A remarkable new butterfly species from western Amazonia
(Lepidoptera: Nymphalidae: Satyrinae).
Una notable nueva especie de mariposa de la Amazonía occidental
(Lepidoptera: Nymphalidae: Satyrinae).
Blanca Huertas1, Gerardo Lamas2, Giovanny Fagua3, James Mallet4,
Shinichi Nakahara5 & Keith Willmott5
1 Natural History Museum London, UK. Email: b.huertas@nhm.ac.uk
2 Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos, Lima, Perú.
3 Universidad Javeriana, Bogotá, Colombia.
4 Harvard University, Cambridge, USA.
5 McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville,
USA.
Abstract
A distinctive new species of butterfly in the subtribe Euptychiina (Nymphalidae: Satyrinae), which is widespread
throughout the upper Amazon in Colombia, Ecuador and Peru, is here described. The species is provisionally placed in the
genus Magneuptychia Forster, 1964, although this is likely to change as the higher level taxonomy of Euptychiina is
resolved and the genus is reviewed in detail.
Keywords: Systematics, Euptychiina, Colombia, Peru, Ecuador.
Resumen
Se describe una nueva especie de mariposa distintiva de la subtribu Euptychiina (Nymphalidae: Satyrinae), distribuida en la
región amazónica de Colombia, Ecuador y Perú. La especie se ubica provisionalmente en el género Magneuptychia Forster,
1964, lo que puede cambiar cuando se haya resuelto la sistemática de Euptychiina y el género se revise en detalle.
Palabras clave: Sistemática, Euptychiina, Colombia, Perú, Ecuador.
Introduction
The taxonomy of the Neotropical satyrine subtribe
Euptychiina remains one of the most unresolved of any
butterfly group, both at the species and higher levels, due
to its high especies richness, cryptic morphology, limited
variation in wing pattern/morphology among some
species, generally dull coloration and, in some clades,
variability in wing pattern and paucity of useful genital
characters (e.g. Peña & Lamas 2005; Murray & Prowell
2005; Peña et al. 2006, 2010; Marín et. al 2011).
Furthermore, the subdued dark wing coloration of many
species has apparently resulted in these butterflies often
being overlooked by collectors and thus being poorly
represented in many collections. Magneuptychia Forster,
1964 is one of the largest genera of Euptychiina (Lamas
2004; Peña et al. 2006), including over 30 described
species, with at least another 10 awaiting formal
description (Lamas 2004). These butterflies inhabit
tropical wet forests throughout the Neotropical region,
from sea level to around 2000m elevation. Some species
can be among the most locally abundant butterflies in the
understorey and edges of primary and secondary forests.
The diversity of Magneuptychia is due, at least in part, to
the genus actually comprising a heterogeneous
assemblage of species. Forster (1964: 125) characterized
it as ‘distinguished by the robust structure and larger
[wing] span, by the fundamental lack of an eye spot on
the upper wing surface…and not significantly, by the
different structure of the male apparatus with an
essentially stronger uncus’ [translation from German].
These
characters
are
clearly
unreliable
as
synapomorphies since revisionary works on related
genera as well as phylogenetic studies have shown that
the genus is not monophyletic (Murray & Prowell 2005,
Peña et al. 2006, Marín et al. 2011, Huertas 2014),
containing perhaps a half dozen or more generic level
clades, and is in need of a comprehensive revision.
Recently, Costa et al. (2016) suggested that all species
apart from the type and two close relatives should be
excluded from Magneuptychia, since they do not share
several putative synapomorphies with those three species
they proposed to retain in the genus. While we agree that
the generic classification of Magneuptychia is in strong
need of revision, Costa et al. (2016) provided no new
generic combinations for excluded species, and in the
absence of a phylogenetic study to clarify the limits of
genera, we continue to treat Magneuptychia in the sense
of authors immediately prior to Costa et al. (2016).
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5
The focus of this paper is to describe a distinctive new
euptychiine species, which we provisionally place in
Magneuptychia. A few specimens belonging to the new
species described herein remained for well over a century
scattered among the various collections at NHMUK and
more recently in other museums without being named.
What are thought to be some of the oldest known
specimens were brought to the former BMNH through
the bequest of the James John Joicey Collection in 1934.
Joicey had purchased the Henley Grose-Smith Collection
in 1910 and the latter included five specimens of which
two were labelled only ‘Ecuador’, one ‘Yahuas Terr,
Peru’ and one without data. The latter, was subsequently
illustrated by D’Abrera (1988: 786-787) and labelled as
‘Euptychia? sp’. However, a further, possibly older
specimen was later found by BH in the MFNB, collected
by the German naturalist Paul Hahnel (1843-1887), who
collected in the Amazon from 1879 to 1887 for Otto
Staudinger (Staudinger 1890; Michael 1926). At MUSM,
there is a large series of specimens collected in the 1990s,
and identified by GL as a new species, that was also
listed by Lamas (2004) as "Magneuptychia n. sp. [6]".
During her revisionary work on Euptychiina, BH found
recently collected specimens with full data in various
collections and from different localities across western
Amazonia. In this paper, we combine all field and
collection data and formally name the new species.
Methods
Magneuptychia specimens were examined in major
public and private collections in Europe and the
Americas. In addition, specimens were collected by GJ &
JM, KW & SN, and GL during field work in Colombia,
Ecuador and Peru, respectively. The following acronyms
are used in the text to denote the museum collections
studied. In a few cases specimens, data and/or
photographs were sent by the relevant curators or other
researchers or field observers, when direct examination
was not possible (see Acknowledgements). Relevant
available online data and published works were also
consulted.
DATR
David Trembath collection, United Kingdom.
FLMNH McGuire Center for Lepidoptera and
Biodiversity, Florida Museum of Natural
History, University of Florida, Gainesville,
USA.
FRPI
Francisco Piñas Rubio collection, Quito,
Ecuador.
JFLE
Jean François Le Crom collection, Bogotá,
Colombia.
KWJH Keith R. Willmott & Jason P. W. Hall
collection, Gainesville, USA.
MECN Museo Ecuatoriano de Ciencias Naturales,
Quito, Ecuador.
MEFLG Museo de Entomología Francisco Luis
Gallego, Universidad Nacional, sede Medellín,
Colombia.
MFNB Museum für Naturkunde, Berlin, Germany.
MPUJ
Museo de Historia Natural de la Pontificia
Universidad Javeriana, Bogotá, Colombia
MUA
Museo de la Universidad de Antioquia,
Medellín, Colombia.
MUSM Museo de Historia Natural de la Universidad
Nacional Mayor de San Marcos, Lima, Perú.
NHMUK Natural History Museum (former BMNH
British Museum of Natural History), London,
United Kingdom.
SHNA
Shinichi Nakahara collection, Gainesville,
USA.
All specimens were photographed using digital cameras
and images of the holotype were processed using
Adobe© Photoshop CS2 version 9.0. Nomenclature for
wing veins and cells follows Comstock & Needham
(1918) and Peña & Lamas (2005). Genitalia terminology
follows Klots (1970). We refer to dorsal forewing, dorsal
hindwing, ventral forewing and ventral hindwing as
DFW, DHW, VFW, and VHW, respectively.
Genitalia dissections were made using standard
protocols, with maceration in a 10% aqueous solution of
potassium hydroxide (KOH). Due to the number of
females available and the lack of complete comparative
material, dissections of females were limited and data
relating to such dissections were not studied as
comprehensively as for the male genitalia.
Systematics
Magneuptychia pax sp. nov.
Huertas, Lamas, Fagua & Willmott
‘Euptychia’? sp.: D’Abrera 1988: 786-787, figs.
(Peru? Ecuador).
Magneuptychia sp. n. 1: Lamas, Robbins & Harvey. [1997]: 65.
Magneuptychia [n. sp.]: Lamas, 2004: 220 (no. 1416).
Magneuptychia sp. 3: Piñas, 2004: 34, figs. 271-272.
Holotype: (Fig. 1A). Male, [Colombia, Putumayo],
n[ea]r. Villa Garzón, Los Naranjos [1˚34'13''N, 75˚43'W],
8 Aug[ust]. 1977. J. Mallet leg. BMNH 2011-63.
BMNH(E) 1720531 (deposited in NHMUK).
Allotype: (Fig. 1B). Female [Colombia, Putumayo],
Puerto Limón, marshy forest [1°2'N, 76°33'W] 26
Apr[il]. 1977. J. Mallet leg. BMNH 2011-63. BMNH(E)
1720530 (deposited NHMUK).
Diagnosis
Magneuptychia pax n. sp. broadly resembles other
species in this genus due to its size, general coloration
and patterning, but it is distinguished from all other
Euptychiina species by the broad, white hindwing
marginal border, which extends from the postdiscal ocelli
to the wing margin, and otherwise dark brown wing
ground colour. The ventral surface wing pattern in
general resembles that of several species currently placed
in Magneuptychia, such as M. fugitiva Lamas, 1997, but
6 Conservación Colombiana – Número 24 – 27 de octubre de 2016
Figure 1. Magneuptychia pax n. sp. A. (above) Male holotype dorsal view (left) and ventral view (right). B. (below) Female allotype
dorsal view (left) and ventral view (right).
this species differs in having two dots, instead of one pale
dot, in the VHW ocelli. The ocelli are similar in this
respect to those of Magneuptychia gera (Hewitson,
1850), but while the latter species and its likely relatives
often have white shading in ventral marginal areas, as in
M. pax, they also have an extra ocellus in VHW cell 2ACu2, which is lacking in M. pax. Preliminary molecular
analyses (M. Espeland et al., in prep.) support the
diagnosis of this new species as being highly distinct
from other Euptychiina.
Description of the male
See Fig. 1A. Head: Eyes hairy, dark brown; palpi
approximately twice as long as head, brush-like black
and white, antennae brown. Thorax: Dark brown.
Abdomen: Dark brown with hair-like scales. Wings:
Forewing mean length 21.2 mm (n=9). DFW brown, with
slightly darker submarginal and two marginal lines.
DHW predominantly brown, except for broad white
marginal band from tornus to M2-M1, broadening in
middle where extending in from wing margin to as far as
middle of cell Cu1-M3; one darker brown submarginal
line and two dark brown marginal lines, latter closely
parallel to wing margin, former more strongly undulate;
indistinct, black submarginal ocellus in cell Cu2-Cu1,
with dark yellow ring. VFW with brown ground colour
except for a whitish grey marginal band enclosing darker
brown submarginal line and two darker brown marginal
lines, latter closely parallel to wing margin, former more
undulate; broad, dark brown discal and post-discal bands
extending across almost all of wing, except at costa;
small, black, dark-centred ocellus with a dark orange
border and small greyish centre present in R5-M1. VHW
with similar brown ground colour except for broad white
marginal band enclosing one darker brown submarginal
line and two marginal lines, as on DHW; row of five
submarginal ocelli, those in cells Cu2-Cu1, M2-M1 and
M1-Rs black with a narrow yellow ring and silver dot at
centre, first two ocelli large, last small, ocelli in cells
Cu1-M2 ovoid, brown with a narrow yellow ring and
large silver spot at centre; broad, dark brown discal and
post-discal bands extending across wing, latter kinked
outwards at vein M1. Abdomen and genitalia: (Figs.
2A-D). As illustrated. Eighth tergite largely membranous,
sclerotized portion reduced to a narrow anterior strip;
uncus relatively long, straight and even in width except
tapering at the very end; subuncus directed dorsally of
uncus and slightly shorter than it, much narrower than
uncus and curved inwards at tip; tegumen small and
mitre-shaped; valvae hirsute, especially on distal and
Conservación Colombiana – Número 24 – 27 de octubre de 2016
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Figure 2. Illustration of structures in Magneuptychia pax n. sp. A. Male abdomen. B-D. Male genitalia E. Female abdomen F-G. Female
genitalia. H. Head capsule of first instar larva.
ventral sides, terminating in double prongs; saccus short,
narrow; phallus slender, lacking cornuti, slightly more
sclerotized in the central part, a little wider proximally
and slightly flared distally at tip in ventral view; no
visible juxta; 'V'-shaped sclerotized band, isolated, on
ventral surface of tuba analis.
Description of the female
See Fig. 1B. As illustrated; similar to male holotype, not
significantly smaller, wings more rounded, ground colour
paler, ventral ocelli showing more clearly on DHW.
Wings: Forewing mean length 20.6 mm (n=4).
Abdomen and genitalia: (Figs. 2E-G). As illustrated.
Eighth tergite relatively reduced in size; intersegmental
membrane between 7th and 8th segments soft, pliable
and expanded greatly ventrally to form a series of
wrinkled ventral folds, also forming small, lightly
sclerotized 'plate' underlying ostium bursae; lamella
postvaginalis an isolated, small, square plate; lamella
antevaginalis a large, rectangular lateral plate, not
meeting ventrally and isolated from 8th tergite; ductus
bursae unsclerotized, leading to a small, rounded corpus
bursae, ductus seminalis origin near ostium bursae.
Wing pattern variation
Some specimens have only a weak ocellus in R5-M1
in the HW dorsal, while in other specimens it is lacking
altogether. Some specimens have white scales on the
base of the antennae. In the HW the black submarginal
ocellus in cell Cu2-Cu1 is variable in size and the dark
yellow ring may vary from distinct to very indistinct.
Immature stages
One dissected female contained a first instar larva. The
head capsule is rounded in anterior view, with two short,
rounded dorsal head 'horns' (Fig. 2H).
Paratypes
Colombia: (n=6♂ 1♀). Amazonas: 1♂, Puerto Nariño,
[3°45'S, 70°22'W], 27 Sep 1994++, J. F. Le Crom leg.
(JFLE); Caquetá: 3♂, Serranía de Chiribiquete, Río
Cuñaré, Verde Biche Bosque, 0°32'4''N, 72°37'57''W, 19
Feb 2001, G. Fagua et al. leg. (MPUJ); 1♂, Solano,
Vereda El Quince, 0°48'11''N, 75°11'58''W, 204m, Jama.
4 Sep 2007, 1505h. C. Sañudo leg. Catálogo 15058 93CP 36- (MEFLG); 1♂, Solano, Vereda El Quince,
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Parcela la Sombra, 0°48'11''N, 75°11'58''W, Jama. 204m,
4 Sep 2007, C. Sañudo & F. Muñoz leg. MUA-INN 11171 (MUA); Putumayo: 1♀, Río Caquetá, nr. José María,
Los Naranjos, [1°34'13''N, 75°43'W], 200m, 3 Aug 1977,
J. Mallet, D. Jackson and P. Garcia-C. leg. FLMNHMGCL-195896 (FLMNH).
Ecuador: (n=5♂ 5♀). Orellana: 1♂, 28 km SE Coca,
track NNE of Pindo 13 oil well, [0°39'40''S, 76°49'W],
260m, 19 Aug 2010, D. A. Trembath, A. Neild leg. No.
22961 (DATR); 1♂, km 21 Coca-Loreto rd., 0°29'31''S,
77°8'19''W, 300m, 8 Mar 1995, K. Willmott & J. Hall
leg. (FLMNH); 1♀, Lagunas de Cuyabeno, La Ormiga,
0°1'S, 76°11'W, 250m, 18 Sep 1996, K. Willmott leg.
(KWJH); Pastaza: 1♂, Río Curaray, Oriente, Lorocachi,
1°37'15''S, 75°59'30''W, 250m, 20 Feb 1996, C. Carpio
leg. (37. Images 271, 272) (FRPI); 1♀, Río Pastaza,
Kapawi village, 2°32'16''S, 76°50'10''W, 260m, 23 Jul
2009, K. R. Willmott & J. P. W. Hall leg. FLMNHMGCL-145665 (FLMNH); 1♀, Yutsuntsa, along trail to
Makusar, 2°21'5.02''S, 76°27'18.3''W, 13 Jul 2014, S.
Nakahara & S. Padrón leg. (SHNA); Sucumbíos: 1♀,
Cuyabeno Lodge, across lagoon, [0°0'18''S, 76°10'23''W],
224m, 7 Dec 2010, J. D. Turner leg. FLMNH-MGCL150974 (FLMNH); No data: 1♂, 'Ecuador', Ex. Grose
Smith 1910, Joicey Bequest Brit. Mus. 1934-20,
BMNH(E) 1720540, B.M. SLIDE 32034. 1♂, same data
BMNH(E) 1720533; 1♀, same data BMNH(E) 1720534
(NHMUK).
Peru: (n=10♂ 20♀). Loreto: 1♂, Arcadia 0°59.37’S,
75°18.55’W, 150m, 5 Nov 1993, G. Lamas leg.; 1♀,
same data but 4 Nov 1993, R. K. Robbins leg.; 3♂, same
data but 5 Nov 1993; 2♀, same data but 4 Nov 1993, G.
Lamas leg.; 1♀, same data but 8 Nov 1993; 1♀, same
data but 7 Nov 1993, R. K. Robbins leg.; 1♀, Castaña,
0°48'S, 75°14'W, 150m, 24 Oct 1993, R. K. Robbins leg.;
1♀, same data but 30 Oct 1993; 1♂, Rio Pucacuro,
Coconilla, 2°42'S, 75°6'W, 160m, 17 Jul 2003, J. J.
Ramírez leg.; 1♂, Explornapo-ACEER, Río Sucusari,
[3°15'28''S, 72°55'3''W], 140m, 22 Sep 1995, G. Lamas
leg.; 1♀, same data but 16 Sept 1995; 1♂, same data but
13 Sept. 1995, J. Grados leg.; 1♂, 1♀, same data but R.
K. Robbins leg.; 1♀, same data but 5 Sept 1995; 1♀,
same data but 19 Sept 1995, A. Caldas leg.; 2♀, Puerto
Almendra, 3°50'S, 73°23'W, 120 m, 3 Sep 1995, D. J.
Harvey leg.; 2♀, Río Aguas Negras, 0°31'24''S,
75°15'24''W, 150m, 3 Mar 1994, G. Lamas leg.; 1♀,
same data but 5 Mar 1994; 1♀, same data but 8 Mar
1994; 1♂, Tierra Hermosa, 3°34'S, 73°13'W, 140m, 8
Oct 2013, J.J. Ramírez leg.; 1♀, Picuroyacu, 3°37'S,
73°16'W, 115m, 19 Oct 2015, G. Lamas leg. (all in
MUSM); 1♀, Pebas [3°19'S, 71°51'W, 120m],
H[a]h[ne]l/Euptych. sp?. nobis ignota. Genitalia vial n. M
9100 by Lee D. Miller (MFNB); 1♀, Pebas [3°19'S,
71°51'W, 120m], Amazons, M. de Mathan leg.,
Rothschild Bequest B.M. 1939-1. BMNH(E) 1719068;
1♀, Yahuas Terr. [3°21'S, 71°59'W], Joicey Bequest Brit.
Mus. 1934-20. BMNH(E) 1720535 (NHMUK). No data:
1♂, ‘Peru’. BMNH(E) 1720532 (NHMUK).
No locality: (1♂). Ex. Grose Smith 1910. Joicey Bequest
Brit. Mus. 1934-20. BMNH(E) 1720534 (NHMUK).
Distribution
Magneuptychia pax n. sp. has been found to date only in
a limited area of western Amazonia, where it is
confirmed from specimens collected in Colombia (north
to Caquetá), Ecuador and Peru (south to Loreto). All
known localities are situated within 4 degrees of the
Equator, suggesting a specialisation to warmer habitats
within western Amazonia. This species is known to range
from 115 to 300 metres above sea level (Fig. 3).
Etymology
The name pax is the nominative singular of a third
declension Latin noun, meaning ‘peace’. The name is
non-variable. Magneuptychia pax n. sp. is dedicated to
the peace process in Colombia and to every person
affected there by a conflict that has lasted more than five
decades, including in the remote forests that this butterfly
inhabits. This dedication is made in the hope that a
lasting peace agreement can be reached and to focus
attention on the need for conservation of Amazon forests
and improvement in conditions for research and scientific
discovery in that region.
Ecology
During fieldwork in Serranía de Chiribiquete, Colombia,
Magneuptychia pax n. sp. was a common species in the
understory of Amazonian forests that were transitional
between flooded and terra firme forest. This species was
found in natural forest edges and clearings, such as near
riverbanks or in the transitional areas from forest to
sandy or rocky areas. It seemed to particularly prefer
white sand forests and forest edges. The species was
collected at this locality only during expeditions in
February, although it has been recorded in other months
of the year elsewhere. However, it was not present in a
recent study of stunted tepui-top habitats in the
Chiribiquete region (Huertas et al. 2015). The ecological
preferences of the new species in Colombia are notable
because many other Magneuptychia are more often found
in the interior of dense lowland forest, with closed
canopy and a sparse understorey. The white markings on
the new species may assist matching to pale substrate in
sand forest edge habitats. In Ecuador, the species is
known only from flat, eastern lowland forest, most often
near black water rivers, where individuals were
encountered flying singly in the understorey of primary
forest from 1100hrs to 1330hrs.
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Figure 3. Map of northern South America showing localities where Magneuptychia pax n. sp. has been recorded. Photo: Orellana, Parque
Nacional Yasuní, Río Yasuní, vicinity of 'Baradero' trail, Ecuador by KW.
Discussion
A priority for a much-needed broader taxonomic revision
of Magneuptychia will be the delimitation of new genera,
which will in turn permit more detailed study of species
relationships.
Work is underway in developing
morphological and molecular datasets (Systematics of
Euptychiina project, unpublished data) to produce a more
robust higher-level classification. A separate challenge
arises from the number of undescribed species in
Magneuptychia. The existence of distinctive (and often
phylogenetically significant) undescribed species such as
M. pax remains a barrier to a proper understanding of the
group. Thus, although taxonomic revisions are our
ultimate goal for a number of Euptychiina genera,
including
Magneuptychia
(see
http://www.flmnh.ufl.edu/museum-voices/euptychiina/),
we believe it is helpful to describe the more distinctive
species that it will be essential to include in higher level
phylogenies, even if their generic placement is
provisional. Other authors have followed this approach
in recent years with the descriptions of several
Magneuptychia species (e.g., Brévignon 2005, Brévignon
& Benmesbah 2012, Zacca et al. 2014).
The relationships of this new species remain unclear.
Preliminary sequence data from several nuclear and
mitochondrial genes (Espeland et al., unpublished data)
suggest that M. pax may not have any very close relatives
and is not closely related to type species M. libye
(Linnaeus, 1767) and so could ultimately require its own
genus. However, at present, description of a genus would
be premature given that work is underway to better
resolve the phylogeny of Euptychiina. Such work will
help inform a more appropriate generic classification for
this and many other species that are currently assigned to
Magneuptychia due to historical reasons or by default.
The restricted region from which Magneuptychia pax is
known remains rather inaccessible and poorly explored,
not only due to innate complexity of the Amazon
landscape but also due to political instability in these
habitats particularly in Colombia. Western Amazonia
supports many other undescribed euptychiine taxa (Fagua
& Sánchez unpublished; von Hildebrand unpublished;
Huertas 2014), and more detailed exploration of remote
Amazonian localities, and white sand forest regions in
particular, is warranted.
Acknowledgements
We thank fellow curators and collectors for facilitating
visits, loans and for sending data and photos from
specimens; BH sincerely acknowledges assistance
received from Dr Wolfram Mey (MFNB), Fernando
Valencia and Alexis Acosta (MUA), Jean François
LeCrom (JFLE), Francisco Piñas Rubio (FRPI), Andrew
Neild and David Trembath. Sincere thanks to NHM staff
Dr Klaus Sattler for helping with German translations,
the butterfly collections’ team of volunteers helping with
curation of Euptychiina, library staff who provided help
with references and the Photo Unit team Harry Taylor
Conservación Colombiana – Número 24 – 27 de octubre de 2016
10
and Jonathan Jackson who provided photos of the type
specimens. KW thanks Jon D. Turner. GF thanks
Fernando Gast Harders (Chiribiquete Project), and
Patricio von Hildebrand (Fundación Puerto Rastrojo) for
their support. The specimens from Serranía de
Chiribiquete (Colombia) were collected during the
initiative “Inventory of the National Natural Park of
Chiribiquete” made by the GEMA group (Group of
Exploration and Environmental Monitoring) with
researchers from the Alexander von Humboldt Institute
(IAvH) and Fundación Puerto Rastrojo, Colombia.
Research on Ecuadorian butterflies and euptychiines in
general was supported in part by the Florida Museum of
Natural History and FLMNH Museum Associates, the
National Geographic Society (Research and Exploration
Grant #5751-96) and the National Science Foundation
(grants #0103746, #1256742). We also thank S.
Villamarín, the MECN and Ecuadorian Ministerio del
Ambiente for arranging the necessary permits for
research in Ecuador. KW also thanks Jason Hall for many
years of collaboration in research on Ecuador's
butterflies, and Julia and Jamie Robinson Willmott for
their fine company in the field. BH thanks Thomas
Donegan for his continuous support and fruitful
discussions to improve this manuscript. Finally, thanks
to Dr David Lees (NHM) and two other anonymous
reviewers for their comments on this paper.
References
Brévignon, C. 2005. Description de nouveaux Satyrinae
provenant de Guyane française (Lepidoptera,
Nymphalidae). Lambillionea 105(3)(1): 393-404.
Brévignon, C. & Benmesbah, M. 2012. Complément à
l'inventaire des Satyrinae de Guyane (Lepidoptera:
Nymphalidae), pp. 36-52, 4 pls., 1 tab. In: Lacomme,
D. & L. Manil (eds.), Lépidoptères de Guyane. Tome
7.
Nymphalidae.
Paris,
Association
des
Lépidoptéristes de France.
Comstock, J.H. & Needham, J.G. 1918. The wings of
insects. American Naturalist 32: 231-257.
Costa M., Viloria, A.L., Attal, S., Neild A.F.E., Fratello
S.A. & Nakahara S. 2016. Lepidoptera del Pantepui.
Parte III. Nuevos Nymphalidae Cyrestinae y
Satyrinae. Bulletin de la Société entomologique de
France 121(2), 2016: 179-206.
D'Abrera, B. 1988. Butterflies of the Neotropical Region,
Part V. Nymphalidae (conc.) and Satyridae. Victoria,
Australia: Hill House. Pp. 679-877.
Forster, W. 1964. Beitrage zur Kenntnis der
Insecktenfauna Boliviens XIX. Lepidoptera III,
Satyridae. Veröffentlichungen der zoologischen
Staatssammlung München 8: 51-188, pls. 27-35.
Hewitson, W. C. 1850. Descriptions of some new species
of butterflies. Annals and Magazine of natural
History (2)6(36):434-440, pls. 9-10 (1 December).
Huertas, B. 2014. Evaluating the conservation status of
Neotropical butterflies and the impact of systematics
on threat assessments. PhD Thesis, University
College London, Division of Biosciences.
Huertas, B., Moorwood, A., Forero, F., Kirby, R.,
Rodriguez, A. & Doyer, T. 2015. What’s the point?
New biodiversity records from a rapid assessment of a
tepuy in PNN Serranía de Chiribiquete during the
filming of the National Geographic documentary
‘Wild Colombia’ and the feature film ‘Colombia
Magia Salvaje’. Conservación Colombiana 23:82-90.
Klots, A.B. 1970. Lepidoptera, pp. 97-110 In: Tuxen,
S.L. ed. Taxonomist’s Glossary of Genitalia of
Insects. Copenhagen, Munksgaard. 359 p.
Lamas, G. 2004. Nymphalidae. Satyrinae. Tribe Satyrini.
Subtribe Euptychiina, pp. 217-223. In: Lamas, G.
(ed.) Checklist: Part 4A. Hesperioidea Papilionoidea. In: Heppner, J. B. (ed.) Atlas of
Neotropical Lepidoptera. Volume 5A. Gainesville,
Association for Tropical Lepidoptera; Scientific
Publishers.
Lamas, G., Robbins, R.K., Harvey, D.J. [1997].
Mariposas del alto Río Napo, Loreto, Perú
(Lepidoptera: Papilionoidea y Hesperioidea). Revista
peruana de Entomología 39: 63-74.
Marín, M.A., Peña, C., Freitas, A.V.L., Wahlberg, N. &
Uribe, S.I. 2011. From the phylogeny of the Satyrinae
butterflies to the systematics of Euptychiina
(Lepidoptera: Nymphalidae): History, progress and
prospects. Neotropical Entomology 40(1): 1-13.
Michael, O. 1926. Erinnerungen aus Süd-Amerika. Dr.
Paul Hahnels letzte Reise nach dem Amazonas!
Frankfurt am Main, Verlag des Internationalen
Entomologischen Vereins. 96 + [1] pp., 4 pls.
Murray, D.L. & Prowell, D.P. 2005. Molecular
phylogenetics and evolutionary history of the
neotropical
satyrine
subtribe
Euptychiina
(Nymphalidae: Satyrinae). Molecular Phylogenetics
and Evolution 34(1): 67-80.
Piñas, F. 2004. Mariposas del Ecuador. Vol. 11b.
Familia: Nymphalidae. Subfamilia: Satyrinae. Quito,
Compañía de Jesús. v + 90 pp. + CD.
Peña, C. & Lamas, G. 2005. Revision of the butterfly
genus Forsterinaria Gray, 1973 (Lepidoptera:
Nymphalidae, Satyrinae). Revista peruana de
Biología 12(1): 5-48.
Peña, C., Nylin, S., Freitas, A.V.L., Wahlberg, N. 2010.
Biogeographic history of the butterfly subtribe
Euptychiina (Lepidoptera, Nymphalidae, Satyrinae).
Zoologica Scripta 39(3): 243-258.
Peña,
C.,
Wahlberg,
N.,
Weingartner,
E.,
Kodandaramaiah, U., Nylin, S., Freitas, A. V.L &
Brower, A.V.Z. 2006. Higher level phylogeny of
Satyrinae butterflies (Lepidoptera: Nymphalidae)
based on DNA sequence data. Molecular
Phylogenetics and Evolution 40(1): 29-49.
Staudinger, O. 1890. Lebesnskizze des Dr. Paul Hahnel.
Deutsche entomologische Zeitschrift “Iris” 3(1): 128132.
Zacca, T., Siewert, R.R., Mielke, O.H.H. & Casagrande,
M.M. 2014. A new species of Magneuptychia Forster,
1964 (Lepidoptera: Nymphalidae: Satyrinae) from
Brazilian savannah. Zootaxa 3795(1): 71-78.
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