ANARTIA, 25 (2013 “2015”): 95 - 160
ISSN: 1315-642X
New satyrine butterflies from
the Venezuelan Andes
(Lepidoptera: Nymphalidae)
Ángel L. Viloria
1
, José R. Ferrer-Paris
2
, Jesús Camacho
3
and Mauro Costa
4
1
Centro de Ecología, Instituto Venezolano de Investigaciones Cientí
fi
cas (IVIC),
Apartado Postal 20632, Caracas 1020-A, Venezuela. aviloria@ivic.gob.ve
2
Centro de Estudios Botánicos y Agroforestales, Instituto Venezolano de
Investigaciones Científicas (IVIC), Av. 8, c. 79-80, Maracaibo, Venezuela.
jferrer@ivic.gob.ve
3
Museo de Artrópodos, Facultad de Agronomía, Universidad del Zulia
(MALUZ), Apartado 526, Maracaibo 4011, Zulia, Venezuela.
jcamacho@fa.luz.edu.ve
4
Res. Las Cumbres, Av. Las Acacias, La Florida, Caracas 1010A, D. C., Venezuela.
mauro13x50@gmail.com
Abstract
Members of the speciose subtribe Pronophilina (Nymphali-
dae: Satyrinae) are Neotropical butterflies predominantly found in
cloud forests. A few species live exclusively in open habitats above
the tree line: paramo and puna biomes. This habit requires special
adaptation to particular physical and climatic conditions of the
oreal zone in the tropics. At least ten species of the endemic genus
Redonda Adams and Bernard inhabit the paramos of the Venezu-
elan Andes. We herein describe seven of them: R. castellana Viloria
and Camacho, n. sp., R. centenaria Viloria and Camacho, n. sp.,
R. chiquinquirana Ferrer-Paris, n. sp., R. frailejona Ferrer-Paris and
Costa, n. sp., R. lathraia Viloria and Camacho, n.sp., R. leukas-
mena Viloria and Camacho, n. sp., and R. lossadana Ferrer-Paris,
urn:lsid:zoobank.org:pub:03463167-9D6B-47B5-AA62-5B8F84369CA0
Viloria et al.
96 ANARTIA, 25 (2013 “2015”): 95 - 160
n. sp., and rank Redonda empetrus bolivari Adams and Bernard to
full specific status.
Keywords: Andes, biogeography, Diaphanos, paramo, Quaternary,
Redonda, Steromapedaliodes.
Nuevas mariposas satíridas de los Andes
Venezolanos (Lepidoptera: Nymphalidae)
Resumen
Los miembros de la riquísima subtribu Pronophilina (Nympha-
lidae: Satyrinae) son mariposas neotropicales que se encuentran
predominantemente en bosques nublados. Pocas especies viven ex-
clusivamente en ambientes abiertos, por encima de la ceja de mon-
taña: biomas de páramo y puna. Este hábito demanda adaptación
especial a condiciones físicas y climáticas particulares de la zona
oreal tropical. Al menos diez especies del género endémico Redonda
Adams y Bernard habitan los páramos de los Andes de Venezuela.
Describimos aquí siete de ellas: R. castellana Viloria y Camacho,
n. sp., R. centenaria Viloria y Camacho, n. sp., R. chiquinquirana
Ferrer-Paris, n. sp., R. frailejona Ferrer-Paris y Costa, n. sp., R.
lathraia Viloria y Camacho, n.sp., R. leukasmena Viloria y Cama-
cho, n. sp., R. lossadana Ferrer-Paris, n. sp., y elevamos Redonda
empetrus bolivari Adams and Bernard a estatus específico.
Palabras clave: Andes, biogeografía, Cuaternario, Diaphanos, pára-
mo, Redonda, Steromapedaliodes.
INTRODUCTION
In his monograph on the genus Pedaliodes Butler, Thieme
(1905) described the Venezuelan species Pedaliodes empetrus (pp. 95,
98-99, tbl. 1, fig. 7), based on a single specimen allegedly collected
in the “Schneebergen von Merida” [snowy mountains of Merida].
He however seemed to have doubts in placing it amongst other
north Andean representatives of the group. Pointing out its bigger
size, slightly different habitus, lighter colour, hindwings elongated
as compared to typical Pedaliodes, unique spatulate antennae and
long labial palpi, Thieme appropriately compared the underside
New butterflies from the Venezuelan Andes 97
wing pattern of his ‘P.’ empetrus with that of ‘P.’ albonotata Godman,
another endemic species of the Venezuelan Andes, and possibly a
closely related taxon. Forster (1964) successfully split Pedaliodes into
several new genera, placing P. albonotata under Steromapedaliodes,
but neglecting P. empetrus, apparently because he had no access to
the type material (one single male, originally in Thieme’s collec-
tion, which has not been located – see below). Huber’s study of
1973 revealed that the biotope occupied by empetrus is the open,
windswept paramo above 3200 m (oreal biome), but failed in ar-
bitrarily assigning the species to the genus Punapedaliodes Forster.
Any way, Huber, not being a butterfly taxonomist, perhaps based
its combination exclusively on the external appearance of the speci-
mens he caught, which is reminiscent of certain high elevation gen-
era from the Equatorial and southern Andes (Altopedaliodes Forster,
Punapedaliodes, Argyrophorus Blanchard). Later, in 1975 and 1977,
Adams and Bernard (1981) researched the butterflies of the cen-
tral Cordillera de Mérida, and obtained a new, good series, of this
“unusual” butterfly species. They studied the morphological struc-
ture of the male genitalia, wing venation and pattern, and some
ecological features of ‘Pedaliodes’ empetrus, finding enough synapo-
morphies to justify the diagnosis of the genus Redonda, to include
Thieme’s taxon and another subspecific one they described from
the southwestern portion of the central Mérida range. We herein
consider the latter as a separate species, Redonda bolivari Adams and
Bernard, stat. nov., based on morphological features, but also on
biogeographical inferences.
In the 1990s and the beginning of the following decade, during
a series of butterfly surveys conducted by the authors and collabo-
rators in the oreal biomes of the Venezuelan Andes (“páramos”,
in Spanish), several undescribed forms of Redonda were discovered
inhabiting separate allopatric paramo units, and contiguous para-
patric altitudinal zones. One species from the western Cordillera de
Mérida, Redonda bordoni Viloria and Pyrcz, was already described in
Viloria et al. (2003), who at that time suggested its female might be
the first known case of brachyptery in butterflies. More ecological
and distribution data of each member, so far detected, of the genus
Redonda have also been gathered along the last 25 years. Extensive
collecting by several people allowed for improving the understand-
ing of their geographical distributions (see Pyrcz 2010a, 2010b).
Viloria et al.
98 ANARTIA, 25 (2013 “2015”): 95 - 160
Distribution studies and biogeographic analyses of this genus will
be published separately.
Seven new species of Redonda are described in this work, and
brief information on their geographic distribution, behaviour and
affinities is presented and discussed for them and for the three taxa
previously known.
MATERIALS AND METHODS
Butterflies were collected by conventional methods (hand nets)
in several sectors of the Andes of Venezuela, between 1991 and
1995, 1999 and 2000, and 2001-2015. Observations on the ecology
of the butterflies and other field notes were taken in situ, during the
course of sampling activities.
The following is a list of localities (from Northeast to South-
west, with dates and names of collectors) from where type speci-
mens for this study were obtained:
– Páramo de Los Nepes, 2450-2850 m, Serranía del Cendé, Trujil-
lo-Lara States borderline [Dinira National Park]: 12.viii.1991,
Ángel L. Viloria, Jesús Camacho, Rosanna Calchi; ii.2000, Á.
L. Viloria, José Rafael Ferrer-Paris.
– Páramo de Las Rosas, 2800-3100 m, Serranía del Cendé, Trujil-
lo-Lara States borderline [Dinira National Park]: 20.viii.1991,
Jesús Camacho; 12/16.i.1994, Á. L. Viloria, J. Camacho, R.
Calchi; iii-1994, Á. L. Viloria, J. Camacho, Mauricio García,
Carlos Fernández; 8-viii-1995, M. García.
– Páramo del Cendé, 3100 m, Serranía del Cendé, Trujillo-Lara
States borderline [Dinira National Park]: iii.1994, Á. L. Vilo-
ria, J. Camacho, M. García, C. Fernández; 8.viii.1995, M.
García.
– Páramo de Jabón, 3000 m, Serranía del Cendé, Trujillo-Lara
States borderline [Dinira National Park]: 25.xii.1989, CEUM
[Centro Excursionista de la Facultad de Agronomía, Universi-
dad Central de Venezuela, Maracay].
– Páramo de La Cristalina, 2800 m, north of Niquitao Massif,
Trujillo State: 14.v. 1986; 14.vi. 1986, John Edwin Lattke.
New butterflies from the Venezuelan Andes 99
– Páramo de Las Moras, 3000 m, north of Niquitao Massif, Tru-
jillo State: 19/20.ii.2007, Mauro Costa.
– Páramo de Ortiz, 2850-3100 m, upper Río Castán, north of
the Niquitao Massif, Trujillo State: 12.ix.1991, Á. L. Vilo-
ria, Rosanna Calchi, Edwin Moscó; 7.i.1992, Á. L. Viloria,
J. Camacho; 16.viii.2003, Á. L. Viloria, Mariana Alarcón,
Wilmer Rojas, Giovanny Fagua, Mónica Higuera.
– Páramo de Cabimbú, 2850-2950 m, Serranía de Niquitao (Niqui-
tao Massif), Trujillo State: 11-ix-1991; Á. L. Viloria, R. Calchi.
– Teta de Niquitao (base), 3325 m, Serranía de Niquitao (Niqui-
tao Massif), Trujillo State: 23.ii.2000, Á. L. Viloria, J. Cama-
cho, J. R. Ferrer-Paris, Freddy García.
– Páramo de Tuñame, 3100-3200 m, between Tuñame and Las
Mesitas, south of the Niquitao Massif, Trujillo State (bor-
derline with Mérida State): 10.ix.1991, Á. L. Viloria, R. Cal-
chi; 15.viii.2003, Á. L. Viloria, Tomasz W. Pyrcz, Mariana
Alarcón, Wilmer Rojas, Giovanny Fagua, Mónica Higuera;
16.ii.2010, M. Costa.
– Páramo de Santo Domingo, 3000-3300 m (including Hotel Los
Frailes and Laguna Victoria), Serranía de Santo Domingo, on
the road between Santo Domingo and Apartaderos, Mérida
State [P. N. Sierra Nevada, in part]: 5.ix.1971, Herbert Huber;
26.ix.1997, Andrew F. N. Neild; 12.iv.2004; 31.iii.2012, M.
Costa; 25.iii.2015, Cecilia Lozano, Leandro Morán.
– Páramo de Mucubají, 3400-3600, upper Serranía de Santo Do-
mingo (including Laguna de Mucubají and Laguna Negra),
Mérida State [P. N. Sierra Nevada]: 28.iii.1992; 20.iv.1992,
T.W. Pyrcz; 25.i.2000, 7 and 16.ii.2000, 13 and 20-21.iii.2000,
J. R. Ferrer-Paris; 1.i.2002, J. R. Ferrer-Paris, Bertha Condori;
14.ix.2004, J. R. Ferrer-Paris, M. Alarcón; 25.iii.2015, J. R.
Ferrer-Paris, Leinny González.
– Páramo de La Culata, 3300-3400 m, upper Río Mucujún (includ-
ing Valle del Muerto), Serranía de La Culata, N of Mérida city,
Mérida State [P. N. La Culata]: 30.i.2000, J. R. Ferrer-Paris, Á.
L. Viloria; 15.ii.2000, J. R. Ferrer-Paris; ii.2008, Pierre Boyer.
– Páramo de Los Conejos, 3400-4300 m, upper Río Albarregas (in-
cluding Pico Campanario, 4300 m), Serranía de La Culata,
Viloria et al.
100 ANARTIA, 25 (2013 “2015”): 95 - 160
NW of Mérida City, Mérida State [P. N. La Culata]: 10.ix.1938,
Tracey; 29.viii.1971, H. Huber; 20.vi.1975; 15.viii.1977, Mi-
chael J. Adams, George I. Bernard; 31.viii.2001, J. Camacho.
– Laguna Verde-Laguna del Suero, 4000 m, above La Mucuy, SE
of Mérida City, Sierra Nevada de Mérida, Mérida State [P. N.
Sierra Nevada]: 12.ii.1985, CEUM.
– Loma Redonda, 3900-4000 m, S of Mérida City, Sierra Nevada
de Mérida, Mérida State [P. N. Sierra Nevada]: 21.iv.1975, M.
J. Adams, G. I. Bernard; 8.ii.2007, T. W. Pyrcz.
– Páramo La Negra, 3200 m, southwestern Cordillera de Mérida,
Mérida State [P. N. Juan Peñaloza]: 30.ix.1951, P. Fenjues;
14.i.1982, Carlos Bordón.
– Páramo El Batallón, 3000-3800 m, southwestern Cordillera de
Mérida, Mérida-Táchira States borderline (including Sumu-
sica, Laguna El Cenegón, Laguna Grande, Pico El Púlpito) [P.
N. Juan Peñaloza]: 26/28-ii-1994, Á. L. Viloria, J. Camacho,
M. García; 16-xii-1994, M. García; 12/14-i-1995, J. Camacho,
M. García; 2/4.iii.1996, J. Camacho, M. García, Janusz Woj-
tusiak, T. W. Pyrcz; 12-13.ii.2000, Á. L. Viloria, J. Camacho,
M. García, J. R. Ferrer-Paris; 2.iv.2000, J. R. Ferrer-Paris.
We set, labelled and examined the butterflies, dissected their
wings and genitalia, and made ink drawings by means of standard
methods, using different kinds of WILD and Leica stereomicro-
scopes and adapted cameras lucidas. Photographs were obtained
with a Canon Camera EOS Rebel 3Ti, with a 100 mm macro lens,
using ring flash lights and several other kind of artificial illumina-
tion like fixwed fluorescent and led ring and conventional reflector
lamps, in the laboratories of the Departament of Biology, Faculty
of Sciences and the Faculty of Agronomy of the University of Zulia
in Maracaibo, and the Centre of Ecology of the Venezuelan Insti-
tute for Scientific Research, Altos de Pipe. Types and other speci-
mens examined are deposited in the following private and public
(institutional) collections (abbreviations cited in text):
AFN: Collection of Andrew Neild, London, England;
BMNH: The Natural History Museum, London, England; IVIC:
Instituto Venezolano de Investigaciones Científicas –Centro de Es-
tudios Botánicos y Agroforestales, Maracaibo, Venezuela; JFLC:
Collection of Jean-François LeCrom, Bogotá, Colombia; MALUZ:
New butterflies from the Venezuelan Andes 101
Museo de Artrópodos, Facultad de Agronomía, La Universidad
del Zulia, Maracaibo, Venezuela; MC: Collection of Mauro Costa,
Caracas, Venezuela; MIZA: Museo del Instituto de Zoología Agrí-
cola, Facultad de Agronomía, Universidad Central de Venezuela,
Maracay, Venezuela; MPUJ: Museo Javeriano de Historia Natural
Lorenzo Uribe, S.J., Pontificia Universidad Javeriana, Bogotá, Co-
lombia; MUSM: Museo de Historia Natural, Universidad Nacional
Mayor de San Marcos, Lima, Perú; MZUJ: Muzeum Zoologiczne
Instytutu Zoologii Uniwersytetu Jagiellonskiego, Kraków, Poland;
ZMHU: Zoologisches Museum Humboldt Universität, Berlin, Ger-
many; ZSBS: Zoologische Staatssammlung München, Germany.
Comparative morphological examinations of preserved butter-
flies and their structures, and their descriptions, were performed at
several stages in the Museo de Biología de La Universidad del Zu-
lia, Maracaibo, Venezuela (MBLUZ: 1991-1995, 1999-2000, 2014),
MALUZ (1991-1995, 2012, 2014), BMNH (1995-1998, 2001, 2003,
2011), MZUJ (1997), ZSBS (1997), the Universität Bayreuth, Ger-
many (1999-2000), and most recently in the Centro de Estudios Bo-
tánicos y Agroforestales (Maracaibo, 2011-2015) and the Centro de
Ecología (Altos de Pipe, 2000-2015), IVIC.
We present our results as taxonomic hypotheses based on com-
parative morphology and biogeography, coupled with the interpre-
tation of phylogenetic essays already published for some members
of the genus Redonda. Our recognition of the specific status of each
butterfly taxa described or treated in this study follows the intuition
of such hierarchy through cummulative evidence: observation and
homological comparison of colour pattern of wings and palpi, as
well as morphological features of different structures of the male
genitalic armature. Our previous experience in the systematics of
satyrine butterflies indicates that closely related species among the
Pronophilina may show subtle but constant morphological differ-
ences with respect to the characters evaluated, although there is
some degree of individual variation, that sometimes is validated
by bionomic information derived from field observations. We have
also followed intuitive interpretation of the discrete distribution of
the taxa (both in altitude and latitude) in the Venezuelan Andes, in
agreement with the evolutionary model of speciation of Michael J.
Adams (1977, 1985), which one of the authors of this contribution
tested through the methods of cladistics and found to be accept-
Viloria et al.
102 ANARTIA, 25 (2013 “2015”): 95 - 160
able, coherent, suitable and reliable as a working hypothesis (Viloria
1998). Adams’ model allows for rational explanations of the diver-
sity and distribution in allopatry, parapatry and sympatry of the dif-
ferent species of Redonda in the Venezuelan Andes.
The ten taxa herein studied are all considered fully developed
species, unable to hybridize among them in natural conditions. Some
very similar, apparently closely related species, inhabit discrete allo-
patric and parapatric areas of paramo that are geographically very
close to each other (for instance, in the páramos of Trujillo state), and
some others that are otherwise morphologically different, can live in
contiguous elevation zones (for instance, in the páramo El Batallón,
Mérida-Táchira states), and are therefore parapatric or even partly
sympatric in some very narrow ecological bands of contact. At least
one case of wind drift recorded in this study indicates that there may
be certain degree of accidental sympatry among species that are in a
strict sense, endemic to separate, disjunct geographic ranges. All of
this sum up to evidence that peculiarities of geographic distribution
within this genus support on one side the assumption that our taxa
are species on their own right and not geographic races (subspecies),
and on the other, that their evolutionary development may have been
mediated not only by historical geographic isolation of their high al-
titude habitats but also by ecological factors intrinsic to each species,
most of which remain unknown.
RESULTS
Genus Redonda Adams and Bernard, 1981
[Pedaliodes Butler; Thieme, 1905: 95, 98-99 (in part)]
[Punapedaliodes Forster; Huber, 1973: 195 (misidentification)]
Redonda Adams and Bernard, 1981: 367-368.
Redonda Adams and Bernard; Adams, 1985: 38; d’Abrera 1988: 871; Viloria,
1990: 218, 219; 1993: 69; 1994: 180, 184, 185; 1997: 15; 1998: 7, 14, 24, 37, 77, 78,
91, 95, 97, 99, 108, 124, 126, 182, 318-320, 409, 414, 415, 427, 445, 464, 467 (tbl.
1), 476 (fig. 1, distribution); 2000: 266, 269, 271; 2002: 178 (tbl. V), 190; 2005: 450;
2007: 1, 2, 9; Viloria et al., 1993: 226-227; Pyrcz, 1999: 354; 2004a: 289, 295, 296;
2004b: 570, 616; 2007a: 40, 41; 2007b: 17-19; 2008: 126; 2010a: 12, 13 (fig. 17, an-
tennal clubs), 38, 45 (fig. 35, females venation), 55, 69, 87 (fig. 82, cladogram), 88,
91, 109, 113, 116, 126, 171, 180 (fig.131, wing area), 181 (figs. 132, dimorphism;
134, habitats), 183, 184 (fig. 136, distribution map),185 (fig. 137, potential distribu-
New butterflies from the Venezuelan Andes 103
tion), 186, 207 (putative hostplants), 214; 2010b: 265, 266 (fig. 1), 267, 268 (fig. 2,
distribution), 270 (fig. 4, habitats), 271 (fig. 5), 272, 273; Ferrer-Paris, 2000: i-xiii, 1,
6, 8, 9, 10, 12, 13, 15, 17, 19, 20, 27 (fig. 3.1), 28, 30, 32, 36, 38, 39 (fig. 3.6), 40 (tbl.
3.5), 47, 48 (fig. 3.11), 49, 50 (tbl. 3.9), 51 (fig. 3.13), 52, 63, 66, 67, 68, 69, 70, 78
(fig. A.5), 81, 89 (tbl. C.5), 94-96 (tbl. C.9); Viloria and Pyrcz, 2001: 1, 2, 5, 6, 12,
15, 17; Ferrer-Paris and Viloria, 2002: 138-139; [2004]: 626, 627, 628 (fig. 1, clado-
gram), 629, 630 (fig. 3, wing area), 631; Viloria et al., 2003: 21, 22 (figs. 1, 2), 23
(fig. 3), e-appendices: [6]; 2007: [23], [24]; Lamas et al., 2004: 215; Orellana, 2004:
5-54; Bálint and Wojtusiak, 2006: 283; Shou et al., 2006: 105; Silva Dias, 2006: 21;
Pyrcz et al., 2009: 508, 525; 2014: suppl. material: [2 (tbl. S1)]; Pyrcz and Viloria,
2007: 46, 47 (tbl. 3); Posso Duque et al., 2010: [185]
[“Pedaliodes” Butler; Descimon, 1986: 510]
Redononda [sic]; Pyrcz and Fratello, 2005: 211.
Type species: Pedaliodes empetrus Thieme, 1905, by original
designation.
Revised diagnosis. Butterflies of medium size, some females
rather smaller than males; colour pattern could be sexually dimor-
phic as well; females strongly sedentary (except for that of R. leukas-
mena, n. sp., which is almost as active in flying like males). Antenna
to two-fifths costa. Antennal club heavily spatulated and concave.
Eyes densely hairy. Palpi twice as long as head; first segment one-
quarter the length of the second; third segment three-quarters the
length of the first. Forewing subtriangular; hindwing suboval, lon-
ger than in Pedaliodes Butler, outer margins rounded and regular,
sometimes slightly scalloped in hindwing (wings generally much
narrower in dimorphic females). Androconial patches absent. Ocel-
lar elements present in postdiscal area of both wings (either dor-
sally or ventrally, or both), but simplified as white spots, which are
sometimes elongated and highly distorted (fusiform, v-shaped, or
in the form of a fleur-de-lys); ventral wing pattern highly cryptical
for grass or sandy habitats, sometimes stereomorphic in the sense
of Schwanwitsch (1938); upperside wing pattern with conspicuous
marks (grey or white) developed in discal area at different degrees.
Venation (Fig. 1): Forewing Sc, R1 and R2 all independent, R3, R4,
R5 originated from common root at distal third of wing; R3 half
nearer to root of R4 than to discal cell, no r-m1, but M1 emerg-
ing from R3-5 (totally independent in Steromapedaliodes Forster
(Fig. 2), Dangond Adams and Bernard (Fig. 3) and Paramo Adams
and Bernard (Fig. 4)) m1-m2 curves inwards, vestigial veinlet (only
in males) entering discal cell in the middle of m1-m2 (closer to root
Viloria et al.
104 ANARTIA, 25 (2013 “2015”): 95 - 160
Figs. 1-4. Male wing venation of the type species of four oreal pronophi-
line butterfly genera endemic to the Northernmost Andes and the Sierra
Nevada de Santa Marta; 1. Redonda empetrus (Thieme, 1905),Venezuela,
Cordillera de Mérida, Río Albarregas, N of Merida, 3400 m (wing prep.
ALV27-1997, in BMNH); 2. Steromapedaliodes albonotata (Godman, 1905),
Venezuela, Cordillera de Mérida, Río Albarregas, N of Merida, 3400 m
(wing prep. ALV07-1997, in BMNH); 3. Dangond dangondi Adams and
Bernard, 1979, Venezuela, Sierra de Perijá, S slope of Cerro Avión, 3450
m (wing prep. ALV11-1997, in BMNH); 4. Paramo oculata (Krüger, 1924),
Colombia, Sierra Nevada de Santa Marta, Cambirumeina, 3950 m –dark
areas represent androconial patches (wing prep. ALV12-1997, in BMNH).
New butterflies from the Venezuelan Andes 105
Figs. 5-7. Female wing venation, and comparative shape and size propor-
tions of selected species of Redonda Adams and Bernard; 5. R. bordoni Vi-
loria and Pyrcz, 2003 (paratype, wing prep. ALVSN0-1997, in MALUZ);
6. R. centenaria Viloria and Camacho, n. sp. (paratype, wing prep. AL-
VSN1-1997, in MALUZ); 7. R. leukasmena Viloria and Camacho, n. sp.
(paratype, wing prep. ALVSN2-1997, in MALUZ).
of M2 in Paramo, right at base of M2 in Dangond, absent in Stero-
mapedaliodes); hindwing humeral vein moderately developed, broad-
ened at extreme (unlike Paramo and Dangond, where it is not broad-
ened, and Steromapedaliodes, where it is vestigial); root of Cu1 closer
to that of M3 than that of Cu2; root of M3 halfway between those
of Cu1 and M2; A2 and A3 independent, m1-m2 long (twice as
long as cu1-cu2; shorter in Steromapedaliodes, Paramo and Dangond)
and moderately curved inwards; discal cell about half the length of
the hindwing. Male genitalia: tegumen bubble-domed, and well dif-
ferentiated from uncus; uncus strong and slightly hooked, as long
as tegumen; gnathi absent or vestigial; saccus short and globular;
aedeagus generally straight, or slightly contorted, relatively short (as
long as tegumen + uncus, or slightly longer in R. bolivari Adams and
Bernard, n. stat.), never totally symmetrical; valvae broad, subtrian-
gular, simple, moderately hairy and devoid of ornamentation.
Comments. Redonda comprises a group of, so far ten, high el-
evation-specialized satyrine butterflies species. A genus endemic to
the easternmost branch of the northern Andes, in Venezuela. Most
of its species are distributed discretely in separate, archipelago-like,
Viloria et al.
106 ANARTIA, 25 (2013 “2015”): 95 - 160
high elevation paramo units (above 2700 m) of the Cordillera de
Mérida, northwest of the Táchira Depression. It has been men-
tioned as a putative example of rapid radiation in small isolated
areas under extreme environmental conditions (see Viloria 1998,
Ferrer-Paris 2000, Viloria et al. 2003, Ferrer-Paris and Viloria 2004,
Pyrcz 2010a, 2010b).
SPECIES ACCOUNT
Redonda bolivari Adams and Bernard, 1981, n. stat.
Figs. 8, 9, 10,11, 12, 13 (males), 46, 47 (male genitalia)
Redonda empetrus bolivari Adams and Bernard, 1981: 368-369, fig. 28;
d’Abrera, 1988: 871, fig. (identification erroneously given in caption as Diapha-
nos huberi Adams and Bernard); Viloria, 1998: 319 (in part misidentification of R.
empetrus); 2000: 269, 270 (last, misidentification of R. empetrus); 2002: 190; 2005:
459; Ferrer-Paris, 2000: 27 (fig. 3.1, distribution [in part misidentification of R.
empetrus]), 36, 37 (fig. 3.5), 91-92 (tbl. C.7); Viloria et al., 2003: 22 (fig. 1c, male), 23
(fig. 3); Ferrer-Paris and Viloria, [2004]: 629 (tbl. 1), 630 (fig. 3); Lamas et al., 2004:
215; Pyrcz, 2007b: 18, 19; 2010a: 87 (fig. 82, cladogram), 180 (fig. 131, wing area),
181(fig. 134B, habitat), 184 (fig. 136, distribution), 244; 2010b: 266 (fig. 1J male),
267, 268 (fig. 2, distribution), 270 (fig 4C, habitat), 272.
[Redonda bordoni Viloria and Pyrcz, 2003: 21-24, e-appendix A: pp. [1-2]
(misidentification, in part)].
Type locality: 4000 m, [paramo of] Loma Redonda, Cordillera
de Mérida [northern slopes of the Sierra Nevada de Mérida, and
south of the city of Mérida, Mérida State], Venezuela. Established
by Adams and Bernard (1981).
Comments. Our recognition of Redonda bolivari as a species
in its own right obeys to several sources of evidence. It is the only
species of the genus known to have an aedeagus longer (average
1.3 times) than the combined length of its tegumen + uncus. None
of the other species of Redonda reach 1.0 in the aedeagus/tegumen
+uncus ratio. Additionally, the aedeagus of R. bolivari is more ro-
bust and straight than in any other species so far recognized within
the genus (notably shorter and thinner, and more asymmetrical in
R.empetrus). Its saccus is also more robust than that of R. empetrus,
and definitely the longer within the genus; the valvae are more ob-
long than in R. empetrus. All individuals of this species which we
have examined are steadily large in wingspan (unlike R. empetrus).
New butterflies from the Venezuelan Andes 107
The wings of Redonda bolivari are basally broader and with a more
rounded profile than those of R. empetrus. Males of R. bolivari (like
in R. bordoni Viloria and Pyrcz and R. leukasmena Viloria and Cama-
cho, n. sp., which are quite distinctive in wing pattern) have no scal-
loping on the hindwing margins. These three species have coinci-
dentially the largest and fastest males. The dark colour of R. bolivari,
as well as its notably reduced ocellar elements are stable characters
(along a broad geographic distribution). Its upperside tends to lack
one of the diagnostic characters established by Thieme (1905) for R.
empetrus: a well defined dark discal stripe on the forewing. Morpho-
logical variation among males cannot be well ascertained due to the
few specimens known, but some individuals have a more marked
distinction between the basal (darker) and the distal half (lighter) of
the forewing on both sides. Females have not yet been discovered,
as they are inconspicuous and probably, much smaller than males,
flightless, and brachypterous, as in some other species of the genus.
The natural area of distribution of Redonda bolivari is the Sierra
Nevada de Mérida (south of the city of Mérida), a mountain range
that contains the highest elevations of Venezuela (reaching nearly
5000 m elevation). All Sierra Nevada records come from the up-
permost level of the paramo between 3900 and 4000 m, around the
highest prominences, the peaks Humboldt and Bonpland (Laguna
Verde-Laguna Suero) and the peaks Bolívar and Espejo (Loma Re-
donda). However, the paramo biome of this area is extensive and
continuous from an unknown ecological limit near the Mucubají
area (Sierra de Santo Domingo, northeast of Mérida) to the Los
Nevados area, right to the south of Mérida. It might well reach
the paramos of Acequias and San Pedro, as well as those ones in
the region of the Pueblos del Sur. This supposition is based on the
eventual finding of two aged, badly worn-out and isolated male
individuals of Redonda bolivari (Figs. 10-13, 47) in the páramo del
Batallón (3200-3400 m) in February 1994 (together with 24 males
and 2 females of R. bordoni, and 2 males of R. lathraia Viloria and
Camacho, n. sp.). This region is orographically and ecologically
separated from the paramos of the Sierra Nevada de Mérida by ca.
60 km of distance across territories whose maximum elevations are
too low for developing paramo biome. El Batallón is the natural
habitat of two endemic, possibly allopatric taxa, Redonda bordoni
and R. lathraia,n. sp., and the two butterflies above referred, cer-
Viloria et al.
108 ANARTIA, 25 (2013 “2015”): 95 - 160
Figs. 8-13. Males of Redonda bolivari Adams and Bernard, 1981, n. stat.;
8. Loma Redonda, Sierra Nevada de Mérida (upperside); 9. Ibid. (undersi-
de); 10. Worn-out specimen (1) from Páramo del Batallón (upperside); 11.
Ibid. (underside); 12. Worn-out specimen (2) from Páramo del Batallón,
13. Ibid. (underside).
tainly endemic to the Sierra Nevada de Mérida, may have reached
this area by action of the strong, trade winds of the dry season, that
notably affect the west of Venezuela between December and March.
This fact raises interesting questions about the possibility of even-
New butterflies from the Venezuelan Andes 109
Figs. 14-15. Adults of Redonda bordoni Viloria and Pyrcz, 2003; 14. Male
holotype (Páramo del Batallón), left upperside, right underside; 15. Female
paratype (same locality), left upperside, right underside.
tual hibridization or successful gene exchange between the taxa in-
volved in accidental meetings of local populations with windswept,
flying, foreign males relocated by force in areas where only females
of other species inhabit. However, we believe that gene flow, if any,
must be extremely small. We are inclined to think that mechanisms
of partner recognition may be different for each species, thus, pre-
cluding the possibility of interspecific mating, even when statisti-
cally plausible.
Material examined. VENEZUELA: 1 male, Estado Mérida, Parque Nacio-
nal Sierra Nevada, Laguna Verde-Laguna Suero, 4000 m, 12.ii.1985, CEUM leg.
(in MIZA); 1 male, Estado Mérida, Sierra Nevada de Mérida, Loma Redonda,
3900 m, 08.ii.2007 (genit. prep. ALV550-15) (in MC); 2 males, Estado Táchira,
Páramo El Batallón, entre Laguna El Cenegón y Laguna Grande, 3200-3400 m;
28.ii.1994; Á. Viloria, M. García and J. Camacho leg. (1 genit. prep. ALV544-15)
Viloria et al.
110 ANARTIA, 25 (2013 “2015”): 95 - 160
(in MALUZ. These unusual records are commented upon above);1 male, Estado
Mérida, S. of Mérida, Loma Redonda, 4000 m, 21.iv.1975, M. J. Adams, AB2,
holotype of Redonda empetrus bolivari Adams and Bernard (in BMNH).
Redonda bordoni Viloria and Pyrcz, 2003
Figs. 5 (female wing venation), 14 (male), 15 (female), 48
(male genitalia)
[Redonda bordoni Viloria and Pyrcz, MS, nom. nud.; 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 and 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 (fig. 6 a male genitalia, b
female genitalia, c female wing venation)] ) (in part misidentifications of R. bolivari
and R. lathraia, n. sp.]; [Anonymous], 2003: 24; Blackman, 2003: 26; Williams,
2003: R467; Lamas et al., 2004: 215; Viloria, 2005: 459; 2008: 278; Bálint and Woj-
tusiak, 2006: 288; Pyrcz, 2007a: 40, 41; 2007b: 17, 18, 19; 2010a: 36 (fig. 17C, an-
tennal club), 45 (fig. 35C, female venation), 87 (fig. 82, cladogram), 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;
³abno, 2007: 104; Hazel and Butler, 2008: 33; Ferrer-Paris et al., [in press]
Type locality: between Laguna El Cenegón and Laguna
Grande, Páramo El Batallón, Estado Táchira, Venezuela, 3200-
3400 m, by original designation.
Comments. Among individuals of the type series of Redonda
bordoni there were actually four males which represent two addi-
tional species (one partly sympatric). They were misidentified by
Viloria and Pyrcz (in Viloria et al. 2003). These four specimens were
collected, together with 24 males and two females of R. bordoni,
in an extense area of paramo in the neighbourhood of Laguna El
Cenegón (3100-3250 m) by Á. L. Viloria, J. Camacho and M. Gar-
cía on 27 february 1994. Two dark individuals, very much worn-out
by age and the action of wind are here unequivocally identified as
belonging to Redonda bolivari, n. stat. (see above for implications of
this discovery). The remaining two, formerly believed to be also a
melanic male form of R. bordoni, are herein recognized as belonging
to a new specific entity, Redonda lathraia Viloria and Camacho, n.
sp., and thus, correspondingly justified.
New butterflies from the Venezuelan Andes 111
In the year 2000 we observed males of R. bordoni in the area
of Sumusica (Táchira state), flying at 7 m/s wind speed and tem-
peratures under 10° C and with similar wind conditions but only
at 2° C in the region of El Cenegón. Their flight was erratical and
low, stopping sometimes on the vegetation with wings open and
oriented to the sun. Seldom, they were found resting motionless
in the shadowy side of some leaves. In some places, male indi-
viduals have been found on flowers, especially those of Berberis
sp. (Berberidaceae) and of ground herbs like Gnaphalium (Astera-
ceae). However, we have never seen them feeding on the flowers of
Espeletia (unlike R. castellana, n. sp., and R. centenaria, n. sp., from
the paramos of Trujillo).
One satyrine larva was discovered on Orthosanthus chimboracen-
sis (Kunth) Baker (Iridaceae), in the paramo of Sumusica, and it was
probably of Redonda bordoni: head capsule 4.10 mm wide, 4.02 mm
high; colour: brown with some dark pigmentation on the sides, no
dorsolateral horns. Body pale brown with black marks on both sides
of segments 4-9, body length when caught: 35.20 mm; ca. 0.5 mm
longer after a week in captivity. With a pair of tail appendices, 0.08
mm long, not specially coloured. It was fed in captivity on O. chim-
boracensis and bamboo leaves. It became inactive after two weeks
and apparently entered the praepupa stadium, but we were unable
to continue its rearing.
Material examined. VENEZUELA: 1 male, Estado Táchira, Páramo El
Batallón, entre Laguna El Cenegón y Laguna Grande, 3200-3400 m; 28.ii.1994;
Á. Viloria, M. García and J. Camacho leg., holotype of Redonda bordoni Viloria
and Pyrcz; 23 males (3 in MUSM, 3 in JFLC), 2 females, same data (1 wing prep.
ALVSN0-1997); 19 males (2 in MZUJ, 2 in BMNH), Estado Táchira, Páramo
El Batallón, entre la Antena CANTV y la Laguna El Cenegón, 3100-3250 m;
27.ii.1994; Á. Viloria, M. García and J. Camacho leg.; 4 males, Estado Táchira,
Municipio Jaúregui, Callejón del Cenegón, 16.xii.1994; M. García leg.; 6 males, 1
female, Estado Táchira, Parque Nacional Juan Peñaloza, Páramo El Rosal, 3000
m, 12/14.i.1995; J. Camacho and M. García leg.; 10 males, Estado Táchira, Pára-
mo El Batallón, Entre El Cenegón y Laguna Grande, 3000-3400 m, 05.iii.1996,
J. Camacho, M. García, T. Pyrcz, J. Wojtusiak leg., paratypes of R. bordoni Vilo-
ria and Pyrcz (all in MALUZ, except where indicated); 1 male, Estado Táchira,
Páramo de La Negra, 30.ix.1951, P. Fenjues leg.; 1 male, Estado Mérida, Páramo
de La Negra, 3200 m., 14.i.1982, C. Bordón leg., paratypes of R. bordoni Viloria
and Pyrcz (in 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 leg.; 5 males, Estado Táchira, Páramo El Batallón, Vía El Cenegón,
Viloria et al.
112 ANARTIA, 25 (2013 “2015”): 95 - 160
04.iii.1996, T. Pyrcz, J. Wojtusiak, J. Camacho, M. García leg., paratypes of R.
bordoni Viloria and Pyrcz (in MZUJ).
Redonda castellana Viloria and Camacho, n. sp.
Figs. 16, 17 (male), 18, 19 (female), 49 (male genitalia)
Synonymy, description, type material, distribution, comments,
and etymology, below.
Type locality: Venezuela: E[sta]do. Trujillo, Páramo de Ortiz,
3170 m, 9° 14’ 5.7 N, 70° 23’ 39.2 W. Herein established.
Redonda centenaria Viloria and Camacho, n. sp.
Figs. 6 (female wing venation), 20, 21 (male), 22, 23 (female),
50 (male genitalia)
Synonymy, description, type material, distribution, comments,
and etymology, below.
Type locality: Venezuela: Estado Trujillo, Páramo de Cabim-
bú, 2900 m. Herein established.
Redonda chiquinquirana Ferrer-Paris, n. sp.
Figs. 24, 25 (male), 26 (female), 51 (male genitalia)
Synonymy, description, type material, distribution, comments,
and etymology, below.
Type locality: Venezuela: Estado Mérida, [Páramo de] Mucu-
bají, 3600 m. Herein established.
Redonda empetrus (Thieme, 1905)
Figs. 1 (male wing venation), 27, 28, 29, 30, 31, 32 (males), 52,
53,54 (male genitalia)
Pedaliodes empetrus Thieme, 1905: 95, 98-99, pl. 1, fig. 7; Thieme; Weymer,
1912: 258, pl. 54, row e; Gaede, 1931: 489; Adams and Bernard, 1979: 99;Desci-
mon,1986: 506, 518; Viloria and Pyrcz, 2001: 3.
Punapedaliodes [Pedaliodes] empetrus (Thieme); Huber, 1973: 195.
Redonda empetrus empetrus (Thieme): Adams and Bernard, 1981: 368 (in part
misidentification of R. chiquinquirana, n. sp., female description of this taxon), figs.
13 (male genitalia), 27 (female of R. chiquinquirana, n. sp.); Adams, 1983: 474;
1984: 93; d’Abrera, 1988: 871 (in part misidentification of R. chiquinquirana, n.
sp.); 2001: 186, 341, pl. 141,fig. 17; Pyrcz, 1995: 523; 2007b: 17, 18; 2010a: 87
New butterflies from the Venezuelan Andes 113
(fig. 82, cladogram), 184 (fig.136, distribution), 244; 2010b: 266 (fig. 1I male), 267,
268 (fig. 2, distribution);Viloria, 1998: 10, 11, 13, 141, 142, 144, 146 (tbl. 1),147
(tbl. 2), 150 (fig. 23, male), 158 (fig. 72, wing venation), 170 (fig. 125, male geni-
talia), 320 (last, in part misidentification of other taxa, but fig. 23 correct); 2000:
269, 270; 2002: 190; 2005: 459; Ferrer-Paris, 2000: 27 (fig. 3.1, distribution [in part
misidentifications of R. bolivari and R. chiquinquirana n. sp.]), 29 (tbl. 3.1), 36, 37
(fig. 3.5), 38, 41 (tbl. 3.6), 91-92 (tbl. C.7); Viloria and Pyrcz, 2001: 12, 13; Viloria et
al., 2003: 22 (fig. 1(b) male); Ferrer-Paris and Viloria, [2004]: 627, 628 (fig. 1), 629
(tbl. 1), Lamas et al., 2004: 215.
Redonda empetrus (Thieme); Viloria, 1990: 223; 1998: 126, 176-181 (figs. 151-
157, cladograms), 343, 474 (tbl. 8); 2003: 247, 248 (figs. 1, 2, cladograms); Díaz
et al., 1997: 284, 811; Ferrer-Paris, 2000: x, 17, 30, 31 (fig. 3.2), 32(fig. 3.3), 33, 34
(tbl. 3.2), 35 (tbl. 3.3), 36 (fig. 3.4), 40, 41, 42 (tbl. 3.7), 43 (fig. 3.7), 44 (fig. 3.8), 45
(fig. 3.9), 46 (fig. 3.10), 47 (tbl. 3.8), 66, 69, 78 (fig. A.5, eggs and larvae, fig. A.6),
84 (tbl. C.1), 85-86 (tbl. C.2), 87 (tbl. C.3), 88 (tbl. C.4), 96 (tbl. C.9); Orellana,
2004: 5-57, 5-63, 5-69, 5-70, 5-1; Shou et al., 2006: 105; Viloria et al., 2007: [23];
Pyrcz, 2007a: 40, 41; 2010a: 46, 82, 134, 182, 183; 2010b: 265, 267, 272, 273; Peña,
2009: 102 (figs. 2, cladogram), 103 (fig. 3, cladogram), 105 (fig. 5, cladogram),
107 (fig. 8, cladogram), 111; Bolaños-Martínez et al., 2010: 89; Padrón Martínez,
2010: 19 (fig. 2-1, cladograms A, B), 26 (tbl. 2-1), 84; Peña et al., 2011: 70 (tbl. 1),
80 (fig. 6, cladogram).
[Redonda empetrus bolivari Adams and Bernard; Viloria, 1998: 319; 2000: 270;
Ferrer-Paris, 2000: 27 (fig. 3.1, distribution) (misidentifications, at least in part)].
Redonda empetrus georgei Pyrcz and Viloria, nom. nud.; Pyrcz, 2007b: 18, 19;
2010a:181 (figs. 132C, male; D, female; 133, female), 182, 184 (fig. 136, distribu-
tion), 244; 2010b: 272. Redonda emperatus [sic] (Thieme); Peña, 2009: 104(fig. 4,
cladogram).
Redonda empetrus ssp. 1; Pyrcz, 2010b: 266 (Figs. 1A male, 1B female), 267,
268 (fig. 2, distribution), 270 (fig. 4D, habitat), 271 (figs. 5C, female, 5D, pair in
copula).
Type locality: Herein restricted to the páramos of the upper
Albarregas river (páramo de Los Conejos, pico Campanario, 3100-
4300 m), westernmost extreme of the Serranía de La Culata, north
of the city of Mérida, Mérida State,Venezuela.
Original description, translated into English from Thieme
(1905), by Á.L.V. (diagnostic characters in bold letters):
“Ped. Empetrus n. sp. mine (Tbl. 1, Fig. 7)
A form of Pedaliodes between albonotata Godm. and albomaculata
Weymer, bigger than both, posterior [wing] prolongued, elongated
(as in albomaculata). Fringe of wings all pale white between veins,
anterior entire, posterior barely scalloped. Antennal club thick-
ened, claviform, palps longer than usual, without a scaled area.
Viloria et al.
114 ANARTIA, 25 (2013 “2015”): 95 - 160
Above diluted yellow instead of brown, not quite unicolor, but
brownish stripe, bent, outerly sharp, internally diffuse, anterior
proximal part [of the wing] separated from the distal part. Also
rotten-gray spot in the forewing median cell, crossed by transverse
veins of discal cell. Also posterior discal veins rake-like and grey-
ish. Anterior points (or spots) greater, six submarginal covertly
white, five posteriorly.
Underside anterior saturate brown, discal stripe the same as above,
but rather thin and sharp, submarginal points (spots) the same, as
above. Posterior not brown, but fuscus, all clouded, injected of ir-
regular spots of “dirty dog”. Five transverse submarginal signs co-
vertly white, like figures of the fleur-de-lys, which, that between
veins U R and O R, apart from the shape almost the same as in
Pedaliodes albonotata Godm.
Bigger than the others in the group. Conspicuous by the pale,
wood-tone upperside, which gets blurred by patch bands and
some dirty stains. It recalls albopunctata Weymer due to the ex-
tended, oblong hindwing, and albonotata Godm. (with which it
shares a common locality) by the underside submarginal design
of the hindwing.
The absence of the recurrent vein from the angle of the M D C
duly belongs to Pedaliodes, but the palps are exceptionally long,
the club thickened as a lobe, a case not known to me in any other
Pedaliodes.*)
*Related to the South American species of Oeneis (Argyrophorus
Blanch.) and recalls Argyrophorus Lamna mine (Berl. Ent. Zeitschr.
Vol. XLIX 1904 p. 160), in characters and design of the under-
side.”
A male in my collection from the snowy mountains of Merida.
Comments. The extraordinary monograph on the genus
Pedaliodes, published by Otto Thieme in 1905 appears to basically
be the result of Thieme’s overtaking of the work started by Otto
Staudinger, the prominent German entomologist and insect dealer,
who died in October 1900. It is evident that the aforementioned
research paper took several years of study of the material deposited
in three German and British collections, namely, Thieme’s own col-
lection, Staudinger collection (transferred in 1907 to the Humboldt
University Zoological Museum–Berlin), and the Walter Lionel
New butterflies from the Venezuelan Andes 115
Rothschild collection at Tring, England (later on, bequested to the
British Museum (Natural History), now the Natural History Muse-
um of London). A significant proportion of the new Pedaliodes spe-
cies described by Thieme had been, at least in part, previously iden-
tified as such by Staudinger. Thieme not only may have had access
to the Staudinger butterfly collection, but also to his unpublished
taxonomic notes. Along his monograph, Thieme duly mentioned
Staudinger’s manuscript names several times, and sometimes he
even applied those names to his new species. This is not the case for
‘Pedaliodes’ empetrus, whose description was based on just one male
specimen, preserved in the private collection of Thieme (black and
white photographic image of its upperside illustrated in Thieme’s
plate 1, figure 7). This individual might have come to Thieme’s
possession via some insect dealer based in Berlin; most probably,
Ernst A. Böttcher, and it seems unlikely that Staudinger had ever
known of this taxon. The type specimen of ‘P.’ empetrus allegedly
came from the snowy mountains of Mérida, but Mérida (1600-1800
m) was, and still is, a small city that lies in the river Chama valley,
between two, parallel, very high mountain chains that were both
‘snowy’ in that time, the second more than the first: the northern
Serranía de La Culata (with 54 peaks above 4300 m, maximum el-
evation at Pico Piedras Blancas, 4737 m) and the southern Sierra
Nevada (with 14 peaks above 4300 m, maximum elevation at Pico
Bolívar, 5010 m). A detached portion of the latter, the Sierra de San-
to Domingo, contains the other two peaks in Venezuela that reach
above 4300 m (maximum elevation at Pico Mucuñuque, 4660 m)
(Silva León 2001).
These geographic considerations are relevant because part of
Thieme’s type material of Pedaliodes sensu lato (a dozen taxa), includ-
ing his only individual of ‘P.’ empetrus, is currently lost or has not yet
been located. Rumors spread about the disappearance or loss of his
collections after WWII, but a note published in a German magazine,
the second half of the last century (Gerardo Lamas, pers. comm.),
indicates that his collection, or part of it, was purchased by Kurt
Beuthan after Thieme’s death (in 1907). Beuthan was apparently an
amateur entomologist established in Weissenfels, where he was, at
some point, the Director of the local historical museum. Beuthan’s
entomological collection –or the largest part of it– was auctioned in
1963. The whereabouts of this potentially very valuable collection
Viloria et al.
116 ANARTIA, 25 (2013 “2015”): 95 - 160
is unknown. In May 2003, two of the authors (Á.L.V. and J.R.F.-
P.) visited the Heimat-Naturgarten in Weissenfels, where there still
exist 20+ entomological drawers containing butteflies, moths and
other insects of the Kurt Beuthan collection. However, there are no
neotropical butterflies among them. Consequently, we have been
forced to consider unavailable for this study the type specimen of
Pedaliodes empetrus Thieme.
In the high mountains around Mérida we recognize four spe-
cies of Redonda, all of them show certain degree of morphological
variability and are externally rather similar to each other. They all
live in paramo environments characterized by open grasslands with
rosettes of Espeletia and/or related genera, that develope as a biome
generally around 3000 m and above, just below the periglacial zone.
However, only some male individuals from the Serranía de La Cu-
lata match well the diagnostic characters highlighted in the transla-
tion of the original description above. The best examples of large
imagos with the characteristic shape of lily flowers (fleur-de-lys) in
the submarginal ocellar elements of the hindwing underside come
from the upper Albarregas river (páramo de Los Conejos and sur-
roundings of the Pico Campanario). Additionally, the recto of those
specimens show the closest similarity to the upperside of the type il-
lustrated by Thieme, especially the shape and size of the submargin-
al white dots of the hindwing. Male individuals of the easternmost
population in La Culata (upper Mucujún river) are very variable in
size (FWL: 25-30 mm) and colour pattern. For instance, the series
we have collected and/or examined from the upper Mucujún river,
represents a range of various degrees of grey dusted over the up-
perside of the forewing cell, and show great variation in length and
shape of the white ocellar elements of the submarginal area of the
hindwing underside (Fig. 27-32). Nevertheless, there are specimens
in the Mucujún population that are indistinguishable from those
from the Albarregas, and have, for example, a well developed “fleur-
de-lys pattern”. We have not been able to detect the diagnostic fea-
tures selected from Thieme’s original description of ‘P.’ empetrus,
either in the Sierra Nevada individuals of Redonda (R. bolivari) or in
those from the Sierra de Santo Domingo (altitudinal parapatrics R.
frailejona, n. sp. and R. chiquinquirana, n. sp.). Therefore, we inter-
pret the identity of Redonda empetrus as applicable to those individu-
als of Redonda from the Serranía de La Culata, but restrict its type
New butterflies from the Venezuelan Andes 117
locality to the area where its phenotype appears to be more stable,
the upper Albarregas river.
We believe the type specimen of ‘P.’ empetrus was obtained by
Salomón Briceño Gabaldón at the end of the XIX Century or begin-
nings of the XX in the Serranía de La Culata, region of the Páramo
de Los Conejos. He was the only active, local butterfly collector and
dealer in the Andes of Venezuela during the time that the first par-
amo endemic satyrine butterflies reached Europe and were conse-
quently described: ‘Oxeoschistus’ opalinus (Staudinger, 1897) [Lecto-
type from Mérida, Bricenno [sic], in ZMHU, examined], ‘Pedaliodes’
albonotata (Godman, 1905) [Lectotype from Mérida, Venezuela, Ex
Staudinger, in BMNH, examined]. His collecting localities in the
Andes around Mérida (La Culata, La Pedregosa, Quintero, etc.)
were all located north of the Chama river. There is no evidence that
he, or any other entomologist before 1905, had collected butterflies
in the highest elevations of the Sierra Nevada de Mérida or in the
Sierra de Santo Domingo, which are south or southeast of the city,
and were far more inaccessible than the Serranía de La Culata (see
Viloria and Pyrcz 2001: 2-3).
Male specimens of R. empetrus from the region of the Páramo
de Los Conejos (upper Albarregas river) westernmost portion of the
Serranía de La Culata, are in average larger (29-30 mm in FWL,
mean 29.5 mm) than those from the upper Mucujún river (25-29.5
mm in FWL, mean 27.06 mm).
Adams and Bernard (1981: 368) obtained specimens of Redon-
da from the Mucubají lagoon area (Serranía de Santo Domingo).
From this material, they described under the name of R. empetrus the
female of R. chiquinquirana, n. sp. (op. cit., fig. 27). One specimen lo-
cated in the Adams and Bernard collection, which one of us (Á.L.V.)
have examined at the BMNH, is unlabelled. It certainly comes from
the Mucubají area (Serranía de Santo Domingo) as it fits the pheno-
type of females collected later on in Mucubají (illustrations in Viloria
et al. 2003: fig. 1B, and Pyrcz 2010b: fig. 1D; both also misidentified).
Females of putative Redonda empetrus from the upper Mucujún river
population have been illustrated by Pyrcz under a different nomi-
nal identity (2010a: 181, figs. 132D, 133; 2010b: 266, fig. 1B, 271,
figs. 5C, D). They are different from R. chiquinquirana, n. sp. Pho-
tographic illustrations show that their antennae are approximately
Viloria et al.
118 ANARTIA, 25 (2013 “2015”): 95 - 160
the same length as those of their corresponding males, reaching only
half length of the costa (vs. 2/3 in males). Thus, females are about
3/4 the size of males. Their wing upperside is metallic yellowish,
almost golden, and their underside is cryptic and similar in general
pattern to the males, but lighter and with reduced ocellar elements.
The few female specimens known did not come from the restricted
type locality herein established by us. They are all in foreign private
collections and were not available for this study. Thus, the female of
Redonda empetrus remains formally undescribed.
Some bionomic notes of this species have been presented by
Adams and Bernard (1981). In wild conditions they behave more
or less similar to other species belonging to its genus. We observed
small groups of males of R. empetrus in the upper Mucujún river
very active under extreme climatic conditions; some were seen at 8
o’clock in the morning, as the first sun rays penetrated the valley of
the river at around 3400 m elevation (refugio Casa de Piedra, on the
way from La Culata to Pico Pan de Azúcar) as the thermometer in
the shading side of the valley marked -3° C.
Material examined. VENEZUELA: 1 male, [Edo.] Mérida, Sierra [de]
La Culata, Páramo [de] Los Conejos, Laguna [de] Los Conejos, 8° 41’ 51” N,
71° 12’ 01” W, 3950 m, 31.viii.2001, J. Camacho leg. (genit. prep. ALV553-15)
(in MALUZ); 1 male, Estado Mérida, N de Mérida, Río Albarregas, 3400 m,
20.vi.1975, M. J. Adams and G. I. Bernard leg. (in MIZA); 2 males, Venezuela,
Edo. Mérida, [Serranía de La Culata] P. N. La Culata, Valle del Muerto (páramo)
[upper río Mucujún], 08° 44’ N, 71° 11’ W, alt. 3350 m, 15.ii.2000, 11:00-12:00,
J. R. Ferrer-Paris leg.; 4 males, same locality, 3325 m, 30.i.2000, 12:00-12:30 (in
IVIC); 1 male, Venezuela, Edo. Mérida, Serranía de La Culata, alto río Mucujún,
3400 m, 01.ii.2008, P. Boyer leg. (genit. prep. ALV551-15); 1 male, Venezuela, Edo.
Mérida, La Culata, 3300, 21.ii.2008, [P. Boyer leg.] (genit. prep. ALV549-15) (in
MC); 1 male, Est. Mérida, Campanario, 13500 ft, 10.ix.1938, J. H. Tracey leg., Brit.
Mus. 1939-114, 2 males, Cordillera de Mérida, N of Mérida, Río Albarregas, 3400
m, 15.viii.1977, M.J. Adams and G. I. Bernard leg., AB2 (1 genit prep. ALV088-
96, wing prep. ALV27-97); 1 male, same data, 3600 m (in BMNH); 1 male, Edo.
Mérida, Tal des Río Albarregas, 2100 m [sic], 29.viii.1971, H. Huber leg. (in ZSBS).
Redonda frailejona Ferrer-Paris and Costa, n. sp.
Figs. 33, 34 (male), 35, 36 (female), 37 (dimorphism), 55 (male
genitalia)
Synonymy, description, type material, distribution, comments,
and etymology, below.
New butterflies from the Venezuelan Andes 119
Type locality: Venezuela, Edo. Mérida, Sto. Domingo –
Apartaderos, 3000 m. Herein established.
Redonda lathraia Viloria and Camacho, n. sp.
Figs. 38, 39 (male), 56 (male genitalia)
Synonymy, description, type material, distribution, comments,
and etymology, below.
Type locality: Venezuela: [Estado] Táchira, P[ára]mo El
Batallón, entre Laguna El Cenegón – Laguna Grande, 3200-3400
m. Herein established.
Redonda leukasmena Viloria and Camacho, n. sp.
Figs. 7 (female wing venation), 40, 41 (male), 42, 43 (female),
57 (male genitalia)
Synonymy, description, type material, distribution, comments,
and etymology, below.
Type locality: Venezuela, Estado Lara, Municipio Morán,
Páramo de La Rosa [sic], 3000 m. Herein established.
Redonda lossadana Ferrer-Paris, n. sp.
Figs. 44, 45 (male), 58 (male genitalia)
Synonymy, description, type material, distribution, comments,
and etymology, below.
Type locality: Venezuela, [Edo.] Trujillo, Páramo de Tuñame,
vía Las Mesitas, 3100 m. Herein established.
DESCRIPTIONS
Redonda castellana Viloria and Camacho, new species
Figs. 16, 17 (male), 18, 19 (female), 49 (male genitalia)
urn:lsid:zoobank.org:act:971ED3B8-355D-4B8F-B718-3BC7EC48591B
[Redonda empetrus ssp. nov. 1; Viloria, 2000: 270 (misidentification) (in part)].
[Redonda empetrus n. ssp.; Viloria and Pyrcz, 2001: 12 (misidentification) (in
part)].
[Redonda empetrus ssp. nov.; Ferrer-Paris, 2000: 27 (fig. 3.1, distribution), 29
(tbl. 3.1), 36, 37 (fig. 3.5), 38, 69, 91-92 (tbl. C.7); Viloria et al., 2003: 23 (fig. 3, wing
Viloria et al.
120 ANARTIA, 25 (2013 “2015”): 95 - 160
Figs. 16-19. Adults of Redonda castellana Viloria and Camacho, n. sp.;
16. Male holotype (Páramo de Ortiz), upperside; 17. Ibid., underside; 18.
Female paratype (Páramo de Las Moras), upperside; 19. Ibid., underside.
area); Ferrer-Paris and Viloria, [2004]: 629 (tbl. 1), 630 (fig. 3) (misidentification)
(in part)].
[Redonda empetrus c) [n. ssp.] Viloria and Pyrcz, MS]; Lamas et al., 2004: 215
(misidentification) (in part)].
[Redonda empetrus ssp.; Viloria, 2005: 459 (misidentification) (in part)].
[Redonda empetrus millenaria] nom. nud.; Pyrcz, 2007b: 18 (misidentification
in part).
[Redonda empetrus centenaria Viloria and Pyrcz] nom. nud.; Viloria, 1998: 320;
Pyrcz, 2010a: 36 (fig. 17A, antennal club), 100 (fig. 86F, male), 180 (fig. 131, wing
area), 182 (fig. 135, polimorphism), 184, 184 (fig. 136, distribution), 244 (misiden-
tifications in part)].
[Redonda empetrus ssp. 4; Pyrcz, 2010b: 268 (fig. 2, distribution) (misidentifi-
cation) (in part)].
Male. Forewing length: 22.5-29 mm; mean: 26.87; n = 167.
Eyes very hairy, dark coffee brown, almost black. Palpi twice the
length of head, whitish in basal segment; silvery white scales all
New butterflies from the Venezuelan Andes 121
along, sparsed, setae predominantly black, inserted with ochra-
ceous. Antennae to two fifths of costa. Body moderately hairy, up-
perside black, underside similar in thorax, but shiny brown on abdo-
men. Forewing triangular (outer margin notably more straight, less
convex than similar species R. centenaria, n. sp., its closest allopat-
ric); upperside uniform dark brown with ochraceous scales finely
and scarcely dusted all over, more densely along costa and over the
forewing discal cell and on its distal extreme; basal and discal areas
darker, but not very distinctly defined, a series of five to six minute
postdiscal yellowish dots (more or less marked in different individu-
als), from cell R5 to Cu2, fringes with alternate yellowish between
veins. Hindwing oval, only slightly scalloped, upperside as in fore-
wing but without a darker discal area; a series of three to five post-
discal yellowish dots (from R5 to Cu1). Forewing underside reddish
brown ground colour, more towards posterior portion of discal area;
outer half of the wing and distal quarter of discal cell densely scaled
with ocher, also along costa, but inserted with coffee brown; coffee
brown spotted alternate along costa and outer margin (fringe), and
limiting both sides of discal area, especially in anterior half. Hind-
wing underside brown, with dense dark coffee scaling mostly on its
basal half; discretly mottled on its distal half; postdiscal-submargin-
al series of five to six conspicuous white-to-yellowish, triangular to
>-shaped, spots (R5 to Cu1or Cu2); anterior in general larger and
variably more elongated than posterior, all more or less surrounded
by dark coffee brown. Genitalia illustrated in figure 49; character-
ized by a very short and very wide saccus (the shortest and widest
of all species so far known in the genus); uncus stylized (more than
in R. centenaria, n. sp.), subunci absent, and valva less deep in lateral
view than that of R. centenaria, n. sp.
Female. Forewing length: 21-27; mean: 22.83 mm; n = 32.
Palpi covered by scales creamy brown, setae shorter than in male,
light creamy brown and black. Smaller and slender than male (and
smaller and slender in average, and lighter in colour, than females
of R. centenaria, n. sp.). Differs from male in having narrower wings,
but forewing still notably triangular (as compared to females of R.
centenaria, n. sp., its closest species both morphologically and geo-
graphically). Upperside golden brown, darker in forewing and bas-
al third of hindwing (including discal cell); forewing lighter areas
along discal-postdiscal border, the distal extreme and the middle of
Viloria et al.
122 ANARTIA, 25 (2013 “2015”): 95 - 160
the discal cell; dots as in males. Forewing underside similar to up-
perside, reddish ochraceous scales along veins, mottled with coffee
brown along costa and on submargin in anterior half and apex of
wing. Hindwing underside reddish ochraceous background; mot-
tled with dark coffee brown on basal two thirds of wing and less in
the submargin; series of five triangular-to >-shaped yellowish spots
very conspicuous (R5 to Cu1), more or less surrounded by dark cof-
fee brown. All fringes yellowish between veins.
Holotype. Male, Venezuela: Estado Trujillo, Páramo de Ortiz,
3170 m, 9° 14’ 5.7 N, 70° 23’ 39.3 W, 16.viii.2003, Á. L. Viloria, M.
Alarcón, W. Rojas leg. (in MALUZ).
Paratypes. 57 males (4 MUSM, 2 MC), 15 females (4 MUSM,
1 MC), same data as holotype; 15 males (2 in MZUJ), 4 females
(1 in MZUJ), Venezuela: Estado Trujillo, Páramo de Ortiz, 2850-
3100 m; 12.ix.1991; Á. L. Viloria and E. Moscó leg.; 30 males (1 in
BMNH), 4 females, same locality, 3100 m; 7.i.1992; J. Camacho
and Á. L. Viloria leg.; (in MALUZ); 37 males, 7 females, same lo-
cality and date as holotype, G. Fagua, M. Higuera leg. (in PUJB);
3 males, Venezuela: Estado Trujillo, Vía Boconó-Páramo La Cris-
talina, 2400-2800 m, 14.vi.1986, J. E. Lattke leg.; 1 male, Venezu-
ela: Estado Trujillo, Páramo La Cristalina, 2800 m, 14.v.1986, J. E.
Lattke leg. (in MIZA); 1 male, Venezuela: Estado Trujillo, Páramo
Las Moras, 3000 m, 19.ii.2007, M. Costa leg.; 22 males, 2 females,
same locality and collector, 20.ii.2007 (in MC).
Distribution. 2800-3170 m. Paramo region between Trujillo
and Boconó, divide of the basins of rivers Castán and Burate, NE
of the Teta de Niquitao. This region comprises the paramos La Cris-
talina, Las Moras, El Atajo, Ortiz, Los Pozos, El Corazón and Filo
La Ovejera. It is just orographically separated from Cabimbú and
La Teta de Niquitao by a small disjunction W of the Páramo El
Corazón.
Comments. A few Swallenochloa clumps were found patchily
distributed only in the Páramo de Ortiz, mainly along creeks. The
butterflies did not appear to be associated with bamboos, but with
grasses. During a visit to the Páramo de Ortiz in the dry season, we
found aggregations of males of R. centenaria “puddling” at midday
and in the early afternoon in humid parts of the soil along a dirt
road, where they were protected from direct sunshine by the shade
New butterflies from the Venezuelan Andes 123
of the neighbouring hills. In such spots they were easy to collect.
Thirteen (out of fifteen) females captured in 2003, laid numerous
eggs free of glue (2 to 47 each, totaling 156), during handling. Eggs
are spherical, light green and shiny. Attemps to rear larvae from
such eggs in laboratory conditions have always been unsuccessful.
This butterfly co-exists with Diaphanos curvignathos Viloria and
Steromapedaliodes albonotata (Godman) (Satyrinae), Colias dimera
Doubleday, Catasticta chrysolopha spectrum Reissinger (Pieridae), Hy-
lephila sp. (Hesperiidae), and several species of high altitude Lycae-
nidae.
Etymology. We respectfully dedicate the name of this new
species of butterfly, Redonda castellana, n. sp., to our friend, Dr.
Rafael Ramón Castellanos; diplomat, academician and historian,
and a prolific writer who, with more than 70 published books, has
contributed much to literature and history of Venezuela, and to
the promotion of the Andean regional history and identity. He is a
native of Santa Ana de Trujillo (b. 1931), a village located not far
from the area of distribution of R. castellana, n. sp. Well known to
Venezuelan intellectuals, readers, and bibliophiles, Rafael Ramón
Castellanos has owned and managed the “Gran Pulpería del Libro
Venezolano” in Caracas for many years, one of the largest and most
important book shops (new, second hand and antiquarian) of Latin
America.
Redonda centenaria Viloria and Camacho, new species
Figs. 6 (female wing venation), 20, 21 (male), 22, 23 (female),
50 (male genitalia)
urn:lsid:zoobank.org:act:53550B6B-35EC-4E71-9943-E7D6604FAFC8
[Redonda empetrus ssp. nov. 1; Viloria, 2000: 270 (misidentification) (in part)].
[Redonda empetrus n. ssp.; Viloria and Pyrcz, 2001: 12 (misidentification) (in
part)].
[Redonda empetrus ssp. nov.; Ferrer-Paris, 2000: 27 (fig. 3.1, distribution), 29
(tbl. 3.1), 36, 37 (fig. 3.5), 38, 69, 91-92 (tbl. C.7); Viloria et al., 2003: 22 (fig. 1d,
male, female), 23 (fig. 3, wing area); Ferrer-Paris and Viloria, [2004]: 629 (tbl. 1),
630 (fig. 3) (misidentification) (in part)].
[Redonda empetrus c) [n. ssp.] Viloria and Pyrcz, MS]; Lamas et al., 2004: 215
(misidentification) (in part)]
[Redonda empetrus ssp.; Viloria, 2005: 459 (misidentification) (in part)].
[Redonda empetrus millenaria] nom. nud.; Pyrcz, 2007b: 18.
Viloria et al.
124 ANARTIA, 25 (2013 “2015”): 95 - 160
[Redonda empetrus centenaria Viloria and Pyrcz] nom. nud.; Viloria, 1998: 320;
Pyrcz, 2010a: 36 (fig. 17A, antennal club), 100 (fig. 86F, male), 180 (fig. 131, wing
area), 182 (fig. 135, polimorphism), 184, 184 (fig. 136, distribution), 244 (misiden-
tifications in part)].
[Redonda camachoi Viloria and Pyrcz, nom. nud.; Pyrcz, 2010a: 45 (fig. 35B,
female venation) (misidentification)]
[Redonda empetrus ssp. 4; Pyrcz, 2010b: 268 (fig. 2, distribution) (misidentifi-
cation) (in part)]
[Redonda empetrus ssp. 5; Pyrcz, 2010b: 266 (figs. 1G, male, 1H, female), 267,
268 (fig. 2, distribution), 270 (fig. 4B, habitat) (misidentification) (in part)]
Male. Forewing length: 25-30 mm; mean: 26.75 mm; n = 11.
Eyes hairy, black. Palpi twice as long as head; silver white at base,
coffee brown all along, with dusting of white scales, mixed setae
black, coffee and cream, more or less in the same proportion. An-
tennae reaching two fifths of costa. Body dark brown, light brown
on abdomen underside. Forewing subtriangular, with outer margin
notably convex (different from that of R. castellana, n. sp., which is
more straight, and from that of R. lossadana, n. sp., which is slight-
ly truncated towards apex). Upperside ground colour dark brown,
rather uniform on both wings; anterior half of discal area of fore-
wing diffusely bordered with lighter brown, white over transverse
veins closing the outer extreme of the discal cell, some light brown
dusting along forewing subcostal region, and very subtle on basal
half of both wings; forewing series of four to five postdiscal-sub-
marginal white dots (from R5 to M3 or Cu1); forewing fringes with
white dots between veins; hindwing series of submarginal white
dots usually reduced to two or three (on M1 and M3, sometimes on
M2). Hindwing more crenulated than those of R. castellana, n. sp.,
and R. lossadana, n. sp. Forewing underside ground colour reddish
brown, mottled with dark brown all over (more towards edges and
base); white speckling over distal half of the wing and along costal
and apical areas, as well as in the middle of discal cell, series of
white dots as on upperside, but dots larger (wider). Hindwing under-
side ground colour dark brown, heavily mottled and speckled with
white, and light brown all over, white mottling more intense on both
sides of discal area; series of five submarginal white >-shaped spots,
from R5 to Cu1; those on R5 and M1 twice as long as the other
three. Genitalia illustrated in figure 50; subunci vestigial.
New butterflies from the Venezuelan Andes 125
Female. Forewing length: 24-26 mm; mean = 25 mm; n = 2.
Smaller and slightly slender than male, but larger and darker than
females of R. castellana, n. sp. Palpi all covered with silvery white
scales, setae predominantly light brown, alternate with black. Wings
upperside golden brown, slightly darker forewing; all fringes dark
brown with white between veins. Forewing postdiscal area lighter,
forewing series of dots as in males, but relatively larger and less
distinct; hindwing series of dorsal dots >-shaped as on underside.
Underside pattern in general very similar to male, but ground colour
lighter, especially on the outer third of wings, mottling and speck-
ling more moderate; forewing submarginal white dots smaller.
Holotype. Male, Venezuela: Estado Trujillo, Páramo de
Cabimbú, 2900 m; 11-ix-1991; Á. L. Viloria and R. Calchi leg. (ge-
nit. prep. ALV548-15) (in MALUZ).
Paratypes. 9 males, 2 females (1 wing prep. ALVSN1-1997),
same data as holotype (in MALUZ); 1 male, Venezuela, Edo. Tru-
jillo, región de Niquitao, M.-N, Teta de Niquitao N 09° 07’, W 10°
24’, alt. 3325 m (páramo), 23.ii.2000, 10:30-12:00, J. R. Ferrer-Paris
leg. (in IVIC).
Distribution: 2900-3325 m. Redonda centenaria, n. sp., appears
to be restricted to the páramo surroundings of the Teta de Niqui-
tao (whose NW portion is called páramo de Cabimbú). This is a
relatively isolated high altitude unit in the state of Trujillo (Briceño-
Valero 1920; highest elevation in La Teta, 4006 m), just separated
from a northern portion of paramo biome where another, exter-
nally similar species occur (R. castellana, n. sp.). The latter paramo
unit divides the valleys of the cities of Trujillo and Boconó (see
distribution of R. castellana, n. sp.). Most of the páramos of the
Niquitao region, have a single, long, wet season (ca. April-August),
which is probably due to the influence of the climatic conditions of
the Llanos, south of the Andes (Monasterio and Reyes 1980). The
local vegetation is characterized by the predominance of Espeletia
and other asteraceous rosettes and bunch grasses (Vareschi 1970,
Cuatrecasas 1979).
Comments. Redonda centenaria, n. sp., is a common insect
throughout the year. Both males and females fly actively during sun-
ny periods. Females can be distinguished by their slightly smaller
size, lighter upperside coloration, and relative weakness in flight.
Viloria et al.
126 ANARTIA, 25 (2013 “2015”): 95 - 160
Figs. 20-23. Adults of Redonda centenaria Viloria and Camacho, n. sp.; 20.
Male holotype (Páramo de Cabimbú), upperside; 21. Ibid., underside; 22.
Female paratype (same locality), upperside; 23. Ibid., underside.
Usually they seem unable to sustain a long flight, which is under-
taken in “steps”, and very close to the grasses, most commonly fol-
lowing the direction of the wind. Resting females are very difficult
to see because of their cryptic underside pattern, but they can be
observed dropping suddenly into the grass after a short flight. Then,
they take another flight-step in the same way. When collected, fe-
males released many spherical eggs from the moment of capture
and throughout several hours whilst kept alive in paper envelopes.
All the eggs were also free of cement.
Etymology. La Universidad del Zulia (Maracaibo, Venezu-
ela), began academic activities on September 11
th
1891, under the
principalship of the notable lawyer Dr. Francisco Ochoa, first Rec-
tor. This new species of Redonda was first noticed by Á.L.V. in the
Páramo de Cabimbú, one hundred years later, on September 11th
1991. We name it R. centenaria in celebration of the centennial of
our Alma Mater. We also want to record our gratitude for the unique
New butterflies from the Venezuelan Andes 127
Figs. 24-26. Adults of Redonda chiquinquirana Ferrer-Paris, n. sp.; 24.
Male holotype (Páramo de Mucubají), upperside; 25. Ibid., underside; 26.
Female paratype (vía Laguna Negra), left upperside, right underside.
opportunity we (A.L.V. and J.C.) have had to develop our profes-
sional careers in this institution.
Redonda chiquinquirana Ferrer-Paris, new species
Figs. 24, 25 (male), 26 (female), 51 (male genitalia)
urn:lsid:zoobank.org:act:F56B4A32-2F3E-430E-9E53-C38A22EF1EE0
[Redonda empetrus empetrus (Thieme); Adams and Bernard, 1981: 363 (fig.
27, female), 368 (description of female); d’Abrera, 1988: 871; Viloria, 1998: 320;
Ferrer-Paris, 2000: 27 (fig. 3.1, distribution); Viloria et al., 2003: 21-23 (figs. 1(b),
[female], 2, 3); Ferrer-Paris and Viloria, [2004]: 630 (fig. 3) (misidentifications, at
least in part)]
Redonda empetrus mikei Pyrcz and Viloria, nom. nud.; Pyrcz, 2007b: 18; 2010a:
180 (fig. 131, wing area), 182, 184 (fig. 136, distribution), 244.
[Redonda empetrus ssp. 2; Pyrcz, 2010b: 266 (figs. 1C male, 1D female), 266,
268 (fig. 2) (misidentification)]
Male. Forewing length: 20.5-26 mm; mean = 24.76 mm, n =
23. Eyes hairy, black. Palpi twice the length of head, white at basal
segment, remaining segments black, with sparse white scales, black
Viloria et al.
128 ANARTIA, 25 (2013 “2015”): 95 - 160
and cream setae alternatively inserted more or less in the same pro-
portion. Antennae to two fifths of costa. Body black, only brown
laterally on abdomen. Forewing subtriangular, outer margin con-
vex, tornus rounded. Hindwing oval and distally elongated as in
R. empetrus, but base wider than the latter, outer margin scalloped.
Dorsal ground colour of wings brown (darker than in R. empetrus,
but lighter than in R. bolivari, R. castellana, n. sp., R. centenaria, n.
sp., and R. lossadana, n. sp.), slightly darker on basal half. Veins
on basal half of wings covered with whitish scales, very notable
on those closing discal cell. Forewing discal area limited by dif-
fuse white on both sides, more distinctly in forehalf, white speck-
ling along two thirds of costal area, distinct white between veins
along fringe, submarginal series of five to six circular dots, as large
as those of R. empetrus, and in general twice to three times the diam-
eter of most dark species from Trujillo (i. e., R. castellana, n. sp., and
R. centenaria, n. sp.) and Mérida (i. e., R. bolivari and R. lossadana, n.
sp.), in cells R3 (minute or missing), R5 to Cu1; forewing marginal
area darker. Hindwing upperside postdiscal-submarginal series of
five white, subtriangular dots, from R5 to Cu1, those in R5 and M1
double the length of the others. Genitalia in figure 51. It has a very
tiny, vestigial subunci attached to tegumen, valvae with distinctive
shape, as they are triangular with a sharp tip, a feature only shared
with Redonda lossadana, n. sp.
Female. Forewing length: 18-20.5 mm, n = unknown (Adams
and Bernard, 1981: 368); 1 female in MALUZ: 19 mm. It has al-
ready been described and illustrated by Adams and Bernard (1981:
363, 368), although they have misidentified it as R. empetrus. An
example is illustrated in figure 26.
Holotype. Male, Venezuela, Edo. Mérida, [Páramo de] Mucu-
bají, 3600 m, 11.x.2010, M. Costa leg. (in MIZA).
Paratypes. 2 males, same data as holotype (1 genit. prep.
ALV554-15) (in MC); 1 male, Venezuela, Edo. Mérida, P. N. Sierra
Nevada, Valle del Mucubají (páramo), 08° 45’ N, 70° 49’ W, alt.
3400 m, 25.i.2000, 11:30-12:30, J. R. Ferrer-Paris leg., 3 males, same
locality, 01.i.2002, 11:00-12:00, J. R. Ferrer-Paris, B. Condori leg.;
3 males, same locality, 14.ix.2004, 8:40-8:55, J. R. Ferrer-Paris, M.
Alarcón leg.; 11 males, Venezuela, Edo. Mérida, P. N. Sierra Ne-
vada, Quebrada de Mucubají, 2ª morrena de retroceso, 25.iii.2015,
New butterflies from the Venezuelan Andes 129
9:00-9:45, J. R. Ferrer-Paris, L. González leg.(in IVIC); 1 male, Ven-
ezuela, Estado Mérida, Parque Nacional Sierra Nevada, Laguna
Negra, 3550 m, 28.iii.1992, T. Pyrcz leg.; 1 male, 1 female, Estado
Mérida, Parque Nacional Sierra Nevada, vía Laguna Negra, 3600
m, 20.iv.1992, T. Pyrcz leg. (in MALUZ); 1 female [no data] [M.
J. Adams and G. I. Bernard, AB1] (in BMNH); 1 male, Venezu-
ela, Edo. Mérida, Zw Laguna Grande und Laguna Negra, 3450 m,
12.ix.1971, H. Huber leg. (in ZSBS).
Distribution. 3400-3600 m. Redonda chiquinquirana, n. sp., is
known from the upper paramos of the Serranía de Santo Domingo,
SE of Sierra Nevada de Mérida. It has been observed and captured
in the surroundings of the Laguna de Mucubají, Laguna Negra and
Laguna Grande. It is altitudinally parapatric with R. frailejona, n.
sp., which is found in a lower level of these paramos (see below,
under that species).
Comments. The flight activity of males of this species seems
to be restricted to short periods of the day, and also limited by ap-
parently constant good weather, as it is the case of most species
of the genus. Four females (not available for this taxonomic study)
were captured, at different days, in Mucubaji during a period of eco-
logical studies pursued in 2000. All were sampled from the ground
vegetation, where they were resting or basking with open wings.
Two of them tried to scape from the net by flying 2 to 3 m, before
hiding again inside the very abundant bunches of grass. Some of
the females kept in captivity laid several individual, loose eggs. They
were spherical, green to yellowish green when fresh, 1.01 ± 0.03
mm in diameter and 0.31 ± 0.03 mg in weight (N = 7). Some of
them became more yellowish or brown after 1 to 2 weeks but did not
hatch. Two larvae of Redonda chiquinquirana, n. sp., were found at
Mucubají, feeding on Calamagrostis sp. and Agrostis trichodes (Kunth)
Roem. and Schult., both Poaceae. These larvae were resting in the
centre of grass bunches. They were kept alive in laboratory boxes
and observed resting between the leaves of grass and paper sheets
during the day. They were active, feeding at night.
Synoptical description of the larvae: Head capsule 3.30 and
3.33 mm wide, 2.99 and 3.01 mm high respectively. Colour: dark
brown with black lines on both sides and the front, no dorsolateral
horns. Body dark brown with black stripes on both sides and on
Viloria et al.
130 ANARTIA, 25 (2013 “2015”): 95 - 160
the dorsum, body length when caught: 27,70 mm and 28,30 mm,
respectively, both with a pair of tail appendices, 0.1 mm long, not
especially coloured.
Etymology. J.R.F.-P. first noticed the occurrence of this new
taxon in the Laguna de Mucubají region in 2000, when pursuing
field work for his Biology Diplom Thesis on the ecology of the but-
terflies of the genus Redonda. He names it R. chiquinquirana after his
father, Dr. José Chiquinquirá Ferrer González –“Chinco” Ferrer–, a
petroleum engineer, Emeritus Professor of the Faculty of Engineer-
ing of La Universidad del Zulia, Venezuela, of which he was its
20
th
Rector (1984-1988). Chiquinquirana is a derivation of the second
name of Professor Ferrer, applied to this butterfly species as a token
of love and gratitude for his continuous and steady support along
the author’s (J.R.F.-P.) life and professional career.
Redonda frailejona Ferrer-Paris and Costa, new species
Figs. 33, 34 (male), 35, 36 (female), 37 (dimorphism), 55 (male
genitalia)
urn:lsid:zoobank.org:act:B5DA0BB7-5774-45C1-A3B2-ACC7E2F01917
Male. Forewing length: 24-29.5 mm; mean: 26.04 mm; n = 12.
Eyes hairy, black. Palpi twice as long as head, white at base, remain-
ing part with scattered white scales ventrally; black setae predomi-
nant over some inserted creamy brown ones. Antennae reaching two
fifths of costa. Body black, except for abdomen underside which is
brown. Forewing subtriangular, outer margin almost straight. Hind-
wing oval, elongated beyond the outer borders of forewing, outer
edge crenate. Upperside ground colour brownish, notably darker on
basal half, especially hindwing; fringes dark, white between veins.
Forewing upperside: distal half more or less discretly separated of
darker basal half by a sinuous division, dusting of whitish scales
along the outer part of this sinuous line, and along subcostal area;
light, whitish dusting above distal extreme and center of discal cell;
submarginal series of five to six large circular white dots with dif-
fuse borders on cells R4 (tiny, sometimes absent) to Cu1, regularly
aligned; costa dark brown. Hindwing upperside with a series of four
submarginal >-shaped white dots, from M1 to Cu1, those in M1
and M2 larger and more distinctly shaped; a similar dot in cell R5,
but discal in relation to the position of the others. Forewing under-
side ground colour ochraceous brown, mottled with dark brown on
New butterflies from the Venezuelan Andes 131
basal half, more distinctly at base and at the border of discal area,
distal half speckled with brown, fine dusting of dark brown and
white towards subapical and apical region; white dots as above, but
less distinct and less rounded; all veins dark in distal half of wing.
Hindwing underside ground colour coffee brown, dusted all over
with white, more densely in the outer half of wing, anal area, and
costa at inner border of discal region; white dots more distinctly
>-shaped, and more or less surrounded by coffee brown, their posi-
tions as above; white specks beside each dot, discal to them. Geni-
talia illustrated in figure 55; vestigial subunci attached to tegumen,
saccus tubular, in the shape of a barrel, aedeagus straight in dorsal
view.
Female. Forewing length: 13; n = 1. Much smaller and lighter
than male. It is so small, and their wings are so narrow, that it has
the aspect of a micromoth. Brachypterous and flightless. Palpi twice
as long as head, devoid of scales (naked), setae shiny golden at basal
third, black in remaining two-thirds of its length. Antennae reaching
two thirds of costa. Body black. Forewing slender, narrower than
hindwing, spatulate, slightly wider in distal half, outer margin con-
vex, costa convex, but almost parallel to anal margin, veins much
closer to each other than in males; hindwig slender and narrow, but
broader than forewing, spatulate, outer margin rounded, not scal-
loped. Fringes with tufts of long scales at the end of the veins. Up-
perside ground colour golden white, base of wings, forewing costa
(and basal half of costal region), and hindwing anal region very
dark brown, almost black. Hindwing bearing a lightly marked se-
ries of five >-shaped, submarginal white spots (R5 to Cu1), M3 and
Cu1 larger than the others. Forewing underside ground colour yel-
low; all mottled with dark chocolate brown, especially along coastal
border, centre of postdiscal area, discal area, and base; series of five
tiny, postdiscal, white dots. Hindwing underside groundcolour cof-
fee brown, speckled and mottled with white, especially on basal and
distal thirds, discal area darker; series of six submarginal white dots
from R5 to Cu2, those in M1, M2, M3 and Cu1 larger and with
characteristic >-shape, all more or less surrounded by dark coffee
brown.
Holotype. Male, Venezuela, Edo. Mérida, Sto. Domingo –
Apartaderos, 3000 m, 31.iii.2012, M. Costa leg. (in MIZA).
Viloria et al.
132 ANARTIA, 25 (2013 “2015”): 95 - 160
Figs. 27-32. Morphological (size and wing pattern) variability among adult
males of Redonda empetrus (Thieme, 1905) from the Serranía de La Culata
(SC); 27. Large individual (FWL: 30 mm) with typical wing colour pattern
as described and illustrated by Thieme, from the páramo de Los Conejos,
upper río Albarregas, SE-SC, upperside; 28. Ibid., underside; 29. Darker,
large size form (FWL: 29.5 mm), with reduced hindwing postdiscal oce-
llar elements, from the upper río Mucujún, NW-SC, upperside; 30. Ibid.,
underside; 31. Smaller individual (FWL: 25 mm) with more marked bluish
grey on the upperside of FW discal cell, from the upper río Mucujún, NW-
SC, same locality as 29-30; 32. Ibid., underside.
New butterflies from the Venezuelan Andes 133
Figs. 33-36. Adults of Redonda frailejona Ferrer-Paris and Costa, n. sp.;
33. Male holotype (Santo Domingo - Apartaderos, 3000 m), upperside;
34. Ibid., underside; 35. Female paratype (Hotel Los Frailes), upperside;
36. Ibid., underside.
Paratypes. 1 male, Venezuela, Edo. Mérida, valle de Santo
Domingo, cerca Hotel Los Frailes, 3000 m, 31.iii.2012, M. Costa
leg. (genit. prep. ALV552-15); 4 males, Venezuela, Edo. Mérida, Los
Frailes, 3100 m, 12.iv.2004, M. Costa; 1 female, same data, 3050 m
(in MC); 1 male, Venezuela, Edo. Mérida, Hotel Los Frailes, 3000
m, 25.iii.2015, 10:30-11:35, C. Lozano leg.; 5 males, Venezuela,
Edo. Mérida, Laguna Victoria, 3000-3050 m, C. Lozano, L. Morán
leg. (in IVIC); 7 males, Venezuela, Mérida, Apartaderos-Sto. Do-
mingo, km 12.5, ca. 3100-3300 m, 26.ix.1997, A. Neild (in AFN)
[not measured]; 19 males, Edo. Mérida, oberes Domingo tal, 3070
m, 5.ix.1971, H. Huber; 2 males, same data, 3050 m [not measured]
(in ZSBS).
Distribution. 3000-3300 m. Known only from a reduced sec-
tor of the Serranía de Santo Domingo (southern slopes). It is the
lower, altitudinally parapatric species of Redonda chiquinquirana, n.
sp., which flies in an upper level of those paramos.
Viloria et al.
134 ANARTIA, 25 (2013 “2015”): 95 - 160
Comments. There is very little bionomic information available
of this species, apart from what is known of its behaviour. Males are
strong flyers and the solitary female known in not capable of flight.
They represent the most outstanding case of sexual dimorphism
within the genus Redonda (Fig. 37).
Etymology. Frailejona, is a feminine derivation of “frailejón”,
the vernacular name applied in Venezuela, Colombia and Ecuador
to numerous species of asteraceous plants of the genus Espeletia
(and some related genera) which grow exclusively in the paramo,
and are characteristic elements of its vegetation adapted to high al-
titude North Andean environments. They have long, wooly, suc-
culent leaves that grow in a rosette-like pattern. Their flowers are
bright yellow, and are very attractive to the paramo butterflies and
other pollinator insects. “Frailejón” is possibly an archaic Castillian
Fig. 37. Proportional images, showing the most marked sexual dimor-
phism among representatives of the genus Redonda: R. frailejona Ferrer-
Paris and Costa, n. sp. (left: female paratype, right: male holotype).
37
New butterflies from the Venezuelan Andes 135
word equivalent to the most modern “fraile” (friar). The application
of this name to Espeletia plants may have derived from the appear-
ance of some columnar forms, that grow as tall as a person, looking
like static friars in the misty landscape of the paramo. Most speci-
mens known of R. frailejona, n. sp., were observed and captured in
the surrounding of the Hotel Los Frailes (The Friars), a well known
touristic destination on the road between Santo Domingo and the
Mucubají Lagoon (Mérida state), established in an emblematic co-
lonial building that once was the dwelling for some hermits, friars
of a religious, Catholic Order. Thus, the name is both related to the
frailejón plants and to the Hotel Los Frailes.
Redonda lathraia Viloria and Camacho, new species
Figs. 38, 39 (male), 56 (male genitalia)
urn:lsid:zoobank.org:act:F5C1ABEA-20EC-448A-A473-2EEC06B92D22
[Redonda bordoni Viloria and Pyrcz, 2003: 21-24, e-appendix A: pp. [1-2]
(misidentification, in part)].
[Redonda kenke Pyrcz and Viloria] nom. nud.; Pyrcz, 2007a: 40; 2007b: 17,
18, 19.
[Redonda veilloni Pyrcz and Viloria] nom. nud.; Pyrcz, 2010a: 36 (fig. 17B, an-
tennal club), 181 (fig. 132E, male; F, female), 182, 184 (fig. 136, distribution), 244.
[Redonda n. sp. Pyrcz and Viloria]; Pyrcz, 2010b: 265.
[Redonda sp.]; Pyrcz, 2010b: 266 (figs. 1M, male, 1N, female), 267, 268 (fig.
2, distribution).
Male. Forewing length: 27-28 mm; mean: 27.5; n = 2. Eyes
hairy, black. Palpi twice as long as head, brown, without scales,
Figs. 38-39. Adult male holotype of Redonda lathraia Viloria and Cama-
cho, n. sp. (Páramo del Batallón); 38. Upperside; 39. Underside.
Viloria et al.
136 ANARTIA, 25 (2013 “2015”): 95 - 160
setae dark brown and creamy brown alternate, some silvery setae
on ventral surface of basal segment. Antennae reaching two fifths
of costa. Body dark brown above, ventrally brown. Forewing tri-
angular, apex acute, outer margin slightly convex. Hindwing oval
not exceeding the extent of forewing (unlike R. bordoni, R. empetrus
and R. frailejona, n. sp.), outer margin slightly scalloped. Upperside
ground colour chestnut, finely dusted with brown scales all over, a
bit more dense above the distal extremity of forewing discal cell;
fringes dark chocolate brown; vestigial submarginal white dots on
forewing (only in paratype). Underside colour brown, finely dusted
with ochraceous yellow, more on anterior half of forewing. Apex
and anterior half of marginal region of forewing dusted with choco-
late brown; same colour softly speckled on the limit of discal and
postdiscal regions; a series of five submarginal white dots, small
and slightly oval, from cell R5 to Cu1. Hindwing underside with
chocolate brown along outer margin, a submarginal undulated, fine
line of the same colour, running parallel to margin; basal half to
two-thirds of hindwing heavily dusted of whitish, especially near
veins and along cell Cu2, which are dark brown; dark chocolate
brown mottled inside discal cell, also running parallel to both sides
of whitish line in cell Cu2, and forming > -shaped discal lines point-
ing basad in cells that also bear submarginal fusiform cream-white
stripe-dots: R5, M1, M2, M3, Cu1. Genitalia illustrated in figure
56. Uncus stout, not well differentiated from tegumen, valvae with
rounded tip, covered with very short setae, aedeagus relatively small.
Female. Unavailable for this study. Illustrated by Pyrcz 2010a
(pp.181, fig. 132F) and 2010b (pp. 266, fig.1N). The aforementioned
figures show that female is slightly smaller and lighter in colour than
male, but generally similar in colour pattern.
Holotype. Male, Venezuela: [Estado] Táchira, P[ára]mo El
Batallón, entre Laguna El Cenegón – Laguna Grande, 3200-3400
m; 28.ii.1994, Á. L. Viloria, M. García and J. Camacho leg. (in
MALUZ) [previously misidentified as Redonda bordoni Viloria and
Pyrcz, and designated as paratype of that species].
Paratype. 1 male, same data as holotype (genit. prep. ALV546-
15) (in MALUZ) [previously misidentified as Redonda bordoni, and
designated as paratype of that species].
New butterflies from the Venezuelan Andes 137
Distribution. 3200-3400 m. Apparently, an endemic to the
lower level of the open páramos of the La Negra-El Batallón range
(southwestern Cordillera de Mérida), where it is possibly an altitudi-
nal parapatric, and certainly, partly sympatric with Redonda bordoni
(3000-3800 m). Most of the area shared by these two endemic taxa
is protected by a National Park (Parque Nacional Juan Peñaloza).
The paramos occupied by R. bordoni and R. lathraia seem to be pris-
tine in a large extension, especially towards the eastern El Batallón
region (in Táchira State), which is wetter than the Páramo de La
Negra (in Mérida State), but we noticed some agricultural activi-
ties and cattle grazing entering the boundary zone of the National
Park in the páramo. This is increasingly depauperating the original
paramo biome in the region of the Pico El Púlpito (near Sumusica).
Vegetation in the Páramo El Batallón is dominated by a dense
cover of grasses such as Cortaderia spp. and Calamagrostis spp. (Poace-
ae) (“pajonal and pastizal paramero”, sensu Bono 1996). The “frail-
ejones” (Ruilopezia jahnii (Standl.), R. atropurpurea (A. C. Smith),
Asteraceae) are outstanding floristic elements all over this páramo,
and rosettes of these plants are profusely scattered from the very hu-
mid intermontane valleys (where glacial lagoons often occur) to dry
and rocky cliffs on the higher peaks. Nearly 20 lagoons have been
recorded in the Park, but there are many other small ones or even
swampy areas that seem to be remnants of recent glacial pools. The
vegetation is noticeably more diverse in the narrow humid valleys,
especially around lagoons or flooded areas, where bamboos (Swal-
lenochloa angustifolia (Soderstrom and Calderón), S. spencei (Ernst),
Poaceae) occassionally occur. There is also a high number of flow-
ering plants in these places (see Vareschi 1970 and Bono 1996).
Comments. We found Redonda lathraia occasionally entering
open grassland areas above the treeline, between 3200-3400. How-
ever, according to Pyrcz (2010b: 265) it is parapatric to R. bordoni,
occupying a lower elevation level of the paramo. Perhaps, its true
elevation preference may not be much higher than 3100 m. Higher
up, there is a total dominance of R. bordoni (up to Pico El Púlpito,
ca. 3950 m). We found only two males among very abundant indi-
viduals of R. bordoni, during the dry season, in February 1994. Even
at that time this páramo was apparently more windy, humid and
cloudy than the neighbouring ones of the Cordillera de Mérida. The
behaviour of males of both species is very similar. They are fast-
Viloria et al.
138 ANARTIA, 25 (2013 “2015”): 95 - 160
flying insects that drift with the wind, but are also capable of flying
strongly against it. Like all species of Redonda, and especially their
males, this one avoid attacks by stalling in flight, thus allowing the
wind to change their flight vector very suddenly.
Etymology. Two males of this butterfly species had been col-
lected in the páramo del Batallón in 1994. They were misidentified
as dark representatives of Redonda bordoni (Viloria et al. 2003), and
so, erroneously included among its type series. As a result of such
a confusion we decided to name this taxon with the epithet lathraia,
derived from laqraίoV, the greek adjective for clandestine, smug-
gled, or surreptitious.
Redonda leukasmena Viloria and Camacho, new species
Figs. 7 (female wing venation), 40, 41 (male), 42, 43 (female),
57 (male genitalia)
urn:lsid:zoobank.org:act:41B76B14-BD5C-4885-B795-159917FD309B
[Redonda sp. nov. 2]; Viloria, 2000: 270; Ferrer-Paris, 2000: 27 (fig. 3.1, dis-
tribution), 29 (tbl. 3.1), 36, 37 (fig. 3.5), 38, 69, 91-92 (tbl. C.7); Ferrer-Paris and
Viloria, [2004]: 628 (fig. 1), 629 (tbl. 1), 630 (fig. 3).
[Redonda n. sp.]; Viloria and Pyrcz, 2001: 15.
[Redonda sp. nov.]; Viloria et al., 2003: 22 (fig. 1e, male, female), 23 (fig. 3,
wing area).
[Redonda [n. sp.] Viloria and Pyrcz, MS]; Lamas et al., 2004: 215.
[Redonda sp.]; Viloria, 2005: 459.
[Redonda camachoi Viloria and Pyrcz] nom. nud.; Viloria, 1998: 319; Ferrer-
Paris, 2000: 96 (tbl. C.9); Pyrcz, 2007a: 40, 41; 2007b: 17, 18, 19; 2010a: 38 (fig.
22D), 45 (fig. 35A, female venation), 87 (fig. 82, cladogram), 111, 180 (fig. 131,
wing area), 181 (figs. 132G, male; H, female; 134A, hábitat), 182,183, 184 (fig.
136, distribution), 244; 2010b: 265, 266 (figs. 1O, male, 1P, female), 267, 268 (fig. 2,
distribution), 270 (fig. 4A, habitat), 273.
Male. Forewing length: 28-33 mm; mean: 30.13 mm; n = 27.
Eyes hairy, reddish brown, tiny black hairs. Palpi twice as long as
head, dark coffee brown, devoid of scales, with long black setae,
some sparse bright brown setae on dorsum. Antennae two fifths
length of costa, 37-38 segments; shaft orange-brown, with some
sparse hairs and light brown scales; club broadened and concave
(but less so than R. bordoni), keel running along the concavity of the
club. Body dark coffee brown, with exception of ventral surface of
abdomen, which is light creamy-brown; ventral surface of body very
hairy (less so than R. bordoni). Forewing triangular, tornus obtuse;
New butterflies from the Venezuelan Andes 139
hindwing oval, outer margin very slightly scalloped. Groundcolour
of dorsal surface of wings dark coffee brown, darker on discal area
of both wings; basal third of wings with greenish sheen; forewing
slightly hairy at base, showing sparse white scales along costal mar-
gin, and much more distinct on outer margin, between veins; po-
lygonal subdiscal silvery white patch within discal cell, distal edge
of which straight and well defined at second third of cell, basal edge
rather diffuse, following vein Cu2; hindwing lacking additional
markings except darker postdiscal line and some white scales flank-
ing margin in cells M1 and M3. Ground colour of forewing ventral
surface chocolate brown, becoming coffee brown towards costal
margin and apex; speckling of dark coffee brown over entire sur-
face of wing, denser in apical region and discal area, which appears
well limited at its edges; white scales dusted along costal margin
and apical region; two postdiscal white dots (one basal, the other
distal, separated by ca. 2 mm) in R5; another dot in M2, parallel to
that distal in R5; all of them irregularly surrounded by dark brown.
Groundcolour of hindwing ventral surface chocolate brown, en-
tirely speckled by black, notably denser on basal half of wing and
in submarginal region of each cell; series of five postdiscal creamy-
yellow >-shaped spots, in cells R5 to Cu1, all pointing basad and
surrounded by irregular “shadows” of dark coffee-brown; heavy
dusting of white scales on inner and outer edges of discal area, as
well as on distal margin and around veins; distal margin flanked
with white between the extremities of veins. Genitalia illustrated in
figure 57, shows a long and very stout uncus, lack of subunci; valvae
broad, with rounded tips, saccus wide but small.
Female. Forewing length: 26.5-29 mm; mean: 27.87 mm;
n = 4. It differs from male by its slightly smaller size and narrower
wings. More pointed forewing apex and tornus. Antennae 42 seg-
ments. Palpi devoid of scales (only very few ventral white scales at
base), brown, setae brown and coffee brown, less abundant than
in male. Dorsal surface of wings less dark and brownish, forewing
showing postdiscal series of 4-5 ochraceous, minute dots, in cells
R5 to Cu1 (sometimes missing in M2). Groundcolour of forewing
ventral surface brown, light brown at apex; postdiscal series of five
white dots in cells R5 to Cu1. Groundcolour of hindwing ventral
surface not chocolate, but pale brown, which gives higher colour
contrast to general design.
Viloria et al.
140 ANARTIA, 25 (2013 “2015”): 95 - 160
Holotype. Male, Venezuela: Estado Lara, Municipio Morán,
Páramo de La Rosa [sic], 3000 m, 20-viii-1991, J. Camacho leg. (in
MALUZ).
Paratypes. 6 males, 2 females (1 wing prep. ALVSN2-1997),
Venezuela: Estado Lara, Parque Nacional Dinira, Páramo La Rosa
[sic], 2800 m; 16-i-1994; Á. [L.] Viloria and J. Camacho leg.; 18
males (1 in BMNH, 1 in MZUJ), 2 females, Estado Lara, Parque
Nacional Dinira, Páramo Cendé, 3100 m; 8-viii-1995, M. García
leg. (in MALUZ); 2 males, Estados Lara-Trujillo, Páramo El Jabón,
3000 m, 25-xii-1989, CEUM leg. (in MIZA).
Distribution. 3000-3100 m. High and humid paramos of the
Cendé Region, easternmost extreme of the Venezuelan Andes,
between the States of Trujillo and Lara. This geographic unit in-
cludes the highlands or paramos of Agua de Obispos, Las Rosas,
Jabón, Guache, Naríz, Los Nepes, Cendé, and El Vigía, among oth-
ers (Briceño-Valero 1920). Monasterio (1980) has defined the main
vegetation there as “rosetal de Espeletia jabonensis” (Asteraceae) be-
cause of the remarkable dominance of this endemic plant, but R.
leukasmena is better associated with the equally common “pastizal
de Swallenochloa-E. jabonensis” (Poaceae-Asteraceae). These pára-
mos are highly influenced by the atmospheric conditions of the
Lake Maracaibo basin, which results in two highly pronounced wet
seasons (Monasterio and Reyes 1980). Most of the paramo area is
protected within the Dinira National Park.
Comments. This species has never been found abundant.
Males and females show similar behaviour, and it is almost impos-
sible to sex individuals in flight, because they share similar wing
patterns and size. Both sexes fly energetically, 1-2 m above the thick
bambusoid grasses (mainly Swallenochloa angustifolia (Soderstrom
and Calderón) and Rhipidocladum geminatum (McClure) (Poaceae),
Viloria 1994) that grow in the Cendé paramos. We failed to find R.
leukasmena in the dry, low Páramo de Los Nepes (2400-2850 m),
where bamboos seem to be partly replaced by tussock grasses such
as Agrostis and Calamagrostis (Poaceae). The first female collected
laid one egg immediately upon capture. It was spherical and pale
green, with no evidence of any adhesive on its surface.
Like most butterflies found at very high elevations in the north-
ern Andes, R. leukasmena only flies in sunshine and is very hard
New butterflies from the Venezuelan Andes 141
to see on the wing in either cloudy or foggy conditions. Then, the
adults tend to hide in the bamboo grasses, preferring the most inac-
cessible basal stems, where they are also thought to spend the night.
Nothing is known about its biology. However, from observations of
the biotopes occupied by the adults, we infer that their larvae feed
on bamboos throughout the paramo. Diaphanos fuscus Viloria and
Steromapedaliodes schuberti Viloria and Pyrcz are two more satyrines
of the paramos of the Cendé region. S. schuberti shows a similar
wing pattern and behaviour, which may suggest a possible mimetic
relationship with Redonda leukasmena, but they have not been found
flying together (see comments in Viloria and Pyrcz 2001).
Etymology. Redonda leukasmena owes its specific name to the
conspicuous white patches on the dorsal surface of its forewings,
which look like bleached portions of the very dark background.
Therefore, we designate this new species with the greek word
leukasmέna, which literally means bleached.
Redonda lossadana Ferrer-Paris, new species
Figs. 44, 45 (male), 58 (male genitalia)
urn:lsid:zoobank.org:act:23E43E42-CA82-4FC3-A37B-2EB9677F684B
[Redonda empetrus ssp. nov. 1; Viloria, 2000: 270 (in part misidentification of
R. centenaria centenaria, n. sp.)]
Redonda empetrus decenia Pyrcz and Viloria, nom. nud.; Pyrcz, 2010a: 83 (fig.
79A), 182, 184 (fig. 136, distribution), 244.
[Redonda empetrus ssp. 3; Pyrcz, 2010b: 266 (figs. 1E male, 1F female), 268
(fig. 2), 270 (fig. 5A male) (misidentification)]
Male. Forewing length: 24-28 mm; mean: 25.87 mm; n = 4.
Eyes hairy, dark coffee brown. Palpi twice the length of costa, white,
dusted with black laterally, setae coffee brown and light brown al-
ternate more or less in equal proportion. Antennae reaching two
fifths of costa. Body dorsally dark brown, ventrally dusted with
white scales, abdomen brown. Forewing subtriangular, apex acute,
outer margin convex. Hindwing oval, about same extent of fore-
wing, outer margin softly scalloped. Upperside ground colour dark
brown, finely dusted with ochraceous scales. Forewing discal area
darker (less ochraceous scales), but also limited on both sides by
more dense light dusting, whitish scaling above veins closing distal
extremity of discal cell; a series of tiny, almost vestigial, white, sub-
marginal dots, from R5 to Cu1; fringes dark, white between veins.
Viloria et al.
142 ANARTIA, 25 (2013 “2015”): 95 - 160
Figs. 40-43. Adults of Redonda leukasmena Viloria and Camacho, n. sp.
(Páramo de Las Rosas); 40. Male holotype, upperside; 41. Ibid., underside;
42. Female paratype, upperside; 43. Ibid., underside.
Figs. 44-45. Adult male holotype of Redonda lossadana Ferrer-Paris, n. sp.
(Páramo de Tuñame); 44. Upperside; 45. Underside.
New butterflies from the Venezuelan Andes 143
Hindwing unicoloured; white marginal dots in cell R5 (diffuse),
M1, M3 (diffuse). Forewing underside ground colour hazelnut, gen-
erally speckled with chocolate brown, more densely at both sides of
discal area; yellow scales dusted irregularly all over, mottled along
costal margin and in subapical region, just beyond the limit of discal
dark area; dense yellow and dark brown dusting inside discal cell;
tiny submarginal white dots as above, but continued with a series
of linear dots towards the outer edge of wing. Hindwing underside
ground colour brown, heavily mottled with dark coffee brown and
white; a series of white elongated dots (sometimes >-shaped), sub-
marginal, from R5 to Cu2, those in R5 and M1 twice the length of
the others, all surrounded by dark brown; white mottling especially
heavy on discal area, which has a lichenous aspect; a white, fine line
along the middle of cell Cu2; marginal area darker. Genitalia as il-
lustrated in figure 58, valvae with sharp tips.
Female. Unavailable for this study. Illustrated by Pyrcz (2010b:
266, fig. 1F). Similar in general appearance to the female of R. castel-
lana, n. sp., but slightly smaller and with broader and less pointed
forewing.
Holotype. Male, Venezuela, [Edo.] Trujillo, Páramo de Tu-
ñame, vía Las Mesitas, 3100 m, 10.ix.1991, Á. L. Viloria, R. Calchi
leg. (in MALUZ).
Paratypes. 2 males, same data as holotype (2 genit. prep.
ALV545-15, ALV547-15) (in MALUZ); 1 male, Venezuela, Estado
Trujillo, Páramo de Tuñame, 3200 m, 16.ii.2010, M. Costa leg. (in
MC)
Distribution. 3200 m. Restricted to the paramo area of Tu-
ñame and Guirigay (border of Trujillo and Mérida states), SW of
the Niquitao massif.
Comments. This butterfly has been found scarce, and was col-
lected by two of the authors (Á. L. V. and M. C.) only once each
(September 1991 and February 2010, respectively). It was not found
in the same places during a visit to the area in August 2003.
Etymology. The University of Zulia in Maracaibo operated as
a novel institution between 1891 and 1904, the year that Cipriano
Castro, President of Venezuela decreed its closure on the ground
of certain academic and administrative recommendations of Ed-
uardo Blanco, Ministry of Public Instruction. It was formally re-
Viloria et al.
144 ANARTIA, 25 (2013 “2015”): 95 - 160
opened only in August 1946 under the national administration of
a Revoultionary Joint of Government. Dr. Jesús Enrique Lossada
(1892-1948) was appointed as the new Rector of the university. Los-
sada was a professional lawyer and educator from Maracaibo. He
was in fact, a polymath, who apart from have taken several gov-
ernment responsabilities, played music, wrote narrative, poems and
theatrical plays, among others heterodox activities. His role in the
reinstauration, consolidation and development of the university,
was a renowned example of academicism, austerity and honesty.
He is well remembered as a scholar; a respectable figure, who lead
a group of colleagues and disciples to establish a modern univer-
sity. His untimely death came not long after he was removed from
the rectorship. The name lossadana is a posthumous homage to this
most noble man, Jesús Enrique Lossada.
CONSIDERATIONS ON THE POSSIBLE ORIGINS OF
REDONDA AND CONCLUSIONS
Preliminary phylogenetic hypotheses suggest that the genus
Redonda is monophyletic (Viloria, 1997, 1998, 2003, 2007, Ferrer-
Paris 2000, Viloria et al., 2003, 2007, Ferrer-Paris and Viloria 2004,
Pyrcz 2007a, 2007b, 2010a, 2010b, Posso Duque et al. 2010). Results
of any robust cladistic analysis of its species should be correlated
with vicariant events in which the current páramo units sequen-
tially fragmented, and alternatively connected, during the cycles of
the Mérida Glaciation (geologically documented for each range in
Schubert 1970, 1972, 1974, 1975, 1982, Schubert and Valastro 1974,
and Schubert and Vivas 1993).
The glaciers in the Venezuelan Andes, and the consequent
elevation of the páramo biome, may have retreated gradually and
simultaneously at the end of the Mérida Glaciation (Salgado-La-
bouriaru et al. 1988, Schubert and Vivas 1993). It is plausible there-
fore that, the narrower the geographic connection of the current
páramo units during the glacial maximum, the sooner a disjunction
would occur as glacial conditions ended. If so, we can predict a most
recent sequential splitting of the páramo units: (Cendé (Batallón-La
Negra (Niquitao + Mérida + La Culata))). Any attempt to unveil
the phylogenetic history of Redonda needs to take the evidence for
these events into account.
New butterflies from the Venezuelan Andes 145
Figs. 46-49. Lateral views of the male genitalia of three species of Redonda.
Valvae and aedeagus have been detached from natural positions. Valvae:
upper, inner side; lower, outer side. Aedeagus: upper, lateral view; lower,
dorsal view; 46. R. bolivari Adams and Bernard, n. stat., Loma Redonda;
47. Same species, Páramo del Batallón; 48. R. bordoni Viloria and Pyrcz,
Páramo del Batallón (paratype); 49. R. castellana Viloria and Camacho, n.
sp., Páramo de Ortiz (paratype).
Viloria et al.
146 ANARTIA, 25 (2013 “2015”): 95 - 160
The problem of the origins of the high montane representa-
tives of the satyrine subtribe Pronophilina has been raised in previ-
ous works (i. e., Brown 1942, Adams 1977, 1985, Adams and Ber-
nard 1981, Descimon 1986, Shapiro 1992, Viloria 1998, 2007, Pyrcz
2010a, 2010b). Adams and Bernard (1981) considered that the oreal
genera Dangond and Redonda were relics “which have their origins
in an early south-temperate pronophiline fauna”. More recent mor-
phological investigations (Viloria 1998, 2003, 2007) and DNA phy-
logenies (Peña 2009, Peña et al. 2011) have shown that the majority
of genera of the Satyrinae in the austral region of South America
might not belong to the same group of Pronophilina than those of
the Tropical Andes. There have not been comparative studies on
the tribe to determine what can be a “relic” or an “early” genus, but
from field observations on the flight behaviour of the butterflies in
question, we hypothesize that dispersal within these pronophilines
Figs. 50-51. Lateral views of the male genitalia of two species of Redonda.
Valvae and aedeagus have been detached from natural positions. Valvae:
upper, inner side; lower, outer side. Aedeagus: upper, lateral view; lower,
dorsal view; 50. R. centenaria Viloria and Camacho, n. sp., Páramo de
Cabimbú (holotype); 51. R. chiquinquirana Ferrer-Paris, n. sp., Páramo de
Mucubají (paratype).
50 51
New butterflies from the Venezuelan Andes 147
Figs. 52-54. Lateral views of the male genitalia of Redonda empetrus (Thie-
me). Valvae and aedeagus have been detached from natural positions. Val-
vae: upper, inner side; lower, outer side. Aedeagus: upper, lateral view;
lower, dorsal view; 52. Typical individual from the Páramo de Los Conejos
(type locality, herein established); 53. Large, dark form individual from the
upper Río Mucujún; 54. Small, grey-marked individual from same locality
and date.
Viloria et al.
148 ANARTIA, 25 (2013 “2015”): 95 - 160
Figs. 55-58. Lateral views of the male genitalia of four species of Redonda.
Valvae and aedeagus have been detached from natural positions. Valvae:
upper, inner side; lower, outer side. Aedeagus: upper, lateral view; lower,
dorsal view; 55. R. frailejona Ferrer-Paris and Costa, n. sp., Páramo de
Santo Domingo (paratype); 56. R. lathraia Viloria and Camacho, n. sp.,
Páramo del Batallón (paratype); 57. R. leukasmena Viloria and Camacho,
n. sp., Páramo de Las Rosas (paratype); 58. R. lossadana Ferrer-Paris, n.
sp., Páramo de Tuñame (paratype).
New butterflies from the Venezuelan Andes 149
must be exceedingly slow. Pronophiline butterflies were most prob-
ably lowland forest dwellers in the past, and continuous forests in
large portions of the South American continent may have favoured
a wide distribution. However, extant genera show very localised dis-
tribution patterns, which also suggests local origins and local spe-
ciation processes that can be better understood by correlation of
phylogenies with the geological history of each area. As the open
habitats of the high Andes are certainly one of the most recently
established biomes in South America, we consider a local, relatively
recent origin for their faunas far more likely than dispersal of an
ancient fauna, and thus explain the occurrence of highly especial-
ised pronophiline butterflies in the oreal zone of the northernmost
extremity of the Andes.
This assertion is reinforced by considering also the results of
Vuilleumier and Ewert (1978) and Vuilleumier (1981), who found
very little evidence for an austral origin of the high Andean birds
in Venezuela. The conclusion is also consistent with the ideas of
Kroonenberg, Bakker and van der Wiel (1990), who suggested that
the recent origin of the Colombian and Venezuelan high Andes
leaves no temporal or spatial possibility for austral-antarctic biotic
colonization, as had been previously conceived. Additionally, the
mountains considered in this study have always been separate from
the Colombian Eastern Cordillera (the only link with the main An-
dean chain) by a major orographical barrier, the so called “Táchira
Depression”
*
(Vuilleumier 1984). To explain this scenario, it is nec-
essary to review the local geological history.
Vigorous mountain uplift to the páramo zones above 3000 m
in the non-volcanic, extreme northern end of the Andes (Sierra Ne-
vada de Santa Marta, Sierra de Perijá, Cordillera de Mérida, Ser-
ranía del Tamá and Sierra Nevada del Cocuy) occurred less than
6 million years ago, in the Late Miocene, long after the establish-
ment of high elevation regions in the rest of the Andean chain
south of the Santander Massif (Kroonenberg et al. 1990). An open
* A geographical depression is defined as a portion of land lying below
the sea level. Although largely accepted in Venezuelan toponymy,
the term “Táchira depression”should not strictly be used in this case,
because it refers to the wide and low intermontane valley of the rivers
Uribante and Táchira.
Viloria et al.
150 ANARTIA, 25 (2013 “2015”): 95 - 160
type of vegetation (preparamo) is thought to have occurred locally
on hilltops before the northern Andes rose to its present level (van
der Hammen and Cleef, 1986). The formation of preparamo areas
would have been more influenced by local edaphic and climatical
conditions (like the savannahs of the Pantepui), than altitudinal
alone, but the theoretical possibility of an early open vegetation at
increasingly higher elevations during the Late Tertiary uplift should
be considered (“embryonic paramillos” sensu van der Hammen and
Cleef). The possible precursor of the paramo vegetation may have
been present in such open areas and developed from floristic ele-
ments of these “paramillos” in the Early Pliocene, when the tree
line was about 2500 m. The upper forest-páramo belts started to
change position when more land was available for the establishment
of true paramo vegetation, and subsequently with the climatic fluc-
tuations of the Quaternary.
Since there are no true pronophiline butterflies inhabiting open
Andean vegetation habitats at low or middle elevations (they occur
only in the cloud forests where their foodplant Chusquea is abun-
dant and diverse; see Clark 1995), we assume that the few existing
pronophiline butterflies of the paramos (and montane savannah)
are exceptions within the tribe. These taxa may have a relatively
recent origin, more or less contemporary with the new high altitude
habitats to which they are exclusively adapted. Therefore, the north
Andean species belonging to endemic genera of the high elevations
Paramo, Dangond, Redonda, and Steromapedaliodes are here consid-
ered very recently evolved animals whose precursors conquered the
protoparamo from the upper cloud forests, most probably at the end
of the Miocene. Other satyrine genera inhabiting the high paramos
of the northern Andes (Diaphanos Adams and Bernard, Idioneurula
Strand, and Sabatoga Staudinger) are morphologically, ethologically
and ecologically different from what we consider true pronophil-
ines, and their must be set apart.
By the time the Cordillera de Mérida had reached its present
elevation, the genus Redonda could already have been differentiated
from the cloud forest fauna. The former occurrence of a continuous
paramo along the different units of the Cordillera de Mérida (north-
east of the “Táchira Depression”, from El Batallón to the Cendé)
suggests that a single species of Redonda inhabited the páramo at
least during the first fragmentation to form the current units dur-
New butterflies from the Venezuelan Andes 151
ing the last Quaternary glacial period (dated 11,100 years ago; Rull
1996). Thus, the ten species of Redonda are probably geographic vi-
cariants, forming a cohesive monophyletic clade.
ACKNOWLEDGEMENTS
This long term research has been undertaken with support of
the División de Investigación and the Decanato de la Facultad Ex-
perimental de Ciencias, La Universidad del Zulia, as well as from
MALUZ through the efforts of M. Quirós. Financial support in
different stages came from Fundayacucho, CONICIT (Venezuela),
British Council, BMNH, King’s College London, The Linnean So-
ciety (UK); Funding from IVIC was instrumental between 2000 and
2015. Thanks to those who contributed in field work and research,
M. Alarcón, T. R. Barros, P. Boyer, R. Calchi, V. Carrizo, B. Con-
dori Ramos, E. Contreras, L. Morán, M. Duarte, G. Fagua, F. Gar-
cía, M. García, L. González, M. González, M. Higuera, N. León
(†), C. Lozano, J. Moscó (†), E. Moscó, P. Mora, N. Pérez, A.Y.
Sánchez-Mercado, J. Suárez, R. Piña, T. W. Pyrcz, W. Rojas, J. Woj-
tusiak (†), L. Zambrano; and the officers of INPARQUES in Lara,
Mérida, and Táchira for providing us with permits for scientific re-
search in National Parks. H. Suárez (IVIC) took the photographs
of the type specimens and set the plates that illustrate this work.
Our gratitude to those who gave support during our work in public
and private collections and other institutions; AFN: Andrew Neild;
BMNH: P. R. Ackery, B. and L. d’Abrera, J. Harvey, B. Huertas; G.
Martin; L. Mitchell, J. Reynolds, J. T. Tennent; R. I. Vane-Wright;
IVIC: R. Chiussi, X. Jayaro, M. Losada, A. Mata, J. Mavarez, R.
Miller, J. Nassar, D. Posso-Duque, B. Quiroz, C. Ríos-Málaver, G.
Rodríguez (†), J. P. Rodríguez, W. Rojas, A. Ruiz, C. Schubert (†),
H. Suárez, Y. Velásquez, E. Wagner; JFLC: J.-F. LeCrom and C.
LeCrom; MALUZ: R. Carvajal, D. Chirinos, I. Dorado, F. Geraud,
E. Inciarte, M. Quirós, E. Rubio, MBLUZ: R, J. Acosta (†), V. H.
Gutiérrez, A. Hernández-Casanova, J. Moscó (†); MIZA: Q. Arias,
C. Bordón (†), A. Chacón, J. Clavijo, J. De Marmels, M. Gaiani,
E. Guerrero, L. J. Joly, L. D. Otero, F. Romero; MPUJ: G. Fagua,
M. Higuera; MUSM: J. Grados, G. Lamas; MZUJ: T. W. Pyrcz, J.
Wojtusiak (†), ZMHU: W. Mey, M. Nuss, ZSBS: U. Buschbaum; A.
Hausmann; Heimatnaturgarten Weissenfels: U. Radestock. A num-
Viloria et al.
152 ANARTIA, 25 (2013 “2015”): 95 - 160
ber of friends assisted and offered hospitality and support during
academic and field trips; Families Dáger, Jaimez and Otero (Méri-
da), Family Duque (La Grita), Family Mora (Trujillo), A. Jasiñski
(Warsaw), T.W. and J. Pyrcz, R. Laskowski, J. Weiner, J. Wojtusiak
(†) (Krákow), Mr. and Mssrs W. Neukirchen (Berlin), C. Häuser
(Stuttgart), G. W. and J. Beccaloni, A.F.N. and A. Neild, B. Huertas
and T. Donovan, J. and P. Banks (London). The work of M. J. Ad-
ams and G. I. Bernard was a source of inspiration; we thank them
for their friendship and encouraging during Á.L.V. studies in the
UK. Long time ago, G. W. Beccaloni, I. Kitching, D. C. Lees, K.
Sattler, M. Scoble, C. Vardy, and R. I. Vane-Wright (BMNH), B. G.
Gardiner (Linnean Society) and K. Fiedler (Universität Bayreuth)
reviewed and edited several parts of earlier versions of this manu-
script. T. R. Barros, G. Rivas, and an anonymous reviewer made
useful commentaries and suggestions to improve the organization
and edition of the definitive article.
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