ZaologicalJoumaLofthe Linnean Society (198 l), 71: 343-372. With 28 figures
Pronophiline butterflies (Satyridae)
of the Cordillera de Mkrida, Venezuela
M . J. ADAMS
Blyanston School, Blandford, Dorset
AND
G. I. BERNARD
Oxford Scientijc Films, Oxford
Acceptedfor publication April 1480
The 36 species of the tribe Pronophilini known from the Cordillera de MCrida, the main Andean
mountain range in Venezuela, are listed, together with three others probably erroneously described
or labelled as from the range. The general ecological characteristics of the tribe are described, using
the Merida species as examples. These include its restriction to high altitudes, endemism in
mountain ‘islands’, species partitioning up the altitude gradient, mimicry within the tribe, and
seasonality. The evolutionary relationships of the 16 endemic species are analyzed. Two new genera,
six new species, two new subspecies and one new form are described. Three new synonymies are
established, three new combinations made, new status is given to two taxa and the status of one taxon
is revised.
KEY WORDS:- Taxonomy - distribution - Satyridae - Pronophilini - Cordillera de MCrida Venezuela - ecology - speciation - mimicry - new taxa new synonyms - mountain islands Pleistocene.
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C 0NTENTS
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Introduction
List of the species
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Species of doubtful status in the Cordillera de MCrida
Acknowledgements
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References
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INTRODUCTION
The main, southern stem of the Andes divides into three parallel ranges in
southern Colombia. One of these, the Eastern Cordillera, diverges again, about
300km further north-east, to form the Sierra de PerijP and the Cordillera de
Merida. An earlier paper (Adams & Bernard, 1979) dealt with the pronophiline
butterflies of the high northern part of the Sierra de PerijP (the Serrania de
343
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18
0 198 1 The Linnean Society of London
344
M. J . ADAMS AND G. 1. BERNARD
Valledupar); the present paper is concerned with the Pronophilini of the MCrida
range in Venezuela.
The Cordillera de MCrida, situated in the west of Venezuela, is about 360 km in
length by 100 km broad, and runs in a SW-NE direction from 70°40‘N to 10°N.
To its east stretch the flat, low-lying ‘llanos’ and to its west, separating it from
the Sierra de Perija, are the tropical plains and the lake of the Maracaibo basin.
The connection with the Colombian Eastern Cordillera, at its south-western end,
is narrow and low, at a maximum altitude of a little over 1OOOm (the TPchira
depression). Here the mountains are arid and largely bare of forest. Similarly, at
its north-eastern end, the Lara depression (which separates it from the Cordillera
de la Costa) supports a mainly semi-desert vegetation, and the maximum
elevation of contact is about 600 m.
The Chama valley divides the highest section of the Cordillera de MCrida into
two parallel ridges, which unite in the centre of the range at the valley’s head.
The densest forests in the cordillera occur on both of these ridges, and thus flank
the city of Mkrida, which lies on a river terrace in the Chama valley at 1600m.
These two tracts of cloud-forest are now isolated from one another because of
deforestation of the upper Chama valley, but with the formation in 1952 of the
‘Simon Bolivar de la Sierra Nevada’ National Park, 190000 ha to the south and
east of MCrida were protected from destruction and colonization. They comprise
extensive forests between 600m (2300m on the north-western slopes) and the
tree-line at about 3 loom, moor-like ‘paramo’ above that altitude, and four
snow-peaks including Venezuela’s highest mountain, Pic0 Sim6n Bolivar
(5007m). The equally dense and even less disturbed forests and paramo to the
north and north-west of MCrida are also conserved, as a Protected Zone.
Otherwise, cloud-forest occurs in the range only in small and scattered areas,
and much of the paramo is under cultivation.
The cloud-forests owe their existence to high annual rainfall and almost
regular daily cloud-immersion. Moisture rising from the forests themselves is
responsible for much of the cloud and rain, but in the wet season between April
and November the flooded llanos to the east provide an extra source of moisture.
The relatively greater strength of the prevailing easterly and north-easterly winds
in the dry season often keeps cloud away from the forested slopes until late in the
day, while in the wet season clouds regularly form over MCrida and ascend the
slopes before midday, coalescing with cloud already engulfing the paramo. In
our experience the slopes to the north and west of MCrida receive more sunshine
than those in the national park to its south and east, as does the paramo at the
less-forested head of the Chama valley. Minimum temperatures recorded in
August 1977 varied between 8.5 and 10°C at 2500m, and between 4.5 and 6.5OC
at 3000m. Only lOkm down the Chama valley from MCrida there is an abrupt
change in the vegetation, and the forests give way to eroded slopes whose sparse
cover consists largely of cactuses and other xerophytes.
The great majority of our time in the Cordillera de MCrida was spent in the
Sierra Nevada national park and the protected zone opposite, both of which are
accessible on foot (and by cable-car in the national park) from MCrida. The 32
species of Pronophilini which we recorded in the range were all obtained in these
two regions, and no additional species resulted from our visits to other sites (the
Santo Domingo valley in Estado Barinas, the Tovar range in the extreme southwest of Estado MCrida, and the road between Las Cruces and La Azulita, to the
PRONOPHILINI OF MkRIDA
345
west and WNW of Mtrida itselfl. All altitudes between 1500 and 4000m were
covered, but with a concentration on elevations above 2300m. A total of nearly
11 weeks were spent in the MCrida range, between April and June 1975 and in
July and August 1977.
Many of the general ecological characteristics of the Pronophilini,
adumbrated by Adams 8c Bernard (1977; 19791, are illustrated by the Mkrida
fauna :
( 1) With very few exceptions, all the northern South American Pronophilini
are denizens of the subtropical and/or temperate cloud-forests, or the piramo.
In the Cordillera de Mkrida we recorded only three species below 1800m, and
the maximum diversity is reached at 2500 m, where we found 2 1 of the species.
Three species live above the tree-line.
( 2 ) Each mountain range has a high proportion of endemic pronophiline
species, being 33.3%in the MCrida range (13 of the probably total of 39). Eight of
these 13 species have their closest relatives in the main Colombian Eastern
Cordillera; of these, five also have close allies in the Serrania de Valledupar, and
three of the five in the Sierra Nevada de Santa Marta as well. Three species have
their nearest relatives in the MCrida range itself, and the remaining two belong to
monobasic, endemic genera. In addition, the range contains endemic subspecies
of four species.
(3) Each pronophiline species is restricted to a well-defined band of altitudes,
no species occupying the whole range utilized by the fauna as a whole. Among
those in the Cordillera de MCrida for which we have adequate data, the width of
the band varies between 200 and lOOOm, and the mean width for 30 of the 32
species we encountered is 537 m. The limits of the bands do not coincide with any
clear ecotones in the vegetation, except at the forest piramo boundary, and in
the different species they occur at 1500 and 4000m and at almost every altitude
between 1800 and 3400m.
(4) The endemic species in each mountain range are concentrated at altitudes
higher than those occupied by the non-endemics. None of the endemics in the
Cordillera de MCrida occurs below 2000m, and their mean lower and upper
limits are 26 17 and 3183 m, respectively; the corresponding altitudes for the nonendemic species are 2078 and 2628m.
( 5 )Many pairs and triads of closely-related species occur together in the same
mountain ranges, but at mutually exclusive altitudes, replacing one another up
the altitude gradient. In the Mbrida range, at least 16 (possibly 17) of the species
belong to such replacing series, comprising four pairs and at least two (possibly
three) trios (named in order of ascending altitude in each case): Altopeduliodes
albarregus sp. nov. and A . albonotata (Godman); Corades pannonia Hewitson and
C . cistene Hewitson; Eretris calisto (Felder 8c Felder) and E . porphyria (Felder 8c
Felder); Pronophila bogotensis Jurriaanse and P . epidipnis Thieme; Corades enyo
Hewitson, C. chelonis Hewitson and C .pax Watkins ;Lymanopoda obsoleta Westwood,
A . dietzi sp. nov. and L. marianna Staudinger; and Pedaliodes manis (Felder &
Felder), P. m o n t u p sp. nov. and possibly P. untigUa sp. nov.
An explanation for the distributions of the Pronophilini of northern South
America, in both the horizontal and the vertical planes, was offered by Adams
( 197 7 ). It involves the hypothesis that the glacial and interglacial periods of the
Pleistocene resulted in a cycle of spread and isolation of pronophiline
populations, and therefore in the tribe’s colonization of mountainous areas
346
M . J. ADAMS A N D G . I . BERNARD
isolated from the main Andes (in Central America, for example) and in its rapid
radiation within the Andes. The series of replacing species up the altitude
gradient are explained by postulating shifts in altitude preferences by closely
related species, after coming into competitive contact during glacial periods. The
inability of the upper members of such series to spread in subsequent cold
periods, because of their high lower-altitude limits, explains the existence of
endemic genera and species, which occur in the greatest number in mountain
ranges having the lowest connections with their neighbours.
( 6 ) Certain pairs of species from different genera or species-groups, living at
the same altitudes in the same places, resemble one another to such an extent
that the evolution of mimicry may be inferred. This is particularly so when the
resemblance is shown by the upperside rather than the underside markings
(because convergent evolution of underside patterns for camouflage on similar
backgrounds has probably occurred commonly in the tribe), and when one or
both of the species has diverged from the normal upperside patterns of its genus
or species-group.
0 ne case exists in the Cordillera de Mkrida which bears many of the hallmarks
of Batesian mimicry. Cheimar opalinus (Staudinger) and Lymanopoda marianna share
a large, circular, light blue patch on their upperside hindwings, visible only in
flight and when the butterflies sun themselves at rest. Lymunopoda marianna is the
only member of its genus with such upperside markings; its altitude range lies
entirely within that of C. opalinus, and the two species were often seen around the
same clumps of bamboo. Where they coexist, opalinus is very abundant, while
marianna is scarce; opalinus has a slow flight, while the smaller marianna always flies
very fast. We consider it possible that opalinus is in some way obnoxious to
prehators, and that it is mimicked by the less obnoxious marianna.
( 7 ) Some species of’ Pronophilini are strongly seasonal, notably in the genus
Penrosada, represented in the Cordillera de Mkrida by P.franciscae sp. nov. This
species was quite common in August 197 7 at the site where a few individuals were
seen in June 1975 and where it was apparently absent in April 1975. Conversely,
the high piramo species, Diaphanos huben‘ sp. nov., was absent in June at the site
where we obtained our first series before the wet season began, in April 1975.
LIST OF THE SPECIES
A total of 36 pronophiline species are known from the Cordillera de MCrida.
We obtained 32, of which two were new and a further three undescribed. The
remaining four are represented at the British Museum (Natural History), two by
specimens labelled as from MCrida, and two by Venezuelan specimens almost
certainly from MPrida rather than the coastal range or the Pantepui mountains in
the south. Although there is no evidence in the literature or in the British
Museum (Natural History) collections, it is also possible that Catargynnis phaselis
(Hewitson), Pedaliodes pisonia (Hewitson) and Pronophila orcus (Latreille) occur in
the Mkrida range.
Three species, all described by Felder 8c Felder (Eretris apulqa, Idioneurula
erebioides and Lasiophila circe) were either described from ‘Venezuela’ or, in the
case of I . erebioides, have one ‘Venezuelan’ representative in the British Museum
(Natural History). Although these are listed at the end, we consider it unlikely
PRONOPHILINI OF MERIDA
341
that any of them do in fact fly in the Cordillera de MCrida. Our reasons are given
in the text.
The specific altitude limits and notes on behaviour and habitats come from
our own records and observations. Anecdotal information on oviposition and
early stages of a few of the species was obtained during our 1977 visit.
The relationships of the endemic species have been judged mainly on the basis
of male genitalia structure, but also on ecological factors such as behaviour and
altitude preferences, where these are known, and on certain aspects of the facies
considered to be the most stable. While many closely-related species cannot be
distinguished surely by comparison of their male genitalia, genera and speciesgroups can be separated on this basis.
All our specimens are the property of the British Museum (Natural History),
except for the four paratypes presented to the Museum of Entomology at the
Universidad Central de Venezuela in Maracay (a male each of Lasiqpfiila mpatoza
meridue subsp. nov. and Lymanopoda dietzi sp. nov., and a pair of Penrosuda
j+anciscae sp. nov.).
The following abbreviations are used : BMNH-British Museum (Natural
History); MNHU-Museum fur Naturkunde der Humboldt-Universitat, Berlin;
and CM-Cordillera de Mkrida.
Altopedaliodes albarregas sp. nov.
(Figs 1, 14, 15)
Male
Forewing length : 22-23 mm. Upperside ground colour dark brown, hindwing
without markings, forewing with white markings as follows: a rectangular patch
across discal cell, extended across base of cell Cul to vein Cu2; small patch near
base of cell R 5 (sometimes absent). No scent brand. Fringes of both wings
chequered dark brown and white.
Underside forewing dark brown with white markings as on upperside, but the
patch in cell R5 extends into M 1, and there is a series of 4 or 5 white postmedian
spots in cells R4 to M2 or M3. Underside hindwing dark brown speckled with
whitish scales, particularly at inner margin; an indistinct, slightly lighter
postmedian band with a series of 3 white spots in cells Cul, M3 and M2, 2 white
>--shaped marks in M1 and Rs, and 2 or 3 flecks at the costa; a small white
streak halfway along costa, and (in some specimens) an additional fairly distinct
costal streak which extends to form a discontinuous whitish line across the discal
cell and as far as the inner margin,
Fernale
Forewing length: 24mm. Upperside as male, but with white speckling along
forewing costa and a small costal streak near the hindwing apex.
Underside forewing with better-developed postmedian spots, yellowish in
colour, and an additional one in cell Cul; apex mottled with grey, dark brown
and chestnut. Underside hindwing dark brown mottled with lighter brown and
lilac; the indistinct postmedian band contains 5 yellow marks, the posterior 3 of
which are > -shaped (cells Cul to M21, and others are >--shaped (cells M 1 and
Ks).
348
M. J. ADAMS A N D G. 1. BERNARD
Holotype $, VENEZUELA: Estado MCrida, Rio Albarregas, 3150m, 16.viii.77
(M.J .A.).
Paratypes, locality as holotype: 1 8 , 3200m, 17.vi.75; 1 9, SOOOm, 19.vi.75; 18,
3200m, 20.vi.75; 18,34?00m, 21.vi.75; 5d, 3100-3150m, 13.viii.77; 1Q,3300m,
14.viii.77; 18,3100m, 15.viii.77; 2 8 , dataasholotype(MJ.A.andG.1.B.).
Altopedaliodes albarregas, named after the valley in which we discovered it, is
evidently very closely related to A . albonotata (Godman),which differs in having its
white forewing patch restricted to the discal cell and to the upperside, and in
having two postdiscal white spots on the upperside forewing. The subunci of
albarregas are shorter, the valvus broader (Fig. 11, and the forewing longer and
more pointed.
3000-3300 m. An endemic species. Altopedaliodes albarregm and A . albonotata are
totally separated ecologically: albonotata lives close to the ground in the paramo,
while albarregas flies around the tops of flowering trees in the very uppermost
forest. It is because such forest extends up sheltered gullies into the paramo that
the altitude ranges of the two species overlap. Both sexes of albarregas occasionally
come to the ground, to drink from moist gravel or to feed on organic matter. It
has a rapid and erratic flight.
Aftopedufiodes ufbonotata ( Godman) comb. nov.
(Fig. 2)
Pedaltodes albonotata Godman, 1905: 189, pl. 10, fig. 11. 28 syntypes,
VENEZUELA: Merida (BMNH), [examinedl.
Pedaliodes albonotata Godman; Thieme, 1905:96; Weymer, 1912 :257 ; Gaede,
1931 :488.
Steromapedaliodes albonotata (Godman); Forster, 1964:148.
Forster ( 1964)erected the genus Steromapedaliodes for this species on the basis of
its male genitalia. His figure of the genitalia of albonotata, however (Forster,
1964: 148, fig. 1721, is incorrect (see Fig. 2). We consider that the species
belongs in Forster's ( 1964) genus Altopedaliodes, with which we synonymize
Steromapedaliodes (syn. nov.). All Altopedaliodes have broadly similar male armatures and (except for A. albarregm) are denizens of the paramo. Within the genus,
albonotata superficially most resembles rezssi Weymer from the Colombian Central
Cordillera, and A . albarregas.
Individuals in which the two white postdiscal spots on the forewing show
through to the underside were named ab. privigna by Thieme (1905 :9 7).
3100-4000m. An endemic species. The commonest of the three pronophiline
species in the CM piramo, both to the north and to the south of MCrida. I t
occurs in colonies in marshy land, flying close to the ground, and forages over
flowery, rocky slopes, resting often on lichen-covered rocks. I t is capable of fast
and erratic flight, and flies even in the briefest of sunny periods in spite of the
cold.
Chimas opalinus (Staudinger)
Oxeoschistus opalinus Staudinger, 1897 : 145, PI. 5, fig. 10. 6 6 syntypes,
VENEZUELA: Sierra Nevada bei Merida (MNHU, Berlin).
PRONOPHILINI OF MLRIDA
349
Cheimas opalinus (Staudinger); Thieme, 1907:176; Weymer, 1912271; Gaede,
1931516.
Three pronophiline species in the CM bear pale blue patches on their
upperside hindwing: C. opalinus (under certain lights its patch appears silveryyellow), Mygona irmina (Doubleday) and Lymanopoda marianna Staudinger. Cheimas
opalinus most resembles M . irmina, and both share a slow and rather feeble flight;
but the male genitalia place opalinus, the only member of its genus, closer to
Lasiophila than to Mygona.
The whitish and yellowish markings on the underside hindwing, most
pronounced in the rarely-encountered female, are variable; the extreme form in
which they are absent was named ab. spoliatus by Staudinger (1897:146).
A few specimens in the BMNH from Cohmbia, and one from Ecuador,
indicate that C. opalinus is not restricted to the CM, but it seems probably that it is
at its commonest there.
2600-3300m. Locally very abundant, especially above 2900 m, flying slowly
around clumps of bamboo and occasionally coming to the ground to feed on
organic matter. Although its habitat extends up gullies, sometimes only a few
metres wide, into the paramo, we have never seen it stray out into the open.
Corades enyo Hewitson
Corades enyo Hewitson, [18491:117, annulosa pl. 4. 2 8 syntypes, VENEZUELA:
Caracas (BMNH),[examined].
Typical C. enyo is known only from the coastal range of Venezuela; ssp. almo
Thieme (1907:222, described as a var.) occurs between Colombia and Bolivia. A
single male referable to almo, labelled ‘Venezuela’, is in the BMNH from the
Fruhstorfer collection, and may have come from the CM. There seems little
reason why it should not fly there, and we failed to find it perhaps because our
collecting was concentrated at altitudes above those at which we would expect it
to occur: in the Sierra Nevada de Santa Marta it flies between 1400 and 2200m
(Adams & Bernard, 1977), and in the Serrania de Valledupar up to 2400m
(Adams & Bernard, 1979).
Corades chelonis Hewitson
Corades chelonzs Hewitson, 1863:71, figs 1, 2. 18 syntype, COLOMBIA: New
Granada (BMNH),[examined].
The population of C . chelonis in the CM was given the name rubeta by Thieme
(1907:223, described as a var.), which may be considered as an endemic
subspecies. I t has more rounded wings and a larger and more oval orange
postdiscal patch on the forewing. Typical chelonis is found in Colombia, and the
race lactefusa Thieme occurs in Ecuador and Peru.
2200-2550m. Usually solitary, fond of resting on foliage two or more metres
above ground, and has a fast and dodging flight.
Coradespax Watkins
(Fig. 3)
Corades pax Watkins, 1939:160. 3 8 syntypes, VENEZUELA: Tachira, Pedregosa
and Quintero 3000m [all CMI (BMNH), [examined].
M. J. ADAMS A N D G. I . BERNARD
350
Thieme (1907:224) noted that among his series of Venezuelan C. chelonis were
two males with an extra orange patch at the tornus of the forewing. These no
doubt represent what Watkins ( 1939) later described as C. pax from three BMNH
specimens. In addition, pax has a larger and less rounded postmedian orange
patch on its forewing, and a greyer underside hindwing than chelonis. The male
genitalia are illustrated in Fig. 3.
Figures 1-6. Male genitalia of the pronophiline species endemic in the Cordillera de Mkrida. Fig. 1.
Alt@daliades albarregm sp. nov. Fig. 2. A . albonotala (Godman) comb. nov. Fig. 3. Corades pax
Watkins. Fig. 4. Diaphrmos huberi sp. nov. Fig. 5. LVnulnopoda dietxi sp. nov. Fig. 6. L. marianna
Staudingrr.
2300-2600 (3IOU?)m. An endemic species, replacing chelonis above 2550m.
Although we did not catch it above 2600m, we recorded a number of individuals
of either chelonis orpux flying high in the trees at 3 loom, and we believe that they
were most likely to have been pax. I t has similar habits to those of enyo and
chelonis; the three species represent an example of a series of closely-allied
congeners replacing one another up the altitude gradient (see Introduction).
Corades medeba Hewitson
Corades medeba Hewitson, 1850:439, pl. 10, fig. 4. 2 6 syntypes, BOLIVIA
( BM NH 1, [examined].
2 1 0 0 - 2 3 5 0 m . Corudes medeba is solitary and uncommon, and most often
encountered settled on the ground, often on moist gravel near streams and
rivers. Like other ground-feeding Corades species, it flies high into the forest
when disturbed, but may return minutes later to the same spot. The species is
known from between Venezuela and Bolivia.
PRONOPHILINI OF M ~ R I D A
35 1
Coradespannonia Hewitson
Corades punnonia Hewitson, 1850:438, pl. 10, figs 1 , 2. Lectotype 8,
VENEZUELA [coast range] (BMNH),here designated [examined].
Corudesjuminalis Butler, 1870:26; Butler, 1874: 182, pl. 62, fig. 8. 19 syntype, [no
locality](BMNH),[examined].
The light-coloured typical race of C. pannonza comes from the Venezuelan coast
range. Two male specimens in the BMNH, on which Hewitson probably based
his description of pannonia, have been designated and labelled lectotype and
paralectotype. Elsewhere in its range (the CM and Colombia), the underside is
greyer. Thieme (1907 :212) described this race as var. ploas; he pointed out that
Butler’sJlurninalisreferred to female plous. Moritz’s previously unpublished name
ichthya was used by Herrich-Schgffer in his illustration (1856 fig. 84); Thieme
(1907) recognized its synonomy with ploas.
2000-2400 rn. A ground-loving species with similar habits to C. medeba and
C . cistene and with a fast and erratic flight.
Corades cistene Hewitson
Corades cistene Hewitson, 1863:72, figs 4, 5. Lectotype 8,BOLIVIA (BMNH),
here designated lexaminedl.
As explained b Thieme ( 1907 :220), Hewitson’s original description of cistene
probably referreJ to a Bolivian specimen, although his figured specimen does
not tally and represents the Peruvian race (var. generosa Thieme, 1907). Two
Bolivian specimens in the BMNH, on which Hewitson may have based his
description of cistene, have been designated and labelled lectotype and
paralectotype. A third race, from the CM, Colombia and Ecuador, was described
by Thieme ( 1907) as var. dymantis.
2500-3000 m. Corades cistene dymantis has habits similar to those of C . pannonia
ploas, to which it is very closely related, and replaces it above
2400-2500 m. We found it most commonly on rocks and gravel near streams and
on the ground at water seepages in the forest.
Diaphanos gen. nov.
Type-species: Diaphanos huba’ sp. nov.
Eyes glabrous, dark chestnut-brown and black. Palpi twice as long as head,
light brown with dark brown and cream-coloured hairs, and white scales; 2nd
segment 2.5 times longer than 1st; 3rd segment very short (three-quarters the
length of the first). Antennae to two-fifths costa; shaft orange-brown, club black;
club blunt with parallel sides, 2.5 times longer than broad, concave. Forelegs
extremely reduced; femur and tarsus each one-third the length of tibia.
venation. Forewing: Sc, R1 and R2 all independent; root of R3 nearer to root of
R4 than to discal cell; no R-M 1 ;M 1-M2 curves only very slightly inwards; M2-M3
cumes outwards osteriorly; M2 enters cell. Hindwing: humeral vein well
developed; root o Cu 1 nearer to root of M3 than that of CUP; root of M3 halfway
between those of Cu 1 and M3.
Forewing tornus obtuse. Both wings rounded.
Uncus blunt and longer than tegumen; subunci long; aedeagus long and
toothed at tip.
P
952
M. J. ADAMS A N D G. I . BERNARD
&
7
f
Figures 7-13. Male genitalia of the pronophiline species endemic in the Cordillera de Mtrida. Fig. 7.
Pedalioderjaphleta Butler. Fig. 8. P. mmtagm sp. nov. Fig. 9. P. antigua sp. nov. Fig. 10. P. m a l a
Grose-Smith & Kirby. Fig. 1 1 . PenroradajirmEircae sp. nov. Fig. 12. Pronophila epidipnis Thieme. Fig.
13. Redonda empefrus(Thieme)comb. nov.
Diuphunos huben’ sp. nov.
(Figs4, 16, 17)
Male
Forewing length: 13.5- 15.5mm. All wings transparent, lightly dusted with
grey scales, particularly in outer half of forewing and over whole of hindwing;
black scales at base of both wings, extending on hindwing posterior of median
vein as far as root of Cu2.
Underside faintly russet-brown along costa of forewing and at base of costa of
hindwing. Fringe brown at outer margin of forewing, white at inner margin;
white around hindwing, grey at tornus.
Female
Forewing length: 13.5mm. Upperside ground colour white, with dusting of
grey-brown scales along costa and over outer third of forewing, and with black
scales at base of both wings.
PRONOPHILINI OF MBRIDA
353
Underside forewing white dusted over whole wing with grey-brown scales; a
discrete light brown patch distally in the discal cell. Underside hindwing dusted
with grey-brown scales, except in the well-defined narrow streaks of the white
ground colour parallel to the veins, from the discal cell to the outer margin, in
cells Cu2 (two streaks), M3 to Rs; the brown colour of the veins themselves is
much less pronounced than on the forewing, except on the median and radial
veins and Sc+ R1; russet-brown coloration at base on costa.
HoloQpe 8,VENEZUELA: Estado MCrida, between Laguna Coromoto and
Laguna Verde, approx. 15km S.E. of MCrida, 3450m, 27.ii.7 1 ( H . Huber).
Paratypes, VENEZUELA : Estado MCrida, between La Aguada and Loma
Redonda, approx. 8km S. of MCrida: 4 8 , 4000m, 19.iv.75; 58, 19, 4000m,
20.iv.75; 28, 3900 & 4000m, 21.iv.75; upper Albarregas valley: 2 8 , 3400 and
3600m, 15.viii.77( M J . A . and G.I.B.).
This, the smallest and most unmarked pronophiline known, was discovered in
February 1971 by Herr H. Huber, after whom we name it. It is clearly a relict
species, whose affinities cannot even be guessed at. Its male genitalia (Fig. 4) are
unlike those of any other pronophiline species we have examined.
3400-4000~2.
An endemic species, confined to the piramo well above the treeline. Other than one male found in a marshy area of short tussocky grass, all the
individuals we have seen flew on well-drained, rocky slopes characterized by the
abundance of a low, whitish-leaved species of Espeletia. It flies like a micro-moth,
rapidly and erratically very close to the ground, and often drops suddenly into
the sparse vegetation when alarmed. On a rare sunny day in the wet season, in
June 1975, it was absent from the site where we had obtained our series two
months before. In June 1975 and August 1977, however, it flew in the mountains
north of MCrida where the weather is less inclement.
Eretris calisto (Felder & Felder)
Pronophila calisto Felder & Felder, 1867 :472. 1 8 syntype, COLOMBIA: Bogota
(BMNH1, [examined].
Eretris calisto (Felder & Felder); Thieme, 1905: 133; Weymer, 1912:263, pl. 56,
row 6 ; Adams & Bernard, 1977:269.
Although we did not obtain this species in the CM, the BMNH has one male
specimen labelled ‘Venezuela’,purchased from Dyson; our failure to find it there
was perhaps a result of its preference for altitudes lower than those at which we
concentrated our collecting: elsewhere it flies between 1400 and 2200m (see
Kruger, 1924;Adams & Bernard, 197 7 ). The species is replaced in the Venezuelan
coast range by E. encycla (FeIder & Felder), and outside the CM it occurs in
Central America and all the main mountains of Colombia.
Eretris porphyria (Felder & Felder)
Pronophila porphyria Felder & Felder, 1867:470. 1 8 syntype, VENEZUELA [:CMl
(BMNH),[examined].
Eretrisporphyria (Felder & Felder); Thieme, 1905: 132; Weymer, 1912:263, pl. 56,
row b.
Eretris porphyria probably forms a monophyletic group with E . perija Adams &
Bernard from the Serrania de Valledupar and E. decorata (Felder & Felder) from
354
M . J. ADAMS AND G . I . BERNARD
PRONOPHILINI OF M ~ R I D A
355
the Colombian Eastern Cordillera, which are the middle-range members of
three-species series of replacing congeners up the altitude gradient (see Adams &
Bernard, 1979; Kruger, 1924). It is distributed in the CM and parts of the main
Colombian Eastern Cordillera.
2200-2800m. It has the typical skipping flight of the genus, and is most
commonly found around clumps of climbing bamboo, rarely coming to within
1 m of the ground. Ova are laid on leaves of a species of Chusquea and, at least at
first, are green in colour.
Lasiophila zapatoza (Westwood)
Pronophila zapatoza Westwood, [18511: 358. 2 d , 1Q syntypes, VENEZUELA
I: coastal range] (BMNH),[examined]; and 'BOLIVIA, [not located-see also
Butler, 1868: 1811.
Lasiophila zupatoza (Westwood); Butler, 1868: 181; Thieme, 1907 :127;Weymer,
1912:2.65;Adams &Bernard, 1979:102.
Lasiophila zapatozu is divided into a number of distinct races in Colombia and
Venezuela. The typical subspecies comes from the Venezuelan coast range, ssp.
semipartita Weymer occurs in the Sierra Nevada de Santa Marta (Adams &
Bernard, 1977), ssp. manaurera Adams & Bernard (1979) in the Serrania de
Valledupar, and the CM has an undescribed race- subsp. meridae.
Lasiophila zapatoza meridae subsp. nov.
(Fig. 18)
Male
Forewing length: 29.5-3 1.5mm. Differs from the typical subspecies in the
following respects: it is larger (cf. typical forewing length: 29-29.5mm); on the
upperside forewing the ground colour is darker; the dark brown distal markings
extend further proximally in cells Cu 1 to ZA, completely enclosing the triangular
russet-brown patch in cell Cul; all the postmedian patches are darker, of the
ground colour, and reduced in extent; the subapical patch is broken into two,
and the posterior patch barely extends into cell MS. Upperside hindwing ground
colour darker; the postmedian series of dark brown circular patches are all
larger, those in cells Rs, M I and M2 coalesced and joined to the marginal band
along the veins, that in M3 also joined to the marginal band; the marginal rustybrown line between anal angle and vein M 3 is less well developed.
The difference in the light brown markings of the forewing are reflected on the
underside; those submarginal in cells Cul and Cu2 do not coalesce; the basal
area of rusty-brown extends further distad in cell M3 and the discal cell, and
enters cell M2. Underside hindwing similar but slightly darker throughout.
Figures 14-23. The left shows the upperside, the right half the underside. From the Cordillera
de Mirida, in the vicinity of Mirida. Fig. 14. Ahjw&liodt?s albarregas sp. nov., male holotype. Fig. 15.
The same, female paratype. Fig. 16. Diuphams hubm'sp. nov., Male holotype. Fig. 1 7 . The same, female
paratype. Fig. 18. Lasiophila znpatour (Westwood) mcridoc subsp. nov., male holotype. Fig. 19.
~ymnopoda&tzi s p nov., male holotype. Fig. 20. The same, female paratype. Fig. 21. Peduhdes
m a p a sp. nov., male holotype. Fig. 22. Pedaliodcs mrtigW sp. nov., male holotype. Fig. 23. The
same, female paratype. (All Figs c. life-size, except Figs 16 & 1 7 , which are c. twice life-size.)
356
IM.J. ADAMS AND G . I . BERNARD
k-emale
Forewing length: 35mm. Forewing similar to that of ssp. semipartita, darker
than in manaurera; hindwing close to that of munaurera, but darker and with larger
postmedian dark brown patches. Larger than the other races (we have not seen
female typical tapatora3.
VENEZUELA: Estado Mtrida, La Montafia, S. of MCri'da, 2350m,
Holotype 8,
12.iv.75 ( M J . A . 1 .
Paratypes, locality as holotype: 1 8 , data as holotype; 78, 2450-2500m,
lO.iv.75; 5 8 , 19, 2350-2450m, 12.iv.75; 1 8 , 2500m, 13.iv.75; 1 8 , l Q , 2500m,
6.vi.75; 18, 2500m, 14.vi.75; 1 8 , 2500m, 15.vi.75; 18, 2300m, 4.viii.77; 128,
19, 2450-2550m, 9.viii.77; 1 8 , 19, 2450m, lO.viii.77; 7 8 , 2450-2550m,
19.viii.7 7 ; 1$, 2450-2550 m, 2O.viii.7 7 ; Estado Mtrida, La Mucuy, near Tabay,
18, 19, 2200--2300m, 16.iv.75 (8Universidad Central, Venezuela); Estado
Merida, Rio Albarregas: 18, 2250m, 16.vi.75; 1 8 , 2250m, 22.vi.75; 5 8 ,
1900-2050 m, 3 1.vii. 7 7 ; 38, 2000, 2 150 and 2350 m, 17 .viii. 7 7 ( M J . A . and
C . 1 . B . ) ;Culata, 18;Quintera, 1 8 , Crowley Bequest BM 1901-78; 'Venezuela',
18,Godman-Salvin Coll. BM 1904-1 (Goering).
1900-25OOm. Locally quite common in the CM, flying slowly in sunny
clearings and settling in foliage, rarely lower than 1 m above the ground, or on
the ground itself where it is well camouflaged against leaf litter.
Lymanopoda obsoleta (Westwood)
Sarromia obsoleta Westwood, [May 18511: pl. 6.7, fig. 5. 18 syntype, BOLIVIA
(BMNH),[examined].
Lymanopoda obsoleta (Westwood);Westwood, [July 185 11:402.
Lymanopoda larunda Hopffer, 1874 : 36 1. 8 syntypes, PERU : Chanchamayo/
VENEZUELA [whereabouts of types unknown]. [Synonymy given by Weymer,
1912 :2481.
Lymanopoda larunda Hopffer; Thieme, 1905:70.
Sarromia obsoleta Westwood; Forster, 1964: 145.
Lymanopoda obsoleta (Westwood); Butler, 1868:168; Weymer, 1912:247; Gaede,
1931:487; Brown, 1943:98; Adams & Bernard, 1979:105.
The genus Sarromia was correctly synonymized with Lyrnanopoda only two
months after it was proposed. The members of' Lymanopoda are diverse in size,
wing-shape and colour pattern, but common to all of them is the bulbous
subuncus which is unique in the Pronophilini (see Brown, 1943).Weymer ( 19 12)
explained the synonymy of larunda with obsoleta.
Lymanopoda obsoleta is distributed between Panama and f)olivia, including
Venezuela, and at least in Colombia and Venezuela where it flies in the lower
cloud-forests, it is replaced by different closely-related congeners at higher
altitudes in each mountain range. In the CM, where the group of Lymanopoda
species which includes L. neuada Kriiger and L. labda Hewitson is absent, a third
member of the obsoleta group occupies the highest cloud-forest zone. The
resulting trio of replacing species is obsoleta, dietzi sp. nov. and marianna Staudinger.
2100-2600 m. Common in the CM, characteristically found skipping around
and between clumps of tall bamboo, usually more than 1 m above ground, but
sometimes feeding on the ground. The pupa is a unicolorous green; we found one
PRONOPHILINI OF MkRIDA
957
suspended under a large bamboo leaf 2 m above ground, from which a female
emerged ten days later.
Lymanopoda dietzi sp. nov.
(Figs 5, 19, 20)
Male
Forewing length : 23-26 mm. Upperside forewing dark brown with triangular
orange basal patch in cells 2A and Cu2 (sometimes entering discal cell and base
of Cull; the patch has a green sheen. Upperside hindwing basal half greeny
golden-brown; outer half dark brown.
Underside forewing chocolate-brown, russet-brown at apex, slightly lighter
brown in partial postmedian band posterior of vein M3, which opens to include
much of cells Cu2 and 2A as far as outer margin, and which contains 2 whitepupilled black spots in cells Cul and M3; 2 submarginal white spots in cells M 1
and M2, and faint whitish subapical streak in R4 and R5. Underside hindwing
russet-brown, dusted with lilac in distal half; postmedian band in 2 separate
parts: first pinkish, dusted with brown and lilac, from costa curving into cell and
continuing to inner margin faint and rusty brown; second dark pink, heavily
dusted with brown and lilac, from submarginally in cell M1 to inner margin; 6
very small, faintly white-pupilled, dark brown spots, 2 submarginal in cell Cu2,
one each postmedian in Cul to M 1 (the anterior 3 and the inner of the 2 in Cu2
are sometimes barely visible).The male genitalia are shown in Fig. 5.
Female
Forewing length: 22.5-25 mm. Upperside forewing dark brown, with
triangular green-shining, golden yellow basal patch larger than in male; whitish
transparent submarginal spot in cell M 1 and trace of another below it in M2; 2
postmedian spots in cells Cul and M3 show through faintly as dark brown,
pupilled with light brown. Upperside hindwing dark brown with greenish sheen
over basal half.
Underside forewing dark rusty brown in base of discal cell and of cells M3 to
Cu2; dark sandy brown over rest ofwing, except for rusty brown outer margin in
apical region; 2 transparent whitish submarginal spots in cells M 1 and M2, and 2
postmedian white-pupilled black spots in Cul and M3; rusty brown patch in
discal cell contiguous with median vein. Underside hindwing almost uniform
grey-brown heavily dusted with lilac, and faintly with turquoise along inner
margin and at base posterior of median vein; very faint indication of double
postmedian band as slight lightening of colour ;white-pupilled dark brown spots
as in male, but extremely faint.
Holotype d,VENEZUELA: Estado Mkrida, above La Montafia, S . of MCrida,
2450-2550m, 2O.viii.77 (A4J.A.).
Paratypes, locality as holotype: 2 8 , data as holotype; 3 8 , 2600-2650m,
10.iv.75 (1 Universidad Central, Venezuela); 2 8 , 2600 8c 3000m, ll.iv.75; 18,
19, 2400m, 13.iv.75; 1 8 , 3100m, 7.vi.75; 4 9 , 3000m, 8.vi.75; 18, 2600m,
14.vi.75; 28, 2600m, 15.vi.75; 1 8 , 2900m, 3.viii.77; 3 8 , 2450-2550m, 9.viii.77;
2 8 , 2450-2550m, 19.viii.77; Estado Mkrida, La Mucuy, near Tabay: 148, 39,
2400-2700m, 16.iv.75; 28, 2100-2300m, 24.iv.75; Estado Mkrida, Rio
Albarregas: 3d, 2800m, 17.vi.75; 4 8 , 2900-3050m, 18.vi.75; 28,2400-2600m,
358
M. J. ADAMS AND G. I. BERNARD
18.vi.75; 18, 2800m, 19.vi.75; 1 8 , 3000m, 20.vi.75; 4 3 , 2850m, 21.vi.75; Id,
3150m, 13.viii.77; 6 8 , 3050-3150m, 14.viii.77;23,2700m, 16.viii.77;2 8 , 2350
& 2600m, 17.viii.77 (A4J.A. and G.Z.B.); ‘Venezuela’, 2 3 , ex Fruhstorfer Coll.
BM 1933-131 8c 1937-285; ‘Merida’, 78, Joicey Bequest BM 1934-120 (Brideno);
El Valle [N.E. of MPridal, l d , Joicey Bequest (Briceno).
We name L. dietzi after Robert Die& IV of New Mexico, U.S.A., to thank him for
having broadened our entomological knowledge and experience, and for his
help and friendship in Venezuela. The species probably belongs to a
monophyletic assemblage composed of the other middle-altitude-level members
of the obsoleta group, L. maletera Adams & Bernard in the Serrania de Valledupar
and L. altis Weymer in the remaining major mountain ranges of Colombia, and
Ecuador.
350-3120 m. (Two isolated individuals recorded between 2100 and 2300 m at
La Mucuy in disturbed forest). An endemic species with habits typical of
members of the obsoleta group of the genus. The butterflies are often found on
animal excrement on the ground, where they may remain until long after the sun
has gone in.
Lymanopoda marianna Staudinger
(Fig. 6)
Lymanopoda marianna Staudinger, 1897 : 146, pl. 5 , fig. 6. 2 8 syntypes,
VENEZUELA: Merida (MNHU, Berlin).
Because they share blue patches on their upperside hindwings, L. marianna and
L. cinna Godman & Salvin from Guatemala resemble one another, but their male
genitalia place them in different species groups. L. marianna belongs with the
obsoleta group, whose members have two long and narrow processes on the valve
(Fig. 6). As far as we know, it is the only member of the group to fly in the very
topmost forests, which in most places in northern South America are occupied
by species of other groups of Lymanopoda, which are absent in the CM. The
resemblance between marianna and Cheimas opalinus, with which it flies, may
represent a case of mimicry (see introduction).
3000-3300m. An endemic species, which rarely descends to within three
metres of the ground. We found it skipping around the. tops of tall bamboo
clumps and occasionally flying at high speed between them, in one small area of
the upper Albarregas valley.
Lymanopoda aibocincta Hewitson
Lymanopoda albocincta Hewitson, 1861:157, pl.9, fig. 5. 1 8 syntype, COLOMBIA:
New Granada (BMNH),[examined].
Lymanopoda issacha Butler, 1870:26. 18 syntype, [ECUADOR?] (BMNH),
[examined].(Synonymygiven by Adams 8c Bernard, 1979: 107).
Lymanopoda albocincta Hewitson; Butler, 1868:171; Weymer, 1912:249; Gaede,
1931:484; Brown, 1943:lOO; Adams &Bernard, 1979:107.
1900-2500m. A mainly ground-loving species, feeding among leaf litter, and
flying with the characteristic lymanopodine skipping action. We found it only in
small numbers. It is distributed between north-eastern Colombia and Peru, and
the CM.
PRONOPHILINI OF MBRIDA
359
Lymanopoda albocincta is one of the few members of the Pronophilini to exhibit
pronounced sexual dimorphism: while the male is little marked but for the white
band across the hindwing underside, the female has large postdiscal ocelli on
both surfaces of the forewing, and a washy underside hindwing with only a few
faint white scales in place of the male’s white band.
Muscopedaliodesjephtha (Thieme)comb.nov.
Pedaliodesjephtha Thieme, 1905:84, pl. 2, fig. 18. 1 8 , 19 syntypes, COLOMBIA:
Antioquia [whereabouts of types unknown].
Pedaliodesjephtha Thieme; Weymer, 1912:255; Gaede, 1931:491.
The genus Muscopedaliodes was erected by Forster (1964:153) for a group of
Bolivian species, previously classified in Pedaliodes, with mottled underside
markings and a narrow aedeagus and rather long saccus. The facies and the male
genitalia ofjephtha place it in this group: the genitalia are close to those of
M. muscosa (Thieme), the type-species, as illustrated by Forster (1964:154,fig. 185).
We obtained a single worn female of M . jephtha at 2500m to the south of
MCrida. Described from the Colombian Western Cordillera, the species occurs
also in the Eastern Cordillera, where we found it in 1977, and this is the only
known record of it from Venezuela. It is clearly a rarity.
Mygona irmina (Doubleday)
Pronophila irmina Doubleday, [18491: pl. 60, fig. 2. 1 8 syntype, VENEZUELA
[coastal range?] (BMNH), [examined].
Pronophila irmina Doubleday; Westwood, [ 18511:358.
Oxeoschistus irmina (Doubleday);Butler, 1867 :268.
Mygona irmina (Doubleday); Thieme, 1907: 167; Gaede, 1931513; Adams &
Bernard, 1979:107.
2000-2500m. Local and usually solitary, flying feebly and slowly in sunny
clearings, normally about 2m above the ground. The species is found in
Venezuela (including the coastal range), Colombia and Ecuador.
Panyapedaliodes panyasis ( Hewitson)
Pronophila panyasis Hewitson, 1862: 7, pl. 3, fig. 22. 3 8 syntypes, VENEZUELA
and BOLIVIA (BMNH),[examined].
Pedaliodespanyasis (Hewitson); Butler, 1868: 179; Thieme, 1905:81.
Panyapedaliodes panyasis (Hewitson); Forster, 1964: 157 ; Adams 8c Bernard,
1977:273; 1979: 107.
2500-2600m. This species, distributed between Venezuela (CM and the coastal
range) and Bolivia, is as rare in the CM as elsewhere in our experience, perhaps
because of its preference for life high in the forest canopy (see Adams 8c Bernard,
197 7 ). We found two individuals only of each sex.
Pedaliodesjaphleta Butler
(Fig. 7 )
Pedaliodes japhleta Butler, 1870: 24. 19 syntype, [VENEZUELA: CMI (BMNH),
[examined].
19
360
M . J. ADAMS AND G . I. BERNARD
Pedaliodes polemon Grose-Smith, 1900: pl.4, fig. 8. 1 8 syntype, VENEZUELA:
Merida (BMNH),[examined].
Pedaliodes juphleta Butler; Butler, 1874: 180, pl. 62, fig. 5 ; Thieme, 1905:54;
Weymer, 1912: 251; Gaede, 1931:491; Forster, 1964: 158.
Butler’s type of P.japhleta is a female, and Grose-Smith’spolemon type is a male
of the same species, as noted by Thieme (1905). Its closest relatives are
P. leucocheilus Godman 8c Salvin h o m the Sierra Nevada de Santa Marta, P.suspiro
Adams & Bernard from the Serrania de Valledupar, P. poesia (Hewitson) from
Colombia to north Peru, and P. piletha (Hewitson) from the Venezuelan coast
range, all of which share similar male genital structure (Fig. 7). The markings of
japhleta are closest to those ofpiletha, and in size it is intermediate between piletha
and its other relatives.
2000-2600m. An endemic species, not uncommon in the CM. It has a robust
flight, normally remaining more than 1m above ground level. We observed one
oviposition, in which a whitish egg was laid on a small, young leaflet, near the
node of a tall cane of Chusquea bamboo, two feet above ground.
Pedaliodes proerna (Hewitson)
Pronophila proerna Hewitson, 1862:9, pl. 4, fig. 29. 1 8 syntype, COLOMBIA:
Bogota (BMNH),[examined].
Pedaliodes proerna (Hewitson); Butler, 1867:267; 1868: 173; Thieme, 1905:60;
Weymer, 1912:252; Gaede, 1931:502; Forster, 1964: 161; Adams & Bernard,
1979: 109.
Pedaliodes proem fumaria Thieme stat. nov.
Pedaliodes fumuria Thieme, 1905:60, pl. 3, fig. 25. 8 syntypes, VENEZUELA:
Merida [MNHU, Berlin?].
Pedaliodesfumaria Thieme; Weymer, 19 12: 252; Gaede, 1931:490.
The only difference between Thieme’s fumaria and Hewitson’s proerna is the
indistinct whitish wedge on the underside hindwing of fumaria. Although this is
absent in typical proerna, in some specimens from the Serrania de Valledupar it is
present in the form of very faint scales. The male genitalia are indistinguishable.
We consider them to be conspecific, and therefore reducefurnuria in status to be a
subspecies of P. proerna (Hewitson).
1500-2100m. An endemic subspecies, rare in our experience. We found all our
specimens on the ground, and it is as wary and fast in flight as typicalproema is in
Colombia.
Pedaliodes perperna (Hewitson)
Pronophda perpernu Hewitson, 1862: 16. 2 8 , 19 syntypes, [VENEZUELA: coastal
range?] ‘SOUTH AMERICA’ (BMNH), [examined].
Pronophila satyroides Felder & Felder, 1867 : 469. 2 8 syntypes, VENEZUELA:
Caracas (BMN H 1, [examined].
Pedaliodes perperna (Hewitson); Butler, 1868: 173; Thieme, 1905:68; Weymer,
1912:253, pl.53, rowe; Gaede, 1931:495.
Thieme (1905) synonymized satyroides with p e r p m . The BMNH has a solitary
female from the CM. It is otherwise known from between Costa Rica and
Colombia, and the Venezuelan coast range. In our experience, it flies in the
PRONOPHILINI OF M ~ R I D A
36 1
lower reaches of the cloud-forest (1400m in the southern Sierra de Periji, and
1500/1600m in the Venezuelan coast range).
Pedaliodes manis (Felder 8c Felder)
Pronophila manis Felder & Felder, 1867 :469. 2 8 syntypes, COLOMBIA: Bogota
(BM N H 1, [examined].
Pedaliodes manis (Felder 8c Felder); Butler, 1868: 174; Forster, 1964: 165; Adams 8c
Bernard, 1977:274; 1979: 109.
Pedaliodespisonia form manis (Felder 8c Felder); Weymer, 1912:254, pl. 53, row e.
The differences between P. manis and P. pisonia were discussed by Adams 8c
Bernard (1979). Pronophila manis is found from Costa Rica to Peru, and in
Venezuela and Guyana.
1500-25OOm. A common species, not as particular in its requirement for lush
forest as the majority of pronophilines living below the tree-line. I t is predominantly a ground-level butterfly.
Pedaliodes mntagna sp. nov.
(Figs 8, 21)
Male
Forewing length: 25-27 mm. Differs from P. manis in the following respects:
upperside ground colour darker, without distal lightening; the postmedian band
on the underside hindwing, and particularly the whitish markings along its basal
edge, are less well defined, and the orange coloration at the tornus is more
subdued; the hindwing is broader, with a straighter costa. Most easily distinguished by its genitalia: main process of valve with extra tooth; secondary
process broader but more pointed, and much less heavily toothed (Fig. 8).
Female
We have been unable with any certainty to distinguish between female P. manis
and females found flying with P. montagna males and assumed to belong to the
latter species. Females are therefore not included among the paratypes.
Holotype 8, VENEZUELA: Estado MCrida, La Montaiia, S. of MCrida,
2450-2550 m, 19.viii.77 (M.J. 14.1.
Paratypes, locality as holotype: 3 8 , 2350-2450m, 12.iv.75; 4 8 , 2450-2550m,
9.viii.77; l d , 2450m, 10.viii.77; 58, data as holotype; 18, 2450-2550m,
2O.viii.77; Estado MCrida, La Mucuy, near Tabay: 1 8 , 2500m, 16.iv.75; Id,
1800m, 24.iv.75 ( M J . A . and G.I.B.).
The difficulty of distinguishing P. montagna from P. manis is compounded by
sympatry in the upper part of the latter’s altitude range and, in our experience,
their similar habits. We have dissected several males, and there can be no doubt
from the structure of the male genitalia that it is a distinct, though
closely-related, species. Perhaps its nearest relative, however, is P. manneja
Thieme, which similarly prefers higher elevations than manis in the Serrania de
Valledupar, and overlaps with it over a considerable range of altitudes (Adams 8c
Bernard, 1979); it also has the most similar male armature. (Two males of
mannqa in the BMNH collection are almost certainly erroneously labelled as
Irorn Mkrida.1
362
M. J. ADAMS AND C. I. BERNARD
I n the CM, P. montagna can also be confused with P. ferratilis Butler, which,
however, lacks the postmedian band on the underside hindwing and has very
distinct genitalia, placing it in another section of the genus.
2350-2550m. Probably an endemic species. The single male caught at 1800m
at La Mucuy was a long way below its usual range, perhaps because the cloudforest there is especially humid and, up to about 2300m, very disturbed. Only in
the upper 50m of montagna’~range have we failed to find manis, but the latter is
infrequent above 2300m. The males are particularly fond of resting on mossy
rocks or exposed earth on steep, moist banks, and have a sturdy flight.
Peduliodes rmtigua sp. nov.
(Figs 9, 22, 23)
Malr
Forewing length: 27 mm. Upperside unicolorous dark brown, lighter in the
outer third of the forewing. Scent brand on the forewing distinct and very dark,
comprising a streak above cell 2A and a triangular patch between cell Cu2 and
vein M2, entering discal cell.
Underside forewing dark brown, with russet-brown at apex and down outer
margin; a postmedian band, a transverse streak midway down the discal cell, and
a parallel streak at the end of the cell, all of a lighter brown. Underside hindwing
basal half brown speckled with dark brown; distal half dark russet-brown
similarly speckled; a lighter brown postmedian band, narrowing in cells Cu2 and
ZA, with a faint white submarginal spot in Cul; a well-defined white costal
streak, narrow and broken into two parts, in cells R 1 and Rs.
Female
Forewing length: 26.5 mm. Differs from the male in being lighter coloured
throughout, and in having a larger and better defined white costal streak on the
hindwing underside.
Holotype 6,
VENEZUELA, Mhrida, 1897, ex Oberthur Coll. BM 1927-3.
Paratypes, locality as holotype: 18, 19, data as holotype; 1 8 , Jokey Bequest
BM 1934-120 (Briceno).
I n spite of its superficial resemblance to P. prosa Staudinger, P. antigua
possesses distinct male genitalia (Fig. 9) which place it in the P. manis group,
which includes the following species : manis, dejecta (Bates),manneja, montagna and
pomponia (Hewitson).
A n endemic species. While the Serrania de Valledupar has two species from the
nrani.r group, manneja tending to live above manis in altitude, and while in Central
America and Ecuador manis is accompanied by dejecta and pomponia, respectively,
the CM has three species. We did not find antigua, and there is no evidence of its
altitude preferences, but it is possible that it replaces rnontagna at the highest levels
in the forest, being the upper member of a three-species replacing series
(rr~anis/montagna/anti~ua).
Pedaliodesferratilis Butler
(Fig. 24)
Pedaliodesferratilis Butler, 1873: 221. 2d syntypes, PERU (BMNH), [examined].
Pedaliodes ferratilis Butler; Thieme, 1905: 101 ; Weymer, 19 12 : 258 ; Gaede,
1931:490; Forster, 1964: 169.
PRONOPHILINI OF MkRIDA
363
Figures 24-28. Fig. 24. Pedalzodes ferrutilis Butler luteocosta form nov., male. Fig. 25. Penrosada
franciscue sp. nov., male holotype. Fig. 26. The same, female paratype. Fig. 27. R~dondaempetrus
empetrus (Thieme)comb. nov., female. Fig. 28. Rcd4ndn mpchuc bdivmi subsp. nov., male holotype.
(All Figs c. life-size.)
Typical ferrutilzs is uniformly dark brown except for a suffusion of dark
chestnut in the anal area of the hindwing underneath, and a rectangle of orange
scales set in this suffusion. In the CM population, both to the north and south of
Mkrida, about half of the individuals bear a yellow costal streak on the underside
hindwing. This streak varies in size, and at its largest it is ‘L’-shaped, 5mm long,
and 2mm wide at the bend of the ‘L’. In the female it is whitish. Although the
BMNH has specimens of this form from the CM, it has none from other parts of
the species’ range (Colombia to Bolivia), and it is probably that this hitherto
unrecognized polymorphism is confined to the Venezuelan Andes. We name the
new form, luteocosta form nov. (Fig. 24). Its habits and behaviour, and the
structure of its male genitalia, leave us in no doubt about its conspecificity with
P. ferratilis.
2300-315Um. A fast and erratic flier, easily disturbed. Although it is often
found settled on the ground, its commonest haunts are among vegetation about
2 m above ground.
Since Dognin ( 1887) described P. morenoi from Ecuador, it has been considered
synonymous with P. ferratilis (Thieme, 1905 ; Weymer, 19 12; Gaede, 193 1). We do
not agree with this: morenoi has a much brighter underside hindwing patch, is
smaller, and has a longer main process on the valve. In addition, ferratilis also
occurs in Ecuador. Peduliodes morenoi is formally reinstated below as a species in its
own right.
364
M. J . ADAMS AND G . I. BERNARD
Pedaliodes morenoi Dognin, 1887: 18, pl. 1, fig. 2. 1d syntype, ECUADOR: Zamora
(BMNH), [examined]. Stat. rev.
Pedaliodes polla Thieme
[Pronophila polusca Hewitson var.; Hewitson, 1862: 10, pl. 4, fig. 32. Misidentification.] [Specimen lost].
Pedaliodes polusca var. polla Thieme, 1905: 103, Gaede, 1931 :50 1.
Pedaliodes polusca form polla Thieme; Weymer, 1912: 258.
Pedaliodespolla Thieme; Forster, 1964: 17 1.
Hewitson ( 1862) figured two butterflies as polusca, but his description clearly
referred to the larger and darker of the two ( 1862: fig. 3 1). The other, described
by Thieme (1905) as var. polla, was given specific status by Forster (1964).
Although the male genitalia of the two taxa are similar, we concur with Forster’s
decision because we found both polla and polusca in the Cocuy area of the
Colombian Eastern Cordillera, polusca replacing its relative above 2700 m on the
same mountain slopes.
Pedaliodes polla is represented in the BMNH by specimens from between
Panama and Ecuador, including Venezuela, and Forster’s (1964) records of it
from Bolivia are doubtful. Other close relatives are P . tyrrheus Godman & Salvin
i n the Sierra Nevada de Santa Marta, and P. tyrrheoides Adams & Bernard in the
Serrania de Valledupar. Live individuals of tyrrheus and polla were illustrated by
Adams (19771, but the statement in the caption that they are not closely related is
an error.
2,500-3000m.Occurs locally, in small numbers, flying deceptively fast at times,
commonly between 1 and 2m above ground. It is often to be found feeding at
ground level.
Pedaiiodes plotina (Hewitson)
Pronophila plotina Hewitson, 1862:4, pl. 2, figs 9, 10. 18 syntype, VENEZUELA
I:coastal range?] (BMNH), [examined].
Pedaliodes plotina (Hewitson); Butler, 1868: 178; Thieme, 1905: 1 1 7 ; Weymer,
1912:260; Gaede, 1931:500;Adams&Bernard, 1979: 110.
Tvpical plotina was described from Venezuela and the type corresponds to the
coastal range population. Populations with similar individuals occur in the
Serrania de Valledupar and the main Colombian Eastern Cordillera (Adams &
Bernard, 1979).Individuals in the CM have darker markings and are placed by us
as ssp. rapha Butler.
Pedaliodes plotina rapha Butler stat. nov.
Pedaliodes rapha Butler, 1870:24. 1 8 syntype, [VENEZUELA: CMI (BMNH),
[examined].
Pronophila phanaspes Hewitson, 1874:pl. 8, fig. 5 5 . 18 syntype, VENEZUELA
[: CMI (BMNH), [examined]. syn. nov.
Pedaliodes plotinella Grose-Smith, 19OO:pI. 4, figs 3, 4. 1 8 syntype, VENEZUELA:
Merida (BMNH),[examined].
Pedaliodes rapha Butler; Thieme, 1905: 117.
Pedaliodes plotina form rapha Butler; Weymer, 1912 : 260.
Pednliodes plotina form pharnmpes (Hewitson);Weymer, 1912 : 260.
PRONOPHILINI OF MkRIDA
365
Although Butler’s rapha referred to an individual whose markings were
particularly dark and small, and Hewitson’s phanzaspes described an individual
which is more representative of the CM population, the former name has
priority. Both clearly came from the same CM population, and are therefore
synonymous. A gradation exists between rapha-like and pharnaspes-like
butterflies, but none in the CM is as brightly marked as typical plotina. The
breaking up of the underside hindwing wedge into a series of crescent-shaped
spots, a feature of the rapha type, is not characteristic of very dark individuals, but
occurs in a proportion of all the butterflies of the subspecies (and indeed of the
typical race also). Thieme (1905) gave the synonymy of Grose-Smith’s plotinella
with rapha; although he classed rapha as a distinct species, he expressed doubt as
to whether it may not be a local form ofplotina.
The valve of plotina is of a shape unusual in the genus, with its blunt dorsal
extension possessed otherwise only by P.phazania Grose-Smith from the Sierra
Nevada de Santa Marta. The two are most probably sister-species.
1800-2600m.The commonest pronophiline between 1800 and 2500m, but an
abrupt drop in numbers occurs above 2500m. I t is a medium fast flier, usually
close to the ground and resting on it or on low foliage. We found a male at night
settled underneath a bamboo leaf about 2 m above ground.
Pedaliodes ornata Grose-Smith & Kirby
(Fig. 10)
Pedaliodes ornata Grose-Smith % Kirby, 1895: [1371, figs 1, 2. ld syntype,
VENEZUELA: Merida (BMNH), [examined].
Pedaliodes phaeaca Staudinger, 1897: 123, pl. 6, fig. 1. 4d, 1Q syntypes,
VENEZUELA: Merida (MNHU, Berlin).
Pedaliodes ornata Grose-Smith & Kirby; Thieme, 1905: 125; Weymer, 1912 : 26 1;
Gaede, 1931:492.
As judged by its facies and male genitalia (Fig. 101, P. ornata probably belongs
within a monophyletic group including P. cesarense Adams & Bernard from the
Serrania de Valledupar, and P. phaea (Hewitson) from the main Colombian
Eastern Cordillera, both of which fly in the same range of altitudes. Pedaliodes
ornata lacks the orange band on the upperside hindwing possessed by the others,
except for a dusty yellowish patch at the apex. Pedaliodes phaeaca Staudinger is a
synonym of onzata (Thieme, 1905).
2300-2900m. An endemic species, erroneously described from Bolivia by
Grose-Smith & Kirby (1895), and not included in the Bolivian list by Forster
(19641, although he figured its male genitalia (1964: 174, fig. 257). It is the
commonest pronophiline at the particularly rich elevations of 2500-2700 m,
normally flying l m or more above the ground, but often resting and feeding
among leaf litter.
Penrosadafianciscae sp. nov.
(Figs 11, 25, 26)
Male
Forewing length: 20.5-2 1.5 mm. Upperside unicolorous dark brown, slightly
lighter distally. The hindwing has a small submarginal white-pupilled black
366
M. J . ADAMS A N D G . I. BERNARD
ocellus in cell Cul , faintly ringed with chestnut-brown, a faint submarginal dark
brown line, and the postmedian yellow band of the underside shows through
very faintly.
Underside forewing as upperside, but with faint darker brown marginal and
submarginal lines, and a series of 5 tiny submarginal white spots in cells R4 to
M3 (sometimes very faint). Underside hindwing dark brown, with a yellow
postdiscal band from costa to inner margin, of uniform width (c. 1.0-1.5 mm)
but tapering in cell 2A and displaced distally in R1; a series of 7 submarginal
spots, as white-pupilled chestnut-ringed black ocelli in cells Cu2 (two),Cul and
M i , as small white dots in M 2 and M3, and as a white dot faintly ringed with
black in Rs (sometimes absent).
Female
Forewing length: 2 1.5-22 mm. Ground colour slightly lighter, forewing
narrower than in male. Upperside forewing with a submarginal white-pupilled
chestnut-ringed black ocellus in cell Cul, larger than the one on the hindwing, a
tiny ocellus in M 1, and a faint dark brown submarginal line. Upperside hindwing
as male, but the ocellus is a little larger, and (in some specimens) there is a
minute black submarginal dot in cell Cu2.
Holotype 8,VENEZUELA: Estado Mkrida, above La Montafia, S. of MCrida,
2500-2600m, 14.vi.75 ( M J . A . 1 .
Paratjpes, locality as holotype: 1 7 8 , 29, data as holotype; 58, 2600m, 15.vi.75
( 1 Universidad Central, Venezuela); 8 3 , 2450-2550 m, 9.viii. 7 7 ; 6$, 3 9 ,
2450-2550m, 19.viii.77 (19 Universidad Central, Venezuela); below La
Montaiia, 18, 2300m, 4.viii.77 (h4J.A. and C.Z.B.1; Mkrida, 18,Jokey Bequest
BM 1934-120 (Brzceno).
Penrosada franciscue is closest in appearance to P. leaena (Hewitson)from Peru,
and P. satura (Weymer) from Ecuador and north Peru. The yellow band on its
underside hindwing is twice as broad as that of P. lanassa (Felder & Felder) from
the Colombian Eastern Cordillera. Parosada leuena has no ocellus on the upperside hindwing, and has smaller and fewer submarginal ocelli on the underside
hindwing (usually 4 white dots, surrounded by black, in cells Cu2 to M2, and a
single one only in Cu2). The ocelli in satura are better developed than infranciscue,
and while the latter’s lie nearer to the outer margin than to the yellow band, in
satura they lie closer to the band. The male genitalia offranciscue (Fig. 11) are
nearest to satura, but at the tip of the valvus there are 5 equal-sized teeth, while
satura has 3 larger ones, the third being especially long and broad; leaena has 4,
and unlike the other two species, it has no other teeth further down the valvus.
Brown ( 1944),who erected the genus Penrosada, erroneously figured the genitalia
of lanasa under leaenu (his fig. 161.31, and of an unnamed species similar to
lunarsa under lanassa (his fig. 1616).
The species is named after Professor Francisco Fernandez YCpez, of the
Department of Agricultural Zoology at the Central University of Venezuela,
Maracay, in gratitude for his help, encouragement and friendship.
2300-260Um. An endemic species. I t was apparently absent in April 1975 from
the site at which we obtained a series in June 1975, and in August 1977 it was
quite common there. Other cases of seasonality in species of Penrosada in the
Serrania de Valledupar were reported by Adams & Bernard (1979). We did not
hidfranciscue to the north of MCrida. The butterfly flies with a skipping action,
PRONOPHILINI OF M ~ R I D A
56 7
usually more than 2m above ground, and rests on foliage. The ova, which are
pink at first, are laid on the brownish petioles of young leaflets, near the nodes of
tall ‘bolted’canes of Chusquea bamboo, between 1 and 4 m above the ground.
Pronophila bogotensis Jurriaanse
Pronophila bogotensis Jurriaanse, 1926: 51. 8 holotype, COLOMBIA: Bogod
(presumablyin Rijksmuseum van Natuurlijke Historie, Leiden).
Pronophila bogotensisJurriaanse; Forster, 1964: 183; Adams 8c Bernard, 1977 :277;
1979: 115.
Pronophila bogotensis has often been confused with P. thelebe Doubleday, which
only occurs in the Venezuelan coast range (and’perhaps in the Venezuelan
Pantepui mountains). It is larger than thelebe, and with a much darker underside;
the white postmedian patch in cell M2 of the forewing is smaller and does not
appear continuous with the main patch in M 1 as in thelebe. P. bogotmis is found in
Colombia, Ecuador and the CM.
1500-2000m. It flies robustly, typically between 2 and 5m above ground level,
rarely descending to the forest floor.
Pronophila epidipnis Thieme
(Fig. 12)
Pronophila epidipnis Thieme; 1907 :204. 4 8 syntypes, VENEZUELA: Merida
[MNHU, Berlin?].
Pronophila epidipnis Thieme; Weymer, 1912: 273, pl. 60, row a ; Gaede, 1931: 5 18.
This species probably forms a monophyletic group with P. pliani Adams &
Bernard from the Sierra Nevada de Santa Marta, and P. orchewitsoni Adams 8c
Bernard from the Colombian Andes and Ecuador, as judged by their underside
markings and altitude preferences. Pronophila epidzjmis has the darkest upperside
forewing, the whitish subapical markings possessed by the other species being
represented by only a faint lightening of the dark brown ground colour. The
male genitalia are shown in Fig. 12, and are as featureless as those ofalf the other
Pronophila species we have examined.
2000-2700m. An endemic species, which replaces its congener P. bogotensis
above 2000m, and has similar habits. In our experience it is uncommon.
Redonda gen. nov.
Type-species: Pedaliodes empetrus Thieme
Eyes hairy, black and chestnut-brown. Palpi twice as long as head, dark brown
with yellow and dark brown hairs, and white scales especially at base; first
segment one-quarter the length of the second; third segment very short (threequarters the length of the first).Antennae to two-fifthscosta; orange-brown shaft,
darkening to dark brown distally; club 2.5 times longer than broad, dark brown
and chestnut-brown, concave.
Venation. Forewing: Sc, R1 and R2 all independent; root of R3 nearer to root
of R4 than to discal cell; no R-M1; Ml-MZ curves only slightly inwards.
368
M. J. ADAMS AND G . I. BERNARD
Hindwing: humeral vein moderately developed; root of Cu 1 closer to that of M 3
than that of Cu2; root of M3 halfway between those of Cu 1 and M2.
Female much smaller than male. Male forewing tornus obtuse, hindwing
rounded with outer margin almost unindented. Female wings narrower, with
more pointed forewing apex.
Gncus equal to or shorter than tegumen; subunci lacking.
Redonda empetrus (Thieme) comb. nov.
(Fig. 13)
Pedaliodes empetrus Thieme, 1905:98, pl. 1, fig. 7. ld syntype, VENEZUELA:
Merida [MNHU, Berlin?].
Pedaliodes empetrus Thieme; Weymer, 1912: 258; Gaede, 1931: 489.
Thieme ( 1905)was clearly doubtful about his placing of empetrus in the genus
Pedaliodes, and he pointed out its superficial resemblance to species of the southtemperate pronophiline genus, Argyrophorus, and in particular to A . lamna
Thieme. The male genitalia of lumna, although they have well-developed subunci,
are somewhat similar to those of empetrus (Fig. 131, and a close, even if not recent,
phyletic relationship between Redonda and Argyrophorus cannot be ruled out. The
only other prmophiline species known to us, in which the female is much
smaller than the male, is Dangond dangondi Adams 8c Bernard from the paramo of
the Serrania de Valledupar; and it is possible that Dangond and Redonda, both
monobasic, are relict genera which have their origins in an early south-temperate
pronophiline fauna.
The species occurs in two distinct races in the CM.
Redonda empetrus empetrus (Thieme)
(Fig. 27)
The typical subspecies was found by us in the Mucubaji region north-east of
Mkrida, and at the head of the Albarregas valley, north of the city. The female
has not been described before; it is illustrated in Fig. 27.
Female
Forewing length: 18-20.5mm (cf. 25-27 mm of males). Upperside ground
colour varies between dull silvery white and light sandy brown, with a dark
brown forewing costa and hindwing inner margin, and outer margins chequered
ground colour and dark brown; submarginal white streaks on both wings
showing through faintly; in the darkest specimen, a narrow lighter postdiscal
band crosses the forewing from costa to vein 2A.
Darkness of underside varies with upperside; markings as in male, but the
submarginal white patches on the hindwing are extended basad.
Redonda empetnu bolivari subsp. nov.
(Fig. 28)
Male
Forewing length: 27-31mm. Differs from the typical subspecies in the
following respects: larger (mean forewing length 28.7mm7 cf. 25.5mm in ssp.
PRONOPHILINI OF MkRIDA
369
empetrus); darker on the upperside; a deeper orange-brown on underside
forewing; darker ground colour on underside hindwing, but with a more clearlydefined narrow whitish postmedian band; the submarginal white spots (in cells
Cu 1 to Rs, and sometimes in Cu2) are almost round and not extended distally to
form -< -shapes on their side as in typical empetrus.
Female unknown.
Holotype 8,VENEZUELA: Estado Mkrida, Loma Redonda, S. of Mkrida,
4000m, 21.iv.75 (MJ.A.1.
Paratypes, locality as holotype: 1 8 , data as holotype; 18,4000m, 18.iv.75; 3 8 ,
3800m, 19.iv.75;48,4000m, 19.iv.75; 68,4000m, 20.iv.75(MJ.A.andG.Z.B.).
??00-4000m. An endemic species, restricted to the paramo. In the areas where
ssp. empetrus flies, which receive relatively low precipitation and more sunshine, it
is mostly confined to marshy habitats. Ssp. bolivari was-found on open, flowery
and rock-strewn mountainsides, resting among grass or on lichen-covered
boulders. It has a fairly slow flight, usually close to the ground. All the females
we saw were in a very marshy area; they were loath to fly even when disturbed,
and after very short flights disappeared among low tussocks of grass.
Steroma bega Westwood
Steroma bega Westwood, [18511:400, pl.66, fig. 6. l d syntype, VENEZUELA
[:coastal range?] (BMNH),[examined].
Steroma zibia Butler, 1870: 23. 19 syntype, VENEZUELA (BMNH),[examined].
Steroma bega Westwood; Weymer, 1912: 241; Gaede, 1931:480; Forster,
1964: 141; Adams &Bernard, 1979: 116.
Adams & Bernard (1979) formally synonymized S. zibia with S. bega. The
Venezuelan Andean population is referable to typical bega with its almost
complete silvery patch at the costa of the underside hindwing: this race is
distributed down the eastern slopes of the Colombian Eastern Cordillera and
perhaps as far as Bolivia, and the CM and the Venezuelan coastal range (where
the type almost certainly came from).
2300-2800m. Predominantly found on the ground or on rocks near the
ground, and in marshy areas or near streams, flying with a skipping action close
over the forest floor.
Thiemeia phoronea (Doubleday)
Pronophila phoronea Doubleday, [ 18491: pl. 60, fig. 1. l d syntype, VENEZUELA
[:coastal range?] (BMNH),[examined].
Pronophilaphoronea Doubleday; Westwood, [ 18511:358 ; Hewitson, 862: 14.
Daedalmaphoronea (Doubleday);Butler, 1868: 183.
Catargynnisphoronea (Doubleday);Thieme, 1907 : 150.
T h i e ~ e i phoronea
a
(Doubleday);Weymer, 1912:267.
Thimeia [sic!] phoronea (Doubledayj; Gaede, 1931:!ill; Forster, 1964: 179.
This butterfly, reminiscent of species of Catargynnis but with very distinct male
genitalia and ‘tails’ on the hindwing, was described from Venezuela (probably
the coastal range), and the BMNH collection contains specimens from the CM,
Colombia and Ecuador. We did not find it.
370
M . J . ADAMS AND G . I . BERNARD
SPECIES OF DOUBTFUL STATUS IN THE CORDILLERA DE MLRIDA
Eretn’s apulqu (Felder & Felder)
Pronophila apuleja Felder & Felder, 1867 : 47 1 . 18 syntype, [ECUADOR?]
‘VENEZUELA’(BMNH ), fexaminedl.
Pronophila phyllalia Hewitson, 1868:pl. 4, figs 21, 22. 18 syntype, ECUADOR
(BMNH), [examined]. syn. nov.
Eretris upuleja (Felder & Felder); Butler, 1868 : 1 7 5 ; Thieme, 905 : 132 ;Weymer,
1912:263, ~ 1 . 5 6rowb;
,
Gaede, 1931:504.
Eretrisphyldzliu(Hewitson);Thieme, 1905: 133.
Eretris apuleja formphyllalia (Hewitson);Weymer, 19 12: 263
Eretris upuleJa var. phyllalia (Hewitson); Gaede, 193 1 :504.
The types of apuleja and phyllalia are similar enough to suggest that they may
have come from the same population. The BMNH has a series of phyllalia from
Ecuador, but no specimens of apulqu from its type-locality of ‘Venezuela’ other
than the solitary type, and we did not find it in the CM or the coastal range. Since
the closely-related E . ochrea Thieme flies in the mountains directly between the
type-localities of phyllalia and apuleja (in the main Colombian Eastern Cordillera),
we consider that the Venezuelan locality for apuleja is an error, and suggest that it
and phyllalia are synonymous for an Ecuadorian species.
Zdioneurula erebioides (Felder & Felder)
Idioneura erebioides Felder & Felder, 1867 : 474. 58 syntypes, COLOMBIA: Bogota
(BMNH 1, [examined].
Zdioneura erebioides Felder & Felder; Butler, 1868:99; Gaede, 1931 : 479.
Zdioneurula erebioides (Felder & Felder); Strand, 1932: 146.
Because Felder & Felder’s Zdioneura is a junior homonym (of a fly genus),
Strand ( 1932) proposed the new name Zdioneurula for this monobasic genus. We
doubt that it occurs in Venezuela, and consider that the single specimen in the
BMNH among a series from Colombia was labelled as from ‘Venezuela’ in error.
In the Colombian Eastern Cordillera we have found it between 2250 and
2850m, in short grass on open land, marsh or near streams, where it flies rather
slowly, close to the ground.
Lmiophila circe Felder & Felder
Laszophila circe Felder 8c Felder, 1859:326. 18 syntype, [no locality] (BMNH),
[examined].
Pronaphila praeneste Hewitson, 1860:pl. 2, fig. 8. Lectotype 8,
COLOMBIA: New
Granada (BMNH), here designated [examined].
Lasiophila circe Felder & Felder; Butler, 1868: 181; Thieme, 1907: 123; Weymer,
1912:265; Gaede, 1931:506.
Hewitson ( 1860) figured two specimens of Pronophila praeneste, believing one
(figs 7 , 9) to be the female of the other (fig. 8). The types (both males!) are in the
BMNH. As Thieme (1907) pointed out, the two specimens belong.to distinct
species, fig. 8 representing L. circe Felder & Felder, and figs 7 and 9 representing
L. cirta Felder & Felder (both described in 1859). We believe thatpraeneste should
be synonymized with circe (following Thieme, 1907 : 1231, since it was described
PRONOPHILINI OF
MBRIDA
37 1
from Colombia where circe is found, while cirtu occurs in upper Amazonian Peru;
to stabilize this synonymy, we hereby designate the circe male of Hewitson’s
syntypic series as the Lectotype of Pronophila praeneste Hewitson.
Adding to this confusion is the fact that the Felders described L. circe from
Venezuela, although the type specimen now has no locality label, and the species
has not been found there since. All the BMNH specimens come from Colombia,
and it is most probably confined to that country. Kruger (1924:35) found it in the
Eastern and Central Cordilleras between 2600 and 3200m, and we obtained it in
the eastern range at 2650 and 2700m. It replaces L. rapatora in the uppermost
forests, and has a similar lazy flight.
ACKNOWLEDGEMENTS
We are indebted to the following institutions and individuals who helped to
fund the third ‘North Colombia Butterflies Expedition’ of 1974/1975:The Royal
Society; Percy Sladen Memorial Fund; Explorers’ Club of New York; Royal
Geographical Society; World Expeditionary Association; Mount Everest
Foundation; Lloyds Bank International; British Museum (Natural History); Mr
& Mrs R. W. Adams; Mr Robert Ginzburg; and Mrs C. Bernard and to
Bryanston School and Mr Robert Ginzburg for loans towards the cost of the 197 7
expedition. We are grateful for the help and hospitality extended to us by
Professors F. Fernandez YCpez, Josk V. Scorza and Pedro Durant, and their staffs
at the Universidad Central in Maracay and the Universidad de 10s Andes in
Mkrida; Ingos. Ignacio Rubio and Jaime Grimaldo, and their colleagues at the
Ministerio del Ambiente y de 10s Recursos Naturales y Renovables in Mkrida;
Robert Dietz IV; Dr Carlos Bord6n; Earl and Gail Aagaard; and the Dolfo
family. For regular access to the collections at the BMNH, the loan of material
and advice on the preparation of this paper, we thank Dr Paul Freeman, R. I.
Vane-Wright, P. R. Ackery and R. L. Smiles. The photographs were produced by
the photographic unit of the BMNH, and the plates were prepared by Ms R.
Arora.
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