Biological Journal ofthe Linnean Society (1988). 33: 95-203 .

With 1 figure

Hostplants and classification: a review of

nymphalid butterflies

P . R . ACKERY

Department of Entomology. British Museum (Natural History). Cromwell Road. London

SW7 5BD

Received 8 April 1987. accepted f o r publication 26 August 1987

I n reviewing the hostplant associations of nymphalid butterflies. particular emphasis is placed on

the intractable problem of nymphalid classification . Although offering few certain conclusions. if
used in conjunction with more formal morphological characters. the data presented should
contribute toward a resolution of the inter-relationships of the many widely recognized groupings
within the Nymphalidae. several of which seem to be broadly characterized by typical host families .
As a direct result of this analysis. the presumed association between larval hostplants and
unpalatability is re.appraised .

KEY WORDS:-Nymphalidae - hostplants - classification - mimicry - unpalatability .

CONTENTS

Introduction . . . . . . . . . . . . . . . . . . . 96
Classification of the Nymphalidae . . . . . . . . . . . . . . 96
Hostplants as ‘characters’ . . . . . . . . . . . . . . . . 98
Nymphalid hostplants . . . . . . . . . . . . . . . . . 98
Brassolinae . . . . . . . . . . . . . . . . . . 99
Amathusiinae . . . . . . . . . . . . . . . . . . 101
Morphinae . . . . . . . . . . . . . . . . . . 103
Satyrinae . . . . . . . . . . . . . . . . . . . 103
Haeterini . . . . . . . . . . . . . . . . . . 103
Biini . . . . . . . . . . . . . . . . . . . 104
Elymniini . . . . . . . . . . . . . . . . . 104
Eritini & Ragadiini . . . . . . . . . . . . . . . 107
Satyrini . . . . . . . . . . . . . . . . . . 107
Calinaginae . . . . . . . . . . . . . . . . . . 123
Charaxinae . . . . . . . . . . . . . . . . . . 123
Pallini & Euxanthini . . . . . . . . . . . . . . . 124
Charaxini . . . . . . . . . . . . . . . . . 124
Anaeini . . . . . . . . . . . . . . . . . . 128
Preponini . . . . . . . . . . . . . . . . . 131
Prothoini . . . . . . . . . . . . . . . . . . 132
Acraeinae . . . . . . . . . . . . . . . . . . . 135
Heliconiinae . . . . . . . . . . . . . . . . . 137
Argynninae . . . . . . . . . . . . . . . . . 140
. Melitaeinae . . . . . . . . . . . . . . . . . . 144
Euphydryini . . . . . . . . . . . . . . . . . 145
Melitaeini . . . . . . . . . . . . . . . . . 146
Phyciodini . . . . . . . . . . . . . . . . . 146
Limenitinae . . . . . . . . . . . . . . . . . . 149
95
0024-4082/88/020095 + 109 $03.00/0 0 1988 T h e Linncan Society of London
96 P. R . ACKERY
Neptini . . . . . . . . . . . . . . . . . . 152
Bihlini . . . . . . . . . . . . . . . . . . 152
Marpesiini . . . . . . . . . . . . . . . . . I54
Ageroniini . . . . . . . . . . . . . . . . . 154
Pseudergolini. . . . . . . . . . . . . . . . . 156
Parthenini . . . . . . . . . . . . . . . . . 157
Euthaliini . . . . . . . . . . . . . . . . . 157
Epicaliini. . . . . . . . . . . . . . . . . . 158
Limenitini . . . . . . . . . . . . . . . . . 160
Nymphalinae . . . . . . . . . . . . . . . . . . 163
Nymphalini . . . . . . . . . . . . . . . . . 165
Colohurini . . . . . . . . . . . . . . . . . 173
Apaturinae . . . . . . . . . . . . . . . . . . 174
Libytheinae . . . . . . . . . . . . . . . . . . 175
Danainae. . . . . . . . . . . . . . . . . . . 177
Ithomiinae . . . . . . . . . . . . . . . . . . 183
Tellervinae . . . . . . . . . . . . . . . . . . 186
Summary of hostplant patterns. . . . . . . . . . . . . . . 186
Hostplants and unpalatability . . . . . . . . . . . . . . . 188
Acknowledgements . . . . . . . . . . . . . . . . . 190
Krfrrences. . . . . . . . . . . . . . . . . . . . 191

INTRODUCTION

The higher classification of the Nymphalidae is one of the major remaining

challenges for butterfly systematists. Such well-known nymphalid groups as the
I thomiinae, Danainae, Heliconiinae and Melitaeinae must still he viewed in
isolation-their inter-relationships remain largely obscure. This problem is only
likely to be resolved by using unorthodox data, such as hostplant relationships,
in addition to classical morphological characters. While highlighting larval
foodplant associations, this paper does not offer a n evolutionary narrative;
instead it aims to present data that might ultimately be incorporated into a
more wide-ranging analysis. T h e Nymphalidae are a n ideal group for such an
approach. Being often large and conspicuous, they have attracted wide
attention, resulting in copious hostplant data. And, of particular importance for
a study of this kind, the Nymphalidae in their widest sense clearly form a
monophyletic group (Kristensen, 1976).
Since Fraenkel’s (1959) insight into the “raison d’etre” of secondary plant
compounds, and Ehrlich & Raven’s ( 1965) coevolutionary study, butterflies
have moved to the forefront of investigations into plant-herbivore interactions.
This has in turn stimulated detailed studies of hostplant associations in several
butterfly groups, particularly the nymphalids (White & Singer, 1974; Benson,
Brown & Gilbert, 1976; Drummond, 1976; Haber, 1978; Aiello, 1984), resulting
in a wealth of new information to supplement the more traditional sources.
Despite this, it should not be forgotten that the literature abounds with
misidentifications, both of the butterflies and their hostplants, and many of these
errors have been compounded by repetition. T h e frequent absence of voucher
specimens of the plants (or even the butterflies) frustrates efforts to eliminate
such errors.

CLASSIFICATION OF T H E NYMPHALIDAE

Kristensen ( 1976) notes four autapomorphies for the family Nymphalidae-

male forelegs always reduced and clawless, pupa usually suspended only by the
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 97

hooks of the cremaster, most antenna1 segments with two ventral grooves, and
the usually very distinct separation of the laterocervico-tentorial muscles into
two bundles. A long-established grouping, it is conventionally placed as the
‘sister’ of the Lycaenidae (Ehrlich, 1958; Kristensen, 1976; Scott, 1985). Figure
1 presents a consensus of the current state of the higher classification within the
family. Although both Ehrlich (1958) and Scott (1985) exclude the
Libytheidae, this appears to render the residual Nymphalidae paraphyletic
(Kristensen, 1976). Greater heresies (DeVries, Kitching & Vane-Wright, 1985)
suggest that the two largest conventionally recognized subfamilies, the Satyrinae
and Nymphalinae (sensu lato), are polyphyletic, but such views need to be tested
across a broader range of species. DeVries el al., did, however, convincingly
demonstrate that the Morphinae sensu Ehrlich (1958) are paraphyletic.

Ithomiinae
98 P. R . ACKERY

Nevertheless, for the purposes of the present paper, which aims to present
hostplant data within the conventions of a widely recognized system, a
conservative approach is adopted.
At the subfamily level, few inter-relationships have been established. DeVries
el al. (1985) group the Amathusiinae, Brassolinae and Morphinae, with the
addition of Antirrhea and Caerois (Satyrinae), on the basis of a larval character
“vertex horns with narrow base, transition to head abrupt”. Ehrlich (1958)
suggests close affinity between the entire Satyrinae and his Morphinae (which
includes the Brassolinae and Amathusiinae); in both groups the valve is
somewhat simplified and dentate, while the larvae have bifid tails and usually
exploit monocotyledons.
T h e free base of forewing vein 3A might associate the Danainae, Ithomiinae
and Tellervinae. Although relatively unusual within the Nymphalidae
(otherwise only occurring in haeterine Satyrinae and Dryas, Kallima, Apaturina
and Taenaris), it is probably a plesiomorphic feature. However, the grouping
still appears justifiable on biological grounds (Ackery & Vane-Wright, 1984).
The Apaturinae, Charaxinae, Calinaginae and Libytheinae, show no obviously
close relationship to each other or any other nymphalid grouping, but the
remaining subfamilies, the Acraeinae, Heliconiinae, Argynninae, Melitaeinae,
Limenitinae and Nymphalinae are usually associated together, generally on the
basis of non-unique features (see Ehrlich, 1958, who includes many of these
somewhat contentious groupings within a single subfamily, the Nymphalinae,
sensu lato).

HOSTPLANTS AS ‘CHARACTERS

Downey’s (1962) observation that hostplants have not been widely used to
establish classifications still largely holds true. Seitz’ ( 1927) pioneering work is
the only outstanding example; elsewhere Igarashi (1984), following the tradition
established by Munroe (1961), utilized hostplant data in establishing his
proposed classification of the Papilionidae, as have both Pierre (1984) and
Ackery & Vane-Wright ( 1984) for the Acraeinae and Danainae, respectively.
But we should be cautious here there is no reason to suppose that hostplant
associations are any less likely to reflect problems caused by inherited ancestral
traits or homoplasy than are traditional characters. Witness Seitz’ (1927)
unacceptable suggestion that the few apocynaceous ithomiines should be
included within the ApocynaceaeeAsclepiadaceae-Moraceae-feeding danaines
rather than the predominantly Solanaceae-feeding Ithomiinae. And as Miller
i1986) concluded, at least for swallowtail butterflies, the range of acceptable
hosts often owes more to chemical similarity than phylogenetic relationship.
Given phytochemical diversity, homoplasy is to be expected.

NYMPHALID HOSTPLANTS

T h e wealth of available data is here re-appraised. Synopses by subfamily

provide information on the principal hostplants utilized, together with brief
discussions of their interrelationships and known chemical similarities. This in
turn is supplemented by outlines of various other topics where they appear
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 99

relevant, particularly systematics, unpalatability and mimicry, broad habitat

preference and oviposition behaviour.
Much of the host data is not verifiable. So, rather than introduce yet more
errors and inconsistencies, I have opted throughout to retain the original
botanical nomenclature. No doubt, this means that many generic names used
here should be subjectively treated as junior synonyms. Conversely, umbrella
genera, such as Aster and Chrysanthemum are now used in a far more restricted
sense than in the past. Such weaknesses should be minimized by considering
broad trends rather than emphasizing seemingly unique but largely
unsubstantiated records.
Tables, which cross-refer butterfly species to foodplant genera, form the basis
of the review. By using numbers in the appropriate column each record can be
traced back, at least to the major works listed in the table legend. Plant genera
are listed alphabetically and generally assigned to family in accordance with
Willis (1980), within a family sequence based largely on Heywood (1979). T h e
order of butterfly genera and species follows a systematic sequence whenever
possible. However, the frequent absence of modern treatments often necessitates
alphabetical listings.
Finally, a purely nomenclatorial note. I n formulating the family-group
names, the practice most widely adopted by Lepidopterists has been followed
throughout; that is, the appropriate endings, -ina, -ini and -inae, are added to
the basic generic stem. I n some cases this results in departures from the more
widely accepted form. Thus, I have utilized Amaurina, Godyrini and
Limenitinae not Amauridina, Godyridini and Limenitidinae.

Brassolinae
While Ehrlich (1958) subsumes the Brassolinae and Amathusiinae within the
Morphinae, Miller (1968) places the brassolines as ‘Satyrids’, with the
Amathusiinae retained within the ‘Morphids’. DeVries et al. (1985) have
confirmed the monophyly of the Brassolinae + Amathusiinae + Morphinae,
but only with the addition of two ‘satyrine’ genera, Antirrhea and Caerois. At the
species level, Stichel’s works (1909, 1925, 1932) provide the taxonomic basis of
the Brassolinae, supplemented by Bristow’s generic revisions ( 1981, 1982 and in
prep.).
T o date, records suggest that members of this exclusively Neotropical group of
80 crepuscular forest species (Fruhstorfer, 1912; Ross, 1976) feed exclusively on
monocotyledons. Overall, there is little correlation between hostplants and
brassoline classification (Stichel, 1925; Miller, 1968). Only exploitation of
bromeliads by Dynastor appears unique (Table 1), while Brassolis and Opsiphanes
utilize members of the Arecaceae. Musa (Musaceae), a widely exploited Old
World introduction, has close affinities with the Heliconiaceae, Zingiberiaceae,
Marantaceae and Cannanaceae, families present in the Neotropics and utilized
as brassoline larval hosts.
Bristow ( 1981) notes wing pattern correlations in various Catoblepia species
and tentatively invokes mimicry. Throughout the host range there are positive
and negative reports of cyanogenesis together with the irregular occurrence of
100 P. K. ACKERY

TABLE 1. Hostplants of the Brassolinae. Aiello & Silberglied, 1978l; Barcant, 19702;Cassagrande,
1979'; Cuberto, 19854; d'Almeida, 192Z5; d'Araujo e Silva el al., 19686; DeVries, 19867;
Fountaine, 19138, unpublishedg; Fruhstorfer, 19121°; Harrison, 1963"; Hayward, 196912; Miles
Moss, unpublished"; Rothschild, 191614; Urich & Boos, 1980I5; Young, 197716, 1986"; Young
& Muyshondt, 1975'*, 198519

Brassol i s . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . .
astyra . . . . . . . . . . . . . 6 . . . . . . . . 6 . . . . . .
.
isthmia . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . .
sophorae . . . . . . . . . . . . . 6 . . . . . . . . 6 . . . . . .
.
Catoblepia . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . .
amphirhoe . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . .
Dynastor . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . .
darius . . . . . . . . . . . . . . . 7 1 6 6 6 . .10 . . . . . .
.
macrosi r i s . . . . .
. . . . . . . . . . . 15 . . . . . . . . . . . . .
napoleon . . . . . . . . . . . . . . . 6 . . . . . . . . . . . . .
.
opoptera . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . .
aorsa . . . . . 6 . . . . . . . . . . . . . . . . . . . . . . . .
staudingeri . . . . . . 4 . . . . . . . . . . . . . . . . . . . . . . .
syne . . . . . . . 6 . . . . . . . . . . . . . . . . . . . . . .
Opsiphanes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
batea . . . . . . . . . . . 6 . . . . . . . . . . . . . . . . . .
bogotanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
cassiae . . . . . . . . . . . . . . . . . . . . . . . 6 . 1 3 . . . .
cassina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
invirae . . . . . . . . . . . . . . . . . . . . . . . 6 . . . . . .
quiteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
tamarindi . . . . . . . . . . . . . . . . . . . . . . . 18 . 18 . . . .
Caligo , . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
arisbe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
atreus . . . . . . . . . . . . . . . . . . . . . . . . .19 . . . .
beltrao . . . . . . . . . . . . . . . . . . . . . . . 6 . . . 6 . 3
eurilochus . . . . . . . . . . . . . . . . . . . . . . 5 9 . 7 . 6 . .
idomeneus . . . . . . . . . . . . . . . . . . . . . . .13 . . . . . .
illioneus . . . . . . . . . . . . . . 6 . . . . . . . . 14 . . . . . .
martia . . . . . . . . . . . 6 6 . . . . . . . . . . . . . . . .
m o n . 6 . . . . . . . . . . . . . . . . . . . . . 11 . ? . . .
oileus . . . . . . . . . . . . . . . . . . . . . . .13 . . . . . .
.
p l ac id i anus . . . . . . . . . . . . . . . . . . . . . . 13 . . . . . .
prometheus. . . . . . . . . . . . . . . . . . . . . . .14 . . . . . .
teucer . . . . . . . . . . . . . . . . . . . . . . . 2 . 1 3 . . . .
Eryphanis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
aesacus . . . . . . . . . 4 . . . . . . . . . . . . . . . . . . . .
automedon . . . . . 2.. . . . . . . . . . . . . . . . . . . . . . .
reevesii . . . . . 6 . 6 . l o . . 6 . . . . . . . . . . . . . . . . .
Dasyophthalma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
rusina . . . . . 6 . . . . . . . . . . . . . . . . . . . . . . . .
Narope . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
cyllastros . . . . . 6 . 6 . . . . . . . . . . . . . . . . . . . . . .

saponins and tanins (Darnley Gibbs, 1974). When investigating the feeding
efficiencies of various herbivores (including Caligo memnon and Opsiphanes
tamarindi),Auerbach & Strong (1981) found no evidence of chemical defences in
Heliconia species.
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 101

TABLE
1. Continued

. . . . . . . . 6 . 6 . . . 6 . . . 6 . 6 6 . . . . . astyra
.
. . . . .. . . . . . . . 17 . 17 . . . . . . . . . . . . isthmia
. . . . .
. . . . 6 6 6 . . 6 6 6 6 6 6 6 6 6 . . . . . sophorae
. . . . .
. . . . . . . . . . . . . . . . . . . . . . . Catoblepia
. . . . .
. . . . . . . . . . 6 . . . . . . . . . . . . amphirhoe
. . . . .
. . . . . . . . . . . . . . . . . . . . . . . Dynastor
. . . . .
. . . . . . . . . . . . . . . . . . . . . . . darius
. . . . .
. . . . . . . . . . . . . . . . . . . . . . . macrosiris
. . . . .
. . . . . . . . . . . . . . . . . . . . . . . napoleon
. . . . .
. . . . . . . . . . . . . . . . . . . . . . . Opoptera
. . . . .
. . . . . . . . . . . . . . . . . . . . . . . aorsa
. . . . .
. . . . . . . . . . . . . . . . . . . . . . . staudingeri
. . . . .
. . . . . . . . . . . . . . . . . . . . . . . syne
. . . . .
. . . . . . . . . . . . . . . . . . . . . . .Opsiphanes
. . . . 6 . . . . . . . . . . . . . . . . . . . . . . . batea
. . . . 7 . . . . . . . . . . . . . . . . . . . . . . . bogotanus
. . . . 8 . . . . . . . . . . . . . . . . . . . . . . . cassiae
. . . . . 7 . . . . . 1 8 1 8 . . 7 . . . . . . . . . 18 . . cassina
. . . . . . . . . . . . . . . 6 6 . . 6 . 6 1 2 4 6 . . 6 invirae
. . . . . . 6 1 2 . 6 . . . . . 16 . . 6 . . . . 4 . . . . q u i t e r i a
. . . . . . . . . . . . . . . . . . . . . . . . . . . . tamrirdi
. . . . . . . . . . . . . . . . . . . . . . . . . . . .Caligo
. 6 . . . . . . . . . . . . . . . . . . . . . . . . . . arisbe
. . . . . . . . . . . . . . . . . . . . . . . . . . . . atreus
. . 3 . . . . . . . . . . . . . . . . . . . . . . . . . beltrao
. . . . . . . . . . . . . . . . . . 6 . . . . . . . . . eurilochus
- 1 2 . . . . . . . . . . . . . . . . . . . . . . . . . . idomeneus
. . . . . . . . . . . . . . . . . . . . . . . . . . . . illio n eu s
. . . . . . . . . . . . . . . . . . . . . . . . . . . . martia
. . . . . . . . . . . . . . . . . . . . . . . . . . . . m o n
. . . . . . . . . . . . . . . . . . . . . . . . . . . . oileus
. . . . . . . . . . . . . . . . . . . . . . . . . . . . placidianus
. . . . . . . . . . . . . . . . . . . . . . . . . . . . pranetheus
. . . . . . . . . . . . . . . . . . . . . . . . . . . . teucer
. . . . . . . . . . . . . . . . . . . . . . . . . . . . Eryphanis
. . . . . . . . . . . . . . . . . . . . . . . . . . . . aesacus
. . . . . . . . . . . . . . . . . . . . . . . . . . . . autunedon
. . . . . . . . . . . . . . . . . . . . . . . . . . . . reevesii
. . . . . . . . . . . . . . . . . . . . . . . . . . . . Dasyophthalma
. . . . . . . . . . . 6 . . . . . . 6 . . . . . . . . . rusina
. . . . . . . . . . . . . . . . . . . . . . . . . . . . Narope
. . . . . . . . . . . . . . . . . . . . . . . . . . . . cyllastros

Amathusiinae
The corresponding Indo-Australian group, the Amathusiinae, also associates
almost exclusively with monocotyledons. Although not comprehensive for
102 P. R. ACKERY

TABLE 2. Hostplants of the Amathusiinae. Bascombe et al., 1987'; Corbet & Pendlebury, 197fI2;
D'Abrera, 19773; Hill et al., 19784; Kirchberg, 19425; Parsons, 19846; Sevastopulo, 1973'

amathusia . . . . . . . . . . . . . . . . . . . . . . . . . . 5 . . . .
Amathusia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
gunneryi . . . . . . . . . . . . . . . 2 . . . . . . . . . . . . . . .
phidippus . . . . . . . . . . . . . 7 . 7 7 . . . . . . . . . . . . . .
Discophora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
lepida . . . 7 7 . . . . . . . . . . . . . . . . . . . . . . . . . .
sondaica . . 1 1 . . 1 . . . . . . . . . . . . . . . . . . . . . . . .
timora . . . 2 . . . . . . . . . . . . . . . . . . . . . . . . . . .
tullia . . 5 5 . . . . . . . . . . . . . . . . . . . . . . . . . . .
Faunis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
aerope . . . . . . . . . . . . . . . . . . . 5 . . . . . . . . . . .
canens . . . . . . . . 2 . . . . . . . . . . . . . . . . . . . . . .
emus . . . . . . . . l . . . . . . . . 4 . 5 . . 1 . . . 4 . . . .
S t ichopht ha lma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
howqua . . . . . 5 . . . . . . . . . . . . . . . . . . . . . . . . .
Taenar is . . . . . . . . . . . . . . . . . . . . . . . . . . . .
catops . . . . . . . 6 . . . 6 . 6 . . . . . . 6 . . . . . . 6
phorcas . . . . . . . . . . . . . . . . . . . . . 6 . . 3 . . . . . .
gorw . . . . . . . . . . . . . . 6 . . . . . . . . . . . . . . . .
onolaus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
WOPS . . . . . . . . . . 6 . . . . . . . . . . . . . . . . . . . .
horsfieldii . . . . . . . . . . . . . . . . . . . . . . . . . . 2 . . . .
butleri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
dimna . . . . . . . . 6 . . . . . . . . . . . . . . . . . . . . . .
artemis . . . . . . . . . . . . . . . 6 . . . 6 . . . . . . . . . . .

species, Kirchberg ( 1942) provides the taxonomic basis for the group-he
recognized 13 genera. As host records only cover one-fifth of the currently
recognized species (Stichel, 1933), no meaningful patterns emerge (Table 2).
Bamboos seem to be particularly favoured by Discophora species, with
Amathusia and to a lesser extent Taenaris focussing on the Arecaceae; otherwise
'Taenaris species most notably feed on Cycads.
Amathusiines, essentially cryptic, crepuscular butterflies, usually inhabit deep
forest (Fruhstorfer, 1911). Exceptionally, Taenaris species, although still
preferring shade (Brooks, 1950), are common, day-flying butterflies, often
possessing conspicuous eyespots. Vane-Wright ( 1974) suggested Taenaris species
may act as primary models in a mimetic complex involving Mycalesis drusillodes
( 0 ), Hypolimnas species, Elyrnnias species, certain 0 forms of Papilio aegeus and two
i s Hyantis. At present, this particular
other amathusines, ~ o r p ~ o ~ a e n a rand
assertion, supported by Parsons (1984), has no certain chemical basis, although
some other Cycadaceae-feeding Lepidoptera are clearly aposematic (DeVries,
1977), sequestering and storing cycasin (Rothschild, Nash & Bell, 1986). Cycads
are toxic and often lethal to cattle (Parsons, 1984). Among amathusiine
hostplants, cyanogenesis has been variously reported along with saponins and
tannins (Darnley Gibbs, 1974).
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 103

Morphinae
I n the sense used here, this group is confined to the Neotropics, typically in
low- to middle-altitude rain forests. A recent cladistic analysis based on the early
stages (DeVries et al., 1985) showed two classical ‘satyrid’ genera, Antirrhea and
Caerois, to cluster consistently with Morpho, confirming Vane-Wright’s (1972)
suggestion that these three genera constitute the true Morphinae.
Le Moult & Real (1962) recognized some 70 Morpho species divided between
eight subgenera. There are obvious correlations between these subgenera and
the larval hosts (Table 3). M . (Iphimedia) hercules and M . ( I . ) richardus feed on
Menispermaceae, a family not otherwise exploited by Morpho. Similarly,
representatives of subgenus Cytheritis are uniquely found on Bambuseae
(Poaceae). Apart from M . anaxibia, which has its own largely unique range of
hosts, other Morphos centre on the Fabaceae, frequently with the addition of
further diverse families.
As larvae, both Antirrhea and Caerois feed on monocotyledons, mainly
Arecaceae; Caerois has been said to also exploit sugarcane (Poaceae-Bambuseae) .
Although Morphos largely utilize the Fabaceae, it seems more parsimonious to
treat dicotyledon-feeding as secondary. The predilection of Morpho (Cytheritis)
species for bamboos, an obvious trend in both the Amathusiinae and
Brassolinae, suggests that this group may constitute the ancestral host for the
entire complex, perhaps also including the Satyrinae (see below). Unlike the
Amathusiinae and Brassolinae, there is no circumstantial evidence for
unpalatability; although the uppersides of many species are brilliant iridescent
blue, the undersides show characteristic crypsis. No morphines appear to be
involved in obvious mimetic complexes (but see Young, 1972a).

Satyrinae
This cosmopolitan subfamily of perhaps 1500 species (Ehrlich & Raven,
1965) concludes the largely monocotyledon-feeding series. Miller ( 1968)
recognized seven subfamilies within the ‘Satyridae’. By down-grading his system
we arrive at the tribes Haeterini, Biini, Elymniini, Eritini, Ragadiini and
Satyrini-the Brassolinae are here retained as distinct (see above). Of the
copious data, most is applicable to the Satyrini and, with a few exceptions,
refers primarily to the Poaceae. The literature is replete with generalized records
for grasses, perhaps an indication of little host specificity. Recent studies suggest
that many Palaearctic satyrids switch freely between different grasses during
larval development (Bink, 1985).

Haeterini
This tropical American group occurs in deep forest (Miller, 1968). Current
information, although scant, suggests atypical hosts (Table 4),with species of
Pierella and Dulcedo on the Arecaceae and Pierella alone on Heliconiaceae and
Marantaceae-the Poaceae are represented by a single record, Pierella on
Panicum (Table 4). Otherwise just Fountaine’s cryptic observation of Pierella
dracontis exploiting “a kind of grass” represents this tribe. Within the Satyrinae
the predilection of the Haeterini for the Arecaceae is unusual but not entirely
unique (see also Elyrnnias, Neorina) .
104 P. R ACKERY
' I ' A B L ~ 3. Hostplants of the Morphinac. Barcant, 1970'; d'Araujo e Silva et al., 19682; DeVrirs,
1986'; Fruhstorfer, 1912-134; Hayward, 196g5; Kesselring, 19756; Miles Moss, unpublished';
Weymer, 19108; Young, 1972a9, 1982'O; Young & Muyshondt, 1972a", b ' * , 1973b"

chor inaeus . . . . . . . . .
. . . . . . . . . . . . . . . .
Antirrhea . . . . . . . . .
. . . . . . . . . . . . . . . .
geryon . . . . . . . . .
. . . . . . . . . . . . . . . .
m i L t iades . . . . . . . . .
. . . . . . . . . . . . . . . .
phi Loctetes . . . . . . . . .
. . . . . . . . . . . . . . . .
pterocopha . . . . . . . . .
. . . . . . . . . . . . . . . .
Morpho . . . . . . . . .
. . . . . . . . . . . . . . . .
(Iphimedeia) . . . . . . . . .
. . . . . . . . . . . . . . . .
hercules . . . . . 2 . . . . . . . . . . . . . . . . . . .
richardus . . . . . 2 . . . . . . . . . . . . . . . . . . .
( I p h i x i bia) . . . . . . . . . . . . . . . . . . . . . . . . .
anaxibia . 4 . 2 . . . 2 . . . 2 . . . . . . . . . . . . .
(Cyther it i s ) . . . . . . . . . . . . . . . . . . . . . . . . .
portis . . . . . . . . . . . . . . . . . . . . . . . . .
aega . . . . . . . . . . . . . . . . . . . . . . . . .
( C y p r i tis) . . . . . . . . . . . . . . . . . . . . . . . . .
rhetenor . . . . . . . . . . . . . . . . . . 7 . . . . . .
(Pessoni a) . . . . . . . . . . . . . . . . . . . . . . . . .
polyphws . . . . . . . . . . . . . . .ll . . . . . . . . .
laertes . . . . . . . . . 2 . . . . . 2 . 2 . . . . . . .
catenarius . . . . . . . . . . . . . 2 . 2 . . . . . . . . .
(trasseia) . . . . . . . . . . . . . . . . . . . . . . . . .
amathonte . . . . . . . . . . . . . . . . . . . . . . . 3 .
didius . . . . . . . . . . . . . . . . . . . . . . . . .
menelaus . . . . . . . . . . . . . . . . . 7 . . . . . . .
(Morpho) . . . . . . . . . . . . . . . . . . . . . . . . .
granadensis . . . . . . . . . . . . . . . . . 10 . . . . . . .
peleides . . . . . . . . . . . . . . 3 9 913 . 9 . 9 . 3 .
achillaena . . . . . . . . . . . . . . 12 2 . 2 . . 12 . 4 12 .
coelestis . . . . . . . . . . . . . . . . . . . . . . 2 . .
a c h i l les . . . . . . . . . . . . . . . 6 . 6 . . . . . 6 .

Biini
Recent studies (DeVries et al., 1985) suggest that the Biini (Biinae of Miller,
1968) are an unnatural group. Of the three included sub-groupings, the
Antirrheina apparently belong within the Morphinae (see above). Foodplant
information is lacking for the monobasic Biina, but the third subtribe, the
Melanitina, appears typically satyrine in host preferences. Records for Gnophodes
usually just refer to general grasses, but Melanitis itself has abundant data,
mainly grasses of the tribes Paniceae and Andropogoneae, with the occasional
addition of bamboos (Table 4).

E b m n iini
Within the Elymniini, Elymnias on Arecaceae is the outstanding feature--
otherwise the Poaceae, and to a lesser extent the Cyperaceae, are predominant.
Miller's ( 1968) four groupings correspond to the subtribes Elymniina,
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 105

3. Continued
TABLE

. . . . . . . . . . . . . . . . . . . . . . 8 . . . . . chorinaeus
. . . . . . . . . . . . . . . . . . . . . . . . . . . .Antirrhea
. . . . . . . . . . . . . . . . . . . . . . . . 8 . . . geryon
. . . . . . . . . . . . . . . . . . . . . . . . . . . 3 miltiades
. . . . . . . . . . . . . . . . . . . . . . . . . . 7 . philoctetes
. . . . . . . . . . . . . . . . . . . . . . . . . 3 . . pterocopha
. . . . . . . . . . . . . . . . . . . . . . . . . . . . Morpho
. . . . . . . . . . . . . . . . . . . . . . . . . . . . (Iphimedeia)
. . . . . . . . . . . . . . 5 . . . . . . . . . . . . . hercules
. . . . . . . . . . . . . . . . . . . . . . . . . . . . richardus
. . . . . . . . . . . . . . . . . . . . . . . . . . . . ( Ip h ixib ia)
2 . . . . . . . . . 2 . . . . . . . . . . . . . . . . . anaxibia
. . . . . . . . . . . . . . . . . . . . . . . . . . . . ( C yt h erit is)
. . . . . . . . . . . . . . . . . . 2 2 . . . . . . . . p o rt is
. . . . . . . . . . . . . . . . . 2 2 . 2 . . . . . . . aega

. . . . . . . . . . . . . . . . . . . . . . . . . . . . ( C yp rit is)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . rhetenor
. . . . . . . . . . . . . . . . . . . . . . . . . . . . (Pessonia)
. . . . . . . 11 . . . . . . . . . . . . . . . . . . . . POlYphefnuS
. . . . . . . . . . . . . . . . . . . . . . . . . . . . laertes
. . 2 . 2 . 2 . 2 . 2 . . . . . . . . . . . . . . . . . catenarius
. . . . . . . . . . . . . . . . . . . . . . . . . . . . (Grasseia)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . a mt hont e
. . . . . . . . . . . . . . . . . . . . . . . . 7 . . . didius
. . . . . . . . . . 2 . . . . . . . . . . . . . . . . . menelaus
. . . . . . . . . . . . . . . . . . . . . . . . . . . . (Morpho)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . granadensis
. . . . . . . . . . . . l . . . . . . . . . . . . . . . pe 1e i des
. . . . . . . . . . . . . . . . . . . . . . . . . . . . achillaena
. . . . . . . . . . . . . . 2 . . . . . . . . . . . . . coelest is
. . . . . . . . . . . . . . . . . . . . . . . . . . . . achi L Les

Footnote: DeVries (1986) additionally notes Caerois gerdrudtus on Socratea (Arecaceae).

Zetherina, Lethina and Mycalesina. Elymniopsis and Elymnias comprise the Old-
World Elymniina, but data are available for Elymnias species alone (Table 5);
they feed exclusively on palms (Arecaceae). The Zetherina similarly include two
genera, Callarge and Zethera, but their early-stage biology is unknown (Vane-
Wright & Smiles, 1975).
The Lethina, a group with maximum diversity in the Orient as well as Afro-
tropical, Palaearctic and Nearctic representation, have a comparative wealth of
data (Tables 5, 6). The only record for Neorina remains tentative (Arecaceae-
Calamus sp? Piepers & Snellen, 1913), but might cast doubt on its present
position. No other representative of the Lethina seems to feed on palms; indeed,
all remaining convincing records are assignable to either the Poaceae (including
the Bambuseae) or Cyperaceae.
 n -.
m v l
. . . . . . . . . . . . . . . . Aateraceae
2
. . . . . . . . . . . . . Elephantopua 0 1 . .
. . . . . . . . . . . . . . . . Poaceae
0
N . . . . . . . . . . . . Genus? P . 0.
. . . . . . . . . . . . . . . . -Bambuaeae
2 2
. . . . m . . . . 0 ) . . . . . . Bambusa
. . . . . . . . .. . . . . . . -0ryeeae
- . a
. . * .. ma.. . . . . . . . Oryza
. . . . . . . . . . . . . . . . -Ehrharteae
2
. . m . . . . . . . . . . . . . Ehrharta
. . . . . . . . . . . . . . . . -Thyaanolaeneae
2
. . . . . A N . . . . . . . . . Thyaanolaena
. . . . . . . -Poeae
. . . . . . 2 . . . . . . . . . Poa
. . . . . . . . . . . . . . . . -Eragroatideae
.a
2 . .
. . . . . . . . . . . . . Eleusine
. . . . . . . . . . . . . . . . -Paniceae
a
. . . . . a * . .
. . . . . . . Brachiaria
2
. . . . . . m . . . . . . . . . Digitaria,
A
. m a n * P N . . . . . . VI . . Panicum
. . . . . . N . . . . . . . . . Paapalum
4
. . . . . . m . . . . . . . . . Penniaetum
N
. .W . . V O . . . . . . . . . Setaria
. . . . . . N . . . . . . . . . Stenotaphrum
. . . . . . . -Andropogoneae
. . . . . . w . . . . . . . . . Apluda
2 -
. . . . . . . 0 2 . . . . . . . Coix
. . . . . . . 2 2 . . . . . . . Capillipedium
. . . . . . . . N . . . . . . . Imperata
. . . . . . . . 2 . . . . . . . Iachaemum
. , * . . . . * . a , . . . . . . Microstegium
2 4
. . . . .o.a.. . . . . . . . Miscanthua
2
a . a . . a . O N . a . . . . . Saccharum
. . . . . . m.. . . . . . . . Sorghum
2 2
. . . . . om.. . . . . . . . Zea
. . . . . . . . .. . . . . . . Cyperaceae
. . . . ...a.. . . . . . . . Genus?
. . . . . . . Heliconiaceae
2
. . . . . . . . . . . w ,
,, . VI . Heliconia
. . . . . . . . . . . . . . . . Marantaceae
. . . . . . . . . , . , . in . Calathea
. . . . . . . . . . . . . . . . Arecaceae
. . . . . . . . .. . . . . \n . Aaterogyne
. . . . . . . . . ,, . . . . . . Euterpe
. . . . . . . . . . , . , . . Geonoma
90 1
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 107

Enodia and Satyrodes represent the Lethina in the Nearctic. Apart from
generalized grasses, all specific Satyrodes records refer to the Cyperaceae. Enodia
species seem more catholic, exploiting several diverse grasses (Table 5). The
hostplants reflect the favoured habitats with the grass-feeding species typically
found in shaded forests while riverine and swampy areas are characteristic of
sedge-feeders (Shapiro & Shapiro, 1973; Howe, 1975).
Likewise, just a few small lethine genera occur in Africa. Aphysoneura
apparently favours bamboo (Arundinaria) while Paralethe and Aeropetes are found
on ‘true’ grasses. I n the characteristically Oriental Lethe and Neope, more
believable hosts are again mostly bamboos-apart from ubiquitous ‘grasses’.
Miscanthus, Microstegium, Apluda and Capillipedium are noteworthy exceptions,
often in association with disturbed habitats.
The Palaearctic genera Pararge, Lasiommata and Lopinga conclude the Lethina.
They take a similar range of grasses (most commonly representatives of the
tribes Poeae and Triticeae-see Table 6), and, rarely, sedges.
I n the Old-World tropics, the Mycalesina are the characteristic satyrines.
Host records represent the Afrotropical Bicyclus and Henotesia, and Mycalesis and
Orsotriaena in the Austro-Orient. Data for Bicyclus and Henotesia are limited and
largely restricted to general ‘grasses’ but thanks largely to Chang (1967), one
species of Mycalesis, M . gotama, is known from a wide range of hosts. It
particularly exploits grasses of the tribes Paniceae, Andropogoneae, Poeae,
Oryzeae and even Bambuseae. These preferences are widely reflected in the
limited data for other Mycalesis species (see Table 6). T h e foodplants of
0. medus, representing the more widespread of the two Orsotriaena species, follow
the same pattern with reports of Oryza (Oryzeae) and Imperata
(Andropogoneae).
The Elymniini, then, appear typically satyrine in host preferences. Most
noteworthy is the ElymniaslNeorina association with the Arecaceae and the
preference in Lethe, JVeope and Aphysoneura for bamboos.

Erititini €5’ Ragadiini

The early-stage biology of the Oriental Erites and Coelites (tribe Eritini)
remains unknown and until recently the same could have been said of the
Ragadiini. However, Fukuda (1983) has now described the life histories of both
Acrophtalmia artemis and Ragadia luzonia, representing the only currently
recognized genera. They feed on clubmosses (Selaginella-Selaginellaceae;
Table 7-see also Euptychia below).

Satyrini
Overall, the available satyrine hostplant data mostly represents the
nominotypical tribe. By adapting Miller’s (1968) system we arrive at ten
constituent subtribes, the Hypocystina, Ypthimina, Euptychiina,
Coenonymphina, Maniolina, Erebiina, Dirina, Pronophilina, Satyrina and
Melanargiina.
In the Australian region the subtribe Hypocystina predominates. Typically,
members feed on grasses of the tribe Poeae, most commonly Poa, although there
are other significant records (see Table 7). With additional representation of the
Cyperaceae, this subtribe is obviously ‘satyrine’ in host preferences.
The Old-world Ypthimina is mainly a group of tropical open country
(Miller, 1968). Most records as expected fall within the Poaceae. Ehrharta (tribe
108 P. R. ACKERY

TABLE 5. Hostplants of the Satyrinae (Elymniini; Elymniina, Lcthina, in part). Bascombe et 01..
1987'; Brown, 1973'; Chang, 19723; Harris, 1972+; Hayward, 196g5; Hill el al., 19786; Howc,
19757; Iwase, 19548; Kim, 19849; Masters, 19711°; Muroya el al., 1967"; Pennington, 197812;
Piepers & Snellen, 191313; Pholboon, 196514; Sevastopulo, 197315; Shapiro & Shapiro, 19731h;
Tani, 198317; Tietz, 197218; Uchida, 198419; van Someren, 1974*O; Wood, 19842'; Woodhousr,
19522*;Wynter Blyth, 195723

agdas . . . . . . . . . . . . . . . . . . . . . . . . .
hypernmestra . . . . . . . . . . . . . . . . . . . . . . . . .
nesaea . . . . . . . . . . . . . . . . . . . . . . . . .
panthera . . . . . . . . . . . . . . . . . . . . . . . . .
singala . . . . . . . . . . . . . . . . . . . . . . . . .
Manatar ia . . . . . . . . . . . . . . . . . . . . . . . . .
hercyna . . . . . 5 . . . . . . . . . . . . . . . . . . .
Aeropetes . . . . . . . . . . . . . . . . . . . . . . . . .
tulbaghia . . . . . . . . . . . .12 . . . . . . . . . 1 2 . .
Paratethe . . . . . . . . . . . . . . . . . . . . . . . . .
dendrophilus . . . . . . . . . . . . 12 . . . . . 12 . . . . . .
Enodia . . . . . . . . . . . . . . . . . . . . . . . . .
anthedon . . . . . . . . . . 1 0 . . . 1 0 .16 . . . . . . . .
creola . . . . 4 . . . . . . . . . . . . . . . . . . . .
portlandia . 18 . . 4 . . . . . . . . . . . . . . . . . . . .
Satyrodes . . . . . . . . . . . . . . . . . . . . . . . . .
appalachia . . . . . . . . . . . . . . . . . . . . . . . . .
eurydi ce . . . . . . . . . . . . . . . . . . . . . . . . .
Aphysoneura . . . . . . . . . . . . . . . . . . . . . . . . .
pigmentaria . . . . 20 . . . . . . . . . . . . . . . . . . . .
Lethe . . . . . . . . . . . . . . . . . . . . . . . . .
c a l l ipteri s . . . . . . . 8 . . . . . . . . . . . . . . . . .
confusa . . . . . . . . . . . . . . . . . . . . . 1 . 6 1
diana . . . . 9 . 9 8 9 . . . . . . . . . . . . . . . .
drypet is . . . . . 15 . . . . . . . . . . . . . . . . . . .
europa . . . . . 6 . . . . . . . . . . . . . . . . . 6 .
insana . . . . 15 . . . . . . . . . . . . . . . . . . . .
manthara . . . . . 13 . . . . . . . . . . . . . . . . . . .
marginalis . . . . . . . . . . . . . . . . . . . . . . . . 8
minerva . . . . . 13 . . . . . . . . . . . . . . . . . . .
rohria . . . . . . . . . . . . . . . . . . . . 1 1 . 1 .
sicelis . . . . . . . 0 . . . . . . . . . . . . . . . . .
sidonis . . . . 15 . . . . . . . . . . . . . . . . . . . .
verma . . . . . . 3 . . . . . . . . . . . . . . . . . .
Neope . . . . . . . . . . . . . . . . . . . . . . . . .
goschkevitschii . . . . . . . 8 17 . . . . . . . . . . . . . . . .
mi rheadi . . . . 1 1 1 . . . . . . . . . . . . . . . . . .
pulaha . . . . . . . 3 . . . . . . . . . . . . . . . . .
yam . . . .2315 . . . . . . . . . . . . . . . . . . .
Neorina . . . . . . . . . . . . . . . . . . . . . . . . .
crishna . . . . . . . . . . . . . . . . . . . . . . . .

Ehrharteae) is the typical host in the Afrotropics, although other grasses are also
widely reported (see Table 8). Representatives of further closely associated
monocotyledonous families Uuncaceae, Zingiberaceae, Cyperaceae and
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 109

TABLE
5. Continued

. . . . . . . . . 21 . . . . . . . . . agondas
. . . . . . 1 15 13 151 15 14 1 . 15 . 11 19 hypernnestra
. . . . . . . . . . 13 . . . . . . . nesaea
13
. . . . . . . . 13 . . . . . 13 . 13 . . panthera
. . . . . 22 . . . . . . . . . . . . . singala
. . . . . . . . . . . . . . . . . . . Manataria
. . . . . . . . . . . . . . . . . . . hercyna
. . . . . . . . . . . . . . . . . . . Aeropetes
. . . . . . . . . . . . . . . . . . . t ulbaghia
. . . . . . . . . . . . . . . . . . . Paralethe
. . . . . . . . . . . . . . . . . . . dendrophi lus
. . . . . . . . . . . . . . . . . . .Enodia
. . . . . . . . . . . . . . . . . . . anthedon
. . . . . . . . . . . . . . . . . . . creola
. . . . . . . . . . . . . . . . . . . portlandia
. . . . . . . . . . . . . . . . . . . Satyrodes
. 16 2 . . . . . . . . . . . . . . . . appalachia
. 7 . 18 . . . . . . . . . . . . . . . eurydice
. . . . . . . . . . . . . . . . . . . Aphysoneura
. . . . . . . . . . . . . . . . . . . pigmentaria
. . . . . . . . . . . . . . . . . . .Lethe
. . . . . . . . . . . . . . . . . . . callipteris
. . . . . . . . . . . . . . . . . . . confuse
. . . . . . . . . . . . . . . . . . . diana
. . . . . . . . . . . . . . . . . . . drypetis
. . . . . . . . . . . . . . . . . . . europa
. . . . . . . . . . . . . . . . . . . insana
. . . . . . . . . . . . . . . . . . . manthara
. . . . . . . . . . . . . . . . . . . marginalis
. . . . . . . . . . . . . . . . . . . minerva
. . . . . . . . . . . . . . . . . . . rohria
. . . . . . . . . . . . . . . . . . . sicelis
. . . . . . . . . . . . . . . . . . . sidonis
. . . . . . . . . . . . . . . . . . . verma
. . . . . . . . . . . . . . . . . . . Neope
. . . . . . . . . . . . . . . . . . . goschkevi tschi i
. . . . . . . . . . . . . . . . . . . muirheadi
. . . . . . . . . . . . . . . . . . . pulaha
. . . . . . . . . . . . . . . . . . . yama
. . . . . . . . . . . . . . . . . . .Neorina
. . . . . . . . . 1 3 . . . . . . . . . crishna

Restoniaceae) appear as minor 'secondary' hosts. Oriental representatives (Xois

and Ypthima in part) centre more exclusively on the Poaceae, notably the tribes
Andropogoneae and Paniceae.
110 P. R. ACKERY

TABLE 6. Hostplants of the Satyrinae (Elymnnini; Lethina, in part, Mycalesina). Bascombe et al.,
1987'; Chang, 19672; Common & Waterhouse, 19723; De Viedma & Gomez Bustillo, 19764;
Fountaine, unpublished5; Gifford, 19656; Gullander, 195g7; Henriksen & Kreutzer, 198Z8; Hill
et al., 1978'; Iwase, 19541°, 1964"; Kim, 198412; Kuzuya, 196713; Langer, 195814; Larscn,
197415; hlamet, 194816; Manley & Allcard, 197017; Pennington, 197818; Piepers & Snellcn,
1913''; Pinhey, 194gZ0;Robert et al., 198321;Rosevear, 197822;Schwarz, 194823;Sevastopulo,
1973". 1975'5; 'Iakahashi, 197426,197527;Valletta, 197Zz8;van Someren, 197429;Woodhouse,
195230

Pararge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
aeger ia . . . . . . . . . . . . . . . 7 . 28 . 15 28 15 . . . 15 . . 15 .
epimeni des . . . . . . . . . . . . . . . . . . . 12 . . . . . . . . 13 .
schrenkii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Lasiunnata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
maera . . . . . . . . . . . . . . . . . . . . 15 . . 15 . . 15 17 15 .
megera . . . . . . . . . . . . . . . 28 . 28 . . 28 . . . . 15 15 23 15 .
petropolitana. . . . . . . . . . . . . . . . . . . . . . . . . 14 7 . . .
Lopinga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
ach ine . . . . . . . . . . . . . . . 7 . . . 1 4 - 4 . . . . . 4 4 .
Henotesi a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
narcissus . . . 16 . . . . . . . . . 16 . . . . . . . . . . . . . . . .
perspicua . . . . . . . . . . . 18 . . . . . . . . . . . . . . . . . . .
sinwnsi i . 6 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
vola . 5 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
B i cyclus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
anynana . . . . . . . . . . . 18 . . . . . . . . . . . . . . . . . .
canpina .25 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
dubi a .25 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
iccius .29 . . . . . . . . . . . . . . . . . . . . . . . . . .
rnedontias . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
miriam .25 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
rhacot is .20 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
saf itza . . . . . . . . . . . 18 . . . . . . . . . . . . . . . . . .
saussurei .25 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Mycalesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
anapi t a - 2 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
gotama . . . .26 . . . 210 . . . . . . . . . . . . . . . . . .27 .
horsfieldi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
janardana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
m i neus . . . . . . 1 . 1 . . . . P . . . . . . . . . . . . . . . .
patnia . . . . . . . . . 24 . . . . . . . . . . . . . . . . . . . .
perseus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
rarna . . .3 0 . . . . . . . . . . . . . . . . . . . . . . . . . .
Sirius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
terminus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
visala .24 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Orsotriaena . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
medus . . . . . . . . . 3 . . . . . . . . . . . . . . . . . . . .

The New-World Euptychiina have many affinities with the Ypthimina

although, unlike their Old-World relatives, they characteristically inhabit woods
or forest (Miller, 1968). Following Forster (1964), 'Euptychia' (sensu luto) is now
divided into innumerable unfamiliar genera. In trying to correlate host records,
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 111

TABLE
6.Continued

. . . . . . . . . . . . . . . . . . . Pararge
8 .. .21 . . . . . . . . . . . . . . . aegeria
. . . . . . 12 . . . . 13. 12 . . . 13 . . epimenides
. . . . . . . . . . . . . 12 . . . 12 . . schrenkii
. . . . . . . . . . . . . . . . . . . . Lasimta
. . . . . . . . . . . . . . . . . . . . maera
. . . . . . . . . . . . . . . . . . . . megera
. . . . . . . . . . . . . . . . . . . . pet ropol i tana

. . . . . . . . . . . . . . . . . . . . Lopinga
. . . . . . . . . . . . . . . . . 14 . . achine
. . . . . . . . . . . . . . . . . . . .Henotesia
. . . . . . . . . . . . . . . . . . . . narcissus
. . . . . . . . . . . . . . . . . . . . perspi cua
. . . . . . . . . . . . . . . . . . . . simonsii
. . . . . . . . . . . . . . . . . . . . vola
. . . . . . . . . . . . . . . . . . . . B i cyclus
. . . . . . . . . . . . . . . . . . . . anynana
. . . . . . . . . . . . . . . . . . . . canpina
. . . . . . . . . . . . . . . . . . . . dubia
. . . . . . . . . . . . . . . . . . . . iccius
. . . . . . . . . . . . . . . . . . . 2 2 medontias
. . . . . . . . . . . . . . . . . . . . miriam
. . . . . . . . . . . . . . . . . . . . rhacotis
. . . . . . . . . . . . . . . . . . . . safitza
. . . . . . . . . . . . . . . . . . . . saussurei
. . . . . . . . . . . . . . . . . . . . Myca Lesi s
. . . . . . . . . . . . . . . . . . . . anapita
. . . . . . 2 10 2 2 . . . 11 . 2 . . . . gotama
. . . . . . . . . . . 1 9 . . . . . . . . horsfieldi
. . . . . . . . 19 . . . . . . . . . . . janardana
. . 1 . . . . . . . . . 9 . 9 . . . . . mineus
. . . . . . . . . . . . . . . . . . . . patnia
. . . . . . . . . . . 19 . . . . . . . . perseus
. . . . . . . . . . . . . . . . . . . . rama
. . . . . . . . . . . 3 . . . . . . . . sirius
. . . . . . . . . . . 3 . . . . . . . . terminus
. . . . . . . . . . . . . . . . . . . . visela
. . . . . . . . . . . . . . . . . . . . Orsotriaena
. . . . . . . . . . . 1 9 . . . . . . . . medus

two important qualifications must be made-misidentifications are likely to be

rife, and Euptychia (sensu stricto) must surely contain a rag-bag of species,
including many not covered by Forster. Otherwise, available data only applies
to T'getis, Satyrotaygetis, Oressinoma and Megisto.
 112 P. K. ACKERY

'rA BLE 7. Hostplants of the Satyrinae (Ragadiini; Satyrini, Hypocystina). Common &
Lt'aterhouse, 1972', 1981'; Craw. 1978'; D'Abrera, 19774; De Baar, 19815; Fukuda, 1983'j;
Gibb5. 1980'; Holloway & Peters, 1976*; Laidlaw, 19709; McCubbin, 1971l o ; Wilson, 1982"

artemis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Ragadia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Luzonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Argynnina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
hobartia . . . . . . . . . . . . . . . . 2 . . . . . . . . . . . . . . . .
Argyrophenga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
antipodum . . . . . . . . . 7 . . . 3 . 3 . 9 . . . . . . . . . . . . . . .
harrisi . . . . . . . . . 7 . . . . . . . . . . . . . . . . . . . . . . .
janitae . . . . . . . . . 3 . . . . . . . . . . . . . . . . . . . . . . .
Aust roypth irna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
petersi . . . 8. . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Dodonidia . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . .
helrnsii . . . .. . . . . 7 . . . . . . . . . . . . . . . . . . . 7 . . .
Erebiola . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . .
butleri . 9 . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ceitoneura . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . .
acantha . . . .
. . . . . . . . . . . . . l o . . . . . . 1 . . . . . . . .
hobartia . I . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . .
klugii . . . . . . . . . . . 2 . . . . . 2 . . . . . . 2 . . . . . . . .
Heteronynpha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
banksii . . . . . . . . . . . . . . . . . 1 . . . . . . . . . 2 . . . . .
cordace . 2 . . . . . . . . . . . . . . . . . . . . . . . . . l . . . . .
merope . . . . . . . . . . . 2 . . . . . 2 . . . . . . 2 . . . . . . . .
rnirifica . 1 . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . .
paradelpha.. . . . . . . . . . . . . . . . 2 . . . . . . . . . . . . . . .
penelope . . . . . . 1 . . . . . . . . 1 . . . .
. . . . 1 . . . . . . . .
solandri . .
. . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . .
Hypocysta . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
adiante . . . . . . . . . . . . . . . . . . . . . 5 . 5 5 . . . . . . . .
euphemia . . . . . . . . . . . . . . . . . . . . . .
4 . . . . . . . . . .
irius . .
. . . . . . . . . . . . . . . . . . . . . l . . . . . . . . .
metirius . . . . . . . . . . . . . . . . . . . 1 . . . 1 . . . . . . . . .
pseudirius. 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Nesoxenica . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Leprea . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . .
Dreixenica . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
corrae . .
. . . . . . . . . . . . . . . 10. . . . . . . . . . . . . . .
kershaui . .. . . -10. . . . . . . . . . 1 . . . . . . . . . . . . . . .
tathonielia . . . . . 1 . . . . . . . . . . . 1 . . . . . . . . . . . . . . .
latialis . . . . . . . .
. . . . . . . . . 2 . . . . . . . . . . . . . . .
orichora . . . . . . . . . . . . . .10 . . . . . . . . . . . . . . .
. . .
ptunarra . 4 . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . .
Parat isiphone . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . .
Lyrnessa . . . . . . . . . . . . . . . . . . . . . . . . . . 8 . 8 . . . .
Percnodaimon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
meruta . .. . . . . . . . . . . . . . .9 . . . . . . . . . . . . . . .
Tisiphone . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
abeona . .. . . . . . . . . . . . . . . . . . . . . . . . . . . l . . .
helena . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . 1 1 . . .
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 113

TABLE 8. Hostplants of the Satyrinae (Satyrini; Ypthimina). Bascombe et al., 1987I; Common &
Waterhouse, 1972*, 19823; Hill et al., 19784; Iwase, 19545, 19646; Kashiwai, 1982'; Pennington,
19788; Piepers & Snellen, 19139; Pinhey, 1949IO; Sevastopulo, 197311, 197512; Woodhouse,
195213

nirmala . . . -11 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Cassionynpha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
cassius . . . . . . . . . . 8 . 8 . . . . . . . . . . . . . . . . 8 . . . .
Coenyra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
aurantiaca . . . . 8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Coenyropsis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
bera . . . . 12 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Mashuna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
mahuna . . . . . . . . . . . . . . . . . .8 . . . . . . . . . . . . . . .
Melampias . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
huebneri . . . . . . . . 8 . . . . . 8 . . . . . . . . . . . . . . . . . . .
Neita . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
durbani . . . . . . . . 8 . . . . . . . . . . . . . . . . . . . . . . . . .
ex t ensa . . . . . . . . 0 . . . . . . . . . 10 . . . . . . . . . . . . . . .
Neocoenyra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
duplex . . . . 12 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Physcaeneura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
leda . . . . 12 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
panda . . . . . . . . 8 . . . . . . . . . . . . . . . . . . . . . . . . .
pione . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 0
Pseudonynpha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
detecta . 8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 . .
hippia . 0 8 . . . . . 8 . . . . . . . . . . . . . . . . . . . . . . . . .
magnus . . . . . . . . 8 . . . . . . . . . 0 . . . . . . . . . . . . . . .
magoides . . . . . . . . 8 . . . . . . . . . . . . . . . . . . . . . . . . .
trimeni . . . . . . . . . . . 8 . . . . . . . . . . . . . . . . . . . . . .
Stygionynpha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
dicksoni . . . . . . . . . . . . . . . . 8 . . . . . . . . . . . . . . . . .
irrorata . . . . 8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
robertsoni . . . . . . . . 8 . . . . . . . . . . . . . . . . . . . . . . . . .
vigilans . 8 . . . . . . 8 . . . . . . . . . . . . . . . . . . . . . . . . .
uichgrafi . . . . 8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Xois . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
arctoa . . . . . . . . . . . . . . . . . . . . . . . . 3 . . . . . . . . .
Ypthima . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
albida . . . . 12 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
argus . . . . . . . . . . . . . . . . . . . . . 5 . . . . . . . . . . . . .
asterope . . . . . . . . . . . . . . . . . .10 . . . . . . . . . . . . . . .
avanta . . . . 13 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
baldus . . . . . . . . . . . . . . . . . . 9...9..4.4......
ceylonica . . . .ll . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
inplra . . . . . . . . 8 . . . . . . . . . . . . . . . . . . . . . . . . .
tisandra . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4 . . . . . . .
mtschulskyi. . . . . . . . . . . . . . . . . . . . 5 . . . . . . . . . . . . .
n i kaea . . . . 11 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
pandocus . . . . . . 9 . . . . . . . . . . . . . . . 9.9....... . .
praenubila . . . . . . . . . . . . . . . . . . . . . . 1 . . 1 . 4 . . . . . .
riukiuana . . . . 6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
sensilis . . . . . . . . . . . . . . . . . . . . . 7 - . - . . . . . . . . .
114 P. R. ACKERY

ellJu!~elaS. . . . . . . . . . . . . . . . . . N . . . . . . . . . . . .
aea,ell Ju!ge,aS. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
s!sdo,Jy,aN. . . . . . . . . . . . . . . . . N . . . . . . . . . . . . .
aea,e,ay,aN . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
e,~laM . .................
~ . . . . . . . . . . . .
emouoa3 . . . . . . . . . . . . . . . . . .
N . . . . . . . . . . . .
ad,aln3 . . . . . . . . . . . . . . . . . .
N . . . . . . . . . . . .
Jea,e,J,v.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
eaqlele3 . N . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
JeJ,elue,eyy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
snd,!,p . . . . . . . . . . . . . . . . .a . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . N .

sn,ad13 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
xa,e3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
.D.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
run3taeqs,os . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . -
N

aeauosodo,puv..
lunledeed
mn,,ued
. . . -- -
. . . . . . . . . . . . . . . . . . . . . . . . . .
.C . . - . . . . . . . . . . . . . . . . . . . . .
. N . . N

snuams!ldo. . . . . . . . . . . . . . . . . . . . . . . . . . .
7 c
I N . .

enqlueuq,l.. . c- . . . .
. . . . . . . . . . . . . . . . . . .
. N . ,

e ~ , e l
..-
~ . . . . . .
~ ~ a ~ ~ ~ ~ ~ . ~

sndouoxv . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .a
ee,a,o,,v. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
aea,!"ed-. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
uopou~3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . OD

aeJp!lolq3-. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
au,="alp . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
N N

mn!ualso[ll>ea. . . . . . . . . . . . . . . . . . . . - . . . . . . . . . . N

aeap!)80J~o,a- . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
eod
. . . . . . . . . . . . . . . . . . . . . . . . . . . Y \ . . m

B!ldl?FcI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 0
r

JeJOd-
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
e,Alo. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
aea,llO-.
eol,ouallemS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N

eanbsnq3.. . . . . . . . . . . . . . . . . . . . . .
. N . . . N . .

esn~mea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
is"ua3.
aeJseod..
.
* - N * m s N . . -
. . . . . . . . . . . . . . . . . .
7
. . . . . .
. N . U . .

. .
U

. . . . . . . . .
- U
F

. . . . . . . . . . . . . . . . . . N . . . . . . . . . . . .
aea>ep!JAX' . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
c

mn!lo,!,~. . . . . . . . . . . . . . . . N . . . . . . . . . . . . . .
r
aeJ,eqed' . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
c a .-
m
a m
a m .- u-x

.-V L
u w 0
 . .......................... Trifolium
. . . . . . . . . . . . . . . . . . . . . . . . . . . Xyrid-ae
. . . . . . . . . . . . . . . . . . . . . . . w . . . XYriS
. . . . . . . . . . . . . . . . . . . . . . . . . . . Poaceae
N L n . . N .
. . . . . . . . . . ul. N .
. . . . . . . Genus?
. . . . . . . . . . . . . . . . . . . . . . . . . . . -Bambuseae
A
. . . . . . L n W . .
* . . . . . . . . . . . . . . . . Bambusa
. . . . . . ,,,. . . . . . . . . . . . . . . . . . . . Chusquea
. .......................... Swallenocloa
. . . . . . . . . . . . . . . . . . . . . . . . . . . -Ob=
. . . . . . . . N . . . . . . . . . . . . . . . . . . Olyra
. . . . . . . . . . . . . . . . . . . . . . . . . . . -Poeae
. . . . . . . . . . . . . . . . . . . . . . . . . . . Dactylis
. . . . . . . . . . . . . . . . - . . . . . . . . . . Poa
. . . . . . . . . . . . . . . . . . . . . . . . . . . -Eragrostideae
. . . . . . . . . . . . . . . . . . . . . . . . . . . Dactylocteniurn
. . . N . . . . . . . . . . . . . . . N . . . . . . . Eleusine
. . . . . . . . . . . . . . . . . . . . . . . . . . . -Chlorideae
. .......................... Cynodon
. . . . . . . . . . . . . . . . . . . . . . . . . . . -Paniceae
A
. . . . . . . . L n . .
. . . . . . . . . . . . . . . . Acroceraa
. . . . . . . . . . . . . . . . . . . . . . . . . . . Axonopus
. . . . . . . . . . . . . . . . . . . . . . . . . . . Digitaria
. . . . . . . . . . . . . . . . . . . . . . . . . . . Ichnanthus
. . . . . . . . . . . . . . . . . . . . . . . . . . . Oplismenus
. . . . . . . . . . . . . . . . . . . . . .
N . . PanicumN .
. . . . . . . . . . . . . . . . . . . . . . . . . . . Paapalum
. . . . . . . . . . . . . . . . . . . . . . . . . . . -Andropogoneae
. . . . . . . . . . . . . . . . . . . . . . . . . . . Sorghaatrum
. . . . . . . . . . . . . . . . . . . . . . . . . . . Cyperaceae
. . . . . . . . . . . . . . . . . . . . . . . . . . . Carex
. . . . . . . . . . . . . . . . . . . . . N . . . . . . Cyporus
. . . . . . . . . . . . . . . . . . . . . . . . . . . . Laparus
. . . . . . . . . . . . . . . . . . . . . . . . . . . Scirpus
. . . . . ..................... .Marantaceae
. . . . . ...................... Calathea
. . . . . ...................... Arecaceae
. . . . . . . . . . . . . . . . . . . . . . . . . . . Euterpe
. . . . . ...................... Geonoma
. . . . . . . . . . . . . . . . . . . . . . . . . . . Welfia
. . . . . . . . . . . . . . . . . . . . . . . . . . . Neckeraceae
. . . . . ...................... Neckeropsis
. . . . . ...................... Selaginellaceae
. . . . . ...................... Selaginella
S317dX?ILLLntl (II7VHdMIAN :NOILV3IdISSV73 (INV SLNV7dLSOH
116 P. R. ACKERY

Within ‘Euptychia’ (sensu lato), Gissia is tentatively redefined by Singer,

DeVries & Ehrlich (1983). Most species occur on grasses of the tribe Paniceae,
with the notable addition of the Arecaceae and Marantaceae for C. confusa.
Otherwise, only C. lube is unusual, exploiting “bambusoid grasses” (DeVries,
1986). Elsewhere in ‘Euptychia’, other than in Euptychia itself, records are scant
and often generalized (Table 9 ) . Notably, Eleusine (tribe Eragrostideae) is quite
widely accepted while Splendeuptychia utilizes Bambusa.
Food plants of the residual Euptychia species probably underline thc
unnaturalness of the group. Outstanding are E. insolata on an epiphytic moss
(.;lreckeropsis undulata) , E. mitchellii on Carex and Scirpus (Cyperaceae), E. cymela on
the barely believable Trzfolium (Fabaceae) together with the more convincing
Xyris (Xyridaceae), and E. mollina (possibly a complex of species) on Selaginella.
In discussing the evolution of this last host preference, Singer, Ehrlich & Gilbert
(1971) drew attention to the habit of several grass-feeding species of ovipositing
off their larval hosts, thus repeatedly exposing early instars to such plants as
mosses and ferns (see Chew & Robbins, 1984). Indeed, Ebner (1970) notes
E. cymela ovipositing at the base of moss-covered tree-trunks. Alternatively, there
is some evidence of chemical similarity between Selaginella and certain grasses
(Singer et al., 1971). Elsewhere in Euptychia, several species are said to exploit
unspecified grasses. Less generalized records refer to the tribes Bambuseae,
Poeae, Eragrostideae, Paniceae and Andropogoneae (see Table 9).
The remaining euptychiines are interesting in picking up various Euptychia
themes. Taygetis favours bamboos, although other grasses have occasionally been
noted. The sole record for Oressinoma cites Cyperus (Cyperaceae), while Xyri.r
(Xyridaceae) alone typifies Megisto, with Satyrotaygetis on true grasses (Panicum).
However, much more information is required to confirm the reality of such
specificity. Overall, the Euptychiina are typically ‘satyrine’, with notable
contributions from the Arecaceae, Marantaceae, Xyridaceae, Poaceae
(Bambuseae) and Selaginellaceae.
In northern temperate areas, the Coenonymphina replace the closely related
Ypthimina (Miller, 1968). Apart from shared records for Poa and Brachypodium,
Coenonympha species and A. hyperantus (the sole representative of Aphantopus) have
a barely overlapping proliferation of poaceous hosts (see Table 10) with the
occasional addition of the Cyperaceae or even Iridaceae.
Rather more limited information suggests that species of the genera Cercyonis,
Hyponephele, Maniola and Pyronia, the Maniolina, are marginally less catholic.
‘This subtribe, closely- related to the preceding, has a similarly Holarctic
distribution, and again the tribe Poeae predominates (Table 10). Just one
record, for C. pegala on sedges (Ferris & Brown, 1981), falls outside the Poaceae.
T h e Erebiina, yet another Holarctic subtribe, are probably monobasic.
Characteristic of alpine and sub-Artic zones, the large genus Erebia lacks obvious
affinities with any other satyrine group, with the possible exception of the
Ypthimina (Miller, 1968). Association with the tribe Poeae (principally Poa and
Festuce-see Mansell, 1985) is even more marked than in the preceding two
groups. Apart from E. oeme on Juncus and Luzula Uuncaceae), grasses and to a
lesser extent sedges dominate the records (see Table 11).
These three Holarctic subtribes, the Coenonymphina, Maniolina and
Erebiina, thus all centre on the poaceous tribe Poeae (most commonly Poa and
Festuca) . Within the Afrotropical Dirina (host records available for Dzra,
 10. Hostplants of the Satyrinae (Satyrini; Coenonymphina, Maniolina). Brooks & Knight, 1982’; De Viedma & Gomez Bus6ll0, 1976*;Ferris &
E
, 198l3; Forster & Wohlfahrt, 19554; Gullander, 195g5; Henriksen & Kreutzer, 1982‘j; Higgins & Riley, 1970’; Howarth, 19738; Howe, 1975’;
1954’O; Langer, 1958’l; Larsen, 197412; Leech, 1892j3; Manley & Allcard, 197014; Robert el al., 198315; Schwarz, 194816; Shapiro & Shapiro,
197317;Tietz, 197218

hyprrntur. . . . . 1 . . . 16 . . . . . . . 8 . . 14 . . . . . 16 . . 16 . . . 14 . . . . . . . . 16 . . . .
ornonymphr... ............................................
rmprlor . 1 8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
rrcrnlr ... . . . .14 . . . . . . . . . .14 - 1 4 . . . . . . . . . . . . . - 1 4 . . . . . . . . . . .
crllfornlr... . . . . . . . . . . . . . . . ..la.. . . . . . . . . . . ..la.. . . . . . . . . .
dorua . . . . . . . . . . . . . . . . . .1 4 . . . . . . . . . M . . . . . . . . . . . . . . . . . .
glycarlon . . . . . 11 . 15 . . . . . . . 11 11 . 15 . 15 . . . . . . . 15 . . . . . . 11 . . . . . . . . . . .
hero . . . . . . . . . .1 6 1 1 6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0 . . . . .
Inornrtr . . . . . . . . . . . . . . . . . . .. l a . . . . . . . . . . . . . . . . . . . . . . .13. .
odlppus . . . . .14 . 1 4 . . . . . . , 1 4 . .14 2 2 . . . . . . . . . . . . . . . . . . . ., 1 3 . . . 2
prmphllur . . . . . . . . . . . . . . . . 14 m12 . 1 2 . . 5 . . . . . . . . . . . . I 1 2 . . . . . . . . .
tullir . . . . . . . . . . . . . . . . . .5 . . . . . . . . . . . . 1 6 . . . . . . . ...8168..
rrcyonfs . . . ............................................
mrdil . 9 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
octue . ................... 9 .
. . . . . . . . . . . . . . . . . . . . . . . . .
pegala . . .18. . . . . . . . . . . . . . . . . . ..18 . . . . . . . . . . . . . . 1 7 . 5 . . . . .
sthenslc . ................... 9 .
. . . . . . . . . . . . . . . . . . . . . . . . .
yponcphelc . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
lycaon . . . . . . . . . . . . . . . . . .14 .14 .
1 1 . . ......................
lupinur . ................... 1 5
. . . . . . . . . . . . . . . . . . . . . . . . . .
sniolr . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
jurtinn . ................... 1 4
. . . . . . . . . . . . 6 . . . . . . . . . . . . . 6
nurag . 7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
yronia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
bathesba . . . . . 1 5 . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Cecilia . . . . . 1 5 . . . . . . . . . . . . . . . .
4 . . . . . . . . . . . . . . . . . . . . . . . .
tithonus 14 14
. . . . . . . . . . . . . . . 11 14 14 8
. . . . . . . . . . . . 14 . . . . . . . . . . . . .
118 P. R. ACKERY
'I'ABLE11. Hostplants of the Satyrinae (Satyrini; Erebiina). De Viedma & Gomez Bustillo, 1976';
Dornfcld, 1980'; Forster & Wohlfart, 19553; Gullander, 1959'; Henriksen & Kreutzer, 19825;
Higgins & Riley, 19706; Howarth, 1973'; Iwase, 1954*; Langer, 19589; Manley & Allcard,
1970'"; Kel, 1982"; Pyle, 1974"; Schwarz, 194813;Scott & Scott, 1980'+; Tabuchi, 195915;
Tietz, 197216

ligea . . . . . . . . . . . . . . . . 4 . . . . . 13 . . . . . a
euryale . . . . . . 10 . . . . . . . . . 9 . . . . . 9 . . . . . .
eriphyle . . . . . . . . 13 . . . . . . . . . 13 . . . . . . . . . .
manto . . . . . . 10 . . . . . . . . . . . . . . . . . . . . . .
claudina . . . . . . . . . . 6 1 3 . . . . . . . . . . . . . . . . .
flavofasciata . . . . . . 3 . . . . . . . . . . . . . . . . . . . . . .
epiphron . . . . . . 10 .
10 . 13 10 . . . . . . . . 7 . . . . . . . .
christi . . . . . . 3 . . . . . . . . . . . . . . . . . . . . . .
pharte . . . . . . 13.13 . . . . . . . . . . . . . . . . . . . .
melenpus . . . . . . . .13 . . . . . . . . . . . . . . . . . . . .
vidleri .12 . . . . . . . . . . . . . . . . . . . . . . . . . . .
aethiops . . . 7 . 13 . . 13 . 13 . . . 13 . . . . . . . . . . . . . .
niphonica . . . . . . . . . . . . . . . . . . . . . . . . . . . .15
t r ia r ius . . . . . . .lo10 . . . . . . . . . . . . . . . . . . . .
&la . . . . . . . . . . 9 5 . . . . . . . . . . . . . . . . 4
magdalena . . . . . . . . . . . . . . . . . . . . . . . . . . . ,14
medusa . . . . . . - 1 3 1 3 . . . . . . .13 . . . . . 1 3 . . . . . .
discoidalis . . . . . . . . 16 . . . . . . . . . . . . . . . . . . . 16
alberganus . . . . . . 13 . 13 . . . . . . . . . . . . . . . . . . . .
epipsodea . 2 . . . . . . . . . . . . . . . . . . . . . . . . . 1 4 .
pluto . . . . . . . . 3 . . . . . . . . . . . . . . . . . . . .
gorge . . . . . . 10 . 1 0 . . . . . . . . . . . . . . . . . . . .
epistygne . . . . . . 1 . I . . . . . . . . . . . . . . . . . . . .
tyndarus . . . . . .13 - 1 3 . . . . . . . . . . . . . . . . . . . .
pronoe . . . . . . . . 10 . . . . . . . . . . . . . . . . . . . .
scipio . . . . . . 11 . . . . . . . . . . . . . . . . . . . . . .
st i r ius . . . . . . . . 3 . . . . . . . . . 3 . . . . . . . . . .
mtanus . 6 . . . . . . . . . . . . . . . . . . . . . . . . . . .
rapateri . . . . . . 1 . 1 . . . . . . . . . . . . . . . . . . . .
neoridas . . . . . . 10 . 10 . . . . . . . . . . . . . 10 . . . . . .
oeme . . . . . . 10 . 13 . . . 10 . . . . . . . . . . . 10 13 . . .
meolans . . . . . . 1 0 10 10 . . . . . . . . . . . . . . . . . . . .
palarica . . . . . . 1 . 1 . . . . . . . . . . . . . . . . . . . .
pandrose . . . . . . 1 0 .10 . . . . . . . . . . . . . . . . . . . .

'Tarsocera and Torynesis) (Table 12) emphasis switches again to the tribes
Ehrharteae and Danthonieae, the groups most typical of African satyrines (see
Mycalesina, Ypthimina and Paralethe). However, if current data are
comprehensive, host partitioning between these three genera appears complete.
All these butterflies are seemingly upland species, lacking obvious habitat
differentiation to account for their hostplant segregation.
In the essentially South American Pronophilina, the poaceous tribe
Bambuseae predominates (Table 12). Pronophilines occur in a range of habitats
 HOSTPLANTS AND CLASS1FI CATION: NY M PHALI D BUTTERFLIES 119

TABLE12. Hostplants of the Satyrinae (Satyrini; Dirina, Pronophilina). Brown & Heineman,
1972'; Cornstock, 19442; Cuberto, 19853; Dethier, 19404; DeVries, 1980a5, 19866; Hayward,
1969'; Pennington, 19788; Riley, 19759; Shapiro, 1982'O; Weymer, 1912"

clytus . . . . . . . . 8 . 8 . . . . . . . . . . . . 0 0 . . . . .
jansei . . . . . . . . 8 . . . . . . . . . . . . . . . . . . . . .
oxylus . . . . . . . . 8 . . . . . . . . . . . . . . . . . . . . .
suanepoeli . . . . . . . . 8 . . . . . . . . .8 . . . . . . . . . . .
Tarsocera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
cassina . . . . . . . . . . . . . . 8 . 8 . . . . . . . . . . . . .
cassus . . . . . . . . . . . . . . . . . . . . . . . . . . 8 . . .
Torynesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
hauequas . . . . . . . . . . . . 8 . . . . . . . . . . . . . . . . .
mintha . . . . . . . . . . . . 8 . . . . . . . . . . . . . . . . .
Argyrophorus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
argenteus . . . . . . . . . . . . . . . . . . . . 10 . . . . . . . . .
Calisto . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
herophile . . . . . . . . . . . . . . . . . . . . 4 . . . . . . 9 . 9
nubi l a . . . . . . . . . . . . . . . . . . . . 2 . . . . . . . . .
pulchella . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 . .
zangis . . . . . . . . . . . . . . . . . . . . 1 . 1 . . . . . . .
D io r i s t e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
cothonides . . . 6 . . . . . . . . . . . . . . . . . . . . . . . . . .
Elina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Lefebvrei . . . 7 . . . . . . . . . . . . . . . . . . . . . . . . . .
Eretris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
suzannae . . . 5 . . . . . . . . . . . . . . . . . . . . . . . . . .
Lymanopoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
sami us . . . 11 . . . . . . . . . . . . . . . . . . . . . . . . . .
Oxeochistus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
puera . . . 3 . . 3 . . . . . . . . . . . . . . . . . . . . . . .
Pedalides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
phrasicla . . . . 6 . . . . . . . . . . . . . . . . . . . . . . . . .
Praepedal i d e s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
phanias . . 7 7 . . . . . . . . . . . . . . . . . . . . . . . . . .
Pseudolnanio l a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
rogersi . . . 6 . . . . . . . . . . . . . . . . . . . . . . . . . .

from tropical forests to southern temperate zones and Andean highlands (Miller,
1968). Exploitation of the bambusoid genus Chusquea is typical of the subtribe.
Species of Calisto and Argyrophorus, however, favour non-bambusoid grasses,
especially Cynodon (tribe Chlorideae) . These preferences for bamboos and non-
bambusoid grasses are uncorrelated with Miller's suggested generic series.
The Holarctic Satyrina and the Palaearctic Melanargiina conclude the
Satyrini. In host preferences, both are typical temperate satyrine groups. The
Satyrina particularly favour the Poeae, Brachypodieae and Brorneae, but wide-
ranging available records indicate a constellation of other poaceous foodplants
(see Table 13). As so often in temperate satyrines, the Cyperaceae are also
TABLE 13. Hostplants of the Satyrinae (Satyrini; Satyrina). De Viedma & Gomez Bustillo, 197G1;
Dornfeld, 19802; Ebner, 19703; Emmel & Emmel, 19744; Ferris & Brown, 19815; Forster &
LVohlfahrt, 19556; Gullander, 1959'; Henriksen & Kreutzer, 19828; Higgins & Riley, 19709;
Howarth, 1973'O; Howc, 19751'; Iwase, 195412; Kim, 198413; Larsen, 197414; Larsen & Larsen,
1980l5; Lccch, 189216; Manley & Allcard, 1970"; Masters & Sorensen, 1969'a; Oehmiglg; Pyle.
1971": Robert et al., 198321;Schwarz, 194822;Scott & Scott, 19802'; Sevastopulo, 1973L4;
ShirGzu, 196Oz5;Tabuchi, I95gz6;Teitz, 197Z2'; Wattison, 192gZ8;M'eymer, 1911 2 9

arcthusa . . . . . . '. . . . . . . . . . 22 22 22 22 . . . . . .
Bcrberia . . . . . . . . . . . . . . . . . . . . . . . . . .
abdelkader . . . . 9 . . . . . . . . . . . . . . . . . . . . .
Brintcsia . . . . . . . . . . . . . . . . . . . . . . . . . .
c i rce . . . . . . . . 17 . 21 . . . . . . 1 7 1 7 . . . 17 . . 21
Chazara . . . . . . . . . . . . . . . . . . . . . . . . . .
briseis . . . . . . . . 22 . . . . . . . . . 22 22 . . . . . .
prieuri . . . . . . . . . . . . . . . . . . .21 . . . . . .
Hipparchia . . . . . . . . . . . . . . . . . . . . . . . . . .
azorim . . . . . . . . . . . . . . . . . 19 . . . . . . . .
caldeircnse . . . . . . . . . . . . . . . . . 19 . . . . . . . .
fagi . . . . . . . . i 7 .2a . . . . . . i 7 . . . . . . .i7
hansi i . 9 . . . . . . . . . . . . . . . . . . . . . . . .
hermione . . . . . . . . 22 . . . . . . . . 22 . . . . . . . 22
m i g w lensi s . . . . . . . . . . . . . . . . . 19 . . . . . . . .
parisatis . 15 . . . . . . . . . . . . . . . . . . . . . . . .
semele . . . . . . . . 22 . . . 17 8 . 22 . 17 . 10 . 17 . . 17 .
statilinus . . . . . . . . 22 . 22 . . . . 22 . 22 22 22 . 6 . . . .
Minois . . . . . . . . . . . . . . . . . . . . . . . . . .
dryas . . . . . . . . 17 . 22 . . . . . 22 22 22 22 . . . 6 . .
nagasauae . . . . . . . . . . . . . . . . . . . . . . . . . .
Yeminois . . . . . . . . . . . . . . . . . . . . . . . . . .
ridingsii . 2 . . . . . . . . . . . . . . . . . . . . . . . .
Oeneis . . . . . . . . . . . . . . . . . . . . . . . . . .
alkrta . . . . . . . . . . . . . . . . -23 . . . . . . . .
asamana . . . . . . . . . . . . . . . . . . . . . . . . . .
bore . . . . . . . . . . . . . . . . . 7 . . . . . . . .
chryxus . 3 . . . . . . . . . . . . . . . . . . . . . . . .
daisetsuzana . . . . . . . . . . . . . . . . . . . . . . . . . .
gl a c i a 1i s . . . . . . . . . . . . . . . . . 22 . . . . . . . .
ivallda . . . . . . . . . . . . . . . . . . . . . . . . . .
jutta . . . . . . .a . . . . . . . . . . . . . . . . . . .
macotmi .29 . . . . . . . . . . . . . . . . . . . . . . . .
melissa .20 . . . . . . . . . . . . . . . . . . . . . . . .
nevadensis . 2 . . . . . . . . . . . . . . . . . . . . . . . .
norna . . . . . . . . . . . . . . . . . . . a . . . . . .
polixenes .5 . . ......................
uhleri .ll . . . . . . . . . . . . . . . . . . . . . . . .
walkyria . . . . . . . . . . . . . . . . . . . . . . . . . .
Pseudochazara . . . . . . . . . . . . . . . . . . . . . . . . . .
hippolyte . 1 . . . . . . . . . . . . . . . . . . . . . . . .
Pseudotergunia . . . . . . . . . . . . . . . . . . . . . . . . . .
fidia . . .21 . . . . . . . . . . . . .21 .21 . . . . . .
Satyrus . . . . . . . . . . . . . . . . . . . . . . . . . .
actaea . . . . . . . .21 .21 . . . . .21212121 . . . . . .
fatua . . . . . . . . . . . . . . . . . . . 14 . . . . . .
ferula . . . - 1 7 . . . . . . . . . . . .17 . . .17 . . . .
hyperantus . . . . . . . . . . . . . . . . . . . 16 . . . . . .
suaha .24 . . . . . . . . . . . . . . . . . . . . . . . .
 TABLE
13. Continued

. . . . . . . . . . . . . . . . . . . . . . . arethusa
. . . . . . . . . . . . . . . . . . . . . . . Berberia
. . . . . . . . . . . . . . . . . . . . . . . abdelkader
. . . . . . . . . . . . . . . . . . . . . . . Brintesia
. . . . . . . . . . . . . . . . . . . . . . . circe
. . . . . . . . . . . . . . . . . . . . . . . Chazara
. . . . . 22 . . . . . . . . . . . . . . . . . briseis
. . . . . . . . . . . . . . . . . . . . . . . prieuri
. . . . . . . . . . . . . . . . . . . . . . . Hipparchia
. . . . . . . . . . . . . . . . . . . . . azorina
. . . . . . . . . . . . . . . . . . . caldeirense
. . . . . . . . . . . . . . . . . . . . . . . fagi
. . . . . . . . . . . . . . . . . . . . . . . hansii
. . . . . . . . . . . . . . . . . . . . . . . herrnione
. . . . . . . . . . . . . . . . . . . . . . . miguelensis
. . . . . . . . . . . . . . . . . . . . . . . parisatis
. . . . . . . . . . . . . . . . . . . . . . . semele
. . . . . . . . . . . . .. . . . . . . . . . statilinus
. . . . . . . . . . . . .. . . . . . . . . . Minois
. 12 22 . . . . . . . . . . . . . . . . . . . . dryas
. . . . . . . . . . . . . . 25 . . . . . . . . nagasauae
. . . . . . . . . . . . .. . . . . . . . . . Neominois
. . . . . . . . . . . . . . . . . . . . . . . ridingsii
. . . . . . . . . . . . . . . . . . . . . . . Oeneis
. . . . . . . . . . . . . . . . . . . . . . . alberta
. . . . . . . . . . . . . . . . . 2 6 . . . asamana
. . . . . . . . . . . . . . . . . . . . . bore
. . . . . . . . . . . . . . . . . . . chryxus
. . . . . . . . . . . . . . . . I 2 . . . . daisetsuzana
. . . . . . . . . . . . . . . . . . . . . glacialis
. . . . . . . . . . . . . . . . . 4 . . . ivallda
. . . . . . . 8 . . . . . . 27 . . 12 . 18 8 j u t t a
. . . . . . . . . . . . . . . . . . macwni
. . . . . . . . . . . . . . . . 2 ? . . . melissa
. . . . . . . . . . . . . . . . . . . . nevadensis
. . . a . . . 8 . . . . . . . . ? . . . norm
. . . . . . . . . . . . . . . . . . polixenes
. . . . . . . . . . . . . . . . . . uhleri
. . . . . . . . . . . . .13. . ualkyria
. . . . . . . . . . . . . . . . Pseudochazara
. . . . . . . . . . . . . . . . . . . . hippolyte
. . . .
. . . . . . . . . . . . . . . . . Pseudotergunia
. . . .
. . . .21 .21 . .
. . . . . . . . fidia
. . . .
. . . . . .
. . . . . . . . . Satyrus
. . . . . . . . . . . . . . . . . . . . . actaea
. . . . . . . . . . . . . . . . . . . . fatua
. . . . . . . . . . . . . . . . . . . . ferula
. . . . . . . . . . . . . . . . . . . . hyperantus
. . . . . . . . . . . . . . . . . . . . suaha
122 P. R . ACKERY
TABLE 14. Hostplants of the Satyrinae (Satyrini; Melanargiina). Forster & Wohlfahrt, 1955';
Fountaine, unpublished2; Higgins & Riley, 19703; Kurentsov, 19704; Manley & Allcard, 19705;
Robert et al., 19836; Schwarz, 1948'

galathea . . . 6 7 . 7 3 . 6 6 . 5 7 . 7 . 6 . 5 . . . .
. .
h a l i d e . . . . . . . . . . . . . . . 4 4 . . . . .
ines . . 6 . . . . . . . . . . . . . . . . . . . . .
occitania. . 2 . . . . . . 6 6 . . . . . . . . . . 6 . 6
russiae . . . . . . . . . . . . 1 5 . . . . . . . . . .

utilized; one species, Oeneis p t t a , supposedly also takes Juncus (Juncaceae). Of

the Palaearctic Melanargiina, represented only by Melanargia, just M . galathea
has abundant data, taking a broad range of grasses (see Table 14). Only
,tl.orcilanica appears remotely comparable to galathea in hostplant range.
Given the proliferation of foodplant records in the subfamily Satyrinae, it is
disappointing that none of the major host groupings-Arecaceae, Poaceae,
Bambuseae, Selaginellaceae or Cyperaceae-characterizes monophyletic
groups, assuming acceptance of current ideas on satyrine classification (Miller,
1968). Generalizing, however, there are some examples of host partitioning,
perhaps most clearly marked in the Neotropics. Here palms seem particularly
characteristic of the Haeterini, the Pronophilina together with Taygetis and
Manataria favour bamboos, while the Euptychiina mainly exploit true grasses
(with the addition of Selaginella). I n the Austro-Orient, three groups especially
focus on true grasses -Mycalesina, Hypocystina and Ypthimina--with Lethe
and Neope favouring bamboos. T h e Arecaceae niche is effectively exploited by
the Elymniina, and Acrophtalmia and Ragadia take Selaginella. In Africa, palms
and bamboos are less important as satyrine hosts. Etymniopsis bammakoo might be
expected to feed on the Arecaceae, and probably only Aphysoneura takes
bamboos. In northern temperate zones satyrines feed almost exclusively on non-
bambusoid grasses and sedges.
Palatability has been demonstrated in certain satyrines, particularly Euptychia
(Brower & Brower, 1964) and Cercyonis (Bowers & Wiernasz, 1979), with
perhaps some evidence of distastefulness in Maniola and Melanargia (Lane,
1957). Spcculating, Wilson (1986) suggests that flavonoids may contribute
toward unpalatability in H. galathea. However, as the archetypal cryptic
butterflies, it is hardly surprizing that satyrines are only rarely involved in
mimicry. Among the most convincing mimics, Elyrnnias species converge on the
patterns of danaines and perhaps Taenaris, and the closely related Elymnzopsis is
obviously involved in the Bamatistes-Acraea association (Eltringham, 1910).
Elsewhere, Zethera species, and probably Penthema, are also remarkably danaine-
like (Vane-Wright & Smiles, 1975); males of Mycalesis drusillodes mimic Telleruo
(see below), while the extraordinary Malagasy endemic Masoura masoura
apparently converges on Mylothris (Pieridae) patterns (R. I. Vane-Wright,
personal communication). Outside these groups, Adams ( 1986) points out
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 123

parallel patterns amongst pronophiline species, favouring Batesian or Miillerian

associations rather than arithmetic mimicry-at present, there is no suggested
chemical basis for these associations.

Calinaginae
The Calinaginae, represented by the Sino-Himalyan genus Calinaga, lack
obvious affinities with any other nymphalid grouping. Although commonly
regarded as monobasic (see Ehrlich, 1958), there are in fact several recognizable
species of which at least four are grossly sympatric in western China (R. I.
Vane-Wright, personal communication).
Ashizawa & Murolya (1967) detail the only known comprehensive life
history-for Calinaga furmusana in Taiwan. The confirmation of cyanogenesis in
Murus species, the larval hosts (see also Uchida, 1985), coupled with the regular
occurrence of saponins, alkaloids and even occasionally tanins, almost certainly
implicates Calinaga species as possible models in mimetic complexes involving
pierids, danaines and zygaenid moths of the genus Agalope. Tremewan (1960)
records Agalupe bifasciata feeding on Aquifoliaceae (Ilex) and Rosaceae (Prunus).
With cyanogenesis widespread in Prunus species, and representatives of the
Zygaenidae known to be cyanogenic (Davis & Nahrstedt, 1979), a further
possible chemical basis for the association is established.

Charaxinae
By downranking Rydon's ( 1971) classification, and incorporating Comstock's
(1961) opinions, we arrive a t six tribes, most of which have characteristic
hostplants: the Pallini (Convolvulaceae) , Euxanthini (Sapindaceae) , Charaxini
(Fabaceae), Anaeini (Euphorbiaceae, Piperaceae, Flacourtiaceae) and
Prothoini (perhaps the Annonaceae) . Only the Preponini lack obviously typical
hosts.

TABLE 15. Hostplants of the Charaxinae (Euxanthini, Pallini). Sevastopulo, 1975', unpublished';
Smiles, 19853;van Sorneren, 19744;van Sorneren & Rogers, 19285; van Son, 1979'j; Vuattoux &
Blandin, 1977'

trajanus . . . . 1 1 . . 5 . . . . . . . .
tiber ius . . . . 1 1 . . . . . . . . . . .
crossleyi . . . 3 1 1 . . . . . . . . . . .
eurinome . . . 3 1 1 . . . . . . . . . . .
uakefieldi . 6 . . 6 2 6 . . . . . . . . . .
Pal La . . . . . . . . . . . . . . . . .
ussheri . . . . . . . . . . 1 . 4 . 1 . .
decius . . . . . . . . . . . . . 7 . . .
violinitens. . . . . . . . . . . . 4 . 1 . 4
124 P. R. ACKERY

Pallini @ Euxanthini
Both monobasic, the Pallini and Euxanthini are confined to the Afrotropical
region. Records for Palla species on the Convolvulaceae appear convincing
(Table 15); others, Toddalia (Rutaceae) and Clerodendrum (Verbenaceae),
probably require confirmation. In Euxanthe, a similarly small genus (Smiles,
1985), the Sapindaceae predominate (Table 15). Further recorded hosts, the
Fabaceae (Afzelia) and Meliaceae (Lovoa) are also quite typically charaxine.

Charaxinz
’Together, the Old-world genera Polyura and Charaxes comprise the Charaxini.
Of the 120 species of Afrotropical Charaxes recognized by van Someren (1975),
five are placed in the subgenus Stonehamia; the rest, divided between 16 species-
groups, form the nominotypical subgenus. T h e 20 or so Indo-Australian species,
however, are not included in this system.
Sapindaceae-feeding (especially on Allophylus species) appears characteristic of
Slonehamia (Table 16). Otherwise only the Melianthaceae (Bersama), closely
related to the Sapindaceae, and Anacardiaceae (Rhus) are recorded.
Representatives of subgenus Charaxes also utilize the Sapindaceae, but more
particularly Phialodiscus and Deinbollia; see also Euxanthe above. Indeed, such is
the bewildering diversity of families exploited by certain Charaxes groups that
few, if any, trends can be confidently viewed as unique.
TO date, no records are available for either the hadrianus or nobilis groups--
otherwise the following outline is in the species-group order proposed by van
Someren (1975). Host records for the nominate species alone represent the
candiope group (Table 16). It commonly exploits the Euphorbiaceae (Croton), a
family also utilized as ‘supplementary’ hosts by many early Charaxes groups; the
only other record, for Pennisetum (Poaceae), at first sight seems unlikely but
monocotyledons often figure among Charaxes hosts.
T h e Fabaceae (especially iifzelia-Caesalpiniodeae) are widely utilized by the
gmihia-group (Table 16); Grzfonia and Albizia alone represent the Mimosoideae
(Darge, 1983). The Poaceae again feature, particularly Oxytenanthern species,
with occasional representation of Arundinaria and Bambusa.
Representatives of the lucretius-group take diverse hosts (Table 16); lucretius is
recorded on the Annonaceae and Linaceae, while the Myrtaceae and
Rhamnaceae include the known hostplants of C. lactetinctus, families that recur
as minor hosts elsewhere in Charaxes. Problems of interpreting such limited
records pale when compared to the discordant data available for the groups
centred on C. jasius and C. tiridates. Records for the jasius-group embrace 20
different families (Table 17)! Three species, jusius, pelias and castor, focus to a
greater or lesser extent on the Fabaceae (mainly the tribes Caesalpinioideae and
Papilionoideae) . Five further species are characterized by the Melianthaceae.
These species again take a range of ‘secondary’ hosts, although perhaps
significantly there is little overlap with the supplementary families utilized by
the ‘Fabaceous’ section (Table 17).
I n diversity the tiridates-group appears marginally less daunting- 1 1 species
on 15 different families of which only six apparently serve as foodplants for more
than one species! T h e Sapindaceae occur most frequently. Elsewhere, records
are shared for the Linaceae, Tiliaceae, Fabaceae (mainly tribes Papilionoideae
and Caesalpinioideae), Ulmaceae and Euphorbiaceae (Table 18).
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 125
TABLE 16. Hostplants of the Charaxinae (Charaxes-in part; subgen. Stonehamia; subgen. Charaxes,
the candiope, cynthia and l u c r e h s groups). Darge, 1983 ; Pennington, 1978*; Pinhey, 194g3;
Sevastopulo, 1975*; van Someren, 19745;van Son, 197g6

varanes . . . . . . . . . . . . . . . . .
fulvescens . . . . . . . . . . . . . . . . .
acuni na tus . . . . . . . . . . . . . . . . .
candiope . . . . . . . . . . . . . . . . .
protoclea . . . . . . . . . 5 2 1 . . . . 5
bueti . . . . . . . . . 2 . . . . . . .
lasti . . . . . . . . . 5 . . 4 4 5 . .
Cynthia . . . 5 . . 1 1 . . . . . . . . .
lucretius . 5 . . . . . . . . . . . . . . .
Iactet inctus . . . . . . . . . . . . . . . . 5

va ranes . . . . . . . 2 2 3 . 2 . . . . . . .
fulvescens . . . . . . . 5 . . . . . . . . . . .
acuninatus . . . . . 5 . 5 . . . . . . . . . . .
candiope . 5 . . . . . . . . . . . . . . . .6
protoc 1ea . . . . . . . . . . . . . . . . .6 .
bouet i . . . . . . . . . . . . . . . 5 1 5 .
lasti . . . . . . . . . . . . . . . . . . .
Cynthia . . . . . . . . . . . . . . . . . . .
lucretius . . . . . . . . . . . . . 4 . . . . .
lactetinctus. . . 4 . . . . . . . . . . . . . . .

*Most of the remaining species-groups exhibit less hostplant diversity, but this
is perhaps a reflection of their size and the paucity of available data. However,
increased emphasis on the Mimosoideae (Fabaceae) is unquestionable, with
widespread records for Acacia, Adenanthera, Albizia, Ambbgonocarpus, Entada, Parkia
and PiPtadenia. Acacia and Entada are noted for C. zoolina (Table 19-the only
species of the zoolina-group with hostplant data). The trend continues with
Albizia species typifying the eupale-group, while C. jahlusa, the only
representative of the jahlusa-group, has been noted on Dalbergia
(Fabaceae-Papilionoideae) and Sapindaceae (Pappea) as well as Acacia.
C. paphianus and pleione, which together comprise the pleione-group, also at
present appear restricted to the fabaceous tribe Mimosoideae (Acacia species).
In exploiting Hugonia (Linaceae), the monobasic zingha-group goes against the
general tenor, but the etesipe-group returns in part at least to the mimosaceous
theme with both penricei and etesipe on Entada (in addition to a range of other
 126 P. R. ACKERY

'TABLE 17. Hostplants of the Charaxinae (Chnruxes-in part; the jnsius group). Darge, 1983';
Fountaine, unpublished2; Gifford, 19653; Haig, 19384; Manley & Allcard, 19705; Ncl, 197g6;
Ouattara et ul., 1977'; Owen & Owen, 1973*; Pennington, 19789; Pinhey, 19491°; Sevastopulo.
1975"; van Someren, 197412;van Son, 197911

a
m
d
Charaxes
I. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
jasius . 2 . 6 . . . . . 9 . . .12 9 1 2 9 . 1 1 4 . 9 . . 4 . . . . . 13 . 5
p e l ia s . . . . . . . . .10 . . . 1 0 1 3 1 0 . . . . . 13 . 9 . 9 . . . . 10 . 9
hansali . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
castor . . . . . . . . . . .ll . . 3 . 9 0 . . 9 . . 0 . . . . . . . . . . . .
brutus . . . . . . .12 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
ansorgei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
phoebus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
pollux . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
druceanus . . . . . . . . . . . . . . . . . . . . . . . . . . . .lo12 . _ . -
eudoxus . . . .ll . . . . . . . . . . . . . . . . . . . . . . . .12 . . . .

Charaxes . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . .
jasius . 12 12 12 . . . . . 9 . . . . . . . . .. . . . . . . . . .712 7 . .
p e l ia s . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . .
hansali . . . . . 1 1 12 . . . . . . . . . . . .. . . . . . . . . . . . . . .
castor . 12 9 1 2 . . . . 9 . . . 9 . . . . . . . . . . . . . . . . .13. .ll
brutus . . . . . . . . . . 12 . . . 12 . 11 . 11 . . . 12 12 9 9 . . . . . . . .
ansorgei . . . . . . . . . . . . . . 12 . . . . . . . . . . . . . . . . .
phoebus . . . . . . . . . . . . . . 12 . . . . . . . . . . . . . . . . . . .
pollux . . . . . . . . . .12 9 ..12 . .12 . . 9 . . . . . . . . . . . . .
druceanus . . . . . . . . . . . . . . 9 . . . . . . . . . . . .ll . . . . . .
eudoxus . . . . . . . . . . . . . . . . . . . . . . . . . . .12 . . . . . .

hosts; Table 19). Contrary to the overriding trend, C. achaemenes exploits

fabaceous genera of tribes Caesalpinioideae and Papilionoideae.
With 42 species, the etheocles-group contains about one-third of Afrotropical
Charaxes, although known larval hosts are restricted to just seven families
(Table 20). Again, the fabaceous tribe Mimosoideae figures prominently, but
additional significant records abound for the Caesalpinioideae, together with
scattered references to the Papilionoideae. T h e Rhamnaceae (Scutia sp.),
previously recorded for only three disparate species, appear as important hosts;
C. karkloof and marieps share records for the Ochnaceae (Ochna species).
Finally, C. nichetes, the sole representative of the nichetes-group, feeds uniquely
on Uapaca (Uapacaceae), and C. laodice, which alone represents the laodice-
group, takes Albizia (Fabaceae-Mimosoideae), Pterocarus, Lonchocarpus
(Fabaceae-Papilionoideae) and Paullinia (Sapindaceae) (Table 20).
Known food plants for the Indo-Australian species remain somewhat sparse.
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 127
TABLE 18. Hostplants of the Charaxini. (Churuxes-in part; the tiridutes group). Darge, 1983';
Haig, 1938*; Henning, 19773; Pennington, 19784; Sevastopulo, 19755; van Someren, 19746; van
Son, 1979'; Vuattoux & Blandin, 19778

violetta . . . . . . . . . . . . . . . . . . . . . 6 . 6 . . .
nunenes . . . 6 . . . . . . . . . . . . . . . . . . . . 6 . .
tiridates . . . 6 . . . 6 . 6 . 6 6 . . 6 . . . 1 . 1 2 . . . .
bipunctatus . . . . . . . . . . . . . . . . . . . . . . . . . . . .
bohemani . . . . . . . . . . . . . . . . . . . . . 6 . 5 . 4 .
smaragdat i s . . . 1 . . . . . . . . . . . . . . . . . . . . . . .
xiphares . 4 . . . . . . . . . . .4 . . . . . . . . . . . . . .
nandina . . . . . . . . . . . . . . . . . . . . . . . . . . .
cithaeron . . . 7 . 6 . . . . . 4 6 4 . . . . 7 4 . 6 . . . . 4
ameliae . . . . . . . . . . . . . . . . . . . . . . . 3 . 3 .
pythodoris . . . . . . . . . . . . . . . . . . . . . . . . . . .

violetta . . . . . . . . . . . . . . . . . . . 6 . . . . .
nunenes . . . . . . . . . . . . . . . . . 5 5 6 5 . 6 . .
tiridates . . . 1 2 . . 5 . . . . . . . . . . 8 . 6 . 6 . .
biprnctatus . . . . . . . . . . . . . . . . . . . .
. . . 6 .
bohemani . . . . 4 4 . . . . . . . . . . . . .
. . . . . 7
smaragdalis . . . . . . . . . . . . . . . . . . . .
. . . . .
xiphares . . . . . . . . . . . . . 4 . 4 . . .
. . . . . .
nandina . . 6 . . . . . . . . . . 6 . . . . .
. . . . . .
cithaeron 4 6 . . . . . . 7 6 4 . . . . . . . . . . . .
. .
ametiae .
. . . . . . . . . . . . . . . . . . . . . . . .
pythodoris.. 6 . . . . . . . . . . . . . . . . . . . . . .

However, records for C. latona and C. bernardus hint at wide ranging hosts
comparable in diversity to those of their African congeners (see Table 21).
In again focusing on the Fabaceae, the Indo-Australian genus Polyura
underlines the dominant theme of the tribe. Of the 26 currently recognized
species (Smiles, 1982), only eight have hostplant data; disappointingly, there is
no obvious correlation with Smiles' proposed cladogram. Although centring on
the Fabaceae, particularly the Mimosoideae and Caesalpinioideae, several other
families are represented in the literature (see Table 21).
Generalizing, hosts for the Charaxini are concentrated in the Rosidae, most
particularly the Fabales, Rhamnales and Sapindales, with additional convincing
records for the closely related Dilleniidae (Malvales, Urticales) . Current data
suggest that the Charaxini take a wider range of hostplants than any other
128 P.R. ACKERY
TABLE 19. Hostplants of the Charaxini (Charaxes- in part; the imlina, eupale, jahlusa, pleznne, zingha
and etesipe groups). Darge, 1983’; Pennington, 19782; Sevastopulo, 19753; van Somcren, 19744;
van Son, 197g5

zoolina . . . . 2 . 2 . . . . . . . . . . . . . . . . . . . . . .
eupale . . . . . 4 . . . . . . . . . . . . . . . . . . 4 . . . .
subornatus. . . . . 4 . . . . . . . . . . . . . . . . . . . . . . .
dilutus . . . . . 4 . . . . . . . . . . . . . . . . . . . . . . .
jahlusa . . . . 4 . . . . . . . . . 5 . . . . . . . . . . . 2 . .
pleione . . . . 4 . . . . . . . . . . . . . . . . . . . . . . . .
Daphianus.. . . 4 . . . . . . . . .’. . . . . . . . . . . . . . .
zingha . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
etesipe . 1 . . . . 5 . 5 . . 3 . . 3 . . . . 3 3 5 3 . . . . . .
penricei . . . . . . 4 . . . . . . . . . 2 . . . . . . . . . . . .
achaemenes.. . . . . . . . 5 4 . 2 . 2 2 . 2 . . . . . . . . . . .

nymphalid group. With the Fabaceae typically utilized, the absence of

leguminous records in Charaxes (subgenus Stonehamia) is surprising. Elsewhere, in
the perhaps polyphyletic or paraphyletic nominate subgenus, hostplant diversity
is most notable in thejasius and tiridates groups. With the criteria adopted by
van Someren in establishing his species-groups remaining unclear, the range of
hostplants adopted by these groups may merely reflect their ‘unnaturalness’.
However, the genus Polyura, apparently monophyletic, exhibits comparable
diversity with no obvious pattern. Overall, in Charaxes itself, the distribution of
Mimosoideae records provides the most obvious trend; widely utilized
elsewhere, they are hardly noted in the candiope-, cynthia-, lucretius- and jusius-
groups. Their utilization in only part on the tiridates-group perhaps casts further
doubt on the status of this grouping. Finally, records for the monocotyledonous
Poaceae are surprising but occur often enough to appear authentic.

Anaeini
Rydon (1971) and Comstock (1961) present contrasting treatments of the
New-World Anaeini. Comstock subsumes many conventionally recognized
genera within Anaeu, while Rydon raises most to tribal level! However, there is
broad agreement on the composition of groups: the differences mainly concern
rankings. Compromising, two subtribes may be distinguished: the Anaeina
(Hypna, Anaea, Polygrapha, Consul, Cymatogramma, Fountainea and Memphis) , and
the Zaretina (zaretis, Siderone and Coenophlebia) .
To date, records suggest that Hypna, Anaea and Polygrapha species are largely
restricted to the Euphorbiaceae (mainly Croton) . Although centred on Crotun,
both Fountainea and Memphis comparatively show a greater range (see Table 22),
with three diverse Memphis species on Piper (Piperaceae) alone, an occasional
host elsewhere in the genus. Memphis moruus is the only anaeine species with a
wide range of known hosts; in addition to the predictable Croton and P$er,
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 129

s!u!ll"ad.. .............................
esa,spu!daS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
sJsdsn . . . . . . . . . . . . . . . . . . . . . . . . . . . . . m
sea,e~odpn' . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
s!qn,S . . . . . . . . . m . . . . m In. .m .m . . . . . . . . .
aeaJsuurayTI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
enual&ty . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Q

asa,e,qss,a3 ..............................
end,sso,ald ...............................
s!r.+all!yy . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Q

gndre,oq,"oi ...............................
s!8,aqlea . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ln Ln b

s,!p"v.. . . . . . . . . . . . . . . . . . . . . . . . . . .
. b

aeap!ouo!l!dad- . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
snpu!,eurej . . . . . . . . . . . . . . . . . . . .* . . . . . . .
Ln yI

wn!qolo,aqd . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
In
wnroydol,ad . . . . . . . . . . . . . . . . . . . . . . . . . . . . pq Q

uo,aauo,aty . C . . . . . . . . . . . . . . . . . . . . . . . . . . . .
s!p,lsu,aqlny . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . - .* . . . . . .
yI

e!uo,J!13
wnalqdojy.+Q'. . . . . . . . . . . . m . . . . . . . . . . . . . . . .
wnpdaso*dfiac . . . . . . . . . . . . . . . . . . . . . . . . . . N . .

q!u!dpsas3 . . m . . . . . . . . . . . . . . . . . . . . . . . . . . .
e!~a)gXqsa,a . . . m . . . . . . . . . . . .m . . . . . . . . . . . .
VI

*!la,,Y.. . . . . . . . . . . . . . . . . . . . . . . . .
. h . .

aeap!o!u!dlesas3- . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
B!uapr.+d!d . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
VI

s ! ~ ~ s d .. . . . . . . . . . . . . . . . . . . . . . . . . . .
. m
..................... . . . . . . . .
Ln

snd,o,ouosllqwv . . . . . . . . . . . .m . . . . . . . . . . . . . . . .
*.I

e!l!q,V.. . . . .
r m N L n . * m * M . . .
. 4 L n h .. . r N

s,aqqua"apv. . . . . . . . . . . . . . . . . . . . . . . . . . . .
. Y )

s!JaJy . m m . . . . . . . . . . . . . . . m . . m . . . . . .
.In
aeap!ogomlty-. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
aaa,sqed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
so,r(dso!a.. . . . . . .*) . . . . . . . . . . . . . . . . . . . . .
aea,suaqa., ............................
'!Ha3.. . . . . . . . . . . . . . . . . . . . . . . . .
. I n . .
asa,swln.. . . . . . . . . . . . . . . . . . . . . . . . . . . . .
suq,o. . . . . ..om . . . . . . . . . . . . . . . . . . . . . .
aea,euq,o.. . . . . . . . . . . . . . . . . . . . . . . . . . . . .
euouuv . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r

aea,suouuv . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
V)
130 P. R. ACKERY

'I'ABLE21. Hostplants of the Charaxini (Oriental Charaxes and Polyura). Bascombe et al., 1987';
Carey Hughes & Pickford, 19772; Common & Waterhouse, 19813; Corbet & Pendlcbury, 1978+;
D'Abrera, 19775;Fountaine, unpublished6; Hill et al., 1978'; Parsons, in prep.R;Pholboon, 1965';
Sevastopulo, 1973'O; Smiles, 1982"; Uchida, 198412; Wood, 198615; Wynter Blyth, 195714

I
(Y

z
0

Charaxes
4. . . . . . . . . . . . . . . . . . . . . . . . . . .
bernardus .1 0 . 2 . 2 . . . . . . . . . . . . 4 4 . . . . . . .
latona . . . . 13 8 . . . . . . . . . . . . . . . . . . . . .
marmax . . . . . . . . . . . . . . . . . . . . . . . . . . .
psaphon . 14 . . . . . . . . . . . . . . . . . . . . . . . . .
solon . . . . . . . . . . . . . . . . . . . . . . . . - .
Polyura . . . . . . . . . . . . . . . . . . . . . . . . . . . .
jupiter . . . . . . . 1 1 . . . . . . . . . . .ll . . . 8 . . . .
satto . . . . . . . . . . . . . . . . . . . . . . . . . . .
setrpronius . . . . . . . . . . . l l . l l . . 311 . l l . . . 1 1 1 1 . 11 .
hebe . . . . . . . . . . . . . . . . .llll . . . . . . . .
athamus . . . . . . . . - 1 1 . . . . . . 11 11 11 11 11 9 . 11 . . . 1
schreiber . . . . . . . . . . . . . . . . . . 11 . . . . . .ll . .
eudamippus . . . . . . . . . . . . . 11 . . . . . . . . . . . . .
narcaea . . . . . . . . . . . . . . . . . . . . . . . . . . .

bernardus . 1 . . . . . . . . . . . . . . . . . . . . 10 . . 7
Mona . . . . 8 . 8 . . . . . . . . . . . . . . . 0 8 . .
marmax . . . . . . . . . . . . 6 . . . . . . . . . . . . .
psaphon .14 . . . . . . . . . . . . . . . . . . . -14 . . .
solon .lo10 . . . . . . . . . . . . . . . . . . . . . . .
Polyura . . . . . . . . . . . . . . . . . . . . . . . . . .
jupiter . . . . . . . . . . . . . . . . . . . . . . . . . .
satco 1 1 . . .. . . . . . . . . . . . . . . . . . . . . .
sempronius . . . . . . . 11 . 11 . . . . . . . . . . 3 . . . . .
hete . . . . . . . . . . . . . . . . . . . . . . . . . .
atharms 1 1 . . .. . . . . . . . . . . . . . . . . . . . . .
schreiter . . 1'1 . . . . . . . . . . . . . 11 . 11 . . . . . . .
eudamippus . . . . . 12 . . . . . . . . 1 1 . . . . . . . . . . .
narcaea . . . . . . . . . . .12 . . . . . . . . . . . . . .

members of the Lauraceae are also widely utilized. Consul is notable for lacking
euphorbiaceous records; instead the Piperaceae predominate, as in some
Memphis species. Cymatogramma, the sole remaining genus of the Anaeina, at
present lacks biological data.
The Zaretina (Siderone and Xaretis-Coenophlebia lacks records) focus on the
Flacourtiaceae (mainly Casearia species). With most exceptions unconfirmed,
the generalization of flacourtiaceous Zaretina and euphorbiaceouslpiperaceous
Anaeina remains supportable (Table 22). Perhaps the records of Memphis on the
Piperaceae, otherwise only widely noted for the closely-related Consul, raise
questions concerning the 'naturalness' of some genera. Overall, the major hosts
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 131

TABLE 22. Hostplants of the Charaxinae (Anaeini). Barcant, 1970l; d'Araujo e Silva et al., 19682;
DeVries, 1986'; Fountaine, unpublished4; Hayward, 196g5; Howe, 19756; Miiller, 188fi7;
Muyshondt, 1973P; 1974a9, blO; 1975a", c L 2 ;1976bI3;Muyshondt et al., 197614;Riley, 197515;
Ross, 197616

marthesia . . . . . . . . . .13 . . 3 . . . . .
ga Lanthis . . . . . . . . . . 15 . . . . . . . .
Zaretis . . . . . . . . . . . . . . . . . . .
itys . . . . . . . 3 . . 0 3 3 . . . . . .
cat L idryas . . . . . . . . . . 13 . . . . . . . .
Hypna . . . . . . . . . . . . . . . . . . .
clytennestra . . . . . . . . . . . . . . . . 2 . .
Anaea . . . . . . . . . . . . . . . . . . .
aidea . . . . . . . . . . . . . . .316..
cubana . . . . . . . . . . . . . . . . . . 15
floridalis . . . . . . . . . . . . . . . . 6 . .
andria . . . . . . . . . . . . . . . . 6 . .
Polygrapha . . . . . . . . . . . . . . . . . . .
suprema . . . . . . . . . . . . . . . . 2 . .
Consul . . . . . . . . . . . . . . . . . . .
fabius . . . . . . . 9 4 . . . . . . . . . .
jansoni . . . . . . . 3 . . . . . . . . . . .
electra . . . . . .13 . . . . . . . . . . . .
F w n t a inea . . . . . . . . . . . . . . . . . . .
eurypyle . . . . . . . . . . 1 . . . . . 10 . .
ryphea . . . . . . . . . . . . . . . .2 . .
glyceriun . . . . . . . . . . . . . . . . 3 . .
cratias . . . . . . . 5 . . . . . . . . 5 . .
Menphis . . . . . . . . . . . . . . . . . . .
nobilis . . . . . . . . . . . . . . . . 14 . .
vert icordi a . . . . . . . . . . . . . . . . 15 . .
artacaena . . . . . . . . . . . . . . . . 3 . .
pithyusa . . . . . . . . . . . . . . . . 12 . .
arginussa . . . . . . . . . . . . . . . .3 . .
forreri . . . . 3 . . . . . . . . . . . . . .
clemstra . . . . . . . 3 . . . . . . . . . . .
mrvus . 7 7 1 1 5 . . 3 . . . . . . . . 5 . .
oenomais . . . . . . . . . . . . . . . . 3 . .
Leonida . . . . . . . 4 . . . . . . . . . . .
otrere . . . . . . . . . . . . . . . . 2 . .
beatrix . . . . . . . 3 . . . . . . . . . . .

of the Anaeini, the Piperaceae (Piperales, Magnoliidae), Flacourtiaceae

(Violales, Dilleniidae) and Euphorbiaceae (Euphorbiales, Rosidae) are widely
diverse; the affinities of the Lauraceae (Laurales, Magnoliidae), convincing
'secondary' hosts for Memphis morvus, remain doubtful but some authorities
suggest weak association with the Piperaceae.

Preponini
Rydon (1971) recognizes five genera of Preponini, the familar Agrias,
Archaeoprepona and Prepona (see Table 23), and the rather obscure Anaeomorpha
132 P. R. ACKERY
TABLE 23. Hostplants of the Charaxinae (Preponini, Prothoini). Barcant, 1970’; Barselou, 1983’;
Corbet & Pendlebury, 197g3; d’Araujo e Silva et al., 19€18~;
DeVries, 1980b5, 19866; Fountaine,
unpublished’; Fruhstorfer, 191G8; Hayward, 19Gg9; Muyshondt, 1973ei0, 1976a”, b l * ;Janzrn,
personal communication‘

cha Ic iope . . . . . . . . 4 . . . . . . . . . . . . . . . . . . .
demophon . 8 . . 1 . 7 . . 6 6 9 . . . . . . . . . . . . . . .ll
meander . . . . . . . . .12 . 6 6 .
. . . . . . . . . . . . .
Prepona . . . . . . . . . . . . . . . . . . . . . . . . . . .
Laertes . . . . . . . . . . . . . . . .1313 . 4 . . . 1 . . .
ompha Ie . . . . . . . . . . . . . . . . . . 10 6 . . . . . . .
Agrias . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
amydon . . . . . . . . . . . . . . . . . . . . . . . . . . 5 . .
c laudi a . . . . . . . . . . . . . .2 . . . . . . . 4 . . . . . .
Pro t hoe . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
f ranck . . 3 . . . . . . . . . . . . . . . . . . . . . . . . . .

and Noreppa which lack foodplant records. Archaeoprepona species typically exploit
the Lauraceae together with the Monimiaceae and Siparunaceae, three
intimately related families. Prepona species are quite characteristically charaxine,
utilizing members of the Fabaceae and Sapindaceae, with the addition of
Hirtella and Licania (Chrysobalanaceae) . Only DeVries’ ( 1980b) record of Agrias
amydon on Erythroxylum (Erythroxylaceae) appears authentic for Agrias; elsewhere
the Quiinaceae (Quizna) and Myrtaceae are implied. These families lack obvious
affinities, either with each other or with those favoured by Prepona and
A rchaeoprepona.

Prohoini
The Prothoini conclude the charaxine section. Rydon (1971) recognizes two
genera, Prothoe and Agalasa. The group is represented by a single record-Prothoe
franck on Freisodielsia (Annonaceae) (Table 23). Members of this family recur as
minor hosts of Charaxes and Prepona.
Generally, the wing patterns of charaxines seldom converge on those of the
more ‘traditional’ distasteful groups. A few show not altogether convincing
similarites to the danaines (Euxanthe, see Smiles, 1985), or the
silvaniform-heliconiine/tiger-stripedithomiine complex (Consul species). Sur-
prising pattern parallelisms between Agrias and Asterope (Callithea auctt., see
Descimon, 1977) have no known chemical basis. Charaxes acraeoides appears to
converge on acraeine patterns.
Large and small Charaxes species frequently show pattern convergences.
Swynnerton ( 1926), investigating the possible defence strategies of Charaxes
species, found that they were protected from a number of bird predators. His
conclusions can probably be regarded as typical of the entire subfamily. When
flying, larger Charaxes seem difficult for birds to seize and reduce to edible
fragments. Swynnerton concluded that bird predators did not associate the
difficulties of making a successful ‘kill’ with the size of their prey, but rather with
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 133

TABLE24. Hostplants of the Acraeinae (Purdopsis, Bemutistes, Acraea, in part). Common &
Waterhouse, 1972l; Fountaine, unpublished2; Gifford, 19653; Owen & Owen, 19724; Paulian,
19565; Pennington, 19786; Pfitzer & Fargher, 1976'; Pierre, 197g8; Pierre & Vuattoux, 19789;
Pinhey, 1949'O; Sevastopulo, 1973", 197512;van Someren, 197413;Wynter Blyth, 1957"

prnctatissima.. . . . . .6 . . . . . . . . . . . . . . . . . . . . . . . . . .
Acraea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
cerasa . . . . . . . . 13 . . . . . 13 . . . . . . . . . . . . . . . . . . .
rogersi . . . . . . . . . . . . . . . . . . 4' . . . . . . . . . . . . . . .
petraea . . . . . . . . . . . . . . . . 6 . . . . . . . . . . . . . . . . .
aMera . . . . . . . . . . 9 . . . . . . . . . . . . . . . . . . . . . . .
violarun . . . . . . . . . . . . . . . . . . . . . . . 6 . . . . . . . . . .
asema . . . . . . . . . . . . . . . . . . . . . . . . . 3 . . . . . . . .
nohara . . . . . . . . . . . . . . . . . . . . . . . 6 . 6 . . . . . . . .
egina . . . . . . . . . . . . . . 2 . . . 4 . . . . . . . . . . . . . . .
acrita
. . . . . . . . . . . . . . . . . . 13 . . 12 . . . . . . . . . . . .
aglaonice . . . . . . . . . . . . . . . . . . . . -10 . . . . . . . . . . . .
braesia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
doubledayi . . . . . . . . . . . . . . . . . .13.. . . . . 2 . . . . . . . .
sykesi . . . . . . . . . . . . . . . . . . 13 . . . . . . . . . . . . . . .
Oncaea . . . . . . . . . . . . . 6 . . . . 1 3 3 . . . . . 6 . 13 . . . . . .
e q u a t o r i a l i s . . . . . 13 . . . . . . . . . . . . . . . 12 . . . . . . . . . . . .
caldarena . . . . . . . . . . . . . . . . . -13 . . . . . . 6 . . . . . . . .
caecilia . . . . . . . . . . . . . . . . . -13 . . . . . . . . . . . . . . .
natalica . . . . . . . . . . . . . . . . . . 6 . . 6 . . . 6 . . . . . . . .
pseudegina . . . . . . . . . . . . . . . . . . . . . 4 . . . . . . . . . . . .
asboloplintha . . . . . . . . . . . . . . . . . . 13 . . 12 . 13 . . . 12 . . . . . .
zetes . . . . . . . . . . . I 2 . . . 4 . - 1 3 . . 41213 . . . . . . .12 . .
a m s a . . . . . . . . . . . . . . . . . . . 6 . . . . . . . 12 . . . . . .
rabbiae . . . . . . . . . . . . . . . . . . 12 . . . . 13 . . . . . . . . . .
ranavalona . . . . . . . . . . . . . . . . . . . . . 2 . . . . . . . . . . . .
qui r i na . . . . . . . . 13 . . . . . . . . . . . . . . . . . . . . . . . . .
igati . . . . . . . . . . . . . . . . . . 5 . . . . . . . . . . . . . . .
horta . . . . . . . . . . . . 6 . . . . . . . . 6 6 . . . . . . . . . . .
andrmche . . . . . . . 7 . . . . . . . . . . 1 . . 1 . . . . . . . . . . . .
neohle . . . . . . .9 . . . . . . . . . .6 . . 6 . 1 3 . . . . . . . . . .
violae . . . .14 . . . . . . . . . . . . . - 1 1 -14 . . . . . . . . . . . .
camaena . . . . . . . . . . . . . . . 9 . . . . . . . . . . . . . 4 . . . .
insignis . 12 . 12 . . . . . . . . . . . . . . 12 . . . . . . . . 13 . . . . . .
endoscota . . . . . . . .8 . . . . . . . . . . . . . . . . . . . . . . . . .
leucographa . . . . . . . . 8 . . . . . . . . . . . . . . . . . . . . . . . . .
ahatha . . . . . . . . 0 . . . . . . . . . . . . . . . . . . . . . . . . .
BematiStes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
macaria . . . . . . . . . . . . . . . . . . 4 . . . . . . . . . . . . . . .
alcinoe . . . . . . . . . . . . . . . . . .4 . . . . . . . . . . . . . . .
aganice . . . . . . . . . . . . . . . . . . 13.66.13 . . . . . . . . . .
quadricolor . . . . . . . . . . . . . . . . . -13 . . . . 2 . . . . . . . . . .
poggei . . . . . . . . . . . . . . . . . . 12 . . . . . . . . 1 2 . . . . . .
tellus . . . . . . . . . . . . . . . . . . 13 . . . . . . . . 12 . . . . . .
epaea . . . . . . . . . . . . . . . . . . 4 . . . . . . . . . . . . . . .
I34 P. R. ACKERY

TABLE
25. Hostplants of the Acraeinae (Acraea, in part). Birket Smith, 1960'; Eltringham, 1912';
Fountaine, unpublishedB; Gifford, 1965+;Guilbot & Pierre, 19785; Muroya el al., 19676;Owen &
Owen, 19727; Paulian, 19568; Pennington, 1978y; Pierre, 1981a1", c' I ; Pierre & Vuattoux,
197812; Pinhey, 194913; Sevastopulo, 197514; ShirBzu, 196015 ; Uchida, 1984Ifi; van Somcrrn,
197417; Wiley & Hudson, 194318; Wynter Blyth, 1957lY

anacreon . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 4 .
vesta . . . . . . . . . . . . . . . . . . . 6 6 . . . 1 6 .15 . . . .
rahira . 9 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
pentapolis . . . . . . . . . . . . . . . . . . .14 . .14 14 . . . . . . .
vespera I i s . . . . . . . . . . . . . . . . . . . . . . . 14 . . . . . . .
orest ia . . . . . . . . . . . . . . . . . . . . . 1 4 . . . . . . . . .
obei r a . . . . . . . . a . . . . . . . . . . . . 9 . . 9 9 . . . . .
eponi na . . . . . . 9.a9127.14a . . . . . . . . . . . . . . . .
rangatana . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
ventura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
acerata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
oberthuri . . . 2 . . . . . . . . . . . . . . . . . . . . . . . . . . .
uvui . . . . . 1717 . . . . . . . . . . . . . . . . . . . . . . . .
bonasia . . . . 7 .17. . . . . . l a . . . . . . . . . . . . . . . . .
cabira . . . . . . 9 . . 9 . . .la.. . . . . . . . . . . . . . . .
soti kensis . . . . . . 14 . . . . . . . . . . . . . . . . . . . . . . . .
excelsior . . . . . . 17 . . . . . . . . . . . . . . . . . . . . . . . .
encedana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
encoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
emedon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
ansorgei . . . . . . . . . . . . . . . . . . . . . . . . . 1 4 . . . . .
disjuncta . . . . . . . . . . . . . . . . . . . . . . . . .14 . . . . .
alcippoides . . . . . . . . . . . . . . . . . . . . . . . . . 17 . . . . .
alciope . . . . . . . . . . . . . . . . . . . . .1211 . . . . . . . .
aurivillii . . . . . . . . . . . . . . . . . . . . .ll . . .ll . . . . .
jodutta . . . . . . . . . . . . . . . . . . .14 . . . .14 . . . . . .
esebria . . . . . . . . . . . . . . . . . . . 14 . 9 . . 17 9 . . . . .
Lycoa . . . . . . . . . . . . . . . . . . . . . 12 . . ? . . . . . .
johnstoni . . . . . . . . . . . . . . . . . . . . . 14 . . 17 . . . . . .
pharsalus . . . . . . . . . . . . . . . .17 . . . . . . . . . . . .
pene 1eos . . . . . . . . . . . . . . . . . . . . . . . . 14 . . . . .
quirinalis . . . . . . . . . . . . . . . . . . . . . . . . 17 . . . . .
parrhasia . . . . . . . . . . . . . . . . . . . . . . . . ? . . . . .
circeis . . . . . . . . . . . . . . . . . . . . . . . . ? . . . . .
oreas . . . . . . . . . . . . . . . .14 . . . . . . .14 . . . . .
amicitae . . . . . . . . . . . . . . . . . . . . . . . . 17 . . . . .
igola . . . . . . . . . . . . . . . . . . . . . . . . 9 . . . . .
penelope . . . . . . . . . . . . . . . . . . . . . . . . 14 . . . . .
mlanoxatha . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
perenna . . . . . . . . . . . . . . . . . . . . . . . . . . . .14 . .

the 'kind'. When presented with a smaller butterfly of the same 'kind', the birds
seemed to 'expect' the same difficulties experienced with a large one.
Swynnerton's ideas have not been developed further for butterflies but
Hespenheide ( 1973) suggests a similar association between various Diptera and
Coleoptera, birds and other visually hunting predators having difficulty in
capturing flies of the size and type mimicked by the beetles.
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 135

TABLE
25. Continued

. 4 . 17 9 . 9 1 4 . . . . . . . . . . . .
. . . . . . . . . . anacreon
. . . . . . . . . . . . . .19 . . .
. . . . . . . . . . . vesta
. . . . . . . . . . . . . . . . . . .
. . . . .17 . . . . . rahira
. . . . . . . . . . . . . . . . . .
. . . . . . . . . . . pentapolis
. . . . . . . . . . . . . . . . . .
. . . . . . . . . . . vesperal i s
. . . . . . . . . . . . . . . . . .
. . . . . . . . . . . orestia
. . . . . . . . . . . . . . . . . .
. . . . . . . . . . . obeira
. . . . . . . . . . . . . . . . .14 . . . . . . . . . . . eponina
. . . 17 . . . . . . 17 17 . . . . . . . . . . . . . . . . . rangatana
. . . . . . - 1 4 . ..................... ventura
. . . . . . . . . . . . . . . . . 9 . . 14 2 1 2 . . 1 . . . 14 acerata
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . oberthuri
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . wui
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . bonasia
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cabira
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . sot ikensi s
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . excelsior
. . . . . . . . 5 . . . . . . . . . . . . . . . . . . . . . encedana
. . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0 . . encoda
. . . . . . . . . . . . . . . . . . . . . . . . . . . 5 . . encedon
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ansorgei
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . disjuncta
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a 1c ippoides
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . alciope
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . aurivillii
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . jodutta
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . esebria
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lycoa
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . johnstoni
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pharsalus
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . peneleos
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . quirinalis
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . parrhasia
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . circeis
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .oreas
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . amicitae
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . igola
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .penelope
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . nelanoxatha
. . . . . . . . . . . . . 1 4 . . . . . . . . . . . . . . . . perema

Acraeinae
Pierre (1984) presents a cladistic analysis of the Acraeinae but, as he
incorporates hostplants as characters, the obvious patterns must be treated with
caution. Nevertheless his results appear supportable even without the inclusion
of hostplant data. Conventionally, the Acraeinae comprise Acraea, Actinote,
136 P R.ACKERY

Bematisles and Pardopsis. Pierre demonstrates the monophyly of Acraea +

Bematzstes + Actinote although Acraea in its current sense is paraphyletic. A basal
dichotomy divides the subfamily in two: a paraphyletic section of Acraea +
Bematistes, and the remainder of Acraea (also paraphyletic) +
Actinote. If Pierre’s
conclusions are accepted, the minimum course would be to subsume Bematistes
within Acraea (in its now restricted sense), while including under Actinote several
conventional ‘Acraeas’. T h e position of Pardopsis remains ambiguous-it seems
to have been treated as an outgroup in Pierre’s analysis. However, for present
purposes, the four widely recognized genera are retained.
T h e subfamily divides into two groups each broadly characterized by
hostplant preferences, a ‘Passifloraceous’ group, and a less firmly defined
‘Urticaceous’ group, including clear Asteraceae and Tiliaceae themes. Although
some records suggest quite a wide range of hosts, no single species feeds
interchangeably between the Urticaceae and Passifloraceae. Passifloraceous
species also utilize other families (Table 24), notably the Turneraceae, a family
with close affinities with the Passifloraceae, and the Vitidaceae. Elsewhere,
ccattered records represent the Flacourtiaceae and Violaceae, intimately related
to the Passifloraceae and Vitidaceae within the Violales, together with the
Acanthaceae, Malvaceae, Theaceae, Asteraceae and Verbenaceae. Bematistes
species, firmly grouped within this section of Acraea, take a similar range of
Passifloras. Two species, B. poggei and B. tellus, are also recorded on Vitis,
reflecting the Passifloraceae-Vitidaceae association found in Acraea.
Most strictly ‘Urticaceous’ species confine themselves to the Urticaceae alone.
Others, although firmly assigned to this subsection by morphological characters,
feed on various groups (see Table 25). In particular, a well-marked subgroup
focusses on the Tiliaceae, Sterculiaceae and Malvaceae, three families closely
associated within the Malvales. Actinote species, the only Neotropical acraeines,
although clearly placed within the urticaceous section, feed largely on the

TABLE
26. Hostplants of the Acraeinae (Aclinote). Barcant, 1970’; d’Araujo e Silva el al., 1968’;
DeVries, 19863; Hayward, 196g4;Jordan, 19135; Miiller, 18866; Ross, 1976’

alalia . . . . . . . . . . 5 . . . . . .
anteas . . . . . . . . . . . . . 6 . . .
equatoria . . . . . . .
. .. 4 . . . . . .
genet r i x. . . . . . .
. .. . . . 2 . . .
lapitha . . . . . . .
. .. . . . 3 . . .
leucomelas. . . . . . .
. .. . . 7 3 . . .
mamita . 4 . . . . .
. 2 . . . . . . . 2
parapheles . . . 5 . . .
. . . . . . 2 . . .
pellenea . . . . . 4 4 . 2 . 1 4 . 4 4 . .
perisa . . . . . . . . . . 4 . . . 4 . .
pyrrha . . . . . 2 2 . . . 4 . . 4 . . .
rhodope . . . . . . . . . . . . . 2 . . .
thalia . . . . . . . . 2 . . . . 2 . . .
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 137

Asteraceae (Table 26)-heliconiines are the principal butterfly herbivores on

the Passifloraceae in the Neotropics.
Although traditionally regarded as models in mimetic complexes (see
Eltringham, 19lo), the chemical basis for acraeine unpalatability remains
poorly understood. Speculating, Rothschild ( 1971) implicated hydrogen
cyanide-the Passifloraceae include cyanogenic species, and Owen ( 1970) found
that the thoracic glands of ‘A. encedon’ give off HCN when exposed to air.
Feeding experiments (Swynnerton, 1915a, b; 1919) confirm the relative
unacceptability of acraeines to various potential predators.
Nahrstedt & Davis (1981, 1983) confirmed cyanogenesis in the Acraeinae,
but favoured synthesis of glucosides by the butterflies themselves. Glucosides
present in Acruea differ from those known in the Passifloraceae, the family
assumed by Nahrstedt & Davis to include the hosts of the tested species.
However, the species seemingly containing the highest concentration of cyanide,
‘A. encedon’ and A. eponina, apparently never feed on Passifloras-eponina is
largely associated with the Tiliaceae (with additional records for the
Sterculiaceae, Malvaceae and Solanaceae), while ‘encedon’ feeds on’
Commelinaceae (with the Fabaceae also being recorded-while the individuals
tested no doubt belonged to the encedon-complex, their precise identity must be
open to doubt). Cyanogenesis has been confirmed in several species of both the
Tiliaceae and Commelinaceae, although many species have also given negative
results (see Darnley Gibbs, 1974). No ‘Urticaceous’ acraeine was tested.
I n hostplant preferences, Actinote species are very much isolated from other
acraeines. I n view of current investigations into the efficacy of pyrrolizidine
alkaloids with respect to distastefulness in danaines (Conner, Eisner, Van der
Meer, Guerro & Meinwald, 1981; Eisner, 1980), the widespread exploitation of
composites is of particular interest. Species of the genera Senecio, Mikania and
Eupatorium, all foodplants of Actinote species, are known to be effective danaine
attractants (Ackery & Vane-Wright, 1984)-the presence of pyrrolizidine
alkaloids has been demonstrated in Eupatorium (Dominguez, 1977) and Senecio
(Robins, 1971).
Heliconiinae
Michener’s (1942) study provides the orthodox generic classification. It has
recently been suggested that this almost exclusively Neotropical group (there is
also weak Nearctic representation) should include the Old-world passi-
floraceous ‘nymphalid’ Cethosia, and perhaps Vindula (Brown, 1981). A further
modification, the separation of Eueides from Heliconius (sensu Michener) is
probably justified (Brown & Mielke, 1972); more contentious is the recognition
of Neruda for the H. aoede-group and the resurrection of the archaic name
Laparus for H. doris (Brown, 1981).
Nearly all heliconiines feed on the Passifloraceae, usually PassiJlora but
occasionally also Mitostemma, Dilkea and Tetrastylis species (see Benson et al.,
1976). Agraulis vanillae is said to also exploit members of the Caprifoliaceae
(Biezanko, 1949) and Tiliaceae (Brown & Heineman, 1972), the latter reflecting
the Tiliaceae-Passifloraceae association in Acraea, while Eueides procula has only
been recorded on Erblichia (Turneraceae) (Janzen, 1983).
As tabulated and discussed by Benson et at. (1976), the hostplant associations
conform to a ‘co-evolutionary’ pattern. The most primitive representatives of
138 P. R. ACKERY

TABLE27. Hostplants of the Argynninae (in part). Brooks & Knight, 1982I; Common &
Waterhouse, 1972*; Corbet & Pendlebury, 19783; D’Abrera, 19774; Ehrlich & Ehrlich, I961 5 ;
Ferris & Brown, I981 6 ; Forster & Wohlfahrt, 19557; Fountaine, unpublished8; Friedrich, 1978g;
Gullander, 1959”; Henriksen and Kreutzer, 1982”; Higgins and Riley, 1970i2;Hill et al.,
1978”; Hoffman et al., 193814; Howe, 197515; Iwase, 195416; Kim, 198417;Kurcntsov, 197018;
Lambkin & Lambkin, 1977”; Langer, 195820; Leech, 1892”; Manley & Allcard, 19702*;
Niculescu, 196523;Perkins & Meyer, 197324;Pyle, 197425;Rancourt, 197gZ6;Schwarz, 194gY7;
Sevastopulo, 197328;Tietz, 19722g;Uchida, 198430;Wiltshire, 19573

anadyomene . . . . . .17. . . . . . . . . . . . . . . . . . .
paphia . . . . 23 . 16 . . . . . . . . . . 232327 . . . . . .
Argyreus . . . . . . . . . . . . . . . . . . . . . . . . . .
hyperbius . . . . . .19. . . . . . . . . . . . . . . . . . .
Argyr- . . . . . . . . . . . . . . . . . . . . . . . . . .
ladice . . . . . .16.. . . . . . . . . . . . . . . . . .
ruslana . . . . . -17.. . . . . . . . . . . . . . . . . .
Brenthis . . . . . . . . . . . . . . . . . . . . . . . . . .
daphne . . . . . -22 . . . . . . . . . . . . . . . . . . .
hecate . . . . . . . . . . . . . . . . . . . . . . . . . .
ino . . . . 21 . . . . . . . . . . . . . . . . . . . . .
BoL or ia . . . . . . . . . . . . . . . . . . . . . . . . . .
aquitonaris. . 7 . . . 7 . . . . . . . . . . . . . . .11 . 7 .
napaea . 612 . . .ll . . . . . . . . . . . . . . . . . 5 .
pales . . . . . .22 . . . . . . . . - 2 3 . . . . . . . . .
sifanica . . 20 . . . 10 . . . . . . . . . . . . . . . . . 20 .
Cirrochroa . . . . . . . . . . . . . . . . . . . . . . . . . .
regina . . . . . . . . 4 . . . . . . . . . . . . . . . . .
thais . . . . . . . . . . 28 . . . . . . . . . . . . . . .
Ctossiana . . . . . . . . . . . . . . . . . . . . . . . . . .
alberta . . . . . . . . . . . . . . . . . . . . . . . . . .
anphilochus. . . . . . . . . . . . . . . . . . . . . . . . . .
astarte . . . . . . . . . . . . . . . . . . . . . . . . . .
bellona . . . . . . 5 . . . . . . . . . . . . . . . . . . .
chariclea . . . . . .26 . . . . . . . 2 6 . . . . . . . . . . .
dia . . . . . . 9 . . . . . . . . . . . . . . . . . . .
distincta . . . . . . . . . . . . . . . . . . . . . . . . . .
epithore . . . . . . 24 . . . . . . . . . . . . . . . . . . .
euphrosyne . . . . . . 20 . . . . . . . . . . . . . . . . . 20 .
freija . . . . . . . . . . . . . . . . . . . . . 5 . 5 2 5 .
frigga . . . . . - 2 9 . . . . . . .5 . . . . . . . . . . .
inprcba . . 1 1 . . . . . . . . . . . 5 . . . . . . . . . . .
iphigenia . . . . . . 16 . . . . . . . . . . . . . . . . . . .
krienhitd . . . . . .15 . . . . . . . . . . . . . . . . . . .
polaris . . . . . . . . . . . . . . . . . . . . . . . . . .
selene . . . . . .17.. . . . . . . . . . . . . . . . 5 .
thore . . . . . . 17 . . . . . . . . . . . . . . . . . . .
titania . 6 5 . . .15 . . . . . . . 5 . . . . . . . . . . .
Cupha . . . . . . . . . . . . . . . . . . . . . . . . . .
erymanthis . . . . . . . . 28 13 . 13 14 . 30 . . . . . . . . . . .
prosope . . . . . . . . 2 . . 2 2 . . . . . . . . . . . . .
Damora . . . . . . . . . . . . . . . . . . . . . . . . . .
segana . . . . . .16 . . . . . . . . . . . . . . . . . . .
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 139

27. Continued
TABLE

. . . . . . . . . . . . . . . . . . . . . . . . . . . . anadyomene
. . . . . ' . 2 3 . . . .2327 . . . . . . . . . . . . . . paphia
. . . . . . . . . . . . . . . . . . . . . . . . . . .Argyreus
. . . . . . . . . . . . . . . . . . . . . . . . . . . hyperbius
. . . . . . . . . . . . . . . . . . . . . . . . . . . Argyronm
. . . . . . . . . . . . . . . . . . . . . . . . . . . laodice
. . . . . . . . . . . . . . . . . . . . . . . . . . . ruslana
. . . . . . . . . . . . . . . . . . . . . . . . . . . Brenthis
. . . . . . . . . . . .2217.. . . . . . . . . . . . daphne
. . . . . . . . . . . . 31 . . 23 . . . 23 . . . . . . . hecate
. . . . . 27 . . 16 . . . 2 0 1 7 . 17 . . . . . . . . . . . ino
. . . . . . . . . . . . . . . . . . . . . . . . . . . Boloria
. . . . . . . . . . . . . . . . . . . . . . . . . . . aquilonaris
napaea
. . . . . . . . . . . . . . . . . . . . . . . . . . .
. . .23.. . . . . . . . . . . . . . . . . . . . . . pales
. . . . . . . . . . . . . . . . . . . . . . . . . . . sifanica
. . . . . . . . . . . . .
. . . . . . . . . . . . . . Cirrochroa
. . . . . . . . . . . . .
. . . . . . . . . . . . . . regina
. . . . . . . . . . . . .
. . . . . . . . . . . . . . thais
. . . . . . . . . . . . .
. . . . . . . . . . . . . . CLossiana
. . . . . .
. 1 8 . . . .
. . . . . . . . . . . . . . . alberta
. . . . . . . . . . . . . . . . . 18 . . . . . . . . . amphilochus
. . . . . . . . . . . . . . . . . 1 5 . . . . . . . . . astarte
. . . . . . . . . . . . . . . . . . . . . . . . . . . bellona
. . . . . . . . . . . . . . . . . . . . . . . . . . . chariclea
. . . . . . . . . . . . 2 7 . . . . . . . . . . . . . 2 7 dia
. . . . . . . . . . . . . . . . . 18 . . . . . . . . . distincta
. . . . . . . . . . . . . . . . . . . . . . . . . . . epithore
. . 1 . . . . . . 2 2 . . . . . . . . . . . . . . . . . euphrosyne
5 . . . . . . . . . . . 15 . 16 . . . . . . . . . . . . f r e i j a
. . . . . . - 1 5 . . . . 5 . . . . . . . . . . . . . . frigga
. . . . . . . . . . . . . . . . . . . . . . . . . . . inproba
. . . . . . . . . . . . . . . . . . . . . . . . . . . iphigenia
. . . . . . . . . . . . . . . . . . . . . . . . . . . krienhild
. . . . . . .20.. . . . . . . . . . . . . . . . . . polaris
. . . . . . . . . 5 . . . . . . . . . . . . . . . . . selene
. . . . . . . . . . . . . . . . . . . . . . . . . . . thore
. . . . . . . . . . . . . . . . . . . . . . . . . . . titania
. . . . . . . . . . . . . . . . . . . . . . . . . . .Cupha
. . . . . . . . . . 8 . . . . . . . . . . .28 . 3 . . erymanthis
. . . . . . . . . . . . . . . . . . . . . 4 . . . . . prosope
. . . . . . . . . . . . . . . . . . . . . . . . . . . Danwra
. . . . . . . . . . . . . . . . . . . . . . . . . . . sagana
140 P. R. ACKERY

each of the five postulated heliconiine radiations (Philaethria, primitive EueideJ,

the Heliconius aoede-group, H. nattereri and H. hermathena) are primarily associated
with the most ‘ancient’ Passifloraceae (Dilkea, Mitostemma, Tetrastylis and
subgenus Astrophea of PassiJora) . Generally, more advanced representatives of
each group associate with the more recently radiated Passifloras. T h e sara-sapho
group provides a notable exception; this advanced group feeds on ‘primitive
passifloras’, argued as a secondary development by Benson et al. (1976). Several
adaptations in Passzjlora seem to be specifically related to exploitation by
heliconiines, particularly modified leaf-hairs (trichomes) (Gilbert, 197 1 ), egg-
dummies, extra-floral nectaries and the extraordinary diversity of leaf shape
(Gilbert, 1975).
Assumed unpalatability in the heliconiines is fundamental to Bates’ ( 1862)
original treatise on mimicry, while the presence of parallel heliconiine patterns
(Brown, Sheppard & Turner, 1974) is explained in ‘Mullerian’ terms.
Papageorgis ( 1975) documents five vertically stratified homochromatic mimetic
rings in the rain forest of Peru; heliconiines are conspicuous elements in four of
the five groupings. Despite this circumstantial evidence, the chemical basis for
heliconiine unpalatability remains poorly understood. Indeed, the Passifloraceae
have often been cited as non-poisonous although cyanogenic glycosides are now
known to be widespread (see Brower & Brower, 1964, and references therein;
Darnley Gibbs, 1974). Nahrstedt & Davis (1981), although demonstrating
cyanogenesis in Heliconius species, favoured de nouo synthesis in preference to
sequestration from the larval hosts. The recent novel suggestion that adult
heliconiines obtain toxins directly or indirectly through pollen-feeding,
particularly on the widely favoured genera Guarania and Psiguria
(Cucurbitaceae), has yet to be tested seriously (Grimshaw, 1983).
Argynninae
Two studies, Warren (1944) and Dos Passos & Grey (1945), provide an
incomplete systematic basis for this largely Holarctic group; unfortunately, each
is somewhat parochial, being respectively concerned with only the European or
North American faunas. Warren recognized 14 genera. To these must be added
Euptoieta and Yramea, and a number of exotic genera conventionally treated as
‘Argynnines’: Cupha, Vagrans, Phalanta, Vindula, Paduca, Cirrochroa and Terinos
(Corbet & Pendlebury, 1978), together with Smerina and Lachnoptera (Carcasson,
1981).
Holarctic representatives mostly exploit Viola (Violaceae), but several other
families, particularly the Ericaceae, Rosaceae, Salicaceae and Polygonaceae,
occur often enough to appear convincing (Tables 27-29). I n Brenlhis, at least,
rosaceous hosts have been noted far more frequently than members of the
Violaceae. However, with argynnines often ovipositing on various substrates
other than their larval hosts (Dethier, 1959), many oviposition observations
have now no doubt become enshrined in the literature as fanciful foodplants.
Remington ( 1952) suggests that each non-violaceous record for ‘Boloria’
probably represents an observation of oviposition.
Argyreus hzperbius, the sole Argyreus species, is the only typically Oriental
argynnine restricted to the Violaceae ( Viola). Otherwise, the Flacourtiaceae
appear most typical of the Orient, mirroring the switch found in Issoria sinha, an
exotic representative of the predominantly Holarctic genus Issoria. However, in
Terinos and Vindula other families predominate (Table 29). Vindula is known
28. Hostplants of the Argynninae (in part). Comstock & Garcia, 19611;d'Araujo e Silva et al., 19682; Ehrlich & Ehrlich, 19613; Emmel &
E
1973*; Hayward, 196g5; Henriksen & Kreutzer, 198Z6; Higgins & Riley, 1970'; Langer, 19588; Mori, 19589; Niculescu, 1965"-',Robert et al.,
1983l I ; Rose, 197612;Schwarz, 194913; Sevastopulo, 197314; Tietz, 197215; van Son, 197916; van Someren, 197417 3

Claudia . 3 - 3 . 1 . 3 . 3 . . . . 3 . 1 5 . . . 3 . 1 5 3 . . . . 4 . 1 . 2 . . . 15 . .
hegesia . . . . . . . . . . . . . . . . 1 2 . . . . . . . . . 15 . . . . . 2 . . . . . .
Fabriciana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
adi ppe . . . . . . . . . a . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
elisa . . . . . . . . . 7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
nerippe . . . . . . . . .9 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
niobe . . . . . . . . .6.... . . . . . . . . . . . . . . . . . . . . . . . . 7
lssoria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
baunanni . . . . . . . . . 17 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
cytheris . . . . . . . . . 5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
hanningtoni . . . . . . . . . 17 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Lathonia . . . . . . . . .ll . . . . . . . . 1 0 . . . . . 1 3 . . . . . . . . . l l l l . . .
sinha . . . . . . . . . . . 1414 . . . . . . . . . . . . . . . . . . . . . . . . . .
smaragdifera . . . . . . . . . 16 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
142 P. R. ACKERY
TABLE29. Hostplants of the Argynninae (in part). Bascombe et al., 1987'; Common &
Waterhouse, 1972'; Corbet & Pendlebury, 19783; D'Abrera, 19774; Durden, 19655; Ebner,
19706; Emmel & Emmel, 19737; Emmel et al., 19708; Ferris & Brown, 19819; Fountaine,
unpublished''; Gifford, 1965"; Harris, 19721Z; Higgins & Riley, 197OI1; Hill et al., 197814;
Hoffman et al., 193815; Hopkins, 192716; Howe, 1975"; Langer, 195818; Mamet, 194819;
Pcnnington, 1978'O; Pinhey, 19492'; Pyle, 1975''; Robert et al., 198323; Samson, 198OZ4;
Sevastopulo, 1975'5; Swezey, 1942bZ6;Tietz, 1972'7; Tilden, 196528; van Someren, 197419;
Woodhouse, 195230

iole . . . . . . . .25 . . . . . . . . .29 . . . . . .

Mesoacidalia. . . . . . . . . . . . . . . . . . . . . . . . .
aglaja . . . . . . 23 . . . . . 23 . . . . . . . . . . . .
Pandoriana . . . . . . . . . . . . . . . . . . . . . . . . .
pandora . . . . . . . . . . . .23 . . . . . . . . . . . .
Phalanta . . . . . . . . . . . . . . . . . . . . . . . . .
alcippe . . . . . . . . . . 30 . . . . . 24 . . . . . . . .
eurytis . . . . . . . . . . . . . . . 20 . . . 29 10 . . . .
exulans . . . . . . . . . . .16 . . . . . . . . . . . . .
phalantha . . . . . . . . . . . . 21 . 30 . . . . 14 29 15 . . .
Proclossiana. . . . . . . . . . . . . . . . . . . . . . . . .
eununia . . . . . 9 1 8 . . . . . 18 . . . . . . . . . . . .
Speyeria . . . . . . . . . . . . . . . . . . . . . . . . .
aphrodite . 27 . 27 . . . . . . . . 6 . . . . . . . . . . . .
atlantis . . . . . . . . . . . . 9 . . . . . . . . . . . .
callippe . . . . . . . . . . . . 7 . . . . . . . . . . . .
coronis . . . . . . . . . . . .22 . . . . . . . . . . . .
cybele . . . . . . . . . . . . 9 . . . . . . . . . . . .
diana . . . . . . . . . . . .12 . . . . . . . . . . . .
eduardsi . . . . . . . . . . . . 9 . . . . . . . . . . . .
egleis . . . . . . . . . . . .28 . . . . . . . . . . . .
hydaspe . . . . . . . . . . . . 7 . . . . . . . . . . . .
idalia . . . . . . . . . . . .27 . . . . . . . . . . . .
mormonia . . . . . . . . . . . .8 . . . . . . . . . . . .
nokunis . . . . . . . . . . . .8 . . . . . . . . . . . .
zerene . . . . . . . . . . . .22 . . . . . . . . . . . .
Terinos . . . . . . . . . . . . . . . . . . . . . . . . .
terpander . . . . . . . . . . . . . . . . . . . . . . . . .
Vagrans . . . . . . . . . . . . . . . . . . . . . . . . .
egista . . . . . . . . . . . . . . . . .2 . . . 2 . . .
Vindula . . . . . . . . . . . . . . . . . . . . . . . . .
arsinoe . . . . . . . . . . . . . . . . . . . . . . . 2 .
dejm . . . . . . . . . . . . . . . . . . . . . . . 3 .
erota . . . . . . . . . . . . . . . . . . . . . . .3030

from several passifloraceous genera (Adenia, Passgora, Modecca-see the

Heliconiinae above), while records for Terinos are limited to T. terpander on
Stilaginaceae (Antidesma).
Species of Phalanta occur throughout much of the Old-World tropics. Despite
scattered records for violaceous hosts, most fall within the families
Flacourtiaceae, Salicaceae and Celastraceae. Finally, the flacourtiaceous theme,
although recurring in the Afrotropical Lachnoptera, is absent in the New World
genus Euptoieta, for which such seemingly diverse families as the Violaceae,
Passifloraceae, Podophyllaceae, Crassulaceae, Fabaceae, Portulacaceae,
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 143

TABLE
29. Continued

. . . . . . . . . . . . . . . . . . . . . . . . iole
. . . . . . . . . . . . . . . . . . . . . . . . Mesoacidal ia
. . . . . . . . . . . . . . . . . . . . . . . . aglaja
. . . . . . . . . . . . . . . . . . . . . . . . Pandoriana
. . . . . . . . . . . . . . 2 3 . . . . . . . . . paKfora
. . . . . . . . . . . . . . . . . . . . . . . . Phalanta
. . . . . 2 4 . . . . . . . . . . . . . . . . . . alcippe
. . . . 20 . . . 25 . . . . . . . . . . . . . . . eurytis
. . . . . . . . . . . . . . . . . . . . . . . . exulans
. . . . 19 14 . 29 25 . 2 . . . . . 1 1 . . . . . 30 phalantha
. . . . . . . . . . . . . . . . . . . . . . . . Proclossiana
. . 13 . . 17 . . . . . . . . . . . . . . . . . . eunomia
. . . . . . . . . . . . . . . . . . . . . . . .Speyeria
2 7 . . . . . . . . . . . . . . . . . . . . . . . aphrodite
. . . . . . . . . . . . . . . . . . . . . . . . atlantis
. . . . . . . . . . . . . . . . . . . 5 . . . . callippe
. . . . . . . . . . . . . . . . . . . . . . . . coronis
. . . . . . . . . . . . . . . . . . . . . . . . cytele
. . . . . . . . . . . . . . . . . . . . . 2 7 . . diana
. . . . . . . . . . . . . . . . . . . . . . . . eduardsi
. . . . . . . . . . . . . . . . . . . . . . . . egleis
. . . . . . . . . . . . . . . . . . . . . . . . hydaspe
. . . . . . . . . . . . . . . . . . . . 2 7 . . . idalia
. . . . . . . . . . . . . . . . . . . . . . . . mormnia
. . . . . . . . . . . . . . . . . . . . . . . . nokmis
. . . . . . . . . . . . . . . . . . . . . . . . zerene
. . . . . . . . . . . . . . . . . . . . . . . .Terinos
. . . . . . . . . . . . 3 . . . . . . . . . . . terpander
. . . . . . . . . . . . . . . . . . . . . . . .Vagrans
. . . . . . . 2 6 . . . . . . . . . . . . . . . . egista
. . . . . . . . . . . . . . . . . . . . . . . .Vidula
2 . . . . . . . . . . . . . . . . . . . . . . . arsinoe
. . . . . . . . . . . . . . . . . . . . . . . . dejone
1 0 . . . . . . . . . . . . . . . . . . . . . . . erota

Menispermaceae, Linaceae, Boraginaceae, Turneraceae, Nyctaginaceae,

Asclepiadaceae, Convolvulaceae and Onagraceae are all recorded (Table 28).
Two closely related families, the Flacourtiaceae and Violaceae, thus appear
typical of the Argynninae. Their distributions (the generally tropical or
subtropical Flacourtiaceae and the cosmopolitan but chiefly temperate
Violaceae) are reflected by the hostplant preferences within the subfamily.
Scattered records for the Passifloraceae, another group of Violales, are of
particular interest in view of the widespread assertion that the passifloraceous
heliconiines are closely related to both the Argynninae and the frequently
144 P. K.ACKERY

passifloraceae-feeding Acraeinae (see Ehrlich, 1958). T h e diverse, less-favoured

families, particularly the Rosaceae, Salicaceae, Polygonaceae, Saxifragaceae
and Plantaginaceae, show little close interrelationship or especial affinity with
the Violales.

Melitaeinae
Higgins ( 1981) recognizes with certainty three melitaeine tribes-the
Holarctic Euphydryini, the largely Holarctic Melitaeini and the New World
.Phyciodini; all four species of the Antillean genus Atlantea, tentatively suggested
to constitute a further tribe, lack hostplant data. With the Euphydryini focusing
very much on the Scrophulariaceae and Caprifoliaceae, the Melitaeini on
Scrophulariaceae and Asteraceae, and the Phyciodini on the Asteraceae and
Acanthaceae, there is considerable overlap in hostplant preferences.

'J'ABLE 30. Hostplants of the Melitaeinae (Euphydryini, in part). Bowers, 1981l; Ehrlich &
Ehrlich, 19612 ; Forster & Wohlfahrt, 1955'; Gullander, 195g4; Harris, 197Z5; Henriksen &
Kreutzer, 19826;Higgins & Riley, 1970'; Kim, 19848; Langer, 19589;Manley & Allcard, 1970"';
Mazel, 1982"; Muller, 1969'*; Robert et d.,198313; Schwarz, 194914; Tietz, 197215

nmturna . . . . . 9 14 . . . . . . . . . . . . . . . .
. 14 14 . . .
intermedia . . . . . . . . . . . . . . . . . . . . . . . .
. . . . .
Cynthia . . . 3 . . . . . . . . 3 . . . . . . . . . .
. . . . . .
gillettii . .. . . . . . . . . . . . . . . . . . . . . .
. . . . .
i duna . .. . . . . . 6 . . . . . . . . . . . . . .
. . . . . .
Euphydryas . .. . . . . . . . . . . . . . . . . . . . .
. . . . . .
phaeton .
. 15 . . . . . . . . . . . 15 . 5 . 15 . . . .
. 5 . . . .
Eurcdryas . .. . . . . . . . . . . . . . . . . . . . . .
. . . . .
aur i ni a . .. . . . . . . . 14 . . . . . . . . . 1 0 . 3 . . . . 9 .
desfontainii. . . . . . . . . . . . . . . . . . . . . . . . . . . .10

maturna . 9 . . . . . . . . . 14 . 9 . 1 4 . . . . . . 4 . . . . . .
intermedia . . . . . . . . . . . . -11 . . . . . . . . . . . . . . .
Cynthia . 3 . . . . . . . . . . . . . . . . . . . . . . . . . . .
gillettii . . . . . . . . . . . . . 15 15 . . . . . . . . . . . . . .
iduna . 6 . . . . . . . . . 6 . . . . . . . . . . . . . . . . .
Euphydryas . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
phaeton . . . 15 2 . 5 1 5 1 2 1 5 1 . . 5 . 5 . . . . . . . . . 15 . . 15
Eurcdryas . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
aurinia . . . . 10 . . . . . 10 . . 14 . . 14 6
10 . 11 10 9 10 . . 14 . .
desfontainii . . . . . . . . . . . . . . . . . . . . 11 7 . . . . 13 . .
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 145

Euphydry ini
The smallest of these three tribes, the Euphydryini, comprises four genera
(Higgins, 1978, 1981); Euphydryas, Hypodryas, Occidryas and Eurodryas (although
others argue that these are only species-groups of a single genus; Ehrlich &
Murphy, 1982). With the exception of the North American gilletti, all species of
Hypodryas are Palaearctic. Available hostplant records suggest no family as
'typical' (see Table 30), but the scarcity of records for the characteristically
melitaeine Scrophulariaceae is surprising; perhaps it relates to competition with
Mellicta and Didymaeformia species which widely utilize this family.
The exclusively Nearctic Occidryas-four species in all, each with known
hosts-focus on the Scrophulariaceae and, to a lesser extent, the Caprifoliaceae
and Plantaginaceae (see Table 31). Euphydryas, in Higgins' (1978) restricted
sense, is now monobasic, accommodating only the eastern Nearctic E. phaeton.
Again, the Caprifoliaceae and Scrophulariaceae predominate (Table 30). Such
weak representation of the Plantaginaceae is surprising, although Bowers ( 1980)
notes post-diapause switching to Plantago, and Stamp (1979) indicates
oviposition on Plantago lanceolata. Other supposed hosts probably require
confirmation.
Four species represent the remaining genus, the exclusively Palaearctic
Eurodryas; only the widespread E. aurinia has abundant records. While sharing
with other 'Euphydriines' a penchant for the Caprifoliaceae, Scrophulariaceae
and Plantaginaceae, it is also widespread on the Dipsacaceae, as is E. desfontuinii
(Table 30).

TABLE31. Hostplants of the Melitaeinae (Euphydryini, in part). Austin & Austin, 1981';
Dornfeld, 19802; Ehrlich & Ehrlich, 19613; Emrnel & Emrnel, 19734; Garth & Tilden, 19635;
Howe, 19756; Murphy & Ehrlich, 1984'; Pyle, 1974"; Singer, 19849; Tietz, 1972IO; Tilden,
1965"; White, 197912; White & Singer, 197413

anicia . 10 . . . . . . 10 . 10 . 6 . . . . 10
chalcedona . . . 10 10 . 10 . . . . . 5 . 10 4 . 4
colon . . . . . . . . . . . . . . . . . .
editha . 10 . . . . . . 10 . 6 . . . . . . 4

anicia . 12 a . . . . . . . 7 3 . . . . 10 . . . . . .
chalcedona . . 4 . 4 10 11 1 4 . . 4 4 10 .4 2 . .
4 10 10 10
colon . . . . . . . . . . . 3 . . . . 6 . . . . . .
editha . . 410 4 . . . . 813 9 . . . . . . . . . . .
I46 P. R. ACKERY

Melitaeini
The second tribe, the largely Holarctic Melitaeini, divides into four groups
(Higgins, 1981), the typically scrophulariaceous Mellicta- and Didymaeformia-
groups, the largely Acanthaceae-Asteraceae-feeding Chlosyne-group, and the
Gnathotriche-group, for which the larval foodplants remain unknown.
Melampyrum, Digitalis, Veronica and Linaria are the principal scrophulariaceous
hosts of the Palaearctic genus Mellicta, the sole representative of the Mellicta-
group; additional significant records occur for Plantago (Plantaginaceae) and
two composite genera, Chrysanthemum and Centaurea (Table 32).
The Didymaeformia-group contains four genera-Melitaea, Poladryas,
Didymaeformia and Cinclidia-and, apart from the north American Poladryas, is
exclusively Palaearctic. With hostplant data generally scant, particularly in
Melitaea and Didymaeformia, few generalizations emerge. Records for Melitaea
centre on the Scrophulariaceae and Plantaginaceae, with the addition of various
‘secondary’ hosts (see Table 32). Known hosts for Didymaeformia, such as they
are, show a similar, typically ‘Melitaeine’ range, with particular emphasis on
the Scrophulariaceae, Plantaginaceae, Lamiaceae and Violaceae, although
D.didyma also takes a range of composites (see Table 32).
The only Nearctic representatives of the Didymaeformia-group, Poladryas
species, are apparently restricted to the Scrophulariaceae (Penstemon-see
Table 32), but with Cinclidia, we begin to move away from the scrophu-
lariaceous theme toward greater emphasis on the Asteraceae (Table 32), thus
far only a minor, but widespread, host family. However, C. phoebe additionally
takes typically ‘Melitaeine’ hosts of the families Scrophulariaceae, Planta-
ginaceae and Dipsacaceae.
The Asteraceae theme is also evident in the New-World Chlosyne-group. With
five included genera, only Microtia lacks foodplant data. Chlosyne species divide
between three species-groups (Higgins, 1960). The exclusively Nearctic harrisii
species-group feeds largely on Asteraceae. I n the second species-group, also
Nearctic, known hosts again centre largely on composites, but apart from Aster
itself, very little obvious overlap occurs in hostplant preferences, either within
the group, or in comparison with the harrisii-group (see Table 33).
Only the third group, the lacinia species-group, extends into the Neotropics.
While the Asteraceae theme is still evident, in the Neotropics emphasis switches
to the Acanthaceae; only narua on Amaranthaceae and hippodrome on Asteraceae
appear exceptional. Reverting to the overall trend, C. lacinia, a widespread
species of both the Neotropics and North America, takes a broad range of
composite plants, particularly Helianthus and Xanthium, with additional
representation of the Amaranthaceae. Of the three genera concluding the
Chlosyne-group, Texola and Dymasia continue to emphasize Acanthaceae-feeding,
while Thessalia species exploit the Scrophulariaceae as major hosts (Table 34).

Phyciodini
With the exception of some species now placed in the Melitaeinae, Higgins’
( 1981) tribe Phyciodini largely corresponds to Phyciodes, sensu Hall ( 1928-30).
Higgins recognizes 12 genera of Phyciodini. Phyciodes, in its now restricted sense,
contains most of the Nearctic representatives of the tribe but also extends
southwards into Central America. Composites are the most widely utilized
hostplants, most particularly Aster and Cirsium. Of the lesser noted families
32.
E Hostplants of the Melitaeinae (Melitaeini, in part). Agenjo, 1975l; Ehrlich & Ehrlich, 19612; Forster & Wohlfahrt, 19553; Gullander, 195g4;
s & Riley, 19705; Howarth, 19736; Howe, 1975'; Iwase, 1954'; Kawazoe, 1956'; Kim, 1984IO; Kurentsov, 1970"; Langer, 1958' 2; Larsen,
13;Manley & Allcard, 197014;Mazel, 198215;Niculescu, 196516;Robert et al., 1983"; Schwarz, 1949"; Wattison, 193019;Wiltshire, 1957*"

athalia . . . . . . . . . . . . . . 6 . 16 . . . 18 . 16 . . . . 12 . . 16 . . . . . . . 1816 . 16 . . . .
deione . . . . . . . . . . . . . . . . . . . 119515 . . . . 15 . . . . . . . . . . . . . . . . . .
parthenoides. . . . . . . . . . . . . . . . 17 . . . . . 14 . . . . 14 . . . . 17 . . . . . 1 . . . . . . .
aurelia . . . . . . . . . . . . . . . . 5 . . . 121512 . . . . 12 . . . . . . . . . . . 12 . . . . . .
britomartis . . . . . . . . . . . . . . . . 4 . . .18 4 3 . . . .ll . . . . . . . . . . . 3 . . . . . .
varia . . . . . . . . . . . 5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
elitaea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
arduinne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20. . . . . . .
cinxia . . .13. . . . . . . . . . . .18 . . . . . . . . . . 1 3 . . . . . . 1 8 . . . 1 8 . . 1 8 . . .13
diamina . . . . . . . . . . . . . . . . 16 . . . . 15 16 . . . . 16 . 9 18 . . . . . . . . . . . . . . .
oladryes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
minuta . . . . . . . . . . . . . . . . . . . . . . . . 2 . . . . . . . . . . . . . . . . . . . . .
arachne . . . . . . . . . . . . . . . . . . . . . . . . 7 . . . . . . . . . . . . . . . . . . . . .
idymaeformia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
didyma . 18 . 18 . . . 18 . 16 . . . 18 13 . 16 . 13 . 17 . 16 . . 18 18 13 . . 18 . 16 . . 18 . 18 16 . . . 18 . . .
deserticola . . . . . . . . . . . . . . . . . .13 . . . . . . . . . . . . . . . . . . . . . . . . . . .
persea . . . . . . . . . . . . . . . . . . . . . 20 . . . . . . . . . . . . . . . . . . . . . . . .
trivia . . . 1 8 .18 . . . . . . . .16 . 1 7 . . . .17 .16 .1313 . . . . . . . . . . . . . . . . . . .
inclidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
phoebe . . . . . . . . . . . . . . . .17 . . . . . . . . .18 . . . . .13 . . .18 13 14 . 8 . . 8 . .
aetherie . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 . . . . . . .
scotosia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .lo . .lo . .
148 P. R. ACKERY

TABLE 33. Hostplants of the Melitaeinae (Melitaeini, in part). Austin & Austin, 19811;DeVries,
19862;Ehle, 19573; Ehrlich & Ehrlich, 1961'; Emmel & Emmel, 19625, 19736;Fountaine, 1913';
Garth & 'Tilden, 19638; Harris, 19729; Hayward, 19691°; Higgins, 196011;Howe, 197512; Iftner,
198311; Kimball, 1965l'; Masters, 196915; Neck, 197316, 1976l'; Pyle, 197418; Scott & Scott,
198Olg;Shields et al., 196gZ0;Tietz, 19722'

. . . . . . . . . . . . . . . . . . .
nycteis . . . . . . . . . . . . . 21 . . . . . . . . . . . 3 - 4 .
palla . . . . . . . . . . . . -21 - 1 2 . . . . . . . . . . . 4 .
gorgone . - - 21 . - - 9 . . - 1 4 . . . . 15 . . . . . . . . .13 4 .
harrisii . . . . . . . . . . . . . . . . . . . . . . . . .2 1 . 4 .
hoffmani . . . . . . . . . . . . . . . . . . . . . . . . . . .1 8 .
gabbi . . . . . . . . . . . . . . . . . 21 . . . . . . . . . . .
danaetas . . - . . - . . . . . . . . . . . . . . . . . . . 8 .
nevnegeni . . . . . . . . . . . . . . . . . . . . . . . . 1 . . 4 .
acastus - . - - - - - - .- . . . . . . . . . . . . . . . . . 11.
janais . - - . - 7 - - . . . . . . . . . . . . . .12. . . . . .
melanarge . . . . . . . . . . . . . . . . . . .2 . . . . . . . . .
lacinia . 10 . . - - - - - . . . . . . . . . . . . . . . . . . 16
californica - . - . - - - - - 21 . . . . . . . . . . . . . . . . . . . .
narva . 2 . . . . . . . . . . . . . . . . . . . . . . . . . . .
gaudielis - . - - - - - . . . . . . . . . . .2 . . . . . . .
hippodram - . . - - . .- - - . . . . . . . . . . . . . . . . . . .
rosi t a - - - . - - - - . . . . . . . . . .I2 . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . 4 . . . . -19. . . -17. . . . .
. . .6 . . . . . . . . . . . . . . . 6 . . . . . .
. . . . . . . . .4.. . . . . . . . . . . . . . .
. . 17 . . . . . - 2 1 . . . . . . . . . . 21 . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . .
. 4 . . . . . 6 4 . 6 . . . . . . . . . . . . . . .
. . . 19 . . . . . . . . . . . . . . . . . . . . . .
. . . . . . ..........2 1 . . . . . . . . .
...........zo..............
. . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . .
2 - - 16 16 12 . 16 16 - - 21 16 . . 16 16 . 16 16 10 16 16 16
. . . . . . . . . . . . . . . . . . . . . 6 . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . 2 . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . .
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 149

(Table 34), the Acanthaceae appear most convincing, particularly in the light
of Kendall’s (1964) laboratory rearing of P. pictus on Siphonoglossa (for which it is
not recorded as a natural hostplant). Further scattered records also suggest the
Verbenaceae, Fabaceae, Convolvulaceae and Plantaginaceae.
Elsewhere in the Phyciodini, records are somewhat scant but in all cases
centre on the Acanthaceae (Table 34). Exceptionally, Ortilia liriope only takes
members of the Verbenaceae, Scrophulariaceae and Asteraceae. Continuing
contrary to the overriding acanthaceous trend, 0. orthia, Tegosa anieta and
Anthanassa texana utilize composites.
Summarizing, utilization of no single plant family is ‘typical’ of the
Melitaeinae; even the preferences of the three tribes are not mutually exclusive.
Exploitation of the Scrophulariaceae, Plantaginaceae, Caprifoliaceae and
Dipsacaceae is characteristic of the Euphydryini and much of the Melitaeini,
although in Cinclidia, Chlosyne, Texola and Dymasia emphasis switches to the
Acanthaceae and Asteraceae, a theme continued throughout the Phyciodini.
While the Euphydryini and Phyciodini perhaps represent little more than single
genera (see Hall, 1928-30; Ehrlich & Murphy, 1982) and are probably
monophyletic, the diverse Melitaeini, showing both euphydriine and phyciodine
hostplant tendencies, may merely be a poly- or paraphyletic residue. Within the
Asteraceae-Acanthaceae-feeding assemblage there is convincing, but not total,
biogeographic correlation, with Nearctic species typically exploiting Asteraceae,
while Acanthaceae-feeding characterizes the Neotropics. All the major
foodplant groups belong to the Asteridae: the Asterales (Asteraceae),
Dipsacales (Caprifoliaceae, Dipsacaceae), Scrophulariales (Scrophulariaceae,
Acanthaceae) and Plantaginales (Plantaginaceae).
Although mimicry is not obviously widespread in the Melitaeinae, Eresia and
Castilia include both ithomiine-like and acraeine-like species (Higgins, 1981 ) .
Higgins also postulates ‘arithmetic mimicry’ to account for remarkable pattern
convergences seen in widely disparate phyciodine species. The results of Bowers’
( 1980, 1981 ) experiments-working with E u p b d y a s phaeton, 0. chalcedona and
0. editha, she demonstrated unpalatability and even emetic properties-were
surprising, although Turner ( 1975, 1977) had suggested aposematic qualities for
the pattern of E. phaeton. Bowers emphasized the widespread presence of iridoid
glycosides in many melitaeine hostplant families, particularly the Plantaginaceae,
Scrophulariaceae, Dipsacaceae, Caprifoliaceae and Lamiaceae, inclining toward
the view that this chemical group makes significant contributions toward both
unpalatability and feeding stimulation (Bowers, 198313) in the species that she
studied. The presence of iridoid glycosides has not been confirmed in either the
Asteraceae or Acanthaceae (Jensen, Nielsen & Dahlgren, 1975)-the
Acanthaceae are known hosts for Eresia and probably Castilia, the most
obviously mimetic phyciodines, suggestive of a ‘Batesian’ role in their
association with the Ithomiinae and Acraeinae. Bowers (1983b) found that the
largely asteraceae feeding Chlosyne harrisii was essentially palatable and almost
certainly a ‘Batesian’ mimic of another melitaeine, the predominantly
scrophulariaceous Euphydryas phaeton.

Limenitinae
In the absence of a clearer idea of their relationships, the remaining
‘nymphalid’ genera can be tenuously split between two major groups, the
150 P. R. ACKERY

TABLE 34. Hostplants of the Melitaeinae (Melitaeini, in part, Phyciodini). Biezanko, 1949l;
d'ilraujo e Silva el al., 19682;DeVries, 19863; Dornfeld, 19804;Ehrlich & Ehrlich, 19615; Emmel
& Emmel, 19736; Ferris & Brown, 1981'; Fountaine, unpublished*; Garth & Tilden, 19639;
Hayward, 1969l"; Howe, 1975"; Kendall, 195912;Kimball, 196513;Lenczewskii, 198014;Leston
el al., 198315; Muller, 188616; Scott, 197417, 197618; Scott & Scott, 198Olg; Shapiro & Shapiro,
1973"; Shapiro et al., 19812'; Tietz, 197222;Young, 1973aZ3

Leanira . . . . . . . . . . . . 6 4 . . . . . . .
theona . . . . . . . . 5 . . . 5 . . . . . . .
12
Texola . . . . . . . . . . . . . . . . . . . . .
elada . . . . . . . . . . . . . . . . . . . . .
Dymasia . . . . . . . . . . . . . . . . . . . . .
dvlnas . . . . . . . . . . . . . . . . . . .5 .
Phyciodes . . . . . . . . . . . . . . . . . . . . .
tharos . . . . . . . . . . 22 . . . . . . . . . .
batesii . . . . . . . . . . . . . . . . . . . . .
canpest r i s . . . . . . . . . . . . . . . . . . . . .
mylittus . . . . . . . . . . . . . . . . . . . . .
pallidus . . . . . . . . . . . . . . . . . . . . .
orseis . . . . . . . . . . . . . . . . . . . . .
pictus . . . 22 . 7 . . . . . . . . . . . . . . .
phaon . . . . . . . 5 . . . . . . . . . . . . .
vesta . . . . . . . . . . . . . . . . . . . . .
Phystis . . . . . . . . . . . . . . . . . . . . .
simiois . . . . . . . . . . . . . . . . . . .10 .
Anthanassa . . . . . . . . . . . . . . . . . . . . .
ptolyca . . . . . . . . . . . . . . . . . . . . .
texana . . . . . . . . . . . . . . . . . . . 5 .
hermas . . . . . . . . . . . . . . . . . . . . .
frisia . . . . . . . . . . . . . . . . . . . . .
Telenassa . . . . . . . . . . . . . . . . . . . . .
teletusa . . . . . . . . . . . . . . . . . . . . 16
Ortilia . . . . . . . . . . . . . . . . . . . . .
liriope . . . . . . . .10 . . . . . .10 . . . . .
orthia . . . . . . . . . . . . . . . . . . . . .
i thra . . . . . . . . . . . . . . . . . . . . .
Tegosa . . . . . . . . . . . . . . . . . . . . .
anieta . . . . . . . . . . . . . . . . . . . . .
Eresia . . . . . . . . . . . . . . . . . . . . .
Lansdorf i . a . . . . . . . . . . . . . . . 1 ..i6
coe la . . . . . . . . . . . . . . . . . . . . .
eutropia .23 . . . . . . . . . . . . . . . . . . .
eunice . . . . . . . . . . . . . . . . . . . .
Cast ili a . . . . . . . . . . . . . . . . . . . .
eranites . . . . . . . . . . . . . . . . . . . .
ofetla . . . . . . . . . . . . . . . . . . 3 .
myia . . . . . . . . . . . . . . . . . . . .

Nymphalinae and Limenitinae. Such minor formal groupings as the Neptini,

Marpesiini, Coloburini, Ageroniini, Parthenini, Pseudergolini, Epicaliini,
Euthaliini and Biblinae are probably for the present best regarded as tribes. In
the Limenitinae, two host families predominate, the Euphorbiaceae and
Sapindaceae, although others, particularly the Rubiaceae, Salicaceae, Flacour-
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 151

TABLE
34. Continued

. . . . . . . . . . . . . . . . . . . . . . leanira
. . . . . . . . . . . . . . . . . . . . . . theona
. . . . . . . . . . . . . . . . . . . . . . Texola
. . .% . . . 5 . 5 . . . . . . . . . . . . elada
. . . . . . . . . . . . . . . . . . . . . . .Dymasia
. . . . . . . 11 . . . . . . . . . . . . . . d m s
. . . . . . . . . . . . . . . . . . . . . . Phycicdes
. . . . . . . . . . 22 . 20 . . 22 . . 19 . . . tharos
. . . . . . . . . . . . 5 . . . . . . . . . batesii
. . . . . . . . . . . 2 2 2 2 . . 22 . . . . . . carrpestris
. . . . . . . . . . . . . . 9 2 1 . 18 . . . . mylittus
. . . . . . . . . . . . . . . . 18 . . . . . . pallidus
. . . . . . . . . . . . . . . . 17 . . . . . . orseis
. . . . . . . . . . . . 5 . . . . . . . . . pictus
. . . . . . . . . . . . . . . . . . . . . . phaon
. . . . . . . 5 . . . . . . . . . . . . . . vesta
. . . . . . . . . . . . . . . . . . . . . . Phystis
. . . 10 10 10 . . . . . . . . . . . . . . . . . sirnois
. . . . . . . . . . . . . . . . . . . . . . Anthanassa
. . . . 3 . . . . . . . . . . . . . . . . . ptolyca
. . . 5 . . .22
5 . 5 . . . . . . . 2 2 . . . texana
. . . . 2 . . . . . . . . . . . . . . . . . hermas
15 . . . 14 14 . . . . . . . . . . . . . . . . frisia
. . . . . . . . . . . . . . . . . . . . . . Te lenassa
. . . 2 . . . . . . . . . . . . . . . . . . teletusa
. . . . . . . . . . . . . . . . . . . . . . Ortilia
. . . . . . . . . . . . . . . . l o . . . . . liriope
. . . . . . . . . . . . 1 1 . . . . . . . . orthia
. . . 10 . 10 . . . . . . . . . . . . . . . . ithra
. . . . . . . . . . . . . . . . . . . . . . Tegosa
. . . . . . . . . . . . . . . . . . .3 . . anieta
. . . . . . . . . . . . . . . . . . . . . . Eresia
. 2 . 2 2 . 2 . . . . . . . . . . . . . . 2 lansdorfi
. . 3 . 3 . . . . . . . . . . . . . . . . . coela
. . . . . . . . . . . . . . . . . . . . . . eutropia
.2 . . . . . . . . . . . . . . . . . . . . eunice
. . . . . . . . . . . . . . . . . . . . . . Casti 1 i a
. . . . 3 . . . . . . . . . . . . . . . . . eranites
. . . . 3 . . . . . . . . . . . . . . . . . ofella
. . . . 3 . . . . . . . . . . . . . . . . . myia

tiaceae, Caprifoliaceae and Moraceae, exemplify certain genera. The Urticaceae

and Acanthaceae clearly characterize the Nymphalinae. Of the tentative tribal
groupings of the Limenitinae, the Epicaliini and Limenitini most likely serve as
catch-all repositories for the less divergent genera; other tribes may well
represent natural groups.
152 P. R. ACKERY
'I'ABLE 35. Hostplants of the Limenitinae (Neptini, in part). Bascombe et al., 1987l; Carey-Hughrs
& Pickford, 1977'; Common & Waterhouse, 19723, 19814; Corbet & Pendlebury, 197g5;
Fountaixie, 19266;Hill et al., 19787;Sevastopulo, 19738;Woodhouse, 195Zg;Wynter Blyth, 1957I "

illigera . . . . . . . . . . . . . . . . . . . . . . . . 6 . . . .
tiga . 5 . . . . . . . . . . . . . . . . . . . . . . . . . . .
viraja . . . . . . . . . . . . . . . 8 . . . . . . . . . . . . .
Pantopria . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
consimilis. . . . . . . . . . . . . . . . 3 . 3 . . . . . . . . . .
hordonia . . . . . . . . . . . . 7 9 7 . . . . . . . . . . . . . .
sandaka . . . . . . . . . . . . . 5 . . . . . . . . . . . . . . .
Phaedyma . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
colunella . 7 . . 1 7 . . . . . . . . .10 . 2 . . . 5 7 . . . 7 . .
shepherdi . . . 3 . . . 4 . 3 3 . . . . . . . . 3 3 . . . . . . . 3

Neptini
Eliot ( 1969) recognized only five neptine genera, Pantoporia, Lasippa, Neptis,
Phaedyma and Aldania. Aldania lacks host data. Otherwise, exploitation of the
Fabaceae is characteristic of the tribe, with the qualification that only Neptis has
extensive foodplant records. While Pantoporia species are restricted to legumes,
Phaedyma and Lasippa appear rather more catholic in their tastes (Table 35),
with convincing records for the Ulmaceae, Clusiaceae, Sterculiaceae and
Bombacaceae.
Eltringham (1921) and Eliot (1969) between them cover the genus Neptis, but
unfortunately neither provides an all-embracing system. Although convinced
that the Afrotropical and Austro-Oriental representatives are congeneric, Eliot
(1969) states that the African fauna is not particularly closely related to any
Oriental elements, and certainly the host data appear to support this-the
Euphorbiaceae and Sapindaceae, the primary hostplant families in Africa
(Table 36), lack representation in any extra-African species (Table 37).
(Similarly, quite extensive records for the Rosaceae are restricted to the
Palaearctic.) Predictably the Fabaceae alone make notable contributions in
both zones, together with minor representation of the Tiliaceae. So, although
Fabaceae exploitation characterizes the group as a whole, the underlying
Sapindaceae/Euphorbiaceae trend remains well-marked in Afrotropical Neptis.

Biblini
Both Eliot (1978) and Carcasson (1981) separate a number of Old-world
genera (Ariadne, Laringa, Byblia, Mesoxantha, Neptidopsis, Eurytela) under the tribe
Biblidini presumably synonymous with the Eurytelini or Ergolini, which
includes the New World Biblis, Mestra and Vila. Of these, only Vila and Laringa
lack host data; all other representatives of the tribe are almost totally restricted
to the Euphorbiaceae (Table 38), one of the most widely utilized limenitine
foodplant families. I n the absence of any divergence from the likely
36.
E Hostplants of the Limenitinae (Neptini, in part). Chang, 1972'; Fountaine, unpublished2; Higgins & Riley, 19703; Iwase, 19544;Kurentsov,
Leech, 1893'j; Mamet, 1948'; Manders, 19088; Niculescu, 19659; Pennington, 1978'O; Pierre-Baltus, 1978' I ; Schwarz, 1949l 2; Sevastopulo,
1975'3; van Someren, 1974'*; van Son, 197915

agoua[e . . . . . . . . . . . . . . .11.. . . . . . . . . . . . . . . . . . . . . . .
dunetorun . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 0 . . . . . . . . . .
frobenia . . . . . . . . 7 . . . . . . . . . . . . . . . . 7 . . . . . . . . . . . . . .
kiriakoffi . . . . . 13 13 . . . . . . . . . . . . . . . . . . 14 . . . . . . . . 14 . . . . .
laeta . . . . . . . . . . . . . . . -1410 . . . . . . .15 . . . . . . . . . . . . . .
lativittata . . . . . . . . . . . . . . . . . . . . . . . .14 . . . . . . . . . . . . . . .
melicerta . . . . . . . . . . . . . . 11 11 . 11 . . . . . . . 10 14 . . . . . . 2 . . . . . .
nemetes . . . . . . . . . . . . . . . . . . . . . . . . . . 14 2 . . . . . . . . . . . .
nyasiodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14 . . . . .
poultoni . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14 .14 . . .
rogersi . . . . . . . . . . . . . . . . . . . . . . . . . . 13 . . . . . . . 1 4 . . . . .
saclava . . . . . 14 13 . . . . . . . . . . . . . . 10 14 . . 14 . . 10 . . . . . . . . . . .
strigata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14 . . .
trigonophera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 . . . . .
troundi . . .ll . . . . . . . . . . . . . . .ll . . . . . . . . . . . . . . I 1 . . . . .
vindo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 . . . . . . . .
aluina . . . . . . . . . . .4 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
ananta . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
pryeri . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . .
rivularis . . . . . . . . . .9 . 6 . . . . . . . . . . . . . . . . . . . . . . . . .1212
sappho . . . . . . . . . . . . . . . . . . 3 . . . . . . . . . . . . . . . . . . . . .
154 P. R. ACKERY
TABLE 37. Hostplants of the Limenitinae (Neptini, in part). Banno, 1984'; Bascombe et al., 1987*;
Chang, 197Z3;Hill ct al., 19784; Iwase, 19545, 19646; Kim, 19847;Kurentsov, 19708; Niculescu,
19659; Sevastopulo, 1973'O; Wynter Blyth, 195711

hylas . . . . . . . 11 11 11 . 11 . 11 . . . . . 2 . . 2
j h h . . . . . 11 . . 11 . . . . 11 . 11 11 . . . . . .
philyra . . . . . . . . . . . . . . . . . . . . 7 . .
phi lyroides . 7 . . . . . . . . . . . . . . . . . . . . .
sappho . . . . . . . . . . . . . . . . . . . . . . .
SOma . . . . . . . . . . . . . . . . . .3 3 . . .
thisbe . . . 8 . . . . . . . . . . . . . . . . . . .
yerburyi i . . . . . . . . . . . . . . . . . . 11 . . . .

hylas . 11 211 . 4 2 6 9 8 . 11 . 4 2 2 . 11 . . 11 . 1 1 . . . .
jvrllah . . . . 11 . . . . . . . 11 . . . . . 11 . 11 . . . 11 . .
philyra . . . . . . . . . . . . . .7 . 7 . . . . . . . . . 7
philyroides . . . . . . . . . . . . . . . . . . . . . . . . . . .
sappho . . . . . . . 5 . 5 5 . . . 1 5 5 . . 5 . . . . . . .
SOma . . . . . . . . . . . . . . . . . . . . . . . . . . .
thisbe . . . . . . . . . . . . . . . . . . . . . . . . . . .
yerburyii . . . . . . . . . . . . . . . . . . . . . . . . . . .

Euphorbiaceae/Sapindaceae limenitine prototype, the hostplants provide no

corroboratory evidence for this grouping. However, J. N. Eliot (personal
communication) believes that, despite the hostplant data, this group should not
be included in the Limenitinae.

Marpesiini
Just three genera represent the Marpesiini, the Old-world Cyrestis and
G'hersonesia, and the New-World Marpesia. Most known hosts belong to the
Moraceae, particularly Ficus and Moms (Table 39). For limenitines, Moraceae
feeding is unusual, but not unique; notably, it is shared with Pseudoneptis.
However, Marpesia (see Comstock & Garcia, 1961), Chersonesia (Hagen, 1896)
and Cyrestis (Fukuda, Harma, Takahashi et al., 1982-83) share a diagnostic
larva, smooth with elongate dorsal tubercles, whereas the Pseudoneptis larva is
typically limenitine (Aurivillius, 1912).

Ageroniini
Hamadryas and Ectima, two essentially Neotropical New-World genera,
together comprise the tribe Ageroniini. The Euphorbiaceae (mainly
Dalechampia, b u t also occasionally Tragia) are the typical host group (Table 38).
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 155

TABLE 38. Hostplants of the Limenitinae (Ageroniini, Biblini). Barcant, 19701;Biezanko, 194g2;
Brown & Heineman, 19723; d'Almeida, 19224; d'Araujo e Silva et al., 19685; DeVries, 19866;
Ehrlich & Ehrlich, 1961'; Ferris & Brown, 19818; Fontaine, 1981b9; Fountaine, unpublishedI0;
Hayward, 1969"; Miiller, 1886l 2; Muyshondt & Muyshondt, 197513;Pennington, 197814;Riley,
197515;Sevastopulo, 197316, 1975l'; van Someren, 197418;van Son, 197919;Woodhouse, 195220;
Wynter Blyth, 195721

ririna . . . . . . . .12 . . . . . . .
rectifascia . . . . . . . . 6 . . . . . . .
Hamadryas . . . . . . . . . . . . . . . .
amphim . 1 . . . . . . 13 . . . . 11 . .
arete . . . . . . . .12 . . . . . . .
arethusa . . . . . . . .5 . . . . . . .
arinane . . . . . . . .10 . . . . . . .
epinome . . . . . 2 . . 5 . . . . . . .
februa . . . . . 2 . .13. - 1 3 . . . .
feronia . . . . . . . . 4 . . . . . . 5
fornax . . . . . . . .5 . . . . . . .
guatemalena . . . . . . . . 13 . . . . . . .
iptheme . . . . . . . .6 . . . . . . .
lacdamia . . . . . . . . 6 . . . . . . .
Ariadne . . . . . . . . . . . . . . . .
albifascia . . . . . . . . 9 . . 9 . . . .
ar iadne . . . . . . .10 .21 .20 . . . .
enot rea . . . . . . . .19.. 9 . . . .
merione . . . . . . . . .16 .21 . . . .
pagenstecher i . . . . . . . . . . . 9 . . . .
personata . . . . . . . . . . 9 . . . . .
Biblis . . . . . . . . . . . . . . . .
biblis . . .10 . . . . . . .12 . . . .
hyperia . . . . . . . . . . . 2 . . . .
Bybl ia . . . . . . . . . . . . . . . .
anvatara . . . . . . . . 19 . . 9 . . . .
ilithyia . . . . . . . . 14 . . 14 . . . .
Eurytel a . . . . . . . . . . . . . . . .
alinda . . . . . . . . . . 9 . . . . .
dryope . . . . . . . . .18 .18 . . . .
hierbas . . . . . . . . 918. 9 . . . .
Mesoxantha . . . . . . . . . . . . . . . .
ethosea . . . . . . . . . . . 9 . . . .
Mestra . . . . . . . . . . . . . . . .
anrymone . . . . . . . .13. . 7 . . . .
cam . . . . . . . .15 . . . . . . .
dorcas . . . . . . . . . . .3 . . . .
hypernnestra . . . . . . . . 1 . . 8 . . . .
Neptidopsis . . . . . . . . . . . . . . . .
fulgurata . . . . . . . . 17 . . . . . . .
oph ione . . . . . . . . . . . 9 . . . .
156 P. R. ACKERY
‘I‘ABLE 39. Hostplants of the Limenitinae (Marpesiini, Pseudergolini, Parthenini). Barcant, 1970’:
Bascombe et al., 1987*; Biezanko, 194g3; Brown & Heineman, 197Z4; Corbet & Pendlebury,
19785; DAbrera, 19776; d’Araujo e Silva et al., 19687; DeVries, 198Ci8; Fountaine, unpublishedy;
Hagen, 189610;Hayward, 1969”; Hill el al., 1978’*; Howe, 197513; Kimball, 1965”; Miles
Moss, unpublishedI5; Riley, 197516; Sevastopulo, 197317, 197518; R. L. Smiles, pers. comm.lg;
Woodhouse, 195Z2”;Wynter Blyth, 195721

rahria . . . . .1D . . . . . . . . . . . . . . . . . . .
Cyrestis . . . . . . . . . . . . . . . . . . . . . . . . .
acilia . . . . . 6 . . . . . . . . . . . . . . . . . . .
camillus . . . . -18. 9 . . . . . . . . . . . . . . . . .
cocles . . . . . . . 9 . . . . . . . . . . . . . . . . .
periander . . . . . . . 9 . . . . . . . . . . . . . . . . .
thyodamas . . . . . 2 . . . . . . . . . . . . . . . . . . .
Marpesia . . . . . . . . . . . . . . . . . . . . . . . . .
chiron . . . 13 8 1 5 3 3 . . . . . . . . . . . . . . 7 . 7
eleucha . . . . . 4 . . . . . . . . . . . . . . . . . . .
petreus . . . . . 1411 1 . . . . . . . . . . . . 16 . 3 . .
D ichorrag i a . . . . . . . . . . . . . . . . . . . . . . . . .
nesimachus . . . . . . . . . . . . . . . . . 12 . . . . . . .
ninus . . . . . . . . . . . . . . . . . . . 1 9 . . . . .
Pseudergolis . . . . . . . . . . . . . . . . . . . . . . . . .
uedah . . . . . . . . .17 . . . . . . . . . . . . . . .
Amsia . . . . . . . . . . . . . . . . . . . . . . . . .
decora . . . . . . . . . . 5 . . . . . . . . . . . . . .
Parthenos . . . . . . . . . . . . . . . . . . . . . . . . .
cyaneus . . . . . . . . . . . . -17 . . . . . . . . . . .
Sylvia . 21 . . . . . . . . . . 2 0 2 1 . 6 . . . . . . . . .

The behavioural and morphological peculiarities of Hamadryas have long been

recognized (see, for instance, Fruhstorfer, 1916a), leading some to give the genus
family status (for review, see Jenkins, 1983), but in hostplant preferences at least
both genera remain characteristically limenitine. However, their mutual
affinities and relationship with the major tribe, the Epicaliini, still require
investigation.

Pseudergolini
The Pseudergolini, in which Eliot (1978) includes four genera, Dichorragia,
Pseudergolis, Stibochiona and Amnosia, and the Parthenini, are the last of the minor
limenitine groupings. Records of Dichorragia feeding on the Meliosmaceae and
Anacardiaceae, both members of the Sapindales, indicate some affinity with the
true limenitines, but the form of the larvae (generally smooth with a pair of
horns on the head, and paired spines on the eighth abdominal segment) is
clearly suggestive of the Apaturinae (see Eliot, 1978). Amnosia decora and
Pseudergolis wedah on the typically nymphaline Urticaceae (Elatostema and
Debregeasia respectively) only add to the confusion surrounding this interesting
‘group’ (Table 39).
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 157

Parthenini
Eliot (1978) includes just two genera within the Parthenini, Parthenos and
Lebadea. Only representatives of Parthenos have known hostplants, most
believably Adenia and Modecca of the Passifloraceae (Table 39; cf. Cethosia and
perhaps Vindula-Old-world passifloraceous nymphalines now considered to be
closely related to the Heliconiinae, Brown, 1981). Larvae of both genera show a
slight variation on the typically limenitine form, having strikingly long spines on
the second and third segments (Eliot, 1978), while the scant host data certainly
seem to suggest a notable shift.

Euthaliini
Their most distinctive larvae, with remarkably elongate lateral spines, readily
characterize this palaeotropical tribe. Eliot ( 1978) includes five Oriental genera,
Tanaecia, Euthalia, Dophla, Lexias and Bassarona: only the latter lacks data, while
the rest feed on no less than 15 families (see Table 40). Of this abundance of
data, only the Loranthaceae, and perhaps Anacardiaceae, appear to have been
recorded independently on several occasions. The Loranthaceae are well-known
parasites of the Anacardiaceae, suggestive of one of the principal evolutionary
pathways for postulated hostplant shifts (Chew & Robbins, 1984).
Among African limenitines, larval form places Catuna, Aterica, Euphaedra,
Euriphene, Hamanumida and Bebearia within the Euthaliini. These six genera are
noted on ten plant families (Table 41)-only two, the Sapindaceae and
Anacardiaceae, being shared with the Oriental representatives. Bebearia species,

TABLE 40. Hostplants of the Limenitinae (Euthaliini, in part). Bascombe et al., 1987’; Corbet &
Pendlebury, 1978*; D’Abrera, 19773; Fountaine, unpublished4; Hill et a/., 197F; Muroya et al.,
19676;Sevastopulo, 1973’; Wynter Blyth, 19578

evelina . . . . . . . . . . . . . . . . .7 . . . . . . . . . . . . . 8 . . .
!uthaLia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
aconthea. . . . . . . . . . 7 7 . . . 7 . . . . . 7 . . . . . 7 . . . 7 7 . .
aetion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
anosia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 . .
formosana. 6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Lubentina.. . . . . . . . . . . . . . . . . . . . . . . . . 5 5 . . . . . . .
monima . . . . . . . . . . . . . . . . . . . . . . . 2 . . . . . . . . . . .
nais . . . . . a . . . . . . . . . . . ? . . . . . . . . . . . . . . . . .
patala . 8. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
phemius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 . . 1 . .
vasanta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 . .
ex ias . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
aeropus . . . . . . . 3 . . . . . . . . . . . . . . . . . . . . . . . . . . .
dirtea . . . . 2 . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . 4
‘anaecia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Lepidea. . . . . . . . . . . . .7.. . . . . . . . . 7 . . . . . . . . . .
pelea . . . . . . . . . . . . . . . . . . . 2 . . . . . . . . . . . . . . .
1j a P. R. ACKERY

TABLE 41. Hostplants of the Limenitinae (Euthaliini, in part). Fountaine, unpublished'; Guilbot
& Guillaumin, 1977*; Hecq, 19803; Owen & Owen, 19734; Pennington, 19785; Sevastopulo,
19756; van Someren, 1974'; van Son, 1979*

galene . . . . . . 7 . . . . . . . . . . . . . . . 6 . . . . . . .
Bebearia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
carshena . . . . . . . . . . . . . . . . . . . . . . . . 6 . . . . .
chrienhilda.. . . . . . . . . . . . . . . . . . . . . . . . . . . 6 .
mardania . . . . . . . . . . . . . . . . . . . . . . . . . . 7 6 7 7
orientis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
sophus . . . 6 . . . . . . . . . . . . . . . . 6 . . . . . . . . .
Catuna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
erithea . . . 4 . . . . . 7 . . . . . . . . . . . . . . . . . . . .
Euphaedra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
coprates . . . . . . . . . . . 7 . 6 6 . 7 . . . . . . . . . . . . .
eleus . . . . . . . . . . . 6 . 6 6 . 7 . . . . . . . . . . . . .
medon . . . . . . . . . . . 6 . 7 6 . 7 . . . . . . . . . . . . .
neophron . . . . . . . . . . . 6 . 7 6 . 6 . . . . . . . . . . . . .
ochracea . . . . . . . . . . . 3 . . . . . . . . . . . . . . . . . .
rezia . . . . . . . . . . . . . . . 2 . . . . . . . . . . . . . .
Uganda . . . . . . . . . . . 7 . 7 . . . . 6 . . . . . . . . . . .
spatiosa . . . . . . . . . . . 6 6 6 7 . 7 . . . . . . . . . . . . .
Euriphene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
achlys . . . 5 . . . . . . . . . 6 . . . . . . . . . . . . . . . .
plautilla . . . 6 . . . . . . . . . . . . . . . . . . . . . . . . . .
Hamanunida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
daedalus . 1 . . . 5 . 8 . . . . . . . . . . . . . . . . . . . . . .

curiously often grouped as a subgenus of Euphaedra (e.g. Carcasson, 1981),

mainly take monocotyledons, mostly Arecaceae but also Marantaceae.
So, although larval structure adequately characterizes the Euthaliini, the
larval hosts offer little to conform the naturalness of the grouping. Exploitation
of the Moraceae and Rosaceae reflect more particular limenitine themes (see the
Marpesiini and Neptini above). Otherwise, the shift to monocotyledons by
Bebearia is most surprising (Charaxes, the only other nymphalid grouping with a
comparable range of hosts, similarly utilizes the Poaceae). Underlying this
diversity, the likely ancestral limenitine/Sapindaceae association continues with
variable emphasis in Euthalia, Euphaedra and Euriphene.

Epical iini
The predominantly New-World tribe, the Epicaliini, in the sense of Miller &
Brown (1981), subsumes such familiar tribal groupings as the Eunicini,
Catagrammini, Callicorini and Catonephelini. Apart from the curious genus
Lucinia, said to feed on Echites (Apocynaceae), the group divides neatly into
two-an overwhelmingly sapindaceous section, quite distinct from the
remaining genera which utilize the Euphorbiaceae. These series cut across the
boundaries of the traditional epicaliine and catagrammine associations. The
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 159

TABLE 42. Hostplants of the Limenitinae (Epicaliini, in part). Barcant, 19701;Biezanko, 194g2;
d’Araujo e Silva et al., 19683; DeVries, 1986*; Fountaine, unpublished5; Hayward, 1969‘j;
Lenczewski, 1980’; Miles Moss, unpublished8; Muller, 18869; Muyshondt, 1973b’O; c1I ; d ’ 2;
1 9 7 4 ~ ’ 1975b14;
~; Muyshondt et al., 197615;Riley, 1975“j

batesi . . . . . . . . . . 8 . . . .
Lepr ieuri . . . . . . . . . . 5 . . . .
Callidula . . . . . . . . . . . . . . .
pyrauus . . . . . . 6 2 . . 6 6 . .
6
Catagrm . . . . . . . . .. . . . .
cynosura . . . . . . . . .. 8 5 . . .
Lyca . . . . . . . . ..4 4 . . .
pitheas . . . . . . . . .. . 4 . . .
pygas . . . . . . 9 . . . . . . . .
Cyc logramna . . . . . . . . . . . . . . .
pandama . . . . . . . . . . .15 . . .
D i aethri a . . . . .. . . . . . . . . .
astala . . . . . . . . 14 . 14 14 . . .
aurelia . . . . 1 . . . . . . . . . .
caodrena . . . 2 . . . . . . . . . . .
clylnena . 3 . 2 7 . . . . . . . . . .
eupepla . . . . . . . . . . . 4 . . .
marchalli. . . . 4 . . . . . . . . . .
Epiphile . . . . . . . . . . . . . . .
adrasta . . . . . . . . 4 . 10 10 10 . .
orea . . . . . . . . . . 9 9 . . .
Nica . . . . . . . . . . . . . . .
flavilla . . . . . . . . 12 . 12 12 . . .
Pyrrhogyra . . . . . . . . . . . . . . .
crameri . . . . . . . . . .4 . . . .
edocla . . . . . . . . . . .4 . . .
neaerea . . . . . . . . . . 13 . . . .
tipha . . . . . . . . . . 1 . . . .
Temenis . . . . . . . . . . . . . . .
Laothoe . . . . . . . . 11 . 11 4 11 . .
Lucinia . . . . . . . . . . . . . . .
sida . . . . . . . . . . . . . . 16

sapindaceous theme of Asterope (‘ = Callithea auctt.), ‘Callidula’ (correctly

Haematera), Catagrarnma, Cyclograrnrna, Diaethria, Epiphele, Nica, Pyrrhogyra and
Ternenis involves just seven genera (see Table 42). Only Diaethriu strays beyond
these narrow confines, with particularly significant exploitation of the
Ulmaceae, a family of the Urticales with no obviously close relationship with the
Sapindales.
I n the euphorbiaceous series, Catonephele, Dynamine, Eunica, Myscelia and
Nessaea, the more believable hosts are again confined to a limited number of
genera (Table 43). Outside the New-World, only the African genus Sallya
represents the Epicaliini; indeed, it is sometimes treated as a subgenus of the
New-World Eunicu (see Carcasson, 1981). Like Eunica, Sallya species take a range
of euphorbiaceous hosts (Table 43).
160 P. R. ACKERY
TABLE
43. Hostplants of the Limenitinae (Epicaliini, in part). Barcant, 1970I; Biezanko, 194g2:
Brown & Heineman, 197Z3; d'Almeida, 19224; d'Araujo e Silva el al., 19685; DeVrics, 198tifi;
Doyle, 197g7; Ehrlich & Ehrlich, 196Is; Fontaine, 1981a9; Fountaine, unpublished'"; Hayward,
1969l Jenkins, 1984' 2 ; 198513; Miles Moss, unpublishedi4; Muller, 1886l 5 ; Muyshondt,
1973aIfi; Muyshondt & Muyshondt, 197517; Riley, 197518; Sevastopulo, 1975l 9 ; van Someren,
197420

acontius . 5 . . .13.. 5 . . . . . . . . . . . . . . . . . . . . .
chrunis . . . . . . . . 13 . . . . . . . . . . . . . . . . . . . . .
mexicana . . . . . . . . . 17 . . . . . . . . . . . . . . . . . . . .
nunilia . 5 . . . . . . 16 . . . . . . . . . . . . . . . . . . 1 . .
nyctinus . . . . . . . . .13.. . . . . . . . . 6 . . . . . . . . .
orites . . . . . . . . 6 . . . . . . . . . . . . . . . . . . . . .
Sabrina . 5 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Dynamine . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . .
agacles . . . . . . . . . 4 . 5 .. . . . 2 2 . . . . . . . . . . .
artemesia . . . . . . . . . 11 . . .
. . . . . . 10 . . . . . . . . . .
dyonis . . . . . . . . . . . . .. . . . . . 7 . . . . . . . . . .
egaea . . . . . . . . . . . . . . . . . . .3 . . . . . . . . . .
g 1auce . . . . . . . . . 6 . . . . . . . . . . . . . . . . . . . .
hopi . . . . . . . . . 6 . . . . . . . . . . . . . . . . . . . .
mylitta . . . . . . . . . 11 . . . . . 18 . . . . . . . . . . . . . .
myrFhina . . . . . . . . . . . 5 . . . . . 2 2 . . . . . . . . . . .
onias . . . . . . . . . 14 . . . . . . . . . . . . . .. . . . . .
salpensa . . . . . . . . .6 . . . . . . . . . . . . . . . . . . . .
tithia . . . . . . . . . 5 . . . . . . . . . . . . . . . . . . . .
Eunica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
margarita . . . . . . . . . . . 5 . . . . . . 2.. . . . . . . . . .
mi ra . . . . . . . . . . . .6 . . . . . . . . . . . . . . . . .
mnim . .
. . . . . . . . . . . . . . . . . . . . 6 1 . 8 . . . .
mygdonia . .
. . . . . . . . . . 6 . . . . . . . . . . . . . . . . .
nyscel ia . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . .
cyaniris . . . . . .12. 6 . . . . . . . . . . . . . . . . . . . .
.
orsis . .
. . . . . . . 15 . . . . . . . . . . . . . . . . . . . .
Nessaea . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . .
aglaura . . . . . . . 6 . . . . . . . 6...... . . . . . . .
.
Sallya . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
.
anulia . . . . . . . . . . . . . . . .9 . . . . . . . . . . . .
.
bengwlae . . . . . . . . . . . . .9 . . . . . . . . . . . . . . .
.
boisduvali . . 19 . . . . . . 19 . . 19 . . . 9 . . . . . . . . . . . 19
.
garega . . . . . . . . . . . . . . . . 20 . . . . . . . . . . . .
.
mrant i i . . . . . . . . . 2 0 . . 2 0 . . .19 . . . . . . . . . . . .
.
natalensis . . . . . . . . . 19 . . 19 . . . 19 . . . . . . . . . . . .
.
occidentaliun . . . . . . . . . . . . . 20 . . . 9 . . . . . . . . . . . .
pechwli . . . . . . . . . . . . . .9 . . . . . . . . . . . . . . .
trimeni . . . . . . . . . . . . . . . . . 9 . . . . . . . . . . . .

Limenitini
A Rubiaceae-Naucleaceae-Urticaceae theme dominates records for the genus
Adelfiha. O n the basis of morphology of the early stages Aiello (1984) presents a
partial classification of this most difficult genus. I n all, she recognizes seven
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 161

species-groups. Two groups particularly centre on the Urticaceae together with

representatives of a narrow range of additional families (Table 44). A strong
Rubiaceae theme characterizes most other groups with the exception of Aiello’s
group three-here the Verbenaceae are dominant.
The genus Basilarchia is of doubtful status (Chermock, 1950, treated Adelpha,
Basilarchia, Ladoga and Limenitis as congeneric). However, its major hostplants
and those of Adelpha have little in common, with Basilarchia species favouring
Salix and Populus (Salicaceae), but in the Rosaceae at least there are apparent
affinities (Table 45).
Of the Old-world genus Limenitis, only L. populi follows the salicaceous trend
so well marked in Basilarchia. Otherwise the Caprifoliaceae (Lonicera) dominate
the records (Table 46). Eliot (1978) considers that the Oriental species are not
true Limenitis, and in again taking up the Rubiaceae-Naucleaceae they appear
quite distinct (see Adelpha above). O n the other hand, Limenitis carnilla, often
distinguished as Ladoga, is found only on the Caprifoliaceae, much like true
Limenitis. So although two broad themes, Rubiaceae-Naucleaceae and
Caprifoliaceae, are well marked in this ‘group’, the hostplant pattern cuts right
across the generic limits as currently understood.
A number of other Old-World genera might be included in the Limenitini by
default-they do not appear to fit happily elsewhere. Superficially this appears
convincing with Athyma (including Parathyma and Tacoraea) and Pandita again
exploiting the Rubiaceae-Naucliaceae, together with strong representation of
the characteristically limenitine Euphorbiaceae (Table 46). If correctly placed,
+
three further genera, Cymothoe ( Harma) , Pseudacraea and Pseudoneptis, each
seems to represent an independent shift from the underlying limenitine themes
(Table 47); conversely, they may merely emphasize the arbitrary nature of the
Limenitini in the sense used here. Pseudoneptis is only recorded on the
Moraceae, the theme family of the Marpesiini from which it appears to be
excluded by larval form. Similarly, Pseudacraea takes up one of the lesser trends
of the Euthaliini, exploitation of the Sapotaceae. Again, larval structure (see
van Son, 1979-the caterpillars are typically limenitine) precludes this genus
from the tribe with which it shares closest hostplant affinity. Finally, Cymothoe
returns to the argynnine theme, widely utilizing the closely related Violaceae
and Flacourtiaceae (Table 47). Only occasional records for the Euphorbiaceae
and Uapacaceae appear remotely limenitine.
Overall, then, of the nine tribes tentatively associated within the Limenitinae,
the Neptini (Fabaceae), Biblini (Euphorbiaceae), Marpesiini (Moraceae),
Ageroniini (Euphorbiaceae), Parthenini (Passifloraceae) and Epicaliini
(Euphorbiaceae/Sapindaceae) have convincing, if not exclusive ‘theme families’.
However, the qualification should be added that utilization of the
Euphorbiaceae/Sapindaceae is quite likely an ancestral limenitine feature, a
contention perhaps supported by the exploitation of both these families as
‘secondary’ hosts in the Neptini and Euthaliini. The inclusion of Cymothoe,
Pseudacraea and Pseudoneptis is problematical given their hostplant preferences.
As noted elsewhere, the limenitines include many ‘Batesian’ mimics, perhaps
most notably in Pseudacraea (mimics of Acraeinae and Danainae), Neptis
(Tellervinae), Aterica (Danainae) and Basilarchia (Danainae and Papilionidae) .
Additionally, Asterope (Callithea auctt.) and Adelpha convergence on to the
patterns of Agrias and Doxocopa respectively, although there are no known
162 P. R.ACKERY

TABLE 44. Hostplants of the Lirnenitinae (Limenitini, in part). Aiello, 1984l; Barcant, 1970';
d'Araujo e Silva et al., 19683;DeVries, 19864;Ehrlich & Ehrlich, 19615 ; Fountaine, unpublishcd'j;
Hayward, 1969'; Miles Moss, 19338; Muller, 18869;Riley, 1975IO; Young, 1974c' '

basilea . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
basiloides . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
boreas . . . . . . . . . . . . . . . . . . . 4 . . . . . . . . .
cocala . . . . . . . . . . . . . . . . . . . . . . . . . . 8 . .
cytherea . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
demialba . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
fessonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
iphicla . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
leucoptha lma . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
me 1ona . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
mythra . . . . . . . . . . . . . . . . . . . . . 6 . . . . . . .
paraena . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
phliassa . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
plesaure . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
pseudococala . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
SYma . . . . . . . . . . . . . . . . . . . . . 7 . . . . . . .
tracta . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
ralmona . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
celerio . . . . . . . . 1 . . . . 1 . . . 1 . . . . . 1 1 . . . .
delphicola . . . . . . .8 . . . . . 8 . . 8 . . . . . . . . . . . .
isis . . . . . . . . . . . . . 9 9 . 9 . . . . . . . . . . . .
lara . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . .
melanthe . . . . . . . . . . 4 . . 4 . 4 . 4 . . . . . . . . . . .
mesentina . . . . . . . . . . . . .8 . . 8 . . . . . . . . . . . .
phylaca . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . .
bredowi . . . 5 . . . . . . . . . . . . . . . . . . . . . . . . .
creton . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . .
donysa . . . l . . . . . .. . . . . . . . . . . . . . . . . . .
abi a . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
cal liphane . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
epizygis . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
jordani . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
naxia . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . .
calliphiclea . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
serpa . . . . . . . . . . . . . . . . . . . . . . . . a . . . 3
boetia . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . .
erotia . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

chemical bases for these associations. With feeding experiments demonstrating

relative palatability in Hamadryas (DeVries, 1983) and Biblis (Brower & Brower,
1964), and larval hostplants including few of the families traditionally associated
with unpalatable butterflies (see Brower & Brower, 1964), their 'Batesian' role
remains supportable. There are exceptions, however-Basilarchia archippus is
known to be somewhat unpalatable, possibly when feeding on Salix babylonica
(Salicaceae) (Platt, Coppinger & Brower, 1971). More recent studies (Ritland
& Brower, 1986) suggest that the long accepted roles of the Viceroy and the
Florida Queen (Danaus gilippus) may need to be reversed, D.gilippus, with its
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 163

TABLE
44.Continued

. . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . basi lea
. . . . . . 1 . 1 . . 1 . . . . l . . . . . . . . . . . . basi loides
. . . . . . . . . . . . . 4 . . . . . . . . . . . . . . . boreas
. . . . . . . . . . . . 4 1 . . . a 4 4 . . . 6 . i . . i cocala
. . . . . . . . . . . . . . . . . . . . . . . a . . . . . cytherea
. . . . . . . . . . . . . . . . . . . . . 4 . . . . . . demialba
. . . .. . . . . . . . . . . . . . . 1 . . . . . . . . fessoni a
. . . .1 . . . 1 9 . 1 0 . 2 1 . . . . . . 1 . 6 . . . 1 iphicla
. . . . .. . . . . . . . . . . . . 11 . . . . . . . . . . leucopthalma
. . . . .. . . . . . . . . . . . 0 . . . . . . . . . . . me lona
. . . . .. . . . . 9 . . . . . . . . . . . . . . . . . . mythra
. . . . .. . . . . . . . . . a . . . . . a . . . . . . . paraena
. . . . . . a . . . . B . . . . . . . . . . . . . . . . . phliassa
. . . . . . . . . . 9 . . . . . . . . . . . . . . . . . . plesaure
. . . . . . . . . . . . . . . . . . . . . . . B . . . . . pseudococala
. . . . . . . . . . . . . . . . . . . . . . . . . . . 3 . SYma
. . . . . 4 . . . . . . . . . . . . . . . . . . . . . . tracta
. . . . . . . . . . . . . . . . . . . . . . . 4 . . . . . ralmona
. . . . . . . . . . . . . . . . . . . . . . . . . . . . ce 1er io
. . . . . . . . . . . . . . . . . . . . . . . . . . . . delphicola
. . . . . . . . . . . . . . . . . . . . . . . . . . . . isis
. . . . . . . . . . . . . . . . . . . . . . . . . . . . lara
. . . . . . . . . . . . . . . . . . . . . . . . . . . . melanthe
. . . . . . . . . . . . . . . . . . . . . . . . . . . . mesentina
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . phy laca
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . bredoui
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . creton
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . donysa
. . . 9 . . . . . . . . . . . . . . . . . . . . . . . . . abia
. . . 3 . . . . . . . . . . . . . . . . . . . . . . . . . cal l i p h a m
. . . 3 . . . . . . . . . . . . . . . . . . . . . . . . . epizygis
. . . a . . . . . . . . . . . . . . . . . . . . . . . . . jordani
. . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . naxia
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . calliphiclea
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . serpa
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . boetia
. 9 . . . . . . . . . . . . . . . . . . . . . . . . . . . erotia

relatively poor cardenolide defence system, being either a Miillerian or Batesian

mimic of the dark form of the Viceroy. Elsewhere in the Limenitinae, the
Moraceae are variously noted as larval hosts. Euploea species (Danainae), which
particularly favour this family, appear to be aposematic both as larvae and
adults (see below).
Nymphalinae
Most of the true nymphalines divide into two series on the basis of hostplants,
the mainly exotic species, largely on the Acanthaceae, and those with a more
164 P. R. ACKERY

TABLE 45. Hostplants of the Limenitinae (Limenitini, in part). Ebner, 1970'; Ehrlich & Ehrlich,
19612;Maddox & Cannell, 19833; Perkins & Perkins, 1975'; Pyle, 19745; Scott & Scott, I98Ofi;
'I'ietz, 19727

archippls . . . . . . 2 . . . . . 2 2 . 7 . . 7 . . 2 2 . . . . 7 . .
arthemis . . . 3 2 . . . 7 . 7 . 1 2 . . . 7 2 . . 7 7 3 .
7 . . . .
astyanax . 7 . . . . 7 . 7 . . . 2 2 . 2 . . 2 7 . 7 7 7 . . . . . 4
lorquini . . . . . . 7 . . . . . 2 2 . . . . 7 . . 5 2 . . 5 . . . .
ueidemeyeri.. . . . . . . . . . . 2 2 . . . 6 . . 6 . . . . . . . . .

TABLE 46. Hostplants of the Limenitinae (Limenitini, in part). Carey-Hughes & Pickford, 1977';
Chang, 19632; Corbet & Pendlebury, 19783; Fountaine, unpublished'; Henriksen & Kreutzer,
198Z5; Hill et al., 19786; Hoffman et al., 19387; Kim, 19846; Kurentsov, 19709; Muroya el af.,
1967"; Niculescu, 1965I I; Sevastopulo, 197312; ShirBzu, 196013; Tanaka, 1978I'; Uchida,
198415;Wynter Blyth, 195716

Camilla . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 5 1 4
Limenitis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
anphyssa . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 . .
calidosa . . . . . . . . . . . . . . . 1212 . . . . . . . . 4 . . . .
doerriesi . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 . .
dudu . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 . .
helmanni . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 . .
nwltrechti . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 . .
populi . . . 8 . . . . . . . . . . . . . . . . . . . . . . . . . .
procris . . . . . . . . . . . . . . . 12 12 3 12 . . 16 12 . 16 3 . . . .
reducta . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11 .
sulpitia . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 . .
sydyi . . . . . 9 9 . . . . . . . . . . . . . . . . . . . . 8 . .
trivena . . . . . . . . . . . . . . . . . . . . . . . . . . .16 . .
Athyma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
cams . . . . . . . . . 15 . . . . . . . . . . . . . . . . . . . .
kanua . . . . . . . . . . . . . . . . . . . . . . . . . 3 . . . .
nefte . . . . . . . . 1 1 2 . . . . . . 12 . . . . . . . . . . . . .
opalina . 1 3 . . . . . . . . . . . . . . 7 . . . . . . . . . . . . .
perius . . . . . . . . .1313 . . . . . . . . . . . . . . . . . . .
renga . . . . . . . . . . . . 1212 . . . . . . . . . . . . . . . .
selenophora . . . . . . . . . . . . . . . . 13 . 10 . 13 . . . . . . . . .
Pandi t a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
sinope . . . . . . . . . . . . . . . . . . . . . . . . . 3 . . . .

temperate range which favour the Urticaceae, two fairly disparate plant
families. Some genera though, most particularly the closely related Hypolimnas
and Salamis, cut right across this arbitrary division.
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 165

TABLE 47. Hostplants of the Limenitinae (Limenitini, in part). Birket-Smith, 19601; Fontaine,
19822, 19833; Fountaine, unpublished4; Gifford, 19655; Pennington, 19786; Sevastopulo, 1975';
van Someren, 1974*; van Son, 197g9

alcimeda . . . . . . . . . . . 6 . . . . . . . . . . . . . . . . . .
beckeri . . . . . . . . 3 . . . . . . . . . . . . .3 . . . . . . .
caenis . . . . . . . . 3 . . . a . . . . . . . . . . . I . . . . .
cocci nata . . . . . 3 . . . . . . . . . . . . . . . . . . . . . . . .
caranus 7 . . . . . . . . . . . g . . . . . . . . . . . . . . . a a
excelsa . . . . . 3 . . . . . . . . . . . . . . . . . . . . . . . .
haynae . . . . . 3 . . . . . . . . . . . . . . . . . . . . . . . .
herminia . . . . . . . . . . 7 . 7 . . . . . . . . . . . . . . . . .
hyarbi t a . . . . . . . . . . . . . . . . . . . . 3 . . . . . . . . .
jodutta . . . . . . . . 3 . . . . . . . . . . . . . . . . . . . . .
Lurida . . . . 7 . . . . . . . . . . . . . . . . . . . . . . . .
reginaeelirabethae. . . . . . 3 . . . . . . . . . . . . . . . . . . . . . . . .
reinholdi . . . . . . . . . 3 . . . . . . . . . . . . . . . . . . . .
sangar i s . . . . . 7 . . . . . . . . . . . . . . . . . . . . . . . .
theobene . . . . . 5 . 2 . . 5 . . . . . . . . . . . . . . . . . . .
Pseudacraea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
boisdwal ii . . . . . . . . . . . . . . a a 6 . . . . . . . . . . . . .
wrytus . . . . . . . . . . . . . . 8 . 8 . . . . . . . . . . . . .
Lucretia . . . . . . . . . . . . . . 6 7 6 7 7 . . . . . . . & . . .
Pseudoneptis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
ianthe . . 81, . . . . . . . . . . . . . . . . . . . . . . . . . .

Nymphalini
Of the Acanthaceae-series-Anartia, Catacroptera, Doleschallia, Kalima, Junonia,
Siproeta and Yoma-just the cosmopolitan Junonia (including Precis auctt.) has
abundant records (Tables 48-50). While the Acanthaceae characterize the
group as a whole, exploitation of members of the Scrophulariaceae, closely
related to the Acanthaceae within the Scrophualariales, and the Verbenaceae
and Lamiaceae (Lamiales) is widespread.
Data for Junonia mirrors these overall preferences although there is some
partial geographical differentiation-in Africa, many species favour the
Lamiaceae while the Scrophulariaceae and Verbenaceae dominate in the New
World. Although host data for African edemic species are quite extensive, only
Junonia natalica takes species of both the Lamiaceae (Plectranthus, Gifford, 1965)
and Acanthaceae. Otherwise, a well-marked division between lamiaceous and
acanthaceous sections (Table 48)corresponds closely to Junonia and Precis in the
sense of van Son, 1979 (but see J. orithya below). Among Oriental endemics, the
scrophulariaceous theme, most dominant in the New World, is quite strongly
developed with an additional range of acanthaceous genera underlining the
overall trend (Table 49).
Two species, Junonia orithya and hierta, extend throughout much of the Afro-
tropical and Oriental regions. Although hierta appears to be restricted to the
Acanthaceae throughout its range, J . orithya exploits the Lamiaceae in Africa,
166 P. R. ACKERY
TABLE 48. Hostplants of the Nymphalinae (Nymphalini, in part). Fontaine, 1985'; Fountaine,
unpublished2; Gifford, 19653; Mamet, 19484;Owen & Owen, 19735;Paulian, 19566; Pennington,
19787;Pinhey, 194g8; Sevastopulo, 19759;van Someren, 19741°; van Son, 1979' I ; Wynter Blyth,
195712

actia . . . . . . . . . :1 . . . . . . . . . . . . . . . .
andremia ja . . . . . 2. . . . . . . . . . . . . . . . . . . . .
anti Lope . . . . . 1 . . . 1 . . . . . . . . . . . . . . . . .
archesia . . . 9 . 7 . 9 . . 7 9 . . . . . . . . . . . . . . .
ceryne . . . . . . . . 1 . 1 . . . . . . . .
7 . . . . . . 9
eurodoce . . . . . 2. . . . . . . . . . . . . . . . . . . . .
octavia . . . 9 .10. 3 81101011 5 . . . . . . . . . . . . .
pelarga . . . . . . . . . . . . s . . . . . . . . . . . . .
1
rauana . . . . . . . . . 1 . . . . . . . . . . . . . . . . .
tugela . . . . . 910 . . . 7 9 . . . . . . . . . . . . . . .
artaxia . . . . . . . . . . . . . . . . . . . 1 . . . . . . .
chorinome . . . . . . . . . . . . . . . .10 9 . . - 1 0 9 . 9 . .
hierta . . . . . . . . . . . . . . . 12 9 9 7 . . 9 9 . 9 . .
limoria . . . . . . . . . . . . . . . . 10 9 . . . 9 9 . 9 . .
natal ica . . . . . . . . . . 3 . . . . .10 9 . 1 . 9 9 7 1 0 . .
oenm . . . . . . . . . . . . . . . .1 9 . . . 9 9 . 9 . .
rhadana . . . . . . . . . . . . . . . . . 4 . . 6 6 . . . . .
soph ia . . . . . . . . . . . . . . . . 9 9 . . . 910.9..
stygia . . . . . . . . . . . . . . . . 9 9...9101 . . .
terea . . . . . . . . . . . . . . . . 7 9 . 1 . 9 9 710..
uestermnni - 2 . . . . . . . . . . . . . . 109 . . . 9 9 . 9 . :

while the Scrophulariaceae and Acanthaceae typify the Arabian and Oriental
populations, cutting across the JunonialPrecis division. Minor hostplant families
of orithya in the Orient are generally characteristic of extra-African species.
In the New World, the taxonomy of Junonia remains unresolved (see Harvey,
in press; Bowers, 1984; Turner & Parnell, 1985). T h e Acanthaceae are still
represented by scattered records, but generally the Scrophulariaceae dominate,
with significant contributions from the closely related Verbenaceae and
Plantaginaceae.
The most outstanding feature of the Acanthaceae-series is the striking
parallel with the Melitaeinae, similarly dominated by the Asteraceae-
Acanthaceae-Scrophulariaceae-Plantaginaceae. Again, iridoid glycosides have
an important function, playing a key role as larval feeding stimulants (Bowers,
1984). The hostplants also shed light on the Precis/Junonia problem, as the names
are currently (but probably wrongly) applied to the African fauna. The
Lamiaceae-feeding species may well form a natural group (equivalent to Junonia
sensu van Son), although it is doubtful if the residual Precis would be
monophyletic. However, such an interpretation discounts Gifford's discordant
record (see above), and leaves Junonia orithya as an open question.
Hypolimnas and Salamis each has quite clearly differentiated urticaceous and
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 167

TABLE 49. Hostplants of the Nymphalinae (Nymphalini, in part). Atkins, 1975l; Bascombe et al.,
1987'; Pholboon, 19653; Chang, 19634; Common & Waterhouse, 19725; Corbet & Pendlebury,
19786; Hill et al., 1978'; Holloway & Peters, 1976"; Hopkins, 19279; Iwase, 1964lO; Larsen &
Larsen, 1980''; McCubbin, 1971''; Samson, 198013; Sankowsky, 197514, 197815; Sevastopulo,
197316; ShirBzu, 1960' '; Uchida, 1984'"; van Someren, 1974' 9; Woodhouse, 1952'O; Wynter
Blyth, 19572'

. . . . . . . . . . . . . . . . . . . . . . . . . . . . .
almana . . 7 . . . . . . . 6 . . . 1 6 . . . . . . . 10 . . . . . . .
atlites . 2 7 . . . . . . . . . . . . . . . . . . . . . . . . . . .
hedonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
iphita . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
lenwnias . . 3 . . .2 1 . 2 1 . . . . . . . . . . . . . . . . . . . . .
orithya . . . . . . . . . . . . . . . . . . . . 21 . 11 . . . 1 9 1 9 . 11
viLlida . . . . 5 . . . . . . . 9 . . . 5 . 1 2 5 1 3 . . a 5 . . . . 5

almana . . . 4 . 17 10 . . . 16 . 16 21 . 21 . 3 7 . . . . 17 . . . . . . . . . .
atlites . . . . . . 2 . . . . . -21 . 2 1 . . 7 . . . . . . . . . . . . . . .
hedonia . . . . . . . . . . . . . . . . . 15 14 . . . . 12 . . . . . . . . . .
iphita . . . . . . . . . . . . .21 . . . . .21 . . . .21 . . . . . . . . .
Lenwnias. . . . . . . . . . . . .21 . 2 2 3 2 . 721 . 3 2 . . . 318 . . . .
orythia . 5 5 . 5 . . . 7 . . . 16 . 5 11. . .
1 4 2 1 21 5 . . . 5 . . . . . . .
v i l l i d a . . 5 . . . . 1 . . . . . . . . . . . . . . . 1 2 . . . . . . . 9 . 5

acanthaceous sections. Their relationship is apparently very close-Fox,

Lindsey, Clench & Miller (1965) placed Salamis as a subgenus of Hypolimnas.
Vane-Wright, Ackery & Smiles ( 1977) summarize the hostplant associations of
Hypolimnas. Exploitation of the Urticaceae is particularly widespread, and, apart
from the ubiquitous misippus and bolina, there are very few 'secondary7 hosts (see
Table 51). H. misippus, bolina and alimena alone utilize the Acanthaceae. These
three species, together with diomea, demonstrably form a monophyletic
group (Hiura, 1983), underlining again the artificiality of the main
Acanthaceae/Urticaceae division. Salamis exhibits a similar pattern, with temora,
parhassus and anacardi exploiting the Acanthaceae alone and cacta restricted to
the Urticaceae (Table 5 1).
Within the remaining genera of the Nymphalini (and the Coloburini) there
are no further records for the Acanthaceae: instead the Urticaceae predominate.
In the absence of a reliable generic classification, the following sequence owes
more to convenience than to intuitative insights into nymphaline classification!
. . . . . . . . . . . . . . . . . . . . . . Galycanthaceae
. . . . . . . . . . . . w . . . . . . . . . Calycanthus
. . . . . . . . . . . . . . . . . . . . . . Polygonaceae
A
. . o,
. . . . . . . . . . . . . . . . . . . Polygonurn
. . . . . . . . . . . . . . . . . . . . . .M or ac e ae
2
. . . . . . . . . . . C.
. . . . . . . . . . Artocarpus
. . . . . . . . . . . . . . . . . . . . . . Urticaceae
2
. . . . . . . . . . . . . . . . . . . . . Girardinia
A
. . . . . . . . . . . . . o,.. . . . . . . Urtica
. . . . . . . . . . . . . . . . . . . . . . Fabaceae
. . . . . . . . . . . . . . . . . . . . . Calliandra
A
. . . . . . . . . . . . . E .
. . . . . . . Erythrina
. . . . . . . . . . . . . . . . . . . . . Pithecollobium
. .
. . . . . . . . . . . . . . . . . . . . Asclepiadaceae
A
. .
. . . . . . . . . . 2
. . . . . . . . . Gomphocarpus
. .
. . . . . . . . . . . . . . . . . . . . Verbenace=
. .. . . . . . . . . . . . . . . 0 ) .
. . . Aloysia A
. . . . . . . . . . . . . . . . . o,.
- . . Lippia
. . . . . . . . . . . . . . . . . . . . . . Lamiaceae
. . . . . . . . . . . . . . . . . - . . c . . Melissa
. . . . . . . . . . . . . . . . . . . Mentha 2 . .
. . . c . . . . . . . . . . . . . . . . . . . Salvia
. . . . . . . . . . . . . . . . . . . . . . Scrophulariaceae
A
. . . . . . . . . . . . . . . . . N . . . . Bacopa
2
. . . . . . . . . . . . . . . . . .,. . . . Linderia
. . . . . . . . . . . . . . . . . . . . . . Acanthaceae
2
. . . . . . . . . . . . .VI. . . . . . . . Aeystaaia
2
0 ) .
. . . . . . . . . . . . . . . . . . . . Barleria
2 2 2
. . . ". . . . . . . . . . . . * * , * . Blechurn
2
. . . . . . . . . . . . . . . . . . .,. . . Dicliptera
2
. . . . . . . . . . . . . . . . . . . . . Eranthernurn
2
. . . . . . . . . . . . . . . . . . . Graptophyllurn
. w . . . . . . . . . . . . . . . . . . . . Hemigraphis
N
. . . . . . . . . . . . . . . . . . . . . Hygrophila
.................... - . Jacobinia 2
. . . N . . . . . . . . . . . * . . .,. . . Justicia
2
. . . . . . . . . . . . . . . . . . . . . Lepidagathis
A
. . . . . . . . . . . . ,, . . . . . . . . PSeuderanthernurn
2
W . . Y!Y.
. . . . . . . . . . A . 0 . 2 . Ruellia
2
0
. . . . . . . . . . . . . . . . . . . . . Stephanophysurn
2 Strobilanthua
. . . . . . N o ,2- . .
. . ;k.. . . . . . .
h X B B 3 V 'X 'd 89 1
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 169

TABLE51. Hostplants of the Nymphalinae (Nymphalini, in part). Bascombe et al., 1987';

Fountaine, unpublished2; Gibson-Hill, 19473; Mamet, 19484; Owen & Owen, 19735; Pennington,
19786; Sankowsky, 19787;Sevastopulo, 19758; Vane-Wright et a/., 19779; van Someren, 1974'O;
van Son, 1979"

alimena . . . . . . . . . . . . . . . . . . . . . . . . . . . .
anomla . . . . . . . . . . . . . . . . . . . . . 9 1 . . . . .
antevorta . . . . . . . . . . . . . . . . . . .9 . . . 9 . . . .
antilope . . . . . . . . . . . . . . . . . . . . . 9 . . . 9 . .
bolina . 9 . . 9 . . 9 . . 9 . 9 . . 9 . . 9 9 . 9 . . 9 . . 9
deceptor . . . . . . . . . . . . . . . . . . . 911.. . . . . .
dinarcha . . . . . . . . . . . . . . . . . . .9 . . . . . . . .
dubius . . . . . . . . . . . . . . . . . . .9 . . .9 9 . . .
errabunda . . . . . . . . . . . . . . 9 . . 9 , . . . . . . . . .
misippls . . 9 . 9 9 . . . 9 9 9 . . .9 . . 1 . . . . . . . . .
monteironis. . . . . . . . . . . . . . . . . . . 9 . . . 9 . . . .
salmacis . . . . . . . . . . . . . . . . . . . 9 . . . 9 . . . .
usanbra . . . . . . . . . . . . . . . . . . .9 . . . 9.. . .
Sa Lamis . . . . . . . . . . . . . . . . . . . . . . . . . . . .
anacardii . . . . . . . . . . . . . . . . . . . . . . . . . . . .
augustina . . . . . . . . . . . . . . . . . . . . . . . . . . . .
cacta . . . . . . . . . . . . . . . . . . . . . . . 10 . . . .
parhassus . . . . . . . . . . . . . . . . . . . . . . . . . . . .
tmra . . . . . . . . . . . . . . . . . . . . . . . . . . . .

alimena . . . . .. 7 . . . 7 . . . . . 9 . . . . . . . . . . . .
anomla . 3 . . .. . . . . . . . . . . . . . . . . . . . . . . .
antevorta . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
antilope . . . . .. . . . . . . . . . . . . . . . . . . . . . . .
bolina . . . 9 1 . 7 . . 9 . . . . . . 9 1 9 . 9 . . 9 9 . . . .
deceptor . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
dinarcha . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
dubius . . . . . . . . . . . . . . . . . . . . . . 9 . . . . . .
errabunda . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
misippls . . . 9 . . 9 9 . . . . 9 . . . 9 . 9 . . . . . . . . . 9
monteironis . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
salmacis . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
usanbra . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Sa L ani s . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
anacardi i . . . . . . 10 . . . . 2 8 . 8 8 . . . . . . . . . . . . .
a w u s t ina . . . . . . . . . . . . . . . . . . . . . . . . . . & . .
cacta . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
parhassus . . . . . . 6 . 5 . . 6 . . 8 8 . . . . . . . . . . . . .
ternora . . . . . - 8 . . . . .10 2 1 0 1 0 . . . . . . . . . . . . .
 170 P. R. ACKERY
'I'ABLE52. Hostplants of the Nymphalinae (Nymphalini, in part). Biezanko, 1949I; Howe, 1975?:
Common & Waterhouse, 197Z3; 19814; DeViedrna & Gornez Bustillo, 19765; Dirnork, 197gh;
Dornfeld, 1980'; Ehrlich & Ehrlich, 1961R;Emrnel & Emrnel, 1973y; Gibbs, 19801"; Hayward,
1969"; Iwase, 195412; Kurentsov, 197013; Larsen, 197414; McCubbin, 1971 15; Niculescu, 196516;
Schwarz, 1949"; Shapiro, 197418; ShirBzu, 194419, 196Ozo;Tietz, 1972?l; Wynter Blyth, 195722

atalenta . . . . . . . . . . . . . 21 . 8 . . 8 . 9 . . 8 . 8 . . 16 . . . .
indica . 20 . . . . . . . . . . . 12 . 1922 . . . . . . 19 . 12 . 13 . . . . .
tameama . . . . . . . . . . . . . . . 2 . 2 . 2 . 2 2 . . . . . . . . . .
Cynthia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
annabella . . . 2 . 2 . 2 . 6 2 6 . . . . . . . . . . . 2 . . . . . . 2 . .
carye . . . . 1 1 1 1 8 1 1 8 . . . . . . . . . . . . . 1 . . . . . . 8 . 1
kershaui . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
myrinna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
virginiensis. . .21 . . . 2 . . . . . . . . . . . . . . .21 . . . . . .21 . .
Bassaris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
gonerilla . . . . . . . . . . . . . . . . . . . . . . .10 . . . . . . . . .
i tea . . . . . . . . . . . . . . . . . . . . 4 . . 3 . . . . . . . . .

atalanta . .
. . . . . . . . . . . . 21 . . . . . . 14 17 . 16 . . . .
irdica . .
. . . . . . . . . . . . . . . . . . . . . . . . . . .
tameama . .
. . . . . . . . . . . . . . . . . . . . . . _ . . . . . .
Cynthia . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . .
atmabet La . .
. 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
carye . .
. . . . . . . . . . . . . . . . . . . . . 1 1 11 . . . . . .11
kershaui . .. . . . . .4 . . . . . 3 . . . 3 15 . . 15 . . 3 . 3 3 3 . .
myrinna . .
. . . . . . . . . . 1 . . . . . . . . . . . . . .
virginiensis. 21 . . . 21 2 . . . 2 . 1 . . 9 8 2 1 . a 5 21 . . 821 . . . 7 .
Bassaris . . . . . . . . . . . . . . . . . .. . . . . . . . . . .
gonerilla . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
itea . . . . . . . . . . . . . . . . . . . . . . . . . .

At least Vanessa, Cynthia and Bassaris probably form a monophyletic group-

prior to Field (1971) they were treated as congeneric. Bassaris appears to be
restricted to the Urticaceae, the only family common to all three genera
(Table 52). Within Cynthia, there are further widespread records for the
Malvaceae, closely associated with the Urticales within the Dilleniidae. Also, the
proliferation of asteraceous records is unrivalled elsewhere in the Nymphalidae
(see also Table 5 3 ) . The Asteraceae show affinity with two other noted families,
the Boraginaceae and Oleaceae. Given the widespread distribution of at least
some Cynthia species, the abundance of less believable records is hardly
surprising.
In the Urticaceae, Asteraceae and Ulmaceae, Vanessa hostplants have much
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 171

TABLE 53. Hostplants of Cynthia cardui. Austin & Austin, 1981I ; Bascombe et al., 19872; Chang,
19633; Emmel & Emmel, 1973’; Howe, 19755; Kim, 19846; Leech, 1892’; Neck, 197Ij8;
Niculescu, 19659; Pennington, 1978’O; Schwarz, 1949”; Scott & Scott, 19801*; Sevastopulo,
197313; Shapiro, 197414; Shirtnu, 194415; Tietz, 197216; van Someren, 197417; van Son, 197g1*;
Wynter Blyth, 195719

Apiaceae Asteraceae Fabaceae

16 13
Eryngiun Laggera Argyr~lobiun’~
10
Asteraceae Lappa Doli~hos’~
Achilles"12 Madial3 G 1yc ine’
Anaphal i s Onopordun7 Lablab13
9 Partheniun16 ~upinusl~
Antennar ia
15 pent z ia1
Arctiun Phaseo1us l3
Arctotis13 ~enecio’~ Zorni a’
serratula16 Lamiaceae
Artemisia13
Berkhe a’l sity b ~ n l ~ Stachys”
Blunea
5; Sonchus13 Kalvaceae
stotxea13 A1 thaea13
Ca 1endul a16
Tussi Lago’’ Halva13
~arduus’
Venidi unl’ Sphaeralceal
Car 1 ina2
14 Boraginaceae Smilacaceae
Centaurea
Aminckia14 Smi Lax6
Chry~anthenun’~
Anchusa13 Ulmaceae
cirsiunl3
Cnicus16 ora ago^ ULrmS6
13 CryptanthaO U r t icaceae
Cynara
cynoglossun 17 Boehmeria13
Dinmrphotheca” 19
Echiun” Debregeasia
Filago13
Chenopodiaceae Girardinia13
Gazania”
Gnapha1 im3 Chenopodiun16 ~aporteal~
He1ianthus’ 0txtial7
16
He1ichrysun13 Parietaria
urtical3

in common with Cynthia, while further records indicate the Cannabidaceae and
Salicaceae. The Cannabidaceae, Urticaceae and Ulmaceae are closely associated
within the Urticales, and in turn show some affinity to the Salicaceae (Salicales)
within the Dilleniidae, but are only distantly related to the Asteraceae
(Asterales).
Of the genera outside this small group, Mynes, Araschnia, Rhinopalpa and
Symbrenthia are all seemingly restricted to the Urticaceae, and were it not for a
single record for A. dimorphica on the Asteraceae (Carduus), Antanartia could also
be added to this list (Table 54). Reflecting preferences seen elsewhere, Inachis,
although common on Urtica and even Laportea, is also occassionally noted on the
Cannabidaceae and Ulmaceae, together with the more diverse Rosaceae
(Table 55). Aglais again conforms to the overall pattern, the Asteraceae,
Cannabidaceae, Salicaceae, Ulmaceae and Grossulariaceae being noted as
secondary hosts-otherwise Urtica, and occasionally Laportea are the typical
hosts of the genus (Table 54).
The closely associated Ulmaceae and Urticaceae again recur together in
tandem within Hypanartia (Table 54), while most of the supplementary hosts of
Antanartia, Inachis, Aglais and Hypanartia are included in the extensive foodplant
data for the next genus, Polygonia (Table 56).
Although Sevastopulo (1973) notes JV. antiopa on Urtica, the absence of any
other urticaceous records for JVymphalis is probably significant. Many of the
172 P. R. ACKERY
TABLE 54. Hostplants of the Nymphalinae (Nymphalini, in part). Biezanko, 1949I; Brown &
Heineman, 19722; Carey-Hughes & Pickford, 19773; Common & Waterhouse, 19814; Corbet &
Pendlebury, 19785; D’Abrera, 19776; d’Almeida, 19227; Hayward, 196g8; Hill el al., 19789;
Iwase, 1954IO; Kim, 1984”; Mamet, 1948’*; Manders, 1908”; McCubbin, 1971 I r ; Miiller,
188615;Pennington, 197816; Riley, 1975l’; Sankowsky, 197518; Schwarz, 194919; Scott & Scott,
19802”; Sevastopulo, 197321, 197522;‘I’ilden, 196523; van Someren, 197424;van Son, 197gZ5;
Young, 197626

cashmirensis. . . . . . . . . . . . . . . . . . . . . . . 21 . . . . . . .
milberti . . . . . . . . . . . . . . . . . . . . . . 20 . 23 . 23 . . 23
20
urticae . . . . . . . . . . . . . .
.19 . . . . .10 . . . . . . .
.10
Araschnia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
burejana . . . . . . . . . . . . . . . . . . . . . . 10 . . . . . . .
10
levana . . . . . . . . . . . . . . . . . . . . . . 11 . . . . . . .
11
prorsoides . . . . . . . . . . . . . . . . . . . . . . .21 . . . . . . .
Hypanartia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
bella . . . 1 . 6. . . . . . . . . . . . . . . . . 8 . . . . . . .
kefersteini. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
26
Lethe . . . 1 7. . . . . . 15 . . . . . . . .
. . . . . . . . . . .
paul lus . 1 7 . . . 2 . . . . . . . . . . . . . . .
. . . . . . . . . .
Mynes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
geoffroyi . . . . . . . . . . . . . . 18 . . 14 . . b . . . . . . . . . .
Rh i nopa lpa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
polynice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
21
Synbrenth i a. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
.
hi ppoc lus . . . . . . . . . . . . . . . 6 . . . . . . . . . .
3 . 2 1 21
liLaea . . . . . . . . . . . 9 . 5 . . 5 . . .
. . . . . . . . . . .
Antanartia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
abyssinica. . . . . . . . . . . . . . .
-22 . .22 .24 . . . . . . .
.24
borbonica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
13
delius . . . . . . . . . . 24 . . . . . . . . . .24 .22 . . . . . . .
dimorphica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 6 .
.
h i ppomene . . . . . . . . . . . . . . . . 24 25 16 . . . . . . .
24 12 16 12
schaeneia . . . . . . . . . -2216. . - 1 6 . . . .
. 2 4 , . . . . . . . .

wide-ranging hosts are shared with Polygonia, representatives of the Salicaceae,

Ulmaceae, Cannabidaceae, Betulaceae, Rosaceae, Tiliaceae, Ericaceae and
Grossulariaceae figuring most prominently (Table 55). Curiously, the
Rhamnaceae (Ceanothus), a family lacking obvious affinity with any other
Nymphalis hosts, are the characteristic hosts of N . californica.
Finally, the monobasic Oriental genus Kaniska, classically treated as a true
nymphaline (e.g. Eliot, 1978) surprisingly only takes monocotyledons, mainly
Smilacaceae but also Dioscoreaceae and Liliaceae, obviously a considerable
hostplant shift (Table 55).
So, although the Urticaceae and Acanthaceae trends are almost universal and
hardly overlapping there is considerable variation on these themes---the
Asteraceae, Scrophulariaceae and Plantaginaceae associate with the
Acanthaceae, and the Cannabidaceae, Ulmaceae, Grossulariaceae and
Salicaceae (among others, including the Asteraceae again) with the Urticaceae.
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 173

TABLE 55. Hostplants of the Nymphalinae (Nymphalini, in part). Howarth, 1973'; Iwase, 1954';
Kim, 19843; Kurentsov, 19704; Langer, 195€i5; Niculescu, 19656; Robert et al., 1983'; Schwarz,
1949*; Scott & Scott, 19809; Sevastopulo, 1973IO; Shapiro & Shapiro, 1973"; Tietz, 1972";
Uchida, 198413

io . . . . 3 . 0 . 3 3 . . . . . . . . . .
Kaniska . . . . . . . . . . . . . . . . . . . .
canace . . . . . . . . . . . . . . . . . . .
Wyophalis . . . . . . . . . . . . . . . . . . .
ant iopa - 9 . 9 9 . 9 . 10 . 1 1 9 2 . . . . . 9
californica . . . . . . . . . . . . . .12 . . . .
polych loros . . . 0 5 . . . . . 7 7 . . . . 7 7 5
vau-albun . 4 . . 4 . . . . . 5 0 . . . . . . .
xanthanelas . . .10 . . . . . . . 1 0 . . . . . . .

c
:
d
Inachis . . . . .
io 8 . . . . . . . . . . . . . . . . . .
Kaniska . . . . . . . . . . . . . . . . . . .
canace . . . . . . . . . . . . . 2 2 2 . 1 3 2
Nynphalis . . . . . . . . .. . . . . . . . . . .
anti- 9 . . . . . . . .. . . . . . . . . . .
californica . . . .12 . -12 . . . .
. . . . . . . .
polychloros . . 1 . . . . . . . . . . . . . . . . .
vau-albun . . . . . . 5 . 0 . . . . . . . . . . .
xanthomelas . . . . . . . . . . . .10 . . . . . . .

However, this basic division may owe more to distribution than relationship,
Holarctic species favouring the Urticaceae, with Acanthaceae-feeding
characteristic of the tropics.
The confirmation of palatability in Anartia (Brower & Brower, 1964;
Silberglied, 1983) and Siproeta (Brower & Brower, 1964, as Metamorpha) supports
the classical contention of nymphaline acceptability to avian predators. Few
have been drawn into mimetic complexes; Hypolimnas is an obvious exception.
Here, many species converge on to the patterns of danaines and acraeines, with
both unimodal and $? sex-limited mimicry occurring. Rarely, both H . misippus
and bolina feed on Ipomoea, species of which may contain cardio-active substances
(Marsh, Clarke, Rothschild & Kellet, 1977). Adult Hypolimnas might therefore
serve as co-mimics (but see also Brower, 1984), but the notion that the
Nymphalinae are essentially palatable remains supportable.

Coloburini
O n the basis of foodplants, the New-World tribe Coloburini (see Miller &
Brown, 1981) is hardly separable from the Nymphalini. Seitz (1914) includes
174 P. R. ACKERY
TABLE 56. Hostplants of the Nymphalinae (Nymphalini, in part). de la Maza & de la Maza,
1977'; Dornfeld, 19802;Ehrlich & Ehrlich, 19613; Harris, 1972+; Honda, 19715; Howe, 19756;
Kim, 19847;Larsen, 1974O; Leech, 18929; Manley & Allcard, 1970'"; Niculescu, 1965"; Robert
et al., 198312;Schwarz, 194913;Scott & Scott, 198014;Tietz, 197215;Tilden, 196516

c-album . . . .. . 7 . 1 2 . . . . . 7 . 7 . .10
c-aureum . . . .. . . . . . . . . . . . . . 9 5
ComM . . . .. . . . . . 15 . 15 . 15 . 3 . . 3
egea . . . .. . 0 . . . . . . . . . 0 . . .
faunus . . . 3 3 . . . . . . . . . . .1 5 . . .
gracilis . . . . . . . . . . . . . . . . . . . .
i n t e r r o g a t i o n i s . 15 . . . . . . . . 4 . . . 3 . 3 . . 4
oreas . . . 15 . . . . . . . . . . . . . . . .
progne . . . .15 . . . . . . . . . . 1 3 . .15
satyrus . . . . . . . . . . . . . . . . . . . 14
s i lvius . . . . . . . . . . . . . . . . . . . .
zephyrus . . . . . . . . . . . . . . . . 3 ..I5

c-album . 7 . 12 . . 0 . . . . . 10 . . . 10 . 7 . .
c-aureun . . . . . . . . . . . . . . . . . . . . .
ComM . 3 . 3 . . . . . . . . . . 15 . 15 . . . 15
egea . .i3 8 . . . . . . . . . . . .ii . a . .
faunus . . .15.. 3 . 3 . 1 4 . . . . . 3 . . . .
gracilis . . . . . . . . . . . . . . . . . . . . 15
interrogationis . 3 . 3 . . . . . . . . . . . . . . . . 15
oreas . . . . . . . . 2 . 2 . . . . . 2 . . . .
progne . . . . . . . . . . . . . .15 . 3 .15 . .
satyrus . . . 14 . 15 . . 16 . 15 . . . . . 15 . . . .
s i lvius . . . . . . . . . . 6 . . . . . . . . . .
zephyrus . . . . . . . . 315 3 . . 1 5 . . 3 . . . .

just seven genera. Most known hosts are confined to the Urticaceae, most
commonly Cecropia species (Table 57). No members of this tribe are obviously
involved in mimetic complexes; DeVries ( 1983) found Historis odius relatively
palatable, as did Magpie-jays! I n view of the temperate/tropical segregation of
true nymphalines between the Urticaceae and Acanthaceae, exploitation of
urticaceous hosts by this typically tropical group is of particular interest.

Apaturinae
O n the basis of larval morphology, Clarke (1947) grouped the Charaxinae
and Apaturinae as a single family, the Apaturidae, the system followed by
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 175

TABLE57. Hostplants of the Nymphalinae (Coloburini). d’Araujo e Silva el al., 1968’; DeVries,
19862; Hayward, 196g3; Muyshondt & Muyshondt, 1976+, 19785, 197g6;Tietz, 1972’

acesta . l . 2 . . . . . . . . . . . .
Coea . . . . . . . . . . . . . . . .
acheronta . . . 6 . . . . . . . . . . . .
Colobrra . . . . . . . . . . . . . . . .
d i rce . . . 4 . . . . 3 3 . 3 . 1 . .
Historis . . . . . . . . . . . . . . . .
odius . . . 6 . . . . . . . . . . . 7
Smyrna . . . . . . . . . . . . . . . .
blmfildia . . . . 3 5 5 . . . . . . . . .
karuinskii . . . . . 5 5 . . . . . . . . .

Miller & Brown (1981). However, Rydon (1971) notes differences between the
form of the eggs in the two subfamilies-he retains them as distinct-and the
prelimanary analysis of DeVries et al. (1985) did not suggest particularly close
relationship. Le Moult (1950) recognized 18 Old-world apaturine genera to
which must be added the American Asterocampa and Doxocopa (Miller & Brown,
1981), the Middle-eastern Euapatura (Ebert, 197 1 = Protapatura Shirbzu and
Saigusa, 197 l ) , and probably the Oriental Timelaea.
To date, nearly all species appear restricted to the Ulmaceae (mainly Celtis;
see Table 58). The ‘true’ Apaturu and Sephisa go against this generalization,
Apatura iris and ilia favouring the Salicaceae and Betulacaceae, while the
Fagaceae characterize Sephisa. As usual, questionable hosts abound. Of the more
convincing, the Ulmaceae (Urticales) and Salicaceae (Salicales) are somewhat
disparate members of the Dillenidae, while the Fagaceae, Betulacaceae and
Carpinaceae (Fagales) form a close assemblage within the Hamamelidae.
A few Oriental apaturines, particularly Diagora and Hestina species, plausibly
mimic danaines of the genera Parantica or Tirumala (Morishita, 1980); elsewhere
female forms of Idrusia (‘Euripus’) nyctelius closely resemble Euploea species
(Nguyen, 1985). Additionally, some members of the South American genus
Doxocopa are superficially similar to limentine Adelphas. Both are said to be
indistinguishable in flight (Brown & Heineman, 1972)) but this presumed
mimetic association has no known chemical basis.

Libytheinae
Although still widely considered as a separate family (e.g. Ehrlich, 1958;
Aoki, Yamaguchi & Uemura, 1982; Scott, 1985), Kristensen (1976) has
demonstrated that the ‘Nymphalidae minus Libytheinae’ are almost certainly
paraphyletic-the libytheines represent an off-shoot from within the family
rather than its sister-group.
Pagenstecher’s ( 1901) outdated revision (supplemented by Shields, 1985) still
provides the taxonomic basis for the subfamily-he recognized ten species
I76 P. R. ACKERY

I'AHLE 58. Hostplants of the Apaturinae. Austin, 1977l; Biezanko, 194g2; Comstock, 19433;
Corhrt & Pendlebury, 19784; d'hraujo e Silva et al., 19685; Hayward, I96g6; Hill et al., 1978';
Kim, 1984H;Kurentsov, 1970'; Manley & Allcard, 197010;Muller, 1886"; Muroya et al., 1967";
Niculcscu, 1965"; Schwarz, 194914; Scott & Scott, 198015; Tietz, 1972Ifi; Uchida, 1984";
Wynter Blyth, 1957IH

Apat ur a . . . . . . . . . . . . . . . . . . . . . . . .
ilia . . . . .I0 . . . . . . . . . . . .lolo . . . .
iris . . . . .13 . . . . . . . . . . . .1414 . . . .
Asterocampa . . . . . . . . . . . . . . . . . . . . . . . .
argus . . . . . . . . . . 3 . . . . . . . . . . 3 . .
celtis . . . . . . . . . .I5 . . . . . . . . . . . . . .
clyton .I6 . . . . . . . -16 . . . . . . . . . . . .16
Lei La . . . . . . . . . . 1 . . . . . . . . . . . . .
Bremeria . . . . . . . . . . . . . . . . . . . . . . . .
nycteis . . . . . . . . . . . . . 9 . . . . . . . . . .
schrenki i . . . 9 . . . . . . . . . 9 a . . . . . . . . .
Chi t o r i a . . . . . . . . . . . . . . . . . . . . . . . .
c h r y s olora . . . . . . . . . . 12 . . . . . . . . . . . . .
ulupi . . . . . . . . . . a . . . . . . . . . . . . .
Diiipa . . . . . . . . . . . . . . . . . . . . . . . .
f e n e s tra . . . . . . . . . . a . . . . . . . . . . . . .
Doxocopa . . . . . . . . . . . . . . . . . . . . . . . .
cyane . . . . . . . . . . 6 . . . . . . . . . . . . .
k a l L ina . . . . . . . . . . 2 . . . . . . . . . . . . .
l a u re . . . . . . . . . . 5 . . . . . . . . . . . . .
laurentia . . . . . . . . . . 6 . . . . . . . . . . . . .
Lauret t a . . . . . . . . . . 11 . . . . . . . . . . . . .
selina . . . . . . . . . . 5 . . . . . . . . . . . . .
Eulaceura . . . . . . . . . . . . . . . . . . . . . . . .
osteria . . . . . . . . . . . 4 . . . . . . . . . . . .
Euripus . . . . . . . . . . . . . . . . . . . . . . . .
consimilis . . . . . . . . . . . . 18 . . . . . . . . . . .
nyctelius . . . . . . . . . . . . . . . 4 . . . . . . .
Helcyra . . . . . . . . . . . . . . . . . . . . . . . .
p lessen i . . . . . . . . . . 17 . . . . . . . . . . . . .
superba . . . . . . . . . . 17 . . . . . . . . . . . . .
Hest ina . . . . . . . . . . . . . . . . . . . . . . . .
assimi l i s . . . . . . . . . . 17 . . . . . . . . . . . . .
japonica . . . . . . . . . . a . . . . . . . . . . . . .
persimilis . . . . . . . . . .la. . . . . . . . . . . . .
Rohana . . . . . . . . . . . . . . . . . . . . . . . .
p a r i s a t is . . . . . . . . . . 7 . . . . . . . . . . . . .
Sasakia . . . . . . . . . . . . . . . . . . . . . . . .
charonda . . . . . . . . . a a . . . . . . . . . . . . .
Seph isa . . . . . . . . . . . . . . . . . . . . . . . .
d i c hroa . . . . . . . 9 . . . . . . . . . . . . . . . .
pr inceps . . . . . . . 8 . . . . . . . . . . . . . . . .
Thaleropis . . . . . . . . . . . . . . . . . . . . . . . .
ioni a . . . . . . . . . . 11 . . . . . . . . . . . . .
T imelaea . . . . . . . . . . . . . . . . . . . . . . . .
macu l a t a . . . . . . . . . . 12 . . . . . . . . . . . . .
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 177

59. Hostplants of the Danainae (Danaini; Amaurina). Ackery & Vane-Wright, 1984’
TABLE

aglea . . . . . . . . . . 1 . . . . . . . . 1 . 1 . . . . .
agleoides . . . . . . . . . . . . . 1 . . . . . . . . . . . . 1
schenkii . . . . . . . . . . . . . . . . . . . . . . . I . . .
aspasia . . . . . . . . . . . . . 1 . . . . . l . . . . . . .
sita . . . . . . . . . 1 . . 1 . 1 1 1 . . l . . . . . . .
melaneus . . . . . . . . . . . . 1 . . . . . . . . . . . . . .
taprobana . . . . . . . . . . . 1 . . . . . . . . . . . . . 1 .
luzonensis . . . . . . . . . . . . . . .1 . . . . . . . . . . .
ldeopsi s . . . . . . . . . . . . . . . . . . . . . . . . . . .
juventa . 1 . . . . . . . . . . . 1 . . . 1 . . . . . . . . .
similis . . . . . . . 1 . . . . . . . . . . . 1 . . . . . . .
vulgaris . . . . . . . . . . . . . 1 . . . . . . . . . . . . .
gaura . . . . . 1 . . . . . . . . . . . . . . . . . . . . .
Amauris . . . . . . . . . . . . . . . . . . . . . . . . . . .
niavius . . . . . . . . . . . . 1 1 . 1 . . 1 1 . . . . . . .
tartarea . . . 1 . . . . . . . . . . . . . . . . . . . . . . .
ellioti . . . . . . . . . . . . 1 1 . 1 . . 1 1 . . . . . . .
phoedon . . . . . . 1 . . . . . . . . . . . . l . . . . . . .
cmrana . . . . . . . . . . . . . . . . . . . 1 . . . . . . .
echeria . . . . . . . . . . . . 1 1 . 1 . . 1 1 . . . . . . .
craushayi . . . . . . . . . . . .1 1 . 1 . . 1 1 . . . . . . .
albimaculata . . . . . . . . . . . . 1 1 . 1 . . 1 1 . . . . . . .
ochlea . . . . . . . . . . . . 1 1 . 1 . . 1 1 . . . . . . .

within a single genus, Libythea. Michener (1943) proposed the generic name
Libytheana for the New-World species, a convention now generally accepted (see
Miller & Brown, 1981). The world-wide distribution of the subfamily is unusual;
there is representation in each major biogeographic region. Few higher butterfly
groups have such low species diversity and such a cosmopolitan range.
To date, all species have been found to feed on Cellis (Ulmaceae); Libythea
g e o f k y is said to also exploit the Sapindaceae and Lauraceae, with Libythea
bachmani recorded on the Caprifoliaceae. Few investigations offer insights into
palatability-in a single experiment Swynnerton ( 1919) notes ready acceptance.
Neither are they obviously involved in mimetic complexes, with the possible
exception of convergence with Pantoporia (see Longstaff, 1912). Cyanogenesis has
been confirmed in certain Celtis species (others gave negative results); tannins
are reportedly absent although saponins may be present (Darnley Gibbs, 1974).

Danainae
Together, the Danainae, Ithomiinae and Tellervinae form a monophyletic
group, although at present the trichotomy appears unresolvable (Ackery &
Vane-Wright, 1984). The danaines divide into two tribes, the Danaini,
convincingly established, and a more tentative grouping, the Euploeini. Even a t
tribal level there are marked host preferences, with euploeines favouring
178 P. R.ACKERY

TABLE
60. Hostplants of the Danainae (Danaini; Danaina). Ackery & Vane-Wright, 1984'

formosa . . . . . . . . . . . . . . . . . . . . . . . .
pet iverana . . . . . . . . . . . . . . . . . . . . . . . .
linniace . . 1 . . . . . . . l . . . . . . . . . . . . .
septentrionis . I . . . . . . . . . . . . . . . . . . . . . 1
hamata . . . . . . . . . . . . . . . . . . . . . . 1 .
Danaus . . . . . . . . . . . . . . . . . . . . . . . .
cleophi l e . . . . . . . . . . . . . . . . . . . . . . . .
plexippus . . . . . . 1 . . . . . 1 . . . 1 . . . 1 . . .
erippus . . . . . . . . . . . . . . . . . . . . . . . .
genut ia . . . . . . . . . . . . . . . . . . . . . . . .
af f i n i s . . . . . . . . . . . . . . . . . . . . . . . .
melani ppus . . . . . . . . . . . . . . . . . . . . . . . .
eresiws . . . . . . . . . . . . . . . . . . l . . . . .
plexaure . . . . . . . . . . . . . . . . . . . . . . . .
gilippus . . . . . . . . . . . . . . . . . . . . . 1 . .
chrysippus . . . . 1 . . . 1 . . . 1 . 1 . . . . . . . . .

formsa . . . . . . . . . . . . . . . . . . . . . . . . . . .
petiverana . . . . . . . . 1 . . . . . . . 1 . . . . . . 1 . . .
linniace . . . 1 . 1 . . . 1 . . . . . 1 1 . . . . . . l . . .
septentrionis . . . . . . . . . . . . . . . 1 . . . . . . . . . . .
hamata . . . . . . . . . . . . . . . . 1 . . . . 1 . 1 . . .
Danaus . . . . . . . . . . . . . . . . . . . . . . . . . . .
cleophile . . . 1 . . . . . . . . . . . . . . . . . . . . . . .
plexippus . 1 1 1 . 1 . . . . . 1 . . . . . . . . . . . 1 1 . .
erippus . . 1 1 . . . . . . . . . . . . . . . . . . . . . . 1
genut ia . . . 1 . . 1 1 . . . . . 1 1 . . . . . . . . l . . .
af f i n i s . . . . . . . 1 . . . . . . . . . . . l . . . . . . .
me1ani ppus . . . . . . . . . . . . . . 1 . . . . . . . . . . . .
eresiws . . . 1 . 1 . 1 . . . . . . . . . . . . . . . . . . .
plexaure . . . 1 . . . . . . . . . . . . . . . . . . . . . . .
gilippus . . . 1 . 1 . . . . 1 . 1 . . . . . 1 . . . . . . . .
chrys i ppus . . . 1 1 1 . 1 . . . 1 . . . . . 1 . 1 1 1 1 1 . 1 .

Moraceae and Apocynaceae, while the danaines most heavily exploit the
-4sclepiadaceae with just occasional apocynaceous records.
Amauris, Ideopsis and Parantica, which together comprise the subtribe
Amaurina (but see also Kitching, 1985), commonly feed on Asclepiadaceae,
with Tylophora, Marsdenia, Cynanchum, Gymnema and Secamone particularly
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 179

TABLE
60. Continued

formosa . . . . . . . . . . . 1 . . . . . . 1 . 1 . . . .
petiverana . . . . 1 . . . . . . . . . . . . . . . . . . . .
linniace . . . . . . . . . . . . . . . . . . . . . . . . .
septentrionis. . . . . . . . . . . . . . . . . . . . . . . . .
hamata . . . . . . . . . . . 1 . . . 1 . . . 1 . . . . .
Danaus . . . . . . . . . . . . . . . . . . . . . . . . .
cleophile . . . . . . . . . . . . . . . . . . . . . . . . .
plexippus 1 . . . . . . . . . . . 1 . . . . . . . . . 1 . .
erippus 1 . . . . . . . . . . . . . . . . . . . . . . . .
genutia . . . . 1 . . 1 . . . . . . 1 1 . . . . . . . . .
aff inis . . . . . . . . . . . . . . . l l . . . . . . . .
melanippus . . . . . . . . . . . . . . . 1 . . . . . . . . .
eresinus . . . . . . . . . . . . . . . . . . . . . . . . .
plexaure . . . . . . . . 1 . . . . . . . . . . . . . . . .
gilippus 1 . . . . 1 . . . 1 . . 1 . . . 1 . . . . . . . .
chrysippus . 1 1 1 1 . 1 . . . 1 1 1 1 . 1 . . . . 1 . 1 . 1

favoured. Most other listed families (see Table 59) probably need confirmation.
A single Parantica record represents the Peripiocaceae-records for this family,
closely associated with the Asclepiadaceae, are rare but widespread throughout
the Danainae.
Three further genera, Danaus, Tirumala and Tirudelphe, constitute the subtribe
Danaina, a largely pan-tropical group most diverse in the Orient. At present,
the rare and monobasic Tirudelphe lacks hostplant records but both Danaus and
Tirumala are found on much the same asclepiad genera as the Amaurina, with
Danaus itself additionally favouring species of Asclepias, Calotrupis and Perguluria,
genera not heavily exploited elsewhere in the Danaini. Scattered records
represent several further families, most importantly the Apocynaceae.
Periplocaceae and Convolvulaceae (Table 60).
Nothing is known of the early-stage biology of the New Guinea endemic
Prutoploeu apatela. However, the other two genera of the subtribe Euploeina- -
Eupluea and Idea--are familiar Oriental butterflies. Eupluea species exploit the
Moraceae, particularly Ficus, although other genera have also been noted
(Table 61). Again the Apocynaceae are well represented but, with the exception
of E. cure, few species exploit asclepiads. I n comparison, Idea species, despite
extensive data, have never been recorded on the Moraceae: known hosts divide
between the Asclepiadaceae and Apocynaceae.
The smallest subtribe, the Neotropical Itunina, comprises Lycurea and Anetia,
in all just eight species. Lycoreas reliably exploit members of the Caricaceae and
Moraceae (Table 61 ); records for Asclepiadaceae and Myoporaceae seem more
doubtful. Host data for Anetia remains very scanty-A. briarea possibly feeds on
Jacquinia (Theophrastaceae) and thirra apparently takes Metastelma
(Asclepiadaceae).
Despite the long-standing assumption of‘danaine unpalatability (see Muller,
180 P. R. ACKEKY

rrABLE 61. Hostplants of the Danainae (Euploeini; Euploeina, Itunina). Ackery & Vane-Wright,
1984'; Scheermeyer & Zalucki, 19852; Uchida, 19843

euphon . . . . . . 1 . . . .
. . . . . . . . . . . . . . 1 . . . . .
Sylvester . . . . . . 1 . . . .
. . . . . . . . . . . . 1 . . . . . . .
rrulciber . 1 . . . . 1 . . . . . . . . . . . . . 1 . 1 . . . . .
1 . .
phaenareta . . . . . . . . . . . . . . . . . . . . . 1 . . . . . 1 . . .
midams . . . . . . 1 . . . . . . . . . . . . . . . . . . 1 . . . 1 .
eunice . . . . . . 1 . 1 . . 1 . . . . . . . . . . . . . . . . . . .
klugii . . . . . . 1 . 1 . . . . . . . . . . . . . . . . . . . . . .
tulliolus . . . . . 1 . 1 . . . . . . . . . . . . . . . . . 1 . . . . .
darchia . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . .
boisduval ii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
algea . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . .
core . . . 1 . . 1 . 1 . . . . . . . . . . 1 1 . 1 1 1 1 1 1 1 . 1
alcathoe . . . . . . 1 . . . . . . . . . . . . . . . . . 1 1 . . . . .
treitschkei . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . .
modesta . . . . . . 1 . . . . . . . . . . . . . . . . . . 1 . . . . .
crameri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
camaralzeman.. . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 .
uallacei . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . .
batesii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
leuinii . . . . . . 1 . . . . . . . . . . . . . . . . . . 1 . . . . .
nechos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
mitra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Idea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
hypermestra.. . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . .
leuconoe . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . .
lynceus . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . .
iasonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
stolli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
malabarica . . . . . . . . . . . . . . . . . .I . . . . . . . . . . . .
Lycorea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
cleobaea . . . . . . 1 . . . . . . 1 1 . . . . . . . . . . . . . . . .
ilione . . . . . . 1 . . . . . . 1 1 . . . . . . . . . . . . . . . .
Anetia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
thirza . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
briarea . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . .

1878, 1879; Eltringham, 19lo), chemical support remained unforthcoming until

Reichstein identified cardenolides in adult D.plexippus and chrysippus and one of
their larval hosts, Asclepias curassavica (Reichstein, 1967; Reichstein, Euw,
Parsons & Rothschild, 1968). Since then, most considerations of danaine
unpalatability focused on these compounds (see Brower, 1984, for a review).
However, current hostplant records (Ackery & Vane-Wright, 1985) suggest
such heart poisons may not be universal features. Indeed, in the Danaini, only
Danaus itself commonly feeds on plants known to contain cardenolides,
especially Asclepias, Calotropis and Pergularia species, but even their presence is no
guarantee of sequestration and storage (Rothschild & Marsh, 1978). Little is
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 181

TABLE
6 1. Continued

. . . . . . . . . . . . . . . . . . . . . . . . . . euphon
. . . 3 . . . . . . . . . 1 . . . . . . . 1 . . . . sylvester
. . . . . . . . . . . . . . . . . . . . . . . . . . rrulciber
. . . . . . . . . . . . . . . . . . . . . . . . . . phaenareta
. . . . . . . . . . . . . . . . . . . . . . . . . . midamus
. . . . . . . . . . . . . . . . . . . . . . . . . . eunice
. . . . . . . . . . . . . . . . . . . . . . . . . . klugii
. . . . . . . . . . . . . . . . . . . . . . . . . . tulliolus
. . . . . . . . . . . . . . . . . . . . . . . . . . darchia
. . . . 1 . . . . . . . . . . . . . . . . . . . . . boisduvalii
. . . . . . . . . . . . . . . . . . . . . . . . . . algea
. 1 2 . 1 1 1 1 . . 1 1 1 1 . . 1 1 1 1 . . . . . 1 core
. 1 . . . . 1 . . . . . . . . . . 1 . . . . . . .
1 alcathoe
. . . . . . . . . . . . . . . . . . . . . . . . .
1 treitschkei
. . . . . . . . . . . . . . . . . . . . . . . . . . nwdesta
. . . . . . . . . . . . . . . . . . . . . . . . . . crameri
camaralzeman
. . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . uallacei
. . . . . . . . . . . . . . . . . . . . . . . . . . batesii
. . . . . . . . . . . . . . . . . . . . . . . . . . leuinii
. . . . 1 . . . . . . . . . . . . . . . . . . . . . nechos
. . . . . . . . . . . . . . 1 . . . . . . . . . . . mitra
. . . . . . . . . . . . . . . . . . . . . . . . . . Idea
hypernnestra
. . . . . . . . . . . . . . . . . . . . . . . . . .
. . 1 . . . . . . . . . . . 1 . . . . . . . . . . . Leuconoe
. . . . . . . . . . . . . . . . . . . . . . . . . . lynceus
. . . . 1 . . . . . . . . . . . . . . . . . . . . . iasonia
. . . 1 . . . . . . . . . . . . . . . . . . . . . . stolli
. . . . . . . . . . . . . . . . . . . . . . . . . . malabarica
. . . . . . . . . . . . . . . . . . . . . . . . . . Lycorea
. 1 . . . . . . . . . . . . . . . . . . . . . . . . cleobaea
. . . . . . . . . . . . . . . . . . . . . . . 1 . . ilione
. . . . . . . . . . . . . . . . . . . . . . . . . . Anetia
. . . . . . . . 1 1 . . . . . . . . . . . . . . . . thirza
briarea
. . . . . . . . . . . . . . . . . . . . . . . . . .

known of the ability of euploeine species to store cardenolides. Only Euploea core
has been studied in detail (Malcolm & Rothschild, 1983)-it seemingly lacks
cardenolides in the adult and pupal stages even when reared on Nerium oleander,
a plant rich in such compounds.
Attention has now switched toward the probable role of pyrrolizidine
alkaloids in danaine unpalatability (Conner et al., 1981; Eisner, 1980; Brower,
1984). These compounds, taken up by adults, are known as precursors of certain
male pheromones (Bopprt, 1984). However, danaine caterpillars are highly
aposematic, suggesting that either sequestration of some protective compounds
from the larval hosts does occur or that de nouo synthesis is involved.
I82 P. R. ACKERY

TABLE
62. Hostplants of the Tellervinae and Ithomiinae (Tithoreini, Methonini, Melinaeirii,
Mechanitini and three further proposed tribal groupings- see Drummond & Brown, 1987).
Ackery, 1987’; DeVries, 19862; Drummond, 19763; Drummond & Brown, 19874; Haber, 19785;
Young, 1978b6

zoilus . . . . . . 1 . . . . . . . . . . . . . . . . . .
Elzunia . . . . . . . . . . . . . . . . . . . . . . . . .
humboldt . 4 . . . . . . . . . . . . . . . . . . . . . . .
pavdnii . . . . . . . . 4 . . . . . . . . . . . . . . . .
Tithorea . . . . . . . . . . . . . . . . . . . . . . . . .
tarricina . . 3 . 5 . . . 5 . . . . . . . . . . . . . . . .
harmonia . . 3 4 . 4 . 4 4 4 3 . . . . . . . . . . . . . .
Aeria . . . . . . . . . . . . . . . . . . . . . . . . .
eurimedea . . 3 . . 4 . . 6 . . . . . . . . . . . . . . .
olena . . . . . . . . 4 . . . . . . . . . . . . . . . .
elara . . 4 . . . . . . . . . . . . . . . . . . . . . .
Athesis . . . . . . . . . . . . . . . . . . . . . . . . .
clearista . . . . . . . . . . . . . . 4 . . . . . . . . . .
Hethona . . . . . . . . . . . . . . . . . . . . . . . . .
confusa . . . . . . . . . . . . . 3 . . . . . . . . . . .
megisto . . . . . . . . . . . . . 4 . . . . . . . . . . .
singularis. . . . . . . . . . . . . 4 . . . . . . . . . . .
themisto . . . . . . . . . . . . . 3 . . . . . . . . . . 3
Placidula . . . . . . . . . . . . . . . . . . . . . . . . .
euryanassa . . . . . . . . . . . . 4 . . . 3 3 . . . . . . .
Olyras . . . . . . . . . . . . . . . . . . . . . . . . .
crathis . . . . . . . . . . . . . . . . . . . . . 4 . 4 .
Eutresis . . . . . . . . . . . . . . . . . . . . . . . . .
hyperia . . . . . . . . . . . . . . . . . . . . . 4 . 4 .
Melinaea . . . . . . . . . . . . . . . . . . . . . . . . .
meme . . . . . . . . . . . . . . . . . . . . . 4 . . .
ludovica . . . . . . . . . . . . . . . . . . . . . 4 . . .
lilis . . . . . . . . . . . . . . . . . . 4 . . 2 . 4 .
marsaeus . . . . . . . . . . . . . . . . . . . . . 4 . . .
menophilus.. . . . . . . . . . . . . . . . . 4 . . 4 . . .
ethra . . . . . . . . . . . . . . . . . . 2 . . . . . .
Thyrldla . . . . . . . . . . . . . . . . . . . . . . . . .
psidi i . . . . . . . . . . . . . . . . 3 . . . . . . . .
Sais . . . . . . . . . . . . . . . . . . . . . . . . .
rosalia . . . . . . . . . . . . . . . . . . . 4 . . . . .
Scada . . . . . . . . . . . . . . . . . . . . . . . . .
zibia . . . . . . . . . . . . . . . . . . . . . . . . 3
batesi . . . . . . . . . . . . . . . . . . . . . . . . 4
reckia . . . . . . . . . . . . . . . . . . . . . . . . 4
Forbestra . . . . . . . . . . . . . . . . . . . . . . . . .
olivencia.. . . . . . . . . . . . . . . . . . . . . . . 3
Hechani t i s . . . . . . . . . . . . . . .
. . . . . . . .
polymia . . . . . . . . . . . . . .
. . 3 . . . 4 . . . 3
menapis . . . . . . . . . . . . . . .
. . . . . . . . . 3
mazaeus . . . . . . . . . . . . . . .
. . . . . . . . . 3
doryssus . . . . . . . . . . . . . . . . . . . . . . . . 4
lysinmia . . . . . . . . . . . . 4 3 . 3 3 3 . . 3 . 4 . 3
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 183

'I'ABLE 63. Hostplants of the Ithomiinae (Oleriini, Napeogenini). Brown, 1980'; DeVries, 19862;
Drummond, 19763; Drummond & Brown, 19874; Haber, 19785; Young, 197g6

v i r g i n iana . . . . . . . . . . 5 5
egra . . . . . 4 . . . . . .
Oleria . . . . . . . . . . . .
v i c t o r i ae . . . . 5 . . . . . . .
vicina . . . . 5 . . 5 . . . .
makrena . . . . . . . 4 . . . .
a g a r is t a . . . . 3 . . 3 . . . .
janar i1 l a . . . . . . . 4 . . . .
zelica . . . . . . . 3 . . . .
rubescens . . . . . . . 5 . . . .
astrea . . . . . . . 4 . . . .
aquat a . . . . . . . 4 . . . .
crispinilla . . . . . . . 4 . . . .
Epityches . . . . . . . . . . . .
eupompe . 4 4 3 . . 4 . 3 . . .
Napeogenes . . . . . . . . . . . .
t o lo s a . . . . 5 . . 5 . . . .
duessa . . . . 4 . . . . . . .
sylphis . . . . 3 . . . . . . .
inachia . . . . 3 . . . . . . .
sulphurina . . . . 4 . . . . . . .
Rhodussa . . . . . . . . . . . .
cant obr ica . . . . 4 . . 4 . . . .
Carsauritis . . . . . . . . . . . .
xanthostola . . . . . . . 4 . . . .
Hyalyr is . . . . . . . . . . . .
o u l it a . . . . . . . 4 . . . .
excelsa . . . . . . . 5 . . . .
Hypothyris . . . . . . . . . . . .
ninonia . . . . . . . 3 . . . .
f Luonia . . . . . . . 3 . . . .
semifulva . . . . . . . 3 . . . .
daphnis . . . . . . . 1 . . . .
mamercus . . . . . . . 4 . . . .
leprieuri . . . . . . . 4 . . . .
euclea . . . . . . . 6 . . . .
ly c a s t e . . . . . . . 2 . . . .

r~~~~iinae
The exclusively Neotropical Ithomiinae feed largely on the Solanaceae,
particularly Solanum and Cestrum. Exceptionally, species of Aeria, Tithorea, Elzunia
and possibly Melinaea, take apocynaceous hosts, while records for the
Gesneriaceae (Haber, 1978) remain unconfirmed. Brown ( 1977) tentatively
identifies the host of Melinaea as Forsteronia (Apocynaceae)-in more recent
papers, Brown (e.g. 1985) favours Drummond's (1976) Juanulloa (Solanaceae)
record. DeVries ( 1986) has confirmed solanaceous hostplants for Melinaea
species, rather surprising given the aposematic danaoid larva (Brown, 1977);
181 P R . ACKERY

‘1 A R L F 64. Hostplants of the Ithomiinac (Ithomiini, Dircennini). DeVries, 1986’; Drummond,

1976L;Drummond & Brown, 1987?; Haber, 1978+;Young, 1973b5

Miraleria . . . . . . . .
v)
. . . .
cymothoe . 3 3 . . 3 3 . . . . .
Ithmia . . . . . . . . . . . .
diasia . . . . . . . 2 . . . 4
patilla . . . . 2 . . 4 . . . 4
Lichyi . . . . 3 . . . . . . .
amarilla . . . . . 3 . . . 2 . .
celemia . . . . 1 3 . . . . . 4
iph ianassa . . . . . 3 . . . . . 3
heraldica . 1 . . 2 3 . . . . . 1
xenos . 4 . . . 3 . . . . . 4
drymo . 3 . . . . . . . . . .
agnosia . 3 . . . .
2 . 3 . . .
derasa . . . . . . . . 3 . . .
Callithmia . . . . . . . . . . . .
hezia . . . . . . . 3 . 4 . .
lenea . . . . . . . . . 3 . .
Velamysta . . . . . . . . . . . .
cruxi f era . . . . . . . 3 . . . .
H ye l e m a . . . . . . . . . . . .
pascua . . . . . . . . . 3 . .
D i rcenna . . . . . . . . . . . .
adi na . . . . . . . . . 3 . .
loreta . . . . . . . . . 3 . .
klugii . . . . . . . . . 2 . .
relata . . . . . . . . . 5 . .
olyras . . . . . . . . . 3 . .
jemima . . . . . . . . . 3 . .
dero . . . 2 . . . . . 2 . .

such caterpillars are usually associated with the Apocynaceae, while solanaceous
larvae are essentially cryptic.
Assuming the tribes proposed by Fox (1961) represent meaningful groups, it is
disappointing that neither of the two major foodplant-associated groupings,
those centred on the Apocynaceae and Solanaceae, constitute monophyletic
groups (Brown, 1985; Drummond, 1986). However, the classification of the
Ithomiinae is currently under review (Drummond & Brown, 1987). Their
modifications particularly effect the Tithoreini and Oleriini. A new tribe is
proposed for the apocynaceous Aeria (previously placed by Fox in the Oleriini),
while the Tithoreini are now restricted to the apocynaceous Elzunia and
Tithorea. Elsewhere, most records for the tribe Godyrini seem to be centred on
the solanaceous genus Cestrum (Table 65). Methona, the only genus of the
Methonini, exploits Brunfelsia (Table 62), with species of the Ithomiini
favouring Witheringia, Capsicum and Cuatresia (Table 64). Apart from the
Melinaeini, records for other ithomiine tribes suggest a range of solanaceous
hosts (Tables 62-65), but most particularly Solanurn. However, the genera
Markea, Solandra and Juanulloa are characteristic of the Melinaeini (Table 62).
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 185
TABLE65. Hostplants of the Ithomiinae (Godyrini, and a further proposed tribal grouping- see
Drummond & Brown, 1987). DeVries, 1986’; Drummond, 19762;Drummond & Brown, 19873;
Haber, 19784; Young, 1974a5, 1978a6

zygi a . . . 1 . . MS0 . . . . . 3
gonussa . . . . . 3 poecila . . . . . 2
zavaleta . . . 2 . 2 tutia . . . . . 1
kcdcnra . . . . .3 Ceratiscada . . . . . .
Pseudoscada . . . . . . canaria . . . . . 2
tinna . . . 2 . . hymen . . . . . 3
erruca . . . . . 3 Prittuitzia . . . . . .
Hyponenitis . . . . . . hymenaea - 2 . 2 . 2
alphesiboea . . . 3 . . Episcada . . . . . .
libethris . . . 3 . . salvinia . . . . . 1
Greta . . . . . . clausina . . 2 . . .
polissena . . . 4 . . Pteronymi a . . . . . .
mrgane . . . 3 . . artena . . . . 1 3
andrmica . . . . . 3 sinplex . . . . . 4
nero . . . 2 . . cotyt to . . . . . 4
annette . . . 4 . 4 agalla . . . . . 1
ncclungia . . . . . . notilla . . 5 5 . 2
salonina . . . 3 . . carlia . . . 3 . 2
Hypoleria . . . . . . fulvescens . . . . . 4
cassotis . . . 4 . 6 fulvimargo . . . . . 3
malea . . . 2 . . latilla . . . . . 3
orolina . . . 3 . . vestilla . . . . . 2
adasa . . . 3 . . hemixanthe . . . . . 3
Heterosais . . . . . . euritea . . . . . 3
nephele . . . 3 . .
edessa . . . 2 . .

Ithomiine unpalatability is fundamental to the historical development of

mimicry theory, from Bates (1862) and Muller (1878, 1979) onwards. Adult
ithomiines, with their aposematic life-style, participate in co-mimetic rings with
sylvaniform heliconiines, acraeines and troidines, as well as being models for
various supposed ‘Batesian’ associations. With the Solanaceae containing a wide
variety of potentially distasteful compounds, including alkaloids, steroidal bitter
principles, saponins and coumarins (Brown, 1985), it is curious that efforts to
isolate such compounds (or their derivatives) from adult ithomiines have met
with universal failure. Brown concludes that solanaceous larval hosts contribute
minimally toward ithomiine unpalatability, the principal protection being
derived from pyrrolizidine alkaloids sequestered by the adults (Masters, 1968;
Negishi, 1971; Pliske, 1975). The obviously cryptic nature of the Solanaceae-
feeding larvae adds further support to this suggestion, although presumably the
aposematic apocynaceous larvae are deriving some protection from their hosts.
But what of Melinaea? Why is its caterpillar aposematic when it apparently feeds
on Solanaceae?
186 P. R. ACKERY

Tellervinae
The concluding subfamily, the Tellervinae, comprises a single genus, Tellervo,
found in the Old-world tropics from Maluku to the Solomons. Although usually
included within Ithomiinae (Fox, 1956), Ackery & Vane-Wright (1984)
preferred to maintain the Danainae-Ithomiinae-Tellervinae trichotomy.
Widely treated as monobasic (Fox, 1956; Comon & Waterhouse, 1981), there
are in fact as many as six species, some of which are broadly sympatric (Ackery,
1987).
The only convincing Tellervo hostplant records cite Parsonsia (Apocynaceae);
suggestions of the Aristolochiaceae certainly need confirmation. With Parsonsia
species containing pyrrolizidine alkaloids, a possible chemical basis for
unpalatability is established. Additionally, adults may also come to pyrrolizidine
alkaloid sources (R. L. Smiles, personal communication). Mimetic complexes
involving males of Mycalesis drusillodes, the praslini species group of Neptis, Mynes
anemone and females of Praetaxila species seemingly centre on Telleruo patterns
(Vane-Wright, 1971, 1974, 1976; Ackery, 1987).

SUMMARY OF HOSTPLANT PATTERNS

Efforts to trace the ancestral larval hostplants of the Nymphalidae by

reference to their sister group, the Lycaenidae (Kristensen, 1976), are unlikely
to yield certain conclusions. Many lycaenids have highly derived feeding
habits-of the eight subfamilies recognized by Eliot (1973), the Liphyrinae and
Miletinae are wholly aphytophagous. Phytophagy is widespread elsewhere, but
unusually for butterflies, lycaenids particularly favour parts of plants rich in
nitrogen, especially buds, shoots, flowers, fruits and seeds (Cottrell, 1984). With
reference to flower-feeding, Robbins & Aiello (1982) suggests that this habit
may free species from the constraints associated with foliage-feeding. Perhaps
this is reflected in the bewildering diversity of host families noted in the
literature (see Ehrlich & Raven, 1965). However, Scott (1985) firmly considers
the Fabaceae as the likely proto-foodplant of the Papilionoidea/Hesperioidea in
their entirety. With legumes widely utilized by many butterfly groups, including
some nymphalids and lycaenids, this seems quite plausible. However, lycaenid
exploitation of the nitrogen-rich seeds and pods of many fabaceous plants may
represent a derived trait rather than a retained ancestral feature. Elswhere,
Scoble (1986) suggests that the conventional ‘geometrid’ group, the Hedyloidea,
may represent the sister-group of the Papilionoidea. Hedylids are only known to
feed on Sterculiaceae (Buettneria), offering no support for the Fabaceae
hypothesis. The fossil record places the origins of the Nymphalidae in the early
Tertiary (40-50 m.y. BP) (Whalley, 1986). During this period the Fabaceae
became well established and dispersed; there are earlier records but only those
for the late Cretaceous (65-70 m.y. B P ) can be accepted with confidence
(Polhill, Raven & Stirton, 1981).
The relationship between the Libytheinae and the remainder of the
Nymphalidae is crucial to in-group comparison. If their implied sister-group
relationship (see Ehrlich, 1958; Scott, 1985) is substantiated, then the Ulmaceae
would be another potential candidate as the ancestral host, with libytheines and
apaturines both on Celtis. Possibly the wide distribution and low species diversity
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 187

of Libythea offers some support for the Ehrlich/Scott hypotheses. Unfortunately,

while species of the Libytheinae have an impressive array of synapomorphies,
the establishment of ‘Nymphalidae minus Libytheinae’ rests weakly on the
contention that reduction in female forelegs, although common to both groups,
is less extreme in libytheine species. Further supposed evidence from Scott’s
evolutionary scenario for larval morphology is hardly unequivocal.
Viewed in isolation, hostplant association offers only limited insights into the
higher classification of the Nymphalidae. Even the most fundamental
‘grouping’, the monocotyledonous Brassolinae-Amathusiinae-Morphinae-
Satyrinae, has not gone unchallenged (DeVries et al., 1985). Based on data
derived from a limited number of taxa, Brassolinae-Amathusiinae-Morphinae
are confirmed as monophyletic (with the addition of the ‘satyrines’ Antirrhea and
Caerois), although the residual Satyrinae appear polyphyletic. It seems likely
that Fabaceae-feeding will characterize a subset within Morpho; elsewhere,
exploitation of bromeliads by Dynastor (Brassolinae) can also be seen as a unique
trend. Utilization of the Musaceae, an Old-world introduction into the
Neotropics, by both the Amathusiinae and the New-World Brassolinae further
underlines their intimate relationship. Disappointingly, the hostplants of the
Satyrinae (in their ‘traditional’ sense) offer few insights. Bamboos appear
characteristic of the pronophilines and Taygetis, Manataria, Aphysoneura, Lethe and
Neope, with the Haeterini and Elymniina favouring palms-neither association
finds support in received ideas in satyrine classification. Limited data suggest
that the Ragadiini are restricted to club-mosses (Selaginella), but even this is not
a unique trend, occurring also in the New-World genus Euptychia (Euptychiina).
The position of Calinaga remains ambiguous; although known to exploit Morus
(Moraceae), it has no obvious affinities with any other Moraceae-feeding
nymphalid group.
Some members of the Charaxinae have a wider range of hosts than any other
nymphalids. Of the six recognized subtribes, the Pallini (on Convolvulaceae),
Euxanthini (Sapindaceae) and Anaeini (Euphorbiaceae, Piperaceae-Anaeina;
Flacourtiaceae-Zaretina) seem well characterized. Although some genera of the
Preponini have only scant hostplant data, exploitation of the Fabaceae is
characteristic of Prepona, with Archaeoprepona species favouring the Lauraceae.
Charaxes itself, and to a lesser known extent Polyura, exhibits a strong underlying
fabaceous trend; other more particular charaxine themes are also amplified,
especially utilization of the Euphorbiaceae and Sapindaceae. Most noteworthy
perhaps is the characteristic range of sapindaceous hosts exploited by Charaxes
(subgen. Stonehamia) and the distribution of mimosaceous (Fabaceae) records in
the nominate subgenus. Overall, the favoured hostplants offer no insights into
the inter-relationships of the six constituent subtribes.
As pointed out by Ehrlich & Raven (1965), the Passifloraceae-Flacour-
tiaceae-Violaceae association supports the Heliconiinae-Acraeinae-Argynninae
as a probable natural grouping, although the position of the Passifloraceae
feeders, Vindula, Cethosia and Parthenos, remains ambiguous. Members of the
Acraeinae widely utilize members of other families, particularly the Asteraceae
and Urticaceae, but in the light of Pierre’s (1984) cladistic analysis these are
seen as derived features characterizing subsets within the subfamily. Within the
Argynninae, preferences for the Flacourtiaceae or Violaceae generally correlate
with broad distribution.
188 P. R. ACKERY

Several of the constituent subtribes placed within the Limenitinae show

convincing, if not mutually exclusive, hostplant correlations-the Neptini
on the Fabaceae, Biblini on Euphorbiaceae, Marpesiini on Moraceae,
Ageroniini on Euphorbiaceae, Parthenini on Passifloraceae and Epicaliini
on Euphorbiaceae/Sapindaceae. Two broad themes, centred on the
Rubiaceae-Naucleaceae and Caprifoliaceae, represent the Limenitini,
preferences which cut right across the limits of the constituent genera as
presently understood. Elsewhere, Bebearia (on monocotyledons, mainly
Arecaceae) , and Cymothoe (Violaceae), Pseudacraea (Sapotaceae) and Pseudoneptis
(Moraceae), if correctly placed within the limenitines, appear convincingly
characterized.
While the Ulmaceae (Celtis) strongly characterize both the Apaturinae and
Libytheinae, the Nymphalinae are much more diverse in hostplant preferences.
The Urticaceae-Acanthaceae association finds no support in current ideas of
plant classification-perhaps here at least a basic chemical similarity might be
anticipated? Although there are many variations on the two major theme
families-utilization of the Asteraceae, Scrophulariaceae and Plantaginaceae is
widespread in the acanthaceous series, with records for the Cannabidaceae,
Ulmaceae, Asteraceae, Grossulariaceae and Salicaceae, representing
‘departures’ from the urticaceous theme-few appear to exclusively characterize
more particular nymphaline groups. Two notable features stand out. Firstly, the
close parallel between melitaeine hostplant preferences and those of the
nymphalines Junonia, Anartia and Siproeta, all of which centre on the
Scrophulariaceae, Acan thaceae and Plantaginaceae. Iridoid glycosides are
implicated in important aspects of the biology of Junonia and certain
melitaeines. Secondly, the exclusively monocotyledous records for the monobasic
Oriental genus Kaniska represents a surprising departure from the dominant
nymphaline trends.
A convincing case can be made, both phylogenetically (Ackery & Vane-
Wright, 1984) and biologically (Edgar, 1984), for ‘proto-Apocynaceae’
representing the theme family for the Danainae-Ithomiinae-Tellervinae.
Exploitation of the Asclepiadaceae, Moraceae and Solanaceae represents
secondary shifts but none exclusively characterizes a subset within the major
grouping, although Moraceae-feeding is restricted to the Euploeinae.
HOSTPLAN’IS AND UNPALATABILITY

Unpalatability in butterflies derives from larval exploitation of poisonous

hostplants; this is the widely accepted chemical basis of unpalatability. Within
the Nymphalidae, various butterfly-plant associations might be cited in support
of this view, notably the I thomiinae-Solanaceae, Danainae-Asclepiadaceae,
Heliconiinae-Passiforaceae and Acraeinae-Passifloraceae. Nonetheless, this
comfortable notion may now need reconsideration-Brown ( 1985) notes that
“contrary to general wisdom . . . the Ithomiinae derive few or no predator-
defense compounds from their larval hostplants”. Similarly, but perhaps more
contentiously, Nahrstedt & Davis (1981) concluded that in heliconiines and
acraeines “the glucosides must be synthesized by the animals themselves and not
sequestered from their food”. Even in the Danainae, it now seems that cardiac
glycosides, derived from larval hostplants, cannot be considered as universal
features (Ackery & Vane-Wright, 1985). Might a paradox be emerging?
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES I89

Members of these groups appear sufficiently protected from predators to permit

an aposematic life-style, but evidence mounts that their hostplants, although
containing various toxic secondary compounds, may not always be the primary
source of unpalatability. Is it possible that in some cases at least, we have put
‘the cart before the horse’; if certain groups of butterflies are unpalatable per se,
could this open up protected plant groups for exploitation?
As an example, consider a theoretical proto-danaine that acquired the trick of
producing self-secreted cardio-active substances. This may not be as far-fetched
as it first appears: Rothschild, Marsh & Gardiner (1978) report such cardio-
active compounds in both Danaus plexippus and Euploea core. As has already been
established, pathways do exist for cardiac glycosides to be passed on to the eggs
of the next generation (Reichstein et al., 1968) so it may not be unreasonable to
assume an analogous situation for our theoretical self-secreted compound. In
this context, the first instar larval habit of eating its own eggshell might be of
particular importance. Could this to some extent ‘imprint’ the gustatory
response of the newly emerged larva, increasing the likelihood of it accepting
potential hosts that also contain cardio-active substances?
The Ithomiinae may possibly most closely match this no doubt over-simplified
scenario. Adult sequestration of pyrrolizidine alkaloids (PAS) appears plausible
as a derived feature of the Ithomiinae-Danainae-Tellervinae infrafamily, with
‘proto-Apocynaceae’ representing the ancestral larval host (Edgar, 1984; Ackery
& Vane-Wright, 1985). Various scattered ithomiine genera, Aeria, Tithorea,
Elzunia, still exploit the Apocynaceae-in all cases the larvae are
characteristically aposematic and danaoid in appearance. Is it possible that
adult tolerance of pyrrolizidine alkaloids has opened up the solanaceous-alkaloid
containing Solanaceae? Adult derived PAS are abundant in ithomiine eggs but
do not occur in larvae or pupae (Brown, 1985). Generally, Solanaceae-feeding
ithomiine larvae are cryptic, suggesting that although they ‘tolerate’ the known
range of pharmacologically active secondary plant compounds, none is stored to
permit an aposematic life-style. Melinaea larvae provide an obvious exception.
They are aposematic and danaoid in form-Brown (1977) recorded the larval
host as ?Forsteronia (Apocynaceae), a determination that he has since abandoned
(Brown, 1985) in favour of records for the Solanaceae (see Drummond, 1976).
Could it be that Melinaea, at least, has developed the ability to exploit the
chemical defences of the Solanaceae, actively storing hostplant derived
compounds that have permitted a return to the ancestral aposematic larval life-
style? Of course, this begs the ultimate question, why not switch to the
Asteraceae or Boraginaceae and have done with adult PA-foraging altogether?
I n danaines at least, larvae are known to be PA tolerant (Rothschild & Edgar,
1978; Edgar, 1982). With Actinote species (Acraeinae) utilizing such composites
as Eupatorium and Senecio, genera known to contain PAS, no doubt a plausible
‘competitive’ story could be concocted. Generally, though, representatives of the
Boraginaceae are not widely exploited, at least by nymphalid butterflies.
Most heliconiines and many acraeines feed on members of the Passifloraceae,
well-known as sources of cyanogenic glycosides (Darnley Gibbs, 1974). Analysis
of adult acraeines (Nahrstedt & Davis, 1981) revealed the highest concentration
of the cyanoglucosides Linamarin and Lotaustralin in ‘Acraea encedon’ (now
known to be a species complex) and A. eponina, with the lowest in A. natalica.
The same glucosides were confirmed in a mixed sample of Heliconius species. As
190 P. R. ACKERY

the identified glucosides differed from those found in cyanogenic members of the
Passifloraceae, the presumed larval hosts, Nahrstedt & Davis concluded that the
butterflies probably synthesize these compounds themselves. I n the light of their
later study (Nahrstedt & Davis, 1983) this conclusion is almost certainly upheld,
despite the fact that the two Acraea species with the highest concentration of
glucosides in fact never feed on the Passifloraceae! T h e most common and
widespread members of the encedon-complex take species of Commelinaceae or
Fabaceae (Guilbot & Pierre, 1978), with A . eponina on the Tiliaceae or
Sterculiaceae. I n fact, only A . natulica (now considered to comprise two
parapatric species, A. pseudegina and A. natalica: Pierre, 1981b) with the lowest
concentration of cyanoglucosides actually feeds on passifloraceous hostplants.
Nahrstedt & Davis (1983) emphasize that many compounds of a toxic nature
found in the Lepidoptera are either identical to, or similar to, secondary
metabolites of plants. However, cyanogenic glycosides can be synthesized by
butterflies themselves and d o not depend upon sequestration during larval
feeding. I n Heliconius itself, and possibly other heliconiines and acraeines,
cyanogenic glycosides are synthesized from amino acids at both larval and adult
stages. Although adult pollen feeding might play a significant role (Grimshaw,
1983), this can only apply to Heliconius and Eueides-Brown ( 1981) treats pollen-
feeding as a synapomorphy for these two genera. Is it posible then, that in both
of these subfamilies exploitation of the Passifloraceae has been facilitated by
tolerance to self-secreted cyanogenic compounds?
Unlike the ithomiines, all known danaine larvae are typically aposematic,
suggesting that considerable protection is derived from their larval hosts.
Although cardenolides play a significant role in the genus Danaus itself, the
factor (or factors) involved elsewhere have yet to be elucidated. At this stage it
should be confessed that many liberties were taken with the opening scenario
relating to ‘proto-Danainae’. Adult sequestered PAS are yet another
complicating factor (see Edgar, Culvenor & Pliske, 1974; Boppri, 1984). While
pyrrolizidine and solanaceous alkaloids have certain affinities, being derived
from the same evolutionary “nitrogen pathway” (see Harborne, 1982),
cardenolides have more in common with the lactones and saponins of the
“terpenoid pathway”. So, while PAS might ‘unlock’ the Solanaceae for
ithomiine exploitation, a comparable scenario seems unlikely for cardenolides.
Whatever the function of self-secreted cardio-active compounds, the danaines
are the one group amongst the Nymphalidae likely to conform most closely to
classical insectlplant biochemical dogma.
A final plea; too often, hypotheses are seen as mutually exclusive-once an
idea comes into vogue it is forced to fit every situation. If, in some instances, a
proposed hypothesis seems likely to fail, its total abandonment need not
automatically follow. Perhaps there is now mounting evidence that the key to
unpalatability in butterflies does not always lie in hostplant chemistry, but it
surely still has a significant role.

ACKOWLEDGEMENTS

Firstly, I am indebted to R. I. Vane-Wright and C. J. Humphries for reading

and commenting on a succession of preliminary drafts. Professors M. S. Blum,
D. H. Janzen and J. A. Harborne kindly commented on the more contentious
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 191

aspects of hostplants and unpalatability; Professor Harborne also assessed the

occasional forays into the murk of phytochemistry. Their generous advice is
much appreciated. Others gave willingly of their time and expertise: Phil
DeVries, Dave Hollis, John Huxley, Dale Jenkins, Cindy North, Mike Parsons,
Malcolm Scoble, the late Bob Silberglied, Bob Smiles, Campbell Smith, Gerry
Tremewan and Paul Whalley; all are acknowledged with thanks.

REFERENCES

ACKERY, P. R., 1987. The danaid genus Telleruo (Lepidoptera-Nympha1idae)-a cladistic approach.
zoological Journal of the Linnean Society, 89: 203-274.
ACKERY, P. R. & VANE-WRIGHT, R. I., 1984. Milkweed ButterJlies, their Cladistics and Biology. London: BM
(NH).
ACKERY, P. R. & VANE-WKIGHT, K. I., 1985. Patterns of plant utilization by danaine butterflies.
Proceedings of the 3rd Congress of European Lepidopterology, 1982: 3-6. Karlsruhe: Societas Europaea
Lepidopteroligica.
ADAMS, M. J., 1986. Pronophiline butterflies (Satyridae) of the three Andean Cordilleras of Colombia.
~oologicalJournal of the Linnean Society, 87: 235-320.
AGENJO, R., 1975. Las Melitaea (Mellictaj deione Gey., 1827-1832, athalia (Rott., 1775) y parthenoides Kef.,
1851, en Esparia (Lep. Nymphalidae). Graellsia, 30: 3-61.
AIELLO, A., 1984. Adelpha (Nymphalidae): deception on the wing. Psyche, 91: 1-45.
AIELLO, A. & SILBERGLIED, R. E., 1978. Life history of Dynastor darius (Lepidoptera: Nymphalidae:
Brassolinaej in Panama. Psyche, 85: 33 1-345.
AOKI, T., YAMAGUCHI, S. & UEMURA, Y., 1982. Satyridae, Libytheidae. In E. Tsukada (Ed.),
Buttmjies of the South-East Asian Islands, 3: 1-628. Japan: Plapae Co. Ltd.
ASHIZAWA, H. & MUROYA, V., 1967. Notes on the early stages of Calinaga buddha formosana. Special Bulletin
of the Lepidopterological Society of Japan, 3; 79-85.
ATKINS, A., 1975. Larval foodplants of some Queensland butterflies. News Bulletin. Entomological Society of
Queensland, 3: 1 17-1 19.
AUERBACH, M. J. & STRONG, D. R., 1981. Nutritional ecology of Heliconia herbivores: experiments with
plant fertilization and alternative hosts. Ecological Monographs, 51: 63-83.
AURIVILLIUS, C., 1911-13. Family: Nymphalidae. In A . Seitz, Die Gross-Schmetterlinge der Erde, Stuttgart, ( 2 )
13: 121-238.
AUSTIN, G. T., 1977. Notes on the behaviour of Asterocampa leila (Nymphalidaej in southern California.
Journal of the Lepidopterists’ Society, 31: 11 1-1 18.
AUSTIN, G. T . & AUSTIN, A. T., 1981. Butterflies of Clark County, Nevada. Journal of Research on the
Lepidoptera, 19: 1-63.
BANNO, H., 1984. The effect of various plants on growth and food utilization of the larvae of Neptis sappho
intermedia W. B. Pryer (Lepidoptera: Nymphalidae). Japanese Journal of Ecology, 34: 9-14.
BARCANT, M., 1970. ButterJlies of Trinidad and Tobago. London: Collins.
BARSELOU, P. E., 1983. The genus Agrias-a taxonomic and illustrated guide (Lepidoptera, Nymphalidae). Venette.
BASCOMBE, M. et a/., 1987. ButterJlies of Hong Kong. [Seen only in draft.]
BATES, H. W., 1862. Contributions to an insect fauna of the Amazon Valley. Transactions of the Linnean Society
of London, 23: 495-566.
BENSON, W. W., BROWN, K. S. JR. & GILBERT, L. E., 1976. Coevolution of plants and herbivores:
passion flower butterflies. Euolution. Lancaster, Pa, 29: 659-80.
BIEZANKO, C. M., 1949. Acraeidae, Heliconiidae et Nymphalidae de Pelotas e seus arredores. (Contibuiccio ao
conhecimento dajsiographia do Ria Grande do Sul. j Pelotas.
BINK, F. A,, 1985. Host plant preference of some grass feeding butterflies. Proceedings of the 3rd Congress of
European Lepidopterology, Cambridge 1982, 23-29. Karlsruhe: Societas Europaea Lepidopterologica.
BIRKET-SMITH, J., 1960. Results from the Danish Expedition to French Cameroons (1948-1950).
XXVII-Lepidoptera (Part 11). Bulletin de l’lnstitut Fransais BAfrique Noire ( A ) , 22: 924-983.
BOPPRE, M., 1984. Chemically mediated interactions between butterflies. In R. I. Vane-Wright & P. R.
Ackery, The Biology of ButterJlies; 259-275. Symposia of the Royal Entomological Society of London, No.
11.
BOWERS, M. D., 1980. Unpalatability as a defense strategy of Euphydryas phaeton. Evolution. Lancaster, Pa, 34:
586-600.
BOWERS, M. D., 1981. Unpalatability as a defense strategy of western checkerspot butterflies (Euphydryarj.
Evolution. Lancaster, Pa, 35: 367-375.
BOWERS, M. D., 1983a. Mimicry in north American checkerspot butterflies: Euphydryas phaeton and Chlosyne
harrisii (Nymphalidae). Ecological Entomology, 8: 1-8.
192 P. R. ACKERY

BOLVERS, h1. D.. 198313. ‘Ihe role of iridoid glycosides in host-plant specificity of checkcrspot hutrerflics.
,7ournal of Cliemical Ecoloxy, 9: 475-494.
BOL1‘ERS. Yl. L).- 1984. Iridoid Rlycosidcs and host-plant specificity i n larvae of the Buckryc huttcrfiy. ,7unonia
coenia (Nymphalidae). Journal oJChemica1 Ecology, 10: 1567-1577.
BOWERS, M.D. & WIERNASZ, D. C., 1979. Avian predation on the palatable butterfly, Cere7cq‘onisp q a l a .
Ecological Entomology, 4: 2055209.
BRISTOW, C. R., 1981, A revison of the brassoline genus Catoblepia (Lepidoptera: Rhopalocera). zoological
h70urnal of the Linnean Society, 72: 117-163.
BKIS‘TOW, C;. R., 1982. A revision of the brassoline genus Selenophanes (Lepidoptera: Rhopalorera). ~oologi.ic-i11
Jouinal o f t h e Lznnean Society, 76: 273-291.
BRIS’I’OW, C. R . ( I n prep.) A revision of the brassoline genus Opsiphanes (Lepidoptera: Rhopalocerai.
BROOKS, C. J., 1950. A revison of the genus Tenaris Hiibner (Lepidoptera: Amathusiidae). 7ranJaction.s c?f/he
Royal Entomolo~gicalSncieb of London, 101: 179-238.
BROOKS, M.& KNIGHT, C., 1982. A Complete Guide to British ButterJlis. London: Jonathan Cape.
BROL\’ER, L. P., 1984. Chemical defence in butterflies. In R. I. Vane-Wright & P. R. Ackrry. The Biology qf
BulterJlies: 109-134. Symposia of the Royal Entomological Society of London No. 1 I .
BROWER, L. P. & BROWER, J. V . Z., 1964. Birds, butterflies and plant poisons; a study in ecological
chemistry. zoologica, New 1To?k, 49: 137-159.
BROWN, F. M. & HEINEMAN, B., 1972. Jamaica and its ButterJlies. London.
BROLVN, K. S. JR, 1977. Geographical patterns of rvolution in Neotropical Lepidoptera: differentiation uf
the species of Melinaea and Mechanitis (Nymphalidae, Ithomiinae). Systemattc Entomology, 2: 161-197.
BROWN, K. S . .JR, 1980. A review of the genus Hypothyris Hiibner (Nymphalidae), with descriptions of thrce
new subspecies and early stages of H. daphnis. Journal of the Lepidopterists’ Society, 34: 152-172.
BROLVN, K. S. J R , 1981. The biology of Helrconius and related genera. Annual Revzew Df Entomolo~py,2fj:
‘+27--456.
BROLVN, K. S. J R , 1985. Chemical ecology of dehydropyrrolizidine alkaloids in adult Ithomiinae
Lepidoptera: Nymphalidac). Revista Brasileira de Biologia, 44: 435-460.
1

BROWN, K. S. J R & MIELKE, 0. H . H., 1972. The Heliconians of Brazil (Lepidoptera: Nymphalidaei.
Part 11. Introduction and general comments, with a supplementary revision of the tribe. zoologica, N m
l u r k , 57; 1 40.
BROLVN, K. S. JR, SHEPPARD, P. M. & T U R N E R , J. R. G., 1974. Quaternary refugia in tropical
America: rvidcnce from race formation in Heliconius butterflies. Proceedings of the Royol Society of London ( B )
187: 369 378.
BROLVN, L. N., 1973. A population of Lethe appalachia (Satyridae) from west central Florida. j‘ournal of the
Lepidopterists’ Soczety, 27: 238--239.
CARCASSON, R. H., 1981. Collins Handguide to the ButterJlzes of Africa. London: Collins.
CAREY-HUGHES, J. & PICKFORD, J. B., 1977. A n Annotated Checklist o f t h e Butteflies of Hang Kong. Hong
Kong: Green Pagoda Press.
CASAGRANDE, M. M., 1979. Sohre Caligo beltrao (Illiger) 1 : Taxonomia, biologia, morphologia das fases
imaturas e distribuiq6,es espacial e temporal (Lepidoptera, Satyridae, Brassolinae). Revista Brasiliera de
~ntomologia,39: 173-193.
CHANG, B. S., 1963. Life-histories of some Formosan butterflies in Kannon district, North Formosa. ButterJlier
and .Uoths, Kyoto, 14: 90-101.
CHANG, B. S., 1967. The immature stages and life-history of Mycalesis gotama nanda Fruhstorfer in Formosa.
Butteflies and Moths, Kyoto, 17: 75-77.
CH.WG, B. S., 1972. Notes on the immature stages and food-plants of some Formosan butterflies. Butterpzrs
and M o t h s , Kyoto, 23: 19-23.
CHERMOCK, R. L., 1950. A generic revision of the Limenitini of the World. clmerican Midland .Vaturalzst, 43:
5 13-569.
CHEW. F. S. & ROBBINS, R. K., 1984. Egg-laying in Butterflies. In R. I. Vane-Wright & P. R . Ackery, T h e
Biolou OJ ButterJlies: 65-79, Symposia of the Royal Entomological Society of London No. 1 1 .
CLARK, A. H., 1947. The interrelationships of the several groups within the butterfly superfamily
Nymphaloidea. Proceedings of the Entomological Soci@ of Washington, 49: 148- 149.
COZIIMON, I. F. B. & WATERHOUSE, D. F., 1972. ButterJlies of Australia. Sydney. [2nd Edition, 1981.1
Sydney: Angus & Robertson.
COMSTOCK, J . A. & GARCIA, L. V., 1961. Estudias de 10s ciclos biologicos en lepidopteros Mexicano\.
h a l e s del Instituto de Biologia, Uniuersidad de Mexico, 31: 349-448.
COMSTOCK, W. P., 1944. Insects of Porto Rico and the Virgin Islands. Lepidoptera (Suborder!
Rhopalorera (Superfamily) Papilionoidea ( l r u e hutterflies) (Superfamily) Hesperioidca (Skippers .
Scientijc Survey qf Porto Rico and the Virgin Islands, .New Yark, 12: 419-622.
COMS’TOCK, W. P., 1961. ButterJlies of the American Tropics. The genus Anaea: a study of the species heretojore
included in the genera Anaea, Coenophlebia, Hypna, Polygrapha, Protogonius, Siderone and Zaretis. New
York: American Museum of Natural History.
CONDAMIN, M.,1965. Contribution a la faune de Congo (Brazzaville) (Mission A. Villiers et A.
Drscarpentries). \’I. LCpidopttres Satyridae. Bulletin de l’lnstitut Frangaise d’dfrique Noire, d 27: 763 -77 I .
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 193
CONNER, W . E., EISNER, T., VAN DER MEER, R. K., GUERRO, A. & MEINWALD, J. 1981.
Precopulatory sexual interaction in an arctiid moth (Utethesia ornatrix): role of a pheromone derived from
dietary alkaloids. Behaviourat Ecology and Sociobiology, 9: 227-235.
CORBET, A. S. & PENDLEBURY, H. M., 1978. T h e ButterJlies of the Malay Peninsula (3rd edition, revised by
J. N. Eliot). Kuala Lumpur.
CRAW, R. C., 1978. Revision of the genus Argyrophenga (Lepidoptera: Satyridae). N e w zealand Journal of
z o o l o u , 5: 751-768.
COTTRELL, C. B., 1984. Aphytophagy in butterflies: its relationship to myrmecophily. ~oologicalJournal of
the Linnean Society, 80: 1-57.
CUBERTO, R., 1985. Notes on the life cycle and natural history of Opsiphanes quiteria quirinus Godman and
Eryphanis aesacus buboculus Butler (Brassolidae). Journal of the Lepidopterists’ Society, New York, 39: 33-43.
D’ABRERA, B., 1977. BufterJlies of the Australian region (2nd edition). Melbourne: Lansdowne Editions.
DALMEIDA, R. F., 1922. Mllanges Lipidoptiro/ogiques. Etudes JUT les Llpidoptlres du BrLssil. Berlin.
D’ARAUJO E SILVA, A. G . , GONCALVES, C. R., GALVAO, D. M., GONCALVES, A. J. L., COMES,
J , , DO NASCIMENTO SILVA, M. & DE SIMONI, L., 1968. Quarto catalogo dos inretos que vivem nas
+
plantas do Brasil ( 2 ) I : xxviii 622 pp. [Corrigenda (2) 2: 231-265.1. Rio de Janeiro: Ministry of
Agriculture.
DARGE, P., 1983. The genus Charaxes Ochs (Lepidoptera, Charaxidae Doherty). Fauna of the United Republic nf
Cameroun, i: 1-136.
DARNLEY GIBBS, R., 1974. Chemotaxonomy of Flowering Plants. 4 vols. Montreal & London: Queen’s
University Press.
DAVIS, R. H. & NAHRSTEDT, A., 1979. Linamarin and Lotaustralin as the source of cyanide in a g a e n a
Jil$endulae L. (Lepidoptera). Comparative Biochemistry and Physiology, 64B: 395-397.
DEBAAR, M., 1981. Host records for Hypocysta adiante adiunte (Hiibner) and Xois arctous arctous (Fab.).
Lepidoptera: Nymphalidae. News Bulletin. Entomological Society of Queensland, 9: 9.
DE LA MAZA, E. R. G . & DE LA MAZA, E. J., 1977. Notas sobre el ciclo de vida de Polygonia g-argenfeum (Dbl,
y Hew.) (Nymphalidae). Revista de la Sociedad Mexicana de Entomologia, 3: 35-41.
DESCIMON, H., 1977. BiogCographie, mimetisme et speciation dans le genre Agrias Doubleday (Lep.
Nymphalidae Charaxinae). In H. Descimon, Biogeographie el Evolution en Amerique Tropicale. Paris:
Publication du Laboratoire de Zoologie de 1’Ecole Normale Superieure No 9.
DETHIER, V. G., 1940. Life histories of Cuban Lepidoptera. Pvche, Cambridge, Massachusetts, 47: 14-25.
DETHIER, V. G . , 1959. Egg-laying habits of Lepidoptera in relation to available food. Canadian Entomologist,
91: 554-561.
DE VIEDMA, M. G. & GOMEZ BUSTILLO, M. R., 1976. Libro Rojo de 10s Lepidopteros Ibericos. Mabrid:
Instituto Nacional para la Conservacion de la Naturaleza.
DEVRIES, P. J., 1977. Eumaeus m i y a s Hubner, an aposematic Lycaenid butterfly. Brenesia, 12/13: 269-270.
DEVRIES, P. J., 1980a. Description, natural history and distribution of a new species of Eretris (Satyridae)
from Costa Rica. Journal of the Lepidopterists’ Society, 34; 146-151.
DEVRIES, P. J , , 1980b. The genus Agrias (Lepidoptera: : Nymphalidae: Charaxinae) in Costa Rica:
description of a new subspecies of Agrias amydon, new records, and natural history observations. Brenesia, 27:
295-302.
DEVRIES, P. J., 1983. Butterflies of Costa Rica: systematics, ecology and field site checklists. In D. H. Janzen,
Costa Rican Natural History. Chicago: University of Chicago Press.
DEVRIES, P. J., 1986. Hostplant records and natural history notes on Costa Rican butterflies (Papilionidae,
Pieridae & Nymphalidae). Journal of Research on the Lepidoptera, 24: 290-333.
DEVRIES, P. J., KITCHING, I. J. & VANE-WRIGHT, R. I., 1985. The systematic position o f Antirrhea and
Caerois, with comments on the classification of the Nymphalidae (Lepidoptera). Qstematic Entomology, 10:
11-32.
DIMOCK, T. E., 1978. Notes on the life cycle and natural history of Vanessa annabella (Nymphalidae).Journal
of the Lepidopterists’ Society, 32: 88-96.
DOMINGUEZ, X. A,, 1977. Eupatorieae-chemical review. In V. H. Heywood, J. B. Harborne & B. L.
Turner, The Biology and Chemistry of the Compositae, 1: 487-502. London: Academic Press.
DORNFELD, E. J., 1980. The Butte$& of Oregon. Oregon: Timber Press.
DOS PASSOS, C. F. & GREY, L. P., 1945. A genitalic survey of the Argynninae (Lepidoptera,
Nymphalidae). American Museum Novitates No. 1296.
DOWNEY, J. C., 1962. Host-plant relations as data for butterfly classification. Systematic <oolog~, Washington,
I l : 150-159.
DOYLE, J. F., 1979. Temporary range extention and larval foodplant o f Dynamine dyonzs (Nymphalidae) in
Texas. Journal of the Lepidopterists’ Society, 33: 20.
DRUMMOND, B. A., 1976. Comparative ecology and mimetic relationships of Ithomiine butterJlies in eastern Ecuador.
Ann Arbor, Michigan.
DRUMMOND, B. A,, 1986, Coevolution of ithomiine butterflies and solanaceous plants. In W. G . D’Arcy,
Solanaceae: Biology and Systematics: 307-327. New York: Columbia University Press.
DRUMMOND, B. A. & BROWN, K. S. Jr, 1987. Ithomiinae (Lepidoptera, Nymphalidae): summary of
known larval foodplants. Annals of the Missouri Botanical Garden, 74 [in press].
194 P. R . ACKERY

DURDEN, C. J., 1965. Speyeria callippe and Arlemesia, a possible foodplant. Journal of the Lepidoptertsts' SOCZP(Y,
19: 186-187.
EBERT, G., 197 I , Drei neue Macrolepidoptera-Arten aus Iran. Beitruge zur naturkundlichen Fortchung In
Sudwestdeutschland, 30: 65-7 1. 1 pl.
EBNER, .J. A., 1970. The butterflies of Wisconsin. Milwaukee.
EDGAR, J. A,, 1982. Pyrrolizidine alkaloids sequestered by Solomon Island danaine butterflies. The feeding
preferences of the Danainae and Ithomiinae. Journal of <oology, 196: 385-399.
EDGAR, J. A. 1984. Parsonsieae: ancestral larval food plants of the Danainae-Ithomiinae? In R . I. Vane-
Wright & P. R . Ackery, T h e Biology of Butterflies: 91-93. Symposia of the Royal Entomological Society of
London No. 11.
EDGAR, J . A,, CULVENOR, C. C. J. & PLISKE, T. E., 1974. Co-evolution of Danaid butterflies and their
hostplants. Nature, London, 250: 646-648.
EHLE, G., 1957. Unusual occurrence of Melitaea nycteis (Nymphalidae) in Lancaster County, Pennsylvania.
Lepidopterists' News, 11: 38-41.
EHRLICH, P. R., 1958. The comparative morphology, phylogeny and higher classification of ihe butterflies
(Lepidoptera: Papilionoidea). Kansas University Science Bulletin, 39: 305-370.
EHRLICH, P. R., 1984. The structure and dynamics of butterfly populations. In R. I. Vane-Wright & P. R.
Ackery, The Biology of Butterflies: 25-40. Symposia of the Royal Entomological Society of London No. 11.
EHRLICH, P. R. & EHRLICH, A. H., 1961. H o w to know the Butterflies. Dubuque, Iowa: W. M. C. Brown.
EHRLICH, P. R . & MURPHY, D. D., 1982. Butterfly nomenclature: a critique. Journal of Heseurch on the
Lepidoptcra, 20: 1-I 1.
EHRLICH, P. R. & RAVEN, P. H., 1965. Butterflies and plants: a study in coevolution. Evolution, Lancasler,
PA, 18: 586-608.
EISNER, T., 1980. Chemistry, defence and survival: case studies and selected topics. I n M. Locke & D. S.
Smith, Insect Biology in the Future: 847-878. London: Academic Press.
ELIOT, .J. N., 1969. An analysis of the Eurasian and Australian Neptini. Bulletin o f t h e British Museum (~Vatural
History) (Entomology) Suppl. 15.
ELIO'I~,J. N., 1973. The higher classification of the Lycaenidae: a tentative arrangement. Bulletin of Ihe British
Museum (Natural History), Entomology, 28: 373-506.
ELIOT, J . N., 1978. Revised (3rd) edition of A. S. Corbet & H. M. Pendlebury, T h e Butterflies o f t h e Malay
Peninsula. Kuala Lumpur: Malay Nature Society.
EUI'RINGHAM, H., 1910. African Mimetic Butterflies. Oxford: Clarendon Press.
EUI'RINGHAM, H., 1912. A monograph of the African species of the genus Acraea, Fab., with a supplement
of those of the Oriental Region. Transactions of the Entomological Society, London, 1912: 1-374.
EEI'RINGHAM, H., 1921. O n the African species of the genus Neptis Fab. Transactions q f t h c Entomological
Socirty of London, 1921: 532-589.
EMMEL, J. F. & EMMEL, T. C., 1974. Ecological studies of the Rhopalocera in a Sierra Nevadan
Community - D o n n a Pass, California. V. Faunal additions and foodplant records since 1962. Journal of the
Lepidopterists' Society, 28: 344-348.
EMMEI,, J. F., SHIELDS, 0. & BREEDLOVE, D. E., 1970. Larval foodplant records of North American
butterflies. Part 2. Journal of Research on the Lepidoptera, 9: 233-242.
EMMEL, T. C. & EMMEL, J . F., 1962. Ecological studies of Rhopalocera in a high Sierran community-~
Donner Pass, California. I . Butterfly associations and distributional factors. Journal o f the Lepidopterists'
Society, 16: 23-44.
EMMEL, T. C. & EMMEL, J. F., 1973. The butterflies of southern California. Science Series, Natural History
Museum o f L o s Angeles County No. 26, i-xii, 1-148.
FERRIS, C. D. & BROWN, F. M., 1981. Butterflies of the Rocky Mountain States. Oklahoma.
FIELD, W. D., 1971. Butterflies of the genus Yanessa and the resurrected genera Bassaris and Cynthia.
Smithsonian Contributions to 3 o l o g y No. 84.
E'ONTAINE, M., 1981a. Biotopes et premiers etats des espkces de Nymphalidae-Eunicinae observkes au Zaire.
Lambillzonea, 81: 50-57.
FONTAINE, M.,1981b. Lep: Nymphalidae Sous-fam. Eurytelinae. Premiers etats des espkces observees a
Isiro (ex. Paulis) et environs immediats Zaire: region du Haut-Zaire, sous-region du Haut-Uele.
Lambillionm, 80: 65-77.
FONTAINE, M., 1982-3. Genre Cymothoe Hiibner Lep. Nymphalidae- S. fam. Nymphalinae. Note sur les
premiers etats. Lambillionea, 82: 63-64, 67-72, 95-98.
FON'I'AINE, M., 1985. Generalites cles de determination des chenilles et classification des espkces des genres
Precis Hbn et Junonia Hbn. (Lepidoptcra, Nymphalidae). Bulletin et Annales de la SociCti Royale Entomologique
dr Belgtque, 121: 3 17-364.
FORSTER, W. S., 1964. Beitrage zur Kenntnis der Insektenfauna Boliviens XIX. Lepidoptera 111. Satyridae.
Veroffentlichungen der xoologischen Staatssammlung Munchen, 8: 53- 188.
FORSTER, W. & WOHLFAHRT, T. A,, 1955. Die Schmetterlinge Mitteleuropas. Ta,&lter Diurna (Rhopalocera
und Hesperiidae). Stuttgart: W. Keller und Co.
FOUNTAINE, M. E., 1913. Five months butterfly collecting in Costa Rica in the summer of 1911.
Entomologist, 46: 189-194, 214-219.
 HOSTPLANTS AND CLASSIFICATION: NY MPHALID BUTTERFLIES 195

FOUNTAINE, M. E., 1925-6. Amongst the Rhopalocera of the Philippines. Entomologist, 58: 235-239,
263-265; 59: 9-11, 31-34, 53-57.
FOUNTAINE, M. E. Unpublished watercolours of larvae and pupae. 4 vols. British Museum (Natural
History) Collection.
FOX, R. M., 1956. A monograph of the Ithomiidae. Part 1. Bulletin ofthe American Museum of Natural History,
111: 1-76.
FOX, R. M., 1961. A check list of the Ithomiidae. 1. Tribes Tithoreini and Melinaeini. Journal of the
Lepidopterists’ Society, 15: 25-33.
FOX, R. M., LINDSEY, A. W. Jr, CLENCH, H. K. & MILLER, L. D., 1965. The Butterflies of Liberia.
Memoirs of the Americun Entomological Society No. 19, frontispiece, i-ii, 1-438.
FRANEKEL, G., 1959. The raison d’etre of secondary plant substances. Science, 129: 1466-1470.
FRIEDRICH, E., 1978. Zur Zucht, Morphologie und Biologie, von Clossiana dia L., unter Besonderer
Beriicksichtigung des Dormanzphanomens (Lep., Nymphalidae). Zeitschrij der Arbeitsgemeinschaft
Osterreichischer Entomologen, 30: 43-48.
FRUHSTORFER, H., 1911-12. Family Amathusiidae. In A. Seitz, Gross-Schmetterlinge der Erde, Stuttgart, (2)
9: 403-452.
FRUHSTORFER, H., i912. Family: Brassolidae. In A. Seitz, Gross-Schmetterlinge der Erde, Stuttgurt, (2) 5:
285-332.
FRUHSTORFER, H., 1912-13. Family: Morphidae. In A. Seitz, Gross-Schmetterlinge der Erde, Stuttgart, (2) 5:
333-356.
FRUHSTORFER, H., 1916a. Genus: Ageronia. In A. Seitz, Gross-Schmetterlinge der Erde, Stuttgart, (2) 5:
537-545.
FRUHSTORFER, H., 1916b. Genus Prepona. In A. Seitz, D i e Gross-Schmetterlinge der Erde, Stuttgart, (2) 5:
550-566.
FUKUDA, H., 1983. Life histories of two Satyrid butterflies feeding on selaginellas. Buttegies and Moths, Kyoto,
33: 132-144.
FUKUDA, H., HARMA, E., TAKAHASHI, A,, TAKAHASHI, M., TANAKA, B., TANAKA, H.,
WAKABAYASHI, M. & WATANABE, Y., 1982-3. Lije Histories of Buttegies in Japan. 2 vols. Osaka:
Hoikusha Publishing Co.
GARTH, J. S. & TILDEN, W. W., 1963. Yosemite butterflies: an ecological survey of the butterflies of the
Yosemite sector of the Sierra Nevada, California. Journal of Research on the Lepidoptera, 2: 1-95.
GIBBS, G. W., 1980. N e w zealand buttegies; identification and natural history. Auckland: William Collins
Publishers Ltd.
GIBSON-HILL, C. A,, 1947. Contributions to the natural history of Christmas Island in the Indian Ocean.
Lepidoptera (Rhopalocera). Bulletin of the Rafles Museum, 18: 74-80.
GIFFORD, D., 1965. A list ofthe butterJles of Malawi. Blantyre: The Society of Malawi.
GILBERT, L. E., 1971. Butterfly-plant coevolution: has PassiJlora adenopoda won the selection race with
heliconiine butterflies. Science, 172: 585-586.
GILBERT, L. E., 1975. Ecological consequences of co-evolved mutualism between butterflies and plants. In
L. E. Gilbert & P. H. Raven, Coeuolution of Animals and Plants: 210-240. Austin: University of Texas Press.
GRIMSHAW, P., 1983. Test releases of Heliconius melpomene, carried out to discover if it is feasible to house this
species in conjunction with a large collection of tropical birds. Newsletter, Exotic Entomology Group. Amateur
Entomologists’ Society, 1983: 9-1 2.
GUILBOT, R. & GUILLAUMIN, M., 1977. Note prtliminaire sur I’devage et le dtveloppement larvaire
d’Euphaedra rezia Hewitson (Lep. Nymphalidae). Bulletin de la SocilC Entomologique de France, 82: 2 1 1-216.
GUILBOT, R. & PIERRE, J. 1978. Etude comparative des premiers Ptats de deux espkes-jumelles de
Ltpidoptkres Acraeides Acraea encedon [Linne] et A . encedana Pierre. Bulletin de la SociIC Entomologique de
France, 83: 163-170.
GULLANDER, B., 1959. Nordens D a d u r i l a r . Stockholm: P. A. Norstedt & Soners.
HABER, W. A,, 1978. Evolutionary ecology of tropical mimetic buttedzes. Unpublished Ph.D. thesis, Minnesota
University.
HAGEN, B., 1896. Verzeichniss van mir auf Sumatra gefangenen Rhopalocera. Deutsche Entomologische
zeitschrijt, Iris, 9: 153-187.
HAIG, E. F. G., 1938. The butterflies of Nigeria V. Nymphalidae Charaxidinae. Nigerian Field, 7 : 61-80.
HALL, A., 1928-30. A revision of the genus Phyciodes Hubn. (Lepidoptera Nymphalidae). Bulletin of the Hill
Museum, Witley, 2 (Suppl.), 1-44; 3 (Suppl.), 45-170; 4 (Suppl.), 171-[205].
HARBORNE, J. B., 1982. Introduction to Ecological Biochemistry (2nd edition). London: Academic Press.
HARRIS, L., 1972. Buttegies of Georgia. Oklahoma: University of Oklahoma Press.
HARRISON, J. O., 1963. On the biology of three banana pests in Costa Rica (Lepidoptera: Limacodidae,
Nymphalidae). Annals of the Entomological Society of America, 56: 87-94.
HARVEY, D. J., in press. Observations on the biology and systematics of Junonia butterflies (Lepidoptera:
Nymphalidae) in Florida and the Bahamas. Journal of Research on the Lepidoptera.
HAYWARD, K. F., 1969. Datos para el Estudio de la Ontogenia de Lepidopteros Argentinos. Tucuman: Instituto
Miguel Lillo.
HECQ, J., 1980. Etudes des Euphaedra (Lepid. nymph, afr.), Note no 17. Lambillioneu, 79: 76-80.
196 P. R. ACKERY
HENNING, G. A., 1977. Observations on the early stages of Ethiopian Charaxes with notes on lifc historics
ymphalidae). Annals of the Transuaal Museum, 30: 2 19-230.
HENRIKSEN, H. J . & KREU’I‘ZER, I., 1982. The Butterflies of Scandinauia in ”Vature. Odensc: Skandinavisk
Bogforlag.
HESPENHEIDE, H. A., 1973. A novel mimicry complex: beetles and flies. Journal of b,‘nntorno/ogy (’4). 48:
49-55.
HEYWOOD, V. H . (Ed.), 1979. Flowering Plants ! / t h e World. Oxford: Oxford University Press.
HIGGINS, L. G., 1960. A revision of the Melitaeine genus Chlosyne and allied spccics (Lcpidoptcra:
Nymphalinae). Transactions of the Royal Entomolo,gical Society oJ. London, 112: 381-467.
HIGGINS, L. G., 1978. A revision of the genus Euphydryas Scudder (Lcpidoptera: Nymphalidac‘.
Entomologist’s Gazette, 29: 109-1 15.
HIGGINS, L. G., 1981. A revision ofPhyciodeJ Hiibner and rrlated gencsa, with a rrvirw of the classification
of the Mrlitaeinae (Lrpidoptera: Nymphalidae). Bulletin u f t h e British Muspurn (uVatural Histo!y) Entomdog>$.
43: 77-243.
HIGGINS, L. G . & RILEY, N. D., 1970. A Field Guide to the Butterflies of Europe. London: Collins.
HILL, D. S., JOHNSTON, G . T. & BASCOMBE, M. J., 1978. Annotated checklist of Hong Kone,
butterflies. Memoirs o f t h e Hang Kong Natural History Socieg, 11: i -iv, 1-62.
HICRA, I., 1983. Systematics in Hypolimnas [in Japanese]. Rhopalocerist’s Magazine, 6: 3-27.
HOFFMANN, b’.E., NG, Y. C. & TSANG, H. W., 1938. Life history studies in nine families of Kwangtung
butterflics (Lepidoptera: Rhopalorera). Lingnun Science Journal, 17: 227-246, 407-424, 5 15-532.
HOLLOWAY, J. D. & PETERS, J. V., 1976. The butterflies of New Caledonia and the Loyalty Islauds.
Journal oj”a/ura/ History, 10: 273-318.
HONDA, K., 197 1. Some observations on the ovipositing habit and the larval food plants o f Polygonia c-aurpnm
I.inne. Butterflies and Moths, Kyoto, 22: 49-52.
HOPKINS, G. H. E., 1927. Butterflies of Samoa and some other neighbouring island groups. Insects of Samoa
and other Samoan Terrestrial Arthropods, Part 3, Fasc. 1, 64 pp, 4 pls.
HOWARTH, 1‘. G., 1973. Colour Identification Guide to British Butterflies. London: Viking.
HOWE, W.H., 1975. The Butterflies OfN07th America. New York: Doubleday.
IFTNER, D. C., 1983. A new foodplant record for Chlogne gorgone carlota (Reakirt) (Nymphalidae). .7ournal vf
the Lepidopterists’ Sociely, 37: 80-8 I .
IGPIRASHI, S., 1984. The classification of the Papilionidae mainly based on the morphology of thc early
stagcs. Buttegies and Moths, hyoto, 34: 41-96.
IIVASE, T,, 1954. Synopsis of the known life-histories of Japanese butterflies. Lepidop/eris/c’ i\‘eu;s, 8: 95 -100.
IW’ASE, I..,1964. Recent foodplant records of the Loochooan butterflies. Journal of the I,epidop/Prists’ Sociec)’. 18:
105- 109.
J.4N%EN, D. H., 1983. Erblichia odorata Seem (‘lurneraceae) is a larval host plant of Eueides procula vulgiformis
(Nymphalidae: Hcliconiini) in Santa Rosa National Park, Costa Riea. Journal of the Lepzdopterists’ Sorze@,
37: 70-77.
JENKINS, D. h’.,1983. Neotropical Nymphalidae. I. Revision of Hamadryas. Bulletin of the Al!yn MuJeum. 81:
lLl46.
JENKINS, D. W., 1984. Neotropical Nymphalidae. 11. Revision of Mysc.elia. Bulletin of the rlllyn Museum. 87:
1 -~64.
JENKINS, D. W., 1985. Neotropical Nymphalidae. 111. Revision of Catonephele. Bulletin qf thr Allyn Mu.\eum,
92: 1-65.
JENSEN, S. R., NIELSEN, B. J. & DAHLGREN, R., 1975. Iridoid compounds, their occurrence and
systematic importance in the angiosperms. Botanisker Notiser, 128: 148-1 80.
JORDAN, K., 1913. Subfamily Acraeinae. In A. Seitz, Die Gross-Schmetter/inge der Erdc, Stuttgart (21 5:
358-374.
KASHIWAI, N,,1982. A new species of the genus Yfiythima Hubner from Mindanao, thc Philippines
(Lepidopterd, Satyridae). Butterflies and Moths, Kyoto, 33: 44-50.
KAU’AZOE, A,, 1956. O n the eggs and young caterpillars of Melitaea diamina protomrdia. Butterflies and Mob,
X j u t o , 7: 26-28.
KENDALL. R. O., 1959. More larval foodplants from Texas. Journal o f t h e LepidopterutJ’ Socze!y, 13: 221-228.
KENDALL. R. O., 1964. Larval foodplants for twenty-six species of Rhopalocera (Papilionoidca) from Tcxas.
Journal of the Lepidopterists’ Sociely, 18: 129-1 57.
KENDALL, R. O., 1978. Larval foodplant, life history notes and temporal distribution of S p l e r ~ d ~ u p ~ ~ ~ . h z ~ ~
kendalli (Satyridae) from Mexico. Journal uf 6he Lepidopterists’ Society, 32: 86-87.
KESSELRING, J . , 1975. The saving of a Morpho forest. A t a h , 3: 29-30.
KIM, C-W., 1984. A study of the distribution of butterflies and their feeding plants in Korea. Eri/ornolo,qicnl
Research Bulletin. Korea liniversity, 10: 35-124.
KIMBALL, C. P., 1965. Lepidoptera of Florida. Arthropods of Fh7ida and neighbourinz land areas, I : i Li,
1-363.
KIRCHBEKG, E., 1942. Genitalmorphologie und naturliehe Verwandtschaft der Amathusiinae und ihre
Rrziehungrn ziir geographischen Verbreitung der Subfamilie. Mitteilungen der Munchener Entomolo,pirhm
(;e.rrll.rchqfi, 32: 44-87.
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 197

KITCHING, I. J., 1985. Early stages and the classification of the milkweed butterflies (Lepidoptera:
Danainae). /=bologicalJournal of the Linnean Society, 85: 1-97.
KRISTENSEN, N. P., 1976. Remarks on the family-level phylogeny of butterflies (Insecta, Lepidoptera,
Rhopalocera). zeitschrft fur <oologische SJlstematik und Euolutiomforschung, 14: 25-33.
KURENTSOV, A. I., 1970. The Butterflies o f t h e Far East U.S.S.R. (ZdentiJication K v ) [in Russian]. Leningrad.
KUZUYA, T., 1967. Natural foodplants of Kirinia epaminondas Staudinger. ButterJIies and Moths, Kyoto, 18:
24-25.
LAIDLAW, W. B. R., 1970. Butterflies of New zealand. Auckland and London: Collins.
LAMBKIN, T. A. & LAMBKIN, K. J., 1977. Observations on the life history of Argynnis hyperbius inconstans
Butler (Lepidoptera: Nymphalidae). Australian Entomological Magazine, 4: 13-16.
LANE, C., 1957. Preliminary note on insects eaten and rejected by a tame sharma (Kittacincla malabarica) with
the suggestion that in certain species of butterflies and moths females are less palatable than males.
Entomologist‘s Monthly Magazine, 93: 172- 179.
LANCER, T. W., 1958. Nordens Dagsommerfugle. Copenhagen: Munksgaard Forlag.
LARSEN, T . B., 1974. Butterflies of Lebanon. Beirut. National Council for Scientific Research.
LARSEN, T. B. & LARSEN, K., 1980. Butterflies of Oman. Edinburgh: John Bartholomew and Son.
LEECH, J. H., 1892-4. Butteflies f r o m China, Japan and Corea. London: Taylor & Francis.
LE MOULT, E., 1950. Revision de la classification des Apaturinae de I’ancien monde suivie d’une
monographie de plusieurs genres. Miscellanea Entomologica (Suppl.).
LE MOULT, E. & REAL, P., 1962. Les Morpho d’AmCrique du sud et centrale: historique, morphologie,
systkmatique. Nouitates Entomologicae (Suppl.).
LENCZEWSKI, B., 1980. ButterJIies of Everglades National Park. South Florida Research Center Report T-588.
LESTON, D., SMITH, D. S. & LENCZEWSKI, B., 1983. Habitat, diversity and immigration in a tropical
island fauna: the butterflies of Lignumvitae Kay, Florida. Journal of the Lepidopterists’ Society, 36: 241-255.
LONGSTAFF, G. B., 1912. Butterfly-hunting in many lands. London.
MADDOX, G . D. & CANNELL, P. F., 1983. The butterflies of Kent Island, Grand Manan, New Brunswick.
Journal of the Lepidopterists’ Society, 36: 264-268.
MALCOLM, S. & ROTHSCHILD, M., 1983. A danaid Miillenan mimic, Euploea core amymone, lacking
cardenolides in the pupal and adult stages. Biological Journal of the Linnean Society, 19: 27-33.
MAMET, R., 1948. A food-plant catalogue of the insects of Mauritius. Bulletin, Department of Agriculture,
Mauritius, 30: 1-74.
MANDERS, N., 1908. The butterflies of Mauritius and Bourbon. Transactions of the Entomological Society of
London, 1907: 429-454.
MANLEY, W. B. L. & ALLCARD, H. G., 1970. A Field Guide to the Butterflies and Burnets of Spain. Hampton:
E. W. Classey Ltd.
MANSELL, G., 1985. Breeding Erebia species and their early stages. Proceeding of the 3rd Congress of European
Lepidopterology, Cambridge, 1982: 114-1 19. Karlsruhe: Societas Europaea Lepidopterologica.
MARSH, N. A,, CLARKE, C . A,, ROTHSCHILD, M. & KELLET, D. N., 1977. Hypolimnas bolina (L.), a
mimic of danaid butterflies, and its model Euploea core (Cram.) store cardioactive substances. Nature,
London, 268: 726-728.
MASTERS, J. H., 1968. Collecting Ithomiidae with heliotrope. Journal of the Lepidopterists’ Society, 22:
108-1 10.
MASTERS, J. H., 1969. Butterflies of Lynch Hollow, Camden County, Missouri. Journal of the Kansas
Entomological Society, 42: 133-141.
MASTERS, J. H., 1971. A note on Lethe anthedon borealis (Satyridae). Journal of the Lepidopterists’ Society, 25:
256-26 1.
MASTERS, J . H. & SORENSEN, J. T., 1969. Field observations on forest Oeneis (Satyridae). Journal o f t h e
Lepidopterists’ Society, 23: 155-1 61.
McCUBBIN, C., 1971. Australian Butterflies. Melbourne: Thomas Nelson.
MAZEL, R., 1982. Exigences trophiques et tvolution dans les genres Euphydyac. et Melitaea sensu lato (Lep.
Nymphalidae). Annales de la SociW Entomologique de France (N.S.), 18: 21 1-227.
McALPINE, W. S., HUBBELL, S. P. & PLISKE, T . E., 1960. The distribution, habits and life history of
Enptychia mitchellii (Satyridae). Journal of the Lepidopterists’ Society, 14: 209-226.
MICHENER, C. D., 1942. A generic revision of the Heliconiinae. American Museum Nouitates No. 1197.
MICHENER, C. D., 1943. Some systematic notes on the Libytheidae (Lepidoptera). American Museum
Jouitates No. 1232.
MILES MOSS, A. Unpublished drawings. British Museum (Natural History) collection.
MILES MOSS, A,, 1933. Some generalizations on Adelpha, a neotropical genus of nymphalid butterflies of the
group Limenitidi. Nouitates Zoologicae, 39: 12-20.
MILLER, L. D., 1968. The higher classification, phylogeny and zoogeography of the Satyridae. Memoirs of the
American Entomological Society No. 24.
MILLER, L. D. & MARTIN BROWN, F., 1981. A catalogue/checklist of the butterflies of America north of
Mexico. Lepidopterists’ Society Memoir No. 2.
MILLER, J. S., 1986. Phylogenetic systematics and chemical constraints on host-plant associations in the Papilioninae
(Lepidoptera: Papilionidae) . Unpublished Doctoral thesis, Cornell University.
I98 P. R. ACKERY

MORI, I., 1958. Life history of Fabriciana nerippe Felder & Felder. Butte$ies and M o t h , Kyoto, 9; 54-56.
MORISHI’I‘A, K., 1980. The Nama group of the genus Hestina (Lep. Nymphalidae). Rhopalocerist’s Magazine, 3
( 3 ) : 28 pp.
.MULLER, F., 1878. Uber die Vortheile der Mimicry bei Schmetterlinge. zoologische Anzezger, I : 54-55.
MULLER, F., 1879. Ituna und Thyridia. Kosmos, 5: 100-108. [English translation by R. Meldola, under the
title Ituna and Thyridia: a remarkable case of mimicry in butterflies, appears in the Proceedin..s OJ the
Entomological Society of London, 1879: xx-xxix.]
MULLER, W., 1886. Sudamerikanische Nymphalidenraupen: Versuch eines natiirlicheo Systems der
Nymphaliden. zoolosixhe Jahrbiicher, I : 41 7-678.
MULLER, J., 1969. A new foodplant for Eupbdryasphaeton (Nyrnphalidae).Journal o f t h e Lepidopterists’ Sociep,
23’: 48.
MUNROE, E., 1961. The classification of the Papilionidae. Canadian Entomologist (Suppl.) No. 17.
MUROYA, Y., KUBO, K., MAEDA, K., ASHIZAWA, H. & OHTSUKA, K., 1967. The early stages of
Formosan butterflies. Special Bulletin of the ~epidopterologicalSociety of Japan, 3: 5 1-60.
MURPHY, D. D. & EHRLICH, P. R., 1984. Biosystematics of the Euphydryas of the Central Great Basin,
with description of a new subspecies. Journal of Research on the Lepidoptera, 22: 254-261.
MUYSHONDT, A,, 1973a. Notes on the life cycle and natural history of butterflies of El Salvador. IA--
Catonephele numilia esite (Nymphalidae-Catonephelinae). Journal of the New York Entomological Society, 81:
164-1 74.
MUYSHONDT, A,, 197313. Notes on the life cycle and natural history of butterflies of El Salvador 1IA.P-
Ep$hile adrasta adrasta (Nymphalidae-Catonephelinae). Journal of the N e w York Entomological Society, 81:
214-223.
MUYSHONDI‘, A., 1973c. Notes on the life cycle and natural history of butterflies of El Salvador IIIA.--
‘remenis laothoe liberia Fabricius (Nymphalidae-Catonephelinae). Journal of the N e w York Entomological
Society, 81, 224-233.
MUYSHONDI’, A,, 1973d. Notes on the life cycle and natural history of butterflies of El Salvador 1VA.--
Pxudonica,flauilla canthara (Nymphalidae-Catonephelinae). Journal of the N e w York Entomological Society, 81:
234-242.
MUYSHONDT, A,, 1973e. Notes on the life cycle and natural history of butterflies of El Salvador. I. Prepona
omphale octauia (Nymphalidae). Journal of the Lepidopterists’ Society, 27: 210-2 19.
MLYSHONDT, A,, 1973f. Notes on the life cycle and natural history of butterflies of El Salvador. 11. Anaea
(zaretis) itys (Nymphalidae). Journal of the Lepidopterists’ Society, 27: 294-302.
.MUYSHONDT, A,, 1974a. Notes on the life cycle and natural history of butterflies of El Salvador. 111. Anaea
(Consul) fabius (Nymphalidae). Journal of the Lepidopterists’ Society, 28: 8 1-89.
MUYSHONDT, A., 197413. Notes on the life cycle and natural history of butterflies of El Salvador. 1V. Anaea
(Memphis) turyale confusa (Nymphalidae). Journal o f the Lepidopterists’ Society, 28: 306-3 14.
MUYSHONDT, A,, 1 9 7 4 ~ .Notes on the life cycle and natural history of butterflies of El Salvador. VX.
Pyrrogyra hypsenor (Nymphalidae-Catonephelinae). Journal of the New York Entomological Sotiely, 82;
163-1 72.
MUYSHONDT, A., 1975a. Notes on the life cycle and natural history of butterflies of El Salvador. V. Anaea
(A4emphis) moruus boisduuali (Nymphalidae). Journal of the Lepidopterists’ Society, 29: 32-39.
MUYSHONDT, A,, 1975b. Notes on the life cycle and natural history of butterflies of El Salvador. V I A -
Diaethria astala Guerin (Nymphalidae-Callicorinae). Journal of the N e w York Entomological Sociely, 83:
10-18.
MUYSHONDT, A,, 1 9 7 5 ~Notes . on the life cycle and natural history of butterflies of El Salvador. VI. Anaea
(Memphis) pithyusa (Nymphalidae). Journal o f the Lepidopterists’ Society, 29: 168- 176.
MUYSHONDT, A., 1976a. Notes on the life cycle and natural history of butterflies of El Salvador. V I I .
Archaeprepona demophon centralis (Nymphalidae). Journal of the Lepidopterists’ Society, 30: 23-32.
MUYSHONDT, A,, 1976b. Notes on the life cycle and natural history of butterflies of El Salvador. V I I I .
iirchaeoprepona antimache gulina, Siderone marthesia, zaretis callidryas and Consul electra (Nymphalidae).Journal of
the Lepidopterists’ Society, 30: 159-168.
MUYSHONDT, A. & MUYSHONDT, A. JR, 1975. Notes on the life cycle and natural history orbutterflies
of El Salvador. IB-IIB. (Nymphalidae-Hamadryadinae). Journal of the New York Entomological Society, 83:
157-169, 170-180, 18 I - 191.
MUYSHONDT, A,, MUYSHONDT, A. J R & MUYSHONDT, R., 1976. Notas sohre la biologia de
lcpidopteros de El Salvador. Reuista de la Sociedad Mexicana de Entomolog.ia, 2: 77-90.
MVYSHONDT, A. J R & MUYSHONDT, A,, 1976. Notes on the life cycle and natural history of butterflies
of El Salvador. IC. Colobura dirce L. (Nymphalidae-Coloburinae). Journal of the Nen‘eut York Entomological
Socieo, 84: 23-33.
MUYSHONDT, A. J R & MUYSHONDT, A,, 1978. Notes on the life cycle and natural history of butterflies
of El Salvador. IIC. Smyrna blomjldia and S. karwinskiz (Nymphalidae: Coloburini). Journal of the
Lepidopterists’ Society, 32: 160- 174.
MUYSHONDT, A. JR & M U Y S H O N D I , A,, 1979. Notes on the life cycle and natural history of butterflies
of El Salvador. IIIC. Historis odius and Coea acheronta (Nymphalidae-Coloburinae), Journal of the
Lepidopterists’ Society, 33: 1 12-1 23.
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 199

NAHRSTEDT, A. & DAVIS, R. H., 1981. The occurrence of cyanoglucosides, linamarin and lotaustralin, in
Acraea and Heliconius butterflies. Comparative Biochemistry and Physiology, 68B: 575-577.
NAHRSTEDT, A. & DAVIS, R. H., 1983. Occurrence, variation and biosynthesis of the cyanogenic
glucosides linamarin and lotaustralin in species of the Heliconiini. Comparative Biochemistry and Physiolou,
75B: 65-73.
NECK, R. W., 1973. Foodplant ecology of the butterfly Chlosyne lacinia (Geyer) (Nymphalidaej. I . Larval
foodplants. Journal of the Lepidopterists’ Society, 27: 22-33.
NECK, R. W., 1976. Lepidopteran foodplant records from Texas. Journal of Research on the Lepidoptera, 15:
75-82.
NEGISHI, K., 1971. Butterflies collecting tour across Venezuela. 11. Attracting Ithomiinae with the herb,
“Rabo de Alcran” in the vicinity of Caracas. Yadoriga, 68: 17-26.
NEL, J., 1979. Une nouvelle plante nourriciere pour Charaxes jasius (Lep., Nymphalidae). Alexanor, 11:
157- 158.
NEL, J., 1982. Contribution a I’ktude des premiers Ctats chez les Erebia. Un Clevage d’Erebia scipio Bdv. (Lep.
Satyridae). Alexanor, 12: 283-287.
NGUYEN, T. H., 1985. Le polymorphisme et le mimPtisme d’ Zdrusia nyctelius Doubleday [Lep.
Nymphalidae]. Bulletin de la Sociiti Entomologique de France, 90: 1083-1 102.
NICULESCU, E. V., 1965. Fauna Repubtica Populare Rotnane. Insecta I1 (fasc. 7) Familia Nymphalidae.
Bucharest: Editura Academiei RPR.
OEHMIG, S., 1983. Hipparchia azorina (Strecker, 1899) (Satyridaej; biology, ecology and distribution on the
Azores Islands. Journal of Research on the Lepidoptera, 20: 136-160.
OUATTARA, S., JOLIET, P. & VAN PARYS, E., 1977. Seconde Lists des Insectes et des Plantes-hiites en Haute-
Volta et dans les Rigions Limitrophes. Bobo-Dioulasso.
OWEN, D. F., 1970. Mimetic polymorphism and the palatability spectrum. Oikos, 21: 333-336.
OWEN, D. F. & OWEN, J., 1972. Systematics and bionomics of butterflies seen and collected in the forest
region of Sierra Leone. Part 1. Introduction, Papilionidae, Danaidae and Acraeidae. Revue de zoologie et de
Botanique Aj’ricaines, 85: 287-308.
OWEN, D. F. & OWEN, J., 1973. Systematics and bionomics of butterflies seen and collected in the forest
region of Sierra Leone. Part 2. Nymphalidae and Libytheidae (Lepidoptera). Revue de zoologie et de
Botanique Africaines, 87: 585-6 13.
PAGENSTECHER, A., 1901. Libytheidae. Tierreich, 14.
PAPAGEORGIS, C., 1975. Mimicry in Neotropical butterflies. American Scientist, 63: 522-532.
PARSONS, M., 1984. Life histories of Taenaris (Nymphalidae) from Papua New Guinea. Journal of the
Lepidopterists’ SocieQ, 38: 69-84.
PARSONS, M., in prep. Early Stages of the Butteflies of New Guinea, in Colour.
PAULIAN, R., 1956. Insectes: Lepidopteres; Danaidae, Nymphalidae, Acraeidae. Faune de Madagascar, 2:
1-102.
PENNINGTON, K . M., 1978. Butteflies of Southern Africa. C. G. C. Dickson and D. M: Kroon, (Eds).
Johannesburg: Ad. Donker.
PERKINS, E. M. & MEYER, W. C . , 1973. Revision of the Boloria epithore complex, with description of two
new subspecies (Nymphalidae). Bulletin of the Allyn Museum No. 11.
PFITZNER, J. C. & FARGHER, R. K., 1976. Butterflies of central Australia. Australian Entomological
Magazine, 2: 117-122.
PHOLBOON, P., 1965. A Host List o f t h e Insects of Thailand. Bangkok: Thailand Department of Agriculture.
PIEPERS, M. C. & SNELLEN, P. C. T., 1909-18. T h e Rhopalocera ofJava. The Hague: Martinus Nijhoff.
PIERRE, J., 1979. Le complexe ultraspkcifique d’dcraea admatha Hewitson, reconnaissance de 6 espkces-
jumelles, description de nouveaux taxons (Lepidoptera Acraeidae). Annales de la Sociiti Entomologique de
France (N.S.). 15: 719-737.
PIERRE, J., 1981a. Une nouvelle esptce d’Acraea dans le complexe ultra-spkcifique ‘encedon’. Bulletin de la
Sociiti Entomologique de France, 86: 79-87.
PIERRE, J., 1981b. Vicariance de deux prospecies Acraea natalica et Acraea pseudegina (Lepid.--Nymphalidae).
Lambillionea, 81: 17-22.
PIERRE, J., 1981~.Deux esptces jumelles confondues sous le nom Acraea alciope Hewitson (Lepidoptera
Acraeidae). Annales de la Sociktk Entomologique de France N S . , 17: 349-357.
PIERRE, J., 1984. Systimatique euolutiue cladistique el mimitisme chet les lipidopt2res de genre Acraea. DSc thesis,
University of Paris.
PIERRE, J. & VUATTOUX, R., 1978. Les Acraea de CBte d’lvoire (Lep. Acraeidae). Bulletin de la Sociiti
Entomolgique de France, 83: 3-22.
PIERRE-BALTUS, C., 1978. Rtsultats d’klevages de Neptis A facies ‘melicerta’ en CBte d’Ivoire; description
de trois nouvelles especes. Lambillionea, 78: 33-44.
PINHEY, E. C. G., 1949. Butterj‘lies of Rhodesia, with a Short Introduction to the Insect World. Salisbury: Rhodesia
Scientific Association.
PLATT, A. P., COPPINGER, R. P. & BROWER, L. P., 1971. Demonstration of the selective advantage of
mimetic Limenitis butterflies presented to caged avian predators. Euolution, Lancaster, PA, 25: 692-
701.
200 P. R. ACKERY

PLISKE, T. E., 1975. Attraction of Lepidoptera to plants containing pyrrolizidine alkaloids. Enuironmrntal
Entomolo~y3 4: 455-473.
POLHILL, R. M., RAVEN, R . H . & STIRTON, C. H., 1981. Evolution and systematirs of thc
Leguminosae. In R . M . Polhill & P. H. Raven, Advances in Legume Systrmatics: 1-26. Kew. London: R(J);:I~
Botanical Gardens.
PYLE, R . M., 1974. Watching Washington buttelflies. Seattle: Audubon Society.
R..\NCOLR'L', D., 1979. I.es Roloria (Lepidoptera: Nymphalidae). Fabreries, 6: 48-52.
R..\!VSOK, G. W., 1976. Notes on the biology and immature stages of the White Peacock butterfly, Annrlia
jairophae guanianamo (Nymphalidae). Journat of the Lepidopterists' Society, 30: 207-210.
REICHSTEIN. T., 1967. Cardenolide (herzwirksame Glyroside) als Abwehrstoffc bci Insekten.
.'laturwissensrhafiliche Rundschau, 20: 499-5 1 1 .
REICHSTEIN, T., EUW, J. V., PARSONS, J. A. & ROTHSCHILD, M.. 1968. Heart poisons in the
Monach butterfly. Science, .Vew York, 161: 861-866.
REMINGTON, C. L., 1952. The biology of Nearctic Lepidoptera I. Foodplants and life-histories of Colorado
Papilionoidea. P y - h e , 59: 61-70.
RILEY, N. D., 1975. Butte$es ofthr West Indies. London: Collins.
RITLAND. D. B. & BROWER, L. P., 1986. A reassessment of the mimicry relationship among Viceroys,
Monarchs and Queens in Florida. Abstracls of the Second Inlernational Monarch Conference, Los Angeles p. 5.
KOBBINS, R. K. & AIELLO, A,, 1982. Foodplant and oviposition records for Panamanian Lycaenidae and
Riodinidae. Journal of the Lepidopterists' Society, 36: 65-75.
ROBERT, J. H., ESCARRE, A,, GARCIA, T. & MARTINEZ, P., 1983. Lepiddpleros Rhopaloceros sus plantas
nutricias y su dislribucion geogrdphica en la Prouincza de Alicante. Alicante: Instituto de Estudios Alicantinoa
ROBINS, D. J., 1971. Senecioneae-chemical review. In V. H . Heywood, J. B. Harhornc Br B. L. Turner,
The Biology and Chemistry o f t h e Compositae, 2: 83 1-850. London: Academic Press.
ROBINSON, G. S., 1975. Macrolepidoptera o f F i j i and Rotuma: a taxonomic study. Faringdon: E. M'. Classey
Ltd.
KOSEVEAR, R., 1978. A few Lepidopterous life-histories. Nigerian Field, 43: 177-181.
ROSS, G. N., 1975-77. An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz, Mexico.
Journal of Research on the Lepidoptera, 14: 103-124, 169-188, 233-252; 15: 41-60, 109-128, 185- 200,
225-240; 16: 87- 130.
ROTHSCHILD, M., 1971. Speculations about mimicry with Henry Ford. In R . Creed, Ecological Genetics and
Euolution: 202-223. Oxford: Blackwells.
ROTHSCHILD, M . & EDGAR, J. A,, 1978. Pyrrolizidine alkaloids from Senecio uulgaris sequestered and
stored by Danaus plexippus. Journal of /=bology, 186: 347-349.
ROTHSCHILD, M . & MARSH, N., 1978. Some peculiar aspects of Danaid/plant relationships. Entomologza
Experimentalis el Applicata, 24: 637-650.
ROTHSCHILD, M., MARSH, N. & GARDINER, B., 1978. Cardioactive suhstancrs in the Monarch
hutterfly and Euploea core reared on a leaf-free artificial diet. Nature, London, 275: 649-650.
RO'I'HSCHILD, .M., NASH, R. J. & BELL, E. A., 1986. Cycasin in the endangered butterfly Eumaeus a t a h
Jorida. Phytochemisly, 25: 1853- 1854.
RO'I'HSCHILD, W., 1916. Notes on Amathusiidae, Brassolidae, Morphidae, etc, with descriptions of new
forms. hbritates Zoologicae, 23: 299-3 18.
RYDON, A. H. B., 197 I . The systematics of the Charaxidae. Entomologist's Record and Journal of Variation, 83:
219-233, 283-287, 310-316, 336-341, 384-388.
SAMSON, C., 1980. The butterflies of Santa Aria Island, with descriptions of new taxa from San Cristobal
Island, Solomon Islands. Butterflies and Moihs, Kyolo, 30: 21 1-236.
SANKOIVSKY, G., 1975. Some ncw foodplants for various Queensland butterflies. Australian Entomolo,qical
.2fagazine, 2: 55-56.
SANKOWSKY, G., 1978. Some new food plants for various Queensland butterflies. Australian Entornolo'qical
Magazine, 5: 77-79.
SCHEERMEYER. E. & ZALUCKI, M. P., 1985. Food plant records for Eubloea core corinna (W. S. Maclca);)
with some notes on larval colouration. dusiralian Entomological Magazine, I I : 87-90.
SCHWARZ, R., 1948-9. Motyli [Butterflies of Czechoslovakia] 2 vols. Prague: Vesmir, Nakladetelska a
\'ydavetelska. [In Czechoslovakian.]
SCOBLE, M. J,, 1986. The structure and affinities of the Hedyloidea: a new concept of the butterflies. Bulletin
of the Briiish Museum (Natural History) Entomology, 53: 25 1-286.
SCO'I'T, J . A,, 1974. 'The early stages and biology of Phyciodes orseis (Nymphalidae). Journal of Re~earchon the
I>epidoptera, 12: 236-242.
SCOTT,J . A., 1976. Early stages of Phyciodes pallida, P. orseis and P. mylztta (Nymphalidae). .fournal of Resenrrh
on the Lepidoplera, 14: 84.
SCOTT. J. A,, 1985. The phylogeny of butterflies. journal of Research on the I.epidoptera, 23: 241-281.
SCOTT, J. A. & SCOTT, G. R., 1980. Ecology and distribution of the butterflies of south central Colorado.
Journal of Research on the Lepidoptera, 17: 73-128.
SEITZ, .4.,1914. Group: Gymaeciidae. In A. Seitz, Gross-Schmetterlinge der Erde, Stuttgart, (2) 5: 470-473
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 20 1
SEITZ, A., 1927. Das System der Schmetterlinge. Entomologische Rundschau, 44: 32, 35-36, 39-40; 44: 47-48.
SEVASTOPULO, D. G . , 1973. The foodplants of Indian Rhopalocera. Journal of the Bombay Natural History
SocieQ, 70: 156-183.
SEVASTOPULO, D. G . , 1975. A list of the foodplants of East African Macrolepidoptera. Part 1. Butterflies
(Rhopalocera). Bulletin of the Amateur Entomologists Society, 34: 84-92, 124-132.
SEVASTOPULO, D. G . (unpublished).East African Lepidoptera. 12 vols. BMNH Collection.
SHAPIRO, A. M., 1974. Butterflies of the Suisan Marsh, California. J o ~ r n a lof Research on the Lepidoptera, 13:
191-206.
SHAPIRO, A. M., 1982. Notas sobre 10s estudios inmaduros de la mariposa plateada, Argyrophorus argenteus
Blanchard (Lepidoptera: Satyridae). Reuista de la Sociedad Mexicana de Lepidopterologica, 7: 29-3 1.
SHAPIRO, A. M., PALM, C. A. & WCISLO, K. L., 1981. The ecology and biogeography of the butterflies
of the Trinity Alps and Mount Eddy, northern California. Journal of Research on the Lepidoptera, 18: 69-152.
SHAPIRO, A. M. & SHAPIRO, A. R., 1973. The ecological associations of the butterflies of Staten Island.
Journal of Research on the Lepidoptera, 12: 65-1 26.
SHIELDS, O., 1985. Zoogeography of the Libytheidae (Snouts or Breaks [sic]). Tokurana, 9: 1-58.
SHIELDS, O., EMMEL, J. F. & BREEDLOVE, D. E., 1969. Butterfly larval foodplant records and a
procedure for reporting foodplants. Journal of Research on the Lepidoptera, 8: 21-36.
SHIROZU, T., 1944. Notes on the foodplants of the larvae of some butterflies observed in the Kyushu
District. Transactions of the Kansai Entomological Society, 14: 48-59.
SHIROZU, T . , 1960. Butterfies of Formosa in colour. Osaka: Hoikusha.
SHIROZU, T. & SAIGUSA, T., 1971. A new genus of the subfamily Apaturinae (Nymphalidae). ButterJIies
and Moths, Kyoto, 22: 7-13.
SILBERGLIED, R. E., 1983. Anartia fatima (Cocinera, Whitebanded Fatima). In D. H. Janzen, Costa Rican
Natural History, 682-683. Chicago: University of Chicago Press.
SILBERGLIED, R., AIELLO, A. & LAMAS, G . , 1979. Neotropical butterflies of the genus Anartia:
Systematics, life histories and general biology. Psyche, 86: 2 19-260.
SINGER, M. C., 1984. Butterfly-hostplant relationships: host quality, adult choice and larval success. In R. I.
Vane-Wright & P. R. Ackery, The Biology of Butterflies: 81-88. Symposia of the Royal Entomological
Society of London, No. 1 1.
SINGER, M. C., EHRLICH, P. R. & GILBERT, L. E., 1971. Butterfly feeding on a lycopsid. Science, N e w
York, 172: 1341-1342.
SINGER, M. C., DEVRIES, P. J. & EHRLICH, P. R., 1983. The Cissia conzusa species-group in Costa Rica
and Trinidad (Lepidoptera: Satyrinae). <oological Journal o f t h e Linnean Society, 79: 101-1 19.
SMILES, R. L., 1982. The taxonomy and phylogeny of the genus Polyura Billberg (Lepidoptera:
Nymphalidae). Bulletin of the British Museum (Natural History) Entomology, 44: 115-237.
SMILES, R. L., 1985. Cladistics and distribution of Euxanthe butterflies (Nymphalidae: Charaxinae). Journal
of Natural History, 19: 1165-1 189.
STAMP, N. E., 1979. New oviposition plant for Euphydryas phaeton (Nymphalidae). j‘ournal of the Lepidopterists’
Society, 33: 203-204.
STICHEL, H., 1909. Brassolidae. Tierreich, 25.
STICHEL, H., 1925. Zur Systematik der Brassolidae. Jdeue Beitrage zur Systematischen Insektenkunde, 3: 58-60.
STICHEL, H., 1932. Brassolidae. Lepidoptorum Catalogus, 27 (51).
STICHEL, H., 1933. Amathusiidae. Lepidoptorum Catalogus, 27 (54).
SWEZEY, 0. H., 1942a. Lepidoptera in Samoa. Proceedings of the Hawaiian Entomological Society, 11: 206-216.
SWEZEY, 0. H., 1942b. Butterflies of Guam. In Insects of Guam, I : 31-38. Honolulu: Bernice P. Bishop
Museum.
SWYNNERTON, C. F. M., 1915a. Birds in relation to their prey. Experiments on wood-hoopoes, small
hornbills and a babbler. Journal of the South Afn’can Ornithologists’ Union, 1915: 22-108.
SWYNNERTON, C. F. M., 1915b. Concluding discussion. In G . D. H. Carpenter (1942), Observations and
experiments in Africa by the late C. F. M. Swynnerton on wild birds eating butterflies and the preferences
shown. Proceedings of the Linnean Society of London, 154: 10-46.
SWYNNERTON, C. F. M., 1919. Experiments and observations bearing on the explanation of form and
colouring, 1908-1913, Africa. Journal of the Linnean Society of London (zoology), 33: 203-385.
SWYNNERTON, C. F. M., 1926. An investigation into the defences of butterflies of the genus Charaxes.
Proceedings of the International Congress of Entomology (3) 2: 478-506.
TABUCHI, Y., 1959. The Alpine Butterflies ofJapan. Tokyo: Hobundo Publishing Company.
TAKAHASHI, M., 1974-5. On the larva and pupa of Mycalesis gotama madjicosa Butler (Lepidoptera:
Satyridae) from Ishigaki Island, the Ryukyus. ButterJIies and Moths, Kyoto, 25: 131-135; 26: 74-76.
TANAKA, B., 1978. Larval food-plants and distribution of Japanese Ladoga (Lepidoptera: Nymphalidae).
ButterJIies and Moths, Kyoto, 29: 35-45.
TANI, S., 1983. Geographic variation in photoperiodic induction of pupal diapause of Neope goschkeuitschi
Mtnktrits (Lepidoptera, Satyridae). Butterflies and M o t h , Kyoto, 33: 157-16 I .
TIETZ, H. M., 1972. A n Index to the Described LZfe-Histories. Early Stages and Hosts of the Macrolepidoptera of
Conjinental United States. 2 vols. Sarasota, Florida: A. C. Allyn.
202 P. R. ACKERY

TILDEN, J . W., 1965. ButterJlies of the San Francisco Bay Region. Berkeley: University of California Press.
TREMEWAN, W. G., 1960. A list of the foodplants of some species of the lepidopterous family Zygaenidar.
Entomologist, 93: 108-1 11.
TURNER, J. R. G., 1975. A tale of two butterflies. Natural History, New York, 84: 28-37.
TURNER, J. R . G., 1977. Butterfly mimicry: the genetical evolution of an adaptation. Euolutionary Biology, 10:
163-206.
TURNER, T. W. & PARNELL, J. R., 1985. The identification of two species of Junonia Huhnrr
(Lepidoptera: Nymphalidae): 3. evarete and 3. genoveva in Jamaica. j‘ournal of Research on the Lepidoptera, 24:
142-153.
UCHIDA, H., 1984. Records of the foodplants of45 Formosan butterfly species. Butterjzes and M o h , kjoto, 3:
152-160.
UCHIDA, H., 1985. Records of the foodplants of 17 Formosan butterfly species. ButterJlies and Moths, ITyoto, 35:
208-210.
URICH, F. C. & BOOS, J. O., 1980. Metamorphosis of Dynastor macrosiris Westw. (Lepidoptera Brassolidae:.
Living World, 1981: 34.
VALLETTA, A., 1972. Butterjies of the Maltere Islands. Malta.
VANE-WRIGHT, R. I., 1971. The systematics of Drusillopsis Oberthiir (Satyrinae) and the supposed
Amathusiid Bigaena van Eecke (Lepidoptera: Nyrnphalidae) . Transactions of the Royal Entomological
Society of London, 123: 97-123.
VANE-WRIGHT. R. I., 1972. Pre-courtship activity and a new scent organ in butterflies. Nature, London, 239:
338-339.
L’ANE-WRIGHT, R. I., 1974. Further observations on the occurrence and mimicry of Mycalesis drusillodes
(Lepidoptera: Nymphalidae, Satyrinae). Journal of Entomology, London ( B ) , 42: 2 13-2 16.
VANE-WRIGHT, R. I., 1976. A new Mynes butterfly belonging to the Telleruo mimicry complex. Journal of
Natural History, 10: 409-413.
VANE-WRIGHT, R . I. & SMILES, K. L., 1975. ‘Ihe spccies of’ the genus Zethera k’eldcr (Lepidoptera:
Nymphalidae, Satyrinae). Journal of Entomology, London ( B ) , 44: 8 1-100.
VL4NE-WRIGHT, R. I., ACKERY, P. R . & SMILES, R. L., 1977. The polymorphism, mimicry and
hostplant relationships of Hypolimnas butterflies. Biological .journal of the Linnean Society, 9: 285-297.
VAN SOMEREN, V. G. L., 1974. List of the foodplants of some East African Rhopalocera, with notes 011 the
early stages of some Lycaenidae. Journal of the Lepidopterists’ Society, 28: 3 15-33 1.
VAN--SOMEREN, V. G. L., 1975. Revisonal notes on the African Charaxes, Palla and Euxanthe (Lepidoptera:
Nymphalidae). Bulletin of the British Museum (Natural History) (Entomology), 32: 65- 136.
VAN SOMEREN, V. G. L. & ROGERS, K.ST A,, 1925-31. Butteflies of Kenya and Uganda. Nairobi: East
African Natural History Society.
VAN SON, G., 1979. The Butterflies of Southern Africa. Part IV Nymphalidae: Nymphalinae. Pretoria:
Transvaal Museum Memoir No. 22.
VUATTOUX, R. & BLANDIN, P., 1977. Les LCpidopttres Charaxinae (Nymphalidae) rCcoltCs a la Station
d’Ecologie Tropicale de Lamto (CBte d’Ivoire). Annales de 1’Universiti #Abidjan (Ecologie), 10: 7-26.
WARREN, B. C. S., 1944. Review of the classification of the Argynnidi, with a systematic revision of the
genus Boloria. Transactions of the Royal Entomological Society of London, 94: 1-102.
WKYIISON, J. T., 1928-30. Lepidopteros de Portugal. Memorias e Estudos de Museu ~oologicode Uniuersidade de
Coimbra No. 29.
WEYMER, G., 1910-12. Family: Satyridae. In A. Seitz, Gross- Schrnetterlinge der Erde, Stuttgart, ( 2 ) 5: 173-283.
WHALLEY, P., 1986. A review of the current fossil evidence of Lepidoptera in the Mesozoic. Biologzral Journal
of the Linnean Society, 28: 253-271.
WHITE, R. H., 1979. Foodplant of alpine Euphydryas anicia (Nymphalidae). Journal of the Lepidopterzsts’ Socieiy,
3.5: 170-173.
WHITE, R. W. & SINGER, M. C., 1974. Geographical distribution of hostplant choice in Euphydryas editha
(Nymphalidae). Journal o f t h e Lepidopterirts’ Society, 28: 103-107.
WILEY, A. J. & HUDSON, J. R., 1943. Some common butterJlies o f t h e Nazrobi district. Nairobi: East African
Natural History Society.
\\’ILLIS, J . C., 1980. A Dictionary of the Flowering Plants and Fe7ns. (8th edition, revised by H. K. Airy Shaw.)
Cambridge: Cambridge University Press.
WILSON, A., 1986. Flavonoid pigments and wing color in Melanargia galathea. Journal of Chemical Ecology, 12:
49.- 67.
WILSON, W. J., 1982. Notes on life histories of some Queensland butterflies. Circular. Entomological Sectzori of
the Royal Sociely of New South Wales, 21: 1-2.
WILTSHIRE, E. P., 1957. The Lepidoptera of Iraq. London: Nicholas Kaye Limited.
WOOD, G. A., 1984. The life history of Elymnias agondas australiana Fruhstorfer (Lepidoptera: Nynphalidarj.
Australian Entomological Magazine, 11: 41-43.
WOOD, G. A., 1986. Some early stages of Charaxes latona Butler (Lepidoptera: Nymphalidae; Charaxiriar\.
‘lustralian Entomological Magazine, 13: 20-2 1.
WOODHOUSE, L. G. O., 1952. The ButterJly Fauna of Ceylon (2nd Edition). Colombo: The Colombo
Apothecaries’ Company.
 HOSTPLANTS AND CLASSIFICATION: NYMPHALID BUTTERFLIES 203
WYNTER-BLYTH, M. A., 1957. ButterJlies of the Zndian region. Bombay: Bombay Natural History Society.
YOUNG, A. M., 1972a. Adaptive strategies of feeding and predator avoidance in the larvae of the
Neotropical butterfly Morpho peleides limpida (Lepidoptera: Morphidae). Journal of the New York
Entomological Society, 80: 66-82.
YOUNG, A. M., 1972b. The ecology and ethology of the tropical Nymphaline butterfly Victorina epaphus, 1.
Life cycle and ndtural history. Journal of the Lepidopterists’ Society, 26: 155-170.
YOUNG, A. M., 1973a. Notes on the biology of Phyciodes (Eresia) eutropia (Lepidoptera: Nymphalidae) in a
Costa Rican mountain forest. Journal of the N e w York Entomological Society, 81: 87-100.
YOUNG, A. M., 1973b. The life cycle of Dircenna relata (Ithomiidae) in Costa Rica. Journal of the Lepidopterists’
Sociey, 27: 258-267.
YOUNG, A. M., 1974a. Notes on the biology of Pteronymia notilla (Ithomiidae) in a Costa Rican mountain
forest. Journal of the Lepidopterists’ Society, 28: 257-268.
YOUNG, A. M., 1974b. On the biology of Hamadryas februa (Lepidoptera: Nymphalidae) in Guanacaste,
Costa Rica. Zeitschrqt f u r Angewandte Entomologie, 76: 380-393.
YOUNG, A. M., 1974c. Notes on the natural history of a rare Adelpha butterfly (Lepidoptera: Nymphalidae)
in Costa Rican high country. Journal of the New York Entomological Society, 82: 235-244.
YOUNG, A. M., 1975. Observations on the life cycle of Heliconius hecale zuleika (Hewitson) in Costa Rica. Pan-
Pa@ Entomologist, 51: 76-85.
YOUNG, A. M., 1976. Notes on the life cycle of the butterfly Hypanartia kefersteini (Nymphalidae:
Nymphalinae: Nymphalini) in Costa Rica. Brenesia, 9: 61-69.
YOUNG, A. M., 1977. Notes on the defoliation of Coconut Palm (COCOS nucifera) by the butterfly Opsiphanes
quiteria quirinus in north eastern Costa Rica. Deutsche Entomologische zeitschrift, 24: 353--365.
YOUNG, A. M., 1978a. Notes on the biology of the butterfly Hypoleria cassotis (Bates) (Nymphalidae) in
northeastern Costa Rica. Brenesia, 14/15: 97-108.
YOUNG, A. M., 1978b. The biology of the butterfly Aeria eurimedea agna (Nymphalidae: Ithomiinae: Oleriini)
in Costa Rica. Journal of the Kansas Entomological Society, 51: 1-10.
YOUNG, A. M., 1979. Weather and the regulation of Hypothyis euclea (Nymphalidae): populations in
northeastern Costa Rica. Journal of the Lepidopterists’ Society, 33: 68-69.
YOUNG, A. M., 1982. Notes on the natural history of Morpho granadensis pobbaptus (Butler) (Lepidoptera:
Nymphalidae: Morphinae), and its relation to that of Morpho peleides limpida Butler. Journal of the New York
Entomological Society, 90: 35-54.
YOUNG. A. M., 1984. Natural history notes for Taygetis andromeda (Cramer) (Satyridae) in eastern Costa
Rica. Journal of the Lepidopterists’ Society, 38: 102-1 13.
YOUNG. A. M., 1986. Natural history notes on Brassolis isthmia Bates (Lepidoptera: Nymphalidae:
Brassolinae) in northeastern Costa Rica. Journal of Research on the Lepidoptera, 24: 385-392.
YOUNG, A. M. & MUYSHONDT, A,, 1972a. Biology of Morpho pohphemus (Lepidoptera: Morphidae) in El
Salvador. Journal o f t h e New York Entomological Society, 80: 18-42.
YOUNG, A. M . & MUYSHONDT, A,, 1972b. Geographical and ecological expansion in tropical butterflies
of the genus Morpho in evolutionary time. Reuista de Biologia Tropical, 20: 231-263.
YOUNG, A. M. & MUYSHONDT, A,, 1973a. Ecological studies of the butterfly Victorina stelenes
(Lepidoptera: Nymphalidae) in Costa Rica and El Salvador. Studies on the Neotropical Fauna, 8: 155-176.
YOUNG, A. M. & MUYSHONDT, A,, 1973b. Notes on the biology of Morpho peleides in central America.
Caribbean Journal of Science, 13: 1-49.
YOUNG, A. M. & MUYSHONDT, A,, 1975. Studies on the natural history of Central American butterflies
in the family cluster Satyridae-Brassolidae-Morphidae (Lepidoptera: Nymphaloidea). 111. Opsiphanes
tamarindi and Opsiphanes cassina in Costa Rica and El Salvador. Studies on the Neotropical Fauna, 10: 19-56.
YOUNG, A. M. & MUYSHONDT, A,, 1985. Notes on Caligo memnon Felder and Caligo atreus Kollar
(Lepidoptera: Nymphalidae: Brassolinae) in Costa Rica and El Salvador. Journal of Research on the
Lepidoptera, 24: 154-1 75.
YUKAWA, J., 1984. Geographical ecology of the butterfly fauna of the Krakatau Islands. Indonesia. Buttegies
and Moths, Kyoto, 35: 47-74.