriately
illustrated in its original description and deposited in the
NHMUK (Fig. 12d.). is specimen is similar to the lec-
totype of Neonympha alcinoe C. Felder & R. Felder, 1867
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
23
(see under this species) but presents an additional ocellus
on the inner margin of VHW. Although we did not nd
another similar specimen which exhibit this additional
ocellus, we think that it represents no more than a varia-
tion (see discussion in M. alcinoe comb. nov.) as it can oc-
casionally occur in the above mentioned species. e pres-
ence of this ocellus, almost totally neglected in previous
comparative studies of buttery wing patterns, probably
represents an ancestral trait still present in Magneupty-
chia and some other species of the Neotropical Satyrinae,
like Euptychia attenboroughi Neild, Nakahara, Fratello &
Le Crom, 2015, E. sophiae Zacca, Nakahara, Dolibaina
& Dias, Vanima lesbia (Staudinger, 1886) and V. palladia
(Butler, 1867), among others.
Figure 10. Male genitalia dorsal view showing intraspecic variation and variabilty within the same specimen. a. Malaveria nebulosa
comb. nov. (MB-0004); b. M. nebulosa comb. nov. (MB-006); c. Malaveria rodriguezi risaralda ssp. nov. (MB-013); d. Malaveria sp.
(MB-014); e. Modestia remypignoux sp. nov. (MB-0039); f. Modestia remypignoux shueyi ssp. nov. (MB-0213); g. Malaveria mimas
comb. nov. (MB-0010); h.M. mimas comb. nov. (MB-0014); i. Malaveria ballo sp. nov. (MB-0072); j. M. ballo sp. nov. (MB-
0073); k. Malaveria duponti sp. nov. (MB-0145); l. M. duponti sp. nov. (MB-0146).
Figure 11. Additional inner margin ocellus on VHW in some species of Euptychiina. a. “Magneuptychia” tricolor; b. “M.” tricolor;
c.Megisto cymela; d. Modestia remypignoux sp. nov.; e. Magneuptychia libye; f. M. libye; g. Euptychia marceli; h. Modestia remypignoux
sp. nov. (Scale: 1 cm).
M. Benmesbah, Á. L. Viloria & J. Murienne
24
DNA analysis and results (Tables 1-3).
General comparison for Modestia gen. nov., Malaveria
gen. nov. and Koutalina gen. nov
We obtained 16S sequences for 101 specimens (of
which only 17 were excluded in the following tables to
avoid space-consuming repetitions of the same sequenc-
es), with a fragment size ranging from 218 to 220 base
pairs. e nal alignment comprised 231 sites among
which 71.43% were invariant. e marker (16S) shows
expected patterns of genetic distances, with intra-specic
distances typically above 97% divergence (i.e., M. r. shueyi
ssp. nov. / M. r. remypignoux sp. nov. at 98.6% similarity).
Genetic similarity among closely related species is below
97% for M. gomezi comb. nov. / M. modesta comb. nov.
(95.9%), M. modesta comb. nov. / M. r. remypignoux sp.
nov. (94.1%), M. mimas comb. nov. / M. duponti sp. nov.
(94.1%). In the same fashion, comparison of inter-generic
distances shows even lower similarity for Cissia (sensu
stricto)/ Koutalina gen. nov. (92.8%), Malaveria gen.
nov. / Koutalina gen. nov. (89.1%), Modestia gen. nov. /
Magneuptychia (93.2%) and Megeuptychia Forster, 1964
/ Euptychoides (88.7%). However, there are also a few in-
stances in the genus Malaveria gen. nov., where the 16S
fragment could not discriminate between closely related
species. Indeed, M. mimula comb. nov., and M. duponti
sp. nov., as well as M. r. rodriguezi sp. nov., and M. bottoi
sp. nov., show exactly the same sequence. In addition, the
cluster M. ballo sp. nov., M. r. rodriguezi sp. nov. / M.
bottoi sp. nov., despite showing few dierences, present a
very high genetic similarity of 99.5%. While a phenom-
enon of introgression is possible, the lack of resolution of
the 16S marker is most probably related to the relatively
small size of the fragment.
SPECIES ACCOUNT
Modestia Viloria & Benmesbah, gen. nov.
Modestia modesta (Butler, 1867), comb. nov.
(Figs. 3a-3d, 12a, 13a-13m)
Type material: Butler described Euptychia modesta
based on an unspecied number of specimens from Pará
(eastern Amazonian Brazil) in the Bates collection, and
provided an illustration. A male syntype in good condi-
tion, which agrees with the original description, is hosted
in the NHMUK (Fig. 12a). To x the identity of this tax-
on, to stabilize its taxonomy and to avoid any further con-
fusion in this group of very similar species, we designate
this syntype as the L of Euptychia modesta: /
Type H.T./Syntype/♂/Type of Species./ Godman-Salvin
Coll. 1904.--1./Euptychia modesta, Butl./♂ Pará modesta
Butler type / Para, L. Amazons. H. W. Bates. / B. M. TYPE
N°. Rh 3222 Euptychia modesta, ♂ Butl./BMNH(E)
#986581/(examined) [NHMUK].
Material examined: 6 males (6 dissected), 13 fe-
males(5 dissected). FRENCH GUIANA (6 males, 13 fe-
males): Grand-Santi: 16.V.2012 1 female (MB-0036*);
Mitaraka: 01.III.2015 1 female (MB-0026*); Mont
Itoupé: 28.XI.2014 1 male (MB-0024*); 27.XI.2014
1 female (MB-0027); 30.XI.2014 1 female (MB-
0030); 02.XII.2014 1 female (MB-0035); Papaïchton:
23.VII.20122 males (MB-0023*, MB-0025*); 23.VII.2012
1 female (MB-0029); Saint-Jean du Maroni: 02.VII.2015
1 female (MB-0038*); Saint-Laurent du Maroni:10.I.2012
1 female (MB-0031*); Saül: 12.IX.2013 1 male (MB-
0020*); 03.IX.2011 1 male (MB-0021*); 30.IX.2012
1 male (MB-0022*); 12.IX.2013 1 female (MB-0028);
11.IX.2013 1 female (MB-0032*); 07.VIII.2012 1 female
(MB-0034); 11.IX.2013 1 female (MB-0033); 07.X.2011
1 female (MB-0037) [MB].
Redescription: Male (Figs. 13a-13b)– FWL: 21-
22mm (6 specimens), average 21.5 mm; DFW: ground
color brown, almost uniform; VFW: ground color brown;
submedian and median lines thin, dark brown, slightly
ferruginous; submarginal band dark brown, discreet;
one small black ocellus at the apex in M1-M2 with one
tiny white pupil, two others reduced and almost invis-
ible in M2-M3 and M3-Cu1; submarginal line thinner,
scalloped, pointing distally in R5 to 2A; marginal line
straight, thin, parallel and close to outer margin. DHW:
ground color uniform, same as DFW; outer margin slight-
ly scalloped; thin marginal area outside marginal line more
clear near tornus. VHW: ground color same as VFW;
submedian and median lines of same width and color as
in VFW, slightly irregular, joining anal margin without
making a curve; submarginal line scalloped, composed
by curved adjacent segments, basally convex from M1 to
2A; marginal dark line thin, parallel to outer margin from
costa to tornus; ve ocelli surrounded with discreet dark
brown in submarginal area, one in each space between Rs
and Cu2,
,
ocelli in Rs-M1, M1-M2 and Cu1-Cu2 round-
ed, well dened, black circled with yellow ring, ocellus in
Cu1- Cu2 larger,
with two white pupils, ocelli in Rs-M1
and M1-M2 smaller with one white pupil, ocelli in M2-
M3 and M3-Cu1 less dened, lacking black central area
and pupils.
Female (Figs. 13c-13d) – FWL: 20-22 mm (12 speci-
mens), average 20,8 mm. Similar to male, but diers by
underside color lighter and greyish, ocelli in M2-M3 and
M3-Cu1 on VFW and VHW well dened, each with two
white pupils.
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
25
Table 3. Data of the specimens of several satyrine taxa selected and sampled for 16S mitochondrial rDNA sequencing in this study (genera Cissia, Euptychoides,
Magneuptychia, Megeuptychia, Modestia gen. nov., Malaveria gen. nov. and Koutalina gen. nov.). Individual codes generated for each voucher specimen in rst
column, GenBank accession codes in fourth column.
M. Benmesbah, Á. L. Viloria & J. Murienne
26
Table 3. (Continuation)
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
27
Male genitalia (6 dissected) (Figs. 13e-13k): char-
acterized by the shape of the valvae, which are long and
gradually pointed distally. In lateral view the valvae have a
particular ampullar ridge (dorsal) with a moderate notch,
just before their tip. is is not seen in other species of the
genus. Uncus lanceolate, twice as long as tegumen; saccus
digitiform, same length as subunci; aedeagus long, about
same length as tegumen + uncus.
Female genitalia (5 dissected) (Figs. 13l-13m): exter-
nal sclerites of the 8th abdominal segment (ventral) very
similar to those of M. remypignoux sp. nov.; papillae anales
prominent and setose, bursa copulatrix bearing two paral-
lel signa, almost as long as the corpus bursae.
Variation: there is slight variation on VHW inthe size
of the ocelli and the distance between submarginal and
marginal lines, but no consistent dierence has been no-
ticed.
Diagnosis: M. modesta comb. nov., diers from M. go-
mezi comb. nov. (Fig. 14a-14c) by: 1. less marked median
and post median lines on the underside, which are brown
Figure 12. Historical type specimens with corresponding labels. a. Lectotype of Euptychia modesta Butler, 1867; b. Lectotype of Ne-
onympha sylvina C. Felder & R. Felder, 1867; c. Lectotype of Neonympha alcinoe C. Felder & R. Felder, 1867; d. Syntype of Euptychia
benedicta Butler, 1877. (Scale: 1 cm; labels not presented to scale).
Figure 13. Modestia modesta comb. nov., imagos and genitalia. a. Male ventral; b. idem dorsal; c. Female ventral; d. idem dorsal; e. Male
genitalia lateral view (one valva removed); f. idem dorsal view; g. Valva lateral inner view; h. idem lateral outer view; i. idem dorsal view;
j.Aedeagus dorsal view; k. idem lateral view; l. Female genitalia lateral view; m. idem ventral view. (Scale imago: 1 cm; scale genitalia: 1 mm).
M. Benmesbah, Á. L. Viloria & J. Murienne
28
and heavily marked in M. gomezi comb. nov.; 2. shape of
the ocelli in M2-M3 and M3-Cu1 on VHW, in which the
yellow ring almost invade the entire ocelli in M. gomezi
comb. nov., the brown center being reduced to only a
dash; 3. distal extremity of the valvae sharper and present-
ing an obvious inner protrusion in M. modesta comb. nov.
(Figs. 13g-13i), while it is more spatulate and with a much
less developed inner protrusion in M. gomezi comb. nov.
(Figs. 14f-14h). Dierences with M. remypignoux sp. nov.,
are treated under that species.
Habitat and behavior: M. modesta comb. nov., has
been captured several times in French Guiana and is prob-
ably more widespread. One of the syntypes is from the
Lower Amazon in Pará state of Brazil. e species seems
to y all year round. It has been encountered as well in
primary forest (Mitaraka Mounts), secondary forest
(Saint-Jean-du-Maroni), close to cassava plantations (Pa-
païchton) or close to human habitation (Saül). At Saül,
the specimens collected were ying in the late morning
at the edge of the trail, together with Cissia penelope (Fa-
bricius, 1775) and species of Hermeuptychia. Its ight is
characteristic for members of the Euptychiina: shy, in the
shade of the understorey, ying for several meters before
resting again, wings closed, at the edge of the trail. Dis-
tinction between M. modesta comb. nov. and Hermeupty-
chia sp. was very dicult on the wing and only systematic
collecting allowed the capture of a few specimens of M.
modesta comb. nov. It has also been observed during the
middle morning (between 9:30 and 10:30am) at the edg-
es of a wide bamboo eld on the slopes of Mount Itoupé
at around 630m. Several specimens were observed over
several days revealing the same behavior. e buttery
had a nervous ight at the summit of the bamboos (2 to
3m high) and rested on leaves for a few seconds to a few
minutes before ying. e specimens appeared when the
angled sunlight lit up this part of the bamboo eld, and
then stopped ying when sunshine became strong. One
specimen was collected at the summit of the same mount
(second highest summit in French Guiana at 750m) in
late evening, close to the ground when it had been ob-
viously disturbed. e species has thus been collected in
French Guiana from 0 to 750m. Nevertheless, it has been
rarely encountered and there are few specimens in recent
French Guianan collections. While reporting several spe-
cies of the Euptychiina hilltopping in the same site in Saül
(French Guiana), neither M. modesta comb. nov., nor M.
remypignoux sp. nov., have been observed among them.
ey were captured a few hundred meters from this hill-
top (Benmesbah et al. 2018).
Discussion: the specimens which we identied as M.
modesta comb. nov., from French Guiana completely
agree with the original description and are similar to the
lectotype of this species. Distance between French Guiana
and Pará state (Brazil) is not excessive and the two regions,
being biogeographically continuous, share various species
of Satyrinae (e. g., several Euptychia, Pareuptychia Forster,
1964, Emeryus, Cissia). Butler also mentions in its descrip-
tion a variety of E. modesta without naming it, with “small-
er ocelli on ventral surface” from Cameta (also a locality
in Pará) in the Bates collection. is latter variety could
refer to M. remypignoux sp. nov., which is sympatric with
modesta in French Guiana and ies also in close Pará state.
ere are two records from the Orinoco Delta in Venezu-
ela (A. Neild, pers. comm.), and we examined some other
specimens very similar to M. modesta from Venezuela,
Brazil, Peru, Colombia, Panama and Belize. ere is some
variation in the ventral color, size of the ocelli on VHW,
shape, distance to margin and width of the submarginal
line on VHW. Unfortunately, we did not have this mate-
rial to hand to compare specimens directly and appreciate
the subtle dierences that could occur. Two males (1 from
Panama and 1 from Belize) did not show consistent dif-
ferences in the male genitalia when compared with those
from French Guiana. ere are probably other similar,
cryptic species, such as M. remypignoux sp. nov., herein de-
scribed. e study of dierent populations of this widely
distributed complex is beyond the scope of this work and
would require larger number of samples, accurate locality
data, numerous dissections and DNA comparative studies.
Modestia gomezi (Singer, DeVries & Ehrlich, 1983),
comb. nov.
(Figs. 14a-14j)
Type material: H male: /Holotype/Costa
Rica/Rothschild Bequest, B.M. 1939-1/Cissia gomezi det.
P. J. DeVries/BMNH(E) #806407/; P(all from
Costa Rica): 2 males, same data as holotype; 1 female, Par-
que Nacional Corcovado, Osa Peninsula, Sirena, ex. larva
24 Sept. 1980 [NHMUK] (examined).
Redescription: Male (Figs. 14a-14b): FWL: 21-
22 mm (2 specimens), average 21.5 mm. DFW: back-
ground brown uniform. VFW: ground color light brown;
submedian and median lines thick, color ferruginous
brown; submarginal band dark brown, well dened from
costa to 2A-Cu2; only one small black ocellus circled by
a yellowish ring in the apex in M1-M2 with one small
white pupil; submarginal line dark brown thinner, scal-
loped, running from radius to inner margin, pointing
distally in M1 to 2A; marginal line straight, thinner,
parallel and close to outer margin. DHW: ground color
same as DFW; marginal and submarginal lines observ-
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
29
able through translucent wing; thin marginal area outside
marginal line lighter near tornus; ocellus Cu1-Cu2
barely
visible. VHW: ground color same as VFW; median and
submedian lines thick, ferruginous brown; median line
joining anal margin with a marked angulation; submar-
ginal line thinner, scalloped, basally convex from M1-M2
to Cu2-2A; marginal dark line thin, parallel to outer mar-
gin from costa to tornus, thicker in Cu2-2A; submarginal
area with ve ocelli, one in each space between Rs and
Cu2, which external counter touching adjacent veins,
ocellus in
Cu1-Cu2
larger, well dened with two tiny
white pupils, ocellus in M1-M2 smaller and ocellus in
M2-M3 even smaller,
well dened, centered by one white
pupil, ocelli in M2-M3 and M3-Cu1 less dened, lacking
black central area and white pupils; all ocelli surrounded
by irregular thin brown area.
Female (Fig. 14c) – FWL: 22 mm (1 specimen). Simi-
lar to male on ventral surface (dorsal surface not exam-
ined).
Male genitalia (1 dissected) (Fig. 14d-14j): chitinized
structures of the genitalia show the characteristic features
for the genus: globular and short tegumen, long lanceolate
uncus, subunci straight, stylized, as long as the digitiform
saccus. Valvae in this species are characterized by a contin-
uous ampullar (dorsal) ridge, convex, with tiny irregular
denticles; aedeagus slightly curved upwards.
Femalegenitalia: not examined.
Variation: the specimens examined showed only slight
variation in wing pattern.
Diagnosis: M. gomezi comb. nov., diers from all oth-
er members of Modestia gen. nov., by having: 1. the wid-
est ocelli on VHW; 2. the widest submedian and median
lines on underside; 3. almost no membrane on the distal
extremity of the valvae of male genitalia which is more
spatulate than in M. modesta comb. nov. and M. remyp-
ignoux sp. nov.
Hostplant: reported by DeVries (1986) as a grass (Poa-
ceae). Early stages described by Singer et al. (1983).
Habitat and behavior: we have not observed this spe-
cies in nature. Museum records show that it ranges from
Costa Rica to Colombia through Panama.
Discussion: Singer et al. (1983) dened the “Cissia con-
fusa-species group” mainly on the basis of early instar mor-
phology and divided it into 4 subgroups. ey assigned M.
gomezi comb. nov., to one of these subgroups, of which it
was the only representative. ey pointed out the dier-
ences observed with respect to other species of the “Cissia
confusa-species group”: larval head, pupae shape, and adult
hindwing ocelli in Rs-M1 and M1-M2 with only one pu-
pil each. As discussed above, ocellar pattern is in our view
diagnostic at generic level. ese dierences support our
decision to remove gomezi from that species-group and
transfer it to Modestia gen. nov. Furthermore, DNA com-
parative analysis shows that M. gomezi is evidently closer
to M. modesta comb. nov. (95.9% similarity), than to ‘Cis-
sia’ myncea (91.4% similarity) (Tables 1 and 2).
Material examined: 3 males (1 dissected), 2 fe-
males: COLOMBIA(1 male): Antioquia: Porce 800m,
27.VIII.2017 1 male (MB-0258*) [MB]; PANAMA
(1male, 1 female): Zona del Canal: Cocolí, 06.IV.1987
1 male (photograph) and 28.XII.1986 1 female (photo-
graph) [JMD]; COSTA RICA (1 male, 1 female): 1 male
Figure 14. Modestia gomezi comb. nov., imagos and genitalia. a. Male ventral; b. idem dorsal; c. Female ventral; d. Male genitalia lateral
view; e. idem dorsal view; f. Valva lateral inner view; g. idem lateral outer view; h. idem dorsal view; i. Aedeagus dorsal view; j. idem
lateral view. (Scale imago: 1 cm; scale genitalia: 1 mm).
M. Benmesbah, Á. L. Viloria & J. Murienne
30
(holotype) no precise locality (photograph) [NHMUK];
1 female Parque Corcovado-Sirena (photograph).
Modestia remypignoux Benmesbah & Viloria, sp. nov.
(Figs. 15a-15m)
http://zoobank.org/urn:lsid:zoobank.org:act:DC4CBBA5-0DA9-4DB7-
8682-315391AAFBAD
Type material: H male (Figs. 15a-15b): /
French Guiana, Maripasoula, Antecume Pata, 13.III.2012/
Holotype of Modestia remypignoux remypignoux Benmes-
bah & Viloria, 2020/MB-0039*/ [MB]; P:
9 males (4 dissected), 13 females (3 dissected) (all from
FRENCH GUIANA): Kaw: Dégrad Escolle, 14.XI.2011
1 male (MB-0041*) [MB] and 1 female (MB-0047*) [MB];
Kaw pk 8 17.IX.2010 1 female (MB-0053*) [MB]; Grand
Santi: 16.V.2012 1 female (MB-0055*) [MB]; 18.V.2012
1 female (MB-0355) [MB]; Kourou: Montagne des Pères
22.VI.2012 1 male (MB-0042) [MB]; 31.I.2013 1 male
(MB-0040*) [MNHN]; Macouria: Matiti 18.VIII.2012
2females (MB-0049) [MB], (MB-0054) [MIZA]; Mari-
pasoula: Antecume Pata 13.III.2012 2 females (MB-
0048*, MB-0050) [MB]; Talwen 15.III.2012 1 male
(MB-0045*)[MB]; Matoury: Rochambeau 20.XII.2008 1
male [LCB]; 29.VIII.2009 1female [LCB]; Rémire: Dé-
grad des Cannes 06.XI.1988 2 males [LCB]; Saint-Lau-
rent-du-Maroni: 07.VI.1992 1male [LCB]; 24.X.2011 1
female (MB-0051) [AN]; 19.I.2012 1 male (MB-0044)
[MIZA], 1 female (MB-0052) [IVIC]; Ouanary II.2019
1 female (MB-0371) [MB]; Sinnamary: pk 17 D21 1 fe-
male 06.IX.2009 [LCB].
Description: Male (Figs. 15a-15b): FWL: Holotype
21 mm, 21–23 mm (7 specimens), average 21.9 mm.
DFW: background light brown uniform. VFW: ground
color light greyish brown; submedian and median lines
thin, ferruginous brown, median line curved outwards,
and basally concave in Cu2-2A; submarginal band thin,
dark brown, discreet; submarginal area with three small
ocelli, one black, circled by a yellowish ring in the apex
in M1-M2, with one small white pupils, two others even
smaller, less dened in M2-M3 and M3-Cu1; submar-
ginal line thinner, dark brown, scalloped, pointing dis-
tally in R5 to 2A; marginal line straight, of same width,
parallel and close to outer margin. DHW: ground color
same as DFW; marginal and submarginal lines observ-
able through translucent wing. VHW: ground color same
as VFW; median and submedian lines thin, ferruginous
brown, discreetly irregular; median line joining anal mar-
gin making a double angulation; submarginal line thin-
ner, scalloped, pointing distally in M1 to 2A; marginal
dark line thin, parallel to outer margin from costa to tor-
nus, thicker in Cu2-2A; submarginal area with ve small
ocelli almost of same size, one in each space between Rs
and Cu2, ocellus in
Cu1- Cu2
slightly bigger, well de-
ned, with two tiny white pupils, ocelli in M2-M3 and
M3-Cu1 with one or two apparent pupils, ocelli in Rs-
M1 and M1-M2 centered by one white pupil; thin brown
area surrounding all ocelli.
Figure 15. Modestia remypignoux remypignoux sp. nov., imagos and genitalia. a. Male ventral; b. idem dorsal; c. Female ventral; d. idem
dorsal; e. Male genitalia lateral view; f. idem dorsal view; g. Valva lateral inner view; h. idem lateral outer view; i. idem dorsal view; j. Ae-
deagus dorsal view; k. idem lateral view; l. Female genitalia lateral view; m. idem ventral view. (Scale imago: 1 cm; scale genitalia: 1 mm).
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
31
Female (Figs. 15c-15d) – FWL: 19–22 mm (8 speci-
mens), average 20,1 mm. Similar to male but diers from
it by the more rounded shape, underside slightly lighter
and greyish, ocelli on ventral side a little bit wider.
Male genitalia (5 dissected) (Figs. 15e-15k): very simi-
lar to that of the preceding two species, but uncus rect-
angular and regular, saccus is not as digitiform (more
conical); the valvae closer in shape to those of M. modesta
comb. nov., but with an indented ampullar ridge and lack-
ing the notch. Aedeagus straight and shorter.
Female genitalia (3 dissected) (Figs. 15l-15m): similar
to that of M. modesta comb. nov.
Diagnosis: M. remypignoux sp. nov., can be dierenti-
ated from M. modesta comb. nov., by the following char-
acters: 1. ocelli on VHW almost of equal size and smaller
than in M. modesta comb. nov.; 2. submarginal line more
distant from marginal line on VFW and VHW than in M.
modesta comb. nov.; 3. median line on VHW making a
double angulation when joining the anal margin, whereas
it is almost straight in M. modesta comb. nov.; 4. uncus
rectangular and regular instead of tapered in M. modesta
comb. nov.; 5. distal extremity of the valvae more regular,
with less membrane than in M. modesta comb. nov. M. re-
mypignoux sp. nov., is dierentiated from M. remypignoux
shueyi ssp. nov., under this taxon. For dierences with M.
gomezi comb. nov., see under that species.
Etymology: we name this species in honor of our friend
Doctor Rémy Pignoux, French physician and naturalist in
love with nature in general and birds in particular, who has
been living in French Guiana for decades, still providing
constant devoted care to the populations of the Maroni
river and instilling great ethical values to his young doc-
toral students. Modestia remypignoux is a masculine noun
in apposition.
Hostplant: unknown to us.
Habitat and behavior: M. remypignoux sp. nov., was
observed on several occasions in French Guiana. Its be-
havior is typical of the Euptychiina (see under M. modesta
comb. nov.). e species is widespread all over the country
and present in dierent types of habitat, from primary to
secondary forest, near human habitation and patches of
low forest within savannah. M. remypignoux sp. nov., is
described from French Guiana, but it is likely to be more
widespread in similar habitats in the Amazonian region.
Discussion: M. remypignoux sp. nov., is sympatric with
M. modesta comb. nov., in French Guiana. is species has
been collected before and was known to local entomolo-
gists. Brévignon (2008) who provided important knowl-
edge on the French Guianan buttery fauna identied this
species as Magneuptychia modesta and provided a picture
of the male and its genitalia. At that time Brévignon (pers.
comm.) considered the specimens of M. remypignoux sp.
nov., to agree with the syntype of Euptychia modesta,of
which he examined photographs. He studied a single fe-
male of M. modesta comb. nov., in the collection of Père
Barbotin captured in Saül in September 1961, but did not
notice the dierence with M. remypignoux sp. nov. In fact,
true M. modesta comb. nov., is much more rarely encoun-
tered than M. remypignoux sp. nov., in French Guiana,
and larger series of this species were only very recently col-
lected there, which allowed clearly separation of these two
species. In a more recent work, M. modesta comb. nov.,
was erroneously identied by Brévignon & Benmesbah
(2012) as Magneuptychia alcinoe.
Apart from M. modesta comb. nov., the species super-
cially closer to M. remypignoux sp. nov., is Neonympha
sylvina C. Felder & R. Felder, 1867, which was dubiously
placed in the genus Paryphthimoides (Lamas 2004). Fol-
lowing the original description, it was described from at
least two specimens (a male and a female) from Bahia
(Brazil), and no illustration was provided by the authors.
A single syntype (probably a male) is known, hosted in the
NHMUK (Fig. 12b). Because of the possible presence of
at least 2 syntypes and the close habitus shared by dier-
ent taxa in this group, for the purpose of nomenclatural
stability, we x here the identity of Neonympha sylvina
and designate as L of this taxon the specimen
with the following labels (Fig. 12b): /Neonympha sylvina
Feld./Bahia Luschnatt type/FELDER COLL
n
/Type/
Type of N. sylvina Feld, very near E. modesta. Butl/Sylvina
n./Rothschild Bequest B.M.1939-1/ [NHMUK]. We did
not have access to this specimen to perform dissections.
To date we are not aware of the existence of another syn-
type. In the original description the male presents ventral
surface ocelli with one pupil, and this corresponds with
the syntype which presents a unique pupil in ocellus Cu1-
Cu2 on VHW. On the other hand, the female description
mentions a trace on DHW (presumably in the tornus) of
a bipupillate ocellus, which should correspond to a VHW
ocellus in Cu1-Cu2 with two pupils. As discussed above
the number of pupils appears to be a discriminating su-
praspecic character not just in Modestia gen. nov. and
Malaveria gen. nov., but also in other genera of the Eupty-
chiina. Plausible morphogenic mechanisms leading some
specimens to present a single pupilled VHW ocellus in
Cu1-Cu2 are also discussed above (Figs. 9a-9f). Actually,
some specimens (of both sexes) of N. sylvina from north-
eastern and mid-western Brazil presenting single pupils
have been found at DZUP and ZUEC collections. ese
specimens agree with the lectotype ofN. sylvina (T. Zacca
pers. comm.). Although it is possible that the lectotype of
N. sylvina represents an aberrant individual (i.e., with sin-
M. Benmesbah, Á. L. Viloria & J. Murienne
32
gle pupil in the Cu1-Cu2 VHW ocellus), it is more likely
to be a valid species, probably belonging to a dierent ge-
nus. A deeper study of this taxon (Zacca et al. in prep.)
should conrm this hypothesis.
Notwithstanding, the examination of the lectotype of
N. sylvina and three additional specimens (2 males and
1female, courtesy of T. Zacca) shows at least three addi-
tional dierences compared with M. remypignoux sp. nov.:
1. ocelli in M3-Cu1 and Cu1-Cu2 on VHW are much
less dened in N. sylvina than in M. remypignoux sp. nov.;
2.ocellus in M3-Cu1 is more ovoid in N. sylvina than in M.
remypignoux sp. nov.; 3. median line on VFW is straight
in Cu2-2A in N. sylvina and neither curved outwards nor
basally concave as in M. remypignoux sp. nov. Finally, N.
sylvina is described from Bahia, which is more than 2,500
km from French Guiana, and both territories represent
two dierent areas of endemism for butteries, compris-
ing various dierent species of the subtribe Euptychiina
(for instance, Euptychia atlantica Nakahara & Freitas [in
Nakahara et al. 2017], Godartiana amadoi Paluch, Zacca
& Freitas [in Zacca et al. 2016], for the Bahia region; T.
Zacca pers. comm.). Based on what precedes we treat M.
remypignoux sp. nov., as a dierent taxon from N. sylvina.
We examined several similar specimens from various
locations in Brazil, Peru and Ecuador. We noticed some
consistent dierences, although subtle, in wing pattern
and genitalia. e study of these specimens is beyond the
scope of this paper and the only other taxon we describe
here is a subspecies of M. remypignoux sp. nov., from Cen-
tral America. More comprehensive morphological studies
and molecular analyses are likely to reveal several cryptic
species (Zacca et al. in prep.).
Modestia remypignoux shueyi Benmesbah & Viloria,
ssp. nov.
(Figs. 16a-16i)
http://zoobank.org/ urn:lsid:zoobank.org:act:99ACCD73-0E4E-4FA9-
B3F1-33FE91C4C7DC
Type material: H male (Figs. 16a-16b): /
Belize, Cayo Dist., Baldy Beacon- 3,5km SSW - 1F5231
logging road in pine/oak, 750m, 23.IX.2016, J. Shuey
- P. Labus, coll./Holotype of Modestia remypignoux
shueyi Benmesbah & Viloria, 2020/MB-0143/ [MB];
P (8 males, all from Belize): same data 1 male
(MB-0144) [MB] and 4 males [JS]; same data 700m,
24.IX.2016 1 male [JS]; same data 750m, 24.IX.2016
2males [JS].
Material examined (not included in the type series):
6 males (3 dissected).- PANAMA: Colón: Cerro Santa
Rita, 225m, 08.I.1987 1 male (MB-0216*) [MB] and
1 male (photograph) [JMD]; 245m, 18.II.2014 1 male
(MB-0213); Cocle: El Valle, 600m, 22.VIII.2015 1 male
(photograph) [JMD]; 800-850m, 02.IV.1987 1 male
(MB-0214*) [MB]; Gatun, West Creek trail, 30.XII.1985
1 male (MB-0215*) [MB].
Description: Male (Figs. 16a-16b): FWL: Holotype
22 mm, 21–22 mm (2 specimens), average 21.5 mm. Male
similar to M. remypignoux sp. nov., but diers in having a
darker ground color on both sides and ocelli M2-M3 and
M3-Cu1 on VHW less dened, lacking the black central
area and the pupils.
Female: not examined.
Male genitalia (5 dissected) (Figs. 16c-16i): similar to
M. remypignoux sp. nov., in the 2 specimens examined.
Female genitalia: not examined.
Variation: there is slight variation in the size of the
ocelli on VHW in the specimens examined.
Diagnosis: subspecies shueyi is very similar to the nom-
inal subspecies, nevertheless it diers from it by the darker
ground color and the less dened ocelli on VHW in M2-
M3 (which is elongated) and M3-Cu1 (well dened black
ocelli centered by white pupil in M. r. remypignoux sp.
nov.). For dierences with M. modesta comb. nov., see
under M. r. remypignoux sp. nov. For dierences with M.
gomezi comb. nov., see under that species.
Etymology: dedicated to John Shuey, a North-Ameri-
can entomologist who is passionately keen on the Hespe-
riidae and buttery fauna of Belize. He collected all the
specimens of the type series. e Latinized subspecies
name shueyiis a masculine noun in the genitive case.
Host plant: unknown.
Habitat and behavior: comments regarding habitat
and behavior are directly quoted from J. Shuey who col-
lected all the specimens of the type series. “e locality
is located at ~ 16
o
58’ 37” x 88
o
46’ 34” and is a narrow
ridgeline that descends from approximately 900m altitude
to 600m. In totality, this ridgeline supports a complex
ecotonal habitat, transitioning from short-statured, open
grassland at 900m to pine dominated forest on eastern fac-
ing ridge lines and steep slopes. Immediately downslope,
riparian broadleaf shrubs line the adjacent stream bot-
toms and adjacent slopes.A dirt logging road follows the
ridgeline southward, ultimately descending into broadleaf
rainforest in the Sibun River Valley 4km to the south.At
the type locality, the ridge line is very narrow and supports
pine woodlands with riparian scrub immediately down-
slope along stream bottoms. is species is very localized
in Belize, and known only from the type locality. Even
here, they were found at only two places along the trail in
two days collecting, and were common at only one of the
sites. Only males were observed, ying very rapidly upslope
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
33
through dense vegetation, and then briey along the ridge-
line before descending back downslope. ey appeared
to be “hill topping” in search of mates and they were not
observed resting on any vegetation. Because of their rapid
ight and because they y through, not around, dense veg-
etation, specimens were dicult to capture. However, they
generally followed set paths along the road edge that al-
lowed a few individuals to be netted while in ight.”
We have records from Belize and Panama suggesting
this subspecies probably ranges through a great part of
Central America.
Discussion: the characters that dierentiate the
two subspecies are stable in the specimens we examined
(16specimens of shueyi ssp. nov. and 22 of ssp. remyp-
ignoux sp. nov.). Although the study of male genitalia
did not evidence dierences, DNA comparative analysis
showed the two specimens from Belize to slightly group
apart from those of M. r. remypignoux sp. nov. with 98.6%
similarity (Tables 1-2). is is likely to agree with a sub-
specic genetical distance. Finally, important geographi-
cal distance separate these two taxa belonging to two dif-
ferent areas of endemism for butteries. For these reasons
we treat this population as a subspecies of M. remypignoux
sp. nov. Specimens from Belize represent the westernmost
sample we could examine. Six other specimens from Pana-
ma (3 dissected) similar to shueyi ssp. nov., were also iden-
tied as this subspecies. We did not include them in the
type series because of the distance separating them from
the type locality. As it was said for the nominal subspecies,
we examined various specimens from Brazil, Peru and Ec-
uador that probably represent undescribed new taxa.
Malaveria Viloria & Benmesbah, gen. nov.
Malaveria nebulosa (Butler, 1867), comb. nov.
(Figs. 17a-17l)
Type material: L male: /Venezuela/Ven-
ezuela Pur. from Dyson 47-9/[NHMUK] (examined).
Material examined: 92 males (8 dissected), 46 fe-
males (2 dissected).- VENEZUELA, Miranda: Altos
de Pipe, IVIC 1,650m 22.VI.2012 1 male (MB-0338*);
18.IX.2013 2 males (MB-0336*, MB-0341*); 19.IX.2013
1 male (MB-0333*); 25.X.2017 1 male (MB-0339) and
21.V.2012 1 female (MB-0337); 22.VI.2012 1 female
(MB-0332); 31.X.2012 1 female (MB-0331); Guaicai-
puro, El Jarillo, uebrada Honda 1,300m, 23.VII.2013
2 males (MB-0002, MB-0006*); 1,700m, 23.VII.2013
1 male (MB-0003); 05.VIII.2013 1 male (MB-0004*);
1,200m 12.VIII.2013 1 male (MB-0334*); 26.VIII.2013
1 male (MB-0007*) and 05.III.2013 3 females (MB-0335,
MB-0340, MB-0342); 06.III.2013 2females (MB-0001*,
MB-0005*)[MB]; Altos de Pipe, IVIC 1,650m 30.VI.2011
2 females; 30.VII.2011 1 female; 22.VI.2012 2 males,
2 females; 29.VI.2012 1 male, 2 females; 30.VI.2012
1 female; 30.VII.2012 2 males, 1 female; 31.VII.2012
1 male; 23.VIII.2012 1 male; 24.VIII.2012 3 males,
5 females; 25.VIII.2012 1 male; 31.VIII.2012 1 male;
Figure 16. Modestia remypignoux shueyi ssp. nov., imagos and genitalia. a. Male ventral; b. idem dorsal; c. Male genitalia lateral view
(one valva removed); d. idem dorsal view; e. Valva lateral inner view; f. idem lateral outer view; g. idem dorsal view; h. Aedeagus dorsal
view; i. idem lateral view. (Scale imago: 1 cm; scale genitalia: 1 mm).
M. Benmesbah, Á. L. Viloria & J. Murienne
34
11.IX.2012 1 male; 16.IX.2012 2 males; 17.IX.2012
3males; 29.IX.2012 2 males, 1 female; 15.X.2012 1male;
17.X.2012 1 male; 19.XI.2012 1 male; 21.XI.2012
3 males; 22.XI.2012 2 males; 23.XI.2012 5males, 2 fe-
males; 24.XI.2012 2 males; 6.XII.2012 2 males; 7.XII.2012
1 male; 8.XII.2012 1 male; 15.I.2013 1 male, 1 female;
16.I.2013 1 male, 1 female; 17.I.2013 2 males; 9.II.2013
2 males; 10.II.2013 2 males; 16.II.2013 1 male, 2females;
17.II.2013 2 males; 17.IV.2013 2males; Guaicaipuro, El
Jarillo, uebrada Honda 1,300m, 6.VIII.2013 18 males,
4females; 3.IV.2013 2 males, 1 female; 30.V.2013 3 males;
22.VII.2013 4 males, 3 females; 23.VII.2013 3 females;
26.VIII.2013 2 females; 14.IX.2013 1 male; 15.IX.2013
1 male; 30.X.2013 4 females; 31.X.2013 1 female [IVIC].
Redescription: Male (Figs. 17a-17b): FWL: 20-22
mm (11 specimens), average 21 mm. DFW: background
dark brown uniform. VFW: ground color dark brown;
submedian and median lines thin, color darker ferrugi-
nous brown, median line curved outwards in Cu2-2A,
submedian line slightly wavy; submarginal area with one
small black ocellus in the apex, circled by a yellowish ring
in M1-M2 with two tiny white pupils; submarginal line of
same width than median line, more scalloped at the apex,
pointing distally in R5 to 2A; marginal dark line thin
and straight, parallel and close to outer margin. DHW:
ground color same as DFW; marginal and submarginal
lines observable through translucent wing. VHW: ground
color same as VFW; median and submedian lines of same
width and color as on VFW, submedian line a little irregu-
lar, median line more irregular, curved to the base when
passing through the cell, making a slight angulation when
joining anal margin; submarginal line of same width and
color, scalloped, composed by straight adjacent segments,
pointing distally in M1 to 2A; marginal dark line thin,
scalloped, parallel to outer margin from costa to tornus;
submarginal area with ve ocelli, one in each space be-
tween Rs and Cu2, ocelli in
M1-M2 and Cu1-Cu2 larger,
of same size, black circled by a yellowish ring, each with
two small white pupils, ocelli in M2-M3 and M3-Cu1
smaller, without black in the center, less dened, ocellus in
Rs-M1 very small.
Female (Figs. 17c-17d) – FWL: 19-21 mm (8 speci-
mens), average 20,4 mm. Similar to male but diers from
it by the more rounded shape, both sides slightly lighter.
Male genitalia (8 dissected) (Figs. 17e-17k): the entire
structure robust; tegumen globular, well dierentiated
from uncus, which is one and a half times longer than tegu-
men, lanceolate, but shorter that in the species of Modestia
gen. nov.; subunci about same length as tegumen, slightly
sinuous in upper view; saccus tubular, broad at base in the
joint with vinculum; valvae well developed, elongated,
ending in a digitiform extreme, slightly hooked, with a
prominent horn-like process at tip and another one half
way the narrowing of the valva, both pointing inwards; ae-
deagus as long as valva, as thick as saccus.
Female genitalia (2 dissected) (Fig. 17l): externally
similar to that of the species of Modestia gen. nov., papil-
lae anales well developed, setose; corpus bursae bearing
two very well developed signa along three quarters of its
length.
Figure 17. Malaveria nebulosa comb. nov., imagos and genitalia. a. Male ventral; b. idem dorsal; c. Female ventral; d. idem dorsal;
e.Male genitalia lateral view (one valva removed); f. idem dorsal view; g. Valva lateral inner view; h. idem lateral outer view; i. idem
dorsal view; j. Aedeagus dorsal view; k. idem lateral view; l. Female genitalia lateral view. (Scale imago: 1 cm; scale genitalia: 1 mm).
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
35
Variation: all specimens examined show very stable
wing pattern.
Diagnosis: M. nebulosa comb. nov., diers from the
other species treated here (except M. duponti sp. nov.,
for which character one is dierent) by: 1. darker ground
color; 2. tiny ocelli on VHW; 3. markedly irregular wavy
median line on VHW, typically curved basally when pass-
ing through the discal cell; 4. distal extremity of the valvae
thin without membranous extension, presence of a spiny
proximal inner projection on the distal portion of the val-
vae. For dierences with M. duponti sp. nov., see under
this species.
Hostplant: unknown.
Habitat and behavior: similar to other euptychiines of
the same size (see M. modesta comb. nov.). is species is
restricted to the Cordillera de la Costa in northern Ven-
ezuela.
Discussion: Nakahara et al. (2015) discuss the taxo-
nomic status of M. nebulosa comb. nov., and although
they compared it to dierent species then identied un-
der Magneuptychia alcinoe, we agree with their ideas. ey
treated Magneuptychia nebulosa as a bona de species,
comparing it with the assumed variable M. alcinoe and M.
mimas, and they designated the lectotype for this species.
In fact, as this study shows, several dierent species have
been considered as M. alcinoe (see under Malaveria alcinoe
comb. nov.) and represent a complex group of dierent
species. All specimens examined of M. nebulosa comb.
nov., showed a remarkably stable habitus with almost no
variation. Being an endemic to the Cordillera de la Costa
in northern Venezuela, it is geographically isolated from
other species of Malaveria gen. nov. (Nakahara et al.
2015), and is the only representative of this genus in this
area (Viloria pers. obs.). e closest record of a member
of the “alcinoe-complex” is in Lara state in the northern
part of the Cordillera de Mérida, and corresponds to M.
alcinoe comb. nov., where M. nebulosa comb. nov., has
not been recorded. Furthermore, in our DNA compara-
tive analysis, the 6 specimens of M. nebulosa comb. nov.,
clearly grouped apart from the other species of the genus.
e wavy median line and the tiny ocelli on VHW of M.
nebulosa comb. nov., are reminiscent of the geographically
distant M. duponti sp. nov. (see under this taxon), but the
2taxa seem not to be so close, as their molecular divergence
is 6.8%. In fact, the similar shape of the distal extremity of
the valva and the closer similarity ratio in the molecular
fragment analyzed (98.6%) support the hypothesis that
nebulosa and alcinoe are closer to each other than to the re-
maining species of the “alcinoe-complex”. Finally the FW
venation of M. nebulosa comb. nov., presents a particular
character also shared by M. alcinoe comb. nov., and not
encountered in the other species of Malaveriagen. nov.:
the origin of R2 is distal to the bifurcation between r2-
5-m1 and r1-r2 (Fig. 4).
Malaveria mimas (Godman, 1905), comb. nov.
(Figs. 18a-18i)
Type material: S Male: /Type H.T./♂/Type of
Species./Coroico. 6500., Bolivia. Garlepp./B.M. TYPE
No. Rh3225. Euptychia mimas, Godm./B.M.(N.H)
Rhopalocera Slide No. 16843./T.G.H. 1953. 16./God-
man-Salvin Coll. 1904.-1. Euptychia mimas, Godm./
BMNH(E) #983007/ [NHMUK] (examined).
Material examined: 6 males (6 genitalia dissected*).-
BOLIVIA (4 males): La Paz: Caranavi 4 males (MB-
0010*, MB-0011*, MB-0012*, MB-0013*)[MB].- PERU
(2 males): Cusco: Cusco, carretera Manu 800-2900m,
October 2015 1 male (MB-0014*) [MB]; 2000-3000m,
1908 1 male (MB-12*) [MNHN].
Redescription: Male (Figs. 18a-18b): FWL: 22-25
mm (6 specimens), average 23.5 mm. DFW: background
brown uniform. VFW: ground color brown, lighter than
upperside; submedian and median lines very thin, darker
brown, curved outwards in Cu2-2A; submarginal band
dark brown, discreet; submarginal area with one small
black ocellus in the apex in M1-M2, circled by a brown-
ish ring with two tiny white pupils; submarginal line of
same width as median line, more scalloped at the apex
and attened from M3 to 2A, pointing distally in M2
and M3; marginal dark line thinner and regular, parallel
and close to outer margin. DHW: ground color same as
DFW; marginal and submarginal lines observable through
translucent wing. VHW: ground color same as VFW;
median and submedian lines of same width and color as
VFW, submedian line making an angulation when join-
ing the anal margin, median line regular joining anal mar-
gin without making angulation; submarginal line of same
width and color, scalloped, composed by straight adjacent
segments, pointing distally from M1 to 2A; marginal dark
line thinner, discreetly scalloped, parallel to outer margin
from costa to tornus; submarginal area with ve ocelli,
one in each space between Rs and Cu2, ocelli in
M1-M2
and Cu1-Cu2 of same size, well dened, black, circled by
a brownish ring, each with two small white pupils, ocelli
in M2-M3 and M3-Cu1 of almost same size, less dened
without black in the center, ocellus in Rs-M1 very small.
Female: not examined.
Male genitalia (6 dissected) (Figs. 18c-18i): Similar to
that of M. nebulosa comb. nov., but base of saccus more
robust; strong narrowing of the valva in its middle part on
lateral view; inner horn-like processes of the valvae more
M. Benmesbah, Á. L. Viloria & J. Murienne
36
developed and prominent; aedeagus thicker and slightly
sinuous in lateral view.
Female genitalia: not examined.
Variation: the 6 specimens examined did not show
signicant variation in wing pattern. Dierences in color-
ation are discussed below.
Diagnosis: M. mimas comb. nov., diers from all the
other species treated here by: 1. background brown color
lighter, almost coppered in specimens from Bolivia (darker
and greyish in specimens from Peru); 2. thinner and more
delicate elements of the underside pattern (bigger and
wider in most other species); 3. submarginal line on VHW
and VFW more attened and less scalloped(more irregu-
lar and scalloped in others); 4. submedian line on VHW
angled when joining anal margin(straight in all others);
5. aedeagus short and stocky (proportionally longer and
thinner in others); 6. strong narrowing of the valva in its
middle part on lateral view giving a markedly angled as-
pect to its dorsal edge.
Hostplant: unknown.
Habitat and behavior: we did not observe M. mimas
comb. nov., in the eld but its behavior is presumably
similar to that of its congeners. Our records range from
northern Bolivia to southern Peru.
Discussion: Euptychia mimas was described by God-
man in 1905 on the basis of 3 specimens from Coroico,
La Paz, Bolivia, collected either by Otto or most probably
Gustav Garlepp at 6,500 . (1,980m). An illustration is
given in the original description and a syntype is hosted in
the NHMUK. e type locality is 20 km from where our
recent specimens come from, in Bolivia. ey fully cor-
respond with the original description. e discriminating
characters in the habitus listed above that dierentiate M.
mimas comb. nov., from other species of Malaveria gen.
nov, are clearly evident in the original illustration and in
the syntype. Nakahara et al. (2015) discussed the possible
conspecicity of M. mimas with M. alcinoe. In this work
we have been able to distinguish between what those au-
thors considered to be variations of M. alcinoe comb. nov.,
and stable wing patterns that we have assigned to dierent
species (see below). We treat here M. mimas comb. nov.,
as a bona de species based on the dierences in habitus
(see diagnosis), the dierences in male genitalia (mainly
the aedeagus), wing venation (see under M. alcinoe comb.
nov.) and DNA sequence analyses in which the specimens
identied as M. mimas (3 from Bolivia and 1 from Peru)
clearly stand apart from other species of the “alcinoe-com-
plex”. From the latter point of view, the closest representa-
tive of the “alcinoe-complex” is M. ballo sp. nov., sharing
95.9% similarity, whereas in M. alcinoe comb. nov., this
is only 93.3%. Furthermore, M. mimas comb. nov., seems
to be restricted to Bolivia and southern Peru, at least. We
are not aware of the presence of M. alcinoe comb. nov.,
or close similar species in southern Peru or in Bolivia. e
ranges of the “alcinoe-complex” and M. mimas comb.
nov., seem not to overlap, but the study of more material
Figure 18. Malaveria mimas comb. nov., imagos and genitalia. a. Male ventral; b. idem dorsal; c. Male genitalia lateral view (one valva
removed); d. idem dorsal view; e. Valva lateral inner view; f. idem lateral outer view; g. idem dorsal view; h. Aedeagus dorsal view;
i.idem lateral view. (Scale imago: 1 cm; scale genitalia: 1 mm).
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
37
from these areas is needed to conrm this hypothesis. We
examined two specimens from Peru. Both are from Cuzco,
one recently collected (2015), the other, much older (col-
lected in 1908). e former diers from the specimens
we examined from Bolivia by its clearly darker coloration,
while the latter shows similar coloration. Godman in his
description mentioned the ground color (probably of the
upperside) as dark-brown and the underside paler. e
illustration seems to present a darker specimen than the
series we examined from Bolivia and the syntype hosted
in the NHMUK. It is likely that these dierences in color-
ation are simply related to the age of the specimens and do
not represent a strong diagnostic character to dierentiate
populations within the species.
Malaveria alcinoe (C. Felder & R. Felder, 1867),
comb. nov.
(Figs. 19a-19m)
Type material: Neonympha alcinoe C. Felder & R.
Felder: L male (Fig. 12c): /Type/FELDER
COLL
N
./Alcinoë Feld./Rothschild Bequest B.M. 1939-1./
Type of N. alcinoe Feld? = E. benedicta, Butl. of w. it. may
be a good local form. Comp. w. type E. benedicta Butler./
BMNH(E)1266958/[NHMUK] (examined); Euptychia
benedicta Butler: S female (Fig. 12d): /Type H.T./
Type of Species./E. benedicta Butler type./B.M. TYPE
No. Rh3227. Euptychia benedicta, ♀ Butl./Sarayacu, Ec-
uador. C. Buckley./Godman-Salvin Coll. 1904.-1. Eup-
tychia benedicta, Butl./BMNH(E)1266955/[NHMUK]
(examined).
Material examined: M. alcinoe comb. nov.,: 4 males
(1 dissected) and 4 females (1 dissected).- VENEZUELA
(2 males, 3 females):Barinas: Barinitas, La Chimenea, La
Soledad, 1450m, 24.X.2008 1 male (photograph) [DT];
Táchira: Siberia 1000m, 26/28.X.2008 1 female (pho-
tograph) [DT]; Lara: Parque Nacional Yacambú,13km
South Sanare, 15-21.I.1979 1 male (MB-0008*)
[MALUZ]; Trujillo: El Boquerón, West of Alto de Es-
cuque 1050-1100m, 17.X.1999 1 female (photograph)
[AN]; El Paramito, 7-11km South-West of Escuque
1550-1600m, 18.X.1999 1 female (photograph) [AN].-
COLOMBIA (1 male, 1 female): Cundinamarca: Bogo-
ta 1male (BMNH(E) 1205423) [NHMUK]; Colegio
430m, 15.VII.1975 1 female (MB-0009*) [MB].- ECU-
ADOR (1 male): Morona-Santiago: Parque Nacional
Sangay, San Isidro 1 male (photograph) (web site: www.
sangay.eu). Malaveria sp.: 22 males (4 dissected), 1fe-
male.- COLOMBIA (15 males): Cundinamarca: Santa
Fé de Bogota 1850 1 male (MB-14*) [MNHN]; Caldas:
Samaná, Florencia 1581m, 21.II.2020 1 male (photo-
graph) [ICRM]; Cesar:San Alberto 300m, 14.VI.2017
1male (MB-0247*) [MB]; Santander:El Hato, Serranía
de los Yariguíes 2100m, 25-28.XI.201910 males (pho-
tograph) [ICRM]; La Mesa-Santos 1500m, 28.XII.2007
1male (MB-0251*); No precise locality: 1 male (MB-11*)
[MNHN].- ECUADOR (6 males): Pastaza: Canelos
1 male (BMNH(E) 1205364) [NHMUK]; Morona-
Santiago: Parque Nacional Sangay 3 males (photograph)
(web site: www.sangay.eu); Tungurahua: Rio Machay
1700m, 04-05.VII.1993 1 male (photograph) [MGCL];
Zamora-Chinchipe: Zamora 3000-4000 1 male (BM-
NH(E) 1025422) [NHMUK].- PERU (1 male, 1 fe-
male): Madre de Dios: Parque Manu, Pakitza, 1 male
400m, 06.X.1990 (photograph), 1 female 340m,
04.X.1991 (photograph).
Redescription: Male – (Figs. 19a-19b): FWL: 21-22
mm (3 specimens), average 21.3 mm. DFW: ground color
brown uniform. VFW: ground color as upperside; sub-
median and median lines thin, darker brown, median line
slightly curved outwards in Cu2-2A; submarginal area
with one small ocellus, black circled by a yellowish ring in
the apex in M1-M2 with two tiny white pupils, two other
ocelli faded in M2-M3 and M3-Cu1; submarginal line of
same width as median line, scalloped from radius to 2A,
running from radius to inner margin, pointing distally in
M1 to Cu2; marginal dark line thinner and straight, paral-
lel to outer margin. DHW: ground color same as DFW;
marginal and submarginal lines observable through trans-
lucent wing. VHW: ground color same as VFW; median
and submedian lines regular, of same width and color as
on VFW, submedian line joining anal margin without
making angulation, median line making a curved angu-
lation when joining the anal margin; submarginal line of
same color and width, scalloped composed with almost
straight adjacent segments, pointing distally from M1
to Cu2; marginal dark line thin, parallel to outer margin
from costa to tornus; submarginal area with ve ocelli, one
in each space between Rs and Cu2, ocelli in
M1-M2 and
Cu1-Cu2 of same size, rounded, bigger, black, circled by a
yellowish ring, with two small white pupils, ocelli in M2-
M3 and M3-Cu1 smaller and less dened, without black
in the center, with two white pupils, ocellus in Rs-M1 very
small, black with two white pupils.
Female – (Figs. 19c-19d): FWL: 20-22 mm (4 speci-
mens), average 21 mm. Similar to male, diers by its more
rounded shape. e specimen illustrated presents a more
wavy median line and a lighter underside coloration.
Male genitalia (1 dissected)(Figs. 19e-19k): similar to
that of M. mimas comb. nov., but valvae clearly broader
(in lateral view) at basal half, consequently, distal half
(digitiform) more dierentiated, horn-like inner processes
M. Benmesbah, Á. L. Viloria & J. Murienne
38
similar in proportional size to those of M. nebulosa comb.
nov.; aedeagus relatively robust, but notably broad and at
at basal extremity.
Female genitalia (1 dissected) (Figs. 19l-19m): setae of
the papillae anales longer than in previous species of Ma-
laveria gen. nov.; corpus bursae apparently longer than
in M. nebulosa comb. nov., signa running parallel to each
other, reaching only three hs of the length of the corpus.
Variation: there is slight variation in the size of ocellus
Cu1-Cu2 that seems to be slightly wider in the northern
part of the range, especially in females.
Hostplant: unknown.
Habitat and behavior: we did not observe M. alcinoe,
comb. nov., in natural conditions, but its behavior is pre-
sumably typical of other similar representatives of the Eup-
tychiina, see M. modesta, comb. nov. For range distribu-
tion see under “Discussion”.
Diagnosis (Figs. 20a-20l): M. alcinoe comb. nov., is,
together with M. nebulosa comb. nov., the smallest species
in Malaveria gen. nov. ey dier from M. bottoi sp. nov.,
by: 1. more rounded shape of HW external marginin M.
bottoi sp. nov.; 2. smaller ocelli on VHW that are all of
almost same size in M. bottoi; 3. more sinuous median line
on VHW in M. bottoi sp. nov.; 4. more attened submar-
ginal line on VFW and VHW in M. bottoi sp. nov.
M. alcinoe comb. nov. diers from M. rodriguezi sp.
nov. by: 1. smaller ocelli on VHW in M. rodriguezi sp.
nov.; 2. more sinuous median line on VFW and VHW in
M. rodriguezi sp. nov.; 3. the median line on VHW joins
the anal margin more straightly and in contact with the
submarginal line in M. rodriguezi sp. nov.
M. alcinoe comb. nov. diers from M. ballo sp. nov.
by: 1. its smaller size (average FWL: 21.3 mm for M. al-
cinoe comb. nov. vs. 24.3 mm for M. ballo sp. nov.); 2. the
dark brown color in fresh specimens of M. ballo sp. nov.;
3. proportionally smaller ocelli on VHWin M. ballo sp.
nov.; 4. the median line on VFW and VHW more sinuous
in M. ballo sp. nov.; 5. the median line joining the anal
margin with a distinct angulation in M. ballo sp. nov.,
whereas it is curved in M. alcinoe comb. nov. For dier-
ences with M. nebulosa comb. nov., and M. mimas comb.
nov., see under those respective taxa.
Discussion: the taxonomic problem set by the “alcinoe-
complex” is dicult to solve. Based on dierences in the
habitus of the imagos, DNA analysis results, wing vena-
tion and biogeographical inference, we describe here three
new species belonging to this group of butteries with
very similar habitus. We hope this comparative study will
help clarify the situation and encourage other entomolo-
gists to investigate deeper the issue of species limits within
this group.
During this study we found that several dierent taxa
have been treated under alcinoe, some even belonging to
dierent genera. For example, DeVries (1987) reported al-
cinoe from Costa Rica ying up to 600m and described the
imago as presenting double pupilled ocelli on VHW, but
the specimen illustrated (pl. 41, g. 1) presents a unique
pupil in ocellus M1-M2, characteristic of Modestia gen.
nov. (in this case, probably M. r. shueyi ssp. nov.). Lamas
(2004) put E. benedicta and E. pamela in synonymy with
Magneuptychia alcinoe. We show here that pamela is clear-
ly dierent and belongs to another entity, Koutalina gen.
nov. Later, Brévignon & Benmesbah (2012) erroneously
identied true M. modesta comb. nov., as Magneuptychia
alcinoe, which is not present in French Guiana. And more
recently, Nakahara et al. (2015) considered M. alcinoe to
be variable regarding the number of pupils in ocellus M1-
M2 on VHW, possibly misidentifying specimens belong-
ing to Modestia gen. nov., as M. alcinoe. is character is
very diagnostic at the generic level, as we discuss it here.
Malaveria alcinoe comb. nov., has been thought to be a
widespread and variable species. In fact, based on the data
we have, members of the “alcinoe-complex” are predomi-
nantly inhabitants of semi-montane areas, from 700 to
2,300m elevation. A few records from lower altitudes do
exist but are all from the Andean foothills (for example M.
ballo sp. nov.). ese species seem to all be restricted to
the Andean slopes from Peru to its northern prolongation
in Colombia (Cordilleras Oriental, Central and Occiden-
tal) and Venezuela (Cordillera de Mérida). We obtained
little data from Peru and Venezuela. It seems replaced
in the south by M. mimas comb. nov., from southern
Peru (Cuzco) and Bolivia, and in the north by M. nebu-
losa comb. nov., in the Venezuelan Cordillera de la Costa.
Samples from the Sierra de Perijá and the Sierra Nevada de
Santa Marta have not been studied in detail in this work.
To date this species-group has not been recorded in the
Pantepui region of Venezuela (M. Costa pers. comm.), nor
in Brazil or the Guianas.
Colombia is the key region where most of the taxa in-
volved occurs, and a more detailed study has begun to take
place (Le Crom et al., in prep.). e diculty to identify
and understand the relationships between them are due to
several problems: 1. complexity of the topography, some
areas being isolated from each other by high mountains,
whereas others are separated by lower elevations that al-
low butteries dispersion (such as in the northernmost
tips of the Cordilleras Oriental, Central and Occidental)
(Fig.21); 2. intraspecic variability of the habitus: some-
times dicult to assess and sort out pertinent diagnostic
characters, larger series of specimens with precise locality
data are needed and they are currently not easily available;
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
39
3. overlapping of distribution ranges (Fig. 21): more than
one taxon can be found at the same place which seemingly
complicates the assessment of the discriminative value of
some habitus characters (as on the eastern slopes of the An-
des in Ecuador and the western slopes of the Cordillera Ori-
ental in Colombia); 4. poor discriminative value at specic
level of male genitalia structures; 5. few museum specimens
available from Colombia with phenotype matching that of
the type specimens of N. alcinoe and E. benedicta (see un-
der M. alcinoe comb. nov.); 6. probable lack of power of
Figure 19. Malaveria alcinoe comb. nov., imagos and genitalia. a. Male ventral; b. idem dorsal; c. Female ventral; d. idem dorsal; e. Male
genitalia lateral view (one valva removed); f. idem dorsal view; g. Valva lateral inner view; h. idem lateral outer view; i. idem dorsal view;
j.Aedeagus dorsal view; k. idem lateral view; l. Female genitalia lateral view; m. idem ventral view. (Scale imago: 1 cm; scale genitalia: 1 mm).
Figure 20. Species of Malaveria gen. nov. and related taxon ‘Paryphthimoides’ grimon, underside pattern. a. Taxonomically diagnostic
elements of the underside pattern (number references in main text); b. M. alcinoe comb. nov. (Lectotype); c. M. alcinoe comb. nov.
(Venezuela); d. M. rodriguezi rodriguezi sp. nov. (Colombia); e. M. rodriguezi risaralda ssp. nov. (Colombia); f. M. ballo sp. nov.
(Ecuador); g.M. bottoi sp. nov. (Colombia); h. M. mimas comb. nov. (Bolivia); i. M. nebulosa comb. nov. (Venezuela); j. M. duponti
sp. nov. (Bolivia); k. M. mimula comb. nov. (Paraguay); l. ‘P.’ grimon (Brazil). (Scale imago: 1 cm).
M. Benmesbah, Á. L. Viloria & J. Murienne
40
the DNA analysis in this study, due to the size of molecular
fragments used from Colombian samples. We examined
40 specimens directly or in photographs, and dissected
13males from dierent locations very likely to represent
several new taxa. Nevertheless, because of what precedes,
we feel more prudent treating this subject in a more com-
prehensive study in the future (Le Crom et al. in prep.) and
describe here only two distinctive new species M. rodri-
guezi sp. nov. (twosubspecies) and M. bottoi sp. nov., from
Colombia. In Ecuador we identied a third distinctive spe-
cies West of the Andes, M. ballo sp. nov., described here-
in. We examined other specimens from the eastern slopes
of the Andes that belong to at least two dierent taxa (one
being probably new). e northernmost record identied
as M. alcinoe comb. nov., is from the northeastern slopes of
the Cordillera de Mérida in the Parque Nacional Yacambú
(Lara state); the southernmost record is from the eastern
slopes of the Andes in Madre de Dios, Peru, and it likely
represents another undescribed species. For the same rea-
sons above mentioned we do not treat these taxa here.
We studied 82 specimens belonging to the “alcinoe-
complex”, 7 from Venezuela, 33 from Ecuador, 40 from
dierent parts of Colombia and 2 from Peru. Among
them 46 were examined directly and the other 36 stud-
ied on high quality photographs. Some of the characters
that could have initially appeared as phenotypic variations
were revealed to be suciently repetitive and stable. eir
occurrence was also coherent with biogeographical dis-
tribution and discriminating DNA analysis results. Wing
venation study was performed on a limited number of
specimens (1 or 2) and provided interesting characters at
specic level that need more investigation.
Twenty ve male genitalia have been dissected and
compared. We focused on the shape of the distal extrem-
ity of the valvae, which was expected to be diagnostic at
specic level. e only specimen of M. alcinoe comb. nov.
from Venezuela (Figs. 18a-18b) we could dissect showed a
valva more similar to that of M. nebulosa comb. nov., than
to those of other species of the “alcinoe-complex”, charac-
terized by the presence of a proximal spiny inner projec-
tion and the absence of membrane in the distal part of
the valva (Figs. 18e-18k). Nevertheless, the examination
of this single specimen is to be interpreted with caution.
Indeed, we found that all the genitalia examined were very
similar and the range of variability could not provide suf-
cient arguments by itself to distinguish all the species.
We performed DNA analysis using the ADN 16S frag-
ment of 218-220 base pairs (Table 1). e specimens sam-
pled were from north-western Ecuador (16 specimens),
western (4 specimens) and eastern (1 specimens) slopes of
the Cordillera Occidental, eastern (3 specimens) slopes of
the Cordillera Central and western slopes (3 specimens) of
the Cordillera Oriental of Colombia (Table 2). e DNA
analysis for the Venezuelan male of M. alcinoe comb. nov.,
failed but the corresponding female from Cundinamarca
(Figs. 18c-18d) surprisingly revealed to be more related to
M. nebulosa comb. nov., than to M. rodriguezi sp. nov., M.
bottoi sp. nov. and M. ballo sp. nov. is unexpected re-
sult and the similarity in the male genitalic valvae suggest
that M. alcinoe comb. nov. could be rather closer to M.
nebulosa comb. nov., than to the other species of the “al-
cinoe-complex”. e results showed that M. ballo sp. nov.,
from western Ecuador, although slightly dierent, grouped
apart, arguing for its specic status. Two specimens from
Cesar and Santander (Colombia) diered by a few basis
from the main group. eir habitus is a little distinct and
potentially they represent dierent taxa that need a deeper
study. All the other specimens did not show dierences in
the DNA fragment analyzed and corresponded to M. ro-
driguezi sp. nov. (two subspecies) and M. bottoi sp. nov.
Finally, although performed on a limited number of
specimens, the wing venation study highlighted an interest-
ing and potentially specically discriminating character: the
origin of R2 on the FW. In M. nebulosa comb. nov., and M.
alcinoe comb. nov., the latter is distal to the bifurcation be-
tween r2-5-m1 and r1-r2, whereas it is proximal to it in M.
mimas comb. nov. and M. rodriguezi sp. nov. In M. ballo
sp. nov., M. mimula comb. nov., and M. duponti sp. nov.,
R2, r2-5-m1 and r1-r2 share the same origin (Figs. 4a-4h).
e similarity in habitus, genitalia and genetic dis-
tance suggests the “alcinoe-complex” is composed of sev-
Figure 21. Map of distribution of species and subspecies of the
Malaveria “alcinoe-complex”.
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
41
eral very close sister species. Based on the examination of
these specimens and our experience in taxonomy of the
Satyrinae, we retained the following habitus morphologi-
cal characters for being useful to separate the dierent taxa
(Fig. 20a-20l). Some of these are strong and clearly dis-
criminating at the specic level: they represent dierences
in the structure pattern itself (A), while others are weaker
and aect qualitatively the pattern without modifying its
structure and composition (B):
- (1) shape of the median line on VHW: straight, sinuous
or irregular (A).
- (2) junction between median line and anal margin on
VHW: straight, slightly angled or strongly angled (A).
- (3) shape of the submarginal line on the underside: at-
tened or more scalloped (A), thin or wide (B).
- (4) background color (A and B).
- (5) proportion between ocelli in Cu1-Cu2 and M1-
M2 on VHW: same size, wider or ratio inverted (B).
- (6) shape of the median line in Cu2-2A on VFW in
some species (A).
- average size of the specimens (A and B).
- size of the ocelli on VHW: small, medium, or large (B).
Using these characters associated with biogeographical
patterns of distribution and, in some cases genitalia morphol-
ogy, venation study, and DNA similarity (genetic distances),
we identied 3 new taxa belonging to the “alcinoe-complex”.
ese taxa are described as new species because the dier-
ences observed are sucient to justify this treatment. A more
complete morphological study gathering series of samples
from various localities and a DNA analysis using longer seg-
ments of nucleotide chains and dierent genes will probably
improve the taxonomic discrimination within this group.
Cajetan and Rudolf Felder described Neonympha al-
cinoe in 1867 based on an unspecied number of speci-
mens from Nova Granada: Bogotá and Venezuela [sic].
A syntype male from Bogotá is hosted in the NHMUK
(Fig. 12c). Another male specimen in the NHMUK is
possibly a syntype from Venezuela, but we could only ex-
amine its underside. Furthermore, the original description
of alcinoe mentions both sexes. It is thus plausible that the
type series comprises at least 3 specimens (2 males and 1
female). A female syntype remains unknown to us. Be-
cause of the various similar taxa involved in the “alcinoe-
complex”, the possible inclusion of more than one taxon
under Felder’s description and for nomenclature stability
purposes, it is necessary to x the identity of this taxon.
erefore, we designate the specimen with the following
labels as the L of Neonympha alcinoe C. Felder
& R. Felder, 1867 (Fig. 12c): /Type/FELDER COLL
N
./
Alci noë Feld/Rothschild Bequest B.M. 1939-1./Type
of N. alcinoe Feld? = E. benedicta, Butl. of w. it. may be
a good local form. Comp. w. type E. benedicta. Butler./
BMNH(E)1266958/[NHMUK] (examined).
e habitus of the lectotype is characterized on VHW
by: 1. regular median line, 2. median line making a slight
angulation when reaching the anal margin at a short dis-
tance from the submarginal line, 3. ocelli of almost the
same size in M1-M2 and Cu1-Cu2, 4. submarginal line
scalloped, composed by almost straight adjacent segments,
5. relatively small size (FWL: 21 mm). We found 4 males
closely matching this habitus: 1 from Bogotá (NHMUK),
1 from Sangay National Reserve in the eastern Andes of
Ecuador (illustrated on the website www.sangay.eu) and
2 from the Cordillera de Mérida (from Barinas and Lara
states) in Venezuela. We examined 3 additional corre-
sponding females from the Cordillera de Mérida (from
Táchira and Trujillo states, Venezuela) very similar to the
female from Cundinamarca (Figs. 18c-18d). We identied
all these specimens as true alcinoe. Unfortunately, we only
had direct access to 2 of these specimens (the male from
Lara state and the female from Cundinamarca) for dissec-
tions and DNA analysis.
e potential syntype from Venezuela diers notably
from the lectotype on VHW by having the ocellus in M1-
M2 bigger than that in Cu1-Cu2 and by the much smaller
ocelli in M2-M3 and M3-Cu1. We encountered a single
specimen hosted in the MNHN, labelled “Colombia”, pre-
senting a similar habitus. e latter specimen is very old, and
for this reason the locality “Colombia” is vague and could
have applied to some part of Venezuela, or even Ecuador.
We believe this particular phenotype is likely to represent
an extreme variation of the true alcinoe pattern or possibly a
dierent taxon whose distribution area is unknown. Ocellus
in Cu1-Cu2 seems to be slightly wider in the northern part
of the distribution of M. alcinoe, especially in females.
Butler described Euptychia benedicta from an unspeci-
ed number of specimens from Sarayacu, Ecuador, and
illustrated the species. A female syntype is hosted in the
NHMUK (Fig. 12d). is specimen corresponds exactly
to Butler’s description and illustration. It has a particular
character indicated in its description: the sixth anal ocel-
lus on VHW. is is very likely to represent an aberrant
character whose presence is rarely encountered (see discus-
sion above). It is likely that Butler based his description on
this unique specimen. e location Sarayacu is in Pastaza
Province in the Amazonian lowlands of Ecuador, east of
the Andes at around 400 m altitude. A few reliable records
from specimens belonging to the “alcinoe-complex” exist
from the lowlands and thus, this altitude record may be
or may be not plausible. One of the labels mentions “C.
Buckley” who was certainly its collector. Clarence Buck-
ley, was a successful British natural history collector in the
M. Benmesbah, Á. L. Viloria & J. Murienne
42
Andes of Colombia, Ecuador and Bolivia (1868-1878),
and a notable supplier of buttery specimens to William
Chapman Hewitson in England. He collected extensively
in localities on the eastern slopes of the Ecuadorian Andes
(Vane-Wright 1991). We then assume the syntype of E.
benedicta came indeed from the Sarayacu region, a locality
also recorded in mammals and birds obtained by Buckley.
e particular characters of alcinoe, already mentioned,
are apparently shared by benedicta and we agree with La-
mas (2004) in temporarily considering benedicta as a ju-
nior synonym of alcinoe. Specimens collected by A. Neild
(pers. comm.) on the eastern slopes of the Andes of Ecua-
dor have a distinctive golden look to the ventral brown,
and probably represent a dierent, new taxon.
e type locality of alcinoe, “Bogotá”, is far from being
precise and could represent various localities in Colom-
bia. e exact distribution range of true alcinoe remains
uncertain (Fig. 21), and further collecting in the area of
Bogotá and the surrounding lower mountains is needed.
All the specimens we identied as true alcinoe are from
the eastern slopes of the Andes in Ecuador to the north-
ern tip of the Cordillera de Mérida in Venezuela, where
the lower mountains probably allow the species to ow on
both slopes. e only specimen located outside this area is
the Colombian female from Cundinamarca (again a large
department mostly covering part of the Cordillera Ori-
ental, including Bogotá) on which some doubts remains
about the precise collecting locality. Nevertheless, given
the absence of other specimens matching the lectotype in
the material examined elsewhere in Colombia, this record
requires conrmation. We hypothesize M. alcinoe comb.
nov., to be restricted to eastern slopes of the Andes of Ec-
uador, the Cordillera Oriental in Colombia and the Cor-
dillera de Mérida in Venezuela.
Although M. alcinoe comb. nov., is likely to be the only
representative of the “alcinoe-complex” in the Cordil-
lera de Mérida, it is probably sympatric with some related
taxa in Colombia and Ecuador. Comprehensive sampling
along the eastern slopes of the Andes and the regions sur-
rounding Bogotá might conrm this hypothesis.
Additional specimens examined that did not match the
species herein described are listed under Malaveria sp.,
and will be studied separately (Le Crom et al. in prep.).
Malaveria ballofi Benmesbah & Viloria, sp. nov.
(Figs. 22a-22m)
http://zoobank.org/ urn:lsid:zoobank.org:act:44958377-2D0A-4DD7-A107-
1A802A967D08
Type material: H male (Figs. 22a-22b): /EC-
UADOR, Pichincha, Nanegalito, 2016/Holotype male
of Malaveria ballo Benmesbah & Viloria, 2020/Raúl
Aldaz collector/MB-0072*/[MB]. P(all from
ECUADOR): 26 males (11 dissected), 5 females (2dis-
sected): 1 female (MB-0075*) and 9 males same data as
Holotype (MB-0017*, MB-0066, MB-0067, MB-0071,
MB-0070*, MB-0074*, MB-0076*, MB-0257*) [MB,
except 0067 in IVIC], (MB-0073*) [MIZA]; Imbabu-
ra, Parambas 3500, 1913 2 females (MB-06, MB-07*)
[MNHN]; and 2 males (MB-08*, MB-09*) [MNHN];
February 1897 2 males [NHMUK]; March 1897 2 males
[NHMUK]; 20.III.18971 male [NHMUK]; 20.III.1897
1 female [NHMUK]; Pichincha, Nanegalito, October
2015 1 male (MB-0015*) [MIZA]; 16.X.2015 2 males
(MB-0016*, MB-0018), 1 female (MB-0370) [MB]; Car-
chi, Rio Baboso 800m, IX.2016 2 males (MB-0068*, MB-
0069); Carchi, El Carmelo, VIII.2017 4 males (MB-0365)
[AN], (MB-0366, MB-0367, MB-0368) [MB].
Description: Male (Figs. 22a-22b): FWL: Holotype
25 mm, 23-26 mm (15 specimens), average 24.3 mm.
DFW: ground color dark brown uniform. VFW: ground
color as upperside; submedian and median lines thin,
brown slightly ferruginous, median line curved outwards
in Cu2-2A; submarginal area with three small ocelli, one
black circled by a yellowish ring in the apex, in M1-M2,
with two tiny white pupils, two others faded in M2-M3
and M3-Cu1; submarginal line same width as median
line, scalloped from radius to 2A, pointing distally in M1
to Cu2; marginal dark line thinner and regular, parallel to
outer margin. DHW: ground color same as DFW; mar-
ginal and submarginal lines observable through translucent
wing. VHW: ground color same as VFW; median and sub-
median line slightly irregular, of same width and color as on
VFW, submedian line joining anal margin without making
angulation, median line making a marked angulation when
joining the anal margin; submarginal line of same color but
slightly wider, scalloped, composed by almost straight adja-
cent segments, pointing distally from M1 to Cu2; marginal
dark line thin, parallel to outer margin from costa to tor-
nus; submarginal area with ve ocelli, one in each space be-
tween Rs and Cu2, ocelli in
M1-M2 and Cu1-Cu2 round-
ed, bigger, of almost same size, black, circled by a yellowish
ring, each with two small white pupils, ocelli in M2-M3
and M3-Cu1 smaller and less dened, without black in the
center, ocellus in Rs-M1 very small.
Female (Figs. 22c-22d) – FWL: 22-23 mm (3 speci-
mens), average 22,7 mm. Similar to male, but diering by
the more clear coloration, and the faded ocelli on under-
side more marked.
Male genitalia (11 dissected) (Figs. 22e-22k): general
structure similar to previous species herein described; the
valvae being more elongated and stylized than those of
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
43
M. nebulosa comb. nov., M. mimas comb. nov., and M.
alcinoe comb. nov., horn-like inner processes less evident;
base of aedeagus broad as in alcinoe.
Female genitalia (2 dissected) (Figs. 22l-22m): no sig-
nicant variation found when compared to the female
genitalia of M. nebulosa comb. nov., the type species of
this genus. Perhaps ne dissections of membranous struc-
tures would allow for detecting dierences in less evident
structures like the spermatheca (and the lagena) and the
dierent ducti.
Variability: all specimens examined are very similar
to each other, and show a slight variability in the shape of
the VHW median line, which is usually a little more wavy
than in the holotype.
Diagnosis: M. ballo sp. nov., is the larger species of
the “alcinoe-complex”. It diers from M. alcinoe comb.
nov., M. bottoi sp. nov. and M. rodriguezi sp. nov., by: 1.
its larger size (FWL average 24.3 mm for M. ballo sp.
nov., 21.0 mm for M. alcinoe comb. nov., 22.0 mm for M.
bottoi sp. nov., 22.9 mm for M. rodriguezi rodriguezi sp.
nov., 23.0 mm for M. rodriguezi risaralda ssp. nov.); 2. the
marked angle made by the median line when reaching the
anal margin on VHW; 3. median line markedly curved
outwards in Cu2-2A on VFW as a dome concave to the
base. Moreover, it diers from M. alcinoe comb. nov.,
by the more wavy median line on VHW, the presence of
membranes and the absence of the inner spiny projection
on the distal part of the valvae; from M. bottoi sp. nov., by
the darker ground color and the more angular submarginal
lines on underside.
Etymology: dedicated to our friend Daniel Ballof, a
French entomologist who is passionately keen on Neo-
tropical and African butteries, in gratitude for his gener-
osity and hospitality. He has travelled to various countries,
but especially French Guiana, where he lived for years and
contributed greatly to the better knowledge of its butter-
y fauna. e species name ballo is a Latinized masculine
noun in the genitive case.
Hostplant: unknown.
Habitat and behavior: we did not observe M. ballo
sp. nov., in the eld, but its behavior is presumably typical
of that of other members of Malaveria gen. nov. See also
Modestia modesta comb. nov. is species seems to be re-
stricted to the western slopes of the Andes in Ecuador, part
of the Chocó area of endemism (Fig. 21). We have no in-
formation on collecting records in southwestern Colom-
bia (departments of Cauca and Nariño), where it could
also occur. ere are records from sea level up to 1,400m.
Discussion: M. ballo sp. nov. is geographically iso-
lated from the other members of the “alcinoe-complex”.
Several endemic species and subspecies of butteries have
been recently described from northwestern Ecuador (Ben-
mesbah et al. 2019). e 32 specimens examined showed
little variation in the habitus, which is clearly dierent
from the other species of Malaveria gen. nov. In our DNA
study, the specimens analyzed clearly grouped apart from
the other species of the “alcinoe-complex” (except M. ro-
driguezi sp. nov. and M. bottoi sp. nov.) with signicant
dierence: 5.5% for M. duponti sp. nov. and M. mimula
comb. nov., 4.1% with M. mimas comb. nov., 2.7% with
Figure 22. Malaveria ballo sp. nov., imagos and genitalia. a. Male ventral; b. idem dorsal; c. Female ventral; d. idem dorsal; e. Male geni-
talia lateral view (one valva removed); f. idem dorsal view; g. Valva lateral inner view; h. idem lateral outer view; i. idem dorsal view; j. Ae-
deagus dorsal view; k. idem lateral view; l. Female genitalia lateral view; m. idem ventral view. (Scale imago: 1 cm; scale genitalia: 1 mm).
M. Benmesbah, Á. L. Viloria & J. Murienne
44
M. nebulosa comb. nov. and 2.3% with M. alcinoe comb.
nov. (Tables 1 and 2). e dierence observed with M. ro-
driguezi sp. nov. and M. bottoi sp. nov., is low (Table2) and
probably due to a lack of power of the DNA test used here
(see above). On the other hand, wing venation structure on
FW is most similar to M. duponti sp. nov. and M. mimula
comb. nov., in which R2, r2-5-m1 and r1-r2 roots share
the same origin at the distal edge of the cell, diering in
this from M. rodriguezi sp. nov., in which the origin of R2
is proximal to the separation between r2-5-m1 and r1-r2
(Figs. 4a-4h). e specimens dissected (12 males) showed
certain homogeneity in the shape of the distal extremity of
the valvae. Conversely to M. nebulosa comb. nov. and M.
alcinoe comb. nov., small irregular membranes are present
at the distal tip of the valvae and the inner spiny projection
is usually absent, or if present, it is very small and more dis-
tally located. All these elements argue for its specic status
and its closer proximity within Malaveria gen. nov., to M.
rodriguezi sp. nov. and M. bottoi sp. nov.
Malaveria rodriguezi Benmesbah & Viloria, sp. nov.
(Figs. 23a-23i)
http://zoobank.org/urn:lsid:zoobank.org:act:AF9CCEA0-B91D-414C-
94E9-853E38BA8318
Type material: H (Figs. 23a-23b) male: /
COLOMBIA, Antioquia, Frontino 2000m, 22.IV.2007/
Holotype of Malaveria rodriguezi Benmesbah & Viloria,
2020/Gabriel Rodríguez leg./MB-0249*/ [MB]. P-
(all from COLOMBIA): 5 males: 1 male Antio-
quia, Amagá 1700m, 17.08.2003 (MB-0248) [MNHN];
1 male Antioquia, Frontino 1900m, 22.IV.2000
(MB-0253) [MB]; 1 male Antioquia, Guarne 2000m,
20.XI.2003 (MB-0254*) [MB]; 1 male Antioquia, La
Mansa, Ciudad Bolívar 2400m, 25.01.2004 (MB-0255*)
[MB]; 1 male Antioquia, Porce 1000m, 09.08.2003 (MB-
0265*) [MB].
Additional material: 8 specimens (1 male dissected)
not included in the type series:
COLOMBIA(8 specimens): Caldas: Manizales, Bar-
rio Los Nogales 1950m, 03-10.IX.2019 6 specimens
(photograph) [ICRM]; Santander: El Hato, Serranía de
los Yariguíes 2100m, 25-28.XI.20191 male (photograph)
[ICRM]; Tolima: Prado 700m, September 2003 1 male
(MB-0256*).
Description: Male (Figs. 23a-23b): FWL: Holotype
22 mm, 22-24 mm (7 specimens), average 22.9 mm.
DFW: ground color brown uniform. VFW: ground color
same as DFW; submedian and median lines thin, brown
slightly ferruginous, submedian line straight, median
line curved inside near the costa and outwards in Cu2-
2A; submarginal area with one small ocellus, black with
two tiny pupils circled by a brownish ring in the apex in
M1-M2; submarginal line of almost same width as me-
dian line darker, scalloped from radius to 2A, pointing
distally in M1 to 2A; marginal dark line thinner and reg-
ular, parallel to outer margin. DHW: ground color same
as DFW; marginal and submarginal lines observable
through translucent wing. VHW: ground color same as
VFW; median and submedian lines slightly irregular, of
same width and color as on VFW, joining anal margin
without making angulation, median line joining the anal
margin at the origin of the submarginal line; submarginal
line of same color but slightly wider, scalloped, pointing
distally from M1 to 2A; marginal dark line thin parallel
to outer margin from costa to tornus; submarginal area
with ve ocelli, one in each space between Rs and Cu2,
ocellus in
M1-M2 the biggest, black circled by a yellow-
ish ring with two small white pupils, ocellus in Cu1-Cu2
similar, more ovoid and slightly smaller, ocelli in M2-
M3 and M3-Cu1 smaller and less dened without black
in the center, ocellus in Rs-M1 the smallest bipupillate
without black in the center.
Female: not examined.
Male genitalia (5 dissected) (Fig. 23c-23i): general
structure similar to other species of the genus. Shape of
the valvae very similar to those of M. ballo sp. nov., but
horn-like inner processes more developed; aedeagus larger
than valva and more robust than in the species previously
studied.
Female genitalia: not examined.
Variation: the specimens from the type series pres-
ent little variation, mainly in the median line on VHW
that can be a little more wavy than in the holotype. Size
of the ocelli on VHW seems to vary slightly, being wider
in specimens from the south (the holotype presenting the
smallest ones).
Diagnosis (Figs. 20a-20l): M. rodriguezi sp. nov., dif-
fers from M. alcinoe comb. nov. and M. ballo sp. nov.,
by the median line on VHW joining the anal margin at
the origin of the submarginal line without making a pro-
nounced angle; in the two latter species the median line
joins the anal margin at a distance from the origin of the
submarginal line and forms a greater angle in M. ballo
sp. nov. and M. alcinoe comb. nov. (a little less in the lat-
ter than the former). M. rodriguezi sp. nov., diers from
M. bottoi sp. nov. by: 1. its darker background color;
2.straighter median line on VHW; 3. wider ocelli on the
underside and 4. submarginal line more angular on VHW
and VFW (attened in M. bottoi sp. nov.). Male genitalia
morphology does not dier evidently from the other spe-
cies of Malaveria gen. nov.
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
45
Etymology: dedicated to Gabriel Rodríguez, a Co-
lombian geologist and entomologist studying the butter-
y fauna of his country, who collected the type series and
kindly shared this study material with us. e name rodri-
guezi is a Latinized masculine noun in the genitive case.
Hostplant: unknown.
Habitat and behavior: we did not observe M. rodri-
guezi sp. nov., in natural conditions. As in the previous
case, we can only presume that its behavior is similar to
other species of Malaveria gen. nov. is species seems to
be distributed at least through the three Cordilleras in the
northern half of Colombia, with the exception of the east-
ern slopes of the Cordillera Oriental (Fig. 21). Recorded
from 700 to 2,400m.
Discussion: although similar in appearance to the
other species of the “alcinoe-complex”, M. rodriguezi sp.
nov. shows particular characters arguing for its specic
status: 1. the most evident is the shape of the median line
on VHW when joining the anal margin (Figs. 20a-20l);
2.the DNA analysis performed here shows important sim-
ilarity with M. bottoi sp. nov. (100%) and M. ballo sp.
nov. (99.5%), from which M. rodriguezi sp. nov., is, never-
theless, clearly dierentiated by the previous morphologi-
cal criteria, pointing out the probable lack of power of the
DNA technique used here. On the other hand specimens
of M. rodriguezi sp. nov., grouped apart from M. mimas
comb. nov., M. nebulosa comb. nov. and M. alcinoe comb.
nov. (Table 1), and showed respectively signicant dier-
ence: (4.6%), (3.2%) and (2.7%) (Table 2); 3. FW wing
venation presents the particularity (shared also with M.
mimas comb. nov.) that R2 origin is proximal to the bifur-
cation between r2-5-m1 and r1-r2 (distal in M. nebulosa
comb. nov. and M. alcinoe comb. nov., and emerging from
the same root in M. ballo sp. nov.) (Figs. 4a-4h).
e description of M. rodriguezi sp. nov., is based on
a series of 6 similar specimens from dierent localities
in Antioquia sharing stable characters. We also examine
specimens directly and photographs of specimens we also
identied as M. rodriguezi sp. nov., from Caldas, Santand-
er and Tolima departments in Colombia. M. rodriguezi sp.
nov., has a large distribution within Colombia, except the
eastern slopes of the Cordillera Oriental, where M. alcinoe
comb. nov., is present. e wide altitudinal range from
where it is recorded may explain its ability for dispersion
over natural barriers as mountains and valleys. Southern
specimens from the “alcinoe-complex” from Risaralda,
Caldas and Valle del Cauca showed some dierenceswith
typical individuals of M. rodriguezi sp. nov., and are de-
scribed below tentatively as a dierent subspecies.
Malaveria rodriguezi risaralda Benmesbah & Viloria,
ssp. nov.
(Figs. 24a-24i)
http://zoobank.org/ urn:lsid:zoobank.org:act:39FBA58D-DF90-458A-
B0BA-A450C040D1B9
Type material: H male (Figs. 24a-24b): /CO-
LOMBIA, Risaralda, Pueblo Rico 1400m, 20.VII.2005/
Figure 23. Malaveria rodriguezi rodriguezi sp. nov., imagos and genitalia. a. Male ventral; b. idem dorsal; c. Male genitalia lateral view
(one valva removed); d. idem dorsal view; e. Valva lateral inner view; f. idem lateral outer view; g. idem dorsal view; h. Aedeagus dorsal
view; i. idem lateral view. (Scale imago: 1 cm; scale genitalia: 1 mm).
M. Benmesbah, Á. L. Viloria & J. Murienne
46
Holotype of Malaveria rodriguezi risaralda Benmesbah &
Viloria, 2020/Gabriel Rodríguez leg./MB-0250*/ [MB].
P: COLOMBIA: Risaralda: 1 males Pueblo
Rico 1400m, 20.VII.2005 (MB-0252*) [MB]; Valle del
Cauca: 1 male 1902 (MB-13*) [MNHN].
Additional material:4 specimens not included in the
type series: COLOMBIA: Caldas: Anserma, Ecoparque
los Tucanes 1907m, 16-19.II.2019 3 males (photo-
graph) [ICRM]; Manizales, uebrada El Águila 1750m,
04.II.2019 (photograph) 1 male [ICRM].
Description: Male (Figs. 24a-24b): FWL: Holotype
23 mm, 22–24 mm (7 specimens), average 22,9 mm.
DFW: ground color brown uniform. VFW: ground col-
or same as upperside; submedian and median lines thin,
brown slightly ferruginous, median line curved inside near
the costa and outwards in Cu2-2A; submarginal area with
three small ocelli, one black, circled by a brownish ring in
the apex in M1-M2, two others faded in M2-M3 and M3-
Cu1, almost not visible; submarginal line of same width as
median line, scalloped from radius to 2A, pointing distally
in M1 to 2A; marginal dark line thinner and regular, par-
allel to outer margin. DHW: ground color same as DFW;
marginal and submarginal lines observable through trans-
lucent wing. VHW: ground color same as VFW; median
and submedian lines broken and irregular, of same width
and color as on VFW, median line making a slight angula-
tion when joining the anal margin at the origin of the sub-
marginal line; submarginal line of same color but slightly
wider, scalloped, composed by straight adjacent segments,
pointing distally from M1 to 2A; marginal dark line thin,
parallel to outer margin from costa to tornus; submar-
ginal area with ve ocelli, one in each space between Rs
and Cu2, ocellus in
M1-M2 the biggest oval with long
axis directed to the base, black, circled by a yellowish ring
with two small white pupils, ocellus in Cu1-Cu2 similar
but slightly smaller, ocelli in M2-M3 and M3-Cu1 even
smaller and less dened without black in the center, ocel-
lus in Rs-M1 the smallest with little black in the center and
two small white pupils.
Female: not examined.
Male genitalia (3 dissected) (Fig. 24c-24i): there is cer-
tain variation between the general aspect of the genital
structures of this subspecies and those of the nominotypi-
cal one; however, it seems to be within the extremes of in-
dividual variation of the specimens examined. e struc-
ture illustrated shows a saccus narrower at base and valvae
with more prominent processes. e aedeagus looks rela-
tively thicker than that of M. rodriguezi rodriguezi sp. nov.
Female genitalia: not examined.
Variation: the shape of the median line on VHW can
be more or less wavy. e two specimens from Risaralda
(holotype and one paratype) present a markedly wavy me-
dian line.
Diagnosis: M. rodriguezi risaralda ssp. nov., diers
from the nominate subspecies on VHW by: 1. larger ocel-
li, especially in M1-M2 and Cu1-Cu2; 2. ocellus in M1-
M2 a little elongated with its long axis pointing towards
the base (rounded in the nominate) and 3. ocellus in M1-
Figure 24. Malaveria rodriguezi risaralda ssp. nov., imagos and genitalia. a. Male ventral; b. idem dorsal; c. Male genitalia lateral view
(one valva removed); d. idem dorsal view; e. Valva lateral inner view; f. idem lateral outer view; g. idem dorsal view; h. Aedeagus dorsal
view; i. idem lateral view. (Scale imago: 1 cm; scale genitalia: 1 mm).
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
47
M2 only slightly bigger than ocellus in Cu1-Cu2 (notably
bigger in the nominate subspecies).
Etymology: in allusion to the type locality, Risaralda
department in Colombia. “risaralda” is a feminine noun
in apposition.
Hostplant: unknown.
Habitat and behavior: we did not observe this subspe-
cies in the eld. e records are located on both slopes of
the Cordillera Occidental in the departments of Risaralda,
Caldas and Valle del Cauca in Colombia (Fig. 21). Re-
corded from 1,400 to 1,907m.
Discussion: subspecies risaralda shares the same di-
agnostic characters that dierentiate M. r. rodriguezi sp.
nov., from M. alcinoe comb. nov., M. ballo sp. nov., and
M. bottoi sp. nov. (see under nominate). is subspecies
ranges south and west from the area of distribution of the
nominate subspecies (Fig. 21). All the records we have
(7males) are from the western (Pueblo Rico, Risaralda)
and the eastern (Anserma y uebrada el Aguila, Caldas)
slopes of the Cordillera Occidental. ese localities are
very close geographically (20 km) and separated by moun-
tains between 2,000m and 2,800m high. e continuity of
the mountains probably allows butteries of this species to
ow between them. e specimen labelled Valle del Cauca
could have originated from either side. e study of male
genitalia, with 5 specimens of the nominate and 3speci-
mens from M. r. risaralda ssp. nov., dissected, did not
show signicant dierences. e DNA analysis performed
here with 7 specimens of the nominate and 2 specimens of
M. r. risaralda ssp. nov., did not show dierence on the an-
alyzed fragment. Furthermore, we found a few specimens
from Manizales, Caldas, with larger ocelli on VHW and
1 from Anserma with smaller ocelli than usual on VHW,
that are probably intermediate between the 2 subspecies.
For these reasons we treat here risaralda as a subspecies of
M. rodriguezi sp. nov.
Malaveria bottoi Benmesbah & Viloria, sp. nov.
(Figs. 25a-25i)
http://zoobank.org/urn:lsid:zoobank.org:act:122E610E-9A01-4D95-A4F5-
4FD1F2D729C9
Type material: H male: COLOMBIA:
Cundinamarca, Colegio 430m/Holotype male of Malave-
ria bottoi Benmesbah & Viloria, 2020/Patrick Botto leg./
MB-0019*/ [MB].
Description: Male (Figs. 25a-25b): FWL: 22 mm
(1specimen). DFW: ground color brown uniform. VFW:
ground color brown lighter than DFW; submedian and
median lines thin, brown slightly ferruginous, submedian
line straight, median line curved outwards in Cu2-2A;
submarginal area with one faded, very small ocellus in the
apex in M1-M2; submarginal line of same width and col-
or as median line, discreetly scalloped from radius to 2A,
pointing distally in M1 to 2A; marginal dark line thinner
and regular, parallel to outer margin. DHW: ground color
same as DFW; marginal and submarginal lines observable
through translucent wing. VHW: ground color same as
VFW; submedian and median lines irregular and wavy, of
same width and color as on VFW, submedian line joining
anal margin without making angulation, median line join-
ing the anal margin making a slight angulation at the origin
of the submarginal line; submarginal line of same color and
width, attened and discreetly scalloped, pointing distally
from M1 to 2A; marginal dark line thin parallel to outer
margin from costa to tornus; submarginal area with ve
ocelli of almost same size, one in each space between Rs
and Cu2, ocelli in
M1-M2 and Cu1-Cu2 black, circled by
a yellowish ring with two small white pupils, ocelli in M2-
M3 and M3-Cu1 less dened, without black in the center,
ocellus in Rs-M1 with a single and wide white pupil.
Female: not examined.
Male genitalia (1 dissected) (Figs. 25c-25i): a single
individual dissected shows genitalic structures closer in
appearance to M. rodriguezi sp. nov., than to its allegedly
sympatric (or altitudinal parapatric) M. alcinoe comb. nov.
Female genitalia: not examined (unknown).
Diagnosis: M. bottoi sp. nov., diers from all the other
species of Malaveria gen. nov., by: 1. submarginal line on
VHW and VFW attened and subtly scalloped, and much
more removed from the marginal line than in other species
(except M. mimas comb. nov.); 2. markedly wavy VHW
median line, especially near the costa; 3. more rounded
outer margin on hindwing (except M. mimas comb. nov.),
and 4. lighter brown underside. For dierences with M.
mimas comb. nov., see under the latter.
Etymology: in honor of our friend Doctor Patrick Bot-
to, a French physician and entomologist who is an enthu-
siast student of Neotropical butteries and in particular
satyrines of the tribe Haeterini. He collected the holotype
of this species and kindly allowed us to study it. e Lati-
nized species name bottoi is treated as a masculine noun in
the genitive case.
Hostplant: unknown.
Habitat and behavior: we did not observe M. bottoi sp.
nov., in its natural habitat. Following indications from the
collector, the holotype was captured in a locality called Co-
legio (430 m) in Cundinamarca. It possibly corresponds
to Mesitas del Colegio, 630 m, in the Bogotá river valley
(A. Neild pers. comm.) (Fig. 21). is locality is consider-
ably lower than those in which M. alcinoe comb. nov., has
been recorded. We think these apparently dierent species
M. Benmesbah, Á. L. Viloria & J. Murienne
48
might y in parapatry, at dierent altitude levels. We have
no other record for this species.
Discussion: the description of M. bottoi sp. nov., is
based on a unique male specimen. is specimen pres-
ents several particular characters that clearly dierentiate
it from the other species of Malaveria gen. nov. (see di-
agnosis). e combination of these characters makes im-
probable that this specimen represents an aberration or a
particular form of M. alcinoe comb. nov., or related spe-
cies. Unsurprisingly, the male genitalia are similar to the
other species of the “alcinoe-complex” and the analyzed
fragment of DNA in our molecular study did not show
signicant dierences with M. rodriguezi sp. nov. (0%)
and M. ballo sp. nov. (0.5%) (see above). Wing venation
has not been examined because we were not able to dissect
the single specimen available.
Malaveria mimula (Hayward, 1954), comb. nov.
(Figs. 26a-26m)
Type material: H male: /Misiones, San Ig-
nacio, 1937 D.149/Holotipo/Euptychia mimula Hayw.
Holotipo ♂ K. J. Hayward det./Holotipo/Prep. Gen
SAT.159 K. J. Hayward det./1079 K. J. Hayward det./ [in
Museo de Ciencias Naturales de la Fundación Miguel
Lillo, Tucumán, Argentina] (photograph examined).
Additional material examined: 11 males (2 dissected)
and 2 females (1 dissected). PARAGUAY(11 males, 2 fe-
males): Cordillera: Compañia Naranjo, Barrio San José,
03.IX.2014 3 males (MB-0349, MB-0350, MB-0351);
Guaira: Colonia Independencia, 07.IX.2014 1 male (MB-
0155); Kaaguasú: 20 km South Repatriacion, 07.IX.2014
1 male (MB-0352); Paraguari: Cerro Acahay, 05.IX.2014
4 males (MB-0153*, MB-0154, MB-0353, MB-0354); La
Colmena, Kaatymi, 08.IX.2014 1 male (MB-0156); Sa-
pucai, 08.IX.2014 1 male (MB-0152*); 08.IX.2014 2fe-
males (MB-0266, MB-0343*) [MB].
Redescription: Male (Figs. 26a-26b): FWL: 20–23
mm (11 specimens); average 22 mm. DFW: ground color
brown uniform. VFW: ground color brown lighter than
DFW, almost greyish, picked with creamy light scales
except on anal margin, giving a smoky aspect to the pat-
tern elements; submedian and median lines thin, dark
brown, submedian line slightly curved outside in Cu2-2A,
median line curved inside near the costa and straight up
to 2A; submarginal area with one faded very small black
ocellus in the apex in M1-M2; submarginal line of same
width and color as median line, markedly scalloped from
radius to 2A, pointing distally in M1 to 2A; marginal dark
line thinner and regular, parallel to outer margin. DHW:
ground color same as DFW; marginal and submarginal
lines observable by transparency. VHW: ground color
same as VFW; submedian and median lines of same width
and color as on VFW, markedly irregular and broken, me-
dian line joining the anal margin at the origin of the sub-
marginal line; submarginal line of same color and width,
markedly scalloped, pointing distally from M1 to 2A;
marginal dark line thin parallel to outer margin from costa
to tornus; submarginal area with ve very small ocelli, one
in each space between Rs and Cu2, ocelli in M1-M2 and
Cu1-Cu2 black, circled by a yellowish ring, with two very
small white pupils, ocelli in M2-M3 and M3-Cu1 almost
non visible represented by a creamy oval dash, ocellus in
Rs-M1 the smallest, represented by a black dash.
Female (Figs. 26c-26d): FWL: 19-21 mm (2 speci-
mens); average 20 mm. Similar to male, diering by its
rounder global shape and lighter color on the underside.
Male genitalia (2 dissected) (Figs. 26e-26k): the gen-
eral armature in characteristic of the genus Malaveria gen.
nov. e uncus is lanceolate but relatively less stylized
(shorter and broader); the shape of the valvae is distinctive
in the basal half, where it has a dorsal sinusoidal ridge, the
digitiform distal half more ornamented with processes; ae-
deagus relatively chubby.
Female genitalia(2 dissected) (Figs. 26l-26m): we can
only distinguish that the corpus bursae is seemingly less
elongated than in other species of the genus, the signa are
thick, almost as long as the corpus, running parallel to
each other.
Variation: specimens examined from Paraguay did not
show signicant variation among them.
Diagnosis: M. mimula comb. nov., diers from all the
other species of Malaveria gen. nov., by: 1. smoky aspect
of the underside pattern (except M. duponti sp. nov.);
2.markedly irregular and broken median line on VHW;
3.very small ocelli on underside and 4. markedly scalloped
and broken submarginal line on underside. For dierences
with M. duponti sp. nov., see under that species.
Hostplant: unknown.
Habitat and behavior: M. mimula comb. nov., has
been observed by one of the authors (MB) in Paraguay.
It was a common species in the region of Asunción, be-
ing collected at various sites. It was mostly encountered in
secondary growth forest in more or less open areas. e
behavior was typical of other common species of satyrine
butteries of the subtribe Euptychiina (see under M. mod-
esta comb. nov.). e species is distributed in Paraguay,
Southeastern Bolivia, Argentina and Southeastern Brazil.
Discussion: Euptychia mimula was described by Hay-
ward in 1954 from two male individuals, one (holotype)
from San Ignacio, Misiones, Argentina; the other (para-
type) from Santa Cruz, Bolivia. e holotype is illustrated
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
49
on the BOA website (http://www.butteriesofamerica.
com/L/t/Yphthimoides_mimula_a.htm), and agrees very
well with the specimens we examined and dissected from
Paraguay. Despite the apparently modied pattern of the
underside with the smoky aspect, the markedly irregular
submedian and median lines, the markedly broken sub-
marginal line and the very small ocelli, we found that this
species shares most of the characteristics dening Mala-
veria gen. nov. Furthermore, the male genitalia are very
similar to those of the other species of Malaveria gen. nov.
Our study found that M. mimula comb. nov., and M. du-
ponti sp. nov., share the same sequence in the DNA frag-
ment analyzed (see discussion under DNA results section,
Table 2) and the same venation (Figs. 4g-4h). Malaveria
mimula comb. nov., represents the sister of the Andean
species of Malaveria gen. nov. ese arguments conrm
Figure 25. Malaveria bottoi sp. nov., imagos and genitalia. a. Male ventral; b. idem dorsal; c. Male genitalia lateral view (one valva re-
moved); d. idem dorsal view; e. Valva lateral inner view, f. idem lateral outer view; g. idem dorsal view; h. Aedeagus dorsal view; i. idem
lateral view. (Scale imago: 1 cm; scale genitalia: 1 mm).
Figure 26. Malaveria mimula comb. nov., imagos and genitalia. a. Male ventral; b. idem dorsal; c. Female ventral; d. idem dorsal;
e.Male genitalia lateral view (one valva removed); f. idem dorsal view; g. Valva lateral inner view; h. idem lateral external view; i. idem
dorsal view; j. Aedeagus dorsal view; k. idem lateral view; l. Female genitalia lateral view; m. idem ventral view. (Scale imago: 1 cm;
scale genitalia: 1 mm).
M. Benmesbah, Á. L. Viloria & J. Murienne
50
the new taxonomic arrangement adopted here by transfer-
ring mimula to Malaveria gen. nov.
Malaveria duponti Benmesbah & Murienne, sp. nov.
(Figs. 27a-27m)
http://zoobank.org/ urn:lsid:zoobank.org:act:4BE287E2-28C0-4C95-AC3C-
D22F053D02D0
Type material: H male (Figs. 27a-27b): /BO
LIVIA: Caranavi/Holotype male of Malaveria duponti
Benmesbah & Murienne, 2020/Daniel Dupont leg./MB-
0147*/ [MB]. P: 3 males: same data as Holotype
(MB-0145*, MB-0146*) [MB], (MB-0148*)[MNHN]; 1
female: same data as Holotype(MB-0149*) [MB].
Description: Male (Figs. 27a-27b): FWL: Holotype
22 mm, 22–22 mm (4 specimens); average 22 mm. DFW:
ground color brown uniform. VFW: ground color brown,
lighter than DFW; submedian and median lines slightly
irregular, thin, brown, slightly ferruginous, submedian
line curved outside in Cu2-2A, median line curved inside
near the costa and straight up to 2A; submarginal area
with a small black ocellus circled by a yellowish ring with
two tiny white pupils in the apex in M1-M2; submarginal
line of same width and color than median line, scalloped
from radius to 2A, pointing distally in Rs to Cu2; mar-
ginal line thinner and regular, parallel and closer to outer
margin. DHW: ground color same as DFW; marginal and
submarginal lines observable through translucent wing.
VHW: ground color same as VFW; submedian and me-
dian lines very irregular and broken of same width and
color as on VFW, median line joining the anal margin
at the origin of the submarginal line; submarginal line of
same color and width, scalloped, pointing distally from
M1 to 2A; marginal dark line thin, parallel to outer mar-
gin from costa to tornus; submarginal area with ve ocelli,
one in each space between Rs and Cu2, ocelli in M1-M2
and Cu1-Cu2 rounded, black, circled by a yellowish ring
with two very small white pupils, ocelli in M2-M3 and
M3-Cu1of same size as the precedents, yellowish without
black or pupils in the center, ocellus in Rs-M1 the smallest,
represented by a black dash circled with a yellowish ring.
Female (Figs. 27c-27d): FWL: 22 mm (1 specimen).
Similar to male, the only female examined diers by the
lighter under and upperside ground color.
Male genitalia (4 dissected) (Figs. 27e-27k): the gen-
eral aspect of the genital structure of this distinctive spe-
cies matches all features described for Malaveria gen. nov.
e tegumen is globular; the uncus lanceolate and propor-
tionally similar to that of the type species of the genus; the
saccus is more stylized (thinner); the valvae have an angled
corner on the dorsal ridge, at the end of its basal half, just
before the constriction in which the digitiform extremity
is formed, the distal horn-like inner process of the valva
is particularly well developed; the aedeagus is roughly the
same length as the valva, and is moderately robust, broader
at its base.
Female genitalia (1 dissected) (Figs. 27l-27m): very
well developed papillae anales, setose; ostium bursae lo-
Figure 27. Malaveria duponti sp. nov., imagos and genitalia. a. Male ventral; b. idem dorsal; c. Female ventral; d. idem dorsal; e. Male
genitalia lateral view (one valva removed); f. idem dorsal view; g. Valva lateral inner view; h. idem lateral outer view; i. idem dorsal view;
j. Aedeagus dorsal view; k. idem lateral view; l. Female genitalia lateral view; m. idem ventral view. (Scale imago: 1 cm; scale genitalia:
1 mm).
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
51
cated immediately below; ductus as long as the corpus
bursae, in which there are two longitudinal, parallel signa,
running from its base to the middle of its length.
Variation: all the specimens examined did not show
signicant variations.
Diagnosis: M. duponti sp. nov., diers from all the
other members of Malaveria gen. nov. (except M. mim-
ula comb. nov.), by the markedly irregular and angled
submedian and median lines on VHW. In this aspect it
is reminiscent of M. nebulosa comb. nov., in which these
two lines are less irregular and more wavy (Fig. 20). Fur-
thermore the distal extremity of the valvae is dierent: 1.
membranes present in M. duponti sp. nov. (absent in M.
nebulosa comb. nov.), and 2. proximal spiny inner projec-
tion present in M. nebulosa comb. nov., is absent in M. du-
ponti sp. nov. DNA analysis showed specimens of M. neb-
ulosa and M. duponti sp. nov., to clearly group separately
within Malaveria gen. nov. (Table 1), and their dierence
is 6.8% (Table 2). M. duponti sp. nov., is very close to M.
mimula comb. nov., but diers by: 1. the ground color of
the underside (brown in M. duponti sp. nov., greyish in M.
mimula comb. nov.); 2. the more clearly dened pattern
of the underside in M. duponti sp. nov.; 3. the bigger ocelli
in M. duponti sp. nov. e latter is also apparently close
to ‘Yphthimoides’ grimon (Godart, [1824]) with which it
shares the markedly irregular and angled submedian, me-
dian and submarginal lines (Fig. 20l). A careful examina-
tion shows the median line drawing is not exactly the same
and diers subtly from each other. Furthermore,the male
genitalia (4 specimens of M. duponti sp. nov., and 2 of ‘Y.’
grimon dissected) are distinct: 1. upperside of the uncus
markedly convex upward in ‘Y.’ grimon (attened and
straight in M. duponti sp. nov.); 2. distal part of the valvae
much shorter in ‘Y.’ grimonand 3. dorsal part of the valvae
much more angular in ‘Y.’ grimon.
Etymology: dedicated to honor our friend Daniel Du-
pont, a French entomologist and collector, who provided
many interesting buttery samples for study from South
America, including the type series of this species. e
name duponti is a Latinized masculine noun in the geni-
tive case.
Hostplant: unknown.
Habitat and behavior: we did not observe this spe-
cies in the eld. All the specimens we could examine are
from Caranavi, Bolivia. ey were included in a mixed
sample of papered Satyrinae collected from that region.
No information for elevation data was provided, but it is
presumably up to 800m as most of the other species were
collected within that range.
Discussion: the examination of male genitalia (4males
dissected) did not show signicant interspecic dier-
ences; as discussed before, male genitalia are very similar
among dierent species in this genus and are not always
diagnostic at specic level. Molecular analysis showed
specimens of M. duponti sp. nov., and M. mimula comb.
nov., to have the same nucleotide sequence in the DNA
fragment analyzed; as discussed above, we think this is due
to the lack of discriminating power of the method used
here as it occurred with M. modesta comb. nov. and M.
remypignoux sp. nov., and with M. rodriguezi sp. nov. and
M. bottoi sp. nov. Given the dierences in habitus and the
distance separating the two species, we treat here M. du-
ponti sp. nov., as a species in its own right. M. duponti sp.
nov. and M. mimula comb. nov., are probably sister taxa
inhabiting in two dierent biogeographical areas, the An-
dean foothill for the former and the Chaco and adjacent
lowlands of Bolivia, Paraguay, Argentina and Brazil (and
perhaps Uruguay) for the latter. Further collecting in Bo-
livia and a more comprehensive DNA comparative study
could provide a better understanding of the relationships
between these 2 taxa and other species of the “alcinoe-
complex”. Although similar in appearance to M. duponti
sp. nov., ‘Yphthimoides’ grimon shows dierent male geni-
talia characters that could be specic of a distinct species
group.
Koutalina Viloria & Murienne, gen. nov.
Koutalina pamela (Hayward, 1957), comb. nov.
(Figs. 28a-28o)
Type material: not located. Hayward (1957) desig-
nated a male holotype (genitalia preparation No. MSS
26) that was deposited at the Instituto de Historia Natu-
ral “Sánchez Labrador” in San Miguel de Buenos Aires,
Argentina (an ephemeral institution that lasted just a few
years mainly due to the tragic death of its founder – Fa-
ther Albino J. Bridarolli– in 1949). According to Williner
(1938, 1950), SEA (1950) and Asúa (2019), the Bridarolli
collection of the mentioned institute was kept in the Cole-
gio Máximo de San José. Bachmann (2003) indicates that
in 1983 the founders of the Instituto de Entomología de
Salta (INESALT) in Rosario de Lerma, received a mini-
mal part of that entomological collection. e other part
(which is the largest) was rst transferred to the Colegio
del Salvador in Buenos Aires, where it was properly cared
for, but nally donated and transferred to the Museo Ar-
gentino de Ciencias Naturales “Bernardino Rivadavia”
(MACN), on February 25, 2003. Some samples remain at
the Colegio del Salvador for educational purposes (Bach-
mann 2003). Barbosa et al. (2018) also failed to locate in
the MACN collection a type specimen of another Boliv-
M. Benmesbah, Á. L. Viloria & J. Murienne
52
ian satyrine species described by Hayward, Euptychia pa-
tricia. Ellenrieder (2009) does not mention any type of
Lepidoptera in the collections of the Museo de Ciencias
Naturales de Salta.
Material examined: 15 males (4 dissected), 1 female
(1 dissected): - BOLIVIA(9 males): La Paz: Caranavi
800-1500m, January 2013 6 males (MB-0057, MB-0058,
MB-0059*, MB-0060, MB-0061, MB-0062*) [MB],
Yungas de la Paz 1000m, 1913 2 males (MB-03, MB-04)
[MNHN]; No precise location: 1902 1 male (MB-01)
[MNHN].- PERU (6 males, 1 female): Cusco: Carretera
Manu 1 male (MB-0063*), uincemil 26km Southwest
1580m, 28-30.VI.2019 2 males (MB-0347, MB-0348)
[MB] and 1 female (MB-0346*) [MB]; Junín: Mina Pi-
chita 2100m, 12.IX.2001 2 males (MB-0064, MB-0065*)
[MB]; Pasco: Huacabamba, Cerro del Pasco, 1913 1 male
(MB-02) [MNHN].
Redescription: Male (Figs. 28a-28d): FWL: 19-23
mm, average 20.7 mm (15 specimens). DFW: ground
color dark-brown; large darker and rectangular androco-
nial patch in the middle of the forewing (Figs. 29a-29d).
VFW: ground color brown lighter than DFW, discreetly
coppered; submedian and median lines slightly irregular,
thin, dark-brown, median line running outside between
Cu2 and 2A; submarginal area with a small black ocellus
in the apex in M1-M2, circled by a yellowish ring, with
two tiny white pupils; submarginal line of same color,
slightly thinner than median line, scalloped from radius to
2A, pointing distally in Rs to Cu2; marginal line thinner
and regular, parallel to outer margin. DHW: ground color
same as DFW; marginal and submarginal lines observable
through translucent wing; ocellus in Cu1-Cu2 black cir-
cled with yellowish ring, with two tiny white pupils in the
center; marginal line broadened at tornus; areas between
submarginal line and outer margin lighter. VHW: ground
color same as VFW; submedian and median lines slightly
irregular, of same width and color as on VFW, median line
making a smooth curve before joining the anal margin at
the origin of the submarginal line; submarginal line slight-
ly thinner, of same color, scalloped, pointing distally from
M1 to 2A, making a double sinuosity towards the base in
M2-M3, and at in Cu1-Cu2; marginal dark line thinner,
parallel to outer margin, broadened at tornus; submargin-
al area with six ocelli, all with two tiny white pupils, cir-
cled by a yellowish ring and surrounded by a discreet dark
brown area, one in each space between Rs to 2A, ocellus
in M1-M2 bigger, rounded and black, touching M1 and
M2, ocellus in Cu1-Cu2 similar, slightly smaller, ocelli in
M2-M3 oval and M3-Cu1 rounded, both yellowish with-
out black in the center, ocellus in Rs-M1 the smallest, anal
ocellus in Cu2-2A black, same size as that one in M3-Cu1.
Female (Figs. 28e-28f ): FWL: 20 mm (1 specimen).
Similar to male but diers by much lighter upper and un-
derside and by the absence of androconial patch on DFW.
Male genitalia (4 dissected) (Figs. 28g-28m): as in the
generic description above.
Female genitalia (1 dissected) (Figs. 28n-28o): ventral
sclerite of the 8th abdominal segment much broader than
Figure 28. Koutalina pamela comb. nov., imagos and genitalia. a. Male from Bolivia ventral; b. idem dorsal; c. Male from Peru ventral;
d. Male from Bolivia ventral; e. Female from Peru ventral; f. idem dorsal; g. Male genitalia lateral view (one valva removed); h. idem
dorsal view; i. Valva lateral inner view; j. idem lateral outer view; k. idem dorsal view; l. Aedeagus dorsal view; m. idem lateral view; n.
Female genitalia lateral view; o. idem ventral view. (Scale imago: 1 cm; scale genitalia: 1 mm).
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
53
in the representatives of Modestia gen. nov., and Malave-
ria gen. nov. Papillae anales proportionally smaller than
in Malaveria gen. nov.; bursa copulatrix with two distinc-
tive signa, running parallel to each other throughout basal
half of the corpus, but notably separated, perhaps twice as
much as in the species of Malaveria gen. nov., ductus as
long as the corpus bursae.
Variation (Figs. 28a-28d): the 16 specimens (15 males
and 1 female) from Bolivia and Peru showed variation in
the average size, specimens from Peru being smaller with
a more rounded wing shape: average FWL 21.2 mm from
Bolivia (9 specimens), 19.8 mm from Peru (6 specimens).
In general specimens from Peru seem to have a more grey-
ish underside and ocelli, median and submedian band
wider on VHW than specimens from Bolivia (see discus-
sion). Only 1 female was examined and it is from Peru.
On DHW, ocellus in Cu1-Cu2 appears with two distinct
pupils or with a single fused wider pupil in the same fash-
ion as it has been discussed above for species belonging in
Modestia gen. nov., and Malaveria gen. nov. e tiny ocel-
lus in Cu2-2A is present in half of the specimens (none
from Caranavi).
Diagnosis: Koutalina pamela comb. nov., is easily dif-
ferentiated from all other genera currently included in the
Euptychiina by the presence of 6 ocelli on VHW, all bipu-
pillate and a large androconial patch (Figs. 29a-29d) on
DFW in males.
Hostplant: unknown.
Habitat and behavior: Á. L. Viloria observed and col-
lected K. pamela comb. nov., in the locality of Mina Pi-
chita (2,100m), Junín, Peru. is buttery was found y-
ing fast and erratically in slightly disturbed, dry, low cloud
forest. Short periods of the buttery resting on the leaves
of the shrubs at clearings under sunshine allowed their
capture, but in general, individuals escaped by skipping
and fast ying upwards. All the specimens we examined
are from the eastern slopes of the Andes from las yungas
(Bolivia) to Pasco (Central Peru). Records are from 800m
to 2,100m.
Discussion: E. pamela was described by Hayward in
1957 apparently based on a single specimen from Chu-
lumani, South Yungas, Bolivia. A meticulous description
is given and the male genitalia are illustrated. Hayward
mentions two ocelli on the DHW tornus (see Fig. 29a).
We found this character to be inconstant, half of the speci-
mens showing only the biggest ocellus in Cu1-Cu2. Al-
though the androconial patches are not clearly mentioned,
Hayward noticed that the ground color of the DFW was
dark brown (almost black), and the distal third of the
wing lighter. Hayward nished his description considering
E. pamela to be close to E. phineus Butler, 1867, a junior
synonym of Optimandes eugenia eugenia (C. Felder & R.
Felder, 1867), the type species of the clearly dierent genus
Optimandes Marín, Nakahara & Willmott, 2019 (in Will-
mott et al. 2019). Optimandes eugenia diers from it in
many aspects: 1. a single pupil in all the ocelli of the VHW;
2. absence of androconial patch on DFW and 3.dierent
morphology of their male genitalia. To date the holotype
of E. pamela has not been found but it is probably pre-
served in a minor Argentinian collection (see above). La-
mas (2004) treated E. pamela as a synonym of N. alcinoe,
which it is clearly not. D’Abrera represented two males or
K. pamela comb. nov., as “Euptychia sp.?” (p.777) from
Peru and Bolivia in the NHMUK, just between E. phineus
and E. mimas, E. alcinoe and E. benedicta.
is species possesses very particular characters. ey
enabled us to erect and describe the new genus Koutalina
gen. nov., of which it is the only known representative.
Figure 29. Koutalina pamela comb. nov. DFW androconial patch. a. Male dorsal; b-d. Details of DFW scaling. (Scale image: 1 cm;
scale detail DFW: 1 mm).
M. Benmesbah, Á. L. Viloria & J. Murienne
54
e original description by Hayward (1957) mentions
4 characters particular to K. pamela comb. nov.: 1.pres-
ence of two ocelli in the tornus on DHW (it varies, some
specimens have only one); 2. lighter area outside the medi-
an line on VHW; 3. wide VHW ocellus in M1-M2 linked
with oval ocellus in M2-M3 which seems compressed by
the former; 4. VHW ocellus in M3-Cu1 oval, with long
axis parallel to external margin. ese characters are pres-
ent in two specimens from Bolivia (Fig. 28d) labelled
“Yungas de la Paz, 1000m” collected in 1913. Aer a care-
ful study of the original description and the original illus-
tration of the male genitalia, we have no doubt Hayward
based its description on a similar specimen, and the name
“pamela” applies to the taxon represented here by one of
these specimens. ey come from a locality 60km south
of Caranavi, in the same region of Las Yungas. ese two
specimens seem to represent an intermediate habitus be-
tween the populations from Caranavi (Bolivia) and Peru.
For the moment, we prefer treating them as the same
taxon.
In general what we identied as pamela is represented
by specimens that seem to show some variation in their
habitus (see “Variation”), and schematically two dier-
ent phenotypes can be distinguished. Five of the 6 males
from Caranavi (Bolivia) showed (Figs. 28a-28b): 1. larger
size; 2. homogenous light brown coloration on the under-
side; 3. relatively small ocelli on VHW; 4. thin median
and post median lines on VHW. e specimens from
south and central Peru diered as follows (Figs. 28c, 28e-
28f ): 1.smaller; 2. with a greyish VHW; 3. bigger ocelli
on VHW and 4. wider median and submedian lines on
VHW. Furthermore, none of the 6 specimens from Ca-
ranavi have an ocellus on DHW in Cu2-2A, whereas 5 of
the 6 specimens from Peru do present it. We did not nd
dierences in male genitalia (2 specimens from Caranavi
and 2 from Peru dissected), nor in DNA analysis (4 speci-
mens from Caranavi and 3 specimens from Peru). Two
specimens from the Yungas (Fig. 28d.), 1 from Caranavi
(Bolivia), and 2 others from Cusco and Pasco (Peru), had
intermediate habitus.
ACKNOWLEDGEMENTS
We want to thank Daniel Dupont (France), Chris-
tophe Faynel (France), John MacDonald (USA), Gabriel
Rodríguez (Colombia) and John Shuey (USA), who pro-
vided crucial study material from Bolivia, Peru, Colom-
bia, Panama and Belize. We are very grateful to Indiana
Cristóbal Ríos Málaver (Colombia), who kindly shared
with us photos of specimens of the “alcinoe group” from
his collection; to amara Zacca (Brazil), for the inter-
esting exchanges on the subject, the realization of most
of the female genitalia dissections and for allowing us to
edit wing venation of Emeryus argulus and Paryphthimoi-
des poltys; to Blanca Huertas (England), for allowing us
to illustrates the type specimens hosted in the NHMUK;
Rodolphe Rougerie (France), who kindly allowed us to
study the specimens under his care at the MNHN; David
Trembath (England) for sharing with us pictures of its M.
alcinoe specimens from Venezuela; Christian Brévignon
(French Guiana) for sharing with us information about
M. modesta comb. nov. and M. remypignoux sp. nov.;
Patrick Botto (France) who collected the holotype of M.
bottoi sp. nov.; and to Raúl Aldaz (Ecuador) for having
collected several specimens of M. ballo sp. nov. We also
extend our deep gratitude to our friends and colleagues
Andrew Neild (England), who critically read this paper
and greatly improved the quality of the English, the co-
herence of its content and its general scientic quality,
and Mauro Costa (Venezuela) for his accurate review
and constructive comments. Yves Cluenot (France) from
CEBA laboratory in Toulouse, was of great help in man-
aging all the DNA samples. M. Benmesbah thanks John
Kotchalka and Carlos Aguilar Julio (Paraguay) for their
assistance and help during his eld study in Paraguay in
2015. Á. L. Viloria thanks Gerardo Lamas (Peru) and
Héctor Suárez (Venezuela) for his valuable friendship and
cooperation during a eld trip to the selva central in Peru
(2001), the latter is also recognized by his photographic
work, permanent innovation at digital processing and
dedication to image production at IVIC. We are indebted
to Tito R. Barros, Gilson Rivas, Neva Mora and especial-
ly María Eugenia Andara (Venezuela) for their valuable
technical assistance, patience in reviewing and editing the
contents of our texts and gures, and for their enduring
enthusiasm and professionalism.
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Taxonomic notes and descriptions of new satyrine butterflies from tropical America
59
APPENDIX.
DATA OF SPECIMENS ILLUSTRATED
Figure 3:
- 3a. Modestia modesta comb. nov., male: French Guiana,
Saül, 12.IX.2013 (MB-0021) (MB).
- 3b-3c. Modestia modesta comb. nov., male: French
Guiana, Saül, 12.IX.2013 (MB-0020) (MB).
- 3d. Modestia modesta comb. nov., male: French Guia-
na, Saül, 30.IX.2012 (MB-0022) (MB).
- 3e. Malaveria nebulosa comb. nov., male: Venezuela,
Miranda, El Jarillo, Guaicaipuro, uebrada Honda
1300m, 23.VII.2013 (MB-0002) (MB).
- 3f-3h. Malaveria nebulosa comb. nov., male: Venezue-
la, Miranda, El Jarillo, Guaicaipuro, uebrada Honda
1300m, 23.VII.2013 (MB-0006) (MB).
- 3i., 3l. Koutalina pamela comb. nov., male: Bolivia, La
Paz, Caranavi 800-1500m, I.2013 (MB-0059) (MB).
- 3j-3k. Koutalina pamela comb. nov., male: Bolivia, La
Paz, Caranavi 800-1500m, I.2013 (MB-0060) (MB).
Figure 4:
- 4b. Malaveria nebulosa comb. nov., male: Venezuela,
Miranda, El Jarillo, Guaicaipuro, uebrada Honda
1300m, 23.VII.2013 (MB-0002) (MB).
- 4c. Malaveria mimas comb. nov., male: Bolivia, La Paz,
Caranavi (MB-0013) (MB).
- 4d. Malaveria alcinoe comb. nov., male: Venezuela,
Lara, Parque Nacional Yacambú, 13km South Sanare,
15-21.I.1979 (MB-0008) (IVIC).
- 4e. Malaveria ballo sp. nov., male: Ecuador, Pichin-
cha, Nanegalito, 16.X.2015 (MB-0071) (MB).
- 4f. Malaveria rodriguezi sp. nov., male: Colombia, An-
tioquia, Frontino 1900m, 22.IV.2000 (MB-0253) (MB).
- 4g. Malaveria mimula comb. nov., male: Paraguay,
Cordillera, Compañía Naranjo, Barrio San José,
03.IX.2014 (MB-0349) (MB).
- 4h. Malaveria duponti sp. nov., male: Bolivia, Caranavi
(MB-0145) (MB).
Figure 5:
- 5a. Magneuptychia libye male: Venezuela, Cordillera de
la Costa, wing prep. ALV0004-14 (ALV).
- 5b-5c. Magneuptychia libye male: French Guiana.
- 5d. based on Emeryus argulus male venation in Zacca et
al. (2020, Fig. 3a).
- 5e-5f. Emeryus argulus magnum Zacca, Casagrande &
Mielke, 2020, male: French Guiana,
- 5g. based on Paryphthimoides poltys male venation in
Zacca et al. (2020, Fig. 68).
- 5h-5i. Paryphthimoides poltys female: Paraguay,
- 5j. based on Cissia penelope male venation in Zacca et al.
(2018, Fig. 3A).
- 5k-5l. Cissia penelope male: French Guiana, Saül, Boeuf-
mort, 03.IX.2011 (MB-0021) (MB).
Figure 6:
- 6a. Modestia modesta comb. nov., male: French Guiana,
Papaichton, 23.VII.2012 (MB-0023) (MB).
- 6b. Malaveria ballo sp. nov., male: Ecuador, Pichin-
cha, Nanegalito, 2016 (MB-0070) (MB).
- 6c. Malaveria ballo sp. nov., male: Ecuador, Pichin-
cha, Nanegalito, 2016 (MB-0076) (MB).
- 6d. Malaveria ballo sp. nov., male: Ecuador, Pichin-
cha, Nanegalito, 2016 (MB-0074) (MB).
- 6e. Malaveria ballo sp. nov., male: Ecuador, Pichin-
cha, Nanegalito, 2016 (MB-0257) (MB).
- 6f. Malaveria ballo sp. nov., male: Ecuador, Pichin-
cha, Nanegalito, 16.X.2015 (MB-0016) (MB).
Figure 7:
- 7a-b. Malaveria mimas comb. nov., male: Peru, Cusco,
Cusco, carretera Manu 800-2900m, October 2015 1
male (MB-0014) (MB).
- 7c-d. Malaveria mimas comb. nov., male: Bolivia, La
Paz, Caranavi (MB-0012) (MB).
- 7e-f. Malaveria mimas comb. nov., male: Bolivia, La
Paz, Caranavi (MB-0010) (MB).
Figure 8:
- 8a-b. Malaveria nebulosa comb. nov., male: Venezue-
la, Miranda: El Jarillo, Guaicaipuro, uebrada Honda
1700m, 05.VIII.2013 (MB-0004) (MB).
- 8c-d. Malaveria nebulosa comb. nov., male: Venezue-
la, Miranda: El Jarillo, Guaicaipuro, uebrada Honda
1200m, 26.VIII.2013 (MB-0007) (MB).
- 8e-f. Malaveria nebulosa comb. nov., male: Venezue-
la, Miranda: El Jarillo, Guaicaipuro, uebrada Honda
1700m, 23.VII.2013 (MB-0003) (MB).
Figure 9:
- 9a-b. Modestia modesta comb. nov., male: French Guia-
na, Saül, 30.IX.2012 (MB-0022) (MB).
- 9c-d. Modestia modesta comb. nov., male: French Guia-
na, Papaichton, 23.VII.2012 (MB-0023) (MB).
- 9e-f. Modestia modesta comb. nov., female: French
Guiana, Itoupé, 30.XI.2014 (MB-0030) (MB).
M. Benmesbah, Á. L. Viloria & J. Murienne
60
Figure 10:
- 10a. Malaveria nebulosa comb. nov., male: Venezue-
la, Miranda, El Jarillo, Guaicaipuro, uebrada Honda
1300m, 05.VIII.2013 (MB-0004) (MB).
- 10b. Malaveria nebulosa comb. nov., male: Venezue-
la, Miranda: El Jarillo, Guaicaipuro, uebrada Honda
1300m, 23.VII.2013 (MB-0006) (MB).
- 10c. Malaveria rodriguezi risaralda ssp. nov., male: Co-
lombia, Valle del Cauca, 1902 (MB-013) (MNHN).
- 10d. Malaveria sp. male: Colombia, Bogotá, Santa Fé
de Bogotá 1850m (MB-014) (MNHN).
- 10e. Modestia remypignoux sp. nov., Holotype
male: French Guiana, Maripasoula, Antecume Pata,
13.III.2012 (MB-0039) (MB).
- 10f. Modestia remypignoux shueyi ssp. nov., male: Pa-
namá, Colón, Santa Rita 245m, 18.II.2014 (MB-0213)
(MB).
- 10g. Malaveria mimas comb. nov., male: Bolivia, La
Paz: Caranavi (MB-0010) (MB).
- 10h. Malaveria mimas comb. nov., male: Peru, Cus-
co, carretera Manu 800-2900m, October 2015 (MB-
0014) (MB).
- 10i. Malaveria ballo sp. nov., Holotype male: Ecua-
dor, Pichincha, Nanegalito, 2016 (MB-0072) (MB).
- 10j. Malaveria ballo sp. nov., male: Ecuador, Pichin-
cha, Nanegalito, 2016 (MB-0073) (MB).
- 10k. Malaveria duponti sp. nov., male: Bolivia, Carana-
vi (MB-0145) (MB).
- 10l. Malaveria duponti sp. nov., male: Bolivia, Caranavi
(MB-0146) (MB).
Figure 11:
- 11a. ‘Magneuptychia’ tricolor male: Peru, Loreto, Sept-
ember 2013 (MB).
- 11b.‘Magneuptychia’ tricolor male: Peru, Loreto, Sept-
ember 2013 (MB).
- 11c. Megisto cymela male: USA, Ohio, Muskingum
County, Zanesville, 03.VI.1978 (MB).
- 11d. Modestia remypignoux sp. nov., female: French
Guiana, Maripasoula, Antecume Pata, 13.III.2012
(MB-0050) (MB).
- 11e. Magneuptychia libye male: Tobago, January 2009
(MB).
- 11f. Magneuptychia libye female: French Guiana, Saint-
Laurent-du-Maroni, 24.X.2011 (MB).
- 11g. Euptychia marceli male: French Guiana, Saül,
05.X.2011 (MB).
- 11h. Modestia remypignoux sp. nov., female: French
Guiana, Saint-Laurent-du-Maroni, 24.X.2011 (MB-
0051) (MB).
Figure 12:
- 12a. Lectotype of Euptychia modesta: /Type H.T./Syn-
type/♂/Type of Species./ Godman-Salvin Coll. 1904.-
-1./Euptychia modesta, Butl./♂ Pará modesta Butler
type / Para, L. Amazons. H. W. Bates. / B. M. TYPE
N°. Rh 3222 Euptychia modesta, ♂ Butl./BMNH(E)
#986581 (NHMUK).
- 12b. Lectotype of Neonympha sylvina: /Neonympha
sylvina Feld./Bahia Luschnatt type/FELDER CO-
LL
n
/Type/Type of N. sylvina Feld, very near E. modes-
ta. Butl/Sylvina n./Rothschild Bequest B.M.1939-1/
(NHMUK).
- 12c. Lectotype of Neonympha alcinoe: Type/FELDER
COLL
N
./Alcinoë Feld./Rothschild Bequest B.M. 1939-
1./Type of N. alcinoe Feld? = E. benedicta, Butl. of w. it.
may be a good local form. Comp. w. type E. benedicta
Butler./BMNH(E)1266958/(NHMUK).
- 12d. Syntype of Euptychia benedicta: /Type H.T./
Type of Species./E. benedicta Butler type./B.M. TYPE
No. Rh3227. Euptychia benedicta, ♀ Butl./Sarayacu,
Ecuador. C. Buckley./Godman-Salvin Coll. 1904.-1.
Euptychia benedicta, Butl./BMNH(E)1266955/(NH-
MUK).
Figure 13:
- 13a-13b. Modestia modesta comb. nov., male: French
Guiana, Saül, 12.IX.2013 (MB-0020) (MB).
- 13e-13k. Modestia modesta comb. nov., male: French
Guiana, Saül, 30.IX.2012 (MB-0022) (MB).
- 13c-13d, 13l-13m: Modestia modesta comb. nov., fema-
le: French Guiana, Mitaraka, 01.III.2015 (MB-0026)
(MB).
Figure 14:
- 14a-14b, 14d-14j. Modestia gomezi comb. nov., male:
Colombia, Antioquia, Porce 800m, 27.VIII.2017
(MB-0258) (MB).
- 14c. Modestia gomezi comb. nov., female: Panamá,
Zona del Canal, Cocoli, 28.XII.1986 (JMD).
Figure 15:
- 15a-15b, 15e-15k. Modestia remypignoux sp. nov., Ho-
lotype male: French Guiana, Maripasoula, Antecume
Pata, 13.III.2012 (MB-0039) (MB).
- 15c-15d. Modestia remypignoux sp. nov., paratype fe-
male: French Guiana, Macouria, Matiti, 18.VIII.2012
(MB-0049) (MB).
- 15l-15m. Modestia remypignoux sp. nov., paratype fe-
male: French Guiana, Kaw, Dégrad Escolle, 14.XI.2011
(MB-0047) (MB).
Taxonomic notes and descriptions of new satyrine butterflies from tropical America
61
Figure 16:
- 16a-16i. Modestia remypignoux shueyi ssp. nov., Beli-
ze. Holotype male: Belize, Cayo Dist., Baldy Beacon-
3,5km SSW - 1F5231 logging road in pine/oak, 750m,
23.IX.2016 (MB-0143) (MB).
Figure 17:
- 17a-17b, 17e-17k. Malaveria nebulosa comb. nov.,
male: Venezuela, Miranda, El Jarillo, Guaicaipuro,
uebrada Honda 1300m, 23.VII.2013 (MB-0006)
(MB).
- 17c-17d, 17l. Malaveria nebulosa comb. nov., female:
Venezuela, Miranda, El Jarillo, Guaicaipuro, uebrada
Honda 1200m, 06.III.2013 (MB-0005) (MB).
Figure 18:
- 18a-18i. Malaveria mimas comb. nov., male: Bolivia,
La Paz, Caranavi (MB-0012) (MB).
Figure 19:
- 19a-19b, 19e-19k. Malaveria alcinoe comb. nov., male:
Venezuela, Lara, Parque Nacional Yacambú, 13km
South Sanare, 15-21.I.1979 (MB-0008) (IVIC).
- 19c-19d, 19l-19m. Malaveria alcinoe comb. nov.,
female: Colombia, Cundinamarca, Colegio 430m,
15.VII.1975 (MB-0009) (MB).
Figure 20:
- 20b. Malaveria alcinoe comb. nov., Lectotype male: /
Type/FELDER COLL
N
./Alcinoë Feld./Rothschild Be-
quest B.M. 1939-1./Type of N. alcinoe Feld? = E. bene-
dicta, Butl. of w. it. may be a good local form. Comp.
w. type E. benedicta Butler./BMNH(E)1266958 (NH-
MUK).
- 20c. Malaveria alcinoe comb. nov., male: Venezuela,
Lara, Parque Nacional Yacambú, 13km South Sanare,
15-21.I.1979 (MB-0008) (IVIC).
- 20d. Malaveria rodriguezi rodriguezi sp. nov., Holo-
type male: Colombia, Antioquia, Frontino 2000m,
22.IV.2007 (MB-0249) (MB).
- 20e. Malaveria rodriguezi risaralda ssp. nov., Holo-
type male: Colombia, Risaralda, Pueblo Rico 1400m,
20.VII.2005 (MB-0250) (MB).
- 20f. Malaveria ballo sp. nov., Holotype male: Ecua-
dor, Pichincha, Nanegalito, 2016 (MB-0072) (MB).
- 20g. Malaveria bottoi sp. nov., Holotype male: Colom-
bia: Cundinamarca, Colegio 430m (MB-0019) (MB).
- 20h. Malaveria mimas comb. nov., male: Bolivia, La
Paz, Caranavi (MB-0012) (MB).
- 20i. Malaveria nebulosa comb. nov., male: Venezuela,
Miranda, El Jarillo, Guaicaipuro, uebrada Honda
1300m, 23.VII.2013 (MB-0006) (MB).
- 20j. Malaveria duponti sp. nov., Holotype male: Boli-
via, Caranavi (MB-0147) (MB).
- 20k. Malaveria mimula comb. nov., male: Paraguay,
Paraguari, Sapucai, 08.IX.2014 (MB-0152) (MB).
- 20m. ‘Paryphthimoides.’ grimon male: Brazil (MB-05)
(MNHN).
Figure 22:
- 22a-22b, 22e-22k. Malaveria ballo sp. nov., Holoty-
pe male: Ecuador, Pichincha, Nanegalito, 2016 (MB-
0072) (MB).
- 22c-22d, 22l-22m. Malaveria ballo sp. nov., Paraty-
pe female: Ecuador, Imbabura, Parambas 3500, 1913
(MB-06) (MNHN).
Figure 23:
- 18a-18i. Malaveria rodriguezi rodriguezi sp. nov., Ho-
lotype male: Colombia, Antioquia, Frontino 2000m,
22.IV.2007(MB-0249) (MB).
Figure 24:
- 24a-24i. Malaveria rodriguezi risaralda ssp. nov., Ho-
lotype male: Colombia, Risaralda, Pueblo Rico 1400m,
20.VII.2005 (MB-0250) (MB).
Figure 25:
- 25a-25i. Malaveria bottoi sp. nov., Holotype male: Co-
lombia: Cundinamarca, Colegio 430m (MB-0019) (MB).
Figure 26:
- 26a-26b, 26e-26k. Malaveria mimula comb. nov.,
male: Paraguay, Paraguari, Sapucai, 08.IX.2014 (MB-
0152) (MB).
- 26c-26d, 26l-26m. Malaveria mimula comb. nov., fe-
male: Paraguay, Paraguari, Sapucai, 08.IX.2014 (MB-
0343) (MB).
Figure 27:
- 27a-27b, 27e-27k. Malaveria duponti sp. nov., Holoty-
pe male: Bolivia, Caranavi (MB-0147) (MB).
- 27c-27d, 27l-27m. Malaveria duponti sp. nov., Paraty-
pe female: Bolivia, Caranavi (MB-0147) (MB).
Figure 28:
- 28a-28b. Koutalina pamela comb. nov., male: Boli-
via, La Paz, Caranavi 800-1500m, January 2013 (MB-
0060) (MB).
M. Benmesbah, Á. L. Viloria & J. Murienne
62
- 28c. Koutalina pamela comb. nov., male: Peru, Cusco,
uincemil 26km SouthWest 1580m, 28-30.VI.2019
(MB-0348) (MB).
- 28d. Koutalina pamela comb. nov., male: Bolivia,
La Paz, Yungas de la Paz 1000m, 1913 (MB-03)
(MNHN).
- 28g-28m. Koutalina pamela comb. nov., male: Boli-
via, La Paz, Caranavi 800-1500m, January 2013 (MB-
0059) (MB).
- 28e-28f, 28n-28o. Koutalina pamela comb. nov., fema-
le: Peru, Cusco, uincemil 26km SouthWest 1580m,
28-30.VI.2019 (MB-0346) (MB).
