Á. L. Viloria
26
ANARTIA
Publicación del Museo de Biología de la Universidad del Zulia
ISSN 1315-642X (impresa) / ISSN 2665-0347 (digital)
Anartia, 32 (junio 2021): 26-52
http://zoobank.org/urn:lsid:zoobank.org:pub:FD77F0F8-4196-4D8F-8654-135ED2F9FAD4
“The Hermeuptychia Papers”
“Los papeles de Hermeuptychia”
Ángel L. Viloria
Centro de Ecología, Instituto Venezolano de Investigaciones Científicas (IVIC), Apartado Postal 20632,
Caracas 1020-A, D.C., Venezuela
Correspondence: aviloria@ivic.gob.ve, sebastianviloriacarrizo@gmail.com
(Received: 15-11-2020 / Accepted: 28-02-2021 / Online: 15-10-2021)
ABSTRACT
Investigations on the nomenclature of the American buttery genus Hermeuptychia Forster, 1964 (Lepidoptera: Nym-
phalidae, Satyrinae) yielded the reinstatement of three species, the proposal of four new combinations, and the recognition
of eight new synonymies; mainly through the study and scrutiny of information published in scientic papers, books, and
web pages; but also by the examination of its diversity represented in biological collections. e novel taxonomic arrange-
ment proposed for this genus is as follows: Hermeuptychia acmenis (Hübner, 1823), comb. nov. (= H. hermybius Grishin,
2014, syn. nov.), Hermeuptychia atalanta (Butler, 1867), Hermeuptychia camerta (Cramer, 1780), sp. restit., comb. nov.,
Hermeuptychia canthe (Hübner, [1811]), sp. restit., comb. nov. (= Neonympha pimpla C. Felder & R. Felder, 1862, syn.
nov. = Euptychia maimoune Butler, 1870, syn. nov. = Euptychia nana Möschler, 1877, syn. nov.), Hermeuptychia cautheus
(Godart, [1824]), sp. restit., comb. nov. (= Hermeuptychia intricata Grishin, 2014, syn. nov.), Hermeuptychia cucullina
(Weymer, 1911), Hermeuptychia gisella (Hayward, 1957) (= Hermeuptychia hermes var. hermesina Forster, 1964, syn. nov.
= Hermeuptychia clara Nakahara, Tan, Lamas & Willmott, 2016, syn. nov.), Hermeuptychia harmonia (Butler, 1867) (=
Euptychia calixta Butler, 1877 = Hermeuptychia callixta Forster, 1964, syn. nov.), Hermeuptychia hermes (Fabricius, 1775)
(= Hermeuptychia hermes isabella Anken, 1994), Hermeuptychia lupita (Reakirt, [1867]) and Hermeuptychia sosybius (Fa-
bricius, 1793) (= Hermeuptychia hermes kappeli Anken, 1993).
Keywords: America, integrative taxonomy, Lepidoptera, nomenclature, Nymphalidae, Satyrinae.
RESUMEN
Investigaciones en la nomenclatura del género de mariposas americanas Hermeuptychia Forster, 1964 (Lepidoptera:
Nymphalidae, Satyrinae), principalmente mediante el estudio y escrutinio de información publicada en artículos, pági-
nas web y libros cientícos, pero también en el examen de su diversidad representada en colecciones biológicas, arrojó la
restitución de tres especies, la propuesta de cuatro nuevas combinaciones y el reconocimiento de ocho nuevas sinonimias.
El nuevo arreglo taxonómico propuesto para este género es el siguiente: Hermeuptychia acmenis (Hübner, 1823), comb.
nov. (= H. hermybius Grishin, 2014, syn. nov.), Hermeuptychia atalanta (Butler, 1867), Hermeuptychia camerta (Cramer,
1780), sp. restit., comb. nov., Hermeuptychia canthe (Hübner, [1811]), sp. restit., comb. nov. (= Neonympha pimpla C.
Felder & R. Felder, 1862, syn. nov. = Euptychia maimoune Butler, 1870, syn. nov. = Euptychia nana Möschler, 1877,
syn. nov.), Hermeuptychia cautheus (Godart, [1824]), sp. restit., comb. nov. (= Hermeuptychia intricata Grishin, 2014,
syn. nov.), Hermeuptychia cucullina (Weymer, 1911), Hermeuptychia gisella (Hayward, 1957) (= Hermeuptychia hermes
var. hermesina Forster, 1964, syn. nov. = Hermeuptychia clara Nakahara, Tan, Lamas & Willmott, 2016, syn. nov.), Her-
meuptychia harmonia (Butler, 1867) (= Euptychia calixta Butler, 1877 = Hermeuptychia callixta Forster, 1964, syn. nov.),
DOI: https://doi.org/10.5281/zenodo.5571293
The Hermeuptychia Papers
27
Hermeuptychia hermes (Fabricius, 1775) (= Hermeuptychia hermes isabella Anken, 1994), Hermeuptychia lupita (Reakirt,
[1867]) y Hermeuptychia sosybius (Fabricius, 1793) (= Hermeuptychia hermes kappeli Anken, 1993).
Palabras clave: América, Lepidoptera, nomenclatura, Nymphalidae, Satyrinae, taxonomía integradora.
2014; Rick 2014) and the most recent in 2016. e genus
has attracted considerable attention of molecular system-
atists during the last ten years, and several interesting and
sometimes lengthy papers have been produced as a result
of their comparative studies. is increasing interest has
been also supplemented by initiatives in comparative mor-
phology, biology and ecology (Seraphim et al. 2014, Cong
& Grishin 2014, Cosmo et al. 2014, Warren et al. 2014a,
2014b, Anken et al. 2015, Tan & Lucky 2016, Nakahara et
al. 2016 and Austin 2018).
is work considered the relevant information avail-
able in the literature about the butteries of the genusHer-
meuptychia, critically emphasizing the evaluation of the
taxonomic hypotheses put forward in publications that
have appeared during the last 30 years.I call these publica-
tions “e Hermeuptychia papers”.
is is not yet a taxonomic revision, which will surely
involve much more comparative, morphological, molecu-
lar, ethological, and biogeographical work. is is just an
exhaustive exercise in nomenclature that any alpha-taxon-
omist should do before attempting a revisionary study or
even the description of new species.
Our resulting taxonomic hypotheses are substantially
dierent from what have been traditionally accepted. ey
bring some questions about how integral is the so called
“integrative taxonomy”, and about its failure in recogniz-
ing the fundamental role of nomenclature and biogeogra-
phy to biological systematics.
MATERIAL AND METHODS
Two phases of research were accomplished in parallel;
the examination, dissection, and comparison of buttery
specimens preserved in entomological collections (mainly
public natural history museums, but also to a lesser degree,
private collections), and the critical and careful reading of
the content of numerous scientic papers, books, and web
pages. Comparative, logical, and coherent criteria were
conceived to aim an objective taxonomic arrangement. It
was a work of comparative zoology. When the synonymies
were built in regard to the species treated in this work,
special attention was paid to the original descriptions and
re-descriptions, the dierent phylogenetic hypotheses in
which they are included, and the congruence of the latter
with basic notion of biogeography.
INTRODUCTION
e genus Hermeuptychia Forster, 1964 (Lepidoptera:
Nymphalidae, Satyrinae) is only found in the American
continent, at tropical and subtropical latitudes, and from
sea level to around 2,500 m, in the equatorial Andes. It
comprises a group of small-sized species of brown butter-
ies, characterized by possessing few external characteristics
that determine their interspecic dierences: basically, the
dorsal surface of their wings is entirely fuscous and of lim-
ited usefulness for specic discrimination; the ventral basic
pattern for all its taxa, which is developed over a brown
background, is characterized by two darker discal lines
(uniform or variably undulate) running more or less paral-
lel to the outer margins, which are also delineated by ne
submarginal and marginal lines. Darker marks develop over
the veins that close the discal cell. ey possess ocelli (usu-
ally black, ringed with ochraceous color, with a single silver
white pupil) in the postdiscal region. ere are six ocelli on
the hindwing – exceptionally ve in some individuals –,
and a variable number on the forewing, from none to ve.
e male genital armature, which has been reasonably well
studied has a dome-like tegumen, a lanceolate uncus, and
well-developed brachia (subunci) that may reach the length
of the uncus but are usually shorter. e saccus is digitiform
and well-developed, and the valvae distinctively elongated,
with the distal half thinner than the basal. e aedeagus is
relatively thick and straight, devoid of particularities of tax-
onomic interest. eir female genitalia, known for two or
three species, have so far been set aside as a very informative
source of characters for comparative morphology.
e taxonomy of the species included in this genus
goes as far back as J. C. Fabricius, who described the rst
known species from Brazil in 1775, Papilio hermes, the
type of the genus erected by W. Forster almost two hun-
dred years later. Species belonging to Hermeuptychia have
a rich taxonomic history, as it will be seen in this work.
eir names have been historically combined within gen-
era like Papilio Linnaeus, 1758, Satyrus Latreille, 1810,
Oreas Hübner, [1807], Euptychia Hübner, 1818, Neonym-
pha, Hübner, 1818, Megisto Hübner, [1819], Cissia Dou-
bleday, 1848 and Yphthimoides Forster, 1964. Most species
were described between the 18th and 19th centuries, but
a few were discovered later, mainly in the 20th century,
with two highly advertised cases in 2014 ([Anonymous]
Á. L. Viloria
28
Many illustrations of Hermeuptychia butteries have
been published, including photographs of most type
specimens known, the habitus of all species (described and
undescribed), and male genital structures. Both drawings
and photographs are from a wide variety of localities and
habitats of the American continent. Good quality data on
the immature stages of at least three species are available,
as well as data on hostplants, behavior, and ecology. e
amount of data is overwhelming. e preparation of this
notes has taken nearly ten years, and a supplement with
the examined material will be timely published. Opinions
on authority, typication, priority, synonymy, and hom-
onymy followed the provisions of the International Code
of Zoological Nomenclature (ICZN 1999).
Abbreviations: comb. nov.: new combination; et al.:
et alii (and others); GLAHM: Hunterian Museum and
Art Gallery, University of Glasgow, Glasgow, UK; IML:
Museo de Ciencias Naturales de la Fundación Miguel
Lillo, Tucumán, Argentina; in litt.: in litteris; MS: manu-
script; NHMUK: e Natural History Museum, London,
UK; nom. nov.: nomen novum, new name; nom. nud.: no-
men nudum, name not available; op. cit.: opere citato, work
cited; sp. restit.: species restored or reinstated; syn. nov.:
new synonym; TL: Type Locality; USNM: United States
National Museum, Smithsonian Institution, Washington,
D. C., USA; ZMHB: Museum für Naturkunde Berlin
(Zoological Collections), Germany.
RESULTS
Nomenclature and synonymy of the species belonging to
Hermeuptychia
e names hermes Fabricius and sosybius Fabricius have
long been erroneously applied to other Hermeuptychia
species (and to other taxa belonging to dierent genera)
in a signicant number of papers and popular books. e
synonymic citations for these two taxa in this article are
not exhaustive.
Hermeuptychia Forster, 1964: 87-91.
Type species: Papilio hermes Fabricius, 1775; by original
designation (Forster, 1964: 87).
Hermeuptychia Forster; Miller, 1968: 93, 168; 1976:
1; Miller & Brown, 1981: 191, 241; Viloria, 1990:
xx, 88; Anken, 1993: 418; 1994: 283, 1995a: 8;
1995b: 237, 238; Poole & Lewis, 1996: 621, 904;
Opler & Warren, 2002: 42; Lamas, 2003: 69; 2004:
220; Piñas Rubio, 2004: 3; Murray & Prowell, 2004:
74, 77, 78; Brown, Jr. et al., 2007: 478; Koçak & Ke-
mal, 2007: 121; 2015: 54; Pelham, 2008: 404, 492,
627; Pulido & Andrade, 2009: 520, 551; Marín et
al., 2009: 237, 239 tabl. 2 (part), 241, 242; 2011:
6, 8, 10; 2012: 209, 212 g. 1C (tRNA structure),
213 g. 2 (comparative sequence alignment), 214
g. 3 (trees) part; Peña et al., 2010: 249, 252, 254;
Matos-Maraví et al., 2013: 55, 64; Seraphim et al.,
2014: 39, 40, 43, 45, 46; Cong & Grishin, 2014: 43,
44, 45, 46, 47, 51, 52, 54, 55, 56, 57, 58, 60, 61, 65,
66, 67, 68, 72, 73, 76, 78, 79, 80, 81, 86, 87, 88; Cos-
mo et al., 2014: 82, 86; Warren et al., 2014a: 83, 85;
2014b: 45, 46, 47, 48, 50; Anken et al., 2015: 157,
158; Nakahara et al., 2016: 77, 78, 80; Marín et al.,
2017: 768, 769, 774, 775, 781; 2019: 91, 98; Austin,
2018: 311; Espeland et al., 2019: 116, 120, 121, 122;
Benmesbah et al., [2021]: 19, 28.
Hermeuptychia acmenis (Hübner, 1823), comb. nov.
[Megisto acmenis Hübner, [1819]: 54], nom. nud.
Megisto acmenis Hübner, 1823: pl. [41], gs. 233 (male
dorsal), 234 (male ventral). [TL: “Baltimore”].
Neonympha acmenis (Hübner); Westwood, 1851: 375;
Morris, 1860: 10; Weidemeyer, 1864: 527; Herrich-
Schäer, 1865: 70; Mombert, 1869: 562.
Euptychia acmenis (Hübner); Butler, 1867: 476-477;
1868: 24; 1877a: 119; Gerhard, 1878: 2; Kirby,
1871: 50; 1880: 296; Staudinger, 1887: 227, pl. 81
[row 3] (misidentication of Godartiana byses (Go-
dart)); Mabilde, 1896: 97 (misidentication); Eimer
& Fickert, 1897: 314; Sanders, 1904: 366 (misiden-
tication); Weymer, 1911: 206, t. 51b [2]; Gaede,
1931: 437; D’Abrera, 1988: 778 g. [row 4] male
ventral.
Megisto acmenis (Hübner); Scudder, 1875: 244; Mös-
chler, 1876: 35; Barnes & Lindsey, 1922: 90.
[Hermeuptychia hermes (Fabricius); D’Abrera, 1988:
777 g. [row 1, g. 3] male verso (misidentica-
tion)].
Yphthimoides acmenis (Hübner); Lamas, 2004: 223.
[Hermeuptychia sosybius (Fabricius); Glassberg, 2007:
139 [row 2, g. 3]; 2012: 257 [g. Hidalgo Co., TX]
(misidentications)].
Yphthymoides [sic] acmenis (Hübner); Teston & Cor-
seuil, 2008: 47 (misidentication).
Hermeuptychia hermybius Grishin, 2014: 44, 49, 50,
68-81, 83 gs 48–59, 60b, e, h, k, 61b, 62a–m, 64q–
z, 67 (distribution map) part, 70, 75 g 63 (mor-
phometry, in part), 76 g. 65, 78 g. 66 (cladogram)
part, 83 g, 70 (life cycle), 84, 85, 86, 87, 88, syn.
nov. [TL: “E of Brownsville, Texas, USA”].
The Hermeuptychia Papers
29
Hermeuptychia hermybius Grishin; Nakahara et al.,
2016: 83; See et al., 2018: 51; Marín et al., 2019: 91.
Megisto acmenis is a small satyrine buttery species that
was originally illustrated by Hübner, who depicted it with
reasonable detail in dorsal and ventral views (1823: pl.
[41], gs. 233, 234). Its type locality is “Baltimore”, pre-
sumably in what corresponds to the great city of the state
of Maryland, founded in 1729 on the Patapsco River, near
its mouth in the Chesapeake Bay (United States of Amer-
ica). For a long time, it has been an ignored, neglected, or
omitted taxon, because historically no other records of a
similar insect have been known, and because most of the
authors questioned its geographical origin from very early
on. Despite many eorts, it has not been possible to relo-
cate the biological material studied by Hübner to illustrate
this species. Hemming (1937) and Calhoun (2018) men-
tioned that at least part of the butteries available to Hüb-
ner to produce his original illustrations were later kept in
the form of wings pasted in scrapbook-like volumes (or
possibly as lepidochromes?), whose whereabouts is cur-
rently unknown.
Butler (1868: 24) pointed out that William Henry
Edwards (in litt.) referred to this species as inhabitant
of the “southern states”. In this way Butler, relying on
Edwards’ authority, seemed to admit that it is reliably a
North American species. However, Scudder (1875: 244)
denounced the opposite: “this buttery in unknown to
American lepidopterists, as it seems very doubtful wheth-
er it can be considered American, more particularly as
three [sic] other Satyrids given by Hübner in this same
work [referring to the “Zütrage zur Sammlung exotischer
Schmettlinge”] under the names of Symphaedra, Alcandra,
Mycalesis, Otrea and Yphthima [sic], Philomela (all said
by him to come aus Georgien in Florida) have since been
discovered to be East Indian species.” However, this as-
sertion is arguable, as Hübner had already identied and
named this taxon as early as 1819, because the name ap-
peared printed in that year in the “Verzeichniss bekannter
Schmettlinge [sic]” (therefore, at that moment a nomen
nudum). It would be interesting to investigate whether
the taxa referred to by Scudder have also been known al-
ready earlier than 1823.
Hübner’s gures of Megisto acmenis are suciently clear
and unequivocal, so that they qualify as iconotypical rep-
resentation of the species. e International Code of Zoo-
logical Nomenclature, allows for the designation of one of
these gures as the lectotype (ICZN 1999: 82, Art. 74.4),
and although I interpret the illustration of the underside
habitus (Hübner, 1823: pl. [41], gs. 234) as such, there is
no intention in this work to allocate a type for this taxon.
Aer extensive comparative studies with numerous
species of Satyrinae from all continents, I consider that ac-
cording to size, wing shape, and coloration pattern; Megis-
to acmenis evidently represents a species of Hermeuptychia
Forster, 1964, whose uniqueness and rarity consists of it
being devoid of ocelli. D’Abrera (1988: 778 g. [Row 4]
male ventral) presented the photographic image of what
in his opinion is the only authenticated specimen (with-
out abdomen) of “Euptychia acmenis” in the collections
of the NHMUK. His illustration represents an insect that
compares favorably with the iconotypic image of Hübner.
I examined this specimen in June 2016 (its detached abdo-
men is kept in a gelatin capsule). It appears to represent a
species of Hermeuptychia by the detection of a few features
in the wing pattern, such as the system of lines or bands,
the small dark dash on the veins that distally close the dis-
cal cells and the punctual vestiges of the ocellar system.
D’Abrera’s identication appears correct, but it was taken
from an old identity label in the drawer, and it must be at-
tributed to the work of a former curator of the collections
in London. is specimen is labeled with an elongated
piece of paper which is folded several times. It bears the
faint shadow of a handwritten line with fountain pen ink,
which is illegible.
e abundance of Hermeuptychia butteries in wild
populations may be usually medium to high (DeVries
1987, Viloria 1990, Peixoto & Benson 2009, Warren et al.
2014a), so it is also its individual frequency (pers. obs. in
Venezuela, 1988-2019). For this reason, it seems unusual
that over almost two centuries since its discovery, Her-
meuptychia acmenis Hübner comb. nov., is not adequately
represented in collections, at least in the form that Hübner
examined it. is suggests that we might be facing a true
case of ecological rarity.
In some common species of the genus Hermeuptychia,
the size and conformation of the ocelli usually vary mark-
edly among individuals of the same population. However,
in entomological collections it is appreciated that a small
proportion of individuals of a species can be represented
by extreme variations in the number and size of ocelli that
are expressed on the ventral side of their wings. Complete
ocellar reduction or suppression appears to be very rare;
that is, infrequent. However, it does indeed occur, and par-
ticularly in some individuals recently recognized through
the extensive comparative studies of Cong & Grishin
(2014). It is unknown whether this particularity has to
do with the proportion of alleles within populations or
it is a phenomenon linked to seasonality, as it occurs in
well-studied cases in African satyrids, mainly mycalesines
and melanitines (Brakeeld & Larsen 1984, Braby 1995,
Dongmo et al. 2018).
Á. L. Viloria
30
e three cryptic taxa detected and discriminated by
Cong & Grishin in their exhaustive work on the genus
Hermeuptychia in the United States of America (op. cit.),
have genetically and unobjectionable specic statuses.
One such species shows a very large morphological varia-
tion in the degree of expression of its ocelli and of the band
system of the ventral surface of the wings. In its most ex-
treme forms, the maximum reduction is evident (Cong &
Grishin, 2014: 70, gs. 53, 55) or the suppression of bands
and ocelli (https://www.butteriesofamerica.com/L/her-
meuptychia_hermybius_types3.htm), in the manner of
the two credited illustrations of Hermeuptychia acmenis
(Hübner) comb. nov., of which I have already spoken. No
other Hermeuptychia appears to express that kind of ex-
treme morphotype. erefore, I propose the consideration
of Hermeuptychia hermybius Grishin as a junior synonym
(syn. nov.) of H. acmenis (Hübner) comb. nov., as the lat-
ter is the oldest available name.
As for the authenticity of the locality “Baltimore” re-
corded by Hübner, the case requires further consideration.
References are found to at least 23 North American his-
torical localities – some now missing – in the states of
Alabama, California, Colorado, Delaware, Georgia, Indi-
ana, Iowa, Kansas (2), Kentucky (2), Maryland, Michigan,
New York, North Carolina (2), Ohio (3), Tennessee and
Vermont (USA), New Brunswick and Ontario (Canada)
(https://roadsidethoughts.com/ga/baltimore-xx-wilkes-
prole.htm). A good proportion of such sites do not
qualify as a possible place of origin of any species of Her-
meuptychia, because they are found in latitudes outside the
biogeographic distribution of this genus, others are place
names applied to villages or hamlets founded aer the
publication of Hübner’s work.
Personally, I am inclined to think that the locality pro-
vided by this author is inaccurate and perhaps it refers
(as seems to happen in many historical cases of doubtful
or mistaken provenance records of plants and animals,
particularly in dates before 1850) to a simple error origi-
nated by the supplier, to the reference of the place where
the sample was acquired or the port where it was shipped
to Europe. e geographic records provided by Cong &
Grishin correspond only to the southeast of the state of
Texas, along the current border with Mexico. Little is
known about its real distribution, both in the USA and
in Mexico. It is worth pointing out that in 1823 and be-
fore, Texas was still part of the Mexican territory. ere
is an unfortunate information gap regarding travelers and
naturalists who explored the northern region of Mexico
in the two decades prior to 1819-1823, the years of the
publications of Hübner’s name and illustration of Megisto
acmenis (Silva 1944, Mayer 1961, Glantz 1982, Morales
1986). Both Hemming (1937) and Calhoun (2018) claim
that the origin of much of Hübner’s North American ma-
terial is unknown.
Hermeuptychia atalanta (Butler, 1867)
Euptychia atalanta Butler, 1867: 474-475, pl. 39, g.
12. [TL: “Venezuela”].
Euptychia atalanta Butler; Butler, 1868: 24; 1877a:
119; Kirby, 1871: 50; Weymer & Maassen, 1890:
17, 74 (misidentications); Weymer 1911: 207, pl.
48a [3] (mistakenly as a synonym of Euptychia fa-
llax (C. Felder & R. Felder)); Riley & Gabriel, 1924:
9 (types 1 male, 1 female); Gaede, 1931: 449 (last
one as a synonym of Euptychia hermes (Fabricius));
Hayward, 1958b: 236 (as a synonym of Euptychia
hermes (Fabricius)).
Euptychia fallax (C. Felder & R. Felder) var. atalanta
Butler; Butler, 1870b: 251.
[Euptychia hermes (Fabricius); Longsta, 1912: 54;
255, 264, 306, 307, 309, 310, 313, 320, 323, 329,
578-579 (misidentications in part)].
[Euptychia hermes fallax (C. Felder & R. Felder); Bee-
be, 1951: 9 (misidentication)].
Hermeuptychia atalanta (Butler); Lamas, 2004: 220;
Seraphim et al., 2014: 42 g. 1 (cladogram) part,
45, 46, 47, suppl. material: gs. S3 [male genitalia],
S4 [underside habitus]: TO02, morphogroup 11;
Cong & Grishin, 2014: 51, 78 g. 66 (cladogram)
part; Cosmo et al., 2014: 82-86, gs. 1A (egg), B (1st
instar), C (2nd instar), D (3rd instar), E (4th instar),
F (prepupa), G-I (pupa), J (adult male, dorsal), K
(adult male, ventral), g. 2 (morphological details
egg, larvae), g. 3 (chaetotaxy 1st instar); Anken et
al., 2015: 157, 159; Nakahara et al., 2016: 81, 83,
84 (all except Lamas, misidentications of Hermeup-
tychia hermes (Fabricius)); Marín et al., 2019: 91;
Ríos-Málaver et al., [2021]: 85, 99.
Two syntypes of Euptychia atalanta Butler, 1867 (male
and female) are recorded in the NHMUK by Riley & Ga-
briel (1924). ey were collected by David Dyson in the
surroundings of Caracas in 1846. Photographs of the male
syntype taken by G. Lamas and N. V. Grishin are available
online (Warren et al. 2012): [Ven / Venezuela pur. from
Dyson 46-75/ B.M. Type No Rh 3228, Euptychia atalanta
♂ Butl. / Syntype, (examined)].
Forster (1964) did not take this taxon into account in
his monograph. It was also missed by D’Abrera (1988). La-
mas (2004) correctly transferred this species to the genus
Hermeuptychia Forster.
The Hermeuptychia Papers
31
Specimens gured by Seraphim et al. (2014, suppl.
material, gs. S3 [male genitalia], S4 [underside habitus]:
TO02, morphogroup 11) under this name belong to Her-
meuptychia hermes (Fabricius). is error is reected in
the entire content of their work; where the type species of
the genus, H. hermes, the most common species found in
southeastern Brazil, is misidentied as H. atalanta.
Cosmo et al. (2014) also misidentied typical individu-
als of H. hermes – whose type specimens come from south-
eastern Brazil –, described and illustrated their life cycle
under the erroneous name of Hermeuptychia atalanta
(Butler). is notorious mistake has been repeated in the
works of Anken et al. (2015) and Nakahara et al. (2016).
Seraphim et al. (2014: 46) considered that “H. atalanta
is highly widespread”. It is in fact one of the several middle
elevation satyrine buttery species, endemic to the moun-
tains of the Cordillera de La Costa, in northern Venezuela
(Ríos-Málaver et al. [2021]). erefore, it is probably the
most geographically conned taxon of the genus. It does
not occur in the neighboring Venezuelan Andes or the
Serranía del Turimiquire.
Longsta (1912) alleged that individuals of H. atalanta
taken in the vicinity of Caracas represented a dry season
form (with reduced ocelli) of the same species he cap-
tured in Panamá, Trinidad and Tobago (probably mainly
H. canthe (Hübner) sp. restit., comb. nov. See below). As
most authors, Longsta misidentied three dierent taxa
under the name of Euptychia hermes.
Hermeuptychia camerta (Cramer, 1780), sp. restit.,
comb. nov.
Pap.[ilio] Nymph.[alis] Gemm.[ata] camerta Cramer,
1780: 10, pl. 313, g. F. [TL: “Surinamsche”, “Suri-
name”].
Papilio camertus [sic] Cramer; Herbst, 1796: 91, pl.
195, g. 8; Godart, [1824]: 495 (as a synonym of Sa-
tyrus sosybius (Fabricius)); Weidemeyer, 1864: 527;
Strecker, 1878: 149; Skinner, 1898: 33 (last three as
a synonym of Neonympha sosybius (Fabricius)).
Satyrus camerta (Cramer); Ménétriés, 1829: 191 (misi-
dentication of H. hermes (F.)); Verloren, 1837: 111,
203 (as asynonym of S. sosibius [sic] (Fabricius)).
Papilio camerta Cramer; Hübner, [1819]: 54 (as a sy-
nonym of Megisto euridice [sic] Linnaeus); Godart,
[1824]: 495 (as a synonym of Satyrus sosybius (Fa-
bricius)); Doubleday, [1845]: 137 (as a synonym of
Euptychia sosybius (Fabricius), misidentications, in
part); Lamas, 2004: 220; Pelham, 2008: 404, 616
(last two as a synonym of Hermeuptychia hermes
(Fabricius)).
Neonympha camerta (Cramer); Westwood, 1851: 375;
Herrich-Schäer, 1865: 70; Prittwitz, 1865: 310-
311 (misidentication of H. hermes (Fabricius));
Strecker, 1878: 149 (as a synonym of Neonympha
sosybius (Fabricius)).
Euptychia camerta (Cramer); Butler, 1867: 462-463;
1868: 16; 1877: 119; Kirby, 1871: 48; Butler & Dru-
ce, 1874: 335; Druce, 1876: 213; Godman & Salvin,
1880a: xxxii, 74, 80, 86-87, 88, pl. 8, gs. 6, 7 (ma-
les); 1880b: 122; Staudinger, 1887: 226, 324, 333,
t. 81 (male identied as E. hermes (F.)) (misidenti-
cations in both cases); Shannon, 1898: 356; Bayern,
1901: 266; 1908: 312 (all misidentications); Go-
dman, 1901: 656 (erroneously as a synonym of E.
hermes (Fabricius), E. sosybius (Fabricius), E. fallax
(C. Felder & R. Felder) and E. maimoune (Butler));
Sanders, 1904: 366-367 (in part misidentications
of H. hermes (Fabricius) and H. canthe (Hübner)
sp. restit., comb. nov.); Clark, 1905: 9; Longsta,
1908: 54, 1912: 255, 578 (both mistakenly as a sy-
nonym of E. hermes (F.)); Weymer, 1911: 207; Dyar,
1913: 635 (in part misidentications of H. canthe
(Hübner) sp. restit., comb. nov., and H. gisella (Ha-
yward)); 1914: 143 (misidentication); Herrera,
1923: 133 (misidentication); Davis, 1928: 59 (mi-
sidentication); Gaede, 1931: 441-442; D’Abrera,
1988: 777 (the latter as a synonym of E. hermes (Fa-
bricius)).
[Euptychia atalanta Butler var.; Butler, 1867: 475
[Pará] (misidentication)].
[Euptychia hermes (Fabricius); Kaye, 1904: 180; Longs-
ta, 1912: 54; 255, 264, 306, 307, 309, 310, 313,
320, 323, 329, 578-579 (misidentications in part)].
Megisto camerta (Cramer); Barnes & Lindsey, 1922: 90
(as a synonym of Megisto euridice [sic] (Linnaeus).
[Hermeuptychia hermes (Fabricius); Seraphim et al.,
2014: 39, 40, 41, 42 g. 1 (cladogram) part, 45, 46
(misidentication in part), suppl. material: gs. S3
(male genitalia), S4 (underside habitus): PA04, mor-
phogroup 10 (misidentication)].
Cramer (1780: 10, pl. 313, g. F) illustrated this taxon
from a Surinamese specimen, which has not been located.
Hermeuptychia camerta (Cramer, 1780) sp. restit.,
comb. nov., is a very distinctive but uncommon species
that is found ying in sympatry with the much more com-
mon H. canthe (Hübner, [1811]) sp. restit., comb. nov.,
in the lowlands of the Guiana Shield (but the latter has
a wider distribution, both latitudinally and altitudinally).
e rst one is recognized by the uniform small size of its
hindwing ventral ocelli, smaller than those of the type spe-
Á. L. Viloria
32
cies of the genus, H. hermes (which apparently does not
occur in northern South America).
e phenotype of the specimen illustrated by Seraphim
et al. (2014: suppl. material, gs. S3 [male genitalia], S4
[underside habitus]: PA04, morphogroup 10), and espe-
cially the male genitalia, with a remarkably long saccus, are
features determined as characteristic of H. camerta (Cra-
mer) sp. restit., comb. nov. In this case the specimen il-
lustrated comes from the state of Pará. I have dissected and
examined specimens with similar genitalia, from south-
eastern Venezuela (Viloria: Los Satyrinae de Venezuela, in
prep.) and French Guiana. e habitus of the specimens
with these peculiar male genitalia ts well Cramer’s pub-
lished illustration of his Papilio camerta, even though it is
markedly stylized.
is taxon is found to be partly sympatric with H.
canthe sp. restit., comb. nov., but H. camerta sp. restit.,
comb. nov., is apparently less frequent.
It is probable that Longsta’s mention of one “dry sea-
son” specimen of “E. hermes” from Trinidad among sev-
eral other “wet season” specimens (Longsta 1912: 579),
represents the only insular record known for this taxon.
e satyrine buttery fauna of Trinidad & Tobago (Cock
2014, 2017) is representative — but less diverse — of what
is known for the Guiana Shield lowlands.
Hermeuptychia canthe (Hübner, [1811]), sp. restit.,
comb. nov.
Oreas Strigata canthe Hübner, [1811]: pl. [87]: male
[gs.] 1, 2; female [gs.] 3, 4, nom. nov. [TL: ?Su-
rinam].
Oread. strig. canthe Hübner, [1819]: 54 (as a synonym
of Megisto euridice [sic] (Linnaeus), erroneous.
Euptychia canthe (Hübner); Westwood, 1851: 373;
Butler, 1867: 474; 1868: 23; [1870]a: 13; 1870b:
251; Weymer, 1911: 207; Gaede, 1931: 449 (the last
six as a synonym of Euptychia hermes (F.)).
Neonympha pimpla C. Felder & R. Felder, 1862: 177,
syn. nov. [TL: Rio Negro].
Neonympha canthe (Hübner); Herrich-Schäer, 1865:
70.
[Neonympha nana Herrich-Schäer, 1865: 70], nom.
nud., syn. nov.
Neonympha pimpla C. Felder & R. Felder; Herrich-
Schäer, 1865: 70; Kirby, 1871: 49 (as a synonym
of Euptychia renata (Cramer [sic]) var.); Hayward,
1958b: 236 (as a synonym of E. hermes (Fabricius)).
Euptychia pimpla (C. Felder & R. Felder); Butler, 1867:
470; 1868: 20; 1877a: 119; Weymer, 1911: 208;
Gaede, 1931: 449.
[Euptychia fallax C. Felder & R. Felder, var.; Butler,
1867: 474 [Venezuela] (misidentication)].
Oreas (strigata) canthe (Hübner); Butler, 1867: 475;
1868: 23; Burmeister, 1878: 210 (all as a synonym of
Papilio hermes Fabricius); McDunnough, 1938: 11,
190 (as a synonym of Megisto hermes (Fabricius));
Hayward, 1958b: 236 (as a synonym of Euptychia
hermes (Fabricius)); Poole & Lewis, 1996: 962; La-
mas, 2004: 220; Koçak & Kemal, 2007: 959; 2015:
1463; Pelham, 2008: 404, 616 (all as a synonym of
Hermeuptychia hermes (Fabricius)).
Euptychia maimoune Butler, 1870b: 251, pl. 1, g. 4.,
syn. nov. [TL: Pebas, E. Peru].
Euptychia maimoune Butler; Kirby, 1871: 643; Distant,
1876: xiii; Butler, 1877a: 119; Weymer, 1911: 207;
Riley & Gabriel, 1924: 34 (type male); Gaede, 1931:
454; Hayward, 1951: 229; 1958b: 236 (as a synon-
ym of E. hermes (Fabricius)); Lamas, 1969: 283;
D’Abrera, 1988: 789.
[Euptychia camerta (Cramer); Butler & Druce, 1874:
335; Druce, 1876: 213; Godman & Salvin, 1880a:
xxxii, 74, 80, 86-87, 88, pl. 8, gs. 6, 7 (males);
1880b: 122; Godman, 1901: 656; Sanders, 1904:
366-367; Clark, 1905: 9; Dyar, 1913: 635 (all, in
part misidentications)].
Euptychia nana Möschler, 1877: 323-324, syn. nov.
[TL: “dem innern Surinams”].
Euptychia nana Möschler; Kirby, 1877: 843; Butler,
1877a: 128; Weymer, 1911: 207; Gaede, 1931: 449;
Hayward, 1951: 229; 1958b: 236 (as a synonym of
H. hermes (Fabricius)); Forster, 1964: 88; Lamas,
2004: 220; Pelham, 2008: 404 (all, except Kirby, as a
synonym of E. hermes (Fabricius)).
[Euptychia hermes (Fabricius); Kaye, 1904: 180 (misi-
dentication, in part); DeVries, 1983: 722-723, g.
(misidentication, at least in part)].
[Euptychia sosybius (Fabricius); Kaye, 1904: 180 (misi-
dentication)].
[Hermeuptychia hermes (Fabricius); Forster, 1964: 88,
g. 60 (male genitalia), 89-90; Barcant, 1970: 143,
160, pl. 13, [g.] 13; Viloria, 1990: xx, xxvii, 88-94,
271, gs. 44 (male dorsal), 45 (male ventral), 46
(male genitalia); Garwood & Lehman, 2011: 274
[gs.]; Gernaat et al., 2012: 242, 243, pl. 38, gs. 7
& 8 (all misidentications)].
Hermeuptychia pimpla (C. Felder & R. Felder); Forster,
1964: 88, g. 62 (male genitalia); Lamas, 2004: 220;
Seraphim et al., 2009: 331; 2014: 42 g. 1 (clado-
gram) part, 44, 45, 46, suppl. material: gs. S3 [male
genitalia], S4 [underside habitus]: [uncoded], mor-
phogroup 02 (misidentication of Hermeuptychia
The Hermeuptychia Papers
33
harmonia (Butler)); Peña et al., 2010: 247, 250 g. 2
(phylogenetic tree) part, 251 g. 3 (cladogram) part,
253 g. 4 (divergence time) part (misidentication);
Cong & Grishin, 2014: 51, 78 g. 66 (cladogram)
part (misidentication); Nakahara et al., 2016: 81
g. 4 (phylogenetic tree) part, 83, 84 (misidentica-
tion); Marín et al., 2019: 91 (misidentication).
Oreas canthe Hübner; Miller & Brown, 1981: 191, 241
(as a possible synonym of H. hermes (Fabricius)).
[Cissia hermes (Fabricius); DeVries, 1987: 258, 276,
277, 298, pl. 41, g. 3 (misidentication)].
[‘Euptichia’ [sic] hermes (Fabricius); Convey, 1990: 169
(misidentication)].
Hermeuptychia maimoune (Butler); Lamas, 2004: 220;
Seraphim et al., 2014: 42 g. 1 (cladogram) part,
45, 46, suppl. material: gs. S3 [male genitalia], S4
[underside habitus]: CO04, morphogroup 08 and
TO01, morphogroup 09; Cong & Grishin, 2014:
51, 66, 78 g. 66 (cladogram) part, 86; Nahakara et
al., 2016: 81 g. 4 (phylogenetic tree) part, 83, 84;
Marín et al., 2019: 91.
Hübner [1811] illustrated male and female (both dorsal
and ventral) of what he might have thought to be good ex-
amples of the Fabrician P. canthus (described from “Amer-
ica boreali”, 1775). e latter was a name preoccupied by
a distinct and dierent Linnean taxon, also from North
America (Linnaeus 1767). us, due to the similarity of
their names, it is reasonable to interpret Hübner’s proposal
of Oreas Strigata canthe as a nomen novum to replace the
Fabrician junior homonym of the Linnean species. e
taxonomic history of Papilio canthus Fabricius has been
well investigated and described by Cardé et al. (1970). e
taxon does exist, and it has an aspect very similar to Hüb-
ner’s O. S. canthe, but obviously its rst name is invalid and
unavailable (see below, under next species).
However, Hübner’s attempt to replace the aforemen-
tioned Fabricius’ homonym, cannot be accepted because
the provenance of the specimens used by Hübner for his
illustrations was later found to be South America, and
therefore they represent a dierent taxon.
In a detailed study of Jacob Hübner’s published works
and manuscripts, Hemming (1937) reproduced an incom-
plete list of localities of the species gured by that author
in his Sammlung exotischer schmetterlinge, taken from
Hübner MS. 35, in the library of the Royal Entomologi-
cal Society of London. Among those, there is a record of
“? Surinam” for “Papilio nymphalis canthe” (Hemming,
1937, vol. 1: 122; vol. 2: 424; Pelham, 2008: 404). Hüb-
ner’s illustrations do indeed represent one male and one
female of the most common species of Hermeuptychia
found not only in Surinam, but also in the entire adjacent
Guyana Shield region and beyond, in a considerable ex-
tension of the Amazon basin, to the lowlands of northern
Venezuela including Margarita island, Trinidad & Tobago
and perhaps northeastern Colombia.
is taxon diers from P. camerta Cramer sp. restit.,
comb. nov., by the presence of ventral forewing subapical
ocelli (visible in Hübner’s ne and trustworthy gures, but
see below under H. sosybius), and much more developed
ocelli on the hindwing underside. In a second mention of
this taxon as Oreas canthe, Hübner ([1819]: 514), intro-
duces another bit of taxonomic noise not only by repeat-
ing the confusion of the Fabrician species with Megisto
euridice [sic] (Linnaeus, 1763a) and Papilio canthus Lin-
naeus, 1767, but also mistakenly adding Papilio camerta
Cramer, 1780 to the synonymic list; the distinctive South
American taxon already discussed above. However, unlike
Fabricius and Herbst, who had confusions regarding the
identity of other satyrine taxa, Hübner recognized Papilio
argante Cramer as a valid, separate species.
ere is one male (?) specimen, allegedly a syntype
of Neonympha pimpla C. Felder & R. Felder, 1862, syn.
nov., in NHMUK, lacking its head and abdomen [Rio
Negro / Felder colln. / Rothschild Bequest B. M. 1939-1
(examined), photographs in Warren et al. 2012]. I do not
recognize it as a possible type of N. pimpla, as its appear-
ance does not entirely t the original description of this
taxon, which seems to depict instead the phenotype of H.
canthe, a true lowland Guianan-Amazonian species. e
male genitalia illustrated by Forster (1964: 89, g. 62) is
erroneous.
Seraphim et al. (2014, suppl. material) misidentied
Hermeuptychia harmonia (Butler) as H. pimpla (C. & R.
Felder) (gs. S3 [male genitalia], S4 [underside habitus]:
[uncoded, San Antonio, Colombia], morphogroup 02).
Euptychia nana Möschler (1877) syn. nov., was de-
scribed from a small sized single male specimen that is not
in good condition. It was collected in the inner land of
Surinam and represents Hermeuptychia canthe (Hübner,
[1811]) sp. restit., comb. nov. Forster (1964: 88) errone-
ously synonymized E. nana with H. hermes claiming to
have had its type in front of him. e referred specimen,
a lectotype designated by L. D. Miller in 1989, is in the
ZMHB (examined). It bears a typical Forster’s label stat-
ing “Präparat Nr. 154 Zool. Staatssammlung München”;
however, judging from the prevalence of its entire abdo-
men it seems that Forster never dissected its genitalia.
e explanation of this confusion can be deducted from
the observation of Forster’s gure of the male genitalia
of H. hermes, which is wrong (Forster, 1964: 88, g. 60).
e representation of the genital structures of true Her-
Á. L. Viloria
34
meuptychia canthe sp. restit., comb. nov., is comparable
to what I have dissected and examined from northwestern
and southeastern Venezuela, Trinidad and French Guiana
(illustrated in Viloria 1990: 90 g. 46, under the wrong
name of H. hermes). Euptychia maimoune Butler (1870b:
251, pl. 1, g. 4) syn. nov., described apparently from a
single male specimen (Riley & Gabriel 1924, Warren et al.
2012), taken at the Amazonian locality of Pebas in Peru
(NHMUK, examined), also represents the same taxon.
Illustrations of male underside habitus and genitalia
under the name of Hermeuptychia maimoune (Butler) syn.
nov., in Seraphim et al. (2014, suppl. material, gs. S3, S4:
CO04, morphogroup 08, but not TO01, morphogroup
09 [08 sic!], which represents H. hermes) are in our view
correct, but they had to be referred to its prior name, Her-
meuptychia canthe sp. restit., comb. nov.
Hermeuptychia cautheus (Godart, [1824]), sp. restit.,
comb. nov.
Papilio canthus Fabricius, 1775: 486, nom. praeoc. (nec
Papilio canthus Linnaeus, 1767; TL: America Sep-
tentrionali). [TL: America boreali].
Papilio canthus Fabricius; Fabricius, 1781: 64; 1787:
31; 1793: 157; 1796: 120; Gmelin, [1790]: 2285 (in
part misidentication of Papilio canthus Linnaeus,
Papilio eurydice Linnaeus and Papilio argante Cra-
mer); Harris, 1862: 306.
Papilio canthus Cramer [sic]; Jones, 1785 [vol. VI]: pl.
38 (misidentication of Papilio argante Cramer,
1779).
Papilio canthus Linnaeus [sic]; Herbst, 1796: 70, pl.
192, gs. 5, 6 (misidentication of Papilio argante
Cramer, 1779); Hübner, [1819]: 54 (in part) (as a
synonym of Megisto euridice [sic] (Linnaeus), erro-
neous); Butler, 1867: 503 (in part).
Satyrus cautheus Godart, [1824]: 465, nom. nov.
Satyrus cantheus [sic] Godart, [1824]: 493-494.
Satyrus canthus (Fabricius); Verloren, 1837: 83 (as a sy-
nonym of Papilio argante Cramer).
Neonympha canthus (Linnaeus) [sic]; Westwood, 1851:
375; Morris, 1860: 10 (both in part misidentica-
tions).
Neonympha cantheus [sic] (Fabricius [sic]); Morris,
1860: 10 (misidentication in part); Weidemeyer,
1864: 527; Herrich-Schäer, 1865: 69; Mombert,
1869: 562.
Papilio (D. F.) canthus (part.) Fabricius; Butler, [1870]
a: 13 (as a synonym of P. argante Cramer).
E.?[uptychia] canthus (Linnaeus [sic]); Kirby, 1871: 55
(misidentication in part).
Satyrus cantheus [sic] Godart; Scudder, 1875: 243 (as a
synonym of Papilio eurydice Linnaeus).
Satyrus canthus (Linnaeus [sic]); Kirby, 1877: 704 (mi-
sidentication in part).
Euptychia canthus (Linnaeus [sic]); Kirby, 1880: 296
(misidentication in part).
Megisto canthus (Fabricius); Barnes & Lindsey, 1922:
90 (as a synonym of Megisto euridice [sic] (Linnaeus)
and also as a synonym of Megisto camerta (Cramer)).
Hermeuptychia intricata Grishin, 2014: 43, 50, 51, 61-
68 gs. 23-31, gs. 32-35, 40-43, 62n, 68 part, 72
gs. 60c, f, i, l, 73 g. 61a, 76 gs. 64i-p, 77 g. 65
part, 78 g. 66 (cladogram) part, 79, 80 g. 67 (dis-
tribution map) part, 81 g. 68 part, 84, 85, 86, 87,
88., syn. nov. [TL: Brazos Bend State Park, Texas,
USA].
Hermeuptychia intricata Grishin; Warren et al., 2014a:
83, 84 g. 1a (habitus dorsal, lacking androconia), c
(habitus ventral), e (habitus dorsal), g (habitus ven-
tral), j (male genitalia), l (female genitalia), n (distri-
bution map) part, 85; 2014b: 44, 45 gs. 1a (habitus
dorsal), c (habitus ventral), e (holotype dorsal), g
(holotype ventral), I (paratype dorsal, in part); 46
gs. 2 b, d, f, h (wing scales), 47 gs. 3, l-n, 48 gs.
4 c-l, u –x, 49 g. 5 (distribution map), 50, 51; Tan
& Lucky, 2016: 3-4, gs. 6 (comparative wing cha-
racters), 7 (male genitalia compared); Austin, 2018:
307-313, g. 1 (partial life cycle), g. 2 (last larval
instar), g. 3 (habitat); Nakahara et al., 2016: 83;
Marín et al., 2019: 91.
e synonymy of the homonym Papilio canthus Linnae-
us, 1767, a dierent and distinct taxon, now considered a
junior synonym of Satyrodes eurydice (Linnaeus, 1763a)
(Nymphalidae: Satyrinae, Elymniini) is not included here.
An explanation for that case and an introduction to the
case of Hermeuptychia cautheus (Godart, [1824]) sp. res-
tit., comb. nov., needs the recapitulation of a fragment of
the work of Cardé et al. (1970) about S. eurydice (pp. 74,
notes by the present author inserted in square brackets):
“Taxonomic History: the Euptychia names. – e ta-
xonomy of L.[ethe] e.[urydice] eurydice is complicated
by confusion with Yphthimoides (= Euptychia) argu-
lus (Godart) [currently, Emeryus argulus (Godart)].
is problem was not noted by dos Passos, and it is
reviewed here.
Fabricius (1775) reworked the description of canthus,
adding “immaculatis” to the upperside diagnosis and
altering various details. e “immaculatis” may have
been inferred from the lack of reference to spots in the
The Hermeuptychia Papers
35
earlier descriptions, but it seems more likely that Fabri-
cius was working from some other insect he confused
with the Linnean one. In 1779 Cramer described and
gured a species from Surinam as Papilio argante. is
name is a junior homonym of Papilio argante Fabricius
1775 (now Phoebis argante, Pieridae). Fabricius synon-
ymized argante Cramer to canthus (Fabricius, 1781),
improperly emending it to arganthe in synonymy. (Ar-
ganthe is not available as a replacement name because
it was proposed in synonymy). He repeated this usage
in 1787 and 1793. His own descriptions of “canthus”
do not t Cramer’s gure well.
Godart (1821 [sic]) recognized that three species were
included in the Fabrician concept “canthus” and at-
tempted to end the confusion by redescribing the true
canthus (translating Linnaeus), and naming two new
entities, argulus and cantheus [sic]. Godart’s argulus is
a replacement name for the preoccupied argante and
is the oldest valid name of this taxon. Cantheus [sic]
is a renaming o the entity Fabricius rst thought was
canthus, theretofore without a valid name. e identi-
ty of this animal cannot be determined if, as appears,
Fabrician specimens o “canthus” do not exist.”
pp. 76 (notes by the present author inserted in square
brackets):
“Cantheus [sic], which is the unknown animal Fabri-
cius confounded with canthus and then with argante,
usually appears in the synonymy of eurydice = canthus,
but its only proper claim there is its mistaken use in sy-
nonymy by Morris (1860). We have removed cantheus
Godart from the synonymies of the other entities and
regard it as a nomen dubium, presumably a species of
Euptychia sens. lat. Its synonymy is:
‡ Papilio canthus (nec Linnaeus 1767): Fabricius 1775
(partim), Syst. Ent.: 486; 1781, Spec. Ins. 2: 64;
1787, Mant. Ins. 2: 31; 1793, Ent. Syst. 3(1): 157.
Satyrus cantheus [sic] Godart 1821, Encyl. Meth. 9:
465, 493; type locality “l’Amerique septentrionale”;
type not investigated, probably never existed.
Godart’s description of cantheus erroneously cites
Fabricius, “Species Insectorum” or “Mantissa Insecto-
rum”. e name is misspelled “cautheus” in the heading
on page 465.”
ese extraordinary observations, written about y
years ago, remain fundamentally valid, but a few facts need
to be corrected or added:
1. It is true that Fabricius (1781) wrongly listed Papilio
eurydice Linnaeus (1767) and Papilio arganthe [sic]
Cramer (1779) as synonyms of his P. canthus. Howe-
ver, he later suppressed this idea in Mantissa insector-
vm, tom. II (Fabricius, 1787: 31). Some contempo-
rary authors not only ignored this signicant change
of criterion, but they introduced further confusion
to the case. For instance, Herbst (1796) applied the
name P. canthus Linnaeus without making any dis-
tinction from its junior homonym P. canthus Fabri-
cius (1775). His illustrations of P. canthus (Herbst,
1796: pl. 192, gs. 5, 6) seem to be copied from the
previously published illustrations of Papilio argan-
te (Cramer, 1779: pl. 204), a very dierent, larger
satyrine buttery described from Surinam, whose
name, as explained by Cardé et al. (1970), was preoc-
cupied and subsequently replaced by Satyrus argulus
Godart ([1824]). Zacca et al. (2020) designated a
neotype for Godart’s taxon and proposed it as the
type species of the genus Emeryus Zacca, Casagran-
de & Mielke, 2020. Members of Emeryus are also su-
percially distinct from any Hermeuptychia species
by having a larger size, and ve, bipupillated ocelli
on the hindwing underside (not six, monopupilla-
ted, as in Hermeuptychia). In his text, Herbst (op cit.,
pp. 70), refers again to the synonymy between Pa-
pilio eurydice Linneaus, 1763a (prioritary, currently
Satyrodes eurydice (L.)) and its replacement name P.
canthus Linnaeus, 1767 (invalid, and not the Fabri-
cian P. canthus), and erroneously repeated Papilio
arganthe [sic] Cramer as another synonym.
2. In 2016, during a consultation with Jeanne Robin-
son, curator of the historical material of Lepidoptera
at the Hunterian Museum in Glasgow (GLAHM),
the present author detected a male individual of a
species of Hermeuptychia associated to a cabinet
label with the manuscript legend “Pap. Canthus /
Fabr. pag 64 N° 288”, current reference number of
the specimen 127581. Its habitus represents reasona-
bly well the physical description published by Fabri-
cius (1775, 1781, 1787, 1793) for this species. It is
known that Fabricius was heavily involved with the
curation of the insect collection of Dr William Hun-
ter during several of his visits to London, particu-
larly between 1780 and 1787 (Fabricius 1784, Hope
1845, Armitage 1958, Tuxen 1967, Vane-Wright
2007, Hancock 2015): “I knew it [Hunter’s collec-
tion] very well because I laid it out myself and contri-
buted to its gradual increase in size” (Hancock 2015:
158, translating the h letter of Fabricius’ Briefe of
1784). us, the cabinet labels of the Hunterian co-
llection document directly the taxonomic identica-
tions by Johann Christian Fabricius, the only autho-
Á. L. Viloria
36
rity in the case of his Papilio canthus. Fabricius used
his own work Species insectorum (1781) as a source
catalogue in his lengthy process of identication and
curation. us, each specimen identied by him was
cross-referenced by the page and species name in
that work. e current labels in the Hunterian cabi-
nets were handwritten by Matthew Baillie, Hunter’s
nephew and inheritor, whose style corresponds with
the same handwriting of the Trustees’ manuscript ca-
talogue signed o in 1785 (Hancock 2015). Because
of its clear provenance and documented association
with Fabricius, this Hermeuptychia specimen quali-
es at least as a Neotype of Papilio canthus Fabricius,
1775 (Viloria & Robinson, MS), which by that re-
cognition is no longer a nomen dubium.
3. P. canthus Fabricius, 1775, not a nomen dubium,
however is unavailable by homonymy (with P.
canthus Linnaeus, 1767). erefore, the species to
which it corresponds should adopt the next avai-
lable name, proposed by Godart ([1824]). Most
authors, if not all – including Cardé et al. (1970) –,
have wrongly recorded the name Satyrus cantheus
Godart, assuming that the appearance of the printed
word “cautheus” in page 465 of Godart’s work must
be taken just as a misspelling of “cantheus” (page
493). e appearance of the rst spelling is not a
mere heading as stated by Cardé et al. (1970: 76),
because it is immediately followed by Godart’s short
diagnosis of the Fabrician taxon: “Ailes entières, d’un
brun-noirâtre et sans taches en dessus: dessous des
inférieures avec six jeux. Fab.”
It should instead be objectively taken as the rst
use of the replacement name, even if it was a printing
error appearing 28 pages prior to a dierent spelling
of the name applied to the same entity followed by
a more extense description (several cases like this are
known among the Lepidoptera, for instance within
the Nymphalidae Satyrinae, Paramacera Butler,
which should have been Paramecera, or Praefaunala
Forster, the true rst spelling of Praefaunula of the
auctorum). In any case, a provision of the ICZN,
called the Principle of the First Reviser (ICZN
1999: 30, Art. 24.2) allows the present author to
make this decision.
4. e putative type specimen of Papilio canthus Fabri-
cius referred above, is a male of a species of Hermeup-
tychia that bears the distinctive external features of
a taxon recently detected in North America. It was
named Hermeuptychia intricata Grishin, 2014, syn.
nov. e Fabrician/Hunterian specimen has been
identied as such by the apparent absence of an an-
droconial patch on the forewing recto, a diagnostic
feature, characteristic of its sibiline and sympatric H.
sosybius (Fabricius), but also by the presence of other
less stable characteristics extensively studied by the
experts (Cong & Grishin 2014, Warren et al. 2014a,
2014b, Tan & Lucky 2016, Austin 2018).
Hermeuptychia cucullina (Weymer, 1911)
[Euptychia cucullina Staudinger, in litt.], nom. nud.
Euptychia calixta Butler f. cucullina Weymer, 1911:
209, pl. 48 c, g. [3]. [TL: wrongly stated by Wey-
mer as Choco, Colombia; it is in fact Chaco (La
Paz), 2-3000 m, Bolivia].
Euptychia calixta Butler var. cucullina Weymer; Gaede,
1931: 441 (as a synonym of Euptychia calixta Butler);
Hayward, 1958c: 64, 65, g. 13 (male genitalia).
Hermeuptychia cucullina (Weymer); Forster, 1964: 88,
89, g. 65 (male genitalia), 91, pl. 30, gs. 8 (male
recto), 9 (male verso); Anken, 1994: 283, 286, 288
(misidentication); 1995a: 8, 9, 11 gs. 1 (habi-
tus dorsal), 2 (habitus ventral) (misidentication);
1995b: 237-239, gs. 1 (habitus dorsal), 2 (habi-
tus ventral) (misidentications of another taxon);
Lamas, 2003: 69, 145, lám. 25, g. 273; 2004: 220;
Piñas Rubio, 2004: 6, 29, gs. 211, 212 (misiden-
tication of H. gisella (Hayward)); Gottsberger &
Silberbaur-Gottsberger, 2006: 75; Seraphim et al.,
2014: 39, 42 g. 1 (cladogram) part, 44, 45, 46, 47,
suppl. material: gs. S3 [male genitalia], S4 [undersi-
de habitus]: PE03, morphogroup 05 (misidentica-
tion in part)]; Cong & Grishin, 2014: 44, 45, 51, 52,
78 g. 66 (cladogram) part, 85, 86; Nakahara et al.,
2016: 81 g. 4 (phylogenetic tree) part, 84; Marín et
al., 2019: 91.
Euptychia ‘hermesina’ Staudinger, nom. nud.; D’Abrera,
1988: 777 gs. [row 2] male recto & verso, Anken,
1994: 286 (misidentications).
Euptychia cucullina Weymer; D’Abrera, 1988: 777 g.
[row 2] male verso (misidentication of H. gisella
(Hayward)); Anken, 1994: 286; 1995a: 9; 1995b:
237; Fagua, 1999: 358 (misidentications).
Hermeuptychia cuculina [sic] (Weymer); Peña et al.,
2010: 247, 250 g. 2 (phylogenetic tree) part, 251
g. 3 (cladogram) part, 253 g. 4 (divergence time)
part.
Hermeuptychia pompilia Marín et al., 2011: 7 g. 3 (mi-
sidentication), nom. nud.
Described from at least two male specimens collected
in 1893-1894 by one of the Garlepp brothers (Gustav and
The Hermeuptychia Papers
37
Otto), in Chaco, at 2,000-3,000 m, [Yungas de] La Paz,
Bolivia. ey are deposited at ZMHB. One of them desig-
nated lectotype by G. Lamas in 1994. e other individual
is very peculiar, as it has only ve ocelli (instead of six) on
its hindwing verso (examined). Photographs of these spec-
imens taken by Lamas and Grishin are available at Warren
et al. (2012).
Male genitalia correctly illustrated by Forster (1964:
89, g. 65).
Seraphim et al. (2014, suppl. material) correctly identi-
ed Hermeuptychia cucullina (Weymer) from Peru (gs.
S3 [male genitalia], S4 [underside habitus]: PE03, mor-
phogroup 05).
Hermeuptychia gisella (Hayward, 1957)
Euptychia gisella Hayward, 1957: 112-113, 119, g. 2.
[TL: Yungas del Palmar, 2000 m, Bolivia].
[Euptychia camerta (Cramer); Dyar, 1913: 635 (misi-
dentication in part)].
Hermeuptychia gisella (Hayward); Forster, 1964: 88, g.
61 (male genitalia), 90; pl. 30, gs. 5 (male recto), 6
(male verso); Anken, 1994: 286; 1995a: 9; 1995b:
237, 238 (in part, misidentications); Lamas, 1997:
217; T. Racheli & L. Racheli, 2001: 326; Matos-Ma-
raví et al., 2013: 60 g. 3A (divergence times) part,
61 g. 4 (ancestral areas of distribution) (misidenti-
cation); Seraphim et al., 2014: 42 g. 1 (cladogram)
part, 45, 47, supplementary material: gs. S3 [male
genitalia], S4 [underside habitus] [MT13 -morpho-
group 06] (misidentication); Cong & Grishin,
2014: 44, 45, 51, 52, 78 g. 66 (cladogram) part,
85, 86 (misidentication); Nakahara et al., 2016: 81
g. 4 (phylogenetic tree) part, 83, 84 (misidentica-
tion); Marín et al., 2019: 91.
Hermeuptychia hermes var. hermesina Forster, 1964: 90
(mistakenly as a synonym of H. hermes (Fabricius)),
syn. nov. [TL: Chaco, Yungas de La Paz, 2-3000 m,
Bolivia].
Hermeuptychia hermes f. hermesina (Staudinger), in
litt.; Forster, 1964: 90 (mistakenly as a synonym of
H. hermes (Fabricius)); Lamas, 2004: 220 (as a sy-
nonym of H. hermes (Fabricius)).
[Euptychia cucullina Weymer; D’Abrera, 1988: 777 g.
[row 2] male verso (misidentication)].
Euptychia gisella Hayward; D’Abrera, 1988: 789; La-
mas, 2004: 220 (as a synonym of Hermeuptychia cu-
cullina (Weymer)).
Euptychia hermesina (Forster); Anken, 1994: 288.
Euptychia hermes hermesina (Forster); Anken, 1994:
288.
Hermeuptychia hermes hermesina Forster; Anken,
1994: 290; 1995a: 9; 1995b: 237, 238 (in part, misi-
dentication); Pelham, 2008: 404 (as a synonym of
H. hermes (Fabricius)).
Hermeuptychia [n. sp.] Lamas, MS; Lamas, 2004: 220
[# 138].
[Hermeuptychia cucullina (Weymer); Piñas Rubio,
2004: 6, 29, gs. 211, 212 (misidentication)].
Hermeuptychia clara Nakahara, Tan, Lamas & Will-
mott, 2016: 77, 78, 79 gs. 1A, B (male, dorsal and
ventral), C, D (female, dorsal and ventral), 80 gs.
2A (male wing venation), B, C (male and female pal-
pi), D (male foreleg), E (female foreleg), 3A, B, C
(male genitalia), D, E, F (female genitalia), 81 g. 4
(phylogenetic tree) part, 82 gs. 5 (map, locality re-
cords), 6 (habitat & habitus), syn. nov. [TL: uimi-
Cóndor Mirador rd., Zamora-Chinchipe, 1000 m,
Ecuador].
Hermeuptychia clara Nakahara, Tan, Lamas & Will-
mott; Marín et al., 2019: 91.
Most of the type specimens of the satyrine buttery
species described by Hayward from Bolivia in 1957 are
missing (Benmesbah et al. [2021]: 51-52). However, the
holotype of Euptychia gisella is in IML (Tucumán, Argen-
tina). Photographs by G. Lamas are available at Warren et
al. (2012).
Hayward (1957) presented a very gross drawing of the
male genitalia of the holotype of E. gisella. It was probably
executed without the aid of a camera lucida (as it appears
to be the case for his many illustrations of buttery genita-
lia). In this case, both the saccus and the aedeagus appear
to be partly cut or lost.
Male genitalia of this species illustrated by Forster
(1964: 88, g. 61) have a very similar aspect of those of
Hermeuptychia clara Nakahara, Tan, Lamas & Willmott
syn. nov. (Nakahara et al. 2016: 80, gs. 3 A-C). How-
ever, they are dicult to compare with the caricaturesque,
inaccurate drawings of Hayward. Forster (1964) seems to
have correctly identied individuals of H. gisella (Hay-
ward) (pl. 30, gs. 5 [dorsal] and 6 [ventral]) from Yungas
del Palmar, its type locality. As the preceding species H.
cucullina (Weymer), H. gisella (Hayward) is a montane
to upper montane (1,000-2,000 m), Andean species, dis-
tributed along the eastern slopes of the main Andes from
southern Colombia to Bolivia, with some morphological
variation along the latitude gradient. Hermeuptychia gisel-
la is not documented by Nakahara et al. (2016).
Illustrations of a Brazilian individual MT13 -morpho-
group 06- in Seraphim et al. (2014, supplementary mate-
rial, gs. S3 [male genitalia], S4 [underside habitus]) do
Á. L. Viloria
38
not represent H. gisella (Hayward). Biogeographically, it
is predictable that this Andean taxon does not occur in
Brazil.
Hermeuptychia harmonia (Butler, 1867)
Euptychia harmonia Butler, 1867: 478, pl. 39, g. 17.
[TL: uito, Ecuador].
[Euptychia harmonia Butler, var.; Butler, 1867: 478
(Ecuador)].
Euptychia harmonia Butler; Butler, 1868: 24; 1870b:
251; 1877a: 120; Kirby, 1871: 50; Godman & Sal-
vin, 1880a: 88; Dognin, 1891: 93; Weymer, 1911:
209, t. 48 b g. [6]; Riley & Gabriel, 1924: 23 (type
female); Gaede, 1931: 448; Lewis, 1973: 58, g. 9;
D’Abrera, 1988: 777 (in text); Manara, 1994: 23
(misidentication of Optimandes eugenia (C. Felder
& R. Felder)).
Euptychia calixta Butler, 1877a: 125, pl. 12, g. 8. [TL:
Bogotá, Colombia] [synonymy established by La-
mas, 2004: 220].
Euptychia calixta Butler; Kirby, 1877: 843; Weymer,
1911: 209, t. 48 c g. [2]; Gaede, 1931: 441; Bee-
be, 1951: 9 (misidentication of Optimandes euge-
nia (C. Felder & R. Felder)); DeVries, 1986: 332;
D’Abrera, 1988: 777 gs. [row 3] male recto & ver-
so; Kochalka et al., 1996: 212; Tobar et al., 2002:
400; García- Pérez et al., 2007: 648.
Euptychia cucullixta Weymer, 1911: 209, 1082, nom.
nud. [synonymy established by Lamas, 2004: 220].
Euptychia cuculixta Staudinger, in litt.; Gaede, 1931:
441 [synonymy established].
Hermeuptychia harmonia (Butler); Forster, 1964: 89,
g. 66 (male genitalia); Anken, 1994: 286; T. Rache-
li & L. Racheli, 2001: 325; Lamas, 2004: 220; Mur-
ray & Prowell, 2004: 69, 72 g. 1 (phylogram) part,
73 g. 2 (phylogenetic tree) part, 75 g. 3 (phyloge-
netic tree) part, 76 g. 4 (phylogeny) part; Emery
et al., 2006: 90 (misidentication); Chacón & Mon-
tero, 2007: lám. 129 [row 3, le, male ventral] (mi-
sidentication); Beccaloni et al., 2008: 334; Pulido
& Andrade, 2009: 541, 551; Seraphim et al., 2009:
331; 2014: 42 g. 1 (cladogram) part, 45, 46, 47,
suppl. material: gs. S3 [male genitalia], S4 [under-
side habitus]: CO36, morphogroup 03; Peña et al.,
2010: 247, 250 g. 2 (phylogenetic tree) part, 251
g. 3 (cladogram) part, 253 g. 4 (divergence time)
part; Garwood & Lehman, 2011: 274 [gs.]; Puli-
do & Parrales, 2011: 198; Marín et al., 2014: 204;
Cong & Grishin, 2014: 51, 78 g. 66 (cladogram),
part; Nakahara et al., 2016: 81 g. 4 (phylogenetic
tree) part, 83, 84; Hanson & Nishida, 2016: 253
g.; Marín et al., 2017: 776 g. 64 (male genitalia),
777 g. 7D (wing venation), 778 g. 8 (cladogram);
2019: 91; Glassberg, 2018: 162 g.
Hermeuptychia calixta Butler; Forster, 1964: 89, g. 67
(male genitalia [of type specimen]); Anken, 1994:
286; Lamas, 1997: 217; T. Racheli & L. Racheli,
2001: 326.
Hermeuptychia callixta [sic] Butler; Forster, 1964: 89,
syn. nov.
Cissia calixta (Butler); DeVries, 1987: 276-277, 298, pl.
41, g. 2; Chacón, 1988: 72; Van den Berghe et al.,
1995: 39; Vega, 2004: 123.
Euptychia harmonica [sic] Butler; D’Abrera, 1988: 777
gs. [row 4] male recto & verso; T. Racheli & L. Ra-
cheli, 2001: 325 (as a synonym).
[Hermeuptychia pimpla (C. Felder & R. Felder); Se-
raphim et al., 2014: 42 g. 1 (cladogram) part, 44,
45, 46, suppl. material: gs. S3 [male genitalia], S4
[underside habitus]: [uncoded], morphogroup 02
(misidentication)].
Riley & Gabriel (1924) indicated the existence of one
female type specimen of Euptychia harmonia Butler in
the NHMUK [uito coll. By M. Bourcier 50 -111 / BM
Type No. Rh 3231 Euptychia harmonia ♀ Butler / Type
(examined)]. e lectotype ♂ of Euptychia calixta Butler,
designated by L. D. Miller in 1989, is in the ZMHB [Bo-
gota Nolcken / Origin / E. calixta Butler type / Ex collect.
Staudinger (examined)]. Both are represented in photo-
graphs taken by G. Lamas and N. V. Grishin in Warren et
al. (2012).
Male genitalia of H. harmonia by Forster (1964: 89,
g. 66) appear correct. Male genitalia of the type of H.
calixta (Forster’s Präparat Nr. 155 Zool. Staatssammlung
München, label pinned with the lectotype) are also illus-
trated by Forster (1964: 89, g. 67).
Seraphim et al. (2014, suppl. material) correctly iden-
tied H. harmonia (Butler) (gs. S3 [male genitalia], S4
[underside habitus]: CO36, morphogroup 03).
is is a montane to upper montane taxon, distribut-
ed in the Northern Andes (Ecuador, Colombia, western
Venezuela) and apparently also in part of the mountains
of Central America (Panama, Costa Rica, Nicaragua). It
shows a perceptible degree of physical variability across its
range of distribution.
Hermeuptychia hermes (Fabricius, 1775)
P. [apilio] D.[anaus] F.[estivus] hermes Fabricius, 1775:
487; Jones, 1785 [vol. V]: pl. 52. [TL: “Brasilia”].
The Hermeuptychia Papers
39
Papilio hermes Fabricius; Fabricius, 1787: 32, 1793:
158; 1796: 123; Gmelin, [1790]: 2285 (in part mi-
sidentication of Papilio antonoe Cramer); Erichson,
[1849]: 600 (as a synonym of Euptychia libye (Lin-
naeus), as gured by Herbst, 1796); Herbst, 1796:
68-69, pl. 192, g. 4 (misidentication of Papilio
antonoe Cramer, 1775); Riley & Gabriel, 1924: 24
(type, male); Cong & Grishin, 2014: 54; Nakahara
et al., 2016: 77.
Euptychia hermessa Hübner, [1819]: 508, nom. nov. (in
part misidentication of Papilio antonoe Cramer,
1775); Lamas, 2004: 220; Pelham, 2008: 404, 627
(both as synonyms).
Satyrus hermes (Fabricius); Godart, [1824]: 463, 487-
488; Verloren, 1837: 36, 203 (as a synonym of Sa-
tyrus antonoë [sic] (Cramer)).
[Satyrus camerta (Cramer); Ménétriés, 1829: 191 (mi-
sidentication)].
Neonympha hermes (Fabricius); Doubleday, [1845]:
138; Westwood, 1851: 375; Herrich-Schäer, 1865:
70 (the latter as a synonym of Papilio antonoe Cra-
mer, 1775).
Euptychia hermes (Fabricius); Erichson, [1849]: 600
(probably misidentications of H. camerta (Cra-
mer) sp. restit., comb. nov., or H. canthe (Hübner)
sp. restit., comb. nov.); Butler, 1867: 475; 1868: 23;
[1870]a: 13, 297; 1870b: 251; 1877a: 119; 1877b:
112 (misidentication); Kirby, 1871: 50; 1880: 296;
Butler & Druce, 1874; 336 (misidentication); Mös-
chler, 1877: 323; Butler, 1877a: 125, 128; Burmeis-
ter, 1878: 210-211; Weymer & Maassen, 1890: 99
(misidentication); Sharpe 1890: 569 (misidenti-
cation); Dognin, 1891: 33 (misidentication); Wey-
mer, 1895: 323; 1911: 207, t. 48 a gs. [4, 5] (misi-
dentication); Kaye, 1904: 180 (misidentication of
H. camerta (Cramer) sp. restit., comb. nov. and H.
canthe Hübner sp. restit., comb. nov.); Longsta,
1908: 54; 255, 264, 306, 307, 309, 310, 313, 320,
323, 329, 578-579 (misidentications of H. camerta
(Cramer) sp. restit., comb. nov., H. canthe Hübner
sp. restit., comb. nov. and H. atalanta (Butler));
Aurivillius, 1929: 158 (misidentication); Ribei-
ro, 1931: 40 (misidentication); Travassos-Filho &
Carrera, 1946: 197; Bryk, 1952: 60, 61; Hayward,
1958a: 168 (redescription); 1958b: 231 g. 30
(male genitalia), 236-237, lám. 4, g. 139; 1960: 77;
Biezanko, 1960: 2; Emmel, 1970: 153-164, g. 1
(ocellar pattern) (misidentication); DeVries, 1983:
722-723; 1986: 331 (misidentications); D’Abrera,
1988: 777 g. [row 1, gs. 1, 2, 3] [1] male recto
(possible misidentication of H. sosybius (Fabri-
cius), [2] male verso (misidentication of H. sosybius
(Fabricius), [3] male verso (misidentication of H.
acmenis (Hübner) comb. nov.); Álvarez et al., 2005:
27-30, g. 3 (last three, misidentications).
Eupt.[ychia] ? hermes (Fabricius); Westwood, 1851:
374.
[Neonympha sosybius (Fabricius); Capronnier, 1874:
30; 1881: 103 (misidentications)].
[Neonympha camerta (Cramer); Prittwitz, 1865: 310-
311 (misidentication)].
[Euptychia fallax (C. Felder & R. Felder); Mabilde,
1896: 98 (misidentication)].
Megisto hermes (Fabricius); McDunnough, 1938: 11,
216 (misidentication).
Hermeuptychia hermes (Fabricius); Forster, 1964: 88,
g. 60 (male genitalia), 89-90 (misidentication);
Barcant, 1970: 143, 160, pl. 13, [g.] 13 misiden-
tications of H. canthe (Hübner) sp. restit., comb.
nov.); Biezanko et al., 1974: 112; Miller & Brown,
1981: 191, 241; Whittaker, 1983: 109 (misidenti-
cation); Brown, 1992: 152; Anken, 1994: 283, 284,
286, 287, 288, 289, 291; 1995a: 9; 1995b: 237, 238;
Poole & Lewis, 1996: 962, 1010; Fagua, 1999: 358
(misidentication); Maes, 1999: 21 (misidenti-
cation); Kaminski et al., 2001: 196; Schantz et al.,
2001: 214; T. Racheli & L. Racheli, 2001: 325 (mi-
sidentication); Opler & Warren, 2002: 42; Tobar
et al., 2002: 397 (misidentication); Romanowski
et al., 2003: 4; Iserhard & Romanowski, 2004: 653;
Lamas, 2004: 220; Murray & Prowell, 2004: 69,
72 g. 1 (phylogram) part, 73 g. 2 (phylogenetic
tree) part, 75 g. 3 (phylogenetic tree) part, 76 g.
4 (phylogeny) part; Piñas Rubio, 2004: 6, 29, gs.
213, 214 (misidentication); Emery et al., 2006: 90;
Marchiori & Romanowski, 2006a: 447, 450; 2006b:
1031, 1032; Brown, Jr. et al., 2007: 473, 478 (mi-
sidentication); Dessuy & de Morais, 2007: 113;
Koçak & Kemal, 2007: 959; 2015: 1463 (in part
misidentications); Chacón & Montero, 2007: lám.
129 [row 3, right, male ventral] (misidentication);
García-Pérez et al., 2007: 648, 651 (misidentica-
tion); Teston & Corseuil, 2008: 47; Beccaloni et
al., 2008: 334 (in part misidentications); Pelham,
2008: 404, 492, 627 (misidentication); Peixoto
& Benson, 2009: 1, 2, 5, 6, 9, 11; Pulido & Andra-
de, 2009: 541, 551 (misidentication); Marín et
al., 2009: 242 (misidentication); Seraphim et al.,
2009: 331; 2014: 39, 40, 41, 42 g. 1 (cladogram)
part, 45, 46 (misidentication in part), suppl. mate-
rial, gs. S3 [male genitalia], S4 [underside habitus]:
PA04, morphogroup 10 (all misidentications of
Á. L. Viloria
40
Hermeuptychia camerta (Cramer) sp. restit., comb.
nov.); Peña et al., 2010: 247, 250 g. 2 (phylogene-
tic tree) part, 251 g. 3 (cladogram) part, 253 g.
4 (divergence time) part (misidentication); 2011:
69, 80 g. 6 (cronogram) part; Garwood & Lehman,
2011: 274 [gs.]; Prado et al., 2011: 3, 4, 5 [ECO01,
ECO02, ECO03], gs. E1 (probably H. acmenis
(Hübner) comb. nov.), E2, E3 (misidentications);
Gernaat et al., 2012: 242, 243, pl. 38, gs. 7 & 8;
Córdoba-Alfaro, [2012]: 123 (misidentication);
Cock, 2014: 11; 2017: 24 (misidentications);
Cong & Grishin, 2014: 43, 44, 51, 52, 54, 66, 73,
78 g. 66 (cladogram) part, 85, 86, 87; Anken et al.,
2015: 157, 159 (misidentication); Tan & Lucky,
2016: 1; Nakahara et al., 2016: 81 g. 4 (phyloge-
netic tree) part, 83, 84 (misidentication in part);
Marín et al., 2017: 778 g. 8 (cladogram); 2019: 91.
Cissia hermes (Fabricius); DeVries, 1987: 258, 276, 277,
298, pl. 41, g. 3 (misidentication); Singer & Ehr-
lich, 1993: 249, 250, 251 g. 1 (part), 252, 253, 254
(misidentication).
‘Euptichia’ [sic] hermes (Fabricius); Convey, 1990: 169
(misidentications H. camerta (Cramer) sp. res-
tit., comb. nov. and H. canthe (Hübner) sp. restit.,
comb. nov.)]).
Hermeuptychia hermes isabella Anken, 1994: 283, 284,
287 gs. 3 a (habitus dorsal), b (habitus ventral),
288 g. 4 (male genitalia), 289-290. [TL:“Barran-
ca do Rio Amambai, Navirai, Mato Grosso do Sul,
Brasil”].
Hermeuptychia (= Cissia) hermes (Fabricius); Van den
Berghe et al., 1995: 39.
Hermeuptychia hermes isabella Anken; Anken, 1995a:
9; 1995b: 237, 238; Lamas, 2004: 220 (synonymy
established); Pelham, 2008: 404, 629; Anken et al.,
2015: 157, 158 gs. 3 (holotype dorsal), 4 (holotype
ventral), 159 (last as a synonym of H. atalanta (C.
Felder & R. Felder)).
Hermeuptychia hermes (Fabricius); Anken, 1995b: 238.
[Hermeuptychia atalanta (C. Felder & R. Felder); Se-
raphim et al., 2014: 42 g. 1 (cladogram) part, 45,
46, 47, suppl. material: gs. S3 [male genitalia],
S4 [underside habitus]: TO02, morphogroup 11;
Cong & Grishin, 2014: 51, 78 g. 66 (cladogram)
part; Cosmo et al., 2014: 82-86, gs. 1A (egg), B (1st
instar), C (2nd instar), D (3rd instar), E (4th instar),
F (prepupa), G-I (pupa), J (adult male, dorsal), K
(adult male, ventral), g. 2 (morphological details
egg, larvae), g. 3 (chaetotaxy 1st instar); Anken et
al., 2015: 157, 159; Nakahara et al., 2016: 81, 83, 84
(misidentications)].
H.[ermeuptychia] isabella Anken; Anken et al., 2015:
158, gs. 3 (holotype dorsal), 4 (holotype ventral)
(as a synonym of H. atalanta (C. Felder & R. Fel-
der).
Hermeuptychia atalanta isabella Anken; Anken et al.,
2015: 157, 159.
For this species Fabricius (1775: 487) stated a “habitat
in Brasilia”. According to Miller & Brown (1981: 241)
this provenance almost certainly corresponds to a local-
ity of the Rio de Janeiro area in southeastern Brazil. Spec-
imens were probably collected by Sir Joseph Banks in
1768 as the naturalist of the Endeavour, during the rst
of James Cook’s circumnavigations. e specimens Fa-
bricius rst examined during his second visit to London
in 1772 were cited as from “Mus. Banks”. Jones’ illustra-
tion of 1785 is also referred as to “Sr Josph Banks”. Butler
([1870]a: 13) indicates the presence of one type speci-
men in the Banksian collection (then already deposited
in the British Museum), as well as referred by Riley &
Gabriel (1924: 24, 1♂). is putative syntype currently
in the NHMUK [BM Type No Rh 5036, Papilio hermes
Fab. / Type / Syntype; cabinet label: Papilio hermes Fab.
Entomol. p. 487 n. 194 (examined); photos in Warren et
al. (2012)] lacks one of its hindwings (right). Another
possible syntype is in GLAHM, which could have come
from Joseph Banks through a contemporary exchange
with William Hunter. It is known from several sources
that J. C. Fabricius promoted the exchange of duplicate
specimens between dierent collections, especially in
London (Hancock 2015). e Glasgow specimen is as-
sociated to a Baillie’s cabinet label: Pap. hermes Fabr. pag
64 No 292; current reference number of the specimen
127582. It lacks its le forewing (Viloria & Robinson,
MS).
Herbst (1796: 68-69), Hübner ([1819]: 508) and Go-
dart ([1824]: 487) erroneously synonymized this taxon
and Papilio antonoe Cramer, 1775 (currently Megeupty-
chia antonoe (Cr.)). ey are widely separated entities. e
synonymic history of Hermeuptychia hermes (Fabricius) is
very rich in misidentications.
Genitalia gured in Forster (1964: 88, g. 60) is incor-
rect. It represents the genitalia of. H. canthe (Hübner) sp.
restit., comb. nov.
Anken (1994: g. 4) illustrated the holotype male geni-
talia of H. hermes isabella (a synonym of H. hermes). Its
general aspect is reminiscent in shape of that of H. gisella
(Hayward) illustrated by Forster (1964: 88, g. 61).
Male habitus and genitalia illustrated in Seraphim et al.
(2014, suppl. material, gs. S3 [male genitalia], S4 [under-
side habitus]: PA04, morphogroup 10, from Carajás, Pará)
The Hermeuptychia Papers
41
do not appear to represent true H. hermes (F). e saccus
of the genital armature photographed is unusually long,
like in typical specimens of H. camerta (Cramer) sp. res-
tit., comb. nov., from Surinam, Guyane Française, Guyana
and southern Venezuela. On the other hand, specimens
TO01, morphogrop 09 (it reads 08 [sic]) and TO02, mor-
phogroup 11, agree much better with H. hermes (not H.
maimoune (Butler) and H. atalanta (Butler), respectively,
as wrongly indicated for each case).
e life cycle of Hermeuptychia hermes was described
by Cosmo et al. (2014), under the erroneous identity of H.
atalanta (a montane species, endemic to the Venezuelan
Cordillera de La Costa).
Hermeuptychia lupita (Reakirt, [1867])
Neonympha lupita Reakirt, [1867]: 331 (female) [TL:
Orizaba, Veracruz, Mexico].
Neonympha lupita Reakirt; Gerstaecker, 1867: 367;
1869: 63; Hayward, 1951: 229 (as a synonym
of E. hermes (Fabricius)); Lamas, 2004: 223;
Llorente-Bousquets et al., 2006: 975.
Euptychia lupita (Reakirt); Butler, 1868: 39; 1877a:
123; DAbrera, 1988: 789.
Zischkaia lupita (Reakirt); Llorente-Bousquets et al.,
2006: 975.
Hermeuptychia lupita (Reakirt); Marín et al., 2019: 91.
e female type specimen of Reakirt’s Neonympha lupi-
ta has not been found, despite several searches and inqui-
ries in dierent museums of the United States of America.
is is probably the largest species of the genus (and not
the one represented as such in Warren et al. 2012). Only
three male specimens of bona de H. lupita are known to
the present author. ey were obtained in 2015 at middle
elevations in the mountains of the Sierra Madre del Sur,
Oaxaca, Mexico (Llorente-Bousquets et al., in prep.).
Hermeuptychia sosybius (Fabricius, 1793)
Papilio sosybius Fabricius, 1793: 219 [Jon. g. pict. 6.
tab. 52. g. 2.] [TL: ?].
Papilio sosybius Fabricius; Fabricius, 1796: 128; Her-
bst, 1796: 148 [cites Jones’ illustrations]; Hayward,
1958b: 236 (as a synonym of E. hermes (Fabricius));
Cong & Grishin, 2014: 43, 52, 54, 56 [lectotype de-
signation of Jones’ illustrations], 58 [neotype desig-
nation], 60. [TL: Savannah, Georgia, USA].
Satyrus sosybius (Fabricius); Godart, [1824]: 465, 495;
Ménétriés, 1829: 191 (mistakenly as a synonym of
Satyrus camerta (Cramer), but also a misidenti-
cation of H. hermes (F.)); Boisduval & Le Conte,
[1835]: pl. 63, gs. 1-4 (early stages).
Satyrus sosibius [sic] (Fabricius); Verloren, 1837: 111,
203.
Neonympha sosybius (Fabricius); Doubleday, [1845]:
137 (misidentications, in part); Westwood, 1851:
375; Morris, 1860: 10; Weidemeyer, 1864: 527; He-
rrich-Schäer, 1865: 70; Prittwitz, 1865: 311 (mi-
sidentication); Boisduval, 1870: 63; Capronnier,
1874: 30; 1881: 103 (misidentications of H. her-
mes (Fabricius)); Edwards, 1877: 229-231(early sta-
ges); 1883: 68, 69; 1884: 288; Strecker, 1878: 149;
Chambers, 1879: 73; French, 1886: 67, 240-242
(life cycle); Skinner, 1898: 33, xiii; Holland, 1898:
204, pl. 25, g. 5 (male); Denton, 1900: viii, 220-
221, gs. (male upper and underside), 360; Sharpe,
1914: 35; Brimley, 1921: 77; Clark, 1932: viii, 244,
331, 335, pl. 56 g. 7 (male dorsal), 8 (male ventral).
Euptychia sosybius (Fabricius); Butler, 1867: 474; 1868:
22; [1870]a: 13, 302; 1877a: 119; Edwards, 1872:
24; Kirby, 1871: 49; 1880: 296; Gerhard, 1878: 2;
Gosse, 1880: 202 (misidentication); Kaye, 1904:
180 (last one misidentication of H. canthe (Hüb-
ner) sp. restit., comb. nov.); A. H. Clark & L. F.
Clark, 1951: 8, 12, 38, 232, 238, pl. 3 g. i (male
ventral); Comstock & Vázquez, 1961: 379.
Euptychia hermes (Fabricius) var. sosybius Fabricius;
Butler, 1870b: 251.
Cissia sosybius (Fabricius); Scudder, 1875: 245; 1889,
vol. I: xxi; vol. III: 1786-1788 (imago and immatu-
re stages); Möschler, 1876: 35; Dyar, 1903: 32;
Grossbeck, 1917: 19.
Euptychia sasybius [sic] (Fabricius); Aurivillius, 1929:
158 (wrongly as a synonym of E. hermes (Fabricius)).
Megisto hermes sosybius (Fabricius); Richards, 1931:
244.
Megisto hermes (Fabricius) f. sosybius (Fabricius); Mc-
Dunnough, 1938: 11, 259 (erroneously as a synon-
ym of Megisto hermes (Fabricius).
Hermeuptychia sosybius (Fabricius); Forster, 1964: 88,
89, g. 64 [male genitalia]; Miller & Brown, 1981:
191; Anken, 1994: 286; Poole & Lewis, 1996:
1095; Calhoun, 1997: 47; Opler & Warren, 2002:
42; Lamas, 2004: 220; Murray & Prowell, 2004: 69,
72 g. 1 (phylogram) part, 73 g. 2 (phylogenetic
tree) part, 75 g. 3 (phylogenetic tree) part, 76 g.
4 (phylogeny) part; Glassberg, 2007: 139 [row 2,
g. 3] (misidentication of H. acmenis (Hübner)
comb. nov.; 2012: 257 [gs.] (misidentications of
H. cautheus (Godart) sp. restit., comb. nov. [Suolk
Co, VA] and H. acmenis (Hübner) comb. nov. [Hi-
Á. L. Viloria
42
dalgo, Co, TX]); 2018: 162 g. (misidentication);
Koçak & Kemal, 2007: 959; 2015: 1463; Pelham,
2008: 404, 492, 646; Seraphim et al., 2014: 42 g.
1 (cladogram) part, 45, 46, suppl. material: gs. S3
[male genitalia], S4 [underside habitus]: EUA03,
morphogroup 04; Cong & Grishin, 2014: 43, 44,
45, 46, 48, 49, 50, 51, 52, 53 gs. 1-9, 54, 55, 56, 57,
58, 59 gs. 10-21, 60, 61, 63, 64 gs. 36-39, 44-47,
66, 67, 68, 69, 72 gs, 60a, d, g, j, 73 g. 61c, 74 gs.
62o-z2, 75 g. 63 (morphometry, in part), 76 gs.
64a-h, 77 g. 65, 78 g. 66 (cladogram) part, 79, 80
g. 67 (distribution map) part, 81 g. 68 part, 82 g.
69 (life cycle), 83, 85, 86, 87, 88 [neotype designa-
ted. TL: Savannah, Georgia, USA]; Warren et al.,
2014a: 83, 84 g. 1b (habitus dorsal, with androco-
nia), d (habitus ventral), f (habitus dorsal), h (habi-
tus ventral), k (male genitalia), m (female genitalia),
n (distribution map) part, 85; 2014b: 44, 45 gs. 1 b
(habitus dorsal), d (habitus ventral), f (neotype dor-
sal androconia), h (neotype ventral), j (androconia),
46 gs. 2 a, c, e, g (wing scales), 47 gs. 3 a-h, 48 gs.
4 a-b, m-t, 49, 50; Anken et al., 2015: 157, 158; Tan
& Lucky, 2016: 1-5, gs. 1 (female habitus), 2 (oce-
llar pattern), 3 (distribution map), 4 (egg and rst
instar larva), 5 (early stages), 6 (comparative wing
characters), 7 (genitalia compared), 8 (pair in copu-
la); Nakahara et al., 2016: 83, 84; Marín et al., 2017:
778 g. 8 (cladogram); 2019: 91; Austin, 2018: 307-
313, g. 2 (last larval instar), g. 4 (imagos); See et
al., 2018: 51.
[Hermeuptychia hermes (Fabricius); D’Abrera, 1988:
777 gs. [row 1, g. 1] male recto with androconia
(possible misidentication)], [row 1, g. 2] male
verso (misidentication)].
Hermeuptychia hermes kappeli Anken, 1993: 418-419,
gs. 2a (male holotype dorsal), 2b (male holotype
ventral). [TL: Lake Okeechopee, Fla., USA].
Hermeuptychia hermes kappeli Anken; Anken, 1994:
288; Calhoun, 1997: 47 [synonymy established];
Lamas, 2004: 220; Pelham, 2008: 404, 630; Cong &
Grishin, 2014: 46, 51, 59 gs. 12 (holotype dorsal),
13 (holotype ventral), 60, 61; Anken et al., 2015:
157, 158, gs. 1 (holotype dorsal), 2 (holotype ven-
tral); Tan & Lucky, 2016: 2 (all as a synonym of H.
sosybius (Fabricius)).
Hermeuptychia hermes sosybius (Fabricius); Whittaker,
1983: 109 g. 2 (capture numbers) (misidentica-
tion); Raguso & Llorente, 1997: 286 (misidenti-
cation?).
Hermeuptychia sosybius kappeli Anken; Cong & Gris-
hin, 2014: 44.
H.[ermeuptychia] kappeli Anken; Anken et al., 2015:
158, gs. 1 (holotype dorsal), 2 (holotype ventral)
(as a synonym of H. sosybius (Fabricius).
In the original description of Papilio sosybius, Fabricius
(1793: 219) indicated that the specimens he examined
were from the collection of Dru Drury (London). He also
referred to illustrations of this taxon by William Jones in
his “Icones” (“Jon. g. pict. 6. tab. 52. g. 2”). Two gures
fully identied with the name sosybius, representing upper
and underside of this buttery species do appear in manu-
script volume V of Jones, with the explicit indication of be-
ing in the collection of Dru Drury (Jones, 1785 [vol. V]:
pl. 52) (available at: http://www.jonesicones.com/. e
referred illustrations were reproduced by Cong & Grishin,
2014: 53, gs. 1-3). An account of the search for surviving
Drury’s specimens of this species in the Macleay Museum
at Sydney, Australia, is given by the latter authors.
Fabricius did not designate a locality of provenance for
this taxon, and its alleged North American origin has been
taken for granted since a very early date. Many authors
have largely associated the name sosybius to a common
species found in great part of the southern United States.
On this assumption, and following a series of reasonable
criteria, Cong & Grishin (2014) established a type locali-
ty (Savannah, Georgia, USA) while designating a neotype
for P. sosybius (deposited in the USNM). But in the same
work, presumably under the rules of Article 74.4 of the
International Code of Zoological Nomenclature (ICZN
1999: 82) they also designated a lectotype [!] for the same
taxon (Jones’ watercolor painting examined and referred
to by Fabricius). Here, there is a blatant contradiction
(Articles 74 and 75 of the Code), as there cannot be two
types for the same taxon. Only one can be the bearer of
the name.
On the other hand, there are several clues that could
point out to a South American origin for the true Papilio
sosybius (Viloria & Robinson, MS), in which case the only
possibility of identity for the insect described by Fabricius
(and illustrated by Jones) as such, is what has been here
denominated Hermeuptychia canthe (Hübner) sp. restit.,
comb. nov. Should a denite proof of such an undesirable
situation emerge, a supported case for the protection of
the name sosybius for the North American buttery spe-
cies traditionally called so, must be referred to the Interna-
tional Commission of Zoological Nomenclature.
According to the current situation, gure 64 in For-
ster (1964: 89) is correct for North American H. sosybius.
Seraphim et al. (2014, suppl. material) correctly identied
Hermeuptychia sosybius (F.) as well (gs. S3 [male genita-
lia], S4 [underside habitus]: EUA03, morphogroup 04).
The Hermeuptychia Papers
43
Edwards (1877), French (1886) and Scudder (1889)
described the early stages of this species in such a detail
that it is possible to separate certain of its characters from
those of H. acmenis comb. nov. (particularly their pupae).
Cong & Grishin (2014) also described and superbly illus-
trated the life cycle of H. sosybius and H. hermybius syn.
nov., but they failed to even cite the foundational works
of those pioneers.
CONCLUSIONS
e current investigations on the nomenclature of the
American genus Hermeuptychia Forster (Lepidoptera,
Nymphalidae, Satyrinae), mainly through the study and
scrutiny of information published in scientic documents,
but also by the examination of its diversity in biological
collections, yielded the reinstatement of three species, the
proposal of four new combinations and the recognition
of eight new synonymies. e novel taxonomic arrange-
ment proposed for this genus is as follows: Hermeuptychia
acmenis (Hübner, 1823), comb. nov. (= H. hermybius
Grishin, 2014, syn. nov.), Hermeuptychia atalanta (Butler,
1867), Hermeuptychia camerta (Cramer, 1780), sp. restit.,
comb. nov., Hermeuptychia canthe (Hübner, [1811]), sp.
restit., comb. nov. (= Neonympha pimpla C. Felder &
R. Felder, 1862, syn. nov. = Euptychia maimoune Butler,
1870, syn. nov. = Euptychia nana Möschler, 1877, syn.
nov.), Hermeuptychia cautheus (Godart, [1824]), sp. res-
tit., comb. nov. (= Hermeuptychia intricata Grishin, 2014,
syn. nov.), Hermeuptychia cucullina (Weymer, 1911),
Hermeuptychia gisella (Hayward, 1957) (= Hermeup-
tychia hermes var. hermesina Forster, 1964, syn. nov. =
Hermeuptychia clara Nakahara, Tan, Lamas & Willmott,
2016, syn. nov.), Hermeuptychia harmonia (Butler, 1867)
(= Euptychia calixta Butler, 1877 = Hermeuptychia cal-
lixta Forster, 1964, syn. nov.), Hermeuptychia hermes (Fa-
bricius, 1775) (= Hermeuptychia hermes isabella Anken,
1994), Hermeuptychia lupita (Reakirt, [1867]), Hermeup-
tychia sosybius (Fabricius, 1793) (= Hermeuptychia hermes
kappeli Anken, 1993).
Any other species historically classied within Her-
meuptychia are not considered members of this genus by
the present author.
ACKNOWLEDGMENTS
ese notes are the result of many years of work, in
which I was obliged to change my mind several times. My
interest in Hermeuptychia butteries emerged as early as
1989 when I began discussing the identity of H. hermes
with the late Lee D. Miller at the Allyn Museum in Sara-
sota, USA. I dedicate this paper to his memory. For their
help in the laboratory and eldwork in Venezuela, Co-
lombia and Peru, I thank Jesús Camacho (Universidad
del Zulia), Héctor Suárez, Wilmer Rojas (IVIC, Altos
de Pipe), Gerardo Lamas, and Juan Grados (Universidad
Nacional Mayor de San Marcos, Peru). For our enriching
discussions, assistance in collections, eldwork, and supply
of entomological samples from Venezuela, I must deeply
thank my colleagues Mauro Costa (Caracas) and Indiana
Cristóbal Ríos-Málaver (IVIC, Universidad de Pamplona,
Universidad de Caldas). My former students José Enrique
Piñango, José Rafael Ferrer-Paris, Mariana Alarcón, María
Eugenia Losada, Oriana Romero, and Abimel Moreno
collaborated with this study at dierent stages, mainly by
helping me with the incipient buttery collection at IVIC
(before and aer the arrival of Cristóbal). I owe much to
G. Lamas for the time we shared years ago in several sce-
narios, talking about satyrids and bibliography, travelling
and collecting (especially in our trip to Chanchamayo in
2001), and for his teachings in nomenclature and system-
atics. José A. Clavijo, Jürg De Marmels, uintín Arias, and
Marco Gaiani (Universidad Central de Venezuela, Ma-
racay) for granting me access to the buttery collections
of the Museo del Instituto de Zoología Agrícola (MIZA-
UCV) and for their valuable assistance. I am very grate-
ful to Jean-François Le Crom and M. Gonzalo Andrade
in Bogotá for their enduring friendship, hospitality and
collaboration. My working visits to the Le Crom collec-
tion and to the Instituto de Ciencias Naturales de la Uni-
versidad Nacional de Colombia allowed me to learn much
about the taxa I dealt with in this paper. I thank also Jorge
Llorente-Bousquets, Armando Luis-Martínez, and Arturo
Arellano for the philosophical disgressions on buttery
systematics, and for the ne opportunity to do eldwork
with them in Mexico in 2015 (rediscovering H. lupita).
For providing information, photographs and specimens,
and for sharing their knowledge, I am indebted to Jim
Córdoba (Costa Rica), Mohamed Benmesbah (French
Guiana and France), and John Morrall (Trinidad & Toba-
go and United Kingdom). Years ago (1995-98) I was ben-
etted by the teachings and advice of R. I. Vane-Wright
(my former supervisor), Phil R. Ackery, Bernard D’Abrera
(with whom I shared the keys of the old type specimens
collection!), David C. Lees, George W. Beccaloni, and
Andrew F. E. Neild (then at NHMUK). I specially thank
Blanca Huertas, now Senior Curator at this institution, for
granting me access to the collections under her care to be
able to check the enigmatic acmenis specimen in 2016. In
the old days (1990s) I was kindly assisted in many ways
by Zine Ajmat de Toledo (IML), Wolfram May, Matth-
ias Nuss (ZMHB), and Walter Neukirchen (Germany).
Á. L. Viloria
44
Fundamental information for this work was received from
Jeanne Robinson (GLAHM), Robert Blackburn (Macleay
Museum, Sydney, Australia), and Marie Meister (Museum
of Zoology, University of Strasbourg, France); I acknowl-
edge their kindness and attention. anks to Edmundo
Rubio (†) for critically reading my earliest manuscript and
suggesting (too) many changes. Tito R. Barros, Gilson Ri-
vas, Ingrid B. Petit, and an anonymous reviewer indepen-
dently read this work, in parts or in full. Many thanks for
their corrections.
Research for this contribution received its last drive
during a sabbatical stay at the Centre of Latin American
Studies of the University of Cambridge (CLAS), UK (Si-
mon Bolivar Chair 2019-2020). For this privileged op-
portunity I was academically endorsed by Professors Paul
Brakeeld (Trinity College) and Chris Jiggins (St John’s
College). I thank Professors David Ibbetson (President,
Clare Hall), Felipe Hernández (Director, CLAS), and Re-
becca Kilner (Director, Museum of Zoology), as well as
Ed Turner, Sridhar Halali (Museum and Department of
Zoology), Julie Coimbra, and Chriselia DeVries (CLAS)
for their hospitality and support.
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Nota bene: is article was already fully corrected and integrated to the other parts of this journal at the time when the following paper
was published on line (March 10th, 2021). I consider that the results presented by its authors are congruent with the systematic arrange-
ment herein proposed, although our nomenclatural approaches are dierent:
Tan, D., A. Parus, M. Dumbar, M. Espeland & K. R. Willmott. 2021. Cytochrome c oxidase subunit I barcode species delineation
methods imply critically underestimated diversity in ‘common’ Hermeuptychia butteries (Lepidoptera: Nymphalidae: Satyri-
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