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 butteries are discussed and dened:
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 denen 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 dierent 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
redenition. Several eorts 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 buttery taxa formerly allocated in this complex
genus.
e discovery in French Guiana of a new species, pre-
viously misidentied as Magneuptychia modesta (Butler,
1867) (Brévignon 2008, Brévignon & Benmesbah 2012)
prompted us to examine similar looking taxa classied 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 butteries 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-Maraet 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-dened 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
identications 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-specic 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 sucient condence 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 specic 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-
tícas, 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 Bvignon 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: DFWdorsal forewing; DHWdorsal hindwing;
FW–Forewing, FWL–Forewing length, HW–Hindwing,
TLType 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; aer 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
reex digital camera. Illustrations were edited using the
M. Benmesbah, Á. L. Viloria & J. Murienne
10
soware 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 buttery 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 identied with good condence 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
classication 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 modica-
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 reection; 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 reection. 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 dened, 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 dened,
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 dened
with one or two pupils, ocellus in
Cu1-Cu2
well dened
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 dened, ocellus in M2-M3 usually ovoid
slightly displaced basally compared to the others, ocellus
in Rs-M1
usually much smaller than precedents, usually
well dened, 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 inuence 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 intraspecic 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 infraspecic 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 amplication
results even from degraded material. Amplication 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-
signicant 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)
Supercially 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 diers 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
dierent 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., diers by m1-m2 less curved
and m2-m3 more curved than in Emeryus. Modestia gen.
nov., diers from Magneuptychia, Paryphthimoides, Cissia
and Koutalina gen. nov., also by the more curved m2-m3
(straight in all these genera). Hindwing venation, although
supercially similar with all the following genera, diers
in the signicantly 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
dierences 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 dierence 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)
Butteries 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