Comment on Braby et al. (2024)45
ANARTIA
Publicación del Museo de Biología de la Universidad del Zulia
ISSN 1315-642X (impresa) / ISSN 2665-0347 (digital)
https://doi.org/10.5281/zenodo.13755846 / Anartia, 38 (junio 2024): 45-51
http://zoobank.org/urn:lsid:zoobank.org:pub:DB9AE6EA-DEF4-4EF6-A605-152F8C713349
Defective taxonomic descriptions and the electronic publication
fashion. A comment on Braby et al. (2024) and a rectification
Descripciones taxonómicas defectuosas y la moda de la publicación electrónica.
Comentario sobre Braby et al. (2024) y una rectificación
Ángel L. Viloria
Centro de Ecología, Instituto Venezolano de Investigaciones Científicas (IVIC), km 11 carretera Panamericana, Altos de Pipe,
estado Miranda 1204, Venezuela.
Correspondencia: sebastianviloriacarrizo@gmail.com
(Received: 31-05-2024 / Accepted: 22-06-2024 / On line: 12-09-2024)
ABSTRACT
Some reflections are presented on the global practice of publishing scientific works in electronic format, its demands and
challenges. In this context, the recently published article by Braby et al. (2024) is discussed in relation to scientific best
practice in the description of new zoological species. That article refers to numerous descriptive works that suffer from
formal defects and irregularities that nullify the availability of the scientific names of the species described or proposed
in them, due to non-compliance with the regulations established in the International Code of Zoological Nomenclature
(ICZN). One such case is the description of Redonda bordoni Viloria & Pyrcz, 2003 (Insecta: Lepidoptera: Nymphali-
dae, Satyrinae), Bordón’s brachypterous butterfly, a species endemic to a high Andean sector of western Venezuela, whose
name would not be available because its description, together with its typification, mention of the repositories of the type
specimens and the explanation of the etymology of the name, were relegated to an electronic appendix presumably inacces-
sible (at least for a period of time). To resolve this irregularity, the species is redescribed, complying with the fundamental
recommendations of the ICZN.
Key words: Lepidoptera, Nymphalidae, Pronophilina, Redonda bordoni, Satyrinae, Satyrini.
RESUMEN
Se presentan algunas reflexiones sobre la práctica global de la publicación de obras científicas en formato electrónico, sus
exigencias y retos. En este contexto se comenta el artículo recientemente publicado por Braby et al. (2024) en relación a las
mejores prácticas científicas en la descripción de nuevas especies zoológicas. En dicho artículo se hace referencia a numero-
sos trabajos descriptivos que adolecen de defectos formales e irregularidades que anulan la disponibilidad de los nombres
científicos de las especies en ellos descritas o propuestas, por incumplimiento de la normativa establecida en el Código In-
ternacional de Nomenclatura Zoológica (ICZN). Uno de tales casos es la descripción de Redonda bordoni Viloria & Pyrcz,
2003 (Insecta: Lepidoptera: Nymphalidae, Satyrinae), la mariposa braquíptera de Bordón, especie endémica de un sector
altiandino del occidente de Venezuela, cuyo nombre no estaría disponible debido a que su descripción, junto con su debida
tipificación, la mención de los repositorios de los especímenes tipo y la explicación de la etimología del nombre, fueron
relegadas a un apéndice electrónico presuntamente inasequible (al menos durante un lapso de tiempo). Para solventar esta
irregularidad se redescribe la especie, cumpliendo con las recomendaciones fundamentales del ICZN.
Palabras clave: Lepidoptera, Nymphalidae, Pronophilina, Redonda bordoni, Satyrinae, Satyrini.
ANARTIA
Publicación del Museo de Biología de la Universidad del Zulia
ISSN 1315-642X (impresa) / ISSN 2665-0347 (digital)
https://doi.org/10.5281/zenodo.13755846 / Anartia, 38 (junio 2024): 45-51
http://zoobank.org/urn:lsid:zoobank.org:pub:DB9AE6EA-DEF4-4EF6-A605-152F8C713349
Defective taxonomic descriptions and the electronic publication
fashion. A comment on Braby et al. (2024) and a rectification
Descripciones taxonómicas defectuosas y la moda de la publicación electrónica.
Comentario sobre Braby et al. (2024) y una rectificación
Ángel L. Viloria
Centro de Ecología, Instituto Venezolano de Investigaciones Científicas (IVIC), km 11 carretera Panamericana, Altos de Pipe,
estado Miranda 1204, Venezuela.
Correspondencia: sebastianviloriacarrizo@gmail.com
(Received: 31-05-2024 / Accepted: 22-06-2024 / On line: 12-09-2024)
ABSTRACT
Some reflections are presented on the global practice of publishing scientific works in electronic format, its demands and
challenges. In this context, the recently published article by Braby et al. (2024) is discussed in relation to scientific best
practice in the description of new zoological species. That article refers to numerous descriptive works that suffer from
formal defects and irregularities that nullify the availability of the scientific names of the species described or proposed
in them, due to non-compliance with the regulations established in the International Code of Zoological Nomenclature
(ICZN). One such case is the description of Redonda bordoni Viloria & Pyrcz, 2003 (Insecta: Lepidoptera: Nymphali-
dae, Satyrinae), Bordón’s brachypterous butterfly, a species endemic to a high Andean sector of western Venezuela, whose
name would not be available because its description, together with its typification, mention of the repositories of the type
specimens and the explanation of the etymology of the name, were relegated to an electronic appendix presumably inacces-
sible (at least for a period of time). To resolve this irregularity, the species is redescribed, complying with the fundamental
recommendations of the ICZN.
Key words: Lepidoptera, Nymphalidae, Pronophilina, Redonda bordoni, Satyrinae, Satyrini.
RESUMEN
Se presentan algunas reflexiones sobre la práctica global de la publicación de obras científicas en formato electrónico, sus
exigencias y retos. En este contexto se comenta el artículo recientemente publicado por Braby et al. (2024) en relación a las
mejores prácticas científicas en la descripción de nuevas especies zoológicas. En dicho artículo se hace referencia a numero-
sos trabajos descriptivos que adolecen de defectos formales e irregularidades que anulan la disponibilidad de los nombres
científicos de las especies en ellos descritas o propuestas, por incumplimiento de la normativa establecida en el Código In-
ternacional de Nomenclatura Zoológica (ICZN). Uno de tales casos es la descripción de Redonda bordoni Viloria & Pyrcz,
2003 (Insecta: Lepidoptera: Nymphalidae, Satyrinae), la mariposa braquíptera de Bordón, especie endémica de un sector
altiandino del occidente de Venezuela, cuyo nombre no estaría disponible debido a que su descripción, junto con su debida
tipificación, la mención de los repositorios de los especímenes tipo y la explicación de la etimología del nombre, fueron
relegadas a un apéndice electrónico presuntamente inasequible (al menos durante un lapso de tiempo). Para solventar esta
irregularidad se redescribe la especie, cumpliendo con las recomendaciones fundamentales del ICZN.
Palabras clave: Lepidoptera, Nymphalidae, Pronophilina, Redonda bordoni, Satyrinae, Satyrini.
Viloria46
INTRODUCTION
The transition from the publication of scientific prints
to the electronic publication of books, journals and special-
ized articles has taken nearly three decades. It has required
the progressive consideration and incorporation of novel
editorial aspects imposed by this fashion, as well as the
implementation of new rules and practices that guarantee
formality and professional ethics in the publication and
dissemination processes, and above all that ensure the per-
petual availability of electronic documents through their
hosting in secure repositories, preferably institutional.
Many difficulties are being overcome to successfully
couple numerous regulations to legitimize scientific pro-
duction with the globalization of the practice of electronic
publishing. However, the reduction of ecological and eco-
nomic costs by decreasing the use of paper, global open-
ness through electronic communication, as well as the re-
duction of time lapses in the editorial process, have been
accompanied not only by the multiplication of electronic
means of disseminating specialized information but also
the number of people (researchers or not) who make al-
most frenetic use of them.
In response to the proliferation of unprofessional and
irregular taxonomic work, Braby et al. (2024) have pub-
lished an interesting, entertaining and necessarily critical
essay on the formal procedures to describe animal species
in the most appropriate way and in full compliance with
the rules established in the International Code of Zoo-
logical Nomenclature (ICZN 1999, 2012). Not only is
it an extraordinary theoretical review of the topic, which
has also been previously treated by other authors, but it
presents a notable number of examples of bad taxonomic
practice, particularly in relation to the description of new
species, providing no few recommendations to minimize
the risks of making errors.
One such example refers to the publication of the first
case of brachyptery in butterflies (Lepidoptera: Papil-
ionoidea) (Viloria et al. 2003), represented in the female
of a then undescribed species of the genus Redonda Ad-
ams & Bernard, 1981, which lives in the high elevations
of the Andean mountains of Venezuela. Brachyptery in
butterflies is a peculiar biological phenomenon, studied
in the field by the authors of that article for one decade,
the information about it was organized fundamentally as
a descriptive ecological study, in which comparative mor-
phology was necessary to describe the species involved,
but also to carry out elementary morphometric calcula-
tions and a statistical analysis designed to mathematically
demonstrate the physical limitations that prevent flight
in females of said taxon, which derive from a disadvanta-
geous relationship between wing surface and body size. In
addition to the corresponding considerations on the eco-
logical conditions of the tropical highland environment
as a source of possible selective forces involved in the loss
of flight capabilities, the work included the diagnosis and
complete taxonomic description of the species.
The resulting study, submitted for consideration for
publication in the Proceedings of the Royal Society B: Bio-
logical Sciences, received approval from its editorial com-
mittee and reviewers. However, making its publication
conditional on a reduction in the length of the text to
accommodate it in the first issue of the supplement to
the Proceedings B, called Biology Letters, established in
2003 and launched as an independent journal from 2005
(https://royalsocietypublishing.org/journal/rsbl). Not
only were the authors committed to reducing the length
of the work to meet the editorial demands of such a presti-
gious journal, but they were also compromised to separate
the diagnosis from the taxonomic description, relegating
the latter, along with some illustrations, to form part of an
electronic appendix. Unfortunately, as has been pointed
out by Braby et al. (2024), at the time, such actions could
have led to the violation of some of the rules established
by the International Code of Zoological Nomenclature
(ICZN 1999) to guarantee the availability of scientific
names assigned to animal species.
Braby et al. (2024) state that the electronic appendix
in which the description of the aforementioned brachyp-
terous butterfly appeared is no longer accessible, so it was
not possible for them to access the data corresponding to
the series of type specimens and their repositories. Conse-
quently, in the opinion of those authors, the provisions of
Article 16.4 of the ICZN have not been complied with,
which is why the name of the species, proposed in 2003,
is not available.
As can be seen in the next section of this work (below),
the experience of the present author regarding the acces-
sibility of electronic documents in question contradicts
the statement of Braby et al. (2024), in that there is an ac-
cess link to them on the Royal Society of London website.
However, it is not possible to assert that the appendices to
the article by Viloria et al. (2003) have been continuously
available during the 21 years since their publication.
Given this situation of relative ambiguity and in the
spirit of rectifying any past, present or future irregular-
ity, in relation to procedures or acts that may affect the
availability of a scientific name proposed more than two
decades ago, and ever since used with relative frequency
by several authors, the decision has been made to repub-
lish the taxonomic description of the butterfly species to
which said name corresponds, and in whose content the
INTRODUCTION
The transition from the publication of scientific prints
to the electronic publication of books, journals and special-
ized articles has taken nearly three decades. It has required
the progressive consideration and incorporation of novel
editorial aspects imposed by this fashion, as well as the
implementation of new rules and practices that guarantee
formality and professional ethics in the publication and
dissemination processes, and above all that ensure the per-
petual availability of electronic documents through their
hosting in secure repositories, preferably institutional.
Many difficulties are being overcome to successfully
couple numerous regulations to legitimize scientific pro-
duction with the globalization of the practice of electronic
publishing. However, the reduction of ecological and eco-
nomic costs by decreasing the use of paper, global open-
ness through electronic communication, as well as the re-
duction of time lapses in the editorial process, have been
accompanied not only by the multiplication of electronic
means of disseminating specialized information but also
the number of people (researchers or not) who make al-
most frenetic use of them.
In response to the proliferation of unprofessional and
irregular taxonomic work, Braby et al. (2024) have pub-
lished an interesting, entertaining and necessarily critical
essay on the formal procedures to describe animal species
in the most appropriate way and in full compliance with
the rules established in the International Code of Zoo-
logical Nomenclature (ICZN 1999, 2012). Not only is
it an extraordinary theoretical review of the topic, which
has also been previously treated by other authors, but it
presents a notable number of examples of bad taxonomic
practice, particularly in relation to the description of new
species, providing no few recommendations to minimize
the risks of making errors.
One such example refers to the publication of the first
case of brachyptery in butterflies (Lepidoptera: Papil-
ionoidea) (Viloria et al. 2003), represented in the female
of a then undescribed species of the genus Redonda Ad-
ams & Bernard, 1981, which lives in the high elevations
of the Andean mountains of Venezuela. Brachyptery in
butterflies is a peculiar biological phenomenon, studied
in the field by the authors of that article for one decade,
the information about it was organized fundamentally as
a descriptive ecological study, in which comparative mor-
phology was necessary to describe the species involved,
but also to carry out elementary morphometric calcula-
tions and a statistical analysis designed to mathematically
demonstrate the physical limitations that prevent flight
in females of said taxon, which derive from a disadvanta-
geous relationship between wing surface and body size. In
addition to the corresponding considerations on the eco-
logical conditions of the tropical highland environment
as a source of possible selective forces involved in the loss
of flight capabilities, the work included the diagnosis and
complete taxonomic description of the species.
The resulting study, submitted for consideration for
publication in the Proceedings of the Royal Society B: Bio-
logical Sciences, received approval from its editorial com-
mittee and reviewers. However, making its publication
conditional on a reduction in the length of the text to
accommodate it in the first issue of the supplement to
the Proceedings B, called Biology Letters, established in
2003 and launched as an independent journal from 2005
(https://royalsocietypublishing.org/journal/rsbl). Not
only were the authors committed to reducing the length
of the work to meet the editorial demands of such a presti-
gious journal, but they were also compromised to separate
the diagnosis from the taxonomic description, relegating
the latter, along with some illustrations, to form part of an
electronic appendix. Unfortunately, as has been pointed
out by Braby et al. (2024), at the time, such actions could
have led to the violation of some of the rules established
by the International Code of Zoological Nomenclature
(ICZN 1999) to guarantee the availability of scientific
names assigned to animal species.
Braby et al. (2024) state that the electronic appendix
in which the description of the aforementioned brachyp-
terous butterfly appeared is no longer accessible, so it was
not possible for them to access the data corresponding to
the series of type specimens and their repositories. Conse-
quently, in the opinion of those authors, the provisions of
Article 16.4 of the ICZN have not been complied with,
which is why the name of the species, proposed in 2003,
is not available.
As can be seen in the next section of this work (below),
the experience of the present author regarding the acces-
sibility of electronic documents in question contradicts
the statement of Braby et al. (2024), in that there is an ac-
cess link to them on the Royal Society of London website.
However, it is not possible to assert that the appendices to
the article by Viloria et al. (2003) have been continuously
available during the 21 years since their publication.
Given this situation of relative ambiguity and in the
spirit of rectifying any past, present or future irregular-
ity, in relation to procedures or acts that may affect the
availability of a scientific name proposed more than two
decades ago, and ever since used with relative frequency
by several authors, the decision has been made to repub-
lish the taxonomic description of the butterfly species to
which said name corresponds, and in whose content the
Comment on Braby et al. (2024)47
species is appropriately typified. Likewise, in compliance
with the requirements of the most recent amendment
to the International Code of Zoological Nomenclature
(ICZN 2012), these acts are registered in ZooBank.
MATERIAL AND METHODS
Electronic Appendix A to Viloria et al. (2003), cur-
rently available at https://royalsocietypublishing.org/
action/downloadSupplement?doi=10.1098%2Frsbl.2
003.0015&file=bl030021_supp01.pdf ), was retrieved
by the author on May 12, 2024. The text of the primary
taxonomic description of Redonda bordoni (Lepidoptera:
Nymphalidae, Satyrinae) contained in this document was
extracted for adequate electronic and printed publication
in the present contribution. In this way, the transcription
of full data of the type material, including its repositories,
as well as the etymology of the name of the species and
its registration in ZooBank solve the problem of the avail-
ability of its name.
Some collector names have been inserted into the origi-
nal abbreviations (in square brackets), and an annotated
synonymy is provided for this taxon.
Abbreviations
A & B: [Michael J.] Adams & [George] Bernard;
BMNH: The Natural History Museum, London, UK;
JFLC: Jean-François Le Crom Collection, Bogotá, Co-
lombia; MALUZ: Museo de Artrópodos, La Univer-
sidad del Zulia, Maracaibo, Venezuela; MHN: Museo de
Ciencias Naturales, Universidad Nacional de Colombia;
MIZA: Museo del Instituto de Zoología Agrícola, Univer-
sidad Central de Venezuela, Maracay, Venezuela; MZUJ:
Zoological Museum of the Jagiellonian University, Kra-
kow, Poland; P & V: [T. W.] Pyrcz & [Á. L.] Viloria; V &
P: [Á. L.] Viloria & [T. W.] Pyrcz.
RESULTS
Family Nymphalidae Rafinesque, 1815
Subfamily Satyrinae Boisduval, 1833
Genus Redonda Adams & Bernard, 1981
Redonda bordoni, sp. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:6D484533-A1B0-49D1-
9D1D-7312461FC2E5
Original figures in Viloria et al., 2003: fig. 1a [male ho-
lotype, female paratype; ventral] (not reproduced here);
e-Appendix A: fig 5 [female paratype, dorsal] (herein re-
produced in Fig. 1), figs. 6a [male genitalia], 6b [female
genitalia], 6c [female wing venation] (herein reproduced
in Fig. 2), and in Viloria et al. 2015: figs. 14 [male, holo-
type], 15 [female, allotype].
Type locality
Between Laguna El Cenegón and Laguna Grande,
3200-3400 m, Páramo El Batallón, Estado Táchira, Ven-
ezuela.
[Redonda bordoni Viloria and Pyrcz, MS, nomen nu-
dum; Viloria, 1998: 319; Ferrer-Paris, 2000: 96
(tbl. C.9)]
[Redonda sp. nov. 1; Ferrer-Paris, 2000: 27 (fig. 3.1,
distribution), 29 (tbl. 3.1), 36, 37 (fig. 3.5), 38,
40, 41 (tbl. 3.6), 69, 91 (tbl. C.7); Viloria, 2000:
269; Ferrer-Paris & Viloria, [2004]: 628 (fig. 1);
629 (tbl. 1), 630 (fig. 3), 631]
[Redonda bordoni Viloria and Pyrcz, in Viloria
et al., 2003: 21–23 (figs. 1a, male, female, 2,
3, 4), e-appendices: [1], [4 (fig. 5, female)], [5
(figs. 6a male genitalia, 6b female genitalia, 6c
female wing venation)]) (in part misidentifi-
cations of R. bolivari Adams & Bernard, 1981
and R. lathraia Viloria & Camacho, 2015];
[Anonymous], 2003: 24; Blackman, 2003: 26;
Williams, 2003: R467; Lamas et al., 2004: 215;
Viloria, 2005: 459; 2008: 278; Bálint & Wojtu-
siak, 2006: 288; Pyrcz, 2007a: 40, 41; 2007b:
17, 18, 19; 2010a: 36 (fig. 17C, antennal club),
45 (fig. 35C, female venation), 87 (fig. 82, cla-
dogram), 111, 179, 180 (fig. 131, wing area),
181 (figs. 132A, male; B, female, 133A, pair in
copula), 182, 183, 184 (fig. 136, distribution),
244; 2010b: 265, 266 (figs. 1K male, 1L female,
267, 268 (fig. 2), 269 (figs. 3A, B, C females),
271 (fig. 5B pair in copula), 272, 273; Pyrcz et
al., 2007: [26] [3 figs., male & female], Łabno,
2007: 104; Davies & Butler, 2008: 33; Viloria
et al., 2015: 97, 105 (fig. 5 [female wing vena-
tion]), 107, 109 (figs. 14 [male, holotype], 15
[female, allotype]), 110, 111, 112, 135 (as misi-
dentification of R. lathraia Viloria & Camacho,
2015), 136, 137, 138, 145 (fig. 48 [male geni-
talia]), 154, 160; Ferrer-Paris et al., 2015: 322;
Pyrcz et al., 2017: 198, 221], nomen nudum
[Steromapedaliodes bordoni (Viloria & Pyrcz); Pyrcz
et al., 2017: 195, 197 (fig. 1B, distribution map),
200 (tbl. 1), 204 (fig. 4A [male wing venation]),
221, 222, 223, 224, 231 (figs. 10C [male], 10D
[female]), 236 (fig. 15C [male genitalia]), 240
(fig. 19A [female genitalia]; Boyer, 2018: 122,
123 pl. 2 (figs. 5 [pair in copula], 6 [female]), 124,
125 pl. 3 (figs. 5, 7 [male], 6, 8 [female]), 129;
2019: 99, 104, 105 pl. 16 (fig. 14 [male], 129)],
nomen nudum
species is appropriately typified. Likewise, in compliance
with the requirements of the most recent amendment
to the International Code of Zoological Nomenclature
(ICZN 2012), these acts are registered in ZooBank.
MATERIAL AND METHODS
Electronic Appendix A to Viloria et al. (2003), cur-
rently available at https://royalsocietypublishing.org/
action/downloadSupplement?doi=10.1098%2Frsbl.2
003.0015&file=bl030021_supp01.pdf ), was retrieved
by the author on May 12, 2024. The text of the primary
taxonomic description of Redonda bordoni (Lepidoptera:
Nymphalidae, Satyrinae) contained in this document was
extracted for adequate electronic and printed publication
in the present contribution. In this way, the transcription
of full data of the type material, including its repositories,
as well as the etymology of the name of the species and
its registration in ZooBank solve the problem of the avail-
ability of its name.
Some collector names have been inserted into the origi-
nal abbreviations (in square brackets), and an annotated
synonymy is provided for this taxon.
Abbreviations
A & B: [Michael J.] Adams & [George] Bernard;
BMNH: The Natural History Museum, London, UK;
JFLC: Jean-François Le Crom Collection, Bogotá, Co-
lombia; MALUZ: Museo de Artrópodos, La Univer-
sidad del Zulia, Maracaibo, Venezuela; MHN: Museo de
Ciencias Naturales, Universidad Nacional de Colombia;
MIZA: Museo del Instituto de Zoología Agrícola, Univer-
sidad Central de Venezuela, Maracay, Venezuela; MZUJ:
Zoological Museum of the Jagiellonian University, Kra-
kow, Poland; P & V: [T. W.] Pyrcz & [Á. L.] Viloria; V &
P: [Á. L.] Viloria & [T. W.] Pyrcz.
RESULTS
Family Nymphalidae Rafinesque, 1815
Subfamily Satyrinae Boisduval, 1833
Genus Redonda Adams & Bernard, 1981
Redonda bordoni, sp. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:6D484533-A1B0-49D1-
9D1D-7312461FC2E5
Original figures in Viloria et al., 2003: fig. 1a [male ho-
lotype, female paratype; ventral] (not reproduced here);
e-Appendix A: fig 5 [female paratype, dorsal] (herein re-
produced in Fig. 1), figs. 6a [male genitalia], 6b [female
genitalia], 6c [female wing venation] (herein reproduced
in Fig. 2), and in Viloria et al. 2015: figs. 14 [male, holo-
type], 15 [female, allotype].
Type locality
Between Laguna El Cenegón and Laguna Grande,
3200-3400 m, Páramo El Batallón, Estado Táchira, Ven-
ezuela.
[Redonda bordoni Viloria and Pyrcz, MS, nomen nu-
dum; Viloria, 1998: 319; Ferrer-Paris, 2000: 96
(tbl. C.9)]
[Redonda sp. nov. 1; Ferrer-Paris, 2000: 27 (fig. 3.1,
distribution), 29 (tbl. 3.1), 36, 37 (fig. 3.5), 38,
40, 41 (tbl. 3.6), 69, 91 (tbl. C.7); Viloria, 2000:
269; Ferrer-Paris & Viloria, [2004]: 628 (fig. 1);
629 (tbl. 1), 630 (fig. 3), 631]
[Redonda bordoni Viloria and Pyrcz, in Viloria
et al., 2003: 21–23 (figs. 1a, male, female, 2,
3, 4), e-appendices: [1], [4 (fig. 5, female)], [5
(figs. 6a male genitalia, 6b female genitalia, 6c
female wing venation)]) (in part misidentifi-
cations of R. bolivari Adams & Bernard, 1981
and R. lathraia Viloria & Camacho, 2015];
[Anonymous], 2003: 24; Blackman, 2003: 26;
Williams, 2003: R467; Lamas et al., 2004: 215;
Viloria, 2005: 459; 2008: 278; Bálint & Wojtu-
siak, 2006: 288; Pyrcz, 2007a: 40, 41; 2007b:
17, 18, 19; 2010a: 36 (fig. 17C, antennal club),
45 (fig. 35C, female venation), 87 (fig. 82, cla-
dogram), 111, 179, 180 (fig. 131, wing area),
181 (figs. 132A, male; B, female, 133A, pair in
copula), 182, 183, 184 (fig. 136, distribution),
244; 2010b: 265, 266 (figs. 1K male, 1L female,
267, 268 (fig. 2), 269 (figs. 3A, B, C females),
271 (fig. 5B pair in copula), 272, 273; Pyrcz et
al., 2007: [26] [3 figs., male & female], Łabno,
2007: 104; Davies & Butler, 2008: 33; Viloria
et al., 2015: 97, 105 (fig. 5 [female wing vena-
tion]), 107, 109 (figs. 14 [male, holotype], 15
[female, allotype]), 110, 111, 112, 135 (as misi-
dentification of R. lathraia Viloria & Camacho,
2015), 136, 137, 138, 145 (fig. 48 [male geni-
talia]), 154, 160; Ferrer-Paris et al., 2015: 322;
Pyrcz et al., 2017: 198, 221], nomen nudum
[Steromapedaliodes bordoni (Viloria & Pyrcz); Pyrcz
et al., 2017: 195, 197 (fig. 1B, distribution map),
200 (tbl. 1), 204 (fig. 4A [male wing venation]),
221, 222, 223, 224, 231 (figs. 10C [male], 10D
[female]), 236 (fig. 15C [male genitalia]), 240
(fig. 19A [female genitalia]; Boyer, 2018: 122,
123 pl. 2 (figs. 5 [pair in copula], 6 [female]), 124,
125 pl. 3 (figs. 5, 7 [male], 6, 8 [female]), 129;
2019: 99, 104, 105 pl. 16 (fig. 14 [male], 129)],
nomen nudum
Viloria48
Figure 1. Redonda bordoni female displaying the silvery dorsal surfaces of its wings [corresponding to original figure 5 in e-Appendix A
to Viloria et al. 2003. Photo: J. Wojtusiak†
].
Figure 2. Morphological features of Redonda bordoni. a. Male genitalia (aedeagus and valvae removed from their natural positions:
Ae, aedeagus; Ss, saccus; T, tegumen; U, uncus; V, valva; Vm, vinculum). b. Female genitalia (A, anus; Cb, corpus bursae; Cg, colat-
eral gland; Db, ductus bursae; Ds, ductus seminalis; L, lagena; Ob, ostium bursae; Od, oviduct; Op, ovipore; PA, post-apophysis; R,
rectum; S, spermatheca; Sd, spermathecal duct). c. Venation of female wings [corresponding to original figure 6 in e-Appendix A to
Viloria et al. 2003. Female genitalia drawing (b) by J. Wojtusiak†
].
Figure 1. Redonda bordoni female displaying the silvery dorsal surfaces of its wings [corresponding to original figure 5 in e-Appendix A
to Viloria et al. 2003. Photo: J. Wojtusiak†
].
Figure 2. Morphological features of Redonda bordoni. a. Male genitalia (aedeagus and valvae removed from their natural positions:
Ae, aedeagus; Ss, saccus; T, tegumen; U, uncus; V, valva; Vm, vinculum). b. Female genitalia (A, anus; Cb, corpus bursae; Cg, colat-
eral gland; Db, ductus bursae; Ds, ductus seminalis; L, lagena; Ob, ostium bursae; Od, oviduct; Op, ovipore; PA, post-apophysis; R,
rectum; S, spermatheca; Sd, spermathecal duct). c. Venation of female wings [corresponding to original figure 6 in e-Appendix A to
Viloria et al. 2003. Female genitalia drawing (b) by J. Wojtusiak†
].
Comment on Braby et al. (2024)49
Male
Forewing length: 26–32 mm; n = 123; mean = 29.56.
Eyes hairy, reddish brown, circled with black and white
scales. Palpi twice as long as head, light brown, flanked
with white, with brown, dark brown and yellow hairs; first
segment quarter length of first. Antennae up to two fifths
length of costa, 32 segments; shaft orange brown, darker
towards club; club 2.5-3 times longer that wide, concave
(spoon-shaped). Body densely hairy, dorsally dark cof-
fee brown, ventrally light brown, almost white on abdo-
men; hairs glossy light brown. Forewing triangular, tornus
obtuse. Hindwing oval. Upperside ground colour shiny
brown, darker towards basal half and marginal region;
wing bases dark coffee brown; crossveins at distal extrem-
ity of discal cells covered by greyish white; series of five
to six submarginal elongate white spots on both wings.
Underside ground colour as above; forewing pattern as on
upperside; hindwing veins greyish white, as well as longi-
tudinal lines in discal and Cu2 cells, the former bifurcated
near base; postdiscal series of five to six fusiform white
ocellar marks; third Schwanwitsch’s externa, first and sec-
ond Schwanwitsch’s media coffee brown (Schwanwitsch
1924); all discontinuous and dislocated to form a system
of longitudinal lines parallel or oblique to veins and white
marks, in basal half of wing and postdiscal-submarginal re-
gion, respectively. Genitalia illustrated in Figure 2a.
Female
Forewing length: 20–29 mm; n = 4; mean = 20.87.
Wings considerably shorter and narrower than in male;
apex pronounced at hindwing vein M2. Upperside ground
colour silvery white; wing bases very dark coffee brown;
forewing densely dusted with dark brown in apical and dis-
cal region, and within discal cell. Underside colour pattern
as in male, dark coffee brown much more dense in basal
half of hindwing; white lines and veins more distinct; fu-
siform white ocellar marks heavily elongate; costal margin
dark coffee brown. Genitalia illustrated in Figure 2b.
Type material
Holotype male, taken between Laguna El Cenegón
and Laguna Grande, Páramo El Batallón, Estado Táchira,
Venezuela, 3200-3400 m, 28-ii-1994, A. Viloria, M. Gar-
cía & J. Camacho colls.; Allotype female, same data as
holotype (MALUZ). Paratypes: 27 males (3 in MHN, 3
in JFLC), 1 female, same data as holotype; 19 males (2 in
MZUJ, 2 in BMNH), VENEZUELA: Estado Táchira,
Páramo El Batallón, entre la Antena CANTV y la La-
guna El Cenegón, 3100-3250 m; 27-ii-1994, A. Viloria,
M. García & J. Camacho colls.; 4 males, Estado Táchira,
Municipio Jáuregui, Callejón del Cenegón, 16-xii-1994;
M. García coll.; 6 males, 1 female, Estado Táchira, Parque
Nacional Juan Peñaloza, Páramo El Rosal, 3000 m, 12/14-
i-1995; J. Camacho & M. García colls.; 10 males, Estado
Táchira, Páramo El Batallón, Entre El Cenegón y Laguna
Grande, 3300-3400 m, 05-iii-1996, J. Camacho, M. Gar-
cía, T. Pyrcz, J. Wojtusiak colls. (MALUZ); 1 male, Estado
Táchira, Páramo de La Negra, 30-ix-1951, P. Fenjues coll.;
1 male, same locality, 3200 m., 14-i-1982, C. Bordón coll.
(MIZA); 51 males, 1 female, Estado Táchira, Páramo El
Batallón, Vía El Púlpito, 3500-3800 m, 02/04-iii-1996, T.
Pyrcz, J. Wojtusiak, J. Camacho, M. García colls.; 5 males,
Estado Táchira, Páramo El Batallón, Vía El Cenegón, 04-
iii-1996, T. Pyrcz, J. Wojtusiak, J. Camacho, M. García
colls. (MZUJ).
Etymology
This butterfly species is named after Carlos Bordón
(b. Trieste, Italy, 1921; d. Maracay, Venezuela, 2012. See
Viloria 2018), prominent Italo-Venezuelan entomologist
who collected the first known individual of this unusual
butterfly.
ACKNOWLEDGEMENTS
I thank J. Llorente-Bousquets for drawing my atten-
tion to the recent publication by Braby et al. L. A. Gómez-
Urdaneta, P. Boyer, S. Attal and M. Costa assisted me in
searching and obtaining bibliography. People who helped
in the description included in this contribution are men-
tioned in the acknowledgements of Viloria et al. (2003).
REFERENCES
[Anonymous]. 2003. In brief. New species of wood nymph
found. Flutterfree lifestyles. New Scientist 2397: 24.
Bálint, Z. & J. Wojtusiak. 2006. Notes on the genus Podanotum
with description of a new species (Lepidoptera: Lycaenidae:
Eumaeini). Genus. International Journal of Invertebrate Tax-
onomy 17(2): 283–289.
Blackman, S. 2003. Flutterless butterfly. BBC Wildlife Magazine
21(8): 26.
Boyer, P. 2018. Introduction à la connaissance des Pronophilini
(Lepidoptera: Nymphalidae; Satyrinae). Lépidoptéres (Paris)
27(71): 118–130, 4 pls.
Boyer, P. 2019. Introduction à la connaissance des Pronophi-
lini (Lepidoptera: Nymphalidae; Satyrinae) – Troisième et
dernière partie. Lépidoptéres (Paris) 28(74): 98–129, pls.
14–21.
Braby, M. F., Y.-F. Hsu & G. Lamas. 2024. How to describe a
new species in zoology and avoid mistakes. Zoological Jour-
nal of the Linnean Society 2024: zlae043.
Male
Forewing length: 26–32 mm; n = 123; mean = 29.56.
Eyes hairy, reddish brown, circled with black and white
scales. Palpi twice as long as head, light brown, flanked
with white, with brown, dark brown and yellow hairs; first
segment quarter length of first. Antennae up to two fifths
length of costa, 32 segments; shaft orange brown, darker
towards club; club 2.5-3 times longer that wide, concave
(spoon-shaped). Body densely hairy, dorsally dark cof-
fee brown, ventrally light brown, almost white on abdo-
men; hairs glossy light brown. Forewing triangular, tornus
obtuse. Hindwing oval. Upperside ground colour shiny
brown, darker towards basal half and marginal region;
wing bases dark coffee brown; crossveins at distal extrem-
ity of discal cells covered by greyish white; series of five
to six submarginal elongate white spots on both wings.
Underside ground colour as above; forewing pattern as on
upperside; hindwing veins greyish white, as well as longi-
tudinal lines in discal and Cu2 cells, the former bifurcated
near base; postdiscal series of five to six fusiform white
ocellar marks; third Schwanwitsch’s externa, first and sec-
ond Schwanwitsch’s media coffee brown (Schwanwitsch
1924); all discontinuous and dislocated to form a system
of longitudinal lines parallel or oblique to veins and white
marks, in basal half of wing and postdiscal-submarginal re-
gion, respectively. Genitalia illustrated in Figure 2a.
Female
Forewing length: 20–29 mm; n = 4; mean = 20.87.
Wings considerably shorter and narrower than in male;
apex pronounced at hindwing vein M2. Upperside ground
colour silvery white; wing bases very dark coffee brown;
forewing densely dusted with dark brown in apical and dis-
cal region, and within discal cell. Underside colour pattern
as in male, dark coffee brown much more dense in basal
half of hindwing; white lines and veins more distinct; fu-
siform white ocellar marks heavily elongate; costal margin
dark coffee brown. Genitalia illustrated in Figure 2b.
Type material
Holotype male, taken between Laguna El Cenegón
and Laguna Grande, Páramo El Batallón, Estado Táchira,
Venezuela, 3200-3400 m, 28-ii-1994, A. Viloria, M. Gar-
cía & J. Camacho colls.; Allotype female, same data as
holotype (MALUZ). Paratypes: 27 males (3 in MHN, 3
in JFLC), 1 female, same data as holotype; 19 males (2 in
MZUJ, 2 in BMNH), VENEZUELA: Estado Táchira,
Páramo El Batallón, entre la Antena CANTV y la La-
guna El Cenegón, 3100-3250 m; 27-ii-1994, A. Viloria,
M. García & J. Camacho colls.; 4 males, Estado Táchira,
Municipio Jáuregui, Callejón del Cenegón, 16-xii-1994;
M. García coll.; 6 males, 1 female, Estado Táchira, Parque
Nacional Juan Peñaloza, Páramo El Rosal, 3000 m, 12/14-
i-1995; J. Camacho & M. García colls.; 10 males, Estado
Táchira, Páramo El Batallón, Entre El Cenegón y Laguna
Grande, 3300-3400 m, 05-iii-1996, J. Camacho, M. Gar-
cía, T. Pyrcz, J. Wojtusiak colls. (MALUZ); 1 male, Estado
Táchira, Páramo de La Negra, 30-ix-1951, P. Fenjues coll.;
1 male, same locality, 3200 m., 14-i-1982, C. Bordón coll.
(MIZA); 51 males, 1 female, Estado Táchira, Páramo El
Batallón, Vía El Púlpito, 3500-3800 m, 02/04-iii-1996, T.
Pyrcz, J. Wojtusiak, J. Camacho, M. García colls.; 5 males,
Estado Táchira, Páramo El Batallón, Vía El Cenegón, 04-
iii-1996, T. Pyrcz, J. Wojtusiak, J. Camacho, M. García
colls. (MZUJ).
Etymology
This butterfly species is named after Carlos Bordón
(b. Trieste, Italy, 1921; d. Maracay, Venezuela, 2012. See
Viloria 2018), prominent Italo-Venezuelan entomologist
who collected the first known individual of this unusual
butterfly.
ACKNOWLEDGEMENTS
I thank J. Llorente-Bousquets for drawing my atten-
tion to the recent publication by Braby et al. L. A. Gómez-
Urdaneta, P. Boyer, S. Attal and M. Costa assisted me in
searching and obtaining bibliography. People who helped
in the description included in this contribution are men-
tioned in the acknowledgements of Viloria et al. (2003).
REFERENCES
[Anonymous]. 2003. In brief. New species of wood nymph
found. Flutterfree lifestyles. New Scientist 2397: 24.
Bálint, Z. & J. Wojtusiak. 2006. Notes on the genus Podanotum
with description of a new species (Lepidoptera: Lycaenidae:
Eumaeini). Genus. International Journal of Invertebrate Tax-
onomy 17(2): 283–289.
Blackman, S. 2003. Flutterless butterfly. BBC Wildlife Magazine
21(8): 26.
Boyer, P. 2018. Introduction à la connaissance des Pronophilini
(Lepidoptera: Nymphalidae; Satyrinae). Lépidoptéres (Paris)
27(71): 118–130, 4 pls.
Boyer, P. 2019. Introduction à la connaissance des Pronophi-
lini (Lepidoptera: Nymphalidae; Satyrinae) – Troisième et
dernière partie. Lépidoptéres (Paris) 28(74): 98–129, pls.
14–21.
Braby, M. F., Y.-F. Hsu & G. Lamas. 2024. How to describe a
new species in zoology and avoid mistakes. Zoological Jour-
nal of the Linnean Society 2024: zlae043.
Viloria50
Davies, H. & C. A. Butler. 2008. Do butterflies bite?. Fascinating
answers to questions about butterflies and moths. New Bruns-
wick, NJ: Rutgers University Press, 240 pp.
Ferrer-Paris, J. R. 2000. Der Genus Redonda (Lepidoptera: Sa-
tyrinae): Fallstudie über Anpassung und Verbreitung in den
Páramos von Venezuela. Bayreuth: Bayreuth Universität, [ii]
+ xiii + 111 pp. [Dipl Biol thesis]
Ferrer-Paris, J. R., A. Cardozo-Urdaneta & Á. L. Viloria. 2015.
Mariposa braquíptera de Bordón, Redonda bordoni. pp. 322.
In: Rodríguez, J. P., A. García-Rawlins & F. Rojas-Suárez
(eds.): Libro rojo de la fauna venezolana. [4 th ed.]. Caracas:
Provita y Fundación Empresas Polar.
Ferrer-Paris, J. R. & Á. L. Vitoria [sic]. [2004]. Mariposas al-
tiandinas (Lepidoptera: Nymphalidae, Satyrinae) y la con-
servación de los páramos en Venezuela. pp. 626–633. In:
Jaramillo, C. A., C. Castaño Uribe, F. Arjona Hincapié, J. V.
Rodríguez & C. L. Durán (eds.): Congreso Mundial de Pára-
mos. Memorias Tomo I. Bogotá: Conservación Internacional
Colombia.
ICZN [International Commission on Zoological Nomencla-
ture]. 1999. International Code of Zoological Nomenclature /
Code International de Nomenclature Zoologique. 4th ed. Lon-
don: The International Trust for Zoological Nomenclature,
xxx + 306 pp.
ICZN [International Commission on Zoological Nomencla-
ture]. 2012. Amendment of Articles 8, 9, 10, 21 and 78 of the
International Code of Zoological Nomenclature to expand
and refine methods of publication. Bulletin of Zoological No-
menclature 69: 161–169.
Łabno, R. 2007. Studenci piszą. Z kraju, gdzie lato trwa wiecz-
nie. Alma Mater. Miesiecznie Uniwersytetu Jagiellońskiego
(Krákow) 98: 104–107.
Lamas, G., Á. L. Viloria & T. W. Pyrcz. 2004. Tribe Satyrini,
Subtribe Pronophilina. pp. 206–215. In: Lamas, G. (ed.):
Checklist. Part 4A. Hesperioidea - Papilionoidea. In: Hep-
pner, J. B. (ed.): Atlas of Neotropical Lepidoptera. Vol. 4.
Gainesville, Fl.: Association for Tropical Lepidoptera / Sci-
entific Publishers.
Pyrcz, T. W. 2007a. Ewolucja motyli z rodzaju Redonda (Le-
pidoptera, Nymphalidae) ze szczególnym uwzględnieniem
przystosowań do wysokogórskich warunków środowisko-
wych Andów. pp. 40–41. In: Polskie Badania Środowiska
Przyrodniczo-kulturowego w Ameryce Łacińskiej [28–30 maja
2007]. Kraków: Instytut Botaniki im W. Szafera Polska Aka-
demia Nauk.
Pyrcz, T. W. 2007b. Ewolucja motyli z rodzaju Redonda (Le-
pidoptera, Nymphalidae) ze szczególnym uwzględnieniem
przystosowań do wysokogórskich warunków środowisko-
wych Andów. pp. 17–19. In: Ogólnopolski Kongres Zoologicz-
ny, Zmienność, Adaptacja, Ewolucja [12-16 września 2007]:
Olsztyn: Polskie Towarzystwo Zoologiczne, Uniwersytet
Warmińsko-Mazurski.
Pyrcz, T. W. 2010a. Wybrane zagadnienia z taksonomii, zoogeo-
grafii I ewolucji faun górskich na przykładzie grupy modelowej
motyli z plemienia Pronophilini (Nymphalidae). Olsztyn:
Wydawnictwo Mantis, 245 pp. + [iii].
Pyrcz, T. W. 2010b. Evolution of butterflies of the genus Re-
donda (Lepidoptera, Nymphalidae, Satyrinae), and their
adaptation to the high Andean environment. pp. 265–273.
In: Mirek, Z., A. Flakus, A. Krzanowska, A. Paulo & J. Woj-
tusiak (eds.): The nature and culture of Latin America. Review
of Polish Studies. Kraków: Szafer Institute of Botany, Polish
Academy of Sciences.
Pyrcz, T. W., J. Lorenc-Brudecka, A. Zubek, P. Boyer, M. C.
Gabaldón & J. Mavárez. 2017. Taxonomy, phylogeny and
distribution of the genus Steromapedaliodes sensu novo in the
Cordillera de Mérida, Venezuela (Lepidoptera: Nymphali-
dae: Satyrinae: Satyrini). Arthropod Systematics and Phylog-
eny 75(2): 195–243.
Pyrcz, T. W., J. Wojtusiak & Á. L. Viloria. [2007]. Pierwszy przy-
padek ewolucji nielotnošci wšród motyli dziennych (Rhopa-
locera): samica Redonda bordoni. pp. [26]. In: Laskowski, R.,
T. W. Pyrcz, J. Weiner & J. Wojtusiak: Dziesięciolecie badañ
bioróžnorodnošci Andów. Kurs ekologii tropikalnej w Wene-
zueli. Krákow: Muzeum Zoologiczne UJ i Instytut Nauk o
Šrodowiscu UJ.
Schwanwitsch, B. 1924. On the ground-plan of wing-pattern in
nymphalids and certain other families of the rhopalocerous
Lepidoptera. Proceedings of the Zoological Society of London
1924: 509–528, 4 pls.
Viloria, Á. L. 1998. Studies on the systematics and biogeography of
some montane satyrid butterflies (Lepidoptera). London: The
University of London (King’s College London) / The Natu-
ral History Museum, 493 pp. [Dr Phil thesis]
Viloria, Á. L. 2000. Estado actual del conocimiento taxonómico
de las mariposas (Lepidoptera: Rhopalocera) de Venezuela.
pp. 261–274. In: Martín-Piera, F., J. J. Morrone & A. Melic
(eds.): Hacia un Proyecto Cyted para el inventario y estimación
de la diversidad entomológica en Ibero-américa: PrIBES-2000.
m3m-Monografías Tercer Milenio, vol. 1. Zaragoza: Socie-
dad Entomológica Aragonesa.
Viloria, Á. L. 2005. Las mariposas (Lepidoptera: Papilionoi-
dea) y la regionalización biogeográfica de Venezuela. pp.
441–459. In: Llorente-Bousquets, J. E. & J. J. Morrone (eds.):
Regionalización biogeográfica en Iberoamérica y tópicos afines.
Primeras Jornadas Biogeográficas de la Red Iberoamericana
de Biogeografía y Entomología Sistemática (RIBES XII.I-
CYTED). México, D. F.: Las Prensas de Ciencias, Facultad
de Ciencias, UNAM.
Viloria, Á. L. 2008. Mariposa braquíptera de Bordón. Redonda
bordoni Viloria & Pyrcz 2003. pp. 278. In: Rodríguez, J. P. &
F. Rojas-Suárez (eds.): Libro rojo de la fauna venezolana. [3rd
ed.]. Caracas: Provita / Shell de Venezuela.
Viloria, Á. L. 2018. Obituario. Carlos Bordón: 1921-2012.
Anartia, Publicación del Museo de Biología de la Universidad
del Zulia 27: 87–94.
Viloria, Á. L., J. R. Ferrer-Paris, J. Camacho & M. Costa. 2015.
New satyrine butterflies from the Venezuelan Andes (Lepi-
Davies, H. & C. A. Butler. 2008. Do butterflies bite?. Fascinating
answers to questions about butterflies and moths. New Bruns-
wick, NJ: Rutgers University Press, 240 pp.
Ferrer-Paris, J. R. 2000. Der Genus Redonda (Lepidoptera: Sa-
tyrinae): Fallstudie über Anpassung und Verbreitung in den
Páramos von Venezuela. Bayreuth: Bayreuth Universität, [ii]
+ xiii + 111 pp. [Dipl Biol thesis]
Ferrer-Paris, J. R., A. Cardozo-Urdaneta & Á. L. Viloria. 2015.
Mariposa braquíptera de Bordón, Redonda bordoni. pp. 322.
In: Rodríguez, J. P., A. García-Rawlins & F. Rojas-Suárez
(eds.): Libro rojo de la fauna venezolana. [4 th ed.]. Caracas:
Provita y Fundación Empresas Polar.
Ferrer-Paris, J. R. & Á. L. Vitoria [sic]. [2004]. Mariposas al-
tiandinas (Lepidoptera: Nymphalidae, Satyrinae) y la con-
servación de los páramos en Venezuela. pp. 626–633. In:
Jaramillo, C. A., C. Castaño Uribe, F. Arjona Hincapié, J. V.
Rodríguez & C. L. Durán (eds.): Congreso Mundial de Pára-
mos. Memorias Tomo I. Bogotá: Conservación Internacional
Colombia.
ICZN [International Commission on Zoological Nomencla-
ture]. 1999. International Code of Zoological Nomenclature /
Code International de Nomenclature Zoologique. 4th ed. Lon-
don: The International Trust for Zoological Nomenclature,
xxx + 306 pp.
ICZN [International Commission on Zoological Nomencla-
ture]. 2012. Amendment of Articles 8, 9, 10, 21 and 78 of the
International Code of Zoological Nomenclature to expand
and refine methods of publication. Bulletin of Zoological No-
menclature 69: 161–169.
Łabno, R. 2007. Studenci piszą. Z kraju, gdzie lato trwa wiecz-
nie. Alma Mater. Miesiecznie Uniwersytetu Jagiellońskiego
(Krákow) 98: 104–107.
Lamas, G., Á. L. Viloria & T. W. Pyrcz. 2004. Tribe Satyrini,
Subtribe Pronophilina. pp. 206–215. In: Lamas, G. (ed.):
Checklist. Part 4A. Hesperioidea - Papilionoidea. In: Hep-
pner, J. B. (ed.): Atlas of Neotropical Lepidoptera. Vol. 4.
Gainesville, Fl.: Association for Tropical Lepidoptera / Sci-
entific Publishers.
Pyrcz, T. W. 2007a. Ewolucja motyli z rodzaju Redonda (Le-
pidoptera, Nymphalidae) ze szczególnym uwzględnieniem
przystosowań do wysokogórskich warunków środowisko-
wych Andów. pp. 40–41. In: Polskie Badania Środowiska
Przyrodniczo-kulturowego w Ameryce Łacińskiej [28–30 maja
2007]. Kraków: Instytut Botaniki im W. Szafera Polska Aka-
demia Nauk.
Pyrcz, T. W. 2007b. Ewolucja motyli z rodzaju Redonda (Le-
pidoptera, Nymphalidae) ze szczególnym uwzględnieniem
przystosowań do wysokogórskich warunków środowisko-
wych Andów. pp. 17–19. In: Ogólnopolski Kongres Zoologicz-
ny, Zmienność, Adaptacja, Ewolucja [12-16 września 2007]:
Olsztyn: Polskie Towarzystwo Zoologiczne, Uniwersytet
Warmińsko-Mazurski.
Pyrcz, T. W. 2010a. Wybrane zagadnienia z taksonomii, zoogeo-
grafii I ewolucji faun górskich na przykładzie grupy modelowej
motyli z plemienia Pronophilini (Nymphalidae). Olsztyn:
Wydawnictwo Mantis, 245 pp. + [iii].
Pyrcz, T. W. 2010b. Evolution of butterflies of the genus Re-
donda (Lepidoptera, Nymphalidae, Satyrinae), and their
adaptation to the high Andean environment. pp. 265–273.
In: Mirek, Z., A. Flakus, A. Krzanowska, A. Paulo & J. Woj-
tusiak (eds.): The nature and culture of Latin America. Review
of Polish Studies. Kraków: Szafer Institute of Botany, Polish
Academy of Sciences.
Pyrcz, T. W., J. Lorenc-Brudecka, A. Zubek, P. Boyer, M. C.
Gabaldón & J. Mavárez. 2017. Taxonomy, phylogeny and
distribution of the genus Steromapedaliodes sensu novo in the
Cordillera de Mérida, Venezuela (Lepidoptera: Nymphali-
dae: Satyrinae: Satyrini). Arthropod Systematics and Phylog-
eny 75(2): 195–243.
Pyrcz, T. W., J. Wojtusiak & Á. L. Viloria. [2007]. Pierwszy przy-
padek ewolucji nielotnošci wšród motyli dziennych (Rhopa-
locera): samica Redonda bordoni. pp. [26]. In: Laskowski, R.,
T. W. Pyrcz, J. Weiner & J. Wojtusiak: Dziesięciolecie badañ
bioróžnorodnošci Andów. Kurs ekologii tropikalnej w Wene-
zueli. Krákow: Muzeum Zoologiczne UJ i Instytut Nauk o
Šrodowiscu UJ.
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