ArticlePDF Available

Abstract and Figures

Phylogenetic analysis of 53 morphological characters for five species of Panacea and Batesia hypochlora supports the separation of the two genera and showed that the monotypic genus Batesia is basal to Panacea. Male genitalia were uniform within Panacea and characters informative for phylogeny reconstruction were restricted to wing coloration. Illustrations of adults and genitalia, a brief diagnosis, and distributions are provided for each species.
Content may be subject to copyright.
By possessing distasteful wings or body fluids, brightly
colored butterflies are generally avoided by many verte-
brate predators in nature. This phenomenon is particu-
larly well known in various genera of Nymphalidae (e.g.,
Acraea, Heliconius, many Danainae and Ithomiinae),
Papilionidae (e.g., Battus, Parides) and Pieridae (e.g.,
Mylothris, Delias, Appias, Perrhybris, Itaballia) among
others (see Poulton 1908, Sywnnerton 1919, Carpenter
1942, Fisher 1958, Chai 1986). Nevertheless, a great
many of these same butterflies are eagerly sought after
and prized by a different group of predators, human col-
lectors. Although collector value may provide a metric of
how garishly colored a particular butterfly might be, it is
often a poor measure of how well we understand that
species. Therefore, when considering biological or evo-
lutionary understanding of particular butterflies, it is
likely that drab ones are equally as well known as those
that are brightly colored. Although well represented in
museum collections, and available as virtual specimens
on the internet, nymphalid butterflies in the genera
Batesia Felder and Felder, 1862 and Panacea Godman
and Salvin, 1883 are good examples of this phenomenon.
The Neotropical genus Batesia occurs from central
Colombia to eastern Ecuador, southeast Peru, western
Brazil, and likely into northeast Bolivia; effectively an
upper Amazonian distribution. On the other hand,
members of Panacea are found from Costa Rica south
across Venezuela and the Guianas, throughout the
Amazon basin, and into Bolivia.
Both Batesia and Panacea were originally described
as monotypic genera, but only Batesia with its single
species, hypochlora Felder and Felder, 1862 has re-
mained so. The history of Panacea is somewhat convo-
luted. Panacea prola (Doubleday, 1848) was initially
designated the type species of Pandora Doubleday,
1848—a name used previously for different insect gen-
era by at least seven different authors, and thus, an in-
valid homonym (see Hemming 1967). In an attempt to
settle this quandary, Kirby (1871) transferred all species
of Pandora to Batesia. Godman and Salvin (1883), how-
ever, felt that all species formerly in Pandora warranted
separation from Batesia, and erected the genus Panacea
to accommodate them—thus providing a panacea to the
Pandora problem. Eight species have been described in
PanaceaP. prola; P. procilla (Hewitson, 1852); P.
regina (Bates, 1864); P. divalis (Bates, 1868); P. chal-
cothea (Bates, 1868); P. lysimache Godman and Salvin,
1883; P. bleuzeni Plantrou and Attal, 1986; and P. bella
D’Abrera, 1987, not all that are currently regarded as
valid species (see synonymies below).
The vicissitudes of nomenclature aside, nearly all
natural history studies suggest that Batesia and
Panacea are distinct, but closely related genera. At
present they are classified in the Biblidini along with
Hamadryas, Ectima, Eunica, Myscelia, Dynamine,
Colobura and other genera (Godman & Salvin 1883,
Seitz 1916, Ackery 1984, Harvey 1991).
Recent observations indicate that Batesia and
Panacea share Caryodendron spp. (Euphorbiaceae) as
host plants, and that their immature stages are very
similar (DeVries et al. 1999). The correspondence of
immature biology, classification, and the fact that these
genera have never been assessed using cladistic meth-
ods led us to ask whether B. hypochlora was separate
from Panacea, or if it represented a derived species
within Panacea. Accordingly, this study tests both hy-
potheses through phylogenetic analysis of five species
of Panacea plus Batesia hypochlora. Based on adult
morphology we show that Batesia hypochlora is basal
to Panacea, and that together they form a mono-
phyletic group. We then present characters to aid in
species identification, and provide notes relevant to fu-
ture work on their taxonomy and natural history.
Journal of the Lepidopterists’ Society
56(4), 2002, 199–215
PHYLOGENETIC ANALYSIS AND REVIEW OF PANACEA AND
BATESIA BUTTERFLIES (NYMPHALIDAE)
RYAN I. HILL
Section of Integrative Biology, University of Texas, Austin, Texas 78712, USA
CARLA M. PENZ1AND P. J. DEVRIES
Milwaukee Public Museum, 800 W Wells St., Milwaukee, Wisconsin 53233, USA
ABSTRACT. Phylogenetic analysis of 53 morphological characters for five species of Panacea and Batesia hypochlora supports the sepa-
ration of the two genera and showed that the monotypic genus Batesia is basal to Panacea. Male genitalia were uniform within Panacea and char-
acters informative for phylogeny reconstruction were restricted to wing coloration. Illustrations of adults and genitalia, a brief diagnosis, and dis-
tributions are provided for each species.
Additional key words: prola, procilla, regina, divalis, bleuzeni, chalcothea, lysimache, bella, hypochlora, Caryodendron, Euphorbiaceae.
1Adjunct professor at Pontifícia Universidade Católica do Rio
Grande do Sul, Av. Ipiranga 6681, Porto Alegre, RS, 90619-900,
Brazil.
MATERIALS AND METHODS
Species studied. Excepting P. chalcothea (see
identification section below), our phylogenetic analysis
included all valid species of Panacea (P. prola, P.
procilla, P. regina, P. divalis, and P. bleuzeni) and Bate-
sia hypochlora (Figs. 1–10).
To assess intra-specific variation in wing pattern and
genitalia, we examined specimens from five distinct lo-
calities. Abundant material from a single site in eastern
Ecuador (P. prola, n = 57; P. divalis, n = 55; P. regina,
n = 43; and B. hypochlora, n = 24) allowed us to eval-
uate morphological and phenotypic variation within a
single population (see DeVries & Walla 2001 for site
description). Whenever possible individuals from dif-
ferent localities were dissected to evaluate morpholog-
ical variation in the genitalia. Although a small number
of specimens were available of P. procilla (n = 4) and P.
bleuzeni (n = 2), these species are phenotypically dis-
tinctive from other Panacea and characters could be
scored with confidence. For P. bleuzeni, one specimen
of each sex was used to score genitalia characters di-
rectly, but wing and body characters were scored using
the description of Plantrou and Attal (1986), the illus-
trations in D’Abrera (1987:487, as P. bella) and pho-
tographs from the private collection of G. Attal. Charac-
ters 22 and 23 were scored as “missing” for P. bleuzeni
due to lack of material. Table 1 lists the examined taxa,
number of dissected individuals, and locality data.
We used Biblis hyperia (Cramer, 1780) and
Hamadryas arinome (Lucas, 1853), H. amphinome
(Linnaeus, 1767), H. laodamia (Cramer, 1777), and H.
feronia (Linnaeus, 1758) as outgroup taxa for phyloge-
netic analysis. Based on larval and adult morphology,
and host plant use (Euphorbiaceae) these taxa are con-
sidered closely related to Batesia and Panacea (Seitz
1916, Ackery 1984, Harvey 1991).
Preparation of material. Genitalia were prepared
with a standard treatment of 10% potassium hydroxide,
examined with a stereomicroscope, and subsequently
stored in glycerol. Illustrations are given in Figs. 11–13.
Characters and terminology. Our character matrix
200 J
OURNAL OF THE
L
EPIDOPTERISTS
’ S
OCIETY
FIG. 1. Batesia hypochlora, dorsal. Top row, males; bottom row, females. Left column, Garza Cocha, Ecuador; right column, Rondonia, Brazil.
includes 53 characters (43 binary and 10 multistate), of
which 24 were derived from males (23 from genitalia,
one from wing coloration), 7 derived from females (6
from genitalia and one from wing coloration), and 22
from both sexes (16 from wing patterns, four from ve-
nation, one from forelegs and one from body scales).
Terminology for adult external morphology follows
Scoble (1992). Terminology for male and female geni-
talia follows Klots (1970) except for the use of hypan-
drium and ramus, which follow the definitions in the
glossary of Tuxen (1970) and Jenkins (1986, 1987,
1990). We use hypandrium to mean “a male subgenital
plate,” and ramus as “lateral or ventro-lateral process of
male eighth sternite, directed posteriorly” (see glossary
in Tuxen 1970; Jenkins 1983, 1986). In character 10 we
follow D’Abrera (1987) where a “complete ocellus”
consists of a spot surrounded by a round ring (e.g., P.
procilla, Fig. 6), and an “incomplete ocellus” is a spot
without a round outer ring (e.g., P. bleuzeni, Fig. 7).
Phylogenetic analysis. We used a heuristic search
in PAUP 3.1 (Swofford 1993) with all characters given
equal weight, multi-state characters unordered, poly-
morphic characters treated as exhibiting both states,
and the search used a TBR branch swapping routine.
Following analysis, Biblis hyperia was used to root the
tree. Branch support was estimated by 500 bootstrap
replicates, and we used MacClade 3.01 (Maddison &
Maddison 1992) to identify character changes along
the branches of the tree. The character list and data
matrix are in Appendix 1 and 2.
RESULTS
Phylogeny
Our analysis indicates that Panacea and Batesia are
monophyletic, sister taxa. The single most parsimo-
nious tree (tree length = 79, CI = 0.82, RI = 0.88) sug-
gests that Batesia hypochlora is a sister species to
Panacea, a relationship supported by four characters
(Fig. 14; Table 2, clade 1). We found 11 autapomor-
phies for B. hypochlora (Table 2, clade 2), and nine
characters that justify the monophyly of Panacea
(Table 2, clade 3). Our analysis also showed that all
members of Panacea are morphologically similar, but
they differ strongly from Batesia hypochlora.
Among Panacea the genital morphology was notably
conservative, and characters providing the basis for in-
ferring species relationships were derived mostly from
wing morphology. Only one male genital character (hy-
V
OLUME
56, N
UMBER
4 201
FIG. 2. Batesia hypochlora, ventral. Left column, Garza Cocha, Ecuador; right column, Rondonia, Brazil.
pandrium, character 28) could be used to distinguish
among Panacea species. However, as it represents an au-
tapomorphy for P. divalis, character 28 was uninforma-
tive for establishing phylogenetic relationships within
Panacea. The grouping of P. regina, P. divalis, P. bleuzeni
and P. procilla was supported by seven characters, all de-
rived from wing pattern morphology (Table 2, clade 4).
One character justified grouping P. divalis, P. bleuzeni
and P. procilla (Table 2, clade 5) and a single character
grouped P. bleuzeni and P. procilla (Table 2, clade 6).
Identification and Taxonomy
Here we provide synonymies, characters for identifi-
cation of the study taxa, approximate geographical dis-
tributions, and comments on phenotypic variation of
the species included in our analysis. For completeness,
we also provide taxonomic notes on P. chalcothea, al-
though we did not examine this taxon directly.
Batesia Felder and Felder, 1862
Batesia Felder and Felder, 1862. Wien. ent. Monats.
6:112.
Batesia hypochlora Felder and Felder, 1862
(Figs. 1, 2, 11, 13)
Batesia hypochlora Felder and Felder, 1862. Wien.
ent. Monats. 6:113
Batesia hypochlora hypoxantha Salvin and Godman,
1868. Ann. Mag. Nat. Hist. (4)2:147
Batesia hypochlora hemichrysa Salvin and Godman,
1868. Ann. Mag. Nat. Hist. (4)2:147
Batesia hypochlora chrysocantha Fruhstorfer, 1915.
Soc. ent. 30(12):66
Batesia hypochlora f. intermedia Michael, 1931. Ent.
Zeit. 44(20):309–312
Species characters. Forewing dorsal surface
dark iridescent blue from basal to submedial areas, a
prominent postmedial red band surrounded by
black, apex iridescent blue. Hindwing dorsal surface
mostly iridescent blue, with a postmedial black band
and an iridescent blue marginal band from apex to
tornus. Forewing ventral surface dark brown from
basal to submedial areas and tornus, postmedial red
band surrounded by brown, subapex yellow. Hind-
202 J
OURNAL OF THE
L
EPIDOPTERISTS
’ S
OCIETY
FIG. 3. Panacea prola, dorsal and ventral. Top row, left, male; right, female. Bottom row, left male; right, female. All from Garza Cocha, Ecuador.
wing ventral surface chalky yellow with a distinct
black postmedial band and yellow marginal band
from apex to tornus.
Distribution. Western Amazonas, Brazil; Ecuador,
Peru (Seitz 1916, D’Abrera 1987, Austin & Emmel
1990, Robbins et al. 1996).
Variation. Judging by the named subspecies (see
synonomic list) the intensity of yellow on the ventral
surface of the HW may vary. However, whether
these names are biologically meaningful remains
uncertain. We found little variation in our samples
from Garza Cocha, Ecuador, although we note that
Ecuadorian and Brazilian material differ in the
respective width of the forewing subapical band
(Fig. 1).
Panacea Godman and Salvin, 1883
Pandora Doubleday, 1848. Gen. Diurnal Lep. p. 300
Pl. 3 fig 5
Panacea Godman and Salvin, 1883. Biol. Centr. Am.
pp. 274–275
Panacea prola (Doubleday, 1848)
(Figs. 3, 11, 13)
Pandora prola Doubleday, 1848. Gen. Diurnal Lep. p.
300 Pl. 3 fig. 5
Panacea prola female f. dubia Kretzschmar 1894.
Deutsche ent. Zeit. “Iris” 6(2):158–160
P. prola zaraja Fruhstorfer, 1912. Ent. Rundschau
29(6):46
P. prola amazonica Fruhstorfer, 1915. Soc. ent.
30(12):66
P. prola prolifica Fruhstorfer, 1915. Soc. ent. 30(12):66
P. prola amazonica f. bronzina Bryk, 1953. Arkiv. Fur
Zool. 5(1):1–268
Species characters. Dorsal surface with broken
blue-green iridescent bands. Forewing dorsal surface
without a subapical line in both sexes, but some fe-
males with a faint greenish-white subapical band.
Hindwing dorsal surface without ocelli or blue sub-
marginal line. Hindwing ventral surface bright red,
generally without black markings, but sometimes with
a faint black submarginal line.
V
OLUME
56, N
UMBER
4 203
FIG. 4. Panacea regina, dorsal. Top row, male; bottom row, female. All from Garza Cocha, Ecuador.
Distribution. Panama, Colombia, Venezuela,
Guianas and upper Amazon basin (Seitz 1916,
D’Abrera 1987, Emmel & Austin 1990, Otero &
Romero 1992, Lamas 1994, Robbins et al. 1996, Neild
1996).
Variation. We found wide variation in wing length,
but little variation in color pattern in large samples
from Garza Cocha, Ecuador. Small individuals appear
to be the result of caterpillars feeding on poor quality
Caryodendron leaves, or those that were semi-starved
(pers. obs.).
Subspecies. Panacea prola zaraja, from Venezuela,
Merida; P. p. amazonica, from the upper Amazon; P. p.
prolifica, from Ecuador.
Panacea regina (Bates, 1864)
(Figs. 4, 5, 11, 13)
Pandora regina Bates, 1864. J. Entom. 2(10):213.
Panacea regina victrix Fruhstorfer, 1915. Soc. ent.
30(12):66.
Species characters. Dorsal surface with broken
blue-green iridescent bands. Forewing ventral surface
with reddish apex and white subapical band but with-
out the distinct red spots outlined by black in discal
cell (see P. divalis). Hindwing dorsal surface with a
blue medial band adorned with incomplete black
ocelli that vary in size, and may reach the distal margin
of the band; submarginal wavy line sometimes faint.
Hindwing ventral surface red with broken submedial
to medial transverse black lines, the most distal start-
ing at Sc + Rs and ending at Cu2; faint post-medial
ocelli in almost all cells; conspicuous black submar-
ginal line. Females often with a short, white longitudi-
nal stripe in ventral hindwing cell M2–M3, nearly at the
center of wing.
Distribution. Western and upper Amazon
(Ecuador, Peru, Brazil) (Seitz 1916, D’Abrera 1987,
Lamas 1994, Robbins et al., 1996).
Variation. In Ecuadorian and Brazilian samples we
found that the medial ocelli on the dorsal hindwing vary
considerably within populations. In females we found
the ventral hindwing ocelli were sometimes incomplete.
Subspecies. Panacea regina victrix, from Ecuador;
see also P. chalcothea (below).
204 J
OURNAL OF THE
L
EPIDOPTERISTS
’ S
OCIETY
FIG. 5. Panacea regina, ventral. Top row, male; bottom row female. All from Garza Cocha, Ecuador.
Panacea chalcothea (Bates, 1868)
(Fig. 6)
Pandora divalis Bates, 1868. Ent. mon. Mag. 4(44):170.
This somewhat obscure taxon figures importantly in
the history of Panacea, and its taxonomic status is un-
resolved. Although we were unable to examine mate-
rial of chalcothea directly, the photo provided by G.
Lamas (Fig. 6) may serve as a starting point for identi-
fying this taxon. Here we excerpt correspondence re-
ceived from G. Lamas that bears directly on the taxo-
nomic interpretation of Panacea chalcothea:
“Bates (1868:170) described chalcothea based on at least 2 spec-
imens, one female (?) illustrated by Hewitson ([1854], Ill. exot. Butts
1: pl. [42], fig. 4), and thought by the latter to be the female of
procilla; and one male from “southern Equador”. Hewitson’s “fe-
male” belonged to the collection of the Entomological Society of
London, and that specimen is almost certainly lost, while Bates’
male would have been in his collection, and should have gone to the
BMNH through Godman and Salvin. There seems to be no Bates
specimen of chalcothea from southern Ecuador at the BMNH.
However, there is a male specimen from Bates’ collection, labeled
chalcothea by Bates himself, but from “N Peru”, and I interpret this
as a possible syntype of chalcothea, agreeing very well with the writ-
ten description of the male given by Bates in his original paper.
Bates may well have confused “S Ecuador” with “N Peru”. Anyway,
that specimen from “N Peru” most probably came from Amazonas
department in Peru. . . . Now, [it] seems to me that chalcothea
(based on Bates’ o.d. and the syntype referred to above) is . . . very
probably a subspecies of regina, or could even be a full species. For
the time being, I’m calling those 2 specimens as Panacea regina
chalcothea, though I wouldn’t be too surprised if they were to rep-
resent a high altitude species distributed from Colombia to N Peru
(if Hewitson’s “New Granada” locality for his specimen is correct,
which is quite doubtful).”
V
OLUME
56, N
UMBER
4 205
FIG. 6. Panacea chalcothea, male, dorsal and ventral, plus label.
This specimen is an apparent syntype (see Identification and Taxon-
omy). Note: whether chalcothea is a subspecies of P. regina or a valid
species remains to be resolved.
TABLE 1. Number of dissected individuals and locality data. Ab-
breviations for source collections are: P. J. DeVries (PJD); G. Austin
(GTA); G. Attal (GA); Los Angeles County Museum (LACM); Mil-
waukee Public Museum (MPM).
Taxa Source of dissected material
Ingroup
Batesia hypochlora 2 males: Brazil (GTA)
8 males: Ecuador, Sucumbios, Garza
Cocha (PJD)
1 female: Brazil (GTA)
1 female: Ecuador, Sucumbios, Garza
Cocha (PJD)
Panacea bleuzeni 1 male: French Guyana (GA)
1 female: French Guyana (GA)
Panacea divalis 5 males: Ecuador, Sucumbios, Garza
Cocha (PJD)
2 males: Brazil, Rondonia (GTA)
3 females: Ecuador, Sucumbios, Garza
Cocha (PJD)
Panacea procilla 2 males: Brazil (n = 1) and Colombia
(n = 1) (LACM)
1 male: Colombia (MPM)
1 female: Colombia (MPM)
Panacea prola 5 males: Ecuador, Sucumbios, Garza
Cocha (PJD)
3 females: Ecuador, Sucumbios, Garza
Cocha (PJD)
Panacea regina 5 males: Ecuador, Sucumbios, Garza
Cocha (PJD)
3 females: Ecuador, Sucumbios, Garza
Cocha (PJD)
Outgroups
Biblis hyperia 1 male: Ecuador, Sucumbios, Garza
Cocha (PJD)
1 female: Ecuador, Sucumbios, Garza
Cocha (PJD)
Hamadryas amphinome 1 male: Ecuador, Sucumbios, Garza
Cocha (PJD)
1 female: Ecuador, Sucumbios, Garza
Cocha (PJD)
Hamadryas arinome 1 male: Ecuador, Sucumbios, Garza
Cocha (PJD)
1 female: Ecuador, Sucumbios, Garza
Cocha (PJD)
Hamadryas feronia 1 male: Ecuador, Sucumbios, Garza
Cocha (PJD)
1 female: Ecuador, Sucumbios, Garza
Cocha (PJD)
Hamadryas laodamia 1 male: Ecuador, Sucumbios, Garza
Cocha (PJD)
1 female: Ecuador, Sucumbios, Garza
Cocha (PJD)
Distribution. Apparently Western Amazonas
(Ecuador, Peru) and Colombia (?).
Panacea divalis (Bates, 1868)
(Figs. 7, 8, 12, 13)
Pandora divalis Bates, 1868. Ent. mon. Mag. 4(44):171.
Panacea procilla divalis Seitz, 1916. Die Gross
Schmetterlinge der Erde p. 537.
Species characters. Dorsal surface with broken
iridescent blue-green bands. Forewing ventral sur-
206 J
OURNAL OF THE
L
EPIDOPTERISTS
’ S
OCIETY
FIG. 7. Panacea divalis, dorsal. Left column, males; right column, females. Top row, Rondonia, Brazil; middle and bottom rows, Garza
Cocha, Ecuador. Note variation in medial bands and submarginal ocelli.
face with reddish apex, white subapical band and dis-
tinct red spots outlined by black in discal cell (see P.
regina). Hindwing ventral surface brownish red with
a faint purple sheen; broken transversal black medial
lines, the most distal starting at Sc + Rs and ending at
1A; postmedial ocelli (black “rings”) on almost all
cells; conspicuous black submarginal line. Females
with a short, white longitudinal stripe in ventral hind-
wing cell M2–M3, nearly at the center of wing. In-
complete ocelli on dorsal surface of hindwing vary in
size, and may be absent in some specimens.
Distribution. Upper Amazon (Seitz 1916), Colom-
bia to Peru (D’Abrera 1987) and western Brazil (Em-
mel & Austin 1990).
Variation. In males the dorsal hindwing marginal
band varies among samples from Brazil and Ecuador;
the dorsal hindwing ocelli vary from diffuse to sharp; a
short, ventral longitudinal stripe may occur in ventral
hindwing cell M2–M3. In females the white, ventral
longitudinal stripe in hindwing cell M2–M3may be dif-
fuse or faintly expanded into the two cells above.
Subspecies. None.
Panacea procilla (Hewitson, 1852)
(Figs. 9, 12, 13)
Pandora procilla Hewitson, 1852. Exot. Butt. 1.
Panacea lysimache Godman and Salvin 1883. Biol.
Centr. Americana p. 275.
P. procilla ocana Fruhstorfer, 1912. Ent. Rundschau
29(6):46.
V
OLUME
56, N
UMBER
4 207
TABLE 2. Characters justifying the groupings of species and genera. MacClade 3.01 was used to map character changes on the most parsi-
monious tree. Characters indicated in bold type were unique to the group they support (independent of reversals).
Clade 1. Panacea and Batesia
(2:0) Fringe of scales in forewing and hindwing outer margin solid dark color
(16:0) Ventral surface of hindwing with black submarginal line that is discrete in anal area and more diffuse toward costal area
(24:0) Thorax: ventral portion completely covered with red-orange scales
(27:1) In lateral view: Hypandrium without anterior rod-like projections
Clade 2. Batesia hypochlora
(8:2) Males: Ventral surface of forewing apex dark, with a yellow band
(19:0) Forewing venation: M1arched toward anal margin
(25:0) Hypandrium: narrow, plate like, with obvious constriction near the middle of its long axis
(29:0) In lateral view, anterior portion of tegumen extremely projected
(30:1) Uncus tip in lateral view sharply hooked
(32:1) Uncus short
(33:0) In lateral/ dorso-lateral view, base of uncus with obvious large dorsal ridges
(34:1) In lateral view, tip of uncus not reaching or extending beyond tip of valva
(37:0) Distal portion of gnathos small and projected ventrally
(38:0) In ventral view, distal portion of gnathos with a rounded invagination
(43:1) Distal portion of valva with small bare chitinous tip
(53:0) Antrum mostly membranous
Clade 3. Panacea
(4:1) Forewing postmedial band expressed dorsally only
(5:1) In dorsal view, forewing subapical white band reduced
(7:0) Ventral surface of forewing with white subapical band
(10:0) Ventral surface of hindwing largely colored red-orange, with or without purplish sheen
(17:0) Ventral surface of hindwing with dark line imposed upon cross-vein m2–m3(at distal edge of discal cell)
(23:0) Foreleg with white scales laterally
(42:0) Distal portion of valva curving ventrally
(44:0) In lateral view, basal portion of valva with large conspicuous ventrally produced rounded projection
(46:1) In lateral view, distal portion of saccus straight to slightly projected upward
Clade 4. Panacea procilla, Panacea bleuzeni, Panacea divalis and Panacea regina
(8:0) Males: Ventral surface of forewing apex uniformly dirty red-orange
(11:0) Ventral surface of hindwing with prominent dark line across basal half of cell Sc + R1
(12:0) Ventral surface of hindwing with prominent dark line across discal cell
(13:0) Ventral surface of hindwing discal cell with two black dots in basal half
(14:0) Ventral surface of hindwing with nearly continuous line through medial area that crosses cells Sc + R1, Rs, M1, M2, M3, Cu1and Cu2
(15:1) Ventral surface of hindwing with dark line not contiguous and line in cell Cu2more apical than line in cell Cu1
(18:0) Female: ventral surface of hindwing with white patch of scales in medial area of cell M2
Clade 5. Panacea procilla, Panacea bleuzeni and Panacea divalis
(5:0) In dorsal view, forewing subapical white band well developed
(6:0) In ventral view, forewing discal cell with two red-orange spots, one at base and one at mid-length
Clade 6. Panacea procilla and Panacea bleuzeni
(3: 0) In dorsal view, male forewing with oblique, diffuse black band encroaching on postmedial blue/green band.
P. procilla salacia Fruhstorfer, 1915. Soc. Ent. 30(12):66.
P. procilla lysimache Seitz, 1916. Die Gross
Schmetterlinge der Erde p. 537.
P. procilla var. marmorensis Hall, 1917. Entomologist
50(651):171–174.
Species characters. Dorsal surface with broken
blue-green iridescent bands. Forewing ventral sur-
face with distinct red outlined by black in discal cell,
reddish apex and white subapical band. Hindwing
ventral surface brownish red with a faint purple
208 J
OURNAL OF THE
L
EPIDOPTERISTS
’ S
OCIETY
FIG. 8. Panacea divalis, ventral. Left column, males; right column, females. Top row, Rondonia, Brazil; middle and bottom rows, Garza
Cocha, Ecuador. Note variation in white stripe centered in cell M2–M3.
sheen; broken transverse medial black lines, the most
distal starting at Sc + Rs and ending at 1A; complete
postmedial ocelli on almost all cells, those on cells
M3–Cu1and Cu1–Cu2with iridescent pupil; conspic-
uous black submarginal line. Dorsal surface of hind-
wing with a medial blue band adorned with black
ocelli; conspicuous submarginal wavy line. Females
with white medial band on ventral forewing, and also
with a white band on ventral hindwing from cell Sc +
R1–Rs to M2–M3, sometimes interrupted on M1–M2.
Distribution. Costa Rica south to Colombia and
throughout the upper Amazon basin and the Guianas
(Kretzschmar 1894, Apolinar 1926).
Variation. We observed some males that have a
short, white longitudinal stripe in ventral hindwing cell
M2–M3, nearly at the center of wing—a pattern similar
to females of P. regina and P. divalis.
Subspecies. Panacea procilla procilla, western
Venezuela (Neild 1996), P. p. ocana, from lower Mag-
dalena River, Colombia (Seitz 1916, D’Abrera 1987);
P. p. salacia, from Colombia (Seitz 1916, D’Abrera
1987); P. p. lysimache from Volcan Chiriqui, Panama,
Finca la Selva, Costa Rica (DeVries 1987, 1989).
Panacea bleuzeni Plantrou and Attal, 1986
(Figs. 10, 12, 13)
Panacea bleuzeni Plantrou and Attal, 1986. Bull. So-
ciété Sciences Nat. 50:23.
Panacea bella D’Abrera, 1987. Butterflies of the
Neotropical Region, part III: p. 487, new syn-
onoym
Species characters. Dorsal surface distinctively
blue or blue-green. Dorsal surface of hindwing with a
blue medial band adorned with large black ocelli; wavy
iridescent submarginal line conspicuous. Ventral
forewing with distinct red outlined by black in discal
cell, reddish apex and white subapical band (similar to
procilla). Ventral hindwing with transverse medial
black line continuous from cell Sc + Rs to vein 1A;
ocelli faint. Females with white marking extending dis-
tally along black medial line from cell Sc + Rs to
Cu2–1A.
V
OLUME
56, N
UMBER
4 209
FIG. 9. Panacea procilla, dorsal and ventral. Left column, male; right column, female. Specimens from Cali, Colombia.
Distribution. Apparently endemic to the Guianas
(Plantrou & Attal 1986). However, it’s overlapping
range with procilla and close relationship to it (Table
2, clade 6) suggest the possibility that this taxon may
be a subspecies of procilla. This point needs critical
evaluation.
Synonymic notes. Examination of the collection
of the BMNH by A. Neild (pers. com.) revealed that
the single female holotype of P. bella is also a
paratype of P. bleuzeni. This, therefore, indicates that
P. bella and P. bleuzeni represent a single species with
bella as a junior synonym of bleuzeni. Comparing the
illustration of the type specimen of bella (in D’Abrera
1987) with photographs of male and female P.
bleuzeni provided by G. Attal confirms this assess-
ment.
DISCUSSION
Our analysis showed that Batesia and Panacea form
a monophyletic group, with B. hypochlora basal to
Panacea. Therefore, despite similarities in early stage
morphology and host plant use, we reject the hypoth-
esis that B. hypochlora is a derived species from within
Panacea. Our study confirms the maintenance of Bate-
sia and Panacea as separate taxa (e.g., Godman &
Salvin 1883, Seitz 1916), and serves as a framework for
future systematic work on both genera. We note that,
without examining material firsthand, P. chalcothea is
presumed to be the sister taxon of P. regina. However,
the phylogenetic position of chalcothea requires con-
firmation, including its taxonomic rank.
Insect genitalia are widely used for phylogenetic
210 J
OURNAL OF THE
L
EPIDOPTERISTS
’ S
OCIETY
FIG. 10. Panacea bleuzeni, female, dorsal and ventral. This figure is reproduced through the kind permission of B. d’Abrera [Butterflies of
the Neotropical Region, part III:487]. It is the type of Panacea bella D’Abrera, 1987.
reconstruction and delimiting species boundaries be-
cause their morphology may diverge rapidly, and
therefore provide informative characters (Eberhard
1985, Porter & Shapiro 1990, Arnqvist 1998). In
Panacea, however, we found that the genitalia were
highly conserved and provided no informative char-
acters for phylogeny reconstruction, or discrimina-
tion among species. Rather, the species-level rela-
V
OLUME
56, N
UMBER
4 211
FIG.11. Male genitalia: hypandrium, lateral view, ventral view (inset: tip of gnathos in ventral view). Panacea procila, P. bleuzeni, and P. divalis.
FIG. 12. Male genitalia: hypandrium, lateral view (inset: uncus in lateral view), ventral view (inset: tip of gnathos in ventral view). Panacea
regina, P. prola, and Batesia hypochlora.
tionships proposed here were derived solely from
characters of wing pattern (Fig. 14, Table 2). Our
study suggests that the most distinctly colored
species, P. pro l a, is basal to other congeners, with re-
maining species groupings justified by differences in
wing patterns.
The distinctive behavior and coloration make
Panacea easily recognizable in the field. However, in
large samples from one Ecuadorian site we found con-
siderable intraspecific variation in both genital mor-
phology and wing color patterns. This concurs with
Seitz (1916) who noted that in some Panacea species
within population phenotypic variation may be greater
than among population variation, indicating that there
may be transitions among species with respect to color
pattern. With the possible exception of P. prola, such
phenotypic variation precludes the notion that sym-
patric Panacea species can be positively identified in
nature without capturing them.
Batesia and Panacea are obvious and often abun-
dant elements of many Neotropical butterfly faunas
and museum collections. Nevertheless, some taxa are
rare in collections, and this study points to several
questions that will require a full taxonomic revision to
resolve, particularly regarding the status of P. chla-
cothea and P. bleuzeni. Although potentially useful
tools for conservation ecology, little has been reported
on the natural history Batesia and Panacea. What we
do know is that adults of both genera show significant
flight height preference in some lowland rainforests,
and that trees in the genus Caryodendron are larval
hostplants (see DeVries 1989, Montoya 1991, DeVries
et al. 1999, DeVries & Walla 2001). We do not know if
all taxa exhibit vertical stratification, if these butter-
flies use other hostplant genera, or if some species are
warningly colored (e.g., P. prola, Batesia) that repre-
sent models in mimicry complexes. We believe that
field studies, in concert with phylogenetic analyses of
Hamadryas, Ectima, Eunica, and related genera is the
next step toward understanding the evolution of Bate-
sia and Panacea, and the diversification of the Bibli-
dini.
212 J
OURNAL OF THE
L
EPIDOPTERISTS
’ S
OCIETY
FIG. 13. Female genitalia: ventral view, Panacea procila, P. divalis, P. regina, and P. prola. Lateral view: P. bleuzeni, and Batesia hypochlora
(insets: genitalia in ventral view). Note differences in the number of ovarioles between P. bleuzeni and B. hypochlora.
ACKNOWLEDGMENTS
We thank Eric Schwartz and the staff of La Selva Jungle Lodge
for supporting this and other aspects of our research. G. Attal
(France), G. Austin (Nevada State Museum), and B. Brown (Natural
History Museum of Los Angeles County) kindly facilitated our work
by loaning specimens. We are sincerely grateful to P. Ackery (The
British Natural History Museum), G. Lamas (Museo de Historia
Natural, Universidad Nacional Mayor de San Marcos, Peru), G. Mc-
Gavin (Oxford University Natural History Museum), A. Neild (Lon-
don), and C. Smith (The British Natural History Museum) for pro-
viding historical information and comparing type specimens that
helped unravel the taxonomic mysteries of P. chalcothea and P.
bleuzeni. We thank B. d’Abrera for permission to reproduce his
figures of Panacea bella. Comments by G. T. Austin and André
Freitas improved the clarity of this manuscript. RIH thanks Lee
Robertson for inspiration. This study was supported in part by the
National Science Foundation (DEB 00-96241) and is dedicated to
the memory of Sonny Clark, Bill Evans, Thelonius Sphere Monk,
and Bud Powell.
LITERATURE CITED
ACKERY, P. R. 1984. Systematic and faunistic studies on butterflies,
pp. 9–21. In Vane-Wright, R. I. & P. R. Ackery (eds.), The biol-
ogy of butterflies. Academic Press, London.
APOLINAR, H. 1927. [Notes] Boletin de la Sociadad Colombiana de
Ciencias Naturales. 16:34–35.
ARNQVIST, G. 1998. Comparative evidence for the evolution of gen-
italia by sexual selection. Nature 393:784–786.
BATES, H. W. 1868. Note on the genus Pandora (Diurnal Lepi-
doptera). Ent. Month. Mag. 4:169–171.
CARPENTER, G. D. H. 1942. The relative frequency of beak-mark
on butterflies of different edibility to birds. Proc. Zool. Soc.
London 111:223–231.
CHAI, P. 1986. Field observations and feeding experiments on the
response of rufous tailed jacamars (Galbula ruficauda) to free-
flying butterflies in a tropical rainforest. Biol. J. Linn. Soc.
29:166–189.
D’ABRERA, B. 1987. Butterflies of the Neotropical Region. Part III.
Brassolidae, Acraeidae, Nymphalidae (partim). Hill House, Vic-
toria.
DEVRIES, P. J. 1987. Butterflies of Costa Rica: and their natural his-
tory. Vol. 1: Papilionidae, Pieridae, Nymphalidae. Princeton
Univ. Press, Princeton.
———. 1989. Notes on Panacea procilla (Nymphalidae) from
Costa Rica. J. Res. Lep.27:141–142.
DEVRIES, P. J. & T. R. WALLA.2001. Species diversity and commu-
nity structure in neotropical fruit-feeding butterflies. Biol. J.
Linn. Soc. 74:1–15.
DeVries, P. J., C. M. Penz & T. R. Walla. 1999. The biology of Bate-
sia hypochlora from an Ecuadorian rainforest (Lepidoptera,
Nymphalidae). Trop. Lep. 10:43–46.
EBERHARD, W. G. 1985. Sexual selection and animal genitalia. Har-
vard Univ. Press, Cambridge.
EMMEL, T. C. & G. T. AUSTIN.1990. The tropical rain forest but-
terfly fauna of Rondonia, Brazil: species diversity and conserva-
tion. Trop. Lep. 1:1–12.
FISHER, R. A. 1958. The genetical theory of natural selection. 2nd
ed. Dover, New York.
GODMAN, F. D. & O. SALVIN.1883. Biologia Centrali-Americana.
Vol. 1. pp. 265–288.
HARVEY, D. 1991. Appendix B: Higher classification of the
Nymphalidae, pp. 255–273. In Nijhout, H. F., The develop-
ment and evolution of butterfly wing patterns. Smithsonian In-
stitution Press, Washington.
HEMMING, F. 1967. The generic names of the butterflies and their
type species (Lepidoptera: Rhopalocera). Bull. Brit. Mus. (Nat.
Hist.) Ent. supplement 9:1–509.
HEWITSON, W. C. 1854. Illustrations of new exotic butterflies. Lon-
don, John Van Voorst.
JENKINS, D. W. 1983. Neotropical Nymphalidae I. Revision of
Hamadryas. Bulletin of the Allyn Museum 81:1–146.
———. 1986. Neotropical Nymphalidae V. Revision of Epiphile.
Bulletin of the Allyn Museum 101:1–70.
———. 1987. Neotropical Nymphalidae VI. Revision of Asterope
(=Callithea Auct.). Bulletin of the Allyn Museum 114:1–66.
———. 1990. Neotropical Nymphalidae VIII. Revision of Eunica.
Bulletin of the Allyn Museum 131:1–177.
KLOTS, A. B. 1970. Lepidoptera, pp. 115–130. In Tuxen, S. L. (ed.),
Taxonomist’s glossary of genitalia in insects. Munksgaard,
Copenhagen.
KRETZSCHMAR, E. 1894. Eine neue Perisama von Columbia, das fe-
male von Panacea prola Doubl. Hew. Und ein Zwitter.
Deutsche entomol. Zeitschrift “Iris” 6:158–160.
LAMAS, G. 1994. List of butterflies from Tambopata (Explorer’s Inn
Reserve), pp. 162–177. In Foster, R. B., J. L. Carr & A. B.
Forsyth (eds.), The Tambopata reserve zone of soueastern Peru:
a biological assessment. Rapid Assessment Program, RAP work-
ing papers 6. Washingtin, D.C., Conservation International.
MADDISON, W. P. & D. R. MADDISON.1992. MacClade: version
3.01. Sinauer, Sunderland.
MONTOYA, D. C. 1991. Aspectos biologicos del gusano cachón del
inchi (Panacea sp. possible prola) Revista Colomb. Ent. (Bo-
gota) 17:41–45.
NEILD, A. F. E. 1996. The butterflies of Venezuela Part I.
Nymphalidae 1. Meridian Publications, Greenwich, London.
OTERO, L. D. & F. ROMERO.1992. Nota sobre la presences de
Panacea prola Dbldy & Hew. (Lepidoptera: Nymphalidae) en el
sector central de Cordillera de la costa Venezulana. Boletin de
Entomologia Venezulana (N.S.) 7:171–172.
PLANTROU, J. & S. ATTAL.1986. Description D’une Nouvelle Es-
pece Du Genre Panacea Decouverte En Guyane Française:
Panacea bleuzeni nov. sp. (Lepidoptera Nymphalidae). Bulletin
de la Société Sciences Nat. 50:23–24.
PORTER, A. H. & A. M. SHAPIRO.1990. The lock-and-key hypothe-
sis: lack of mechanical isolation in a butterfly hybrid zone. En-
tomol. Soc. Amer. 83:107–114.
POULTON, E. B. 1908. Essays on evolution. Clarendon Press, Oxford.
V
OLUME
56, N
UMBER
4 213
FIG. 14. Single most parsimonious tree obtained from the
analysis of 53 characters for 11 species (tree length = 79, CI = 0.82,
RI = 0.88). Numbers above and below tree branches represent boot-
strap values and the number of unambiguous changes respectively.
ROBBINS, R. K., G. LAMAS, O. H. H. MIELKE, D. J. HARVEY & M. M.
CASAGRANDE.1996. Taxonomic composition and ecological
structure of the species-rich butterfly community at Pakitza,
Parque Nacional de Manu, Peru, pp. 217–252. In Wilson, D. E.
& A. Sandoval (eds.), Manu, the biodiversity of southeastern
Peru. Washington, D.C., Smithsonian.
SCOBLE, M. J. 1992. The Lepidoptera: form, function and diversity.
British Museum (Natural History), London.
SEITZ, A. 1916. 52. Genus Panacea. 53. Genus Batesia, pp.
536–537. In Seitz, A. (ed.), Macrolepidoptera of the World. Vol.
5. Alfred Kernen, Stuttgart.
SWYNNERTON, C. M. F. 1919. Experiments and observations bear-
ing on the explanation of form and colouring, 1908–1913,
Africa. J. Linn. Soc. (Zoology) 33:203–385.
SWOFFORD, D. L. 1993. PAUP: phylogenetic analysis using parsi-
mony, version 3.1.1. Sinauer, Sunderland.
TUXEN, S. L. 1970. Taxonomist’s glossary of genitalia in insects.
Munksgaard, Copenhagen.
Received for publication 19 June 2002; revised and accepted 3 July
2002.
214 J
OURNAL OF THE
L
EPIDOPTERISTS
’ S
OCIETY
APPENDIX 1. Character list used in the phylogenetic analysis. Relevant figures are noted, and comments are included when needed. Defini-
tions are in the Characters and Terminology section.
Wing Characters:
1. Forewing outer margin: concave (0), straight (1), convex (2).
2. Fringe of scales in the outer margin of wings: solid dark color (0), dark interspersed with white sections (1).
3. In dorsal view, male forewing with oblique, diffuse black band encroaching on postmedial blue-green band (0); devoid of such a pattern (1).
Note: P. bleuzeni was scored using original description, illustration in D’Abrera and photos provided by G. Attal.
4. Forewing postmedial band expressed dorsally and ventrally (0); expressed dorsally only (1); absent or reduced (2). Note: H. laodamia and P.
procilla were polymorphic for this character because of differences between the sexes.
5. In dorsal view, forewing subapical white band well developed (0); reduced (1); absent (2).
6. In ventral view, red-orange spots on forewing discal cell: two spots present, one at base and one at mid-length (0), one spot present, at mid-
length (1), absent (2).
7. Ventral surface of forewing with white subapical band (0); devoid of such pattern (1).
8. Males, ventral surface of forewing apex: uniformly dirty red-orange (0); dark, same color as medial area (1); dark, with a yellow band (2).
9. Dorsal and ventral sides of hindwing consistently with four complete ocelli (0); dorsal side of hindwing with five incomplete ocelli (lacking
outer ring) and clearly separated from any black lines (1); ventral side of hindwing with four to six complete ocelli (2); devoid of such pat-
terns (3). Note: To understand the variation in this character a large number of specimens were examined, and we found no exceptions to
the patterns described here (see Methods, Species studied).
10. Ventral surface of hindwing largely colored red-orange, with or without purplish sheen (0); devoid of such a pattern (1). Note: although the
presence of a purplish sheen has been used to separate P. procilla and P. divalis, we found this character to be present in both these species
and variable within each of them.
11. Ventral surface of hindwing with prominent dark line across basal half of cell Sc + R1(0); devoid of such a pattern (1).
12. Ventral surface of hindwing with prominent dark line across discal cell (0); devoid of such a pattern (1).
13. Ventral surface of hindwing: discal cell with two black dots in basal half (0); devoid of such a pattern (1). Note: of the 57 P. prola specimens ex-
amined, three had two dots, 22 had one dot, and 32 lacked dots; in P. divalis, four of the 53 specimens had dots merged into a single marking.
14. Ventral surface of hindwing with: nearly continuous line through medial area that crosses cells Sc + R1, Rs, M1, M2, M3, Cu1and Cu2(0); de-
void of such a pattern (1).
15. Ventral surface of hindwing with: dark line in cell Cu2and cell Cu1contiguous (0); dark line not contiguous and line in cell Cu2more apical
than line in cell Cu1(1); dark line not contiguous and line in cell Cu2more basal than cell Cu1(2); dark line absent from cell Cu2(3).
16. Ventral surface of hindwing with black submarginal line which is discrete in anal area and becomes more diffuse toward costal area (0); de-
void of such a pattern (1). Note: P. bleuzeni was scored using the illustrations in D’Abrera (1987) and photos from the collection of G. Attal.
17. Ventral surface of hindwing with dark line imposed upon cross-vein m2–m3(at distal edge of discal cell) (0), devoid of such a dark line (1).
Note: in P. prola, three of 53 specimens lacked the dark line.
18. Female, ventral surface of hindwing with white patch of scales in medial area of cell M2(0); devoid of white patch (1). Note: two males of P.
procilla had similar white patch. In P. divalis one of 12 lacked the patch, and in P. r egina two of 14 lacked the patch.
19. Forewing venation: M1arched toward anal margin (0); devoid of such a pattern (1).
20. Forewing venation: M2arched toward anal margin (0); devoid of such a pattern (1).
21. Forewing cross-vein m2–m3+ cu1: joins M3+ Cu1at or distal to the fork M3and Cu1(0); proximally to the fork M3and Cu1(1); absent (2).
Note: M3+ Cu1denotes the combination of vein M3and Cu1proximal to the fork where they split.
22. Forewing cross-vein r–m1, and the base of M1and M2: inflated (0); not inflated (1).
Body Characters:
23. Foreleg with white scales laterally (0); devoid of white scales (1).
24. Thorax: ventral portion completely covered with red-orange scales (0); devoid of such a pattern (1).
Male Genitalia Characters:
25. Hypandrium: narrow, plate like, with obvious constriction near the middle of its long axis (0); broad, curling laterally, without a constriction (1).
26. In lateral view, hypandrium with long ramus projecting posteriorly (0); devoid of projections (1).
27. In lateral view, hypandrium with anterior rod-like projections (0); devoid of such a pattern (1).
28. In lateral view, posterior corner of hypandrium extended into an obvious lobe-like process that projects dorsally (0); less lobe-like and not as
projected dorsally (1).
29. In lateral view, anterior portion of tegumen extremely projected (0); devoid of such a pattern (1).
30. In lateral view, uncus tip: pointed (0); sharply hooked (1).
APPENDIX 1. Continued.
31. Uncus: bifid (0); entire (1).
32. Uncus: elongate (0); short (1).
33. In lateral/ dorso-lateral view, base of uncus with obvious large dorsal ridges (0); with small ridges (1); devoid of such a pattern (2).
34. In lateral view, tip of uncus reaching or extending beyond tip of valva (0); devoid of such a pattern (1).
35. Uncus with obvious, long setae dorsally (0); devoid of setae (1).
36. Distal portion of gnathos: completely fused (0); bifid (1).
37. Distal portion of gnathos: small and projected ventrally (0); large and projected posteriorly (1).
38. In ventral view, distal portion of gnathos: with a rounded invagination (0); invaginated in a perfect “V” (1).
39. Valva: with dentate process approximately 2/3 from its base (0); without such a process (1).
40. Process of valva: projecting dorsally (0); projecting medially (1).
41. Process of valva: with setae (0); without setae (1).
42. Distal portion of valva: curving ventrally (0); curving dorsally or straight (1).
43. Distal portion of valva with large bare chitinous tip (0); with small bare chitinous tip (1); devoid of such patterns (2).
44. In lateral view, basal portion of valva: with large conspicuous ventrally produced rounded projection (0); devoid of such a pattern (1).
45. In lateral view, rod-like projections of juxta: large (0); small (1).
46. In lateral view, distal portion of saccus: strongly projected upward (0); straight to slightly projected upward (1).
47. In lateral view, vinculum with obvious dentate process along anterior margin (0); process shaped as a bump, not dentate (1).
Female Genitalia Characters:
48. Signa: present (0); absent (1).
49. Sterigma: present (0); absent (1).
50. Lamella antevaginalis: continuous across ventral surface (0); split (1).
51. Lamella antevaginalis: fused to edge of eighth sternite (0); not fused (1).
52. Ductus seminalis connecting to ductus bursa: very near corpus bursa (0); far from corpus bursa, and near ostium bursa (1).
53. Antrum: heavily sclerotized (0); mostly membranous (1).
APPENDIX 2. Character Matrix.
Ingroup
Batesia hypochlora 1010221231 1111301101 0110011?01 1101110000 1111001111 111
Panacea prola 0011100130 1111300111 0100111010 101011111? ?000011110 110
Panacea procilla 000(0,1)000000 0000100011 0100111010 101011111? ?000011110 110
Panacea divalis 0011000030 0000100011 0100111110 101011111? ?000011110 110
Panacea regina 0011120020 0000100011 0100111010 101011111? ?000011110 110
Panacea bleuzeni 0002000010 0010000011 ??00111010 101011111? ?000011110 110
Outgroups
Biblis hyperia 2112221131 1111311111 2111110?10 0020101?1? ?101001100 110
Hamadryas laodamia 211(0,2)221131 1111311111 1011100?10 1010011?01 0121100011 000
Hamadryas arinome 1110221131 1111311110 1001100?10 1011011101 0121100011 000
Hamadryas amphinome 11102?1130 1111310110 1001100?10 10?1011101 0121100011 000
Hamadryas feronia 111?211?01 0110210?10 1001100?10 1011011101 0121110011 000
V
OLUME
56, N
UMBER
4 215
... Research on Biblidinae included descriptions of the chorion of some species and genera of the subtribe Ageronina sensu Lamas (2004) , which formed a monophyletic group with two clades (sister groups): Batesia-Panacea and Hamadryas-Ectima (Hill et al. 2002;Garzón-Orduña, 2012). According to generic characters, including the exochorion, the two clades may represent a tribe (Nieves-Uribe et al. 2015, 2016d, 2017aDias et al. 2019). ...
... The globose foam type of chorion is characteristic of all species of Hamadryas and Ectima, compared to other Biblidinae (see Nieves-Uribe et al. 2015, 2016d, 2017bSalik et al. 2016). This same differentiation (globose foam) is expected to occurs with Batesia and Panacea, the other two genera of Ageroniini, which are the sister group of Hamadryas + Ectima (Hill et al. 2002) We provide exochorionic descriptions for 11 of the 20 recognized species of Hamadryas and a third of its subspecies (11/33) by adding results obtained from previous studies (Nieves-Uribe et al. 2015, 2016d (Table 3). This sample comprises four of the five endemic Mexican taxa (H. ...
... In his hypothesis, the blue Hamadryas (laodamia group = subgenus Peridromia) are closer to Panacea and Batesia, and the gray Hamadryas (februa and feronia group = subgenera Ageronia and Hamadryas) are more related to Ectima (Nieves-Uribe et al. 2017a, Fig. 1 B). The results of Hill et al. (2002) are also discordant with those of Murillo-Hiller (2012), who proposed Panacea rather than Batesia as the most basal taxon. ...
Article
Full-text available
We studied the chorionic morphology of six species of Hamadryas, and together with previous studies, we compared our results with previously published phylogenies for the genus. Samples were obtained from 19 females collected between 2013 and 2017 whose abdomens were sectioned and preserved for later dissection. Eggs were extracted from those dissections and used for the descriptions and illustrations of the chorion. The Hamadryas egg is of the globose type; it is quasi-spheroidal and has multiple polygonal grids with differentiation in specific zones/regions, and knolls with macrocells in their summits that arise in the apical third. These characteristics are very different from those found in the majority of Biblidinae and for those reported in the literature for Batesia and Panacea, which belong to the same subtribe as Hamadryas (Ageroniina, now Ageroniini). Chorionic characters support a previously suggested division of the genus (februa, feronia and laodamia groups) and they agree with the phylogenetic proposal based on morphological characters. Our study expands previous morphological work focused on this genus and compiles all the information available to date about the exochorion of Hamadryas, which now includes data for 10 species and that of Ectima thecla thecla, the putative sister group of Hamadryas.
... Andrade-C 2002, Núñez Bustos 2009, Salazar et al. 2010, en ocasiones son parte de taxones en estudios comparativos (Monje-Nájera 1991, Burd 1994, Monje-Nájera et al. 1998, o bien por ser especies de interés en conservación y bioturismo (Torrealba et al. 2002, Vázquez et al. 2012. Sobre sus relaciones filogenéticas, tanto Hill et al. (2002) como Murillo-Hiller (2012) concluyen que Ageroniina constituye un grupo monofilético con dos clados hermanos: Batesia-Panacea y Hamadryas-Ectima. Respecto a trabajos sobre morfología del corion, solo hay descripciones breves del corion en Batesia, Panacea, y los mencionados previamente de Hamadryas. ...
... Se asocian a áreas de bosque tropical perennifolio y subcaducifolio, aunque también se puede encontrar en bosques riparios y áreas de crecimiento secundario (Jenkins 1985). Comparte una venación alar similar con Batesia (Hill et al. 2002); el mismo comportamiento de posar de cabeza en troncos de árboles con Panacea y Hamadryas (Keiper 1969, Baker 1972, Murillo-Hiller y Córdoba-Alfaro 2013), y Dalechampia Linnaeus 1753 (Euphorbiaceae) como planta de alimentación principal para las larvas junto con Hamadryas (Beccaloni et al. 2008, Vázquez et al. 2012. No hay filogenia propuesta para este género, tal vez debido a los escasos ejemplares que se encuentran en campo o en colecciones, por la amplitud de variación morfológica dentro de la misma especie, y por considerarse un grupo compacto que no se relaciona fácilmente con algún otro. ...
... Los resultados coriónicos en los géneros Hamadryas y Ectima, respaldan la consideración de ambos géneros como grupos hermanos, que previamente se realizó con base en caracteres de larvas, pupas e imagos (Reverdin 1914, Jenkins 1985, Monje-Nájera 1988, Hill et al. 2002, Freitas y Brown 2004, Murillo-Hiller 2012, Salik et al. 2015. Estos dos géneros presentan un corion de tipo globoso, con formación de lomos (o protolomos), presencia de retícula de fondo poligonal múltiple y macroceldas en las cúspides de los lomos. ...
Article
Full-text available
The chorion of Ectima thecla thecla was examined and compared with the chorionic morphology of Hamadryas spp. An illustrated glossary of the terminology used in chorionic descriptions in Ageroniina was created. The chorion of both genera was typified as globose and unique for the clade -with plesiomorphic structures in Ectima as compared to Hamadryas. The need for clearer and more detailed chorionic descriptions in Batesia and Panacea is noted, as previous publications recognized other types of chorion related to different Biblidinae subgroups, when perhaps a globular chorion would be expected for the entire Ageroniina subtribe.
... ( Fruhstorfer, 1915) es una mariposa tropical distribuida en la Amazonia del Ecuador y Perú ( Robbins et al., 1996;Hill et al., 2002;Checa et al., 2009), y observaciones reciente por Salazar et al., 2010 indican su presencia en Colombia. La única planta hospedera encontrada para Panacea prola amazónica en la amazonia del Perú ha sido Caryodendron orinocense Karst. ...
... Observaciones recientes indican que las larvas de Batesia y Panacea son muy similares y tienen como planta hospedera a Caryodendron spp. ( DeVries et al., 1999;Hill et al., 2002;Daniels et al., 2008). A pesar que en la amazonia peruana Panacea prola es considerada como plaga en plantaciones de Caryodendron orinocense ( Correa & Vásquez, 2007;Gonzales & Torres, 2010); esta mariposa posee potencial para los bionegocios, siendo comercializada como artesanía en países como Estados Unidos, con precios que superan los 25 dólares por cuadro. ...
Article
Full-text available
Los resultados del estudio muestran que Panacea prola amazonica copula entre el tercer y séptimo día de la emergencia. La oviposición, se realiza en hojas y ramas, está relacionada con la aparición de brotes de la planta hospedera “metohuayo” Caryodendron orinocense Karst. Su capacidad promedio de oviposición es de 262.80 ± 119.0 huevos. La duración del ciclo, desde huevo a adulto fue de 26 a 37 días, con 6 estadíos larvales.Los adultos nacen entre las 8.00 a 11.00 am, los machos sobreviven de 6 a 9 y las hembras aproximadamente 30 días en cautiverio. Se encontraron siete enemigos naturales, un parasitoide de huevo, tres parasitoides de larvas, un parasitoide de pre pupa, un depredador de larva, un depredador de adulto y un hongo entomopátogeno de pupa.
... Según [24], Hamadryas está dentro de la subfamilia Biblidinae, en la tribu Biblidini y en la subtribu Ageroniina, la que comparte con los géneros Batesia, Panacea y Ectima. La estrecha relación entre estos tres géneros esta también respaldada por los análisis de [4,25,13]. ...
... Los estudios de campo y análisis filogenéticos de Hamadryas, Batesia, Panacea, Ectima y otros géneros relacionados son el próximo paso para comprender la evolución y la diversificación de la tribu Biblidini [10,4]. Diferentes especies de mariposas se posan en diferentes lugares, posiblemente dependiendo de su habilidad de camuflarse con el tronco [12,10,6]. ...
Article
Full-text available
A revisión of the Hamadryas Hübner (1806) genus was conducted by mainly considering its taxonomic history as a function of the morphological, ethological, ecological, and phylogenetic aspects.
... Adults are born from 8.00 to 11.00 am, the males survive between 6 a 9 days and females about 30 days in captivity. We found seven natural enemies, an egg parasitoid, three larvae parasitoids, a pre pupal parasitoid, a larvae predator, an adult predator and a pupal entomopathogenic fungus., 1915) es una mariposa tropical distribuida en la Amazonia del Ecuador y Perú (Robbins et al., 1996; Hill et al., 2002; Checa et al., 2009), y observaciones reciente por Salazar et al., 2010 indican su presencia en Colombia. La única planta hospedera encontrada para Panacea prola amazónica en la amazonia del Perú ha sido Caryodendron orinocense Karst. ...
... Observaciones recientes indican que las larvas de Batesia y Panacea son muy similares y tienen como planta hospedera a Caryodendron spp. (DeVries et al., 1999; Hill et al., 2002; Daniels et al., 2008). A pesar que en la amazonia peruana Panacea prola es considerada como plaga en plantaciones de Caryodendron orinocense (Correa & Vásquez, 2007; Gonzales & Torres, 2010); esta mariposa posee potencial para los bionegocios, siendo comercializada como artesanía en países como Estados Unidos, con precios que superan los 25 dólares por cuadro. ...
Data
Full-text available
Los resultados del estudio muestran que Panacea prola amazonica copula entre el tercer y séptimo día de la emergencia. La oviposición, se realiza en hojas y ramas, está relacionada con la aparición de brotes de la planta hospedera “metohuayo” Caryodendron orinocense Karst. Su capacidad promedio de oviposición es de 262.80 ± 119.0 huevos. La duración del ciclo, desde huevo a adulto fue de 26 a 37 días, con 6 estadíos larvales. Los adultos nacen entre las 8.00 a 11.00 am, los machos sobreviven de 6 a 9 y las hembras aproximadamente 30 días en cautiverio. Se encontraron siete enemigos naturales, un parasitoide de huevo, tres parasitoides de larvas, un parasitoide de pre pupa, un depredador de larva, un depredador de adulto y un hongo entomopátogeno de pupa.
... First, species of Brassolis are remarkably uniform in morphology, making it difficult for us to find informative characters. Genitalic morphology may often yield informative characters for phylogeny reconstruction (e.g., , Penz 2008; however, interspecific variation in genitalia within Brassolis was negligible, as found in other butterfly genera (e.g., Caldas 1997, Hill et al. 2002. Second, the few coloration characters that varied among species also varied within species. ...
Article
Full-text available
This study examines the phylogenetic relationships among species of the butterfly genera Dynastor and Brassolis using 57 characters from adult morphology and wing coloration. We provide evidence for the monophyly of both genera. The relationships among Dynastor species are well resolved, but we were unable to find informative characters that could resolve the relationships inside Brassolis. We provide diagnoses for Dynastor and Brassolis and all species included in these genera, including illustrations that show geographical variation in wing color. The status of one subspecies is changed to species; Brassolis dinizi d'Almeida, 1956, NEW STATUS.
... First, species of Brassolis are remarkably uniform in morphology, making it difficult for us to find informative characters. Genitalic morphology may often yield informative characters for phylogeny reconstruction (e.g.,, Penz 2008); however, interspecific variation in genitalia within Brassolis was negligible, as found in other butterfly genera (e.g.,Caldas 1997, Hill et al. 2002). Second, the few coloration characters that varied among species also varied within species. ...
Article
Full-text available
This study examines the phylogenetic relationships among species of the butterfly genera Dynastor and Brassolis using 57 characters from adult morphology and wing coloration. We provide evidence for the monophyly of both genera. The relationships among Dynastor species are well resolved, but we were unable to find informative characters that could resolve the relationships inside Brassolis. We provide diagnoses for Dynastor and Brassolis and all species included in these genera, including illustrations that show geographical variation in wing color. The status of one subspecies is changed to species; Brassolis dinizi d’Almeida, 1956, NEW STATUS.
Article
Full-text available
Based on comparative morphology of adults, a phylogeny is proposed for the butterfly tribe Amathusiini (Nymphalidae, Satyrinae). The dataset includes 92 characters scored for 45 species in 12 genera, representing the most comprehensive phylogenetic analysis for this group. Parsimony analyses produced a well-resolved strict consensus tree where genera were divided in three main groups: (clade 1) Stichophthalma; (clade 2) Aemona, Faunis, Melanocyma and Taenaris; (clade 3) Enispe, Discophora, Thaumantis, Thauria, Amathusia, Amathuxidia, and Zeuxidia. While genera in clades 1 and 2 were found to be morphologically homogeneous, clade 3 showed remarkable morphological divergence between and within genera. The monophyly of most genera was recovered with variable levels of support, but Melanocyma and Taenaris nested within Faunis. Therefore, here Melanocyma NEW SYN. is subsumed within Faunis, and Taenaris STAT. REV. is regarded as a subgenus of Faunis. Mimicry likely evolved a single time within the Faunis-Taenaris assemblage, as species of Taenaris formed a monophyletic group. Results are compared to early classifications and recent DNA-based analyses, and points of agreement and conflicts are discussed.
Article
The neotropical butterfly genus Hamadryas Hübner comprises 20 species that exhibit an intriguing variation in their natural history traits. Although revised in 1983, no phylogenetic hypothesis was presented: the first phylogenetic hypothesis is estimated here based on 93 characters and including species from the three other genera in the tribe Ageroniini. The phylogeny is used to test the monophyly of the genus, establish the sister group of Hamadryas and identify its apomorphies. The tree allows the inference of patterns of character change in sound production and sexual dimorphism. Implied weights show that Hamadryas is monophyletic and corroborate Ectima Doubleday as a sister genus. Previously suggested subgenera for Hamadryas were non‐monophyletic, with the exception of the laodamia clade, supported by the presence of a complete sterigma. Sound production is inferred to be a derived condition in Hamadryas that has been lost in the laodamia clade. This, plus the presence of androconial organs and sexual dimorphism in the laodamia clade, suggests a shift in sexual recognition signalling. Furthermore, the phylogeny indicates that the colour pattern of males in the laodamia clade is novel, supporting a Darwinian origin of sexual dimorphism.
Article
Full-text available
The present work intends to contribute to the dried material of butterflies deposited in the biological collections of the Centro de Museos (N.H.), Universidad de Caldas and Julián Salazar’s collection (CJS). It contains some of the groups that belong to the superfamily Nymphaloidea, with related information on its history and with taxonomic notes on their composition and distribution.
Article
Full-text available
The “lock-and-key“ hypothesis of mechanical isolation predicts little if any natural hybridization between genitalically differentiated taxa. Chilean Tatochila mercedis (Eschscholtz) differs from Argentine T. sterodice Staudinger and T. vanvolxemii (Capronnier) in the degree and direction of a twist of the aedeagus. We report data on the genetic bases for these traits and use these data for an analysis of the degree of hybridization and introgression of genitalic traits in contact zones in the field. The direction of twist is based on one or a few genes, whereas the degree of twist is probably polygenic; the two characters are statistically independent in the laboratory. The ease of laboratory pairing and the high incidence of hybridization in nature clearly indicate that mechanical isolation based on these traits is untenable. There is a lack of strict concordance of clines involving these characters in the field; introgression of twist direction occurs farther eastward than twist degree, but this is a likely consequence of the different genetic bases for these traits. We reject the lock-and-key model for this group and suggest that arthropod systematists be cautious in recognizing species based on simple genitalic characters in parapatry.
Article
Full-text available
Notes on the adult population biology and early stages of Batesia hypochlora are presented for the first time. The general natural history, population abundance, warning coloration, behavior, geographical distribution and hostplant use of Batesia hypochlora is discussed. Notes on the taxonomic relationship of Batesia and Panacea are also provided. and Panacea are also provided
Article
Full-text available
To test the veracity of previous studies and illuminate major community patterns from an intact community, a guild of nymphalid butterflies was sampled at monthly intervals for five consecutive years by trapping in the canopy and understorey of five contiguous forest plots in the same rainforest. Significant numbers of species belonged to either the canopy or understorey fauna, confirming fundamental vertical stratification, and showing that sampling in one vertical position is a poor estimator of diversity. Significant monthly variation showed that intermittent or short-term sampling would underestimate diversity, and significant variation among years and areas showed that diversity was strongly influenced by sampling year. Even when the underlying communities were the same, temporal interactions strongly affected species diversity in both horizontal and vertical dimensions. An unprecedented seasonal inversion of species richness and abundance was detected between the canopy and understorey that occurred at the onset of all rainy seasons. This investigation suggests that long-term studies evaluating spatial and temporal patterns of species diversity among many sites may be required for a better understanding of tropical communities and how best to conserve them.
Neotropical Nymphalidae VI. Revision of Asterope (=Callithea Auct
———. 1987. Neotropical Nymphalidae VI. Revision of Asterope (=Callithea Auct.). Bulletin of the Allyn Museum 114:1–66.
Aspectos biologicos del gusano cachón del inchi (Panacea sp. possible prola) Revista Colomb
MONTOYA, D. C. 1991. Aspectos biologicos del gusano cachón del inchi (Panacea sp. possible prola) Revista Colomb. Ent. (Bogota) 17:41-45.
52. Genus Panacea. 53. Genus Batesia
  • A Seitz
SEITZ, A. 1916. 52. Genus Panacea. 53. Genus Batesia, pp. 536-537. In Seitz, A. (ed.), Macrolepidoptera of the World. Vol.
Nymphalidae (partim) Hill House
  • Acraeidae Brassolidae
Brassolidae, Acraeidae, Nymphalidae (partim). Hill House, Victoria.
  • F D O Salvin
GODMAN, F. D. & O. SALVIN. 1883. Biologia Centrali-Americana.