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195
ISSN 1863-7221 (print)
|
eISSN 1864-8312 (online)
© Senckenberg Gesellschaft für Naturforschung, 2017.
75
(2): 195 – 243
8. 9. 2017
Taxonomy, phylogeny and distribution of the genus
Steromapedaliodes sensu novo in the Cordillera de
Mérida, Venezuela (Lepidoptera: Nymphalidae:
Satyrinae: Satyrini)
T W. P *, 1, 5, J L-B 5, A Z
1,
P B 2, M. C G 3 & J M *, 4
1
Entomology Department, Institute of Zoology and Biomedical Research, Jagiellonian University, Gronostajowa 9, 30 – 387 Kraków, Poland;
Tomasz W. Pyrcz * [tomasz.pyrcz@uj.edu.pl] —
2
7 Lotissement l’Horizon, 13610 Le Puy Sainte Réparade, France —
3
Centro de Ecología, In-
stituto Venezolano de Investigaciones Científicas, Apartado 20632, Caracas 1020 – A, Venezuela —
4
Laboratoire d’Ecologie Alpine, UMR 5553
CNRS – Université Grenoble Alpes, BP 53 2233 Rue de la Piscine, 38041 Grenoble, France; Jesús Mavárez * [jesus.mavarez@univ-grenoble-
alpes.fr] —
5
Zoological Division, Nature Education Centre, Jagiellonian University, Gronostajowa 5, 30 – 387 Kraków, Poland — * Correspond-
ing authors
Accepted 17.ii.2017.
Published online at www.senckenberg.de/arthropod-systematics on 30.viii.2017.
Editors in charge: Karl Kjer & Klaus-Dieter Klass
Abstract
We revise the taxonomy and phylogeny of the genera Steromapedaliodes Forster, 1964 and Redonda Adams & Bernard, 1981, both
subparamo and paramo endemics of the Cordillera de Mérida (Venezuela). Redonda is hereby synonymized with Steromapedaliodes, a
decision supported by a comparative study evidencing several synapomorphies of both genera in external characters and genitalia, and the
lack of salient synapomorphies specic to each genus in morphological, biogeographical and ecological traits. Also, Steromapedaliodes
mavarezi sp.n. presents several intermediate characters in relation to the species traditionally assigned to Redonda and Steromapedali-
odes. Furthermore, a molecular phylogenetic analysis with the use of one nuclear and three mitochondrial markers shows no support for
reciprocal monophyly between Redonda and Steromapedaliodes but provides strong support for the monophyly of Steromapedaliodes
sensu novo, with low levels of genetic divergence among species. Two new species of Steromapedaliodes are described, S. kahlua sp.n.
and S. mavarezi sp.n. We describe ve new subspecies of S. albonotata (Godman, 1905), two of S. albarregas (Adams & Bernard, 1981),
and the females of S. empetrus empetrus (Thieme, 1905) comb.n. and S. lathraia Viloria & Camacho, 2015 comb.n. Several new status
and synonymies are also established. Accordingly, Steromapedaliodes contains nine species, including four that are highly polytypic.
Adult Seromapedaliodes share several striking morphological and genital similarities with another north Andean paramo genus, Dangond
Adams & Bernard, 1979 from the Sierra de Perijá. However, our phylogenetic analysis shows that the two are not sister taxa and that
they originated in separate clades of the speciose Pedaliodes complex. This shows that convergence in butteries affects not only external
morphology traits directly correlated with habitat selection but also genitalia. Steromapedaliodes has undergone an adaptive radiation in
geographically isolated paramos resulting in unusual morphological and behavioural characters such as brachyptery and ightlessness of
the females of S. bordoni Viloria & Pyrcz, 2003 comb.n. and S. empetrus (Thieme, 1905) comb.n., a unique adaptation among the world’s
butteries.
Key words
Adaptive radiation, Andes, convergence, endemism, species description, generic revision, paramo, Steromapedaliodes, Redonda.
P et al.: Taxonomy and phylogeny of Steromapedaliodes
196
1. Introduction
Montane forests cover most of the areas in the Northern
Andes, but the altitudinal belt above 3000 m corresponds
to a tundra-like or grasslands ecosystem known as pára-
mo (hereafter paramo, Salgado-labouriau et al. 1988).
This ecosystem is therefore found between the continu-
ous tree line and the snow line and is characterised by
a very high proportion of giant rosette plants, alongside
shrubs and grasses (luteyn 1999). Paramo habitats are
very young, because although the Northern Andes started
to rise in the late Oligocene, they were ridges and small
mountains smaller than 700 m in elevation by mid-Mi-
ocene, attained 30 – 40% of their modern elevation by
5.3 Mya early-Pliocene) and reached present-day heights
through a rapid nal uplift by 2.7 Mya (late-Pliocene)
(Van der Hammen et al. 1973; Van der Hammen & Cleef
1986; Van der Hammen 1989; Kroonenberg et al. 1990;
andrieSSen et al. 1992; HelmenS & Van der Hammen
1994; gregory-WodziCKi 2000; HoogHiemStra et al.
2006). Besides, paramos are also very fragmented; struc-
tured as many small “islands” of cold grassland habitats
surrounded by a “sea” of warm forest. This geological
and ecological set-up has had a profound impact on the
biodiversity of the paramo ecosystem, making it the one
with the highest diversity and the highest proportion of
plant endemics among the world’s high-altitude habitats.
For instance, paramos are home to no less than 20 en-
demic plant genera and an unknown but large number of
endemic monophyletic groups within other taxa (luteyn
1999; Sklenář et al. 2011).
In Venezuela, most paramos are found in the high-
elevation areas of the Cordillera de Mérida, which is a
fairly isolated branch of the Northern Andes (Fig. 1A).
Here too, evolution has favoured the rise and further di-
versication of several endemic or quasi-endemic taxa
such as a monophyletic clade of ~ 60 spp. in the sub-
tribe Espeletiinae (Asteraceae) (Pouchon et al. unpubl.
data), and seemingly monophyletic groups within Pen-
tacalia (Asteraceae) (lapp 2014), Hinterhubera (Aster-
aceae) (Karaman 2006), Valeriana (Valerianaceae)
(Xena de enreCH 1993) and many others. In animals, a
few groups include endemic genera in the Cordillera de
Mérida, such as the mouse genus Aepeomys and its two
described species (VoSS et al. 2002) and the grasshopper
genus Meridacris, also with two species (robertS 1937).
However, the largest number of endemic animal genera
in the Cordillera de Mérida occurs in the Lepidoptera,
particularly among the better-studied butteries (Papilio-
noidea; neild 1996, 2008; pyrCz et al. 2010), and most
notably among the montane subtribe Pronophilina (Nym-
phalidae, Satyrinae), for which at least four genera en-
demic to this range have been described: Cheimas Wey-
mer, 1912 (1 sp.), Diaphanos Adams & Bernard, 1981 (3
spp.), Ste romapedaliodes (sensu Viloria & Pyrcz, 2001)
(2 spp., but see below) and Redonda Adams & Bernard,
1981 (10 spp., but see below), all of them restricted to
the forest-paramo ecotone and paramo grassland. This is
outstanding in comparison to other isolated mountainous
massifs of northern South America or Central America,
which harbour one endemic buttery genus at most: Par-
amo Adams & Bernard, 1977 (1 sp.) in the Sierra Nevada
de Santa Marta (other two genera originally described as
endemic by adamS & bernard (1977) were subsequently
synonymized or found to be more widely distributed),
Dangond (1 sp.) in the Sierra de Perijá and Drucina But-
ler, 1872 (2 spp.) in Panama /Costa Rica (Sierra de Tala-
manca and Meseta Central) and southern Mexico.
It must however be acknowledged that further pro-
gresses in the understanding of the biogeography and
evolution of endemic taxa from the Cordillera de Mérida,
and the Pronophilina in particular, have been hampered
by a lack of studies on their phylogenetic relationships
and taxonomy. Only one molecular phylogenetic analy-
sis including samples of the genera discussed herein has
been conducted so far, and it was designed more to depict
the high level phylogeny of the subfamily Satyrinae rath-
er than to establish relationships among genera (peña et
al. 2011). In consequence, the phylogenetic relationships
of the endemic Pronophilina from the Cordillera the Mé-
rida remain largely unstudied, which also contributes in
some degree to the taxonomic ambiguity associated with
some of these taxa. For instance, the highly distinctive
Diaphanos, with three described species, appears distant-
ly related to Lymanopoda Westwood, [1851] (peña et al.
2011), but its position within the Pronophilina remains a
matter of controversy (pyrCz 2010a). On the other hand,
the phylogenetic afnities of the monobasic genus Chei-
mas are even more dubious, as the analysis of peña et al.
(2011) indicates that it is only loosely related to Corades
Hewitson, 1849, within a highly polytomic clade that
also includes the genera Pronophila Doubleday, [1849]
and Lasiophila C. Felder & R. Felder, 1859 among oth-
ers. However, as for Diaphanos, the generic status of
Cheimas remains unquestioned given its characteristic
external morphology (adamS & bernard 1981).
The phylogenetic afnities of the two remaining en-
demic genera from the Cordillera de Mérida, Steroma-
pedaliodes (sensu Viloria & Pyrcz) and Redonda, are
particularly interesting because these are the two most
speciose taxa and Redonda includes the only known ex-
amples of brachyptery and loss of the ability to y among
the world’s butteries (Rhopalocera) (Viloria et al. 2003;
pyrCz 2010a,b). However, although the positions of both
taxa within the speciose (> 300 spp.) Pedaliodes Butler,
1867 complex, to which they belong, have been subject
to some discussion in the past (adamS & bernard 1981),
their phylogenetic relationships have not been rigorously
explored. The validity of the two genera as separate enti-
ties has not been challenged despite that they appear to be
closely related sister taxa (peña et al. 2011). This is sur-
prising when we consider the lack of good synapomor-
phic characters specic to each genus together with the
existence of good synapomorphic characters linking both
of them into a single group. Thus, Steromapedaliodes
(sensu Viloria & Pyrcz) and Redonda share highly simi-
lar ecological requirements in terms of habitat used, male
197
ARTHROPOD SYSTEMATICS & PHYLOGENY — 75
(2) 2017
genitalia and most notably colour patterns, particularly
on the hindwing venter, in particular the conguration of
median, postdiscal and submarginal lines and eyespots,
parts of the so-called nymphalid groundplan (nijHout
1991), considered as a valuable character for phyloge-
netic relationship among the Pronophilina (pyrCz et
al. 2008). This combination of ecological and morpho-
logical features is exclusive to Steromapedaliodes and
Redonda among the speciose Pedaliodes complex. Re-
donda does show some characteristics that at rst glance
make it look different from Steromapedaliodes (sensu
Viloria & Pyrcz), notably the major reduction or absence
of the subuncus in the male genitalia and a tendency for
larger males than females. Yet the systematic value of
such traits has never been evaluated properly and there
are good reasons to raise doubts about them. For ex-
ample, reduced or absent subunci are also found in the
genus Dangond and in Panyapedaliodes stellata Pyrcz,
2004 (pyrCz 2004). Both taxa are most probably phylo-
genetically distant from Redonda, but they share with it
a life in the paramo, suggesting that these characters of
male genitalia are relatively plastic and able to evolve
repeatedly in response to the extreme conditions of high
elevation habitats. In regards to the relative size of males
Fig. 1. Maps. A: Schematic representation of the land area above 1500 m and above 2500 m (dark grey) in the Northern Andes of Colom-
bia and Venezuela. B: Sampling sites of Steromapedaliodes chiquinquirana (3 ssp.), S. empetrus (4 ssp.), S. leukasmena, S. lathraia and
S. bordoni. C: Sampling sites of Steromapedaliodes albonotata (8 ssp.). D: Sampling sites of Steromapedaliodes mavarezi, S. kahlua and
S. albarregas (3 ssp.).
P et al.: Taxonomy and phylogeny of Steromapedaliodes
198
and females, there are considerable differences among
species within both Steromapedaliodes (sensu Viloria &
Pyrcz) and Redonda, as it seems also to be the case in
other genera of Pronophilina such as Dangond, Paramo
and Punapedaliodes Forster, 1964 (adamS & bernard
1977, 1979; pyrCz 2004, 2007a,b; Viloria 2007).
This study is mainly focused on investigating the phy-
logenetic relationship between Steromapedaliodes (sensu
Viloria & Pyrcz) and Redonda. Two main scenarios are
tested: i) Steromapedaliodes (sensu Viloria & Pyrcz) and
Redonda form two well-dened and reciprocally mono-
phyletic groups, in which case their lack of respectively
synapomorphic characters is somehow a reect of a slow-
er rate of morphological evolution; ii) Stero mapedaliodes
(sensu Viloria & Pyrcz) and Redonda do not form recip-
rocally monophyletic groups, in agreement with the lack
of morphological synapomorphies and in which case the
two genera should be unied into a single one, Steroma-
pedaliodes sensu novo. In addition, we explore the phy-
logenetic relationships of both taxa with other Neotropi-
cal Satyrinae, in particular other groups found in oreal
habitats such as some lineages in the Pedaliodes complex
from the Cordillera de Mérida and Dangond from the Si-
erra de Perijá.
The assessment of these scenarios and relationships
will certainly provide another example of the utility of
molecular phylogenies in disentangling of the complex
patterns of buttery systematics associated with pre-cla-
distic nomenclature (WaHlberg et al. 2005; peña et al.
2006). For Steromapedaliodes (sensu Viloria & Pyrcz)
and Redonda, the abandonment of traditional taxonomic
usages could be a necessary consequence of an advanced
knowledge about their mutual relationship and evolu-
tionary history. In any case, the result will be an updated
lexicon of generic, specic and sub-specic names that
will facilitate a more effective communication among re-
searchers and enthusiasts of Neotropical butteries.
2. Materials and methods
2.1. Taxa studied
The genus Steromapedaliodes was raised by forSter
(1964) for a single species found in the Venezuelan An-
des, Pedaliodes albonotata Godman, 1905, mostly based
on male genitalia. adamS & bernard (1981) rejected
this action, placed albonotata in another genus, Altope-
daliodes Forster, 1964 and described another apparently
closely related species, Altopedaliodes albarregas Ad-
ams & Bernard, 1981. Viloria & pyrCz (2001) demon-
strated that the abdomen studied by Forster was glued to
the thorax and that it belonged to a species of the genus
Manerebia Staudinger, 1987. They sustained however
the validity of Steromapedaliodes and described two new
taxa at the specic level, S. schuberti Viloria & Pyrcz,
2001 and S. sanchezi Viloria & Pyrcz, 2001.
The genus Redonda was erected by adamS & ber-
nard (1981) for a single species, originally described as
Pedaliodes empetrus by tHieme (1905), based on male
genitalia, wing pattern, and some ecological features.
They additionally described one new subspecies, R. em-
petrus bolivari Adams & Bernard, 1981. A second spe-
cies, R. bordoni Viloria & Pyrcz, 2003 was also described
(Viloria et al. 2003). Recent eld studies show that the
genus is much more diverse, and brought to light the
existence of complex biogeographic patterns within the
Cordillera de Mérida (pyrCz 2010a; Viloria et al. 2015).
The genus Dangond was described as monobasic for
the monotypic species Dangond dangondi Adams & Ber-
nard, 1979 (adamS & bernard 1979). Its taxonomy and
phylogeny have not been studied since.
2.2. Study area
Butteries in the genera Steromapedaliodes (sensu
Viloria & Pyrcz) and Redonda are distributed along the
Cordillera de Mérida in Venezuela, which is a north-
eastern extension of the Andes (Fig. 1A). It runs south-
west – northeast between the Colombian Eastern Andean
branch (Cordillera Oriental) and the Venezuelan coast-
al range, but it is separated from them by the Táchira
depression and the Lara depression, respectively. The
Cordillera de Mérida is indeed composed by two ranges
of peaks; the Sierra de la Culata to the north and the
Sierra Nevada de Mérida to the south, which includes
the highest peak in Venezuela, Pico Bolívar (4981 m).
Tributaries of the Orinoco River drain its south-eastern
slopes, while the streams that drain the north-western
slopes empty into Lake Maracaibo. Dangond dangondi
is found in the Sierra de Perijá, which is a mountain
range that represents a northward extension of the Co-
lombian Eastern Andean branch (Cordillera Oriental),
between Colombia and Venezuela, and ending in the
Guajira Desert. It is separated from the Cordillera Orien-
tal by the Catatumbo river in the Maracaibo basin. The
highest point is Cerro Pintado (3660 m), followed by
Cerro de Las Tetas (3630 m) and Cerro Irapa (3540 m).
The mountains of the Cordillera de Mérida and the Si-
erra de Perijá are separated from the Colombian Eastern
Cordillera by a major orographical and biogeographical
barrier, the so-called “Táchira Depression” (pyrCz &
Viloria 2007).
2.3. Field work
Field work was carried out by TP, PB and JM throughout
the Cordillera de Mérida from 1991 – 2015, and by TP in
the Sierra de Perijá in 2012. The list of sampled localities
is given in Table 1. Collecting was conducted on each
locality with standard entomological nets. Van-Someren
Rydon baited traps were used only to a limited degree as
they have proved to be inefcient. They only sporadical-
ly attract adult butteries in paramo grassland, contrary
199
ARTHROPOD SYSTEMATICS & PHYLOGENY — 75
(2) 2017
to cloud forests (pyrCz et al. 2009). Field observations
of adult behaviour were carried out in all the sampled lo-
calities with an emphasis on diurnal activity pattern, mate
locating strategies and feeding behaviour. Specimens in-
tended for molecular analyses were preserved in absolute
ethanol or kept dry in glassine envelopes.
2.4. Morphological data
The following abbreviations were used in the text: FW,
forewing; HW, hindwing; D, dorsum; V, venter. Further
abbreviations are given in the legends of Figs. 4 (wing
venation), 12 (male genitalia), and 16 (female genitalia).
The morphological characters used in analysis include:
wing venation and colour pattern, microstructure of an-
tennae, eyes, labial palpi and tegulae as well as male
and female genitalia. Wing slides were made by remov-
ing scales in warm 10% KOH solution. Genital prepa-
rations were done using standard procedures, by soak-
ing in warm 10% KOH solution for 10 min. Scales and
internal organs were cleaned out in distilled water. Fe-
male genital organs were stained with chlorazole black.
Genital organs were examined and measured, alongside
other morphological microstructures, under an Olympus
SZX9 stereomicroscope equipped with a Nikon Digital
sight Ds – Fi1 camera. Male and female genital prepara-
tions are preserved in glycerol vials and pinned under the
specimens. Genital morphology follows mostly KlotS
(1970) and razoWSKi (1996). We chose to use the term
“subuncus” instead of “gnathos” mostly, because subun-
cus (plural: subunci) was used extensively by adamS &
bernard (1977, 1979, 1981) in their revisional papers to
designate this structure, and by all subsequent authors
dealing with Pronophilina taxonomy. Several taxono-
mists dealing with neotropical Satyrinae, especially with
Euptychiina, prefer the term gnathos (freitaS et al. 2013;
neild et al. 2014). There is some controversy whether the
two structures are homologous (KlotS 1970), although
this incertainty does not seem apply to Pronophilina and
Euptychiina.
Adults were photographed with an Olympus E – 500
digital camera equipped with 50 mm macro lens. Colour
plates were composed using Adobe PhotoShop version 9,
and their quality was enhanced by CombineZ software.
A total of over 400 adult specimens of the genera Ste-
romapedaliodes (sensu Viloria & Pyrcz), Redonda and
Dangond dangondi were examined (listed under each
taxon). Type and other specimens deposited in the follow-
ing institutional and private collections were examined
(acronyms are cited in the text): AME: Allyn Museum of
Entomology, Sarasota, USA; BMNH: The Natural Histo-
ry Museum (formerly British Museum of Natural Histo-
ry), London, United Kingdom; JFLC: collection of Jean
Francois Le Crom, Bogota, Colombia; MALUZ: Museo
de Artrópodos, Facultad de Agronomía, La Universidad
del Zulia, Maracaibo, Venezuela; MHN-UN: Museo de
Historia Natural, Facultad de Ciencias, Universidad Na-
cional, Bogotá, Colombia; MIZA: Museo del Instituto de
Zoología Agrícola, Facultad de Agronomía, Universidad
Central de Venezuela, Maracay, Venezuela; AFN: col-
lection of Andrew F. Neild, London, UK; CEP-MZUJ:
Centrum Edukacji Przyrodniczej (formerly Muzeum
Zoologiczne) Uniwersytetu Jagiellońskiego; ZSM: Zoo-
logische Staatssammlung München, Germany; PBF: col-
lection of Pierre Boyer, Le Puy Sainte Réparade, France;
MCC: collection of Mauro Costa, Caracas, Venezuela;
TWP: collection of Tomasz W. Pyrcz, Warsaw, Poland
(incorporated into MZUJ).
2.5. Molecular data and analyses
DNA was obtained from dry or ethanol preserved speci-
mens listed in Tables 1 and 2 using QIAGEN’s DNeasy
kit and following the manufacturer’s protocol. In general
DNA was extracted from two legs of each specimen but
in the case of poor yields, the whole body was used for
the extraction. Both mitochondrial and nuclear regions
were sequenced in this study using PCR primers de-
scribed in Electronic Supplement Table S1.
Mitochondrial. Sequences from three mitochondri-
al regions were amplied: Cytochrome Oxidase I gene
(COI), tRNA – Leu and Cytochrome Oxidase II gene
(COII). PCR reactions were performed in a 25 µl reaction
volume with 5 – 100 ng of DNA template, 0.5 μM of each
primer, 0.2 mM of each dNTP, 1 U of GoTaq DNA poly-
merase (Promega), 1X GoTaq DNA polymerase buffer
and 1.5 mM MgCl2. The PCR cycling prole comprised
an initial denaturation at 95°C for 5 min followed by 40
cycles of 95°C for 30 s, 50°C for 30 s, 72°C for 1 min,
and a nal extension period of 72°C for 10 min.
Nuclear. Sequences of the Glyceraldehyde-3-Phos-
phate Dehydrogenase gene (GAPDH) were amplied
using primers HybFrigga and HybBurre (Electronic Sup-
plement Table S1). PCR reactions were performed as
above, but with an annealing temperature of 55ºC.
All amplications were performed in a PTC – 100
Thermal Cycler (MJ Research Inc.) or in an Eppendorf
Mastercycler ep thermal cycler (Perkin – Elmer Corp.).
The PCR products were puried with the MagneSil
Green sequencing clean-up reagent (Promega) and se-
quenced using ABI Big-Dye v3.1 terminator chemistry
on an ABI 3130 DNA Sequencer (Applied Biosystems)
following the manufacturer’s protocols.
Chromatograms were edited and base calls checked
using GeneMapper v4.1 (Applied Biosystems). We
searched for reading frame errors and unexpected stop
codons by translating the nucleotide sequences to pep-
tides using Mega5 (tamura et al. 2011). Sequences were
aligned with Clustal W (tHompSon et al. 1994) under de-
fault settings, and then manually adjusted. The alignment
included also GenBank sequences of some taxa within
the Pedaliodes complex (Pedaliodes, Altopedaliodes,
Pu na pedaliodes and Panyapedaliodes Forster, 1964) and
of several genera within the Pronophilina used as out-
group taxa (Manerebia, Lymanopoda, Ianussiusa Pyrcz
& Viloria, 2004, Pronophila Doubleday, [1849] and
P et al.: Taxonomy and phylogeny of Steromapedaliodes
200
Eteona Doubleday, 1848) (accession numbers given in
Table 1). In this study, we cannot designate the closest
relative to the Steromapedaliodes sensu nov. taxon, there-
fore we use a series of genera belonging to the subtribe
Pronophilina. To investigate potential conict between
the partitions of the mitochondrial and nuclear datasets,
we performed a test of Congruence Among Distance
Matrices (i.e. CADM, Campbell et al. 2011) as imple-
mented in the R-package ape (paradiS et al. 2004) and
using pairwise distances calculated with the Maximum
Composite Likelihood model (MCL, tamura et al. 2004)
in Mega5. Bayesian phylogenetic analyses of the mito-
chondrial, nuclear and mitochondrial + nuclear datasets
were performed using MrBayes 3.2 (ronquiSt et al.
2012), applying the most appropriate substitution model
for each region as estimated with the Bayesian Informa-
tion Criterion (BIC) implemented in Mega5 (tamura
et al. 2011). Four Mar kov chains were run simultane-
ously, each one was started from a random tree and run
for one million generations, sampling a tree every 100
generations. All sample points prior to reaching station-
arity (~ 1000 trees) were discarded as burn-in samples.
Data remaining after burn-in (9000 trees) were used to
generate a majority rule consensus tree, with branch sup-
ports calculated as the proportion of trees that contained
the clade and re presenting the posterior probability of
the existence of that clade given the data and model of
evolution (HuelSenbeCK & ronquiSt 2001). Probabilities
> 0.95 were arbitrarily considered indicative of signi-
cant support. Maximum likelihood (ML) analyses were
also performed in Mega5 (tamura et al. 2011), with
alignments partitioned by region (mitochondrial, nuclear
and mitochondrial + nuclear), assigning a separate model
of nucleotide substitution to each partition. Statistical
support was calculated from 100 nonparametric boot-
strap replicates under the same partitioning scheme.
Table 1. A list of the localities, altitudes, coordinates and GenBank accession numbers for Steromapedaliodes sensu novo and outgroup
taxa (Dangond dangondi, Pedaliodes plotina, Diaphanos fuscus and Tamania jacquelinae) sequenced in this study. Accession numbers:
left column: COI-COII, right column: GAPDH.
Species Locality Altitude (m) Longitude Latitude Accession Numbers
S. leukasmena Páramo de Las Rosas 3000–3200 70°07’07’’ 09°34’36’’ KF157845 KR476346
S. chiquinquirana chiquinquirana Laguna de Mucubají 3550–3650 70°48’46’’ 08°47’19’’ KF157848 KR476348
S. chiquinquirana castellana Páramo de Cabimbú 2850–2950 70°27’26’’ 09°10’34’’ KF157846 KR476347
S. chiquinquirana lossadana Teta de Niquitao 3200–3250 70°27’46’’ 09°25’37’’ KF157847 KR476349
S. empetrus empetrus La Culata 3100–3300 71°02’40’’ 08°45’59’’ KF157850 KR476340
S. empetrus bolivari Loma Redonda 3850–4000 71°04’38’’ 08°32’52’’ KF157843 KR476339
S. empetrus frailejona Los Frailes 2850–3100 70°46’37’’ 08°48’45’’ KF157849 KR476338
S. bordoni above Las Antenas 3250–3500 71°54’18’’ 08°07’44’’ KF157844 KR476344
S. lathraia Mesa Alta 2700–2800 71°52’45’’ 08°09’29’’ KF157851 KR476341
S. albarregas albarregas Qda. La Boba 3200–3250 71°16’36’’ 08°37’41’’ KF157857 KR476355
S. albonotata schuberti Páramo Guaramacal 3100–3150 70°11’05’’ 09°14’18’’ KF157853 KR476352
S. albonotata pratense El Potrero 2900–2950 70°53’15’’ 08°59’59’’ KF157856 KR476353
S. albonotata borealis Páramo Cendé 3000–3050 70°07’24’’ 09°39’40’’ KF157852 KR476351
S. albonotata josefinae Los Frailes 2900–2950 70°45’53’’ 08°49’09’’ KF157854 KR476354
S. mavarezi Páramo de Guirigay 2800–3000 70°25’41’’ 09°03’10’’ KY851302 KY851303
Dangond dangondi Casa de Vidrio 3200–3250 72°54’01’’ 10°27’36’’ KF157859,
KF157860
KR476356,
KR476357
Pedaliodes plotina plotina Colonia Tovar 1900–1950 67°16’53’’ 10°24’56’’ KF157858 KR476358
Diaphanos fuscus Páramo Cendé 3100–3150 70°07’09’’ 09°34’33’’ KF157861 KR476336
Tamania jacquelinae Betania 2350–2400 72°25’39’’ 07°27’54’’ KF157862 KR476337
Table 2. Pronophilina outgroup taxa extracted from GenBank. Accession numbers: left column: COI, right
column: GAPDH.
Outgroup taxa GenBank Locality Accession numbers
Pedaliodes ewelina Laguna Uspacocha, 3668 m, Peru DQ338856 EU528407
Pedaliodes ampayana Laguna Angascocha, 3270 m, Peru GQ357239 GQ357489
Punapedaliodes flavopunctata Cerro de Pasco, 4318 m, Peru DQ338861 GQ357493
Pedaliodes phrasiclea Quebrada 7 Jeringas, 1700 m, Peru GQ357238 GQ357488
Panyapedaliodes drymaea Laguna Angascocha, 3270 m, Peru DQ338855 GQ357486
Altopedaliodes sp. Cerro de Pasco, 4318 m, Peru GQ357223 GQ357464
Lymanopoda rana Pampa Hermosa, 1300 m, Peru DQ338853 GQ357479
Manerebia lisa Quebrada Malambo, 2600 m, Peru GQ357233 GQ357480
Ianussiusa maso Runtun, 2600 m, Ecuador GQ861947 GQ861918
Eteona tisiphone Extrema, 1000 m, Brazil DQ338849 GQ357473
Pronophila thelebe Quebrada 7 Jeringas, 1700 m, Peru DQ338859 EU528410
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3. Results
3.1. Genus-level comparative morphology
The most striking result of the comparative morphologi-
cal analysis of the adults of Steromapedaliodes (sensu
Viloria & Pyrcz) and Redonda is the absence of synapo-
morphies of either genus for any of the morphologic fea-
tures analysed.
Body size. In both groups the females are marginally, no-
ticeably or considerably smaller than males (Figs. 5 – 11).
However this feature is not necessarily phylogenetically
informative since this pattern is seen in several other par-
amo taxa in the Pedaliodes complex, including the gen-
era Dangond (Fig. 7A,B), Paramo, and Punapedaliodes,
probably related to the adaptation to paramo conditions,
as discussed by pyrCz (2010b). Males of the species until
now associated with Redonda are generally larger than
the males of Steromapedaliodes (sensu Viloria & Pyrcz),
but with some overlap. For example, larger males of some
subspecies of S. albonotata are similar in size to males of
some smaller species of Redonda. Moreover, the males,
and particularly the females of one of the new species of
Steromapedaliodes described in this study (see descrip-
tion 4 below), are larger than the males of any other spe-
cies known so far in both genera.
Wing patterns. There is one striking character present
in all the species of Steromapedaliodes (sensu Viloria &
Pyrcz) and Redonda: ocellar elements in postdiscal area
of both wings (either dorsally, or ventrally, or both), but
simplied as white, sometimes highly distorted spots, v-
shaped and elongated (fusiform) (Figs. 5 – 11). This fea-
ture is very diagnostic, unique to these two taxa (absent
in Dangond, Fig. 10A,B), and is therefore a strong syna-
pomorphy within the Pedaliodes complex. On the other
hand, there is a FWD white discal patch in most taxa of
Steromapedaliodes, except for two new species described
herein, one of which has an entirely brown dorsum (see
description 3 below) and the other has instead a white
patch in the discal cell of the hindwing (see description
4 below). The latter represents a unique feature not only
for this group of species but also for the entire Pedali-
odes complex. Other similarities in the colour patterns
between Steromapedaliodes (sensu Viloria & Pyrcz) and
Redonda consist in the elaborate markings of the HWV
imitating grass, sandy habitat or lichens.
Genitalia. Male genitalia in Steromapedaliodes (sensu
Viloria & Pyrcz), Redonda and Dangond present several
common characters, particularly a stout, slightly hooked
uncus, and a small, globular saccus (Figs. 12 – 15). The
aedeagus is straight or slightly arched, but rather short
(compared to other species of Pedaliodes complex), with-
out the apical crest or lateral serration present in most spe-
cies of Pedaliodes complex. The valvae are subtriangular
have a smooth dorsal surface without any trace of dorsal
or apical processes, again, a feature characteristic to most
species of the Pedaliodes complex (e.g. Fig. 15E: Panya-
pedaliodes panyasis (Hewitson, 1862), Fig. 15F: Pedali-
odes plotina (Hewitson, 1862)). The only apparent differ-
ence is the presence of subunci (gnathos) in S. albonotata
and S. albarregas, compared to absent or strongly reduced
subunci in Redonda. Subunci are also reduced in Dangond
(Fig. 13F), with the difference that in this genus, there is
a stout base of subunci and a wide breach between it and
the base of uncus, whereas subunci in Redonda, when pre-
sent, are short tips adhering to ventral surface of uncus.
The presence or absence of subunci in male genitalia of
the species of Pedaliodes complex was strongly empha-
sized by adamS & bernard (1977, 1979, 1981), as one
of the key diagnostic characters at the genus level. It is a
fact that subunci are present in the overwhelming major-
ity of > 300 species of the Pedaliodes complex, with the
exceptions mentioned just above. However, pyrCz (2004)
described a species from northern Peru with no subunci,
Panyapedaliodes stellata, and questioned the phyletic va-
lidity of this character. Interestingly, that species is also a
paramo denizen, which suggests that the atrophy of sub-
unci may have some adaptive role to oreal grassland habi-
tats. Also, as we will see below, the absence of subunci
in both Dangond and Redonda brings no phyletic value,
since the two groups are very distantly related.
Female genitalia in Steromapedaliodes (sensu Viloria
& Pyrcz), Redonda and Dangond (Fig. 19D: Dangond dan-
gondi), contrary to other genera of the Pedaliodes com plex
(Fig. 19E: Panyapedaliodes drymaea (Hewitson 1858),
Fig. 19F: Pedaliodes plotina), show no signa on bursa co-
pulatrix. However, this is again hardly a phyloge netically
informative character, but most likely a plastic adaptation
to life in oreal habitats. In some species of Altopedaliodes
signa are also lacking or are extremely reduced, as is also
the case in another paramo-puna genus, Pu na pedaliodes.
Ductus bursae is somewhat longer in Ste ro mapedaliodes
(sensu Viloria & Pyrcz) than in the other two genera, but
in all three ductus bursae is weakly sclerotized compared
to other taxa in the Pedaliodes complex, including Peda-
liodes, and to an even more degree compared to Panya-
pedaliodes, Corderopedaliodes Forster, 1964, Praepe-
daliodes Forster, 1964 and also Punapedaliodes, being the
latter a paramo-puna genus with a wide, slat-like strongly
sclerotized lamella. Furthermore, in all the species of
Ste romapedaliodes (sensu Viloria & Pyrcz) and Redonda,
lamella postvaginalis forms a wide shield covering the an-
trum, much narrower and less prominent than in Dangond.
This character appears to be quite variable among different
species of Pedaliodes and other species of the Pedaliodes
complex, although some phylogenetically informative
characters can be found in the shape of lamellas, for exam-
ple in Praepedaliodes (Pyrcz et al. in press).
3.2. Molecular phylogeny
The mitochondrial dataset consisted of 2226 aligned
nucleotides (COI: 1507 bp, tRNA – Leu: 67 bp, COII:
P et al.: Taxonomy and phylogeny of Steromapedaliodes
202
652 bp), of which 600 (26.95%) were variable and 453
(20.35%) were phylogenetically informative (excluding
outgroup taxa). Maximum likelihood and Bayesian anal-
yses resulted in almost identical tree topologies under
the selected model of sequence evolution, GTR + G + I.
The nuclear dataset consisted of 710 aligned nucleotide
sequences of the GAPDH gene, of which 128 (18.03%)
were variable and 172 (10.14%) were phylogenetically
informative (excluding outgroup taxa). Again, maximum
likelihood and Bayesian analyses resulted in almost the
same tree topology under the selected model of sequence
evolution, GTR + G + I. The mitochondrial and nuclear
datasets were found to be congruent based on the CADM
test (p = 0.18) and a simultaneous phylogenetic analysis
using both markers was therefore performed. As before,
maximum likelihood and Bayesian analyses of the com-
bined mitochondrial + nuclear dataset resulted in basical-
ly the same tree topology, which was very similar to the
phylogenetic trees obtained with single DNA regions and
particularly with mitochondrial data, although some dif-
ferences are observed (see below). In all the analyses, re-
gardless of the partition and phylogenetic approach used,
Steromapedaliodes (sensu Viloria & Pyrcz) and Redonda
form a clade, with bootstrap and posterior probability
values equal to 1 (Figs. 2, 3). The Bayesian and likeli-
hood analyses of the combined mitochondrial + nuclear
dataset indicate that S. albonotata is the sister group of
the remaining taxa, with the newly described species (de-
scription 4 below) being associated with S. albarregas.
In all the analyses, Redonda appears to be nested within
Steromapedaliodes (sensu Viloria & Pyrcz), sometimes
with quite high statistical support (posterior probability
> 0.85). For instance, the combined analysis indicates
that Redonda forms a clade with S. albarregas and one of
the newly described species of Steromapedaliodes (see
description 4 below).
Interestingly, two species previously associated with
the genus Redonda, R. empetrus and R. chiquinquirana
Ferrer-Paris, 2015, are extremely similar morphological-
ly, yet they are never found in sympatry although some-
times they can be found ying in localities geographical-
ly close. The molecular phylogenetic analysis indicates
Fig. 2. Molecular phylogenetic hypothesis of the genus Steromapedaliodes sensu novo and some high-altitude tropical Pronophilina gen-
era. Combined mtDNA (COI, tRNA–Leu and COII) and nuclear (GAPDH) data set. Values at the left of the slash (./) represent posterior
probabilities from the Bayesian analyses, while values at the right of the slash (/.) represent bootstrap values (0 to 1 scale) from the maxi-
mum likelihood analysis.
203
ARTHROPOD SYSTEMATICS & PHYLOGENY — 75
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that they are genetically quite distant, which suggests
that their morphological similarities are not the result of
a recent common ancestry but probably a convergence
somehow due to the adaptations to the same ecological
conditions.
In all analyses the clade Steromapedaliodes (sensu
Vi loria & Pyrcz) + Redonda appears nested within the
Pe da liodes complex, although more closely related to a
clade comprising some Pedaliodes (P. plotina, P. ampa-
yana Lamas, Viloria & Pyrcz, 2008, P. ewelina Pyrcz,
2008), Dangond and some genera belonging to Pedali-
odes complex (e.g. Altopedaliodes, Punapedaliodes).
Further taxonomic/systematic work will be necessary in
order to establish with precision the phylogenetic rela-
tionships of the clade Steromapedaliodes (including Re-
donda) within the large Pedaliodes complex.
In summary, the high phylogenetic resolution provided
by the analyses of mitochondrial and nuclear DNA mark-
ers, altogether with the morphological analyses described
above, suggest that there is no valid reason to maintain
Steromapedaliodes (sensu Viloria & Pyrcz) and Redonda
as two separate genera. We therefore propose formally to
synonimize Redonda syn.n. as a subjective junior syno-
nym of Steromapedaliodes. Thus, Steromapedaliodes
sen su novo represents a monophyletic clade composed by
two species previously associated with Steromapedalio des
(sensu Viloria & Pyrcz), ve species previously associated
with Redonda and two species described below.
3.3. Systematic overview
Steromapedaliodes Forster, 1964, sensu novo
[Steropoda Staudinger, in litt.] nomen nudum.
[Steropoda Thieme, 1905: 95, 141 (as a synonym of Pedaliodes
Butler)] nomen nudum, Synonymy established by Viloria &
pyrCz 2001].
Steromapedaliodes Forster, 1964: 148; miller 1968: 117; Huber
1973: 195, 199; adamS & bernard 1977: 273; 1981: 348 (the
latter as a synonym of Altopedaliodes); lamaS [1997]: 49 (as a
synonym of Altopedaliodes); Viloria & pyrCz 2001: 7; lamaS
et al. 2004: 215.
[Altopedaliodes Forster; adamS & bernard 1981: 348; adamS
1986: 246; deSCimon 1986: 506; Viloria & pyrCz 1994: 347;
lamaS [1997]: 49 – 50 (in part)].
[Redonda Adams & Bernard, 1981: 367 – 368, syn.n.; adamS 1983:
474; 1984: 93; adamS 1985: 38; d’abrera 1988: 871; Viloria
1994: 180, 184, 185; 2000: 266, 269, 271; pyrCz 1999: 354;
Viloria & pyrCz 2001: 1 – 2, 5 – 6, 12, 15, 17; Viloria et al.
2003: 21 – 23; pyrCz 2010a: 36, 38, 45, 179 – 186, 244; 2010b:
265 – 273.]
[Punapedaliodes Forster; Huber 1973: 195.]
[“Pedaliodes” Butler; deSCimon 1986: 510.]
Fig. 3. Molecular phylogenetic hypothesis based exclusively on the mtDNA data, otherwise as specied for Fig. 2.
P et al.: Taxonomy and phylogeny of Steromapedaliodes
204
Type species: Pedaliodes albonotata Godman, 1905 (by
original designation).
Generic diagnosis. Adults: Sexual dimorphism vari-
able, slight and expressed only in a slightly smaller size
of the females, to extreme, with very small, brachypter-
ous females. Antenna: reaching ⅔ of FW costa, mostly
naked, except for some basal agellomeres, club vari-
able between species, between attened and 2 × than
shaft to spoon like and three times thicker. Venation
(Fig. 4: FW characterized by Sc, R1 and R2 independ-
ent, R3, R4 and R5 originating from a common root near
or at base of m1 – m2; R3 ½ – ⅔ the distance from discal
cell to R4; R3 – 5 and M1 originating from the same root
or marginally separated; m1 – 2 and m2 – m3 slightly in-
curved basally; root of M2 emerging half way between
root of M1 and M3, very slightly closer to M2; a ves-
tigial discal veinlet occasionally present in some species
(S. chiquinquirana (Ferrer-Paris) comb.n., S. empetrus
(Thieme) comb.n.) but generally absent. HW venation
with humeral vein present; root of M1 much closer to
Rs than to M2; m1 – m2 bent basally; m1 – m2 2 – 2.5 ×
as long as cu1 – cu2; 1A2 and 1A3 independent. No an-
droconial scales on the FW upperside. Wings colour pat-
terns predominantly of variable shades of brown, with
some discrete patterns of yellowish submarginal dots on
the upperside, and white patches in forewing discal cell
present in most species; a diagnostic row of hindwing
underside submarginal milky white or snow white dots
shaped as arrow-heads pointing basally, the latter char-
acter being, in our opinion, the strongest qualitative syn-
apomorphy of the genus. Male genitalia: saccus globu-
lar and small; aedeagus straight or slightly arched with
no apical crest or lateral serration; valvae subtriangular
with a smooth dorsal surface without any trace of dorsal
or apical processes. Female genitalia: no signa; ductus
bursae weakly sclerotized; postvaginal lamella wide,
slat-like and strongly sclerotized, forming a wide shield
covering the antrum. Preimaginal stages have not been
entirely investigated for any extant species of this genus.
(1a) Steromapedaliodes albonotata albonotata
(Godman), stat.rev.
(Figs. 5A,B, 12A, 17F)
[Steropoda diplogramma Staudinger, in litt.] nomen nudum.
[Pedaliodes diplogramma Thieme, 1905: 96 (synonymy given);
gaede 1931: 488; forSter 1964: 148; lamaS et al. 2004: 215
(all as synonym of P. albonotata)] nomem nudum.
Pedaliodes albonotata Godman, 1905: 189, pl. 10, g. 11; tHieme
1905: 95, 96 – 97; Weymer 1912: 257, pl. 54, row d; riley &
gabriel 1924: 6; gaede 1931: 488, d’abrera 1988: 856, g.;
lamaS [1997]: 49.
Steromapedaliodes albonotata (Godman); forSter 1964: 148, g.
172 (male genitalia, erroneous); Huber 1973: 195; Viloria &
pyrCz 2001: 10; lamaS et al. 2004: 215.
Altopedaliodes albonotata (Godman); adamS & bernard 1981:
345, 348; adamS 1983: 474; 1984: 93; 1987: 38, g. 4; Viloria
1994: 187.
Material examined. Lectotype, ♂, of Pedaliodes albonotata God-
man [designated by Viloria & pyrCz 2001: 11], VENEZUELA:
Mérida, Ex Staudinger, G – S, BMNH type No. Rh. 3991; 1 ♂:
same data; 5 ♂ and 2 ♀: Mérida, Bricenno [sic] ZMHB; 1 ♂: Cor-
dillera de Mérida, S. of Mérida, La Aguada, 3500 m, 07.viii.1977,
M.J. Adams & G.I. Bernard, AB2; 3 ♂: same data, 10.viii.1977;
4 ♂: Cordillera de Mérida, S. of Mérida, SE of La Aguada, 3450
m, 20.viii.1977, M.J. Adams & G.I. Bernard, AB2; 1 ♂: Cordil-
lera de Mérida, S of Mérida, Above La Aguada, 3550 – 3650 m,
10.viii.1977, M.J. Adams & G.I. Bernard, AB2; 1 ♂: same data,
3500 m BMNH; 2 ♂ and 1 ♀: Venezuela, Edo. Mérida, P. N. Sierra
Nevada, La Aguada, 3400 – 3450 m, 01.ii.2007, T. Pyrcz leg. (male:
prep. genit. 08/18.07.2013 J. Lorenc-Brudecka, female: prep. genit.
318/12.01.2016 J. Lorenc-Brudecka), MZUJ.
Type locality: Mérida, Venezuela.
Fig. 4. Examples of wing venation of Steromapedaliodes sensu novo. A: Steromapedaliodes bordoni, Páramo del Batallón. B: Stero-
mapedaliodes chiquinquirana castellana, Páramo de Las Moras. C: Steromapedaliodes albonotata schuberti, Boconó – La Vega. — Ab-
breviations: Sc Subcostal; R Radial; M Median; CuA Cubital; A Anal; H Humeral; DC Discal cell.
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ARTHROPOD SYSTEMATICS & PHYLOGENY — 75
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Redescription. MALE (Fig. 5A): Head: Eyes glabrous,
lustrous, chocolate brown with irregular seal brown
patches, covered with long and dense black hair. Labial
palpi 2 × length of head, covered mostly with blackish
brown hairy scales, except for some basal sandy yellow
ventral hair, ventrally three times longer than dorsally,
and adorned with some sparse snow-white scales. Frons
with a tuft of long blackish brown hair. Antennae slen-
der, composed of 33 agellomeres, russet brown, mostly
naked except for some snow white and grey scales on
basal and sub-clubbal segments; club 2 × wider than
shaft, attened dorso-ventrally, composed of 10 ag-
ellomeres, ventrally lighter, orange brown. Thorax:
black, dorsally covered with blackish brown and navy
blue scales on metathorax, and sparse golden brown
hair, denser on patagia; ventrally covered with dense
brown, grey and sandy yellow hair; mid and hind legs
russet brown, femorae mostly naked except for a crest
of brown and grey hair, tibiae and tarsi densely covered
with steely grey hair, numerous ventral spines on tarsi.
Wings: FW (19 – 20 mm) with a subacute apex, slightly
convex distal margin and gently bent at tornus; fring-
es dark grey at vein ends and snow white in between.
FWD uniform dark chocolate brown, except for sparse
snow white scales along costa and in apical area, a snow
white elongated mid discal patch, and two snow white
submarginal dots in M1 – M2 and CuA1 – CuA2. FWV
uniform chocolate brown, lustrous, a shade lighter than
on the upperside, with snow white scaling denser than on
the upperside along costa and in apical area, and along
distal margin to M3; discal cell white patch not appar-
ent; a row of submarginal snow white minute dots from
R5 – M1 to CuA1 – CuA2, somewhat more prominent in
M1 – M2. HW oval with a slightly undulated distal mar-
gin; fringes longer than on the FW, mostly dark grey,
white scales in intravenal spaces, scarcer than on the
FW. HWD uniform dark chocolate brown, covered with
hairy scales in median half and along anal margin. HWV
ground colour blackish brown and bistre brown with a
heavy and regular overcast of snow white scales, with a
few concentrations producing a more denite pattern, a
short mid-costal streak, and a row of submarginal dots,
two arrow-head like in Rs – M1 and, a much longer one
in M1 – M2, and three half-moon shaped in M2 – M3,
M3 – CuA1 and CuA1 – CuA2. Abdomen: dorsally and
laterally covered with black, and basal segments, lus-
trous navy blue scales and dense, golden brown hair, ven-
trally with densely packed up scales of different colours,
brown, grey and sandy yellow, and sparser hair. Male
genitalia (Fig. 12A): Uncus aligned to tegumen shoul-
der, approximately the same length, slightly arched, and
with a blunt tip; subunci ⅔ the length of uncus, slender
and compressed laterally; pedunculus short, subacute;
vinculum s-shaped; saccus wide but shallow; valvae the
length of tegumen + half uncus, elongated and slender,
with a delicately irrorated dorsal surface and a subacute
tip; aedeagus tubular, slightly arched, not contorted, with
the proximal opening half the entire end, and an acute
tip, without any crest or lateral teeth. FEMALE (Fig.
5B): Sexual dimorphism slight, lighter brown, especially
on ventral surface, HWV duller, with sparser snow white
pattern restricted to the costal streak and submarginal
dots, with a noticeable median darker brown band. Fe-
male genitalia (Fig. 17F): As illustrated.
Remarks. godman (1905), in the original description of
Pedaliodes albonotata, referred to four males, two la-
belled Mérida in his collection (acquired from Stauding-
er) and two in the BMNH (one labelled La Culata and
one La Pedregosa). Accordingly, Viloria & pyrCz (2001:
gs. 1, 2) designated the Lectotype from Godman’s col-
lection. However, their treatment of S. albonotata was
somewhat simplistic. They considered that the species is
monotypic, and that the differences presented by several
populations of S. albonotata from central and southern
Cordillera de Mérida are negligible. They also failed to
point out a detail that reveals crucial for the identica-
tion of the nominotypical subspecies. In fact, only the
nominotypical subspecies of S. albonotata has conspicu-
ous FWD white submarginal spots. The nominotypical
subspecies of S. albonotata, as far as currently known,
is restricted to the northern (El Chama) slopes of the Si-
erra Nevada, where it was collected along the La Mucuy
– Laguna Verde and La Aguada – Loma Redonda trails.
(Fig. 1C)
(1b) Steromapedaliodes albonotata thiemeiana Pyrcz,
repl. name
(Figs. 7G,H, 12B, 17B)
Pedaliodes albonotata ab. privigna Thieme, 1905: 96 – 97.
Altopedaliodes albonotata ab. privigna (Thieme); gaede 1931:
488; adamS & bernard 1981: 348 (as synonym of Altopedali-
odes albonotata).
Steromapedaliodes albonotata ab. privigna Thieme; Viloria &
pyrCz 2001: 10; lamaS et al. 2004: 215 (as synonym of Stero-
mapedaliodes albonotata).
Material examined. Paralectotype, ♂, of Pedaliodes albonotata
Godman, VENEZUELA: Pedregosa; Paralectotype, ♂, of Pedali-
odes albonoatata Godman: Culata, CB; 1 ♂: Quintero, [S.] Brice-
ño, RB; 1 ♂: Quintero, 14.iv.1898, [S.] Briceño, RB; 1 ♂: Quintero,
04.v.1898, [S.] Briceño, RB; 1 ♂: no data, RB; 3 ♂ and 1 ♀: Méri-
da, Ex Grose Smith, 1910, JB; BMNH; 1 ♂: E. Mérida, Campo Ella
[sic] Dist., Páramo de Conejos, 13000 ft., 09.ix.1938, J. Hanbury
Tracey, Brit. Mus. 1939 – 117; 1 ♂ and 1 ♀: E. Mérida, Páramo
Campanario, 13500 ft., 18.ix.1938, J. Hanbury Tracey, Brit. Mus.
1939 – 117; 1 ♂: Cordillera de Mérida, N. of Mérida, Río Albarre-
gas, 3300 m, 14.viii.1977, M.J. Adams & G.I. Bernard, AB2; 10 ♂:
same data, 3400 m, 15.viii.1977; 1 ♂: same data, 2650 m; 1 ♂:
same data, 3650 m; 2 ♀: same data, 3450 m. BMNH; 1 ♂: Vene-
zuela, x.23 von Dr. Lück, Ex Coll. Fruhstorfer, Collection v. Rosen,
Praparät Nr. SA 439, ZSM; 1 ♂: Edo. Mérida, Laguna Negra, 3480
m, 12.ix.1971, H. Huber, ZSBS; 7 ♂ and 1 ♀: Edo. Mérida, Parque
Nacional La Culata, Alto Mucujún, 3400 – 3500 m, 17.iv.1996, T.
Pyrcz leg. (male: prep. genit. 03/18.07.2013 J. Lorenc-Brudecka);
5 ♂: same data but 2900 – 2950 m; 1 ♂: same data but 2950 – 3050
m; 5 ♂: same data but 3350 – 3400 m, 25.XII.1991; 1 ♂: same
data but 29.XII.1991; 5 ♂: same data 3000 – 3050 m, 01.iv.1992,
T. Pyrcz leg. (1 prep. genit. 01/18.07.2013 J. Lorenc-Brudecka);
12 ♂ and 2 ♀: same data but 3200 – 3250 m, 03.iv.1992; 1 ♂:
Edo. Mérida, Parque Nacional La Culata, Mucujún, 01.iv.1992, T.
P et al.: Taxonomy and phylogeny of Steromapedaliodes
206
Pyrcz leg.; 2 ♂: Edo. Mérida, locality?; 1 ♂: Edo. Mérida, Parque
Nacional La Culata, Monte Zerpa, 2200 – 3000 m, 20.ii.1996, T.
Pyrcz leg.; 1 ♂: same data but 2950 – 3000 m, 31.v.1992; 1 ♂:
same data but 31.v.1991, prep. genit. 1010/18 T. Pyrcz; 1 ♂: same
data but 3250 – 3300 m, 13.ii.1996, J. Wojtusiak leg.; 1 ♂: Edo.
Mérida, Parque Nacional La Culata, Alto Mucujún, 3400 – 3500
m, 06.ii.2007, T. Pyrcz leg.; 7 ♂: same data but 01.ii.2008; 1 ♂:
Mérida, Quebrada Achotón, 3200, 09.i.1992, T. Pyrcz; 5 ♂ and
2 ♀: Edo. Mérida, Parque Nacional La Culata, Sector Monter-
rey, 3000 – 3050 m, 06.iv.2006, T. Pyrcz leg.; 1 ♂: Edo. Mérida,
Parque Nacional La Culata, Jaji – Las Gonzales, Qda. La Boba,
3200 – 3250 m, 18.vii.2006, T. Pyrcz leg.; MZUJ; 5 ♂ and 2 ♀:
Edo. Mérida, La Culata, “Cabaña de los Curas”, sector Monter-
rey, 3100 – 3150 m; 06.iv.2006, P. Boyer leg.; 8 ♂ and 1 ♀: Edo.
Mérida, La Culata, 3000 – 3200 m, 3.iv.2006, P. Boyer leg., PBF.
Type locality: Mérida, Venezuela.
Diagnosis. MALE (Fig. 7G): This subspecies differs
from the nominate S. albonotata in several aspects. First
of all, its FWD discal cell white patch, although quite
variable in size, is always wider, approximately the width
and shape of S. albonotata australis ssp.n., whose patch
has a characteristic steely sheen. In most examined indi-
viduals there are two, whitish FWD submarginal dots but
they are invariably very small or obsolete, compared to
conspicuous dots in S. albonotata albonotata. Important-
ly, the HWV is dull, and snow-white scaling is sparse,
compared to the nominate subspecies or indeed to S. al-
bonotata australis ssp.n. restricted to one or two costal
streak and the submarginal dots, with the two arrowhead
shaped much shorter and thinner. Male genitalia (Fig.
12B): Do not differ from the nominate subspecies except
for the straight uncus. FEMALE (Fig. 7H): It is approxi-
mately the size of the male, typically slightly lighter col-
oured on the upperside and paler on the underside, espe-
cially on the FWV which has a rusty sheen, with a sandy
yellow suffusion on the HWV, however not as prominent
as in some other subspecies, in this respect similar to S.
albonotata borealis ssp.n. Female genitalia (Fig. 17B):
As illustrated.
Etymology. This subspecies is dedicated to the eminent
German Lepidopterist Otto Thieme, author of excellent
systematic monographs of the Pronophilina.
Remarks. In the original description of S. albonotata
godman (1905) noticed that the specimens from la Culata
and La Pedregosa have very small, or obsolete two sub-
marginal (white) spots on the FWD compared to the type
specimens. tHieme (1905) called these subsequently as
an aberration privigna. adamS & bernard (1981) obvi-
ously misunderstood the description of privigna, as they
stated that this aberration corresponds to the individuals
in which two white postdiscal spots on the forewing show
through on the underside[!]. Viloria & pyrCz (2001) in
their treatment of Steromapedaliodes (sensu Viloria &
Pyrcz) did not deal with the name privigna and simply
quoted its reference. None of the consulted authors had
associated the expression of the white spots with any par-
ticular population. It is however evident from extensive
sampling throughout the range of S. albonotata, that the
individuals whose white spots are faint or absent, cor-
responding with the diagnosis of privigna, represent a
geographic form. Nevertheless, the name privigna was
clearly created as an infrasubspecic epithet (ICZN arti-
cle 45.6.2). For this reason, a new name is proposed. S.
albonotata thiemeiana occurs on the SE Chama valley
slopes of the Sierra de La Culata (La Pedregosa, Monte
Zerpa, La Culata, Mucujún, Qda. Achotón) and locally in
the watershed of Chama (Mucubají) (Fig. 1C).
(1c) Steromapedaliodes albonotata australis Pyrcz &
Boyer, ssp.n.
(Figs. 5C – H, 12C, 17C,D)
Material examined. Holotype, ♂, VENEZUELA: Edo. Mérida /
Táchira border, El Batallón, Mesa Alta, 2900 – 2950 m, 03.xii.2005,
T. Pyrcz leg., MZUJ, to be deposited in MIZA; Paratypes (126 ♂
and 21 ♀): 14 ♂ and 2 ♀: Edo. Mérida, San José, Páramo de San
José, 3000 – 3050 m, 05.xii.2007, T. Pyrcz leg. (1 male genit.
15/09.03.2006 T. Pyrcz); 1 ♂: same locality but 2800 – 2850 m,
11.ii.2007, T. Pyrcz leg.; 14 ♂: same data but 12.ii.2007; 6 ♂
and 6 ♀: same locality but 2950 – 3000 m, 20.ii.2008, T. Pyrcz
leg. (prep. genit. 09/18.07.2012 J. Lorenc-Brudecka); 5 ♂: same
locality but 2900 – 2950 m, 28.ii.2010; 21 ♂ and 3 ♀: Edo. Mé-
rida, Tovar – Guaraque, Las Antenas, 2550 – 2600 m, 04.XII.2005,
T. Pyrcz leg. (1 prep. genit. 02/18.07.2013 J. Lorenc-Brudecka);
16 ♂ and 1 ♀: Edo. Mérida / Táchira border, El Batallón, Mesa
Alta, 2900 – 2950 m, 03.XII.2005, T. Pyrcz leg. (1 prep. genit.
04/18.07.2013 J. Lorenc-Brudecka); 4 ♂: Edo. Táchira, Páramo El
Batallón, Antena – Cenegón, 3300 – 3350 m, 01.ii.2008, T. Pyrcz
leg.; 3 ♂ and 1 ♀: Edo. Táchira, Páramo El Batallón, vía El Púlpito,
3500 – 3800 m, 03.iii.1996, T. Pyrcz; 4 ♂: Edo. Táchira, Páramo
El Rosal, 2950 – 3050 m, 01 – 02.iii.1996, T. Pyrcz (1 prep. genit.
08/18.07.2013 J. Lorenc-Brudecka); 10 ♂ and 1 ♀: same locality
but 3000 – 3050 m, 01.xii.2005; 1 ♂: Edo. Mérida, locality? MZUJ;
7 ♂ and 1 ♀: Edo. Mérida, San José vers Mucutuy km11, 3000,
5.xii.2005, P. Boyer leg.; 2 ♀: Edo. Mérida, Páramo de San José,
3100 m, 20.ii.2008, P. Boyer leg.; 7 ♂ and 1 ♀: Edo. Mérida, Mesa
Alta, La Grita-Pregonero km3 3 – 5, bifurcation Pregonero vers
Bailadores km 5 – 6, 2850 m, 29.xi.2005, P. Boyer leg.; 1 ♂: Edo.
Mérida, Mesa Alta, Bailadores-Pregonero km 33,5, 5,5 km après
la jonction de la Grita, 2750 m, 02.xii.2005, P. Boyer leg.; 8 ♂
and 1 ♀: Edo. Táchira, Páramo El Rosal, La Grita vers San José
de Bolivar km 22, 3350 m, 01.xii.2005, P. Boyer leg.; 1 ♀: Edo.
Táchira, Antenas, via a Cenegón, Páramo El Batallón, 3200 – 3300
m, 19.ii.2008, P. Boyer leg.; 9 ♂ and 1 ♀: Edo. Mérida, Las Ante-
nas, Tovar vers Guaraque km 22, 04.xii.2005, P. Boyer leg., PBF.
Type locality: Mesa Alta, El Batallón, Estado Táchira –
Estado Mérida border, Venezuela.
Diagnosis. MALE (Fig. 5C,E,G): This subspecies is
slightly smaller (FW length: 18 – 20 mm) than most other
subspecies, probably except for the nominate and S. al-
bonotata borealis ssp.n., although the infrasubspecic
variation is appreciable, with the smallest individuals
of all reported from the Páramo de Guaraque. The FWD
discal cell white patch is also rather variable across the
subspecies range and within populations, but it is larger
than in the nominate and also from S. albonotata thie-
meiana, in some individuals marginally entering the base
of CuA1 – CuA2 and CuA2 – 1A1B. It has a character-
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ARTHROPOD SYSTEMATICS & PHYLOGENY — 75
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istic steely sheen not apparent in any other subspecies.
In most individuals there are two vestigial submarginal
white dots on the FWD, but in some a third one shows
in M3 – CuA1. The HWV is more brightly patterned than
in S. albonotata thiemeiana, with more whitish scaling,
and comparable to the nominate, but the arrowhead sub-
marginal spots are larger than in the latter subspecies,
except for space M1 – M2. Male genitalia (Fig. 12C):
Do not differ noticeably from the nominate subspecies.
FEMALE (Fig. 5D,F,H): Slightly smaller than the male,
duller and slightly lighter brown on the upperside, lighter
brown on the FWV and considerably lighter with a sandy
yellow suffusion on the HWV. Female genitalia (Fig.
17C,D): As illustrated.
Etymology. The epithet of this subspecies refers to its
southerly distribution relative to other populations of S.
albonotata.
Remarks. S. albonotata australis ssp.n. has a wider
range than any other subspecies and is probably also the
most variable across its range. However, there are some
morphological and genetic traits that allow its identica-
tion. It is usually very common wherever it occurs. It ies
in humid lower paramo, and in some areas it occurs at
much lower elevation than any other subspecies. In the
Páramo de Guaraque area it can be found down to 2450
m, which is also due to the fact that the timberline is lo-
cally situated at a lower elevation than elsewhere in the
Cordillera de Mérida (Fig. 1C).
(1d) Steromapedaliodes albonotata sanchezi Viloria &
Pyrcz, stat.n.
(Figs. 6C, 12D)
Steromapedaliodes sanchezi Viloria & Pyrcz, 2001: 12, 13, gs. 5,
6, 11; lamaS et al. 2004: 215.
Material examined. VENEZUELA: 1 ♂: Estado Mérida, Parque
Nacional Sierra Nevada, Páramo del Tisure, 3300 – 3700 m, 28/29.
xii.1994, J. Camacho & M. García (Holotype of Steromapedali-
odes sanchezi Viloria & Pyrcz); 14 ♂: same data as the holotype
(Paratypes of S. sanchezi Viloria & Pyrcz) MALUZ; 18 ♂ and
1 ♀: Venezuela, Estado Barinas, Parque Nacional Sierra Nevada,
Los Morritos, 3000 – 3050 m, 06.i.2006, M. Costa leg. (1 prep.
genit. 15/18.07.2013 J. Lorenc-Brudecka); 1 ♂: same data but
04.i.2006 (prep. genit. 0⅔0.05.2006 T. Pyrcz); 2 ♂: same data but
17.ii.2010, T. Pyrcz leg. (1 prep. genit. 12/18.07.2013 J. Lorenc-
Brudecka) MZUJ; 1 ♂ and 1 ♀: Edo. Barinas, Los Morritos, 3000
m, 06.i.2006, M. Costa leg., PBF.
Type locality: Páramo El Tisure, Parque Nacional Sierra
Nevada, Estado Mérida, Venezuela.
Diagnosis. MALE (Fig. 6C): This subspecies can be
recognized immediately from any other, except for S. al-
bonotata pratense ssp.n., by the very large FWD snow
white discal cell patch which penetrates into the base of
CuA1 – CuA2, and extends widely into CuA2 – 1A/1B as
far as vein 1A/1B and basally nearly to wing base. In
this respect this subspecies is almost identical to the next,
S. albonotata pratense ssp.n. However, in S. albonotata
sanchezi ssp.n. the presence of some chestnut and sandy
yellow scales and denser hair give it a lighter appearance
than in S. albonotata pratense ssp.n. There are no FWD
submarginal white dots, or they are vestigial, whereas
in S. albonotata pratense ssp.n. there is usually a small
CuA1 – CuA2 white dot. Another difference between the
two subspecies is the lighter chocolate brown colour of
the FWV in S. albonotata sanchezi ssp.n. compared to
blackish brown in S. albonotata pratense ssp.n. Male
genitalia (Fig. 12D): Do not differ noticeably from the
nominate subspecies except for the more prominently
arched aedeagus and the slightly lifted uncus tip. FE-
MALE: Not examined.
Remarks. This taxon was described as a separate species.
Morphological differences between S. albonotata san-
chezi ssp.n. and the nominotypical S. albonotata affect
mostly the signicantly larger wingspan in S. a. sanchezi
ssp.n., and its considerably larger size of the FWD post-
basal white patch. Male genitalia of S. a. sanchezi ssp.n.
and S. a. albonotata are, however, almost indistinguish-
able. Considered the lack of sympatry, slight morpholog-
ical differences and molecular phylogeny evidence, the
two are considered here as subspecies of the same spe-
cies. S. albonotata sanchezi ssp.n. is found on the east-
ern, Llanos slopes of the Sierra Nevada. It has been found
only in a geographically restricted area in the north of the
Sierra Nevada and southern extreme of the Serranía de
Santo Domingo, however it is most probably much wider
widespread to the south where, however, the Sierra Ne-
vada eastern slopes have no trail access (Fig. 1C).
(1e) Steromapedaliodes albonotata pratense Pyrcz &
Boyer, ssp.n.
(Figs. 6A,B,D, 12E, 16B)
Material examined. Holotype, ♂, VENEZUELA: Edo. Mérida, El
Aguila – Piñango, Sector Potrero Km 26, 2900 – 2950 m, 09.iv.2006,
T. Pyrcz leg. (to be deposited in MIZA), (prep. genit. 03/30.05.2006
T. Pyrcz); Paratypes (17 ♂, 5 ♀): VENEZUELA: Edo. Mé-
rida, El Aguila – Piñango, Sector Potrero Km 26, 2900 – 2950 m,
09.iv.2006, T. Pyrcz leg., (1 prep. genit.: 03/30.05.2006 T. Pyrcz);
(19 ♂, 4 ♀): same locality but 3100 – 3150 m, 04.iv.2006, (2 prep.
genit.: 14/18.08.2013 J. Lorenc-Brudecka, 18/18.07.2013 J. Lorenc-
Brudecka); 1 ♂: same locality but 08.iv.2006; (2 ♂), same locality
but 3300 – 3350 m, 03.ii.2008; (5 ♂, 1 ♀), no data (1 prep. genit.
02/12.10.2011 J. Lorenc-Brudecka); 1 ♀: Edo. Trujillo, Timotes –
Piñango, Alto de Tafayes, 2650 – 2700 m, 26.ii.2006, M. Costa leg.,
MZUJ; (10 ♂, 8 ♀): Edo. Mérida, vía El Aguila – Piñango km 26,
Potrero, 3200 – 3300 m, 08.iv.2006, P. Boyer leg.; 1 ♂: Edo. Mé-
rida, col de la route de Tafayes à Piñango 3100 m, 30.vii.2009 P.
Boyer leg.; 1 ♂: Edo. Mérida, Potrero, route de Piñango, 3200 m,
30.vii.2009, P. Boyer leg., PBF.
Type locality: Sector El Potrero, vía Pico Aguila –
Piñango, Estado Mérida, Venezuela.
Diagnosis. MALE (Fig. 6A,B): This subspecies can
be recognized immediately from any other, except for
P et al.: Taxonomy and phylogeny of Steromapedaliodes
208
S. albonotata sanchezi ssp.n., by the large size (FW
length 23 – 25 mm), and a very large FWD snow white
discal cell patch which penetrates into the base of
CuA1 – CuA2, and extends widely into CuA2 – 1A/1B as
far as vein 1A/1B and basally nearly to wing base. How-
ever, in S. albonotata pratense ssp.n. the HWD is darker
and less hairy than in S. albonotata sanchezi ssp.n. There
is usually one FWD vestigial submarginal white dot in
CuA1 – CuA2. FWV is blackish brown, darker that the
chocolate brown of S. albonotata sanchezi ssp.n. HWV
ground colour is almost black and snow white pattern is
prominent, with a heavy white scale suffusion over the
entire wing surface, a mid-costal streak generally longer
than in the nominate, extending as far as vein M1, simi-
larly in this respect to S. albonotata sanchezi ssp.n. All
the submarginal dots are arrow-head shaped but the larg-
est of all, the one in M1 – M2, is slightly longer than in
the nominate subspecies, also similar to S. albonotata
sanchezi ssp.n. and considerably larger and longer than
in S. albonotata thiemeiana. Male genitalia (Fig. 12E):
Not differing form the nominate subspecies except for
the slightly lifted uncus tip. FEMALE (Fig. 6D): Smaller
than the male; wings colour pattern paler and duller dor-
sally and ventrally; HWV submarginal arrowhead white
dots considerably smaller and shorter. Female genitalia
(Fig. 16B): Papillae anales membranous in apical half,
gently rounded in lateral view, covered with short, deli-
cate and rather dense hair, sclerotized in basal half, with
developed, apohyses posteriors short; sternite viii, in lat-
eral view, as sclerotized as the basal part of papillae ana-
les, without spiracle; viii fused with tergum viii in lateral
view; postvaginal lamella sclerotized, wide, entirely cov-
ering the antrum; ductus bursae narrow, membranous, ½
the length of bursa, gradually opening into an oval bursa
copulatrix, without signa; ductus seminalis connecting to
bursa near the opening of ductus.
Etymology. This subspecies epithet is derived from the
type locality, El Potrero, meaning meadow, thus pratense
in Latin.
Remarks. This is a puzzling subspecies. It is apparently
most closely related to S. albonotata sanchezi ssp.n. as
indicated by both morphological and molecular data.
However, the two are widely separated geographically
by the high central part of the range, as they occur on
the opposite slopes of the Cordillera. In between, another
quite distinctive subspecies is found, S. albonotata jose-
nae ssp.n. Considered the above, it seems plausible that
they differentiated in the process of allopatric divergence
through extinction in the central part of the range, nowa-
days extremely dry but apparently connected by an eco-
logical corridor during a warmer and more humid phase.
For the time being, S. albonotata pratense ssp.n. is known
only from the type locality situated in NW, Lago de Ma-
racaibo slopes of the Sierra de La Culata, however it is
most probable that its distribution extends southwards
where there are very few access trails to the subparamo
(Fig. 1C). S. albonotata pratense ssp.n. has been found
only ying in humid meadows within the forest paramo
ecotone at 2900 – 3150 m approximately. Male genitalia
differ marginally from the nominate subspecies, as in S.
a. sanchezi ssp.n. the uncus tip is slightly more lifted, and
valvae are even narrower than in other subspecies.
(1f) Steromapedaliodes albonotata josenae Pyrcz &
Boyer, ssp.n.
(Figs. 7C – F, 12F, 17E)
Material examined. Holotype, ♂, VENEZUELA: Edo. Mérida,
El Baho, El Hatico, 3000 – 3050 m, 19.XI.2005, T. Pyrcz leg. (cur-
rently in MZUJ, to be deposited in MIZA); Paratypes (69 ♂ and
12 ♀): 6 ♂ and 1 ♀: Edo. Mérida, Mucubají – Los Frailes, Qda.
Km 4, 3200 – 3250 m, 22.xi.2005, T. Pyrcz leg.; 3 ♂: same locality
but 3400 – 3450 m, 02.ii.2008; 4 ♂: Edo. Mérida, Mucubají – Santo
Domingo, Los Frailes, 3000 – 3050 m, 08.vii.2006, T. Pyrcz leg. (1
prep. genit. 16/18.07.2013 J. Lorenc-Brudecka); 2 ♂: same data
but 31.vii.2009; 1 ♂: Parque Nacional Sierra Nevada, Mucubají,
3500 – 3550 m, 28.iii.1992, T. Pyrcz leg., prep. genit. 11/18.07.2013
J. Lorenc-Brudecka; 1 ♂ and 1 ♀: Edo. Mérida, El Baho, El Ha-
tico, 3000 – 3050 m, 19.xi.2005, T. Pyrcz leg.; 6 ♂: same data but
21.xi.2005; 3 ♂: same data but 24.xi.2005; 1 ♂: same data but
14.xi.2005; 4 ♂: Edo. Mérida, El Baho, via paramo, 3050 – 3150
m, 26.xi.2005, T. Pyrcz leg.; 1 ♂ and 1 ♀: Edo. Mérida, El Baho,
La Cienaga, 2900 m, 26.xi.2005, T. Pyrcz leg.; 2 ♂: Edo. Mérida,
Parque Nacional Sierra Nevada, El Baho, 3000 – 3050 m, 28.i.2008,
T. Pyrcz leg.; 5 ♂ and 2 ♀: Edo. Mérida, Pueblo Llano – Tuñame,
Qda. Rancheria, 3000 – 3050 m, 27.xi.2005, T. Pyrcz leg., (1 fe-
male genit. 330/19.01.2016 J. Lorenc-Brudecka); 2 ♂: Edo. Tru-
jillo, Tuñame – Esnujaque, La Morita, 2900 – 2950 m, 16.ii.2010,
T. Pyrcz leg.; 1 ♂: Edo. Trujillo, Timotes SE, Antenas, 2850 – 2900
m, 29.vii.2009, T. Pyrcz leg.; 1 ♂: no data; 2 ♂: Edo. Mérida, La-
guna Negra, 28.iii.1992, T. Pyrcz leg., MZUJ; 5 ♂ and 1 ♀: Edo.
Mérida, Oberes Santo Domingo-tal, 2800 m, 21.iii.1971, H. Huber;
1 ♂ and 1 female, same data, 3050 m, 05.ix.1971, (paratypes of
S. sanchezi Viloria & Pyrcz) ZSM; 1 ♂ and 1 ♀: Mérida, km 12.5,
Apartaderos – Santo Domingo, ca. 3100 – 3200 m, 26.ix.1997, A.
Neild [AFN]; 2 ♂: Edo. Mérida, El Baho, Santo Domingo vers
Apartaderos km 4, 3300 m, 24.xi.2005, P. Boyer leg.; 4 ♂ and 3 ♀:
Edo. Mérida, El Baho, Santo Domingo vers Apartaderos km 4,
2850 – 3100 m, 21.xi.2005, P. Boyer leg.; 2 ♂ and 2 ♀: Edo. Mérida,
El Baho, Santo Domingo vers Apartaderos km 4, 2850 – 3100 m,
26.xi.2005, P. Boyer leg.; 1 ♂: Edo. Mérida, El Baho, Santo Domin-
go vers Apartaderos km 4, 2800 – 3000, 19.xi.2005, P. Boyer leg.;
5 ♂ and 1 ♀: Mérida, Ranchería vers Niquitao km7, (route Pueblo
Llano – Niquitao), 3050 m, 27.xi.2005, Pierre Boyer leg.; 1 ♂: same
data but 23.xi.2005; 1 ♂: Mérida, SE de Timotes, près de la route
de Pueblo Llano, 2700 – 2800 m, 29.vii.2009, Pierre Boyer leg.;
6 ♂ 1 ♀: Edo. Mérida, Mucubají vers Los Frailes km 4, 3400 m,
22.xi.2005, P. Boyer leg.; 1 ♂: same locality but 3200 m, 02.iv.2006;
1 ♂: Edo. Mérida, Los Frailes, Santo Domingo vers Apartaderos km
10, 2900 – 3000 m, 19.iv.2006, P. Boyer leg., PBF.
Type locality: Estado Mérida, Santo Domingo Valley, El
Baho.
Diagnosis. MALE (Fig. 7C,E): This is a large subspe-
cies (FW length: 20 – 22.5 mm), about the same size as
S. albonotata sanchezi ssp.n., S. a. pratense ssp.n. and
S. a. schuberti., considerably larger than the nominate,
S. a. thiemeiana and S. a. australis ssp.n. Its FWD discal
cell snow white patch is smaller than in S. a. sanchezi
ssp.n. and S. a. pratense ssp.n., does not extend into wing
base and penetrates only marginally into base of cells
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ARTHROPOD SYSTEMATICS & PHYLOGENY — 75
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CuA1 – CuA2 and CuA2 – 1A/1B, which is still more
than S. a. schuberti, which has an even smaller white
patch, narrower, rectangular rather than square, and
never extending, not even marginally into bases of the
two cells. Some individuals present a tiny white submar-
ginal dot in cell CuA1 – CuA2. FWV is blackish brown,
similar to S. a. pratense ssp.n. and S. a. schuberti, but
darker than the chocolate brown of S. a. sanchezi ssp.n.
HWV, although quite variable, is generally less speck-
led with white scales, mid – costal streak extends to vein
M2, and the white arrow-head shaped submarginal dots
are considerably smaller and shorter than in the above
mentioned two subspecies. Male genitalia (Fig. 12F):
similar to the nominate subspecies except for the straight
uncus, as in S. a. sanchezi ssp.n., and a longer, gently s-
shaped aedeagus, valvae are slightly longer than in other
subspecies, nearly approaching the length of tegumen +
uncus. FEMALE (Fig. 7D,F): Slightly smaller than the
male, FW and HWD ground colour lighter brown; FWV
slightly lighter brown and duller; HWV with a sandy yel-
low overcast and a noticeable darker brown median band.
Female genitalia (Fig. 17E): As illustrated.
Etymology. This subspecies is called after Josena
Balzam, a Venezuelan citizen of Polish origin, the owner
of the area of El Baho where it was discovered, in rec-
ognition for her help during the carrying out of this re-
search.
Remarks. This subspecies is geographically squeezed
within the west-east oriented S. a. pratense ssp.n. and
S. a. sanchezi ssp.n. on the one hand, and north-south
oriented S. a. thiemeiana and S. a. borealis ssp.n. on the
other. It is also morphologically intermediate between
the four as far as its wing size and the diagnostic FWD
discal cell snow patch size and shape are concerned. It
seems to be limited to the small geographical region
comprised within the Santo Domingo valley (Fig. 1C).
This is a similar distribution pattern as for the sympatric
Lymanopoda marianna valentinae Pyrcz, 2009 (pyrCz
et al. 2009). S. a. josenae ssp.n. occurs in humid gul-
lies amidst bamboo clumps but also in open, and drier
paramo dominated by Espeletia.
(1g) Steromapedaliodes albonotata schuberti Viloria
& Pyrcz, stat.n.
(Figs. 7A,B, 13A, 17A, 21C)
Steromapedaliodes schuberti Viloria & Pyrcz, 2001: 13 – 14, gs.
7, 8, 12; lamaS et al. 2004: 215.
Material examined. Holotype, ♂, of Steromapedaliodes schuberti
Viloria & Pyrcz, VENEZUELA: Trujillo, Parque Nacional Guara-
macal, vía Boconó – Guaramacal, 1500 – 2700 m, 29.viii.1991, J.
De Marmels; 2 ♂: same data, Paratypes of S. schuberti Viloria
& Pyrcz MIZA; 9 ♂ and 3 ♀: Estado Trujillo, Parque Nacional
Guaramacal, Páramo del Guaramacal, 2800 – 3100 m, 24.viii.1995,
J. Camacho & M. García; 7 ♂ and 1 ♀: Estado Lara, Parque Na-
cional Dinira, Páramo del Cendé, 3100 m, 08.viii.1996, M. García,
Paratypes of S. schuberti Viloria & Pyrcz; 1 ♂: Páramo de Ortíz,
2850 – 3100 m, Estado Trujillo, 12.xi.1991, A. Viloria & E. Moscó,
MALUZ; 17 ♂: Trujillo, Parque Nacional Guaramacal, Boconó La
Vega, 2900 – 2950 m, 15.ii.2007, T. Pyrcz leg.; 11 ♂ and 1 ♀: same
data but 16.ii.2007; 2 ♂ and 1 ♀: Parque Nacional Guaramacal, via
Las Antenas, 3000 – 3050 m, 08.xii.2005, T. Pyrcz leg.; 16 ♂ and
2 ♀: Parque Nacional Guaramacal, Paso – Las Antenas, 2900 – 2950
m, 26.iv.2007, T. Pyrcz leg. (2 prep. genit. 05/18.07.2013 J. Lor-
enc-Brudecka, 07/18.07.2013 J. Lorenc-Brudecka); 1 ♀: Parque
Nacional Guaramacal, via La Vega de Guaramacal, 2800 – 2850
m, 22.xii.2004, M. Costa leg., MZUJ; 5 ♂ and 3 ♀: Edo. Trujillo,
Massif du Guaramacal, Las Antenas, 2700 – 2800 m, 26.vii.2009, P.
Boyer leg.; 2 ♂males, 1 ♀: Edo. Trujillo, Boconó vers Guaramacal,
Les Antennes, 2850 m, 08.xii.2005, P. Boyer leg.; 1 ♂: Edo. Tru-
jillo, route Boconó vers La Vega, Guaramacal, 2700 m, 17.ii.2007,
T. Pyrcz leg., PBF.
Type locality: Vía Boconó – Guaramacal, Parque Na-
cional Guaramacal, Estado Trujillo, Venezuela.
Diagnosis. MALE (Fig. 7A): This subspecies is larger
(FW length: 19.5 – 22 mm) than the nominate or S. a.
thiemeiana but marginally smaller than S. a. pratense
ssp.n. and S. a. josenae ssp.n. The FWD discal cell
snow-white patch is generally marginally smaller than
in S. a. josenae ssp.n., rectangular, compact and usu-
ally with sharp edges, unlike the latter subspecies whose
basal edge is often blurry. It never presents any trace of
FWD submarginal white dots, contrary to more southerly
subspecies. FWV is blackish brown with white scaling
restricted to costal and apical area. HWV ground colour
is very dark, nearly black, and densely speckled with
white scales, similarly to S. a. pratense ssp.n. but its ar-
rowhead shaped white patch in M1 – M2 is consistently
shorter. Male genitalia (Fig. 13A): Slightly different
from the nominate subspecies in the wider and stouter
subunci and wider valvae, not narrowing gradually but
tapering at a blunt tip. FEMALE (Fig. 7B): Approxi-
mately the size of the male, very slightly lighter on the
upperside and duller, without the olive sheen typical of
the male; FWV duller with more prominent yellowish
scaling in apical area; HWV with a heavy sandy yellow
overcast, and smaller arrow-head submarginal whitish
dots. Female genitalia (Fig. 17A): As illustrated.
Remarks. This taxon was originally described as a spe-
cies, as was S. a. sanchezi ssp.n. However, it is not mor-
phologically distinct enough to deserve a separate status.
Also, it clearly forms an internal clade of S. albonotata.
(1h) Steromapedaliodes albonotata borealis Pyrcz &
Boyer, ssp.n.
(Figs. 6E – H, 13B, 16A)
Material examined. Holotype, ♂, VENEZUELA: Edo. Trujillo,
Parque Nacional Dinira, Páramo Jabón (South), 3100 – 3150 m,
07.ii.2008, T. Pyrcz leg. (in MZUJ, to be deposited in MIZA);
Para types (57 ♂ and 3 ♀): 4 ♂ and 1 ♀: same data as the Holotype;
31 ♂: Edo. Trujillo, Boconó – Burbusay, Cabimbu de San Miguel,
2900 – 2950 m, 19.ii.2007, T. Pyrcz leg.; 4 ♂: Edo. Trujillo, Páramo
Las Moras, 2950 – 3050 m, 21.ii.2007, T. Pyrcz leg. (1 prep. genit.
13/18.07.2013 J. Lorenc-Brudecka); 2 ♂ and 1 ♀: Edo. Trujillo,
via Páramo Ortiz, Qda. Ortiz, 2900 – 2950 m, 05.ii.2008, T. Pyrcz
P et al.: Taxonomy and phylogeny of Steromapedaliodes
210
leg. (1 prep. genit. 17/18.07.2013 J. Lorenc-Brudecka); 3 ♂: Edo.
Trujillo, Parque Nacional Dinira, Páramo Cendé, 3000 – 3050 m,
13.ii.2010, T. Pyrcz leg., MZUJ; 4 ♂: Edo. Trujillo, Páramo de Or-
tiz, 2900 m, 05.ii.2008, P. Boyer leg.; 2 ♂: Edo. Trujillo, Cabimbù
de San Miguel, nord de Boconó, 3000 m, 19.ii.2007, T. Pyrcz leg.;
5 ♂ and 1 ♀: Edo. Trujillo, Páramo de Jabón, NE de Carache, Mas-
sif du Cendé, 3100, 07.ii.2008, P. Boyer leg., PBF.
Type locality: Páramo de Las Rosas, Parque Nacional
Dinira, Estado Trujillo, Venezuela.
Diagnosis. MALE (Fig. 6E,G): S. albonotata borealis
ssp.n. is smaller (FW length: than the subspecies found
directly southwards, and similar in size to S. a. thiemei-
ana. The FWD discal cell snow-white patch is slightly
smaller than in S. a. schuberti. As with the latter sub-
species, there is no trace of any FWD submarginal white
dots. FWV is blackish brown with rather prominent whit-
ish speckling along costa and in apical area, slightly more
prominent than in S. a. schuberti. HWV ground colour is
blackish brown dusted densely with white scales. Sub-
marginal white dots are less conspicuous than in S. a.
schuberti and in some individuals, especially in the more
northerly populations, barely noticeable, the arrow-head
shaped M1 – M2 spot is generally smaller and shorter than
in S. a. schuberti. Male genitalia (Fig. 13B): differing lit-
tle from the nominate subspecies, but even less so from S.
a. schuberti with which it shares similarly shaped valvae,
not narrowing apically as in other subspecies, and rather
stout subunci. Aedeagus is slender and nearly straight.
FEMALE (Fig. 6F,H): Slightly lighter and duller on both
the upper and underside; sandy overcast on the HWV less
prominent than in other subspecies, in some individuals
not apparent, arrow-head submarginal dots larger and
longer than in the male, especially in the subapical area.
Female genitalia (Fig. 16A): As illustrated.
Etymology. The subspecic name of this taxon refers to
its northerly distribution compared to other geographical
races of S. albonotata.
Remarks. This is the northernmost subspecies of S. al-
bonotata distributed between the Niquitao massif and the
Páramo de Cendé (Fig. 1C). It is found at similar eleva-
tions as other subspecies and occurs in a mosaic of for-
est and paramo and in the lower paramo, occasionally
sympatrically with local populations of other Steromape-
daliodes sensu novo. Its morphology and molecular data
indicate it is most closely relate to S. a. schuberti, an en-
demic subspecies found only in the Guaramacal Massif.
(2a) Steromapedaliodes albarregas albarregas
(Adams & Bernard)
(Figs. 8A,B, 13C, 16C)
Altopedaliodes albarregas Adams & Bernard, 1981: 345, 347 – 348,
gs. 1 (male genitalia), 14, 15.
Altopedaliodes albarregas Adams & Bernard; adamS 1983: 474;
1986: 303; 1987: 38, g. 4.
Pedaliodes albarregas (Adams & Bernard); d’abrera 1988: 864.
Steromapedaliodes albarregas (Adams & Bernard); Viloria &
pyrCz 2001: 9 – 10; lamaS et al. 2004: 215.
Material examined. Holotype, ♂, of Altopedaliodes albarregas
Adams & Bernard, VENEZUELA: Estado Mérida, Cordillera de
Mérida, Río Albarregas, 3150 m, 16.viii.1977, M.J. Adams, AB2;
2 ♂: same data; M.J. Adams & G.I. Bernard, AB2; 1 ♀: same lo-
cality, 3300 m, 14.viii.1977; 5 ♂: same locality, 3100 – 3150 m,
13.viii.1977; 1 ♀: same locality, 3000, 19.vi.1975; 1 ♂: same lo-
cality, 3200, 17.vi.1975; 1 ♂: same locality, 3200, 20.vi.1975;
1 ♂: same locality, 3200 m, 21.vi.1975; 1 ♂: same locality, 3100,
15.viii.1977; 1 ♂: Estado Mérida, Cordillera de Mérida, La Aguada
+ La Montaña, 2500 m, 03.viii.1977 [locality erroneous?], AB2,
Paratypes of A. albarregas A. & B. BMNH; 2 ♂ and 2 ♀: Edo.
Mérida, Páramo de los Conejos, Las Gonzales, Quebrada de la
Boba, 3150 m, 19.ii.2010, T. Pyrcz leg., PBF; 13 ♂ and 1 ♀: Edo.
Mérida, La Culata – Jají, Qda. De La Boba, 3200 – 3250 m, 18 – 19.
ii.2010, T. Pyrcz leg. (5 ♂ prep. genit.: 08/21.09.2010 T. Pyrcz,
20/18.07.2013 J. Lorenc-Brudecka, 04/27.04.2013 J. Lorenc-
Brudecka, 19/18.07.2013 J. Lorenc-Brudecka, 01/18.07.2013 J.
Lorenc-Brudecka) MZUJ.
Type locality: Cordillera de Mérida, upper Río Albarre-
gas valley.
Redescription. MALE (Fig. 8A): Head: glabrous,
chocolate brown with dark brown irregular patches,
covered with dense black hair; collar golden yellow; la-
bial palpi 2 × length of head covered with longer black-
ish brown and shorter golden yellow hair, ventrally three
times longer than dorsally; frons with a tuft of long, grey
and brown hair. Antennae slender, reaching ⅔ the length
of costa, composed of 33 agellomeres, club 2 × width
of shaft, formed gradually, compressed dorso-ventrally,
blackish brown dorsally, golden brown ventrally, most-
ly naked, except for some sparse white scales on basal
segments. Thorax: black, dorsally covered with golden
brown hair, thicker on patagia, ventrally covered with
dense brown, grey and sandy yellow hair; mid and hind
legs brown, femorae covered with brown hair, tibiae and
tarsi densely covered with grey and brown, and sparse
milky white scales, numerous ventral spines on tarsi.
Wings: FW (21 – 22 mm) triangular with a subacute
apex, straight distal margin and tornus at a straight an-
gle. Fringes mostly snow white and black at vein ends.
FWD black brown, except for some sparse white scales
along costal margin and a snow white oblique mid dis-
cal cell band of some 2 mm width throughout, extend-
ing from Sc to just beyond vein CuA2. FWV blackish
brown, glossy, slightly lighter than on the upperside
turning dark chocolate brown distally; white scaling
along costa; snow white oblique mid discal cell shaped
as on the upperside; a straight row of ve, half-moon
shaped, milky white, small submarginal dots from
R4 – R5 to M3 – CuA1, the largest of which in R5 – M1.
HW oval sharply angles at apex and tornus with a slight-
ly undulated distal margin. Fringes white and black at
vein ends. HWD uniform blackish brown, basal ⅔ hairy.
HWV ground colour blackish brown and chocolate
brown with rather sparse dusting of silver scales slightly
denser only in median area where forming a mid-costal
streak and anal wedge; a row of ve arrow-head shaped,
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ARTHROPOD SYSTEMATICS & PHYLOGENY — 75
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milky white submarginal dots, the largest of which in
Rs – M1. Abdomen: dorsally and laterally covered with
black, and basal segments, lustrous navy blue scales and
dense, golden brown hair, ventrally with densely packed
up scales of different colours, brown, grey and sandy
yellow, and sparser hair. Male genitalia (Fig. 13C): Teg-
umen dorsum gently arched; uncus ¾ the length of tegu-
men shoulder, slender, nearly straight with gently down
curved blunt tip; subunci ⅖ the length of uncus, massive
at base, narrowing to a sharp tip, compressed laterally;
pedunculus little developed, blunt; vinculum gently s-
shaped; saccus attened dorso-ventrally, shallow; val-
vae the length of tegumen + half uncus, massive at base,
then gradually narrowing to a blunt tip, with a delicate-
ly irrorated dorsal surface; aedeagus tubular, slightly
arched, not contorted, with the proximal opening half
the entire end, and an acute tip, without any crest or lat-
eral teeth. FEMALE (Fig. 8B): Lighter brown on both
the upper and underside, with more prominent sandy
yellow speckling on the underside, especially on FWV
subapical area and on the HW; arrow-head submarginal
milky white dots more prominent. Female genitalia
(Fig. 16C): Papillae anales membranous in apical half,
attened in lateral view, covered with short, delicate and
rather dense hair, sclerotized in basal half, with devel-
oped, short apohyses posteriores; sternite viii, in lateral
view, as sclerotized as the basal part of papillae anales,
without spiracle; viii fused with tergum viii in lateral
view; postvaginal lamella sclerotized, massive, strongly
sclerotized, entirely covering the antrum; ductus bursae
wide, membranous, the length of bursa, suddenly open-
ing into a rounded bursa copulatrix, without signa; duc-
tus seminalis connecting in the middle of ductus bursae.
Remarks. This species was described based on a series
of individuals found in the uppermost valley of the river
Albarregas in Sierra de La Culata. For many years it was
known only from the type locality until its rediscovery
in the southern extremity of the same La Culata range.
As pointed out correctly by adamS & bernard (1981),
S. al barregas and S. albonotata are separated ecologi-
cally. Even though they may occur locally in the same
area, in the uppermost cloud forest at the paramo bor-
der, S. al bonotata keeps close to the ground level, ying
above marshy meadows, whereas S. albarregas ies in
the canopy. S. albarregas is also a much more energetic
ier, and actively patrols along apparently well-estab-
lished pathways. In the Quebrada la Boba S. albarregas
was found to be quite common in February of 2010, and
it appears that the species is highly seasonal.
(2b) Steromapedaliodes albarregas mauroi Pyrcz,
ssp.n.
(Fig. 8E)
Material examined. Holotype, ♂, VENEZUELA: Edo. Barinas,
Parque Nacional Sierra Nevada, Los Morritos, 3000 m, 06.i.2006,
M. Costa leg. (in MCC, to be deposited in MIZA).
Type locality: Venezuela, Cordillera de Mérida, Los
Morritos.
Diagnosis. MALE (Fig. 8E): This subspecies differs
from the nominate in a similar way as S. albarregas
prietoi ssp.n. that is in the wider, approximately 3 mm
wide FW mid discal cell snow white band, which how-
ever does not enter cell CuA1 + CuA2. As in S. a. prietoi
there is a whitish FW postdiscal streak. There are four
fully developed moon shaped milky white spots from
R4 – R5 to M2 – M3 and two tiny dots in M3 – CuA1 and
CuA1 – CuA2. This subspecies differs from S. albarregas
prietoi ssp.n. most of all in the more prominent ventral
white dusting, on the FWV apical area, and especially
on the HWV where white scales densely cover the entire
wing surface. Also, the arrowhead shaped submarginal
spots are considerably larger than in S. a. prietoi and
milky white, similarly to the nominate subspecies. Male
genitalia: Not examined. FEMALE: Not known.
Etymology. This subspecies is dedicated to its discover-
er, an amateur lepidopterist from Caracas, Mauro Costa,
in recognition for his support to the rst author over sev-
eral years of research visits in Venezuela.
Remarks. This subspecies is known only from the type
locality, the valley of the Río Gavidia, one of the tributar-
ies of Apure, on the eastern slopes of the Sierra Nevada
(Fig. 1D). It is the same locality as for S. albonotata prat-
ense ssp.n., which is locally sympatric, but contrary to S.
albarregas occurs also at higher elevations in the open
paramo as far as the Páramo de Tisure. In morphologi-
cal terms it is intermediate between the nominate found
on the southern slopes of the Sierra de La Culata and S.
albarregas prietoi ssp.n. detected on the north-western
slopes of the range.
(2c) Steromapedaliodes albarregas prietoi Pyrcz,
ssp.n.
(Figs. 8C,D, 13D, 16D)
Material examined. Holotype, ♂, VENEZUELA: Edo. Mérida,
via Piñango, Sector El Potrero, 3200 m, 08.iv.2006, M. Cos-
ta leg. (in MCC to be deposited in MIZA); Paratypes (1 ♂ and
1 ♀): 1 ♂: Edo. Mérida, El Aguila – Piñango, Sector Potrero (Km
26), 2900 – 2950 m, 09.iv.2006, T. Pyrcz leg. (male: prep. genit.
09/21.09.2010 A. Zubek, female: prep. genit. 332/19.01.2016 J.
Lorenc-Brudecka), MZUJ.
Type locality: Sector El Potrero, Piñango Road, NW
slopes of the Sierra de La Culata, Estado Mérida, Vene-
zuela.
Diagnosis. MALE (Fig. 8C): FW length 21 mm. This
subspecies differs immediately from the nominate by
the consistently wider FW oblique snow white band,
some 3 mm wide, not penetrating into CuA2 – 1A/1B, in
one of the examined individuals with an irregular outer
edge, occasionally penetrating deeper into base of cell
P et al.: Taxonomy and phylogeny of Steromapedaliodes
212
CuA1 – CuA2. There is a diffused FW whitish postdiscal
streak extending from Sc to M1 on the upperside, and to
M2 on the underside. FWV submarginal dots are smaller
and not moon shaped, white instead of milky white. Also
on the HWV submarginal arrowhead shaped dots are
white not milky white, and slightly more compact. Male
genitalia (Fig. 13D): Differing marginally from the nom-
inate subspecies, in the slightly shorter tegumen, notice-
ably longer saccus and slightly shorter valvae. FEMALE
(Fig. 8D): FW length: 23 mm. Sexual dimorphism lit-
tle marked, female a shade lighter on the upperside, and
considerably lighter on the HWV with a heavy sandy
yellow suffusion, especially in the postdiscal area; arrow
head white spots slightly lighter, especially the three to-
wards apex. Female genitalia (Fig. 16D): Differing from
the nominate subspecies only in the longer and narrower
ductus bursa, however given that these two are weakly
sclerotized the apparent differences may be an artefact of
dissection and photography technique.
Etymology. This subspecies is dedicated to the Colom-
bian lepidopterist specializing in the taxonomy and ecol-
ogy of Lycaenidae, Carlos Prieto, in recognition for his
numerous valuable contributions to the knowledge of
Andean butteries, and support to the rst author during
eld work in Colombia.
Remarks. S. albarregas prietoi ssp.n. is known so far
only from the type locality. It is morphologically more
related to S. albarregas mauroi ssp.n. from the opposite
slopes of the Cordillera than to the nominate subspecies
found southwards in the Sierra de La Culata.
(3) Steromapedaliodes kahlua Pyrcz & Boyer, sp.n.
(Figs. 8F, 16E)
Material examined. Holotype, ♀, VENEZUELA: Edo. Mérida,
El Baho – via Páramo, La Ciénaga, 2900 – 2950 m, 26.xi.2005,
T. Pyrcz leg. (prep. genit. 03/12.10.2011 J. Lorenc-Brudecka) (in
MZUJ, to be deposited in MIZA).
Type locality: El Baho – via páramo, La Ciénaga, Edo.
Mérida, Venezuela.
Diagnosis. This species is recognized immediately from
S. albarregas and S. albonotata by the fact that the FW
is entirely blackish-brown lacking any white patch in the
FW or HW discal area, it is also smaller than most taxa in
the Steromapedaliodes sensu novo.
Description. FEMALE (Fig. 8F): Head: Eyes glabrous,
lustrous, chestnut brown, covered with long and dense
black hair. Labial palpi 2 × the length of head, covered
with black, grey and sandy yellow hairy scales, ventrally
three times longer than dorsally. Crest composed of grey-
ish hair and basal golden green scales. Antennae slender,
dorsally chestnut, ventrally orange, covered with sparse
black scales, club two time wider than shaft, widening
gradually, attened dorso-ventrally, composed of 10 ag-
ellomeres. Thorax: black, dorsally covered with sparse
black scales on metathorax, and sparse golden brown
hair, somewhat denser on patagia; ventrally also mostly
naked, with some sparse brown, grey and sandy yellow
hair, chiey on legs base; mid and hind legs brown, lus-
trous, femorae covered with rather sparse brown and grey
hair, tibiae and tarsi more densely covered with grey and
sandy yellow hair. Wings: FW (length: 23 mm) roughly
triangular, with a subacute apex, slightly convex distal
margin and gently bent at tornus; fringes short, dark grey,
intermittently dirty white in the interveins. FWD uniform
dark brown. FWV almost uniform brown, pale, lighter
than on the upperside, especially towards outer margin
and apex, with sparse sandy yellow scaling in subapical
area and along costa, and a short sinuate blackish line
in submarginal area extending from costa to vein M2; a
series of ve minute yellowish submarginal dots. HW
rounded with a slightly undulated distal margin; fringes
short except for vein ends, mostly dark grey. HWD uni-
form dark brown, same shade as on the FW. HWV ground
colour brown, with a narrow light brown postbasal band,
and wide blackish brown median band; postdiscal area
light brown suffused with whitish scales especially to-
wards apical area; a sinuate submarginal line dening
a dark brown marginal area; and a row of ve minute
milky white spots, three of which, towards apex, short
arrow-headed, slightly larger than the remaining. Abdo-
men: dorsally and laterally black, covered with black,
and laterally with grey, and ventrally with densely packed
up scales of different colours, brown, sandy yellow and
predominantly grey. Female genitalia (Fig. 16E): Papil-
lae anales membranous in apical half, attened in lateral
view, covered with short, delicate and rather dense hair,
slightly sclerotized in basal half, with developed, short
apophyses posteriores; sternite viii, in lateral view with-
out spiracle, fused with tergum; postvaginal lamella scle-
rotized, massive, strongly sclerotized but smooth, as a
wide pocket entirely covering the antrum; ductus bursae
wide, membranous, half the length of bursa, suddenly
opening into a rounded bursa copulatrix, without signa;
ductus seminalis connecting in the middle of ductus bur-
sae. MALE: Hitherto unknown.
Etymology. This specic epithet is an allusion to the al-
coholic beverage of Caribbean origin, kahlua, which is
characterized by brownish colour, similar to this species
colour patterns, and incidentally is dedicated to author’s
brown furred cat, Kahlua who frequently accompanied
the rst author while setting butteries.
Remarks. This species is described based on a single
female collected in the forest paramo ecotone above El
Baho (Fig. 1D). Despite numerous attempts no further
individuals were obtained. Nonetheless, the collected in-
dividual is attributed with condence a separate specic
status considered its distinctive morphological charac-
ters. In particular, its wing shape, and the ventral colour
pattern is not attributable to either S. albarregas or S. al-
bonotata. The latter species is sympatric with S. kahlua
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ARTHROPOD SYSTEMATICS & PHYLOGENY — 75
(2) 2017
sp.n., the former has a wide geographical distribution in
the Cordillera de Mérida, and is immediately recognized
from other related congeners by the more acute FW apex.
S. kahlua sp.n. is either extremely localized, which can
be the case with some representatives of Steromapedali-
odes, or/and highly seasonal as is S. albarregas.
(4) Steromapedaliodes mavarezi Pyrcz sp.n.
(Figs. 8G,H, 13E, 16F)
Material examined. Holotype, ♂, VENEZUELA: Edo. Barinas,
Páramo de Guirigay, 2800 – 3000 m, 01.xi.2015, J. Mavárez leg.
(prep. genit. 315/11.01.2016 J. Lorenc-Brudecka) (in MZUJ, to
be deposited in MIZA); Paratype, 1 ♀: same data, (prep. genit.
316/11.01.2016 J. Lorenc-Brudecka), MZUJ.
Type locality: Páramo de Guirigay, Cordillera de Mé-
rida, Venezuela.
Diagnosis. Immediately recognized from other conge-
ners by its larger size, expressed in the wider wings, and
the white discal patch on the hindwing dorsum.
Description. MALE (Fig. 8G): Head: Eyes glabrous,
lustrous, chestnut brown with irregular blackish brown
patches, covered with long and dense black hair. Labial
palpi 2 × length of head, covered with black, grey and
sandy yellow hairy scales, ventrally three times longer
than dorsally, and adorned with snow-white scales near
base. Frons with a tuft of long blackish brown and golden
hair. Antennae slender, dorsally blackish brown, ventral-
ly chestnut, naked; club slightly wider than shaft, widen-
ing gradually, attened dorso-ventrally, composed of 11
agellomeres, ventrally orange brown. Thorax: black,
dorsally mostly naked, covered with sparse blackish
brown and navy blue scales on metathorax, and sparse
golden brown hair, somewhat denser on patagia; ven-
trally also mostly naked, with some sparse brown, grey
and sandy yellow hair, chiey on legs base; mid and hind
legs brown, lustrous, femorae covered with rather sparse
brown and grey hair, tibiae and tarsi more densely cov-
ered with grey and sandy yellow hair, numerous ventral
spines on tarsi. Wings: FW (length: 30 mm) roughly
triangular, with a subacute apex, slightly convex distal
margin and gently bent at tornus; fringes short, dark grey,
intermittently snow white towards apex. FWD lustrous,
uniform black. FWV almost uniform blackish brown,
pale, lighter than on the upperside, especially towards
outer margin, with sparse snow white scaling in subapi-
cal area and along costa. HW rounded with a slightly un-
dulated distal margin; fringes short, mostly dark grey and
white. HWD blackish brown, lighter towards anal mar-
gin, with an olive sheen, with some sparse hairy scales
in basal area; a large snow-white patch covering most of
discal cell, except for its basal part. HWV ground colour
blackish brown, a shade darker, wide median band; post-
discal line edged distally with some lighter, dirty white
scales concentrated into a faint band, sharply bent at vein
M1 producing a short mid-costal streak; and a row of
ve minute milky white submarginal arrow-headed dots,
none of which is considerably larger than the remaining;
a shade darker submarginal line. Abdomen: dorsally and
laterally black, covered with black, and laterally with
grey, and ventrally with densely packed up scales of dif-
ferent colours, brown, grey and, predominantly sandy
yellow. Male genitalia (Fig. 13E): Tegumen dorsum
gently arched; uncus the length of tegumen shoulder,
slender, with a gently down curved blunt tip; subunci
absent; pedunculus very short, blunt; vinculum gently
s-shaped; saccus attened dorso-ventrally, shallow; val-
vae the length of tegumen + half uncus, massive at base,
then gradually narrowing to a blunt tip, with a delicately
irrorated dorsal surface in apical ⅓; aedeagus tubular,
slightly arched, not contorted, with the proximal opening
half the entire length, and an acute tip, without any crest
or lateral teeth. FEMALE (Fig. 8H): Sexual dimorphism
slight. Slightly smaller than the male (FW length 26 mm),
and a shade lighter, especially on ventral surface. HWV
duller, with a more prominently marked median darker
brown band, arrow-headed dots slightly larger. Female
genitalia (Fig. 16F): Papillae anales membranous in api-
cal half, attened in lateral view, covered with short, deli-
cate and rather dense hair, slightly sclerotized in basal
half, with developed, short apohyses posteriores; sternite
viii, in lateral view, as sclerotized as the basal part of pa-
pillae anales, without spiracle; viii fused with tergum viii
in lateral view; postvaginal lamella sclerotized, massive,
strongly sclerotized but smooth, as a wide pocket entirely
covering the antrum; ductus bursae wide, membranous,
the length of bursa, suddenly opening into a rounded bur-
sa copulatrix, without signa; ductus seminalis connecting
in the middle of ductus bursae.
Etymology. This species is dedicated to Jesús Mavárez,
eminent Venezuelan biologist, entomologist and geneti-
cist who contributed signicantly in the studies of neo-
tropical butteries and paramo plants.
Comments. This intriguing species is known so far only
from a small area situated in the Páramo de Guirigay SW
of the locality of Las Mesitas (Fig. 1D). It was spotted
while ying low in a subparamo habitat constituted by a
mixture of bunch grasses, rosette plants and cloud forest
pockets. Its external morphology stands apart from any
other species of Steromapedaliodes sensu novo. Its wing
shape and the presence of the whitish patch in the HW
discal area are strongly reminiscent of Cheimas opalinus
(Staudinger, 1897), which occurs in the same habitat, and
these species are possibly involved in a mimicry relation-
ship (Pyrcz in prep.). Male genitalia are very indicative
of its phylogenetic position as they show intermediate
characters of Steromapedaliodes (sensu Viloria & Pyrcz),
and in particular of S. albarregas, namely the shape of
tegumen and uncus, but has atrophied subunci similarly
to the species previously associated with the genus Re-
donda.
P et al.: Taxonomy and phylogeny of Steromapedaliodes
214
(5a) Steromapedaliodes empetrus empetrus Adams &
Bernard, comb.n.
(Figs. 9A – C, 14C, 18B)
Pedaliodes empetrus Thieme, 1905: 95, 98 – 99, pl. 1, g. 7.
Pedaliodes empetrus Thieme; Weymer 1912: 258, pl. 54, row e;
gaede 1931: 489; adamS & bernard 1979: 99; deSCimon
1986: 506, 518.
Punapedaliodes empetrus (Thieme); Huber 1973: 195.
Redonda empetrus empetrus (Thieme): adamS & bernard 1981:
368, gs. 13, 27; adamS 1983: 474; 1984: 93; d’abrera 1988:
871 (misidentied); 2001: 186, 341, pl. 141, g. 17 (misidenti-
ed); Viloria 2000: 269, 270; Viloria & pyrCz 2001: 12, 13;
Viloria et al. 2003: 22 (misidentied); pyrCz 2010a: 180 – 185;
2010a: 265 – 273.
Redonda empetrus (Thieme); Viloria et al. 2015: 112 – 118.
[Redonda empetrus georgei nomen nudum Pyrcz, 2010a: 179 – 185,
244.]
[Redonda empetrus ssp.n.; Pyrcz, 2010b: 265 – 273.]
Material examined. VENEZUELA: 1 ♂: Edo. Mérida, La Culata,
3400 m, 26.vii.2010, J. Wojtusiak leg., MIZA; 2 ♂: Cordillera de
Mérida, N. of Mérida, Río Albarregas, 3400 m, 15.viii.1977, M.J.
Adams & G.I. Bernard, AB2; 1 ♂: same data, 3600 m; 1 ♂: Estado
Mérida, N de Mérida, Río Albarregas, 3400 m, 20.vi.1975, M.J.
Adams & G.I. Bernard; 1 ♂: Edo. Mérida, Tal des Río Albarregas,
2100 m, 29.vii.1971, H. Huber, ZSBS; 1 ♂: Estado Mérida, Parque
Nacional La Culata, Mucujún, 3600 – 3700 m, 02.i.1992, T. Pyrcz
leg., MALUZ; 5 ♂: (1 in MALUZ), same data, 11.iv.1992; 9 ♂:
same data, 09.i.1992; 2 ♂: same data, 29.xii.1991; 3 ♂: same data,
09.ii.1996; 2 ♂: same data, 17.iv.1996, MZUJ; 24 ♂: P. N. La Cu-
lata, Alto Río Mucujún, 3400 – 3450 m, 01.ii.2008, T. Pyrcz leg.,
(prep. genit. 07/18.10.2011 J. Lorenc-Brudecka), TWP; 13 ♂: same
locality but 3200 – 3250 m (incorrect altitude data), 03.iv.2006,
TWP; 9 ♂ and 2 ♀: same data but 3400 – 3450 m, 06.ii.2007, T.
Pyrcz (prep. genit. 02/21.11.2011 J. Lorenc-Brudecka), TWP;
15 ♂: Mérida, La Culata, 3000 – 3200 m, 3.iv.2006, P. Boyer leg.,
PBF; 12 ♂: Mérida, La Culata, 3400 m, 01.ii.2008, P. Boyer leg.,
PBF; 1 ♀: Mérida, La Culata, 3400 m, 21.ii.2008, P. Boyer leg.,
PBF.
Type locality: [“snow mountains of Merida”], Sierra de
la Culata, Cordillera de Mérida, Venezuela.
Redescription. MALE (Fig. 9A,B): Head: Eyes choco-
late brown with some black patches, lustrous, covered
with dense, long, black hair; antennae ⅖ the length of
costa, orange brown, naked except for a few basal seg-
ments covered with white scales, shaft slender, club
spatulate, made of 11 segments, with two lateral, ventral
bruises, blackish brown; labial palpi two 2 × length of
head, covered with sandy yellow, brown and black hair,
ventrally three times as long as dorsally, and with some
snow white dorsal scales; frons with a tuft of long black-
ish hair. Thorax: dorsally black, mostly naked, except for
some sparse velvet bluish-black scales, patagia and base
of prothorax covered with long and dense golden brown
hair, meso and metathorax only with sparse golden brown
hair; ventrally black densely covered with tufts of brown
chestnut hair; mid and hindlegs chestnut, femora and tib-
iae covered with sparse, brown hair, tarsi with grey and
whitish scales. Abdomen: black, lustrous, covered with
brown, russet and sandy yellow hair, dorsally sparse, ven-
trally dense, and additionally laterally with some whitish
scales. Wings: FW (FW length: 23 – 30 mm, mean: 27.2
mm, n = 69) apex subacute, outer margin slightly con-
cave; fringes made of elongated scales, alternately medi-
um brown and sandy yellow in the interveins. HW oval,
outer margin gently undulated; fringes made of elongated
scales, alternately chestnut and sandy yellow in the in-
terveins. FWD covered with hair along inner margin,
otherwise chestnut, liberally dusted with sandy yellow
scales, denser towards distal edge of a faint, irregularly
shaped postdiscal line which divides wing’s surface into
a slightly darker basal and lighter distal half; a row of ve
sandy yellow submarginal rounded spots, from R5 – M1
to Cu1 – Cu2, rather similar in size, occasionally a sixth,
small spot in R3 – R4; in some individual a faint lilac or
pale white, roughly rectangular patch in outer ⅓ of discal
cell. HWD covered with long chestnut hair in basal and
postbasal area, and along inner margin; blackish in basal
area, otherwise chestnut, liberally dusted with sandy yel-
low scales, a row of ve sandy yellow, oval or subtrian-
gular submarginal spots, in Rs – M1 to Cu1 – Cu2. FWV
predominantly sandy yellow although quite variable and
turning orange brown in some individuals, darker basal-
ly from postdiscal line where more heavier dusted with
brown scales; a row of submarginal sandy yellow dots,
noticeably smaller than on dorsum; subapical and apical
area dusted with somewhat more prominent sandy yellow
and milky white scales. HWV dark brown, sandy yel-
low and milky white, with a wide darker band between
median and postdiscal lines, and extending from costal
to inner margin; a row of ve submarginal milky white
triangular patches, from Rs – M1 to Cu1 – Cu2, pointing
basally, somewhat variable in size but invariably more
prominent than on dorsum. Male genitalia (Fig. 14C):
Very similar to the S. empetrus frailejona, and differing
from other subspecies in the wide, blunt apex of the val-
vae. FEMALE (Fig. 9C): Much smaller than the male
(FW length 16 mm, n = 3) with slightly deformed wings,
especially the folded FW outer half (antennae reaching
half the length of costa); FWD and HWD almost uni-
form lustrous golden yellow, except for the barely visible
row of tiny, whitish FWD subapical dots; outer margin
brown; FWV and HWV colour pattern similar to that of
the male. Female genitalia (Fig. 18B): Papillae anales
gently rounded in lateral view, covered with long, rather
sparse hair (some of them in ventral half transformed into
thicker spines); lamella postvaginalis wide, with a single
shallow concavity; lamella antevaginalis with a single,
little prominent, blunt protrusion; ductus bursae short,
approximately ⅓ the length of bursa copulatrix, wide, tu-
bular, without any sclerotized structures, gently opening
into bursa copulatrix, with ductus seminalis originating
at its middle; bursa copulatrix oval, without any visible
signa.
Comments. The identication of the nominotypical sub-
species of S. empetrus is crucial for the stability of the
nomenclature in the light of the existence of several local
populations of this species associated with separate sub-
species. Viloria et al. (2015) dedicated a lot of attention
to this issue. They concluded that the types of Thieme
215
ARTHROPOD SYSTEMATICS & PHYLOGENY — 75
(2) 2017
were collected in the La Culata range, and designated it
as nominotypical Redonda empetrus. We are inclined to
agree, tentatively with their decision although the ulti-
mate recognition of the exact origin of the Thieme taxon
is not rendered easy by the fact that the type specimen
is missing, as is most of the Satyrinae type collection
of Thieme (for details see Viloria et al. 2015), and that
the original illustration in tHieme (1905), a fairly good
black and white photograph, shows the upperside of a
male individual, whereas the original descriptive text is
not accurate enough as to sort out denitely between the
subspecies. Individual variation among the individuals of
the La Culata population is admittedly impressive and
affects the size, upperside and underside colour patterns.
In our opinion, arguably the most important feature indi-
cated by tHieme (1905) which would point out to the La
Culata population is the presence of a, usually faint, sil-
very or silvery patch in the FWD discal cell, not apparent
in the Mucubají, Los Frailes or Loma Redonda popula-
tion individuals, which otherwise correspond closely to
the upperside of the specimen illustrated by Thieme. It is
not impossible that further sampling in the Cordillera de
Mérida, and more precisely in the La Culata range, which
has been very supercially and only locally sampled for
paramo butteries will reveal a population which even
more closely matches the type of Thieme than the speci-
mens examined from the upper valleys of Río Albarregas
(collected by Adams & Bernard) or Río Mucujún. The
non-designation of a neotype of Pedaliodes empetrus by
Viloria et al. (2015) was therefore a sound decision. In-
terestingly, adamS & bernard (1981) did not recognise
the populations occurring in La Culata and Sierra Nevada
as representing a separate subspecies and misidentied it
with S. empetrus bolivari, which is found across the Cha-
ma. The two differ in size, upper and underside colours,
although admittedly individual variation of S. empetrus
empetrus is impressive.
Steromapedaliodes empetrus empetrus is found in the
Sierra de La Culata, it has been evidenced from the upper
valley of Río Albarregas in the south, to Pico El Aguila in
the north (Fig. 1B). S. empetrus empetrus is known from
several localities, however the best investigated, and also
the most accessible is the one inhabiting the upper Río
Mucujún valley just above the large moraine dominating
the landscape of the La Culata valley. Contrary to S. em-
petrus bolivari, numerous individuals can be observed on
the wing every year being most abundant in January and
February. It could indicate that locally this subspecies
takes only one year to achieve complete development.
Although rather common, the Mucujún population never
attains the level of abundance of S. empetrus frailejona
in Los Frailes. Also, contrary to the latter subspecies it
is rather widespread in the La Culata range and is not
as sedentary. Wandering individuals of S. empetrus em-
petrus have been observed by TP far from boggy para-
mos in inhospitable high altitude desert in the Páramo de
Piedras Blancas at 4300 m. Also, a single individual was
seen by TP and PB north of the Pico El Aguila at 4200 m
in a paramo dominated by tall Coespeletia timotensis in
an area that was previously visited several times without
noticing the presence of Steromapedaliodes. Studies of
the Mucujun daily activity pattern were made by ferrer-
pariS (2000). The rst known female was collected by
K. Casner. Another female was found by TP only minutes
later, and the third a year later in the very same spot as the
previous two. All the females were observed in the early
morning at around 9.00. They were receptive, sunning on
the leaves of Espeletia with wings wide open exposing
a shining golden yellow upperside. An attempt of mat-
ing was observed, and whilst one male was engaged in
copulation with the female, another male approached and
stood up to the competition. Our observations strongly
indicate that females of brachypterous S. empetrus are
active in the sunny mornings. Despite dozens of visits to
the site, no other female was observed later in the day.
(5b) Steromapedaliodes empetrus bolivari (Adams &
Bernard, 1981), comb.n., stat. reinst.
(Figs. 10G,H, 14B)
Redonda empetrus bolivari Adams & Bernard, 1981: 368 – 369, g.
28.
Redonda empetrus bolivari Adams & Bernard; Viloria 2000: 269,
270; pyrCz 2010a: 179 – 185, 244; 2010b: 265 – 273.
Redonda empetrus Adams & Bernard; d’abrera 1988: 871 (er-
roneous photograph caption, as Diaphanos huberi).
Redonda bolivari Adams & Bernard; Viloria et al. 2015: 106.
Material examined. Holotype, ♂, VENEZUELA: Estado Mérida,
S of Mérida, Loma; Redonda, 4000 m, 21.iv.1975, M.J. Adams
leg., BMNH; 15 ♂: Estado Mérida, P. N. Sierra Nevada, below
Loma Redonda, 3900 – 3950 m, 03.ii.2007, T. Pyrcz leg., (prep.
genit. 04/18.10.2011 J. Lorenc-Brudecka), TWP; 12 ♂: same data
but 08.ii.2007, TWP; 1 ♂, Loma Redonda, 3900 m, 08.ii.2007, Es-
tado Mérida, Venezuela, T. Pyrcz leg., MCC.
Type locality: Loma Redonda, South of Mérida, Cordil-
lera de Mérida, Venezuela.
Redescription. MALE (Fig. 10G,H): Differs from other
subspecies in the larger size (FW length: 26 – 30 mm,
mean: 28.12 mm, n = 24), although some individuals of
the nominate are equal in size to S. empetrus bolivari;
FWD and HWD colour pattern is little contrasting com-
pared to other subspecies; ground colour has a golden
sheen; submarginal dots are tiny and faint. FWV ground
colour is fairly variable but has a prominent golden or
golden brown sheen, and is quite uniform, except for the
darker basal half, submarginal spots are small and faint.
HWV is also rather uniform, predominantly grey brown
with a darker basal half and some whitish scales in distal
half; submarginal dots are small, milky white triangles.
Male genitalia (Fig. 14B): Differs from the nominate in
the slightly shorter valvae with a more acute apex, and
from both the nominate and S. chiquinquirana by the ¼
longer aedeagus. FEMALE: So far unknown.
Comments. This subspecies is distributed on the north-
western slopes of the Sierra Nevada (Fig. 1B). Currently
P et al.: Taxonomy and phylogeny of Steromapedaliodes
216
known only from the type locality below Loma Redonda
where it occurs at 3800 – 4000 m and the trail from La
Mucuy to the Humboldt and Bompland glaciers. Its en-
tire distribution is unknown and possibly extends over
large portions of the Sierra Nevada including the south-
eastern slopes, which are mostly inaccessible and have
not been sampled for butteries so far. Males are larger
than other subspecies of S. empetrus except particularly
large individuals of S. empetrus empetrus. They are very
active iers and behave in a way very reminiscent of S.
bordoni. The female is still unknown and we suspect that
it is brachypterous. S. empetrus bolivari ies at higher
elevations than other subspecies, except some reports of
S. empetrus empetrus.
(5c) Steromapedaliodes empetrus frailejona (Ferrer-
Paris & Costa), comb.n., stat.n.
(Figs. 9D,E, 14A, 18A)
Redonda empetrus empetrus (Thieme); pyrCz 2010a: 180 – 185,
2010a: 265 – 273 (misidentied).
Redonda frailejona Ferrer-Paris & Costa (in Viloria et al. 2015):
118, 130, gs. 33 – 37 (adults), 55 (male genitalia).
Material examined. VENEZUELA: 7 ♂: Mérida, Apartaderos –
Sto. Domingo, km 12.5, ca. 3100 – 3300 m, 26.ix.1997, A. Neild,
AFN; 19 males: Edo. Mérida, oberes Domingo tal [upper valley
of “River Santo” Domingo], 3070 m, 05.ix.1971, H. Huber; 2 ♂:
same data, 3050 m, ZSBS; 17 ♂: Edo. Mérida, Mucubají – Santo
Domingo, Los Frailes, 3000 – 3050 m, 08.vii.2006, T. Pyrcz leg.
(prep. genit. 04/27.10.2014 J. Lorenc-Brudecka); 44 ♂: same data
but 19.iv.2006; 3 ♂: same data but 12.iv.2004; 1 ♂: same data but
02.ii.2008; 2 ♂: same data but 31.vii.2009; 1 ♂: same data but
24.ii.2010, MZUJ; 39 ♂: Mérida, Los Frailes, Santo Domingo vers
Apartaderos km 10, 2900 – 3000 m, 19.iv.2006, P. Boyer leg., PBF;
4 ♂: same data but 8.iv.2006, P. Boyer leg., PBF; 1 ♂: Mérida, Los
Frailes, Santo Domingo vers Apartaderos, 2900 m, 31.vii.2009,
P. Boyer leg., PBF; 1 ♂: Mérida, Apartaderos – Santo Domingo
km 12.5, 3100 – 3200, 26.vii.1997, A. Neild leg., PBF; 1 ♀: Edo.
Mérida, Los Frailes, 3050 m, 12.iv.2004 [collecting year probably
erroneous, 2014?], M. Costa leg., (prep. genit. 05/04.10.2012, J.
Lorenc – Brudecka), MCC.
Type locality: Santo Domingo – Apartaderos, Estado
Mérida, Cordillera de Mérida, Venezuela.
Redescription. MALE (Fig. 9D): Differs from the nomi-
nate and S. empetrus bolivari in the smaller size (length:
24 – 28 mm, 24.72 mm, n = 69); otherwise wings dorsal
colour patterns are closely similar to the nominate sub-
species, except that postdiscal yellow dots are usualy
larger, and on the HW rarther triangular than oval, a ea-
ture which more obvious o the HWV; light milky white
or lilac FWD discal patches sometimes apparent in the
nominate subspecies are absent or nearly so; FWV is
darker than in the nominate, orange brown instead of
sandy yellow, but usually lighter than in S. empetrus bo-
livari. Male genitalia (Fig. 14A): Uncus stout, aligned
with and approximately the length of dorsum of tegu-
men; subunci as very small tips adhered to base of un-
cus; pedunculus medium sized; saccus wide and shallow;
valvae the length of tegumen + uncus, massive, gradually
narrowing towards a gently rounded apex, ventrally with
a noticeable subapical notch, dorsally smooth; aedeagus
the length of saccus + valve, slender and gently arched,
terminated with a small bulbous membrane with tiny cor-
nuti. FEMALE (Fig. 9E): The only known specimen is
much smaller than the male (FW length: 12 mm) with nar-
row, deformed wings (antennae reaching half the length
of costa); FWD and HWD almost uniform lustrous silver
yellow, except brown costa and the barely visible row of
tiny, whitish spearhead triangular subapical dots on the
HW; outer margin brown; FWV and HWV colour pattern
similar to that of the male except fo the disproportionally
large spearhead submarginal white spots. Female geni-
talia (Fig. 18A): Papillae anales gently rounded in lateral
view, covered with long, rather sparse hair; lamella post-
vaginalis wide, with a single shallow concavity; lamella
antevaginalis with a single, little prominent, blunt protru-
sion; ductus bursae short, approximately ⅓ the length of
bursa copulatrix, tubular, without any sclerotized struc-
tures, gradually widening and gently opening into bursa
copulatrix, with ductus seminalis originating at its mid-
dle; bursa copulatrix oval, without any visible signa.
Comments. It is worth pointing out that S. empetrus
frailejona is known so far only from a very small area
concentrated around a bog located above the main Mu-
cubají – Santo Domingo road at 2850 – 3100 m (Fig.
1B). Despite extensive sampling carried out by TP and
co-workers in apparently appropriate habitats at slightly
higher elevations towards the Mucubají Pass or above the
village of El Baho at different seasons of the year, no
other population of S. empetrus was detected in the Santo
Domingo valley. It is highly alarming and suggests that
the Los Frailes bog may be effectively the only extant
locality of this subspecies. S. empetrus frailejona occurs
locally at high density at the peak of its ight season.
Simultaneously up to 10 adults can be observed uttering
around above Espeletia spp. inorescences. Occasionally
individuals are seen crossing the asphalt road, overy-
ing Blechnum ferns thickets, and entering the gardens
and yards of the “Los Frailes” hotel facilities. Observa-
tions carried out by the rst author during consecutive
years from 2003 – 2010 in the Santo Domingo valley in-
dicate that S. empetrus frailejona larvae take two years
to achieve their development. In 2006 and 2008 high
ight period took place in February – March whereas
next year’s March only sporadic individuals were ob-
served. It appears that the ight season of S. empetrus
frailejona corresponds with the late blooming period of
the local Espeletia schultzii (Asteraceae), the source of
nectar for adults, which falls at the end of the rainy sea-
son and through most of the dry season from November
to March.
(5d) Steromapedaliodes empetrus ssp.
Material examined. VENEZUELA, 1 ♂: Est. Mérida, Campanario,
13500 ft, 10.ix.1938, J. H. Tracey, BM 1939 – 114, BMNH; 1 ♂:
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ARTHROPOD SYSTEMATICS & PHYLOGENY — 75
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Estado Mérida, P. N. Sierra Nevada, Laguna Verde Laguna Suero,
4000 m, 12.ii.1985, CEUM leg., MIZA; 1 ♂: Venezuela, Ch.
Blanchier, MHNG.
Comments. Three specimens of S. empetrus could not be
attributed to any subspecies. They are left without sug-
gesting any taxonomical status for one, collected in the
Laguna Suero, is badly damaged, while the other has no
exact collecting locality data. Nevertheless, they present
some morphological characters that let us suppose they
could represent yet another undescribed subspecies of
Steromapedaliodes empetrus. This needs to be corrobo-
rated by more sampling.
(6a) Steromapedaliodes chiquinquirana (Ferrer-Paris),
comb.n.
(Figs. 9F,G, 14D, 18C)
[Redonda empetrus empetrus Adams & Bernard; Viloria et al.
2003: 22 (female misidentied).]
[Redonda empetrus mikei nomen nudum Pyrcz, 2010a: 179 – 185,
244.]
[Redonda empetrus ssp.n.; Pyrcz, 2010b: 265 – 273.]
Redonda chiquinquirana Ferrer-Paris (in Viloria et al. 2015): 112,
127 – 130, gs. 24 – 26 (adults), 51 (male genitalia).
Material examined. VENEZUELA: 1 ♂: Estado Mérida, Parque
Nacional Sierra Nevada, Laguna Negra, 3550 m, 28.iii.1992,
T. Pyrcz; MZUJ; 1 ♂ and 1 ♀: Estado Mérida, Parque Nacional
Sierra Nevada, vía Laguna Negra, 3600 m, 20.iv.1992, T. Pyrcz
(male: prep. genit. 02/18.10.2011 J. Lorenc-Brudecka; female:
prep. genit. 04/07.11.2011 J. Lorenc-Brudecka), MALUZ; 8 ♂: Es-
tado Mérida, Parque Nacional Sierra Nevada, Laguna de Mucubají,
3600 m, 05.x.1975, M. Rański; 1 ♂: same data, 01.ix.1988, R.
Manrique; 9 ♂ and 2 ♀: same data, 20.iv.1992, T. Pyrcz, MZUJ;
1 ♂: Edo. Mérida, zw. Laguna Grande und Laguna Negra, 3450 m,
12.ix.1971, H. Huber, ZSBS; 1 ♀: no data [M.J. Adams & G.I. Ber-
nard, AB1], BMNH; 1 ♂: Mérida, PN Sierra Nevada, Páramo de
Mucubají, 3500 – 3600 m, 5.x.1975, J. Rański, PBF; 1 ♂: Mérida,
Mucubají, vía a la Laguna Negra, 3400 m, 19.iv.2006, P. Boyer,
PBF.
Type locality: Páramo de Mucubají, Serranía de Santo
Domingo, Cordillera de Mérida, Venezuela.
Redescription. MALE (Fig. 9F): Head: Eyes chocolate
brown and black, lustrous, covered with dense, long,
brown and whitish hair; antennae ⅖ the length of costa,
orange brown, naked except for a few basal segments
covered with white scales, shaft slender, club spatulate,
made of 11 segments, with two lateral, ventral bruises; la-
bial palpi 2 × length of head, covered with sandy yellow
and brown hair, ventrally three times as long as dorsally,
and laterally with white; frons with a tuft of long sandy
yellow hair. Thorax: dorsally black, base of prothorax
covered with long and dense chestnut and sandy yellow
hair, meso and metathorax only with sparse brown hair;
ventrally black densely covered with tufts of brown and
chestnut hair; mid and hindlegs femora covered with
sandy yellow hair, tibiae and tarsi with brown and silvery
scales. Abdomen: dorsally and laterally covered with
black scales, ventrally with silvery scales and sandy yel-
low hair. Wings: FW length: 23 – 26 mm, mean: 24.8 mm,
n = 19; FWD ground colour varies between chestnut and
olive brown, submarginal dots are generally richer yellow
and fainter. HWD ground colour is similarly variable as
on the FW, submarginal dots, although quite variable are
generally elongated, instead of rounded, and richer yel-
low. FWV ground colour is richer yellow with a brown or
golden shade, submarginal dots are fainter. HWV is con-
spicuously lighter due the presence of prominent sandy
yellow scaling; submarginal dots are considerably elon-
gated, and this is the most immediate diagnostic feature,
which allows the identication of this compared to other
subspecies. Male genitalia (Fig. 14D): Very similar to
S. empetrus except for the slightly more acute apex of the
valvae. FEMALE (Fig. 9G): Much smaller than the male
(FW length 18.5 – 19 mm); FW and HWD light chestnut
with a strong silvery overcast, in some individuals nearly
all shinning silvery; FWD median half and apical area
generally slightly darker with more prominent chestnut
pattern; minute submarginal whitish barely noticeable
on HWD, slightly more prominent on FWD; FWV and
HWV similar to male, with characteristic HV submar-
ginal elongated milky white streaks, even longer than
in the male particularly in Rs – M1 and M1 – M2; silver
scaling on HW less prominent than in the male, in some
specimens nearly absent. Female genitalia (Fig. 18C):
Papillae anales gently rounded in lateral view, covered
with long hair (some of them in ventral half transformed
into thicker spines); lamella postvaginalis wide, with two
shallow concavities; lamella antevaginalis with a single,
little prominent, blunt protrusion; ductus bursae short,
approximately ⅓ the length of bursa copulatrix, wide, tu-
bular, without any sclerotized structures, gently opening
into bursa copulatrix, with ductus seminalis originating
at its middle; bursa copulatrix oval, without any visible
signa.
Remarks. S. chiquinquirana chiquinquirana differs from
all the subspecies of S. chiquinquirana by the slightly
smaller size; FWD submarginal dots are generally richer
yellow and fainter; HWD submarginal dots, although
quite variable, are generally elongated, instead of round-
ed, and richer yellow; FWV is richer yellow with a brown
or golden shade, submarginal dots are fainter; HWV is
conspicuously lighter due the presence of prominent
sandy yellow scaling; submarginal dots are considerably
elongated, and this is the most immediate diagnostic fea-
ture which allows the identication of this compared to
other subspecies. adamS & bernard (1981) thought that
the population found in the Mucubají area represents the
nominate S. empetrus and failed to recognize it as new.
There are seemingly two generations per year, the rst
emerging in January and the second in October. Males
are strong iers, perhaps as energetic as the other species.
Females, which are much smaller than males as recorded
previously by adamS & bernard (1981), are “very loath
to y even when disturbed, and after very short ights
disappeared among low tussocks of grass”.
P et al.: Taxonomy and phylogeny of Steromapedaliodes
218
(6b) Steromapedaliodes chiquinquirana lossadana
(Ferrer-Paris), comb.n., stat.n.
(Figs. 9H – J, 14E,F, 18E,F)
[Redonda empetrus decennia nomen nudum Pyrcz, 2010a: 179 –
185, 244.]
[Redonda empetrus ssp.n.; pyrCz, 2010b: 265 – 273.]
Redonda lossadana Ferrer-Paris (in Viloria et al. 2015): 119,
141 – 144, gs. 44, 45 (adults), 58 (male genitalia), 141 – 144.
Redonda centenaria Viloria & Camacho (in Viloria et al. 2015):
112, 123 – 127, gs. 6 (female wing venation), 20 – 23 (adults),
50 (male genitalia), syn.n.
Material examined. VENEZUELA: 1 ♂: Edo. Mérida, Las Mesi-
tas, Páramo Guirigay, 3100 – 3150 m, 27.vii.2009, T. Pyrcz leg.,
MZUJ; 15 ♂ and 3 ♀: Edo. Mérida, Las Mesitas, Páramo Guirigay,
3100 – 3150 m, 27.vii.2009, T. Pyrcz, TWP; 29 ♂ and 1 ♀: Edo. Mé-
rida, Tuñame, Pueblo Llano crossroad, 3200 – 3250 m, 25.xi.2005,
T. Pyrcz, TWP (male: prep. genit. 01/27.10.2014 J. Lorenc-Bru-
decka; female: prep. genit. 06/18.10.2011 J. Lorenc-Brudecka;
7 ♂: Edo. Mérida Las Mesitas, vía Teta de Niquitao, 3300 – 3400
m, 27.vii.2009, T. Pyrcz, TWP; 8 ♂ and 1 ♀: Edo. Trujillo, Páramo
de Guirigay, 3000 – 3100 m, SE de Las Mesitas, 3000 – 3100 m,
27.vii.2009, P. Boyer, PBF; 12 ♂: Edo. Trujillo, Jonction Niquitao/
pueblo Llano × Niquitao/Tuñame vers Pueblo Llano km5, 3150 m,
23.XI.2005, P. Boyer, PBF; 16 ♂: Edo. Trujillo, Jonction Niquitao/
pueblo Llano × Niquitao/Tuñame vers Pueblo Llano km7, 3300 m,
25.XI.2005, P. Boyer, PBF; 20 ♂ and 1 ♀: Edo. Trujillo, Jonction
Niquitao/pueblo Llano × Niquitao/Tuñame, 3150 m, 25.XI.2005, P.
Boyer, PBF; 3 ♂ and 1 ♀: Edo. Trujillo, Las Mesitas vers Teta de
Niquitao, 3200 – 3300 m, NW de las Mesitas, 27.vii.2009, P. Boyer,
PBF.
Type locality: Páramo de Tuñame, Estado Trujillo, Cor-
dillera de Mérida, Venezuela.
Redescription. MALE (Fig. 9H): Differs from the nomi-
nate subspecies in the considerably darker, chocolate
brown FW and HWD; in a faint and diffused reddish
brown postmedian band; smaller submarginal dots on FW
and particularly on HW, where barely noticeable. FWV
is suffused with reddish brown, a diagnostic character of
this subspecies. HWV is blackish brown, darker than in
the nominate and more contrasting due to the presence of
white scales in basal, postbasal and postdiscal areas; sub-
marginal triangular dots are larger than in S. empetrus but
shorter than in the nominate S. chiquinquirana. Average
size is similar to the nominate (FW length: 24 – 29 mm,
mean: 26.2 mm, n = 52). Male genitalia (Fig. 14E,F):
Differs from the nominate and other subspecies in the
slender valvae. FEMALE (Fig. 9I,J): Considerably
smaller than the male (FW length: 21.5 – 22.5 mm, mean:
22 mm, n = 3); FW and HWD light beige with a silvery
sheen; FWD with a slightly darker brown median band
and submarginal line; a row of 5 submarginal subapical
yellow FW dots and HW spearhead shaped. FWV and
HWD similar to the male but generally more contrast-
ing due to prominent yellow dusting of FW, instead of
reddish – yellow of the male, and heavier whitish speck-
ling of HW. Female genitalia (Fig. 18E,F): very similar
to nominate S. chiquinquirana chiquinquirana. Papillae
anales gently rounded in lateral view, covered with long
hair (some of them in ventral half transformed into thick-
er spines) and with short, strongly sclerotized apophyse
posteriores; lamella postvaginalis wide, with two shal-
low concavities; lamella antevaginalis with a single, lit-
tle prominent, blunt protrusion; ductus bursae short, ap-
proximately ⅓ the length of bursa copulatrix, and wide,
slightly sclerotized, with ductus seminalis originating at
its middle; bursa copulatrix large, oval, no signa.
Comments. This subspecies occurs in the Tuñame me-
seta area, on the southeastern slopes of Teta de Niquitao,
and Guirigay massif (Fig. 1B). Habitats vary between
dry, rocky slopes dominated by rosetal paramo commu-
nities of Espeletia tenorae, and bogs. At the peak of the
ying season, this subspecies can be quite common and
at least 5 – 10 individuals can be observed on the wing
simultaneously. Monitoring is needed and the establish-
ment of small but protected areas should be considered.
Other species found in the bogs include Diaphanos
curvignathos Viloria, 1994 and some endemic Espeletii-
nae (Viloria 1994). Redonda centenaria described from
the Teta de Niquitao (Viloria & CamaCHo 2015) is con-
sidered here as the junior synonym of S. chiquinquirana
lossadana. The types and the individuals collected by the
authors of this article in the type locality of centenaria do
not differ from S. chiquinquirana lossadana.
(6c) Steromapedaliodes chiquinquirana castellana
(Viloria & Camacho), comb.n., stat.n.
(Figs. 11A – F, 15A, 18D, 21B)
[Redonda empetrus ssp.n. 1 ; Viloria, 2000: 270; Viloria & pyrCz
2001: 12; Viloria et al. 2003: 22; pyrCz 2010b: 265 – 273.]
[Redonda empetrus centenaria nomen nudum Pyrcz, 2010a: 36,
179 – 185, 244.]
Redonda castellana Viloria & Camacho (in Viloria et al. 2015):
112, 119 – 123, gs. 16 – 19 (adults), 49 (male genitalia).
Material examined. VENEZUELA: 1 ♂: Estado Trujillo, Ortíz,
Páramo Ortíz, 3100 – 3150 m, 05.ii.2008, T. Pyrcz (prep. genit.
03/27.10.2014 J. Lorenc-Brudecka), MZUJ; 1 ♂: Estado Trujillo,
Páramo de Cabimbú, 2900 m; 11.ix.1991, A. Viloria & R. Calchi,
MALUZ; 9 ♂ and 2 ♀: same data as holotype, MALUZ; 15 ♂ and
4 ♀: Estado Trujillo, Páramo de Ortíz, 2850 – 3100 m; 12.ix.1991,
A. Viloria & E. Moscó leg., (2 ♂ and 1 ♀ in TWP); 30 ♂ and 4 ♀:
same locality, 3100; 07.i.1992, J. Camacho & A. Viloria, MALUZ
(1 ♂ in BMNH); 3 ♂: Estado Trujillo, Vía Boconó-Páramo La
Cristalina, 2400 – 2800 m, 14.vi.1986, J.E. Lattke, MIZA; 1 ♂:
Estado Trujillo, Páramo La Cristalina, 2800 m, 14.v.1986, J.E.
Lattke, MIZA; 54 ♂ and 3 ♀: Estado Trujillo, Boconó – Trujillo,
Páramo Las Moras, 2950 – 3050 m, 21.ii.2007, T. Pyrcz (prep. gen-
it. 02/07.11.2011 J. Lorenc-Brudecka), TWP; 10 ♂ and 2 ♀: Tru-
jillo, Páramo de Ortíz, 3100, 5.ii.2008, P. Boyer, PBF; 3 ♂: Trujillo,
Route Ortiz vers Boconó, Páramo las Moras, 3000 m, 21.ii.2007, T.
Pyrcz, PBF; 2 ♀: Páramo Las Moras, 3000 m, 20.ii.2007, M. Costa,
MCC; 1 ♀: Páramo de Ortíz, 3170 m, 18.viii.2003, A. Viloria and
M. Alaracón, MCC.
Type locality: Páramo de Ortíz, Estado Trujillo, Ven-
ezuela.
Redescription. MALE (Fig. 11A,C,E): FW (length:
25 – 31 mm; n = 57; mean: 27.38 mm). Differs from the
nominate and S. chiquinquirana lossadana in more elon-
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ARTHROPOD SYSTEMATICS & PHYLOGENY — 75
(2) 2017
gated wings, and somewhat more acute FW apex; also
differs from the nominate in the darker brown FW and
HWD, but most examined individuals are a shade lighter
than S. chiquinquirana lossadana; also compared to S.
chiquinquirana lossadana, dorsal colour pattern is more
lustrous, and uniform, apart from a shade darker median
band, and has no reddish brown postmedian suffusion;
FW and HWD submarginal dots are as small and faint
as in S. chiquinquirana lossadana. FWV ground colour
is quite variable, but in most individuals dark chestnut,
much darker than in the nominate, and considerably
duller and less contrasting than in S. chiquinquirana los-
sadana, without the diagnostic reddish brown overcast
covering the whole wing, instead distal part of the wing
is lighter, sandy yellow occasionally with some orange
shade; submarginal dots are small and faint, similar as
in S. chiquinquirana lossadana. HWV ground colour is
grey brown, dull and generally little contrasting; postdis-
cal darker line is always noticeable, in most individuals
edged distally with some sandy yellow scaling; submar-
ginal sandy yellow dots are the same size and shape as in
S. chiquinquirana lossadana, and generally larger than
in the nominate. Male genitalia (Fig. 15A): Differs from
the nominate in the slightly shorter valvae terminated
with an acute apex, similarly to S. chiquinquirana los-
sadana whose valvae are however slender and longer.
FEMALE (Fig. 11B,D,F): Considerably smaller than
the male (FW length: 21 – 26 mm; n = 12; mean = 22.50
mm), greatly variable, some being larger and more pig-
mented than others; in melanic specimens differences
with S. chiquinquirana chiquinquirana are exaggerated,
FWD ground colour ochraceous brown, with some ar-
eas creamy-yellow (for example, distal extremity of
discal cell, and edges of discal region); postdiscal dots
appear more discrete and small, in contrast to darkened
groundcolour; HWD similar to typical subspecies but
densely dusted with ochraceous brown, more thickly at
base and margins; HWV both groundcolour and speck-
ling darker; speckling much more dense (as in males);
FW submarginal dots reduced, and some bluish-white
scales on apex; hindwing with much thicker and more
distinct dark-coffee speckling, which is heavier in basal
half; white speckling in general denser than in typical
subspecies; postdiscal arrow-head shaped white spots
less elongated. Female genitalia (Fig. 18D): Very simi-
lar to the preceding subspecies. Papillae anales gently
rounded in lateral view, covered with long hair (some
of them in ventral half transformed into thicker spines);
lamella postvaginalis wide, with two shallow concavi-
ties; lamella antevaginalis with a single, little prominent,
blunt protrusion; ductus bursae short, approximately ⅓
the length of bursa copulatrix, and wide, little sclerotized
gently widening and opening into bursa copulatrix, with
ductus seminalis originating at its middle; bursa copula-
trix large, pear-like, without any visible signa.
Comments. This subspecies is restricted to the paramos
of the Niquitao region in Trujillo State. It comprises the
localities of La Cristalina, Cabimbú, and northern slopes
of La Teta de Niquitao (4006 m) (Fig. 1B). Most of these
paramos, particularly those located in the Río Boconó
basin, have a single long wet season (ca. April – August),
which is probably due to the inuence of the climatic
conditions of the Llanos, south of the Andes (monaS-
terio & reyeS 1980). Within its area of distribution, S.
chiquinquirana castellana is an abundant insect through-
out the year. Both males and females y actively dur-
ing sunny periods, but females can be distinguished by
their smaller size, paler upperside coloration, and relative
weakness in ight. Usually they seem unable to sustain
a long ight, which is undertaken in “steps”, and very
close to the grasses, most commonly following the direc-
tion of the wind. Resting females are very difcult to see
because of their cryptic underside pattern, but they can be
observed dropping suddenly into the grass after a short
ight, then they take another ight-step in the same way.
When collected, most females released many spherical
eggs from the moment of capture and throughout several
hours whilst kept alive in paper envelopes. Two females
placed together in the same envelope produced 167 oliva-
ceous green eggs in a few hours; a further four specimens
produced 113 ova (some of them pale green to yellowish)
and died with many eggs still within the abdomen. All the
eggs were also free of cement.
The paramos in the Niquitao region differ from the
areas inhabited by other subspecies in the heavy presence
of Chusquea which constitutes a co-dominant element of
oreal plant communities alongside Espeletia rosettes,
Puya bromeliads and bunch grasses (VareSCHi 1970; Cu-
atreCaSaS 1979). A few Swallenochloa clumps are found
patchily distributed only in the Páramo de Ortiz, mainly
along creeks. This buttery co-exists with Diaphanos
curvignathos Viloria and Steromapedaliodes albonotata
borealis, Colias dimera Doubleday, 1847, Catasticta
chry so lopha spectrum Reissinger, 1972, Hylephila sp.,
Rhamma sp., Podanotum andrewneildi Bálint, 2001 and
Rhamma commodus (C. Felder & R. Felder, 1865).
(7) Steromapedaliodes leukasmena (Viloria &
Camacho), comb.n.
(Figs. 11G,H, 15B, 19C)
[Redonda sp.n. 2; Viloria 2000: 270.]
[Redonda sp.n.; Viloria & pyrCz 2001: 15; Viloria et al. 2003:
22].
[Redonda camachoi nomen nudum; Pyrcz, 2010a: 38, 45, 180 – 185,
244; pyrCz 2010b: 265 – 273.]
Redonda leukasmena Viloria & Camacho (in Viloria et al. 2015):
119, 138 – 141, gs. 7 (female wing venation), 40 – 43 (adults),
57 (male genitalia).
Material examined. VENEZUELA: 1 ♂: Estados Trujillo – Lara,
P.N. Dinira, Páramo Jabón (south slopes), 3100 – 3150 m, 07.ii.
2008, T. Pyrcz; 1 ♂: Estado Lara, Municipio Morán, Páramo de La
Rosa [sic], 3000 m, 20.viii.1991, J. Camacho leg., MALUZ; 6 ♂
and 2 ♀: Estado Lara, Parque Nacional Dinira, Páramo La Rosa
[sic], 2800; 16.i.1994; A. Viloria & J. Camacho; 18 ♂ (1 in BMNH,
1 in MZUJ) and 2 ♀: Estado Lara, Parque Nacional Dinira, Páramo
Cendé, 3100 m; 08.viii.1995, M García, MALUZ; 1 ♂: Estado
Trujillo, P. N. Dinira, Pmo. Cende, 3100 – 3150 m, 07.ii.2008, T.
P et al.: Taxonomy and phylogeny of Steromapedaliodes
220
Pyrcz (male: prep. genit. 06/18.10.2011 J. Lorenc-Brudecka); 2 ♂:
Estados Lara – Trujillo, Páramo El Jabón, 3000 m, 25.xii.1989,
CEUM, MIZA; 10 ♂ and 2 ♀: Estados Trujillo – Lara border, P.N.
Dinira, Páramo Jabón (south slopes), 3100 – 3150 m, 07.ii.2008,
T. Pyrcz (female: prep. genit. 01/07.11.2011 J. Lorenc-Brudecka);
2 ♂: same locality but 3000 – 3050 m, 13.ii.2010, MZUJ; 8 ♂: Tru-
jillo, Páramo de Jabón, NE de Carache, massif du Cendé, 3100 m,
07.ii.2008, P. Boyer, PBF.
Type locality: Páramo de La Rosa, Municipio Morán,
Estado Lara, Cordillera de Mérida, Venezuela.
Redescription. MALE (Fig. 11G): Head: Eyes choco-
late brown with black patches, lustrous, covered with
dense, long, black hair; antennae 2/5 the length of costa,
orange brown, naked except for a few basal segments
covered with white scales, shaft slender, club 2 × as
wide, elongated, composed of 10 segments, with a deep
ventral cavity; labial palpi 2 × length of head, covered
with sparse sandy yellow and brown hair, ventrally three
times as long as dorsally, frons with a tuft of long black-
ish brown hair. Thorax: dorsally black, covered with
black scales and russet hair; ventrally black densely
covered with tufts of brown and russet hair; mid and
hindlegs dark brown, with dense dark brown spines and
silver scales somewhat dense on tarsi and tibiae. Abdo-
men: black, covered with brown scales sandy yellow
hair on ventral side. Wings: FW (length: 27.5 – 30.5
mm, mean: 28.8 mm, n = 13) apex subacute, outer mar-
gin slightly concave, tornus obtuse; fringes alternately
dark brown brown and white in the interveins. HW oval,
outer margin regular; fringes dark brown. FWD slightly
hairy at base, lustrous dark brown, slightly darker in
distal half of discal cell, with a large, polygonal subdis-
cal silvery white patch within discal cell, distal edge of
which is straight and well dened at ⅔ of cell, basal edge
rather diffuse, following vein Cu2, and a row of faint,
sometimes not apparent, ve sandy yellow submarginal
rounded dots, from R5 – M1 to Cu1 – Cu2, basal ⅓ with
greenish sheen. HWD covered with long chestnut hair in
basal and postbasal area, and along inner margin; black-
ish in basal area, otherwise almost uniform lustrous dark
brown. FWV chocolate brown, duller and becoming
coffee brown towards costal margin and apex, lustrous
and with a delicate chestnut sheen in basal half; a row
of faint, submarginal sandy yellow dots; subapical, api-
cal and marginal area speckled with dark coffee brown.
HWV chocolate brown speckled with white scales some-
what denser along distal edge of median and postdiscal
line, and black, notably dense on basal half of wing and
in submarginal region of each cell; a row of ve sub-
marginal milky white roughly triangular patches, from
Rs – M1 to Cu1 – Cu2, pointing basally, all of them ir-
regularly surrounded by dark brown; a faint submarginal
darker blackish line. Male genitalia (Fig. 15B): Simi-
lar to S. chiquinquirana except for the considerably, ⅓
longer uncus, completely atrophied subunci and the
longer and slender valvae, which resembles mostly that
of S. chiquinquirana lossadana, but sharing with all the
subspecies of S. chiquinquirana the characteristic ven-
tral notch; saccus wider than in S. chiquinquirana with
singularly folded upwards sides. FEMALE (Fig. 11H):
Smaller than the male (FW length: 25 – 28.5 mm; mean:
26.8 mm; n = 5), with narrower wings, more pointed
apex and tornus of FW; FW and HWD surface less
dark and brownish, forewing showing postdiscal series
of 4 – 5 ochraceous and minute dots, in cells R5 to Cu1
(sometimes missing in M2). FWV groundcolour surface
brown, light brown at apex; submarginal series of ve
white dots in cells R5 to Cu1; HWV groundcolour not
chocolate, but pale brown, which gives higher colour
contrast in general design. Female genitalia (Fig. 19C):
Papillae anales gently rounded in lateral view, covered
with long, thin hair; lamella postvaginalis wide, with
two shallow concavities; lamella antevaginalis with a
single, little prominent, blunt protrusion; ductus bursae
very short, approximately 1/5 the length of bursa copula-
trix, strongly sclerotized laterally and compressed in the
middle, then opening into a wide, tubular second section
which opens into bursa copulatrix, with ductus seminalis
originating at the compression of ductus bursae; bursa
copulatrix large, rounded, without any visible signa.
Comments. This species differs from other species of
Steromapedaliodes, and most particularly those hitherto
associated with Redonda, in the dark, nearly black dorsal
wing surface and a snow white FWD discal cell patch.
Such a dorsal colour pattern is strongly reminiscent of
Steromapedaliodes albonotata, however the much larger
wingspan and rounded wings immediately allow its as-
sociation with S. chiquinquirana and S. empetrus.
It occurs in high and humid paramos of the Cendé
Region, north-easternmost extreme of the Venezuelan
Andes, between the States of Trujillo and Lara (Fig. 1B).
This geographic unit includes the highlands or paramos
of Agua de Obispos, Las Rosas, Jabón, Guache, Naríz,
Los Nepes, Cendé, and El Vigía. monaSterio (1980) has
dened the main vegetation there as “rosetal de Ruilo-
pezia jabonensis” (Asteraceae) because of the remark-
able dominance of this endemic plant. But S. leukasmena
is better associated with the equally common “Swalle-
nochloa – Ruilopezia jabonensis grassland” (Poaceae-
Asteraceae). These paramos are highly inuenced by
the atmospheric conditions of the Lake Maracaibo ba-
sin, which results in two highly pronounced wet seasons
(monaSterio & reyeS 1980).
This species has never been found abundant. The
behaviour is similar in males and females, and as the
wing patterns are almost identical, it is almost impos-
sible to recognize the sex of ying individuals. They y
energetically, 1 – 2 m above the thick bambusoid grasses
(mainly Swallenochloa angustifolia and Rhipidocladum
geminatum (Poaceae), which grow in the Cendé paramos
(Viloria 1994; judzieWiCz & riina 2005). We failed to
nd S. leukasmena in the dry, low Páramo de Los Nepes
(2400 – 2850 m), where bamboos seem to be partially re-
placed by tussock grasses such as Agrostis and Calama-
grostis (Poaceae). The rst female collected laid one
egg immediately upon capture. It was spherical and pale
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ARTHROPOD SYSTEMATICS & PHYLOGENY — 75
(2) 2017
green, with no evidence of any adhesive on its surface.
Like most butteries found at very high elevations in the
northern Andes, S. leukasmena only ies in sunshine and
is very hard to nd in either cloudy or foggy conditions.
Then, the adults tend to hide in the bamboo grasses, pre-
ferring the most inaccessible basal stems, where they
are also thought to spend the night. Interestingly, adults
are active early in the morning on sunny days, at 9 – 11
AM, afterwards their activity nearly stops. This differs
from S. chiquinquirana castellana, which can be seen
on the wing even in the afternoon at 2 – 3 PM. Nothing
is known about its biology. However, from observations
of the biotopes occupied by the adults, we infer that the
larvae feed on bamboos throughout the páramo. Diapha-
nos fuscus Viloria, 1994 and Steromapedaliodes albono-
tata borealis are two more satyrines of the paramo of
the Cendé region (Viloria 1994). Steromapedaliodes
albonotata borealis shows a similar wing pattern and be-
haviour, which may suggest a possible mimetic relation-
ship with Steromapedaliodes leukasmena, but they have
not been found ying together (see comments in Viloria
& pyrCz 2001).
(8) Steromapedaliodes bordoni (Viloria & Pyrcz),
comb.n.
(Figs. 10C,D, 15C, 18A, 21A)
[Redonda sp.n. 1; Viloria 2000: 269.]
Redonda bordoni Viloria & Pyrcz, 2003: 21 – 23 (in Viloria et al.
2003).
Redonda bordoni Viloria & Pyrcz; pyrCz 2010a: 36, 45, 179 – 185,
244; 2010b; pyrCz 2010b: 265 – 273; Viloria et al. 2015:
110 – 112, gs. 5 (female wing venation), 14 – 15 (adults), 48
(male genitalia).
Material examined. Holotype, ♂, VENEZUELA: Estado Táchira,
Páramo El Batallón, entre Laguna El Cenegón y Laguna Grande,
3200 – 3400 m; 28.ii.1994, A. Viloria, M. García & J. Camacho,
MIZA; 27 ♂ and 2 ♀: same data (3 ♂ in MHN, 3 ♂ JFLC); 19 ♂:
Estado Táchira, Páramo El Batallón, entre la Antena CANTV y
la Laguna El Cenegón, 3100 – 3250 m, 27.ii.1994, A. Viloria, M.
García & J. Camacho, (2 in TWP, 2 in BMNH); 4 ♂: Estado Táchira,
Municipio Jáuregui, Callejón del Cenegón, 16.xii.1994; M. García;
6 ♂ and 1 ♀: Estado Táchira, Parque Nacional Juan Peñaloza,
Páramo El Rosal, 3000 m, 12 – 14.i.1995, J. Camacho & M. García;
10 ♂: Estado Táchira, Páramo El Batallón, entre El Cenegón y La-
guna Grande, 3300 – 3400 m, 05.iii.1996, J. Camacho, M. García,
T. Pyrcz and J. Wojtusiak, MALUZ; 1 ♂: Estado Táchira, Páramo
de La Negra, 30.ix.1951, P. Fenjues; 1 ♂: same locality, 3200 m,
14.i.1982, C. Bordón, MIZA; 51 ♂ and 1 ♀: Estado Táchira, Pára-
mo El Batallón, vía El Púlpito, 3500 – 3800 m, 02 – 04.iii.1996, T.
Pyrcz, J. Wojtusiak, J. Camacho, M. García (female: prep. genit.
05/07.11.2011 J. Lorenc-Brudecka); 5 ♂: Estado Táchira, Páramo
El Batallón, vía El Cenegón, 04.iii.1996, T. Pyrcz, J. Wojtusiak,
J. Camacho and M. García, MZUJ (all Paratypes); 4 ♂: Estado
Táchira, Páramo El Batallón, Páramo El Rosal, 3300 – 3350 m,
02.xii.2005, T. Pyrcz; 22 ♂: Estado Táchira, Páramo El Batallón,
Antenas – Cenegón, 2900 – 2950 m, 01.ii.2008, T. Pyrcz, MZUJ;
1♂: same data but 3300 m, 19.ii.2008, (prep. genit. 01/07.10.2014 J.
Lorenc-Brudecka); 20 ♂ and 1 ♀: Tachira, Antenas, vía a Cenegón,
Páramo el Batallon, 3200 – 3300 m, 19.ii.2008, P. Boyer, PBF; 15 ♂:
Tachira, Páramo el Rosal, La Grita vers San Jose de Bolivar km 22,
3150 m, 01.xii.2005, P. Boyer, PBF; 1 ♂: Tachira, vía El Pulpito,
3500 – 3800 m, 03.iii.1996, T. Pyrcz, PBF.
Type locality: Between Laguna El Cenegón and Lagu-
na Grande, 3200 – 3400 m, Páramo El Batallón, Estado
Táchira, Venezuela.
Redescription. MALE (Fig. 10C): Head: Eyes choco-
late brown, lustrous, covered with dense, long, black
hair; antennae 2/5 the length of costa, orange brown, na-
ked except for a few basal segments covered with white
scales, shaft slender, club spatulate, made of 11 seg-
ments, with two lateral, ventral bruises; labial palpi 2 ×
length of head, covered with sandy yellow and blackish
hair, ventrally three times as long as dorsally, and with
whitish scales laterally; frons with a tuft of long orange
and chocolate brown hair. Thorax: dorsally black, base of
prothorax covered with long and dense russet hair, meso
and metathorax only with sparse brown hair; ventrally
black densely covered with tufts of brown chestnut hair;
mid and hindleg femora brown densely covered with
sandy yellow hair, tibiae brown with a gorget of yellow
hair and sparse whitish hair, tarsi medium brown with
dense spines. Abdomen: black, lustrous, dorsally cov-
ered with black scales, laterally with sandy yellow and
ventrally with whitish and brown scales and yellowish
hair. Wings: FW (length: 25 – 32.5 mm, mean: 29.05 mm)
apex subacute, outer margin slightly concave; fringes
chestnut and whitish in the interveins. HW oval, smooth;
fringes mostly whitish intermixed with some chestnut
scales. FWD lustrous, chestnut, with fairly variable pat-
tern of darker grey brown, in some specimens covering
most of basal half and forming a zigzagging median line,
in others faint and barely noticeable; a row of ve sandy
yellow submarginal dots, from R5 – M1 to Cu1 – Cu2,
apparent in all the specimens. HWD covered with long
chestnut hair in basal and postbasal area, and along in-
ner margin; chestnut, lustrous, less patterned than FWV,
in most specimens median half slightly darker, grey
brown; submarginal yellow streak visible in most speci-
mens in spaces Cu1 – Cu2 and Cu2 – 1A, rarely in other
spaces. FWV varying between light chestnut and sandy
yellow, with a postmedian zigzagging grey brown line,
and some darker dusting on the apex; a series of yel-
low submarginal dots as on the upperside. HWV ground
colour chestnut with greyish sheen; dark brown areas
forming well dened intravenal stripes in most individu-
als merging into a zigzagging postdiscal line; a series of
ve submarginal milky white streaks, from Rs – M1 to
Cu1 – Cu2, somewhat variable in size but extending over
at least 3 – 6 mm. Male genitalia (Fig. 15C): Very simi-
lar to S. chiquinquirana chiquinquirana in most sclerites,
uncus, aedeagus and saccus and especially in the massive
valvae terminated bluntly, also closely resembling nomi-
nate S. empetrus, differing only in the subunci, which are
small but fully developed not atrophied. FEMALE (Fig.
10D): Brachypterous and ightless, much smaller than
the male. FW (length 21 – 22 mm, mean: 21.5 mm, n = 5)
and HW elongated, FW apex acute. FWD almost uni-
form lustrous silver, except for the row of tiny, whitish
subapical elongated dots, and some brown basal, costal
and apical dusting; HWD silver with some faint slightly
P et al.: Taxonomy and phylogeny of Steromapedaliodes
222
darker greyish areas in discal cell, along anal and outer
margin; FWV and HWV colour pattern similar to that
of the male but ground colour lighter, whitish. Female
genitalia (Fig. 19A): Papillae anales gently rounded in
lateral view, covered with long, thin hair; lamella post-
vaginalis wide, with two shallow concavities; lamella
antevaginalis with a single, little prominent, blunt pro-
trusion; ductus bursae short, approximately ⅓ the length
of bursa copulatrix, narrow, tubular, little sclerotized, ap-
proximately the same width throughout its length, with
ductus seminalis originating in its middle; bursa copula-
trix large, balloon-like, without any visible signa.
Comments. This species is restricted to the open paramo
of the La Negra – El Batallón range (southwestern Cor-
dillera de Mérida), which is now protected as a national
park (Parque Nacional Juan Peñaloza) (Fig. 1B). Stero-
mapedaliodes bordoni only inhabits open areas above the
treeline, from 3000 m upwards (the highest altitude in the
region is the Pico El Púlpito, about 3950 m). We found
males very abundant during the dry season in February
and March, but even at that time this paramo was ap-
parently more windy, humid and cloudy than the neigh-
bouring ones of the Cordillera de Mérida. Fast-ying,
very active males exploited short periods of sunshine
around midday to travel medium distances across the
paramo, ying erratically over grasses. They drifted with
the wind, but were also able to y strongly against it. In
that case (as in other paramo species, which usually y
in windswept places) the butteries avoided attack (for
instance, from collectors) by stalling in ight, thus allow-
ing the wind to change their ight vector very suddenly.
During a very foggy morning with occasional rain and
strong wind some males were found resting in the centre
of rosettes of Ruilopezia sp. (Asteraceae), in which they
were difcult to disturb.
Females, with reduced and deformed wings, do not
y under natural conditions and spend most of their time
resting on grasses, where they are effectively protected
from predators by highly cryptic underside wing pattern
(Viloria et al. 2003). All females were collected directly
from the ground and bunch grasses by hand, two of them
were observed walking and jumping on the ground, and
after exposure to strong sunshine for ve or six minutes
(for observation), one made a very weak ‘ight’ of about
3 m, probably the limit of its capabilities. An explanation
for this very unusual behaviour was presented by Viloria
et al. (2003) and pyrCz (2010b). One female violently ex-
pelled ve mature eggs when captured, a reaction already
recorded for other species of Steromapedaliodes and two
species of the genus Diaphanos, which probably resulted
from the stress caused by handling (Viloria 1994). As
the eggs are always free of any adhesive, it is obvious
that these butteries do not oviposit on a particular sub-
strate but scatter the eggs over the continuous grassland.
An unusual behaviour was observed by Mauro Costa in
December 2007. Males of S. bordoni were seen when
overying a large puddle, and from time to time throw-
ing themselves against the water, possibly engaging in
interactions with their reection on the water surface. S.
bordoni was never seen mud puddling or drinking water
from the ground.
(9) Steromapedaliodes lathraia (Viloria & Camacho),
comb.n.
(Figs. 10E,F, 15D, 19B)
[Redonda casneri nomen nudum; pyrCz 2010a: 36, 180 – 185]
Redonda lathraia Viloria & Camacho (in Viloria et al. 2015): 119,
135 – 138, gs. 38 – 39 (adults), 56 (male genitalia).
Material examined. VENEZUELA: 1 ♂, Estados Mérida/Táchira
border, El Batallón, Mesa Alta, 2700 – 2800 m, 03.xii.2005, T. Pyrcz,
(prep. genit. 02/07.10.2014 J. Lorenc-Brudecka); 32 ♂ and 1 ♀: Es-
tados Mérida/Táchira border, El Batallón, Mesa Alta, 2700 – 2800
m, 03.XII.2005, T. Pyrcz, (female: prep. genit. 03/07.11.2011 J.
Lorenc-Brudecka) TWP; 3 ♂: same data but 10.ii.2007, TWP; 17 ♂
and 2 ♀: Mérida, Mesa Alta, Bailadores vers Pregonero km 33,5,
5,5 km après jonction de la Grita, 02.XII.2005, P. Boyer leg., PBF.
Type locality: Entre Laguna El Cenegón and Laguna
Grande, Páramo El Batallón, Estado Táchira, Cordillera
de Mérida, Venezuela.
Redescription. MALE (Fig. 10E): Head: Eyes chocolate
brown, lustrous, covered with dense, long, black hair; an-
tennae 2/5 the length of costa, orange brown, naked ex-
cept for a few basal segments covered with white scales,
shaft slender, club spatulate, composed of 11 segments,
with two lateral, ventral bruises; labial palpi 2 × length
of head, covered with sparse sandy yellow and brown
hair, ventrally three times as long as dorsally, frons with
a tuft of long chocolate brown and chestnut hair. Thorax:
dorsally black, base of prothorax covered with long and
dense russet hair, meso and metathorax only with sparse
brown hair; ventrally black densely covered with tufts of
brown chestnut hair; mid and hindlegs chestnut, covered
with grey brown and silver scales, denser on tarsi. Ab-
domen: black, lustrous, covered with grey brown scales,
somewhat lighter ventrally. Wings: FW (length: 23 – 26.5
mm, mean: 24.83 mm, n = 28) apex subacute, outer mar-
gin slightly concave; fringes chestnut except for some
sandy yellow scales in the interveins. HW oval, outer
margin gently undulated; fringes chestnut and sandy yel-
low in apical area. FWD lustrous, chestnut with a greyish
sheen, a row of ve sandy yellow submarginal minute
dots, from R5 – M1 to Cu1 – Cu2, in some specimens not
apparent. HWD covered with long chestnut hair in ba-
sal and postbasal area, and along inner margin; uniform
chestnut, lustrous. FWV chestnut, slightly darker brown
along costa and along outer margin; a row of ve small
submarginal sandy yellow elongated dots. HWV ground
colour chestnut with darker brown areas forming diffused
intravenal stripes and a series of lunular spots along outer
margin; a series of ve submarginal milky white patches,
from Rs – M1 to Cu1 – Cu2, pointing basally, somewhat
variable in size but invariably forming elongated spear-
heads pointing basally extending over 2 – 5 mm. Male
genitalia (Fig. 15D): Most closely resembling S. bordoni
223
ARTHROPOD SYSTEMATICS & PHYLOGENY — 75
(2) 2017
in the shape of massive valvae terminated with a blunt
apex, and S. leuksamena in the completely atrophied
subunci fused with ventral surface of base of uncus, and
the longer uncus than in any subspecies of S. empetrus
or S. bordoni. FEMALE (Fig. 10F): Sexual dimorphism
inconsiderable except for the slightly smaller size of the
female (FW length 23 mm, n = 3), somewhat lighter up-
perside, with a characteristic FW and HW golden sheen
and particularly lighter underside due to a yellowish suf-
fusion of FW and the presence of dense, ne milky white
dusting especially in outer ⅓. Female genitalia (Fig.
19B): Papillae anales gently rounded in lateral view,
covered with long, thin hair; lamella postvaginalis wide,
with two shallow concavities; lamella antevaginalis with
a single, little prominent, blunt protrusion; ductus bursae
short, approximately ⅓ the length of bursa copulatrix,
narrow, tubular, with a prominently corrugated and more
noticeably sclerotized distal half, gently opening into
bursa copulatrix, with ductus seminalis originating in its
middle; bursa copulatrix small, roughly oval, without any
visible signa.
Comments. This species differs from S. bordoni in the
smaller size, more elongated FW, and the much darker,
chestnut with a greyish sheen FW and HWD. In this re-
spect it resembles closely S. chiquinquirana lossadana
from which it however differs in the much elongated
HWV submarginal milky white markings and the more
acute FW apex.
Steromapedaliodes lathraia occurs in boggy paramo
just above timberline overgrown with dwarf Chusquea,
which is possibly its host plant, whereas S. bordoni inhab-
its high paramo grasslands devoid of Chusquea. Margin-
ally however, for example along the Antenas – Cienagas
trail, at some 3100 – 3200 m the two species are syntopic.
This fact mislead previous authors who included several
specimens of S. bordoni in the paratype series of this spe-
cies, which they tentatively identied as an individual
“grey” form of S. bordoni.
4. Discussion: evolution and syste-
matics of Steromapedaliodes
sensu novo
Our decision on considering Redonda as a subjective
junior synonym of Steromapedaliodes sensu novo is sup-
ported by molecular and morphological evidence. Mo-
lecular analyses clearly show that although the species
associated previously with Redonda form a clade, this
group and Steromapedaliodes (sensu Viloria & Pyrcz)
never appear to be reciprocally monophyletic in our phy-
logenetic analyses. Indeed, Redonda is always nested
within the genus Steromapedaliodes (sensu Viloria &
Pyrcz), which makes the latter paraphyletic. Also, our
point of view is strengthened by the discovery of a new
species, S. mavarezi, whose genital characters appear in-
termediate between Steromapedaliodes (sensu Viloria &
Pyrcz) and Redonda, but being placed phylogenetically
in a clade with S. albarregas and Redonda. From a cla-
distic perspective, the retention of Redonda as a sepa-
rate entity would require the inclusion of S. mavarezi
and S. albarregas within Redonda, and the restriction of
the other species to Steromapedaliodes. However, such
scheme would be in strong contradiction with the mor-
phological results shown above. We opted for a simpler
scheme, with only one supra-specic category and with-
out negative effects on the catalogue of biodiversity as
measured by the number of species.
The decision to consider a single genus, Steromape-
daliodes sensu novo, is strongly supported by two results
from the morphological analyses. On the one hand by the
absence of any salient synapomorphies in adult morphol-
ogy of the two genera previously considered (ex Stero-
mapedaliodes (sensu Viloria & Pyrcz) and ex Redonda),
and on the other by the existence of synapomorphic
characters of the genus Steromapedaliodes sensu novo
in male and female genitalia and colour patterns. This
agrees with the view that the comparative analysis of
genital structures provides valuable data, and can be cru-
cial in assessing phyletic afnities of taxa of butteries
in general (turner et al. 1961; lorkovič 1993; freitaS
& broWn 2004), and those belonging to the subtribe
Pronophilina in particular (Viloria 2000; pyrCz 2008;
pyrCz & Viloria 2008; pyrCz et al. 2010). In particular,
the pre sence of the singular arrowhead HWV submar-
ginal dots, unique among all the species of the speciose
Pedaliodes complex, is considered here as a strong syn-
apomorphy. Our results provide further support to the
idea that colour patterns, particularly on the hindwing
venter, are good characters to consider in the phyloge-
netic reconstruction of relationships among butteries
(pyrCz et al. 2008).
Our results indicate that putative ecological differ-
ences between ex Steromapedaliodes (sensu Viloria &
Pyrcz) and ex Redonda pointed out in some previous pa-
pers do not stand (pyrCz 2010a; Viloria et al. 2015). In
fact, although all the species previously associated with
ex Redonda are paramo dwellers, this is by no means an
exclusive feature, as several subspecies of S. albonotata
are also found above timberline, living in association
with the high-altitude tropical grassland characteristic
of the paramo habitat. On the other hand, the habitat of
S. albarregas is conned to dense uppermost forests,
where it ies above canopy, and it does not occur in open
grassland or even in forest-paramo patches, as do some
populations of S. albonotata or S. mavarezi. We believe
that S. albarregas could have retained several plesiomor-
phic and less specialised characters than other congeners,
which can be related to its preferred habitat, typical of
most Pronophilina and particularly of most Pedaliodes.
Accordingly, S. albarregas females are only marginally
smaller than the males and are vigorous iers.
It seems however plausible that some ecological pro-
cesses are at play in the divergence and further mainte-
nance of species differences in Steromapedaliodes sensu
P et al.: Taxonomy and phylogeny of Steromapedaliodes
224
novo. Most species show rather restricted geographic dis-
tributions with almost no overlap with other taxa, indicat-
ing that mechanisms of allopatric or parapatric speciation
must have played a role during the initial phases of di-
vergence in the genus. Thus, S. leukasmena is distributed
allopatrically in the north (Cendé massif), S. empetrus,
S. chiquinquirana and S. albarregas, are distributed
parapatrically in the main central ranges of the Cordil-
lera de Mérida (Culata and Sierra Nevada massifs), and
S. bordoni and S. lathraia, are distributed sympatrically
but rarely syntopically in the southern paramos (Batallón
and La Negra). Very little is known about the geographic
distributions of S. mavarezi sp.n. (Guirigay massif) and
S. kahlua sp.n. (Santo Domingo range), but both seem
to be distributed parapatrically in regards to S. chiquin-
quirana and S. albonotata, respectively. There are only
two exceptions to this pattern, the sympatry between S.
bordoni and S. lathraia mentioned above, and S. albono-
tata, a species with a very large distribution covering al-
most entirely the Cordillera de Mérida and that has been
found in sympatry with S. albarregas and S. chiquin-
quirana in the main central ranges and with S. leukas-
mena in the north (Cendé massif), but never in syntopy.
Indeed, as mentioned above, S. albonotata is generally a
ground-level ying species in the forest-paramo ecotone
that seems to be ecologically separated from other locally
sympatric species by microhabitat use: above the forest
canopy at a 4 – 5 m ight-height (S. albarregas) and true
paramo (S. chiquinquirana and S. leukasmena). The only
known case of syntopy in Steromapedaliodes sensu novo,
although rather marginal, is therefore found in S. bordoni
and S. lathraia. Yet, again, these are two ecologically di-
vergent species that also differ greatly in almost any other
respect, including male genitalia and male wing colours,
but most notably on female wing shape (S. bordoni are
brachypterous, S. lathraia are normal-shaped and good
yers). No hybrid individual between S. bordoni and S.
lathraia has ever been found, which provides support for
the idea of a presumably strong isolation between these
two species. Unfortunately, nothing is known about the
reproductive biology of any species of Steromapedali-
odes sensu novo so the relative importance of pre-zygotic
mechanisms (i.e. ecological isolation) and post-zygotic
mechanisms (i.e. hybrid sterility or inviability) for spe-
cies isolation cannot be stated at this time.
In any case, the ensemble of genetic, ecologic and
geographic data suggest that the divergence within Stero-
mapedaliodes sensu novo in the Cordillera de Mérida oc-
curred rst parapatrically along a habitat axis, i.e. eln
forest forest canopy (S. albarregas) – forest-paramo eco-
tone (S. mavarezi, S. albonotata, S. lathraia, and perhaps
S. kahlua) – grassland paramo (S. bordoni, S. chiquin-
quirana, S. empetrus and S. leukasmena), and then most-
ly allopatrically within similar habitat zones separated
geographically. These ndings provide additional sup-
port for what is emerging as a general model of buttery
diversication in the high-altitude habitats of the Andes.
For example, in the genus Lymanopoda (Nymphalidae:
Satyrinae: Pronophilina) the earlier divergences of lin-
eages are associated with large changes in altitudinal
distribution while recent speciation events appear to be
linked to geographic shifts within the same elevational
band (adamS 1985; CaSner & pyrCz 2010). Similar evo-
lutionary patterns were considered for the genera Theope
Doubleday, 1847, Hypanartia Hübner, 1921 (Nymphali-
dae: Nymphalinae), and Eois Hübner, 1818 (Geometri-
dae: Larentiinae) (Hall & Willmott 1996; Willmott et
al. 2001; Strutzenberger & fiedler 2011). It is interest-
ing to note that a similar scenario might also be at work
in other taxa, such as the emblematic paramo endemic
plants of the subfamily Espeletiinae (Asteraceae), in
which divergence has produced genera specically asso-
ciated with habitat types / altitudinal limits, i.e. Carram-
boa and Libanothamnus in the forest-paramo ecotone,
Espeletia and Espeletiopsis in the paramo proper and
Coespeletia in the desertic superparamo (CuatreCaSaS
2013).
The approach towards the species-level taxonomy of
ex Redonda presented by Viloria et al. (2015) reects a
“splitter” perspective, as all the subspecies of the poly-
typic S. chiquinquirana and S. empetrus are treated as
specically distinct. Such a position is not supported
by our analysis of morphologic and genetic data. Puta-
tive morphological differences separating the various
taxa, particularly in male genitalia, are weak and merely
quantitative, referring to somewhat larger or longer scle-
rits such as uncus, subunci or valvae. Our comparative
studies of genital organs show that individual variation in
male genitalia is important, and there is extensive overlap
in most characters, especially among the subspecies of
S. chiquinquirana. Also, female genitalia were not stud-
ied by Viloria et al. (2015). There is on the other hand
evidence based on eld observations suggesting that male
individuals of S. chinquinquirana and S. empetrus are
particularly active and have been seen several kilometres
away from the nearest established populations, in inhos-
pitable superparamo habitat such as in the Pico El Aguila
or in the Páramo de Piedras Blancas at about 4300 m. We
propose that S. chiquinquirana, S. empetrus, S. albonotata
and S. albarregas are four species with signicant poly-
typic variation. The remaining taxa, S. bor doni, S. kahlua
sp.n., S. lathraia, S. leukasmena and S. ma varezi sp.n.,
are sufciently distinctive from the genetic, morphologic
and biogeographic standpoints so that their specic status
is supported. Indeed, they all represent divergent phylo-
genetic lineages, show non-overlapping morphologies in
regards to variation in wing size, shape, colours, and at
a certain degree on genital traits, and generally exhibit
disjunct geographic distributions.
Our study emphasizes the convenience and impor-
tance of analyses that combine different sources of in-
formation, particularly morphological and molecular,
while assessing phylogenetic relationships. Morphologi-
cal data, especially male and female genitalia, originally
suggested the presence of as a highly polytypic species
(S. empetrus, 6 ssp.) in the central part of the Cordillera
de Mérida. However, molecular data favour the identi-
cation of two different and somehow distantly related
225
ARTHROPOD SYSTEMATICS & PHYLOGENY — 75
(2) 2017
species (S. empetrus, 3 ssp., and S. chiquinquirana,
3 ssp.). Their distribution is locally adjacent in the Santo
Domingo area without apparent overlapping. It seems
therefore that life in the same or very similar habitats has
led to the development of highly similar wing patterns
in these two species. Convergence driven by adaptation
appears therefore to be a common feature of Steroma-
pedaliodes sensu novo butteries. The newly described
species, S. mavarezi, which bears striking resemblance
to the syntopic Cheimas opalinus seems to be another
evident example of this phenomenon.
As stated earlier, the somehow morphologically simi-
lar Dangond is not the sister taxon of Steromapedaliodes
sensu novo, and they are in fact distantly related within
the Pedaliodes complex. This suggests that the similari-
ties between the two taxa are the product of some sort
of morphological convergence probably associated with
life in the high altitudes of the Cordillera de Mérida and
Perijá, respectively. Our study indicates therefore that
the evolutionary responses to similar extrinsic factors
may induce morphological convergence not only affect-
ing discrete colour pattern and other external characters,
but also male and female genitalia. It is difcult at the
moment to establish the origin or the processes behind
such convergences for genitalia since there are no obvi-
ous ecological correlations between life in the paramo
and, for example, the loss of subunci or signa. However,
we notice that the members of Steromapedaliodes sensu
novo and Dangond occupy habitats with no other conge-
ners and therefore there are no chances of interspecic
signal jamming in species recognition and courtship,
which in turn may lead to the disappearing of some sexu-
ally related characters associated with species recogni-
tion. This may explain the presence of simple aedeagus
in the males coupled with short ductus bursae in the fe-
males. On the other hand, the inefciency of chemical
communication in windswept open grassland may induce
the disappearing of specialized androconial scales. In
fact, several other paramo species of Pedaliodes complex
associated with the genus Altopedaliodes also have short
and straight aedeagus, and no specialized androconial
scales in Punapedaliodes (pyrCz & Viloria 1999; Vilo-
ria 2000), although they may occur syntopically with one
or two congeners. However, this remark seems to apply
to Satyrinae only, since within some members of paramo
dwelling Lycaenidae, such as the genus Penaincisalia
Johnson, 1990, scent organs are fully developed and dif-
fer between sympatric species (prieto 2008).
Certain important issues deserve further considera-
tion, although some of them remain yet unsolved due to
insufcient data. First, the closest relative of the genus
Steromapedaliodes sensu novo clade is still to be identi-
ed. Morphological data clearly point out that it should
be a member of the Pedaliodes complex, but our mo-
lecular phylogenetic analysis failed to identify a clear
relationship between the genus Steromapedaliodes sensu
novo and any taxa in the set used as outgroup taxa, either
of our own or from GenBank. It must be acknowledged
however that taxa in the Pedaliodes complex with avail-
able mitochondrial and nuclear sequences in GenBank
represent a very small fraction of this large group, both
in terms of diversity (i.e. 25 taxa out of ~ 300) and geo-
graphic distribution (i.e. only Peruvian taxa although the
group is distributed from Mexico to Argentina). The ge-
nus Steromapedaliodes sensu novo must therefore remain
incertae sedis until a thorough phylogenetic analysis us-
ing a much larger taxonomic and geographic sampling
within the Pedaliodes complex could be achieved.
5. Acknowledgements
TWP research projects in the Venezuelan Cordillera de Mérida were
supported by research grants of the Jagiellonian University – BW/
IZ/13/2005 (Kalibracja danych molekularnych w oparciu o dane
paleoekologiczne i aktualne wzorce rozmieszczenia gatunków z
rodzajów Steromapedaliodes i Redonda w Kordylierze Meridy) and
WRBW/BiNoZ/IZ/73/2007 (Fauna motyli z plemienia Pronophilini
górnego piętra lasu chmurnego oraz paramo masywu Cendé, Wen-
ezuela). Part of the research was nanced by the National Center of
Science, resolution number DEC-2013/ 09NNZ8/03219. JM would
like to thank both Maribel Gonzalez at the Smithsonian Tropical
Research Institute in Panama (STRI) and the staff at the Unidad de
Ecología Genética of the Instituto Venezolano de Investigaciones
Cientícas (IVIC) for technical help. Most of the work of JM in this
project was completed with the support from STRI, IVIC and the
Laboratoire d’Ecologie Alpine (LECA) of the Université Grenoble
Alpes in France. The following persons contributed in different
ways in eld research, M. Costa, M. Garcia, K. Casner, R. Calchi,
E. Moscó, P. Mora, J. Suarez, A.L. Viloria and R. Piña. Venezue-
lan National Park Authorities (INPARQUES) in Lara, Mérida and
Táchira provided permits for scientic research in National Parks.
J. Wojtusiak (late) has collaborated very actively during the later
years of this research producing a large amount of useful informa-
tion along with the accurate female dissections and drawings, and is
here warmly acknowledged. Further genital dissections and micro-
scopic photographs were executed by the staff of MZUJ in Kraków,
in particular A. Czekaj, and the distribution map was composed by
K. Sattler, D. Lees, G. Beccaloni, R.I. Vane-Wright and R. Gar-
lacz reviewed and edited several parts of the manuscript at different
stages. Valuable opinions and suggestions to the text were provided
by K. Casner. The authors have no conict of interest.
P et al.: Taxonomy and phylogeny of Steromapedaliodes
226
Fig. 5. Entire adults (left: dorsum; right: venter). A: Steromapedaliodes albonotata albonotata ♂, La Aguada. B: S. albonotata albono-
tata ♀, La Aguada. C: S. albonotata australis ♂, Páramo de San José. D: S. albonotata australis ♀, Páramo de San José. E: S. albonotata
australis ♂, Mesa Alta. F: S. albonotata australis ♀, Mesa Alta. G: S. albonotata australis ♂, Páramo de Guaraque. H: S. albonotata
australis ♀, Páramo de Guaraque.
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Fig. 6. Entire adults (left: dorsum; right: venter). A: Steromapedaliodes albonotata pratense ♂, El Potrero, Piñango. B: S. albonotata
pratense ♂, El Potrero, Piñango. C: S. albonotata sanchezi ♂, Los Morritos. D: S. albonotata pratense ♀, El Potrero, Piñango. E: S. al-
bonotata borealis ♂, Páramo de las Rosas. F: S. albonotata borealis ♀, Páramo de las Rosas. G: S. albonotata borealis ♂, Páramo de las
Rosas. H: S. albonotata borealis ♀, Páramo de las Rosas.
P et al.: Taxonomy and phylogeny of Steromapedaliodes
228
Fig. 7. Entire adults (left: dorsum; right: venter). A: Steromapedaliodes albonotata schuberti ♂, Páramo de Guaramacal. B: S. albonotata
schuberti ♀, Páramo de Guaramacal. C: S. albonotata josenae ♂, El Baho. D: S. albonotata josenae ♀, El Baho. E: S. albonotata jose-
nae ♂, El Baho. F: S. albonotata josenae ♀, El Baho. G: S. albonotata thiemeiana ♂, Páramo La Culata. H: S. albonotata thiemeiana ♀,
Páramo La Culata.
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Fig. 8. Entire adults (left: dorsum; right: venter). A: Steromapedaliodes albarregas albarregas ♂, Quebrada de La Boba. B: S. albarregas
albarregas ♀, Quebrada de La Boba. C: S. albarregas prietoi ♂, El Potrero, Piñango. D: S. albarregas prietoi ♀, El Potrero, Piñango. E:
S. albarregas mauroi ♂, Los Morritos, Río Gavidia. F: S. kahlua ♀, El Baho, la Ciénaga. G: S. mavarezi ♂, Páramo de Guirigay. H: S.
mavarezi ♀, Páramo de Guirigay.
P et al.: Taxonomy and phylogeny of Steromapedaliodes
230
Fig. 9. Entire adults (left: dorsum; right: venter). A: Steromapedaliodes empetrus empetrus ♂, Páramo La Culata. B: S. empetrus em-
petrus ♂, Páramo de la Culata. C: S. empetrus empetrus ♀, Páramo La Culata. D: S. empetrus frailejona ♂, Los Frailes. E: S. empetrus
frailejona ♀, Los Frailes. F: S. chiquinquirana chiquinquirana ♂, Laguna Mucubají. G: S. chiquinquirana chiquinquirana ♀, Laguna de
Mucubají. H: S. chiquinquirana lossadana ♂, Páramo de Tuñame. I: S. chiquinquirana lossadana ♀, Páramo de Tuñame. J: S. chiquin-
quirana lossadana ♀, Páramo de Tuñame.
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Fig. 10. Entire adults (left: dorsum; right: venter). A: Dangond dangondi ♂, Páramo el Avión. B: Dangond dangondi ♀, Páramo el Avión.
C: Steromapedaliodes bordoni ♂, Páramo El Rosal. D: S. bordoni ♀, Páramo El Rosal. E: S. lathraia ♂, Mesa Alta. F: S. lathraia ♀, Mesa
Alta. G: S. empetrus bolivari ♂, Loma Redonda. H: S. empetrus bolivari ♂, Loma Redonda.
P et al.: Taxonomy and phylogeny of Steromapedaliodes
232
Fig. 11. Entire adults (left: dorsum; right: venter). A: Steromapedaliodes chiquinquirana castellana ♂, Páramo de Cabimbú. B: S. chiquin-
quirana castellana ♀, Páramo de Cabimbú. C: S. chiquinquirana castellana ♂, Páramo de Las Moras. D: S. chiquinquirana castellana ♀,
Páramo de Las Moras. E: S. chiquinquirana castellana ♂, Páramo de Las Moras. F: S. chiquinquirana castellana ♀, Páramo de Las Moras.
G: S. leukasmena ♂, Páramo de Las Rosas. H: S. leukasmena ♀, Páramo de Las Rosas.
233
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Fig. 12. Male genitalia (in lateral view, aedeagus removed from its natural position). A: Steromapedaliodes albonotata albonotata, La
Aguada. B: S. albonotata thiemeiana, Páramo La Culata. C: S. albonotata australis, El Batallón, Mesa Alta. D: S. albonotata sanchezi,
Los Morritos. E: S. albonotata pratense, El Potrero, Piñango. F: S. albonotata josenae, El Baho. — Abbreviations: un uncus; subun
subuncus; va valva; aed aedeagus; teg tegumen; ped pedunculus; vin vinculum; sa saccus.
P et al.: Taxonomy and phylogeny of Steromapedaliodes
234
Fig. 13. Male genitalia (in lateral view, aedeagus removed from its natural position). A: Steromapedaliodes albonotata schuberti, Páramo
de Guaramacal. B: S. albonotata borealis, Páramo de las Rosas. C: S. albarregas albarregas, Quebrada de La Boba. D: S. albarregas
prietoi, El Potrero, Piñango. E: S. mavarezi, Páramo de Guirigay. F: D. dangondi, Sabana Rubia.
235
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Fig. 14. Male genitalia (in lateral view, aedeagus removed from its natural position). A: Steromapedaliodes empetrus frailejona, Los Frail-
es. B: S. empetrus bolivari, Loma Redonda. C: S. empetrus empetrus, Páramo La Culata. D: S. chiquinquirana chiquinquirana, Laguna de
Mucubají. E: S. chiquinquirana lossadana, Páramo de Tuñame. F: S. chiquinquirana lossadana, Páramo de Tuñame.
P et al.: Taxonomy and phylogeny of Steromapedaliodes
236
Fig. 15. Male genitalia (in lateral view, aedeagus removed from its natural position). A: Steromapedaliodes chiquinquirana castellana,
Páramo Las Moras. B: S. leukasmena, Páramo de Las Rosas. C: S. bordoni, Páramo El Rosal. D: S. lathraia, Mesa Alta. E: Panyapeda-
liodes panyasis, Acjanaco – Pillcopata. F. Pedaliodes plotina plotina, Colonia Tovar.
237
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Fig. 16. Female genitalia (in lateral view). A: Steromapedaliodes albonotata borealis, Páramo de las Rosas. B: S. albonotata pratense, El
Potrero, Piñango. C: S. albarregas albarregas, Quebrada de La Boba. D: S. albarregas prietoi, El Potrero, Piñango. E: S. kahlua, El Baho,
la Ciénaga. F: S. mavarezi, Páramo de Guirigay. — Abbreviations: pap.a papillae anales; apo.p apophyses posteriores; ll.p postvaginal
lamella; du.bu ductus bursae; du.sml ductus seminalis; bu.cop bursa copulatrix.
P et al.: Taxonomy and phylogeny of Steromapedaliodes
238
Fig. 17. Female genitalia (in lateral view). A: Steromapedaliodes albonotata schuberti, Páramo de Guaramacal. B: S. albonotata thiemei-
ana, Páramo La Culata. C: S. albonotata australis, Páramo San Jose. D: S. albonotata australis, El Batallón, Mesa Alta. E: S. albonotata
josenae, El Baho. F: S. albonotata albonotata, La Aguada.
239
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Fig. 18. Female genitalia (in lateral view). A: Steromapedaliodes empetrus frailejona, Los Frailes. B: S. empetrus empetrus, Páramo de la
Culata. C: S. chiquinquirana chiquinquirana, Laguna de Mucubají. D: S. chiquinquirana castellana, Páramo Las Moras. E: S. chiquinqui-
rana lossadana, Páramo de Tuñame. F: S. chiquinquirana lossadana, Páramo de Tuñame.
P et al.: Taxonomy and phylogeny of Steromapedaliodes
240
Fig. 19. Female genitalia (in lateral view). A: Steromapedaliodes bordoni, Páramo El Rosal. B: S. lathraia, Mesa Alta. C: S. leukasmena,
Páramo de Las Rosas. D: Dangond dangondi, Sabana Rubia. E: Panyapedaliodes drymaea, Chachapoyas. F: Pedaliodes plotina plotina,
Colonia Tovar.
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Electronic Supplement File
at http://www.senckenberg.de/arthropod-systematics
File 1: pyrcz&al-nymphalidae-steromapedaliodes-asp2017-elec
tronicsupplement.doc – Table S1. Primers used to amplify mito-
chondrial and nuclear markers.
Zoobank registrations
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9B55-730A271DA021
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F96B-4272-9AB7-BC83376378D3
Steromapedaliodes albonotata pratense Pyrcz & Boyer, 2017:
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E3DC-4073-A421-F3720AD6DB04
Steromapedaliodes albonotata thiemeiana Pyrcz, 2017: http://
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44A5-BB80-875E0D949799
