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Updating The Butterflies of the Malay Peninsula

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 Note
John Eliot died on 11 April 2003. At the time of his death, he
was working on an update of the fourth edition (1992) of Corbet
& Pendlebury's 'The Butterflies of the Malay Peninsula'. Both
this, and the third (1978) edition had been prepared for
publication by him.

Before his death he requested that funds be set aside from his
estate to facilitate publication, and to ensure that
every remaining copy in stock of the L992 edition should be
accompanied by a copy of this update.

The Editorial Committee of the Malaysian Nature Society decided

that the best way to reach those who had already purchased a
copy of the fourth edition was to publish the update as part of the
Malayan Nature Journal.

By chance it so happened that this issue also contained Pertz,

DeVries and Kirton's paper on the early stages of Xanthotaenia
busiris Westwood, which was clearly of relevance to Eliot's final
work. For this reason the paper has been bound into the Eliot
offprint.
Malayan Nature Journal 2006,59(f), I- 49

Updating The Butterflies of the Malay Peninsula

J.N. ELIOTI

Abstract: A summary is given of new information on the classification, distribution, early stages and
food plants of Peninsular Malaysian butterflies that has become available since publication of the
4th edition of the standard work, The Butterflies of the Malay Peninsula,in 1992.Apostscript gives
some personal reminiscences by the late author about his life, and his work on butterflies. The
following new formal taxonomic changes are introduced: Pieris malayica Martin, 1909 (Pieridae), is
a subjective synonym of Papilio canidia Linnaeus, 1768 (syn. n.). The subspecies taxon lxias pyrene
alticola Pendlebury (Pieridae) is considered to represent a distinct species, Ixias alticola Pendlebury
1933 (stat. n.). The genus Lebadea Felder (Nymphalidae: Limenitidinae) is moved from tribe Parthenini
to tribe Limenitidini. The monotypic genus Bhagadatta Moore (Nymphalidae) is re-assigned to the
Limenitidinae: Parthenini. The gents NeomyrinaDistant(Lycaenidae: Lycaeninae: Theclini) is removed
from the subtribe Loxurina to the Arhopalina. The genera Ancema Eliot and Pseudotajuria Eliot
(Lycaenidae: Lycaeninae: Theclini) are transferred from subtribe Remelanina to subtribe lolaina.

Key words: Butterflies, Malaysia, taxonomy, classification, distribution, new records, life histories.

INTRODUCTION
Since the 4th edition of The Butterflies of the Malay Peninsula by Corbet &
Pendlebury (revised by Eliot and hereafter referred to as C&P4) was published in
1992, a considerable body of new knowledge has been acquired. In particulal a
new higher classification of the Nymphalidae has been proposed by Harvey ( 1991),
and this has since been modified, notably by Ackery et al. (1999) and Wahlberg
et al. (2003). Mention should also be made of discussions in Freitas & Brown
(2004) and Wahlberg et al. (2005), that do not, however, replace Wahlberg et al.
(2003). In addition, in two superb volumes, The Life Histories of Asian Buttedlies
by Igarashi & Fukuda (1997,2000), more than 600 species have been described
and illustrated, including a greatmany from the Malay Peninsula whose life histories
were omitted, or noted as unknown in C&P4.
It seems appropriate to incorporate this new knowledge in a review, take the
opportunity to coffect some mistakes in C&P4, and to list all records made since
1992. Most of these recent records have already been published (mainly byArshad

I Formerly: Upcott House, Bishop's Hull, Taunton, Somerset TA4 lAQ, England. John Eliot died
on l lth April 2003, shortly after completing the first draft of this paper. This version of the paper
has been edited, enlarged and prepared for publication by Henry Barlow, Richard Eliot, Laurence G.
Kirton and R.I. Vane-Wright.
Correspondence: H.S. Barlow, PO Box 10139, 50704 Kuala Lumpur <hsbar@pcjaring.my>
et al. 1996, 1999,2000,2001,2002), but in publications such as the Malaysian
Naturalisr, ofwhich readers outside Malaysia may be unaware or have difficulty in
acquiring. The Addenda and Corrigenda to Corbet & Pendelb.rry 0992), although
issued simultaneously, were not widely publicised, and the opportunity is also taken
to repeat them here (with some further additions), as an Appendix.
In the entries that follow the page numbers cited are those of C&P4.

NOMENCLATURE
In the most recent fourth edition of the International Code of Zoological
Nomenclature (ICZN, 1999) it has been formally prescribed that the suffix to
denote subtribes is -ina. In previous Codes little mention was made of subtribes,
and as a result there was no stability in the use of suffixes, Americans often using
-iti, which was also used in C&P4, whilst Europeans more often used -ina. Now,
throughoutC&P4, subtribes ending in -iti must have their endings changed to -ina.

CLASSIFICATION OF THE NYMPHALIDAE

Harvey ( I 99 1 ) divided the Nymphalidae into the following 1 3 subfamilies: Danainae,
Ithomiinae, Tellervinae,Calinaginae, Morphinae, Brassolinae, Satyrinae, Heliconiinae,
Nymphalinae, Limenitidinae,Apaturinae, Charaxinae, Libytheinae. With the exception
of Calinaginae, Ithomiinae, Tellervinae and Brassolinae, which are not repr6sented
in Malaysia, and the Limenitidinae (see below), all these subfamilies are applicable
to the similarly-named subfamilies or tribes in C&P4. In passing, it should be noted
that Ackery et al. (1999) downgraded the Ithomiinae and Tellervinae as tribes
within the Danainae, and the Brassolinae likewise as a tribe within the Morphinae,
thus reducing Harvey's system of 13 subfamilies to just 10. With a further set of
changes introduced by Wahlberg et al. (2003), in which the former Limenitidinae
are divided into three subfamilies, the number of subfamilies recognised is set to
rise again,to 12.
In the case of the Satyrinae, Harvey (1991; see also Ackery et al., 1999)
simply downgraded Miller's subfamilies to tribes. The genus Xanthotaenia,in which
the discal cell is closed by a non-tubular vein, occupies an anomalous position,
belonging to none of Miller's (1968) groups. It appears to form a link between the
Satyrinae and Morphinae (tribe Amathusiini - in which it was included by Harvey,
1991), and probably deserves to be placed in a tribe by itself. Its position in C&P4,
following Ethope, is inappropriate and it would be better placed at the end of the
Satyrinae and immediately before Morphinae. The food plant is Hedychium
coccineum (Zingiberaceae), not the palm Calamus as stated in C&P4; according
to Robinsonet al. (2001), it has also been recorded from Zingiber (Zingiberaceae).
In the Limenitidinae, Harvey included 4 tribes: Limenitidini (corresponding
broadly with "Limenitiditi" and "Adoliaditi" in C&P4), Coloburini (wholly
Neotropical), Biblidini (holo-tropical), and Cyrestini. On the basis of marked
differences in the early stages, Eliot (in Pinratana & Eliot, 1996) proposed placing
the last three in a separate subfamily Biblidinae, in which the eggs are longitudinally
ridged and the larvae bear mid-dorsal scoli (usually only apartialrow), whereas in
Limenitidini the eggs are hexagonally-pitted with spines at the corners of the pits,
and the larvae never have mid-dorsal scoli. Since then, several authors have
recognised Colobura to belong to a sub-group of the Nymphalinae, now included
under the tribal name Coeini (Wahlberg et a1.,2003). Wahlberg et al. (op. cit.),
largely on molecular evidence, propose to promote the Cyrestini as well as the
Biblidini to subfamily status, separate from the Limenitidinae sensu stricto. The
analysis of morphological characters (early stages and adults) by Freitas & Brown
(2004) independently confirms the status ofBiblidini.
The subfamilyApaturinae is now widely recognised, and this includes the five
genera from Rohana to Hestinalis, as listed in C&P4 under Nymphalinae on p. 409.
In Pinratana & Eliot (1996),I included the Pseudergolini (formerly part of the
Limenitidinae) as a second tribe within the Apaturinae. Here, however, I follow
Wahlberg et al. (2003) in placing Pseudergolis and its relatives as a tribe of the
Cyrestinae.
The Heliconiinae of Harvey comprised two tribes: Acraeini, which included
Cethosia, and Heliconiini (including Argynnina). He listed as "subtribe uncertain"
the genera Phalanta to Tbrinos (pp. 151-156 in C&P4). He apparently overlooked
that this group had already been designated by Fruhstorfer (1912,in Seitz) as
"Issorididi". However, following the earlier error of Reuter (1896), Fruhstorfer
misused the type-genus Issoria (actually a genus of fritillaries: type species Papilio
lathonia L., the Queen of Spain Fritillary) for the genus now known as Vagrans .In
these circumstances a replacement name for Issorididi was needed, and Eliot (in
Pinratana & Eliot, 1996) substituted Vagrantiti (recte Vagrantina as a subtribe,
Vagrantini as a tribe). However, there is some doubt as to whether this is valid, as
the Commission was not approached as it should have been in the case of a
misidentified type genus. The earlier tribal name Phalantini Teshirogi ( 1990), which
might have been available for this grouping, is invalid, as its introduction was not
accompanied by a diagnosis (G. Lamas and R.I. Vane-Wright, pers. comm.).
Recently a well-argued paper by Penz & Peggie (2003) has been written, of
which Carla Penz kindly sent me an advance copy. The main difference from
Harvey's classification is that Cethosia is removed from the Acraeini and placed in
Heliconiini, thus restoring it to the position in which it was placed by Eliot (in Corbet
& Pendlebury 1978). Penz & Peggie simply subdivide the Heliconiinae into four
tribes, the Acraeini (including the African Pardopsis), the Heliconiini (including
Cetho s ia), Vagrantini and Argynnini.
The main results of all these changes can be summarised by altering the
dendrogram on p. l0l of C&P4. The branch leading to Charaxinae, Acraeinae and
Nymphalinae should be re-drawn to correspond with the following topology:

((Charaxinae * Apaturinae) ((Biblidinae + Cyrestinae * Limenitidinae) +

(Nymphalinae + Heliconiinae)))

In more detail, a preferable order of treatment for C&P4 from pp. 146-167
would be as follows. This order, broadly in agreement with Harvey as amended
above, should be adopted to revise the Check List in C8.P4.

S ugpar\arlv LtvtpNInomle

TribeLimenitidini- Moduza, Lebadea, Athyma, Pandita, Sumalia

TribeNeptini- Neptis, Phaedyma, Lasippa, Pantoporia
Tribe Parthenini Parthenos
TribeAdoliadini- - Tanaecia, Euthalia, Dophla, Bassarono, Lexias

Susrat\atrv BtsI-loNaE

Tribe Biblidini Ariadne, Laringa

-
SuspaNatrv Cvnrsrnqns

Tribe Cyrestini Cyrestis, Chersonesia

-
Tribe Pseudergolini Amnosia, Stibochiona, Dichorragia
-
Su gpaNarlv Nvupual-rN,q.E

TribeNymphalini- Vanessa, Kaniska, Symbrenthia

TribeKallimini- Rhinopalpa, Yoma, Hypolimnas, Doleschallia, Kallima,
Junonia

S uspaNatl-Y HpltcoNmlas

Tribe Acraeini Acraea

-
Tribe Heliconiini- Cethosia
Tribe Vagrantini Phalanto, Cupha, Vagrans, Vindula, Algia (: Paduca),
- Cirrhochroa, Vindula, Terinos
TribeArgynnini Argyreus
-
Finally, I think it is unlikely that the last word has yet been said on nymphalid
classification. But it can be said with certainty that the alrangement set out above
is much preferable to that used in C&P4.

NEW RECORDS AND IMPROVEMENTS TO THE TEXT

p.2 Recent cladistic analyses (de Jong et al.,1996; Wahlberg et a1.,2005) have
shown that the Papilionidae and Pieridae do not arise from a single stem
(see also Kristensen, 1976). The Riodinidae are sister to the Lycaenidae
and, thus, are not at the same taxonomic level as the subfamilies of the
Lycaenidae. For this reason they are now often regarded as forming a family
on their own. The dendrogram should therefore be re-drawn to correspond
with the topology (Papilionidae + (Pieridae + (Nymphalidae +
(Lycaenidae+Riodinidae)))). Other parts of the text in C&P4 affected by
this change are on pages 203 and 410.
p. 58 Troides.It is considered that Trogonoptera and Ornithoptera should be
re-instated as full genera. It is interesting that the structural colour scales of
both are of different types (Tilley & Eliot, 2002).

p. 69 Papilio prexospes. The male lacks the androconial scales present on the
forewing in the helenus group.

p.73 Tribe Teinopalpini, penultimate sentence, add: but there is a dense fringe of
short hairs from vein la to the dorsum on the underside of the hindwing in
Meandrusa, as in Leptocircini.

p. 87 Pieris canidia. Careful examination of Singapore examples with the nominate

subspecies flying in Hong Kong reveals no constant differences. It seems
certain that there has not yet been sufficient time for the evolution of
significant differences sufficient to justify a separate name for the Singapore
population. I therefore estabhsh Pieris malayica Martin, 1909, as syn. n. of
Papilio canidia Linnaeus, 1768.In future the Singapore population should
be designated Pieris canidia canidia (Linnaeus, 1768) (: malayicaMartin,
1e0e).

p. 88 Phrissura. The Malaysian taxon cynis is no longer considered to be

conspecific with aegis (Yata, 1981; Eliot & Kirton, 2000). Change references
to Phrissura aegis cynis to Phrissura cynis cynis and Phrissura aegis
pryeri to Phrissura cynis pryeri, and amend paragraph on the genus as
follows:

The genus comprises two species, P cynis and P aegis. The male of P. aegis,
which occurs in the Philippines (including Palawan) and Sulawesi, has a
ventral abdominal hair tuft anterior to the saccus. In the Neomalayan species,
P cynis, this tuft is absent; the species was formerly separated as Udaiana
Distant.

p.92 lxias. The provisional placement in C&P4 of three taxa under Ixias pyrene
Linnaeus, 1764, requires re-examination. The male of the nominate
subspecies, flying from N.E. India to S.E. China (type locality Canton: Honey
& Scoble, 2001), has a clear yellow ground colour and a bright orange
subapical band on the black apex of the forewing. As the range extends
southwards into Indo-China the ground colour becomes increasingly creamy-
white, outwardly tinged with lemon yellow, in a cline beginning in South Burma,
Thailand and Laos, the orange subapical band becoming wider and paler,
and tending to become clouded with black scales. The most extreme form of
this cline is found in Peninsular Thailand and Kedawi, and this population
was named by Fruhstorfer salangana. The type locality, Salang, often
expanded to Jang Salang which, in the days of sail was a port of call for
merchantmen trading to the Far East and known to British sailors as Junk
 Ceylon, is now better known as the popular tourist resort of Phuket. Anumber
of names have been given to the intermediate populations, of which the
earliest is verna H. Druce, 1824 (type locality Siam). In the past this was
treated by some authors, including Corbet (1937), as a good species distinct
from pyrene.In view of the gradual change from pyrene to verna (as can
be seen by good coloured figures in Osada et al.,1999), this view cannot be
maintained. The difference between Thai and Kedawi populations is
insufficient to justify the use of salangqna, and the Kedawi population should
continue to be known as I. pyrene verno, in the interests of nomenclatural
stability.

Females of the more northerly population are polymorphic. Typical females

from N.E. India and S. China have a white ground colour and white subapical
band, and in the wet season form may be so infuscated that they appear
almost black. Other females may have a white or yellow ground colour with
the subapical band of the same colour or orange. As the population changes
from pyrene to vernq, the female becomes monomorphic with a pale yellow
ground colour and orange subapical band.

It has hitherto been supposed that the Malaysian taxon I. pyrene birdi Distant
had a monomorphic all-yellow female (C&P4, Plate 9: fig. 20), but recently
Chong has found a colony of birdi in Trengganu (Fig. l, S and Fig. 2, t)
where a hitherto unknown female morph (Fig. 2) with a yellow ground colour
and orange subapical band occurs together with the all-yellow morph, but
more commonly.

The third Malaysian taxon, I. pyrene alticola, with a pure white ground
colour in both sexes, was originally described by Pendlebury as lxias
ludekingii alticola. The Sumatran ludekingii Vollenhoeven is generally
treated by all authors as a good species in its own right, and I consider that it
cannot be conspecific with alticola. The male has a creamy white ground
colour and a broad pale yellow subapical band dusted with black scales, and
bears some resemblance to the male of verna, of which it may be the
Sumatran descendant. Throughout the lxias pyrene complex there are no
males with a pure white ground colour. The separation of alticola from
birdi is so narrow both in altitude and vegetation that it is hardly conceivable
that no hybrids would ever have been found should the two be conspecific. I
therefore raise the available name olticolo to species status, as the distinct
species lxias alticola Pendlebury, 1933, stat. n.

p.95 Dercas. The Sundanian populations should be separated as a distinct species,

D. gobrias Hewitson, 1864, separate from the continental D. verhuelli.
The larval food plant is Dalbergia (Fabaceae) (Schulze & Fiedler, 1997).
p. 98 Eurema. Yata (1989) has shown that Eurema should be divided into two
subgenera: Eurema with a predominantly New World distribution, and Tbrias
Swainson with an Old World distribution. All the Malaysian species, with the
exception of E. (E.) brigitta,belone in subgenus Terias.

p.I24 Lethe delila is now considered to be a subspecies of L. chandica.

p. 133 Ragadia. Until recently it was thought that there was only a single black and
white species in Malaysia. This was the submontane insect designated
R. crisilda critolina Evans, 1923 and figured as such in C&P4 atPlate I7,
Figs. 8-10. This species had already been described and named from Dawna
Hills in Burma as R. critolaus by de Nicdville, 1893. It is now known that a
second lowland species flies in Kedawi (Arshad et ol., 1996). The main
difference between the two species lies in the pale submarginal band. In the
former (critolaus) it is white near the dorsum, becoming gradually infuscated
towards the apex. In the second, which is usually a little smaller and has a
more rounded forewing, this band is infuscated throughout and this species
has been universally accepted as the true crisilda Hewitson,1862. Evans, in
his brief description of critolina, included the phrase: upperside forewing
submarginal band may be obsolete. From this it is apparent that Evans had
confused the two species as one and it became necessary to decide to which
species critolina should belong. Evans did not speciff a type and Eliot could
find nothing suggestive of his type series in the collection of The Natural
History Museum (BMNH). Eliot & Kirton (2000), acting as first revisers,
therefore decided to restrict critolina to R. crisilda, in the interests of
maintaining maximum nomenclatural stability. In consequence the species
treated as R. crisilda critolina in C&P4 must in future be known as R.
critolaus de Nicdville, and the legend of Plate 17 (figs. 8-10) should be
altered accordingly, while the species that flies in Kedawi should instead be
known as R. crisilda critolina (Figs. 3, 4).
Ragadiini. A family group name based on Ragadia was first introduced by
Herrich-Schdffer (1864), not Miller (1968).

Key for the separation of the species of Raceora:

| (4) Upperside with a prominent white band, running from near the apex of the forewing
to the inner margin of the hindwing.
2 (3) Upperside forewing submarginal band uniformly blackish, contrasting only weakly
with the black ground colour. Usually smaller (d forewing l8 mm) and wings more
rounded. R. crisilda
3 Upperside forewing submarginalbandwhite attomus, becoming increasinglyinfuscated
towards the apex; usually larger (d forewing22.5 mm). R. critolaus
4 Upperside without a white band, although the pale transverse stripes on the underside
are visible by transparency. R. makuta
 In the final parugraph of the text (p. 133) substitute R. critolaus deNic6ville
1893 for R. crisilda critolina Evans, and delete the penultimate sentence
and replace with the following:

A further species, R. crisildo critolina Evans, 1923, occurs sympatrically

with R. critolaas throughout most of its range but at lower levels. In the
Peninsula it has only been found in Kedawi (for differences, see Key). Its
larva in Vietnam is green with oblique white stripes and feeds on Selaginella
delicatula (Igarashi & Fukuda, 2000).

p. 138 Faunis. The key to species should be amended as follows:

2 (3) 6f forewing termen rounded; underside hindwing with dark median band clearly
traceable from costa to veinlb. t underside hindwing with median band regular and
evenly curved, not bent outwards in space 6. F canens
3 6 forewing termen straight; underside hindwing dark median band obscured by dark
tornal area below vein 3. t underside hindwing median band irregular, serrate in places
on either side and sharply bent outwards in space 6. F. kirata

p. 145 Discophora. The key to the species should be amended as follows:

3 (4) 6f upperside forewing with a few obscure whitish subapical and submarginal spots.
t upperside forewing termen very nalrowly chrome-edged; hindwing plain brown
with termen not, or only very narrowly near apex, chrome-edged. D. timora
4 d upp..side forewing with blue intemrpted subapical band and blue submarginal
a pale
spots in spaces 1b to 5. f
upperside termen prominently chrome-edged and hindwing
with obscure dusky orange spots, submarginal in spaces 2-5 andpost-discal in space
6. D. necho

p.146 Acraeinae is reduced to a tribe of Heliconiinae.

Insert between the first two sentences: The male deposits a seal, termed a
sphragis, over the ostium bursae of the female after mating, so that a second
coupling is impossible.

Insert at the end of the second paragraph: "and Urticaceae" and follow by a
new paragraph:

Acraea violae (Fabricius,1775) (Fig. 5) is a butterfly that has expanded its

range from Thailand as far south as the Kuala Lumpur area (L.G. Kirton,
pers. comm.) on the mainland and to the Langkawi Islands (Arshad et al.,
1996), and will probably continue its spread further southwards. It is easily
recognised by its chrome colour and sparsely black-spotted wings. The larva
feeds on species of Passifloraceae.
 Lastparagraph: delete the opening phrase and add atthe end: The gregarious
larva of A. issoria feed on Boehmeria and other species of Urticaceae.

p. 146-148 See above for the proposed new affangement of subfamilies and tribes
previously included in Nymphalinae. To summarise, the new subfamilies and
tribes in Malaysia are as follows:

Nymphalinae, with tribes Nymphalini (Vanessa, Kaniska and Symbrenthia)

and Kallimini (Junonia and Rhinopalpa to Kallima)

Heliconiinae, with tribes Acraeini (Acraeo), Heliconiini (Cethosia),

vagrantini (cupha to krinos) and Argynnini (only Arg,treus)
Limenitidinae, with tribes Limenitidini, Neptini, Parthenini andAdoliadini
Biblidinae, with tribe Biblidini (Ariadne and Laringa)
Cyrestinae, with tribes Cyrestini and Pseudergolini

Apaturin ae (Rohana to Hestinalis)

The keys on pages !47 and 148 are still usable for identification to tribal
level.

p. 150 Key to subtribes of "Heliconiiti". Upgrade to Heliconiini andArgynnini. At

end of line l, add: except in wndula with long head-horns.

p.152 Paduca must be replaced by the earlier name Algia Herrich-Schiiffq 1864,
of which it becomes a synonym.

p. l54As pointed out above, Vindula and Tbrinos should be removed to Vagrantini.

p. 157 Tribe Nymphalini. As already pointed out, Harvey raises this taxon to subfamily
Nymphalinae, which he subdivides into two tribes: Nymphalini including
Vanessa, Kaniska and Symbrenthia,with hairy eyes and palpi with projecting
scales; and Kallimini with smooth eyes and palpi. A further" significant
difference is in the larval food plants: always Acanthaceae in Kallimini, and
usually Urticaceae in Nymphalini; but Asteraceae in the case of the Painted
Lady, Vcardui.In addition, the distribution of the filiform setae on mature
larvae differs.

Cethosia. The key to species should be amended as follows, to accommodate

C. cyane (Figs. 6-8), newly recorded from the Langkawi Islands (Arshad
et al., 2000).

3 (4) Upperside forewing without a series of white subapical spots; underside hindwing
without a white submarginal band. C. hypsea
 4 (5) Upperside forewing with subapical white spots; underside hindwing with a narrow
white submarginal band outwardly defined by short black stripes.
5 (6) Upperside j t rich orange red and underside hindwing submarginal white band
continuous. C. penthesilea
6 Upperside d much paler, i becoming whitish. Underside hindwing submarginal
white band much intemrpted. C. cyane

p.166 Doleschallia bisoltide. In the Langkawi Islands two currently recognised

subspecies, D. b. pratipa C & R Felder, 1860, and D. b. siamensis
Fruhstorfer,l9l2, occur together, and so far no intergrades have been found
(Arshad et al., 2000). ln siamensis the female more or less resembles the
male, with a detached, yellowish, subapical band on the black apex of the
forewing. Only time will tell if these two overlapping taxa have evolved into
distinct species.

In addition, a new form has been found in Selangor, flying alongside pratipa,
and also in Singapore, by Yong & Rosli Hashim (2002). This form diflers
frompratipa inhaving five small white spots on the black apex of the forewing,
lacking the large white spots on the underside that are usually present in
pratipa, and most significantly in the larva, which these authors have bred
on Asystasia intrusa (Acanthaceae), which exactly resembles the larva of
the Papuan and Australian D. b. australis C & R Felder, with red spots
(absent in pratipa) at the base of the lateral scoli. However, australis is an
inappropriate name for this newly discovered taxon, since the adult phenotype
differs so strongly. The origin of the new taxon is unknown. Rather similar
forms occur in the Philippines, but a Philippine origin seems most unlikely. It
is tentatively suggested that it might be a variety of nominate D. bisaltide
from Java (type locality) which may have reached Malaysia "under its own
steam" by u process of island-hopping. To ensure that its presence is not
overlooked it is included in the Check List as D. bisaltide ?bisaltide var.

p.176 Athyma. The Athymo ranga group has recently been shown to comprise
three species, and some changes in their status and nomenclature have been
made (Tsukada & Kaneko, 1982; Eliot & Kirton,2000). Thus, the species
given as Athyma rango malaya in C&P4 should be known as A. matanga
while the species given as A. abiasa clerica should be known as A. clerica
clerica. Couplets 10 and l1 of the key should be modified as follows:

l0 (l0B) Abdomen white spotted; upperside hindwing upper spot of discal white band
detached. A. matanga
l0B Abdomen not white spotted; upperside hindwing discal white band undivided.
I I (l lB) Upperside forewing small white spot in space 3 placed against the inner edge of
the large white spot in space 2. A. clerica
l lB Upperside forewing spot in space 3 larger and placed centrally above the spot in
space 2. Java, Sumatra, Linggi Islands. A. abiasa

l0
p. 178 Last paragraph referring to Athyma rango group under Athyma nefte
subrata, change A. ranga malaya to A. matanga and A. abiasa clerica to
A. clerica clerica; delete last line and add: Athyma abiasa, a species quite
similar to A. clerica, flies in Sumatra and the Linggi Islands as the race
lingana Fruhstorfer, and may yet occur in the Peninsula. Moore erected the
genus Kironga for A. abiasa and the continental species, A. ranga.

p. 180 Tribe Parthenini. The genus Lebadea should be removed to Limenitidini on

the basis of its early stages, which broadly resemble those of Athyma. ln
tribe Parthenini, which apart from Parthenos now only includes the sino-
Burmese genus BhogadottaMoore, the highly irregular pupae typical of the
Limenitidini are replaced by smooth pupae. The food plant of Bhagadatta rs
Mammeo siamensis (Guttiferae).

p. 1 81 Subtribe "Adoliaditi" should Adoliadini. This tribe has usually

be raised to tribe
been called the Euthaliini Moore, 1895, in the literature, butAdoliadini Guen6e,
1865, has priority and is now accepted.

p. 184 Thnoecia clathrataviolaria. There is a possibility that the ? type of violoria

(Fig. 9), supposed to be from Singapore, may be a mislabelled Tanaecia
munda from Sumatra. (T. Yokochi, pers. comm.; the type is also figured
poorly in Flemin g,197 5;1983). If this is eventually shown to be the case, the
Peninsular Malaysian taxon may become known as P clathrata nicevillei
Distant, 1884, which is currently treated as a synonym of violaria.
Tanaecia coelebs. The Mal ayan race should now be known as Z heringi
regalis Tsukada. Yokochi (1999) has found that heringi Niepelt, 1935, and
coelebs Corbet, 1941, are the same taxon, described from the same sex and
same locality. An application by Yokochi (1999c) to conserve the name
coelebs in favour of heringi was rejected by the International Commission
on Zoological Nomenclature (2002) and, as such, the species should now be
known as Thnaecia heringi Niepelt , 1935. The female mentioned in C&P4
is likely to be a local variant of T. julii. A more probable female of the
species (Fig. 10) resembles munda tin its markings, but the hindwing is
uniformly brown up to the termen (Arshad et al-,1996).

p. 186-1 87 Euthaliakey. Euthalia agnis (Vollenhoeven) and E. tinna Fruhstorfer

have recently been shown to be two distinct species occurring sympatrically
(Yokochi & Matsuda, 1999), as originally proposed by Fruhstorfer (1906).
They occur in the Peninsula as the subspeci es Euthalia agnis hiyamoi
Yokochi (Figs. ll,12, d holotype; Fig. 13, ? paratype; Plate 27: ftg.31 ?),
and E. tinna paupera Fruhstorfer (Plate 27 , frg. 32,6) (the female of the
latter appears to be unknown). Couplets 18 and 19 of the key to Peninsular
Malaysian Euthalia should be revised as follows:

11
 l8 (le) Forewing postdiscal white spot in spaces 2 and 3 always present, the spot in
space 3 slightly basad of the spot in space 2, such that a line through their
centres does not meet the forewing dorsum perpendicularly.
r88 (18C) d forewing postdiscal white spots in spaces 24 ananged in an oblique, dead
straight line.t hindwing without a whitish postdiscal band. E. agnis
18C 6f forewing postdiscal white spots not in a dead straight line, the spots in
spaces 2 and 3 shifted slightly basad. t unknown from Peninsula (but
t of all
other ssp. with a whitish postdiscal band on the hindwing). E. tinna
t9 Forewing postdiscal white spots in spaces 2 and 3, if present, with their centres
forming a line that is perpendicular to the forewing dorsum. E. aconthea

The Malayan population of Euthalia eriphylae was recognised as a distinct

race but was unnamed. It has now been described and named E. eriphylae
elioti Yokochi, 2002 (t holotype also figured inC&P4,Plate 27: frg.27).lt
differs from Indo-Chinese races in having smaller discal spots on the
upperside forewing of the female.

The forewing cell spots of the Peninsula E. lubentina chersonesia

Fruhstorfer are at most very weakly reddened and may not be reddened at
all. In Langkawi the nominate subspecies with reddened cell spots occurs.
[See also note below re p. a08.]

A preferable key to the species ofthe Eutholia lubentina group, from couplet
25 onward, would run:

25 (28) d with fore tarsi reddened, upperside forewing with subapical white spots not in
line, the spot in space 6 shifted basad. t upperside forewing subapical spot in space
6 basadofspot in space 5.
26 (27) d upperside forewing cell spots not reddened, upperside hindwing red band not
dislocated. t upperside hindwing with broad white band. E. adonia
27 6( upperside forewing cell spots may or may not be reddened; upperside hindwing
red band slightly dislocated, with red spot in space 4 shifted slightly basad of spots
in spaces 5 and 3. ! upperside hindwing band red, with spot in 4 shifted slightly
basad. E.lubentina
28 j fore tarsi not reddened; forewing apical spots in spaces 4-6 inline and directed to
apex. ! forewing subapical spots out of line, but that in space 6 not basad of that in
space 5; hindwing with red band.
29 (30) 6f upperside forewing cell spots reddened. t forewing with white post-discal spots
beyond cell adjacent to end-cell bar, and spot at base ofspace 3 large.
E. malaccana
30 6f upperside forewing cell spots not reddened. ? forewing u'ith white post-discal
spots beyond cell well separated from end-cell bar and spot at base of space 3 small
or absent. E. whiteheadi

p. l9l Pseudergolini. First paragraph: there is only a single abdominal keel on the
pupa.

t2
p.zll Poritia key. The recent discovery of a new species, P. ibrahimi Eliot &
Kirton, 2000, necessitates a modification to line 12 inthe key, as follows:

12 Upperside forewing blue; hindwing cell mainly black.

l2B (12C) Upperside forewing with linear black streaks above cubitus and vein lb; small
species, forewing 12.5-14.0 mm. P pleurata
l2C Upperside forewing blue above cubitus and vein lb, without any black streaks;
larger species, forewing 17.0 mm. (Unique, female unknown.) P ibrahimi

p.213 Poritia. The final paragraph should be amended. Add the first sentence to
preceding paragraph, and then replace remainder of the paragraph as follows:

Of the remaining species the most frequently encountered is P. sumatrae,

which is easily recognised by the green colour and broad unspotted black
marginal and central black border filling the cell in the male, and the pale
purple female with the hindwing veins overlaid by reddish brown scales.

The two species which may be most difficult to separate are P. phama
rajota (Plate 32, figs. 13-15) and P. hewitsoni taleva. The males of both
species are greenish-blue, that of hewitsoni being of a greener shade, which
is accentuated when the wings are tilted. In addition, in hewitsoni there is
often a black mark in space lb of the forewing that is never found inphama.
The females are very similar, but the blue colour of hewitsoni is slightly
deeper and there is a narrow black streak in the cell above the cubitus. For
other differences see key. Two rare species in which the males are blue
without a green tinge even when the wings are tilted are P. pleurata and the
newly described P. ibrahimi Eliot & Kirton,2000 (Figs. 14, 15). The former
is usually the smallest species in the genus and on the forewing has a black
streak above the cubitus and a black streak in space 1b. These markings are
absent in the larger ibrahimi.

The very rure P. manilia is rather similar above to P hewitsoni, but the
nominate subspecies flying in Malayaproper is greener and on the hindwing
the discal cell is all black. Both sexes are most easily identified by the off-
white underside with pale cinnamon markings. The male of the subspecies
P. m.evansi is much bluer than subspecies manilia, but the wings become
green with an indigo tint when tilted.

The largest species, P promula elegans, is fairly easily recognised by size.

In both the green male and the pale blue female the broad black borders are
unspotted and do not enter the forewing discal cell.

p. 214 A new female morph of Simiskina phalena, inwhich the upperside is brown
and the underside resembles the typical female morph, has been named
f. nox Seki, 1991.

13
 S.proxima also has two new, apparently unnamed morphs, which on the
' upperside resemble the typical ? morph and f. abisarina of S. phaleno,but
, they are easily separated from the latter species by their very different
undersides.

p.228 Curetis. For Eliot, in press, read Eliot, 1990. The details are given in the
references.

p.245 Zizeeria. S.K. Khew (pers. comm.) has discovered in Singapore apopulation
of Z. maha serica (C. Felder, 1862) wet season form (Figs. 16, l7). This
taxon has also been found recently in Luzon. In both instances it is virtually
certain that its arrival is due to unplanned human agency, probably from
Hong Kong, which is its type locality. The food plant of the lawa is Oxalis
corniculata (Oxalidaceae) (Bascombe et a1.,"1999). Some authors place
maha in a separate genus, Pseudozizeeria Beuret, 1952. However, I think
this is an unnecessary splitting, and propose to retain it in Zizeeria in the
interests of stability of nomenclature.

p.275 Vane-Wright & Gaonkar (2006) have shown that the correct name for
Arhopala pseudocentaurus nakulo (Felder & Felder, 1860) should be
Arhopala centaurus nalaila (Felder & Felder, l g60).

p.279Arhopala denta. Toshforce captured two females, unfortunately

unaccompanied by a confirmatory male, on Tioman Island. These were
identified by Storey as A. denta (Arshad et ql., 20Ol). After careful
measurement of the tails I think this identification is correct, and so it should
be included in the checklist of C&P4 as a new record for the Malay peninsula.

p.292 Zinaspa. The taxon occurring in Malaya is not Z. todara karennia, in which
thedis tailed, but an unnamed taxon with a taillessdwhich also flies in
Sumatra, and was figured in Corbet & Pendlebury Q978: Plate 44,ftgs. l,
2). lt is not certain whether this taxon is a variant of todara or a distinct
species. The early stages, described later, suggest that Zinaspa should at
best be treated as a subgenus of Surendra.

p.298 Neomyrina. The discovery by Igarashi & Fukuda (2000) of the early stages
ofN. nivea, including its flattened onisciform lawa,confirms that this g.not
belongs tothe Arhopala section ofArhopalina, ofwhich it is a slightly abirrant
member. This position is also justified by the forewing venation, with vein 5
much closer to vein 6 than to vein 4.

p.307 Subtribe Iolaina. It has been found that the males of Tajuria sunia and,
T. inexpectata have a male-like five-segmented tarsus (L.G. Kirton, pers.
comm.). This may be due to crossing over in the sex chromosomes. The

t4
 sentence beginning in line 10 of the first paragraph under subtribe Iolaina
should read:

Male foretarsus generally fused to a single, stubby-tipped segment, except in

some species such as Pratapa deva, sukidion inores, Tajuria sunia and
T. inexpectata.

p. 31I Tajuria key. The description of T. sunia d is incorrect. Line 22 of the key
should read

22 d upperside forewing border expanding in an almost even curve from tomus to cell
t paler with a much
apex; ground colour of a slightly deeper blue than T. ister.
narrower black border on forewing. T. sunia

p. 313 Tajuria. The female of T. inexpectata is, as yet, unknown in the Peninsula.
The specimen identified by Eliot as a female of inexpectata from Malaya,
with broader and inwardly straighter black forewing borders than females
from Sumatra (Kirton & Kirton, 1983), is actually a worn male of I ister
(L.G. Kirton pers. comm., by dissection). The two sentences beginning line
10 following the reference to T. sunia plate and figure should read:

Above, the black borders are more inwardly curved in the male, and much
naffower in the female, in comparison to T. inexpectata. Both T. inexpectata
and T. sunia are very rare.

p.314 Eliotia. The genus name should revert to RachanaEliot,and the first sentence
should read:

The name Eliotia Hayashi was erected for this genus, and has a few weeks'
priority over Rachana Eliot. However, the former is preoccupied and has
been replaced by EliotianaKogak,lgg6,which can be considered a subgenus
of Rachana for a small group of Philippine species with different secondary
sexual characters (Eliot & Kirton, 2000).

p.320 Hypolycaena.Line 4 of the key should be modified to include an additional

species, as follows:

4 (5) Underside hindwing outer portion of vein 2 not orange; black tornal spot not edged
with silvery scales in space lb. Forewing 12-15 mm.
48 (4C) Underside hindwing apical region at most barely tinged orange; always with a
prominent black marginal spot in space 6. d upperside shining ultramarine blue
without any violet tinge. H. amabilis
4C Underside hindwing apical region orange tinged; black marginal spot in space 6
absent or ill-defin ed. 6 upperside violet. H. phemis

Last paragraph, first three sentences, replace with the following and retain
the last two sentences on H. merguia:

t5
 Three small species, H. amabilis, H. phemis and H. merguia,can be regarded
as typical ofthe genus, although in many works, including the second edition
of this book, this group has been placed in Chliaria.ln H. amabilis lisba
(Corbet), the male is shining ultramarine blue above, with a diffuse black
border on the forewing, and the female brown with a white tornal area on
the hindwing. Eliot & Kirton (2000) have pointed out that the rather similar
taxon, phemis H.H. Druce, usually treated as a Bornean subspecies of
H. amabilrs, occurs sympatrically with the latter at all elevations in the
Peninsula without intermediates and is, therefore, best treated as a separate
species, H. phemis (Plate 49, figs. 25, 27 6 , 26 ?; genitalia, fig. 3 l6). The
male differs from H. amabilis in being violet on the upperside; other
differences are given in the key above.

p.337 Choaspes. The taxa C. plateni and C. stigmota are not conspecific, both
being good species; nor are hemixanthus and furcatus which have totally
different early stages, as discovered by Igarashi & Fukuda (1997).A further
species that should be looked for in Malaysia is C. xanthopogon, with the
upperside of the male brown. Its nominate subspecies, occurring throughout
the Indo-Burmese area, has an underside similar to C. benjaminii and
C.furcatus, but the Sumatran subspecies xanthopogon coro Evans has
the orange tornal area yellower. Evans (1949) incorrectly and inexplicably
described cora as a subspecies of C. hemixanthals Rothschild.

p. 340 Tapena. The last sentence of the first paragraph and the second paragraph
should be replaced with the following:

A light and dark form occur in Neomalaya (Eliot & Kirton, 2000). In the dark
form (Plate 53, fig. 13 6 ), the ground colour is dark brown with the darker
markings barely contrasting, and with the white spots on the forewing very
small. The pale form is a lighter shade of brown, with the darker markings
more contrasting, and with the white spots sometimes larger. Additionally, a
spot may be present at the cell-end of the hindwing. This form bears some
resemblance to the more heavily spotted subspecies, miniscula, occrtrring
from Burma to Indochina.

The female of 7l thwaitesi bornea has additional white spots on the forewing
atthe cell-end, in spaces 2,3 and sometimes space lb, as well as awhite
spot at the cell end on the hindwing.

p.341 Coladenia. The Malayan race of C. agni listed as an unnamed subspecies

(see checklist of C&P4) has been named C. agni sundoe de Jong &
Treadaway, 1992.It also occurs in Sumatra (the type locality).

p.353 Koruthaialos. Add K. frena after K. sindu in reference to shining white

specialised scales.

16
p.354 Koruthaialos key. Amend as follows:

I (6) Forewing with a red to orange-yellow discal band.

2 (3) Palpi third segment long, thin and conspicuously protruding. Upperside forewing
discal band often not reaching the costa (forewing length 14.0-17.0 mm). K. rubecula
3 Palpi third segment short and just protruding. Upperside forewing discal band always
reaching the costa.
4 (5) Upperside forewing discal band orange to orange-red, and relatively narrow, its width
not greater than the distance between the subapical edge of the band and the forewing
apex. Smaller species (fi^/l 15.0-19.0 mm). K. sindu
5 Upperside forewing discal band paler, orange to orange-yellow, and wider, its width
usually greater than the distance between the subapical edge of the band and the
forewing apex. Larger species (fw120.011.0 mm). K. frena
6 Wings dark brown, with a plum purple suffusion and unmarked; underside forewing
dorsum broadly pale pinkish brown (fuI 16.5-18.0 mm). K. butleri

Koruthaialos sindu sindu. Add to the beginning ofthe second lastparagraph:

Koruthaialos frena Evans, 1949, occurs rather scarcely in the highlands of
the Peninsula, above 2,500 feet. The species is generally similar to K. sindu,
but differs in being larger and in having a broader forewing band (5-6 mm
wide) that is also paler and yellower. The Peninsular Malaysian population
(Figs. 18, 19) has, on average, a smaller wing length (forewing: male 20 mm,
female 21 mm) than the nominate Bornean race in which the forewing length
ranges from 2l-22 mm in the male and24-25 mm in the female (Maruyama,
1991), and probably represents a separate subspecies.

p.361 Suostus. Replace first to third sentence of the third paragraph with the
following (see also life history section):
Suastus gremius gremius (Fabricius) (Plate 54, frg.61 ?) appears to have
been confined in the past to the Butterworth area, Perlis and the Tioman and
Redang Islands, but has increased its range in recent years, perhaps by
human agency, through transportation of the many species of ornamental
palms that serve as its host plant (see life history section). The species is
now widely established in garden and open areas in the Peninsula. The
remaining species are rare in Malaya.

p.368 Salanoemia. The following should be inserted before the last sentence of
the text on Salanoemia:

A distinctive species with cream to yellowish markings, Salanoemia shigerui

Maruyama (2000), was recently described from southeast Johore.

The key should be amended as follows from the fourth line onwards:

4 Upperside hindwing with a yellowish to cream coloured discal area.

5 (8) Upperside forewing cell spots subequal.
6 (7) Upperside hindwing discal area yellow; underside hindwing yellow without a brown
marginalband. S. tavoyana

t7
 Upperside hindwing discal area mainly cream coloured; underside hindwing pale
yellow to cream coloured, with a wide, brown marginal band from tornus to termen.
S. shigerui
Upperside forewing upper cell spot absent or very small. S. similis

p.371 Zela. The genus has been revised by Kirton & Eliot (2004), who have
subdivided the genus into three subgenera and described three new species
(2. steineri, Z. mars and Z. storeyi), all of which occur in the Peninsula.
The section on Zela in C&P4 should therefore be revised as follows:

Genus Zela de Nicdville

The genus is chiefly distinguished by the very short hindwing cell. All the
species also have yellow or orange tornal cilia on the hindwing and iridescent
hair scales on the thorax, which are developed to varying degrees. In addition,
they share a number of characteristics in the male genitalia.

Kirton & Eliot (2004) have recently revised the genus and consider it to
comprise three subgenera, that is, subgenus Zela de Nic6ville, 1895, the
monotypic subgenus Matapoides, H.H. Druce, 19T2, and subgenus Zampa
de Nic6ville, 1895. In subgenus Zela, there are hyaline spots in the cell,
space 2 and space 3 which are usually absent in the other subgenera. The
males have a straight forewing dorsum and long hairs on the hindwing disc,
which do not form the dense patch found in the other subgenera. The subgenus
comprises two rather distinct groups. In the onorq group, comprisingZ. onora
(Butler) (Yellow Palmer) (Plate 57, frg. 15 6) and Z. excellens (Staudinger)
(Plate 57, figs. 16 6 ,17 ?),the hindwing has a yellow bar and male secondary
sexual characteristics are absent (2. onara) or poorly developed
(2. excelleres), comprising only a weak discal stigma in space 2 on the
forewing upperside. In the zeus group, comprising Z. zeus de Nicdville
(Redeye Palmer) (Plate 57,fig. 18 d) and two newly described species,
Z. steineri Kirton & Eliot (Fig. 20) and Z. mars Kirton & Eliot (Figs. 2t , 22),
the hindwing is brown without a yellow bar and the forewing upperside has
a black stigma in spaces lb and 2, which is broken in mid space lb and at
vein 2. Zela zeus occurs as the nominate race in the central and southern
parts of the Peninsula, and as the race optima in Kedawi and Langkawi.
The latter differs from subspecies zeus in having diffrrse yellowish bars at
the cell-ends on the upperside of both wings and a brownish orange patch
above the dorsum on the forewing underside.

Subgenus Zampa is characterised primarily by the existence of a dorsal

extension of the tegumen in the male genitalia, which has been termed the
tegumenal apophysis (Kirton & Eliot,2004). The shape of this structure is
diagnostic for the different species. Males have a bowed forewing dorsum
and a dense hair patch on the upperside of the hindwing, which corresponds
with a polished area on the underside of the forewing dorsum. Zela cowani

t8
Evans, Z. zenon (de Nic6ville) (Tufted Redeye Palmer) (Fig. 23 6) and the
recently described Z. storeyi Kirton & Eliot (Fig. Z4 3; Plate 57, frg.19 ?)
form the zenon Broup, in which the males lack a discal stigma on the upperside
of the forewing. The zero group, comprising Z. elioti Evans (Plate 57,
frg. 21 8 ) and Z. zero Evans (Plate 57 , fig. 20 6), have a discal stigma, and
also have enlarged tegumenal apophyses as well as an additional discal patch
of specialised scales within the polished dorsum on the underside of the
forewing. These scales are easily detached. Zela elioti exhibits geographic
variation in the width and colour of the forewing brand in the male.

Subgenus Matapoides is represented by a single striking species,

Z. smoragdino (H.H. Druce) (Tufted Redeye). It shares in common with
subgenus Zela the absence of a tegumenal apophysis, but has all the other
male characteristics of subgents Zampa more highly developed. The male
valva also has a number of unique characteristics not found in the other
subgenera.

The genus is distributed fromAssam to Sundaland, with a centre of diversity

in Peninsular Malaysia and Borneo. All the species are very rare.

Key for the separation of the species of Zeyli

I (10) Forewing usually with one or more hyaline spots in the cell and also in spaces 2 and
3 (may be faint in some females). Male upperside hindwing with long hairs clothing
the disc, but not forming a dense, localised patch. Slbgenus Zela
2 (5) Hindwing with a large yellow patch or band. (onara group)
3 (4) Underside hindwing yellow band from costa to termen. Z. onara
4 Underside hindwing yellow band from costa to vein lb, continuing below vein lb
only as a dark-dusted, yellow suffusion. Z. excellens
5 Hindwing without a yellow patch or band. (zeus group)
6 (7) Forewing usually with a small spot in space 6. Z. zeus
7 Forewing without a small spot in space 6.
8 (9) Male underside hindwing overlaid with orange brown scales; uncus widest at its
midpoint. Larger, forewing 20.5-22.5 mm. Z. mars
Male underside hindwing brown, without orange brown scaling; uncus widest at its
base. Smaller, forewing 19.0-19.5 mm. Z. steineri
10 Forewing usually without a hyaline spot in the cell; the spots in spaces 2 andlor 3
sometimes absent. Male upperside hindwing with long hairs forming a dense,
localised patch near the base of the wing; upperside hindwing costa and underside
forewing dorsum with a polished area.
l1 (12) Head, thorax and wing bases conspicuously clothed with iridescent greenish hair
scales that, in the female, may appear indigo; male hindwing with a large patch of
black hairs contrasting strongly against a paler ground colour. Subgenus Matapoides.
Z. smaragdina
12 Head, thorax and wing bases not heavily clothed with iridescent greenish or indigo
hair scales; male hindwing with a small patch of dark grey or brown hairs that are
not darker than the ground colour. Subgenus Zampa
l3 ( l8) Male upperside forewing without a discal stigma. Female with more acute forewing
apex; generally smaller, forewing less than22mm. (zenon group)
14 (15) Upperside forewing unmarked. Z. cowani

t9
 15 Upperside forewing with one or more hyaline spots.
16 ( 17) Upperside forewing usually with well developed, hyaline spots in spaces 2 and 3,
the spot in space 2 sometimes reduced. Male hair patch on upperside hindwing
short, reaching to about half the length of the hindwing at the end of vein 4.
Z. zenon
17 Upperside forewing usually only with a small round or oval hyaline spot in space
3, the spot in space 2, if present, much reduced. Male hair patch longer, reaching
to about two-thirds the length of the hindwing at the end of vein 4. Z. storeyi
l8 Male upperside forewing with a discal stigma. Female with more quadrate forewing
apex; generally larger, forewing at least 2l mm, upperside forewing no spot in
space 2. (zero group)
19 (20) Forewing with a hyaline spot in space 3. Z. elioti
20 Forewing without a hyaline spot in space 3. Female unknown. Z. zero

The legends for Plate 57 should be changed as follows: 15. Zela onara
onaro 6,19. Zela ztoreyi l.

p.373 Erionota. Igarashi and Fukuda(1997) have found by breeding that "Erionota"
hormachis Hewitson, 1878, is the hitherto unknown female of (Jnkana
mytheca (Hewitson,1877), of which it becomes a junior synonym. Second
parugtaph under Erionota, delete "except in harmachis" and add "Antennal
club wholly or partly whitened." Key, delete lines 1 and 10. P. 374, delete
last paragraph on ^E harmachis and replace with the explanation above.

p.376 tJnkana.In view of the above, the first paragraph should read:

Adults rather large, and with elongate wings (forewing 2l mmand21,-25 mm

respectively in the male and female of U. mthyeca;27 and 3l mm respectively
in the male and female of U. ambasa). The wings are dark brown, the
forewing with hyaline spots, which are pale yellow in the males, white in the
female of U. ambasa and pale orange in the female of U. mytheca.

An amended key is given below:

| (2') d underside hindwing with a broad white discal band extending from vein lb to the
costa. t hindwing brown, forewing with coalesced pale orange spots; spot in space
3 reaches the base of the space and forms, with the spot at the cell-end and in spaces
2 and lb, a broad pale orange band. U. mytheca
2 d, ! underside hindwing with a whitened area extending from the base to the termen.
tupperside hindwing with a large white discal area; upperside forewing spots white
and widely separated, no spot in space lb. U. ambasa

Last paragraph under the heading Unkana ambasa batara, replace as

follows:

The very rare U. mytheca mytheca (Hewitson) (Plate 58, figs. 12, 13 6,
l8 ?) is easily recognised by the broad white discal band on the hindwing
beneath in the male and the coalesced pale orange spots on the forewing in

20
 the female. It appears to be confined to lowland forest in Malaya. The female
was, until recently, thought to be a species of Erionola because ofthe extreme
sexual dimorphism, which is unusual in the Hesperiidae. The species ranges
from Burma to Neomalaya and Nias. The genus Zea Distant was founded
on U. mytheca.

Plate 58, fig. 18, change from Erionota hormachls t to Unknna mytheca
mytheca l.

p.382 Potanthus hetaerus. The taxon serina Pl6tz is now considered a distinct
species as it occurs sympatrically withhetaerusMabille in Tawi Tawi (de Jong
& Treadaway, 1993). References to P. hetaerus serina should read
P. serina.

p.393 Checklist. The checklist should be updated as given in Eliot & Kirton (2000),
and should also incorporate the taxonomic information given in this update.
In addition, the following changes should be made:

Additional records from Singapore supplied by S.K. Khew are:

Graphium doson evemonides (Pulau Ubin)

Petralea dana (Pulau Ubin)
Bibasis sena
Tapena thwaitesi
Taractrocero archias quinta

Euthalia chersonesia (p.408) is given as a synonym of E. malaccana,but

this is incorrect. The name chersonesia is based on a female lubentina, and
is not a synonym of malaccono (Yokochi, l999a,b).

The order of species in the checklist for the genus Zela (Hesperiidae) should
follow the order given in the key in this update.

NEW INFORMATION ON EARLY STAGES

Unless otherwise stated, all the information in this section is based on discoveries
made by Suguru Igarashi, mainly working in the Malay Peninsula. Only the egg
(where known) and final instar lawa are here very briefly described, and the food
plant listed. Readers wishing to know more should consult the outstanding volumes
by Igarashi & Fukuda (1997,2000) wherein, in most cases, all stages from egg to
adult are figured in superb coloured plates and elaborated in the text, in both Japanese
and English.

p. 58 The larva of Troides aeacus has 6 pairs of fleshy spines per abdominal
segment (two subdorsal and four lateral), all tipped with pink, and a prominent

2t
 white saddle mark astride segments A3 and ,A,4. The subdorsal spine on A4
is white, tipped with pink.

p.60 Trogonoptera brookiana albescens. The food plant is Aristolochia

foveolata (Aristolochiaceae) (Goh, 199\; see also Igarashi & Fukuda(1997).

p. 6l Parides nox. The larva is similar in form to that of Troides aeacus, blt
differs in having the spines on,{3, A4 andAT white; food plant Aristolochia
tagala (Aristolochiaceae). The larva of Parides yoruna is very similar.

p.62 Add at end of line 1: and is almost indistinguishable from that of Parides
nox.

Parides sycorax. The tubercle on segment A7 is white, not that on A8.

The hostplants of P sycorax are Aristolochia coadunota (Aristolochiaceae)
(Igarashi & Fukuda, 1997), and a species of Thottea.

p. 65 Chilasa slateri. The larva is mostly blackish and is covered all over with
large and numerous yellow spots.

p.66 Chilasa agestor Add at end of the paragraph: and is very similar to that of
Chilasa clytia.

p. 68 Papilio demolion The lawa is green dorsally and laterally and dirty whitish
ventrally, with five irregularly spaced whitish-yellow bands.

Popilio nephelus. The larva is rather similar to that of P demoleus, but in

common with other related species has a pair of prominent, lateral eye-spots
on thoracic segment T3. The larval food plant is Micromelum minutum and
other species of Rutaceae (see Robinson et al., 2001).

p.69 Papilio prexospes. The green, more or less unmarked larva shows that it
belongs totheparzs group, which Igarashi separates into the genus Achillides
Hiibner, 1819.

p.73 Papilio palinurus. The green larva has also been found on Clausena
excavata and Tbddalia asiatica (both Rutaceae).

p.77 Footnote, add: The larva of Pathyso megorus is orange-brown with many
black bands bearing pale blue spots.

p. 84 Delias belladonna. The early stages in Malaysia are unrecorded. In the

Himalayas the mature lawa is blackish with a brown lateral band, and is

22
 covered with long but fairly sparse white hairs. The pupa is dark, with irregular
brown and white patches. The food plants are species of Loranthus and
aLso D endrotrophe granulata (Santalaceae).

p. 90 The hostplant of Appias nero has been recorded as Capparis (Capparaceae)

and Drypetes (Elphorbiaceae) in the Philippines (Robinson et al., 2001),
and Drypetes rn Java (Dupont & Scheepmaker, 1936).

p. 91 The larva e of Appias paulina and A. albina are most often found on Drypetes
spp. (Euphorbiaceae).

The larva of Saletara liberia, which closely resembles that of Appias

paulina, has been found in Palawan on Drypetes littoralis (Euphorbiaceae)
(Igarashi & Fukuda,2000: pl. 101, in error as Appias albina).

p. 100 Gandaca harina. The larva has also been found on Monocarpia marginalis
and Mitrephora maingayi (both Annonaceae). Robinson et al. (2001),
however, record only Connaraceae and Rhamnaceae as food plants for
Gandaca.

p. 104 The hostplant of Danaus melanippus is Cynanchum ovalifutium

(Asclepiadaceae) (Kirton & Kirton, 1987).

p. 106 Parantica. After line 1 add: the larvae have tentacles on T2 andA8 and are
generally blackish with a complicated pattern of spots, the most prominent of
which are usually yellow. The pupae are green, marked with black spots.

p. 107 First paragraph (Parantica). The "white" dots are pale blue and the larval
food plant is confirmed as Gymnema (Asclepiadaceae). Second paragraph:
Parantica agleoides has been bred in the Peninsula on Telosma cordata
(Asclepiadaceae).

Ideopsis. Penultimate sentence, add: The larvae of subgenus Radena are

very similar to those of Paranticabut those of subgenus ldeopsis are black
and white banded.

p. 108 Ideopsis juventa sitah. The alleged food plant Piper longum needs
confirmation. Ideopsis gaura has been bred in the Peninsula on an
unidentified vine and Hoya sp. (both Asclepiadaceae); the pupa is yellow
with black spots.

p. 109 ldea lynceus. Third paragraph: the larva of I. lynceus has been found and
bred on fu[yriopteron extensum (Asclepiadaceae).

23
p. 113 Euploea camoralzemon. In addition to Strophanthus dichotomrzs, Igarashi
& Fukuda (1997) record breeding this species on Cerbera manghas
(Apocynaceae). They show alarvathat is very similar in overall appearance
to that of E. phaenareta, and with filaments onT2, T3 and A8, and lacking
a pair on A2 - and thus different from the filament formula reported in
C&P4.

p. 115 Euploea mulciber. The larvae (illustrated in Igarashi & Fukuda, 1997) have
also been found on a number of species ofAsclepiadaceae andApocynaceae.

p. l16 Euploea phaenareta. ln the Peninsula, the larva is pale yellowish brown
with numerous naffow white bands outlined with black, and the filaments on
T2, T3 and A8 are tipped with white. The food plant is Cerbera manghas
(Apocynaceae).

p. 119 Melanitis. M. leda has been bred on Miscanthus, and M. zitenius on

Dendrocalamus giganteus (both Gramineae).

p. 120 Elymnios. The larvae bear rather short stout head horns bearing a few
branches or spines and the tail points are longer than usual for the subfamily.
The pupae are green, and often ornamented with red and yellow spots and
stripes.

p. l22The larva of Elymnias panthera generally resembles that of

E. hypermnestra. It is green, with two yellow dorsal lines that expand at
intervals, which are marked with an orange dot at the point of expansion
(Steiner, 200La). The head is patterned black and white, and the forked
posterior process is orange with black tips. The pupa is green, with yellow
dots and red-bordered white dots. The host plants are Calamus monan
(Steiner, 2001a), Archontophoenix olexandrae and Caryota urens (Igarashi
& Fukuda,1997).

Elymnias penango has been found feeding on a species of rattan thought to

be Plectocomiopsis geminiflorzs (Steiner, 2001a).

p. 123 Elymnias esoco. The larva appears to be unique in Satyrinae in feeding on

orchids (Aeriopsis) - but see also the note below on the morphine Faunis
conens.It is green with three dorsal blackish blotchy spots with relatively
short white tail points, and the head capsule bears many spines in addition to
the horns.

p.124 Lethe.ln L. europa and L. rohria the larval head horns are fused, produced
forward, and taper to a point, and the tail points are relatively long. In the
other Malayan species the head horns are separate and may be directed
forwards (L. chandica, L. verma, L. vindhya) or may be semi-erect

24
 (L. confusa). L. chandica has been bred on Dendrocalamus giganteus
(Gramineae).

p. 126 Neorina lowii. The early stages are now known. The egg is white, smooth
and nearly round. The fulI grown larva is dirty greenish. The head capsule
bears apair of long, smooth, adjacent head horns that are directed forwards,
so that the larva looks similar at both ends. The pupa is pale brown with long,
adjacent horns on the head. The food plant is Dendrocalamus giganteus
(Gramineae).

Xanthotaenia. The hostplants are species of Zingiberaceae, and notCalamus

(see earlier discussion under Classification of the Nymphalidae, and the
description of the life history given by Penz et aL.,2006.)

p. 127 Mycalesis. The larval head capsule bears very short, stout head horns and
the tail points are short.

p. l3l Orsotriaena. The larva of O. medus differs from Mycalesis in having long,
forwardly directed head horns and longer tail points.

p.I32 Erites angularis. The egg is very pale green, smooth and nearly spherical.
The full grown lawa is greenish white with shortish head horns directed
forwards, and feeds on species of bamboo.

The larva of Coelites epiminthia is green, with two dorsal yellow lines
(Steiner, 2001a). The two head horns are conical and are comparably long.
The pupa is green with yellowish-brown edging. The species has only been
found feeding on Calamus monan (Palmae).

The larva and pupa of Coelites euptychioides resemble that of Coelites

epiminthiablttend more towards a greyish colour (Steiner,200la). It has
been found feeding on Colamus castaneus (Palmae), and the larva is figured
in Davison (1995).

p. 133 See note in previous section on lawa and hostplant of Ragadia critolous.

p.134 Ypthima. The larvae may have smooth head capsules, or bear short, stout
head horns. The tail processes are short.

p.137 Amathusiini, last para1raph. Add: The eggs are smooth andnearly spherical,
and are usually whitish or pale green, with reddish brown marks showing
through the chorion as the larva develops within.

p. 138 Faunis canens. The description of the larva applies to the 4th instar. In the
5th and final instar the larva becomes blackish-brown, dorsally paler and

25
 with a yellowish lateral line, and it is clothed with tufts of short hairs as well
as with many long hairs. Additional food plants in Malaysia are Caryota
mitis, Pinanga scortechinii, Elaeis guineensrs (all Palmae) (Steiner, 2001a)
and a species of orchid (L. Kirton, pers. comm.).

p. 139 Eggs of Melanocyma faunula are laid in dense batches of several rows
(Steiner, 2001a). The gregarious larvae are deep red with white bands and
are covered with white hairs. The pupa is yellow and unmarked. The host
plant rs Orania silvicola (Palmae).

p.l4l Eggs of Amathusia ochraceofusco ochraceofusca are laid in batches in

one or more rows on the lower surface of leaves (Steiner, 2001a). The
gregarious larvae are brown in the final instar, with darker lateral and dorsal
lines (Maziah et o1.,1992; Steiner, 2001a). There is a shorter naffow white
line on the dorsum, and the whole body is covered with long hairs.The pupa
occurs in a green and pink-brown form. The food plants arc Calamus monon,
Salacca zalacca and possibly also Myrialepis paradoxa (all Palmae).

The gregarious lawae of Amathusia masina arc reddish-grey above in the

final instar, pale red laterally and dark red sublaterally, and are covered in
long brown hairs (Steiner, 2001a). The pupa resembles that of
A. ochraceofusca. The food plant rs Calamus manon (Palmae).

p.143 The final instar larva of Zeuxidia amethystus is green with a yellow band
behind the head and light red dots laterully, emphasised by long body hairs
(Steiner, 2001a). There are conspicuous lateral tufts ofwhite hair on a thoracic
and posterior abdominal segment. The head and horns at the anal segment
are green, and the horns on the head are orange-yellow. The pupa is green
with red lines and two red patches. The species feeds on Daemonorops
angustifulia, D. calicarpa and another species of rattan thought to be
D. hysterzr (Palmae).

The final instar lawa of Zeuxidia doubledayi is green with a black, yellow-
bordered band behind the head (Steiner, 2001a). It somewhat resembles that
of Zeuxidia amethystus blut the terminal tufts of white hair are especially
conspicuous. The pupa is reddish, marked with green lines. The species
feeds on Calamus monan and C. ornatus (Palmae).

p. 144 The larvae of Thaumantis klugius are solitary. They are covered with long
brown hairs, which are very dense in the last two instars, obscuring the body
colour (Steiner, 2001a). The head is black with conspicuous pink horns, and
the pupa is dull green with darker lines and dots. The species has been found
feeding on Calamus monan and C. caesius (Palmae).

26
p. 153 The larva of Cirrochroa tyche is blackish dorsally with a dorsal grey band
bearing black spots, and laterally grey. The branched spines are black. The
pupa is greenish-white with spiny processes that are shorter than those of
Cupha and Vagrons. The food plants are species of Flacourtia and
Hy dno c arpus (Flacourtiaceae).

p. 155 Theheadhorns ofkrinosareshort,beingmuchshorterthanthoseof Wndula;

additional larval food plants for Terinos are Homalium species
(Flacourtiaceae).

p.157 The larva of Cethosia biblis is red, with intersegmental bands of black,
edged with white.

p. 162 The larva of Symbrenthia hippoclus is black with two dorsal bands of white
spots, with lateral smaller white spots arranged irregularly, and four pairs of
scoli per segment. Food plants are Boehmeria glomerulifera, Oreocnide
trinervis and O. pedunculata (all Urticaceae).

p. 163 Rhinopalpa polynice. The resemblance of the larva to that of Vindula

must be superficial as the latter lacks the mid-dorsal scoli that are present in
Rhinopalpa. In India the larva has been found on Conocephalus
(Urticaceae) (Ackery 1988), but Igarashi & Fukuda (1997) record this
butterfly from Dendrocnide sinuata (also Urticaceae). Other records are
from Poikilospermum (Cecropiaceae).

Yoma. A larval food plant is Ruellia (Acanthaceae).

p. 167 Kallima limborgii. Additional food plants in the Peninsula are Gendarussa
vulgaris and S. pedunculosus (both Acanthaceae).

p. 168 Last paragraph. The larva of Cyrestis cocles is green dorsally and brown
laterally, with 5 lateral white spots and black dorsal spines. The pupa is
green with a long process protruding from the base of the thorax. The food
plant was an unidentified species of Moraceae.

p. 170 The larva of Chersonesia risabears oblique lateral white and blackish stripes
and has a continuous brown mid-dorsal band. The head is furnished with a
pair of long horns, and there are single dorsal, black-tipped processes on,{2
and A8. The striking pupa has two long curved processes protruding
downwards from the head and a very large irregularventral-projecting process
from the leading segments of the thorax. The food plants in Malaysia are
Ficus rocemoso, F. ischnopoda, F. pleyteana and E pustulata (Moraceae).
The larva and pupa of C. rahria are very similar, and the species has been
bred on E rocemoso.

27
p. 172 The larva of Neptis hylas,like those of all Sundanian Neptis larvae except
N. leucoporos, is broadly bicoloured, laterally darker and dorsally with a
long, paler 'saddle mark'extending from T2 toA8 and deepest about segments
A5 and A4, whence it decreases in width rearwards. This pattern is also
shared with Phaedymo columella and most species of Pantoporia.

The larva of N. leucoporos, which follows the alternative feeding pattern

mentioned on p.171, is green with very oblique paler lateral stripes, and does
not have a saddle mark; it also lacks processes on A2. The larva closely
resembles the buds of the food plant, Gironniera hirto (Ulmaceae).

p.175 Pantoporia hordonia. The points on T3 are not longer than those on 42,
and the usual paler saddle-mark is present; recorded food plants are Acacia
intsia and Porkia speciosa (both Leguminosae).

p.176 The pupae of Athymo are irregularly shaped with a prominent thoracic keel
and a large horn-like abdominal process; these sometimes meet to enclose a
circular hollow space. Rather long outwardly-curved processes protrude from
the head.

p. 177 Athyma asuro. The larva is blue-banded and spotted with yellow. The
fubercles are short and much branched, and yellow except on A2 which are
red. The species has not been bred in the Peninsula, but in Palawan the food
plants are Elaeocarpus elmeri (Elaeocarpaceae) and Chionanthus
ramiflorus (Oleaceae).

Athyma nefte. Larvae bred in the Peninsula are green with a large brown
patch on ,A.5, not A6 as stated in C&P4.

p. 178 The larva of Athyma nefte may also be all pale yellowish-green except for
segments A4 and ,{5, which are pale brown. The larvae and pupae of A. cama
and A. selenophoro are very similar to those of A. nefte and their main food
plant is also Glochidion (Euphorbiaceae).

p. 180 The structure of the larva and pupa of Lebadea martha is similar to that of
Athyma. The food plant is Mammea siamerzszs (Guttiferae). Lebadea is
transferred from Parthenini to Limenitidini.

p. 181 First paragraph. Final instar larrae of Parthenos sylvia bred in the Peninsula
by Igarashi are brown with longitudinal dorsal bands of apale reddish colour.
The spiny tubercles onT2 and T3, and on A7 andA8, are longer than those
on ,{1-A6. The green pupa is smooth and without any of the projections
characteristic of Athyma and Lebadea. Simllar early stages are shared by
the monotypic S.E. Asian genus Bhagadatta Moore and this genus should
replace Lebadea in the Parthenini.

28
 of I pelea but
p. 184 Tanaecia iapis. The green larva is generally similar to that
the oval dorsal spots are yellow rings. Food plants include Melastoma
malabathricum and Astronia macrophylla (Melastomataceae), Eurya
acuminata (Theaceae), and Mallotus subpeltatzs (Euphorbiaceae).

p. 186 The larvae of the Euthalia aconthea group are some shade of green with a
nalrow, unbroken yellow or whitish dorsal line. Those of the lubentina group
have large detached brownish spots enclosing a white dot on segments A2,
A3, and A5-A8, and more obscure spots on T1 andT2.

p. 188 Euthalia monina. The larva also feeds on Mallotus subpeltatus and
Macaranga hullettii (both Euphorbiaceae).

p. 189 Dophla evelina. The larvae resemble those of the lubentina group of
Euthalia except that there is an additional dorsal spot onA4. Food plants in
Malaysia are Lithocarpus falconeri (Fagaceae) and Antidesma salicinum
(Euphorbiaceae).

p. 190 Lexias dirtea has been bred in the Peninsula on Cratoxylum maingoyi
(Hypericaceae). The early stages of Lexia pardalis are, like the adults,
almost identical with those of L. dirtea; thelarvae in Laos feed on Hypericum
mono gynum (Hypericaceae).

p. 191 The description of the pupae of Pseudergolini in C&P4 is incorrect. Those

of Pseudergolis and Dichorragia have single thoracic and abdominal keels
which meet, or very nearly so, to enclose a circular hole; in other respects
the pupae are fairly smooth. The pupae of Stibochiona and Amnosia lack
keels and bear short, blunt tubercles on thorax and abdomen. The larvae of
Pseudergolini feed mainly on species of Urticaceae.

p. 192 The lawa of Stibochiona nicea is brown with a dorsal green 'saddle-mark'
extending from A2-A6. The long stout head horns are clubbed at their tip
and bear a few short spines. The spines on A8 are yellow. In India the food
plant is Boehmeria macrophylla (Urticaceae) but in Malaysia it was Ficus
lepicorpa (Moraceae).

Amnosiq decora.In Borneo the larvae have been bred on Pelionia scabra
(Urticaceae).

Dichoruagia. The larval food plants in Japan and Taiwan are species of
Meliosma (Sabiaceae). In Malaysia the food plants have not been discovered.

p. 193 The larvae of Rohana parisatis are green with a yellowish dorsal band
running from head to tail; they feed on Celtis spp. (Ulmaceae). The taxon

29
 ambico belongs to the genus MimathymaMoore, 1896. Its larvae and those
of other Mimathyma spectes feed on Ulmaceae while those of Apatura
feed on Salicaceae.

p. 194 Eulaceurq osteria. The larval head horns bear particularly long branches.
The larva is green with a whitish subdorsal stripe. An additional food plant is
Gironniera hirta (Ulmaceae).

Herona marathus. The smooth green lawa has irregular yellow dorsal
markings on most ofthe segments. The foodplants in India are Celtis species
(Ulmaceae).

p. 195 Euripus nyctelius. The green larva has three pairs of white flap-like
subdorsal tubercles onA2, ,{4 andAS and has been bred in the Peninsula on
Trema orientolis (Ulmaceae).

Sephisa chandra. The green larva differs from that of Euripus nyctelius
by lacking the tubercles onA2. The food plant in Malaysia is not known, but
elsewhere the species feeds on Querczs species (Fagaceae).

p. 196 Hestina. H. mimetica is usually considered to belong to the separate genus

Hestinalis Bryk, 1938. The early stages of Hestinalis mimetica have not
been recorded, but the lawa of Hestinalis nama has subdorsal pairs of short
branched tubercles on segments T2-A8 and feeds on Archiboehmeria
(Urticaceae).

Prothoini. The egg is spherical, slightly flattened at the top and almost smooth,
but with a few very inconspicuous vertical ridges. The larvae are long and
slender with a single pair of stout head-horns and feed on species of
Annonaceae.

p.197 Prothoefranck.Thelarva is long, thin and smooth, mostly pale brown, with
lateral blunt protuberances on segment 42. The food plant in Malaysia is
Annona muricata (Annonaceae).

Agatasa calydonio. In shape the larva is generally similar to that of Prothoe

franck, but the head horns are longer and end in a brown knob, and the
protuberances on ,{2 protrude much further. The final instar larva is dark
green, speckled with white dots, dorsally pale greenish-grey on segments
A7-A 10, and with a white sub-lateral patch on A6. The food plant is Desmos
chinens is (Annonaceae).

p. 199 The larva of Polyura hebe is light green with oblique bands of dark green,
and has been bred in Malaysia on Adenanthera pavonino (Leguminosae).

30
 The larva of Polyura delphis is green with two broad, irregular whitish
bands straddling segments ,A.3 and .A.5, and a pair of whitish, very broad,
dorso-ventrally flattened tail points. The foodplant rs Pithecellobiumjiringa
It also feeds on Intsia palembanica (H.S. Barlow pers. comm.), both
Leguminosae.

p.207 Abisara. The fine hairs on the larva are much longer than in other riodinid
genera and, in addition, there are brushes of hairs that protrude laterally
from the base of each segment. These brushes are particularly long and
thick in A. sovitri and its green pupa has a single dorsal black spot and is
oddly-shaped with Y-shaped lateralprocesses extending outwards from the
abdominal segments. The food plant is Embelia dasythyrsa (Myrsinaceae).
The pupae of other species, at least A. kausambi and A. saturata, lack
lateral processes and spots

p.227 Spalgis. The larvae also feed on other scale insects and are always covered
by the empty skins and white waxy substance secreted by their prey. This
covering is probably designed as aprotection against ants. The pupa is famously
similar in appearance to a monkey's face in miniafure.

p.230 Curetis. The larva of C. regula (described in error as C. saronis by Igarashi

& Fukuda, 2000) is reddish brown with white lateral patches on ,A.5 and
subdorsal whitish streaks; the food plant is Adenanthera povonina
(Leguminosae).

p.236 Pithecops corvus. The larva feeds on the flowers, fruits and young leaves
of Desmodium species (Leguminosae).

p.240 Acytolepis puspa. The larva seems to be polyphagous and has also been
bred in Malaysia on Glochidion zeylanicum, Bridelia whitmorei and
Antidesma leucocladon (all Euphorbiaceae).

p.247 Chilades pandava. The larva also feeds on Moullava spicata

(Leguminosae).

The larva of C. lajus feeds on Citrus (Rutaceae) and several other plants.

p.252 Jamides. The larva of J. malaccanus feed on Millettia sericea

(Leguminosae). The larvae of J. virgulatus and J. caeruleus feed on
Harpullia cupanioides (Sapindaceae), and of J. alecto on Zingiberaceae
(Igarashi & Fukuda,1997;2000). Jamides cyta feeds in the buds of Eugenio
diospyrifulia (Myrtaceae) (Kirton & Kirton, 1987).

p.253 The reddish-brown larva of Jomides bochus feeds on a wide range of

Leguminosae.

3l
p.268Arhopala. Evans gave the popular name oakblues to this genus, and this
name is suitable for nearly all the species since they feed on Quercus,
Lithocarpus and other genera of Fagaceae. However, the amphimuta
subgroup differs in that all the species whose life histories are known feed
on Macarango (Euphorbiaceae). Other known hostplants of Arhopala
belong to the family Dipterocarpaceae (L. Kirton, pers. comm.).

p.280 Arhopala inornata has been bred on Lithocqrpus cantleyanus, A. muta

on Lithocqrpus falconeri, and A. antimuta on Lithocarpus macphailii
(Fagaceae).

p.283 Arhopala zambra. H.S. Barlow (pers. comm.) has found the onisciform
larvae feeding on young leaves of Shorea macrophylla (Dipterocarpaceae),
a species introduced to Peninsular Malaysia from Borneo. The larvae and
pupae are attended by the ant Polyrhachis bellicosa. Fully-grown larvae
and pupae make a faint ticking noise to summon ants. When fully grown the
lawae follow a pheromone trail left by the ants to guide them to a camp, on
the groundwhere, tendedby ants, theypupate, up to 6-8 together. In captivity
the larvae can be bred through to adults without the presence of ants.

p.285 The larvae of the Arhopala eumolphus group, so far as known, feed on
Lithocarpzs species.

p.292 Zinaspa todara. The green larva has short brown and white oblique, lateral
stripes and feeds on Acacia intsio.

p.293 lroota. The eggs in this genus, as is the case with Neomyrina andthe following
genera up to Ancema excluding Catapaecilma acupicta, Acupicta and
Neomyrino, are dome-shaped and usually indented with comparatively large
hexagonal pits (sometimes with short spines at the corners of the pits) or, in
a few cases, smooth. In addition the pupae are nornally ungirdled. ln lraota
the eggs are white, the larvae some shade of brown. In Malaysia,.L distanti
has been bred on Ficus heterophylla and Ficus ischnopoda (Moraceae).

p.295 Catapaecilmo. Lawae of C. evansi feed on the durian tree, Durio

zibethinus (Bombacaceae) (L. Kirton, pers. comm., based on specimens in
the Malaysian Agricultural Research and Development Institute).

p.298 Neomyrino. The early stages of N. nivea are all white, from the egg to the
lawa,which has small lateral black dots on Al-A8. The smooth pupa, which
is pale brownish-orange, and the flattened onisciform lawa, confirm the
position ofthis genus close to Arhopala,aspointed outby Igarashi & Fukuda
(2000). In Thailand the species has been bred on Kurrimia paniculata

32
 (Celastraceae). The hostplant in Peninsular Malaysia is chengal
(Neobalanocarpus heimii Dipterocarpaceae). (L. Kirton, pers. comm.)

Loxura.In this genus and in Yasoda the eggs are smooth.

p. 300 Thamala. The white egg is rather shallowly dome-shaped and hexagonally
pitted. The brownish-red larvahas white bands onA2 and4.6 and has paired
pointed subdorsal and lateral processes, those on segments T3, A2 and A6
being particularly long. The smooth pupa is green, irregularly marked with
brown ventrally, and is suspended without a girdle. The food plant in Laos is
Semecarpus reticulata (Anacardiaceae).

p. 301 Cheritra. The larva of C. freja seems to be polyphagous. Other recorded

food plants in Malaysia are Monocarpia marginalis (Annonaceae), Deruis
elliptica and Adenonthera pavonina (both Leguminosae), and
Mischocarpus sundaicus (Sapindaceae). The egg is hexagonally pitted.
The pupa usually lacks a girdle, but one may be present (K. Fiedler, pers.
comm.)

p.304 Drupadia rovindra. Additional food plants are Schleichera oleosa

(Sapindaceae) (Igarashi & Fukuda,1997), Shorea maxima (Osawa & Kirton,
1997) and Hopea helferi (L. Kirton, pers. comm.) (the last two
Dipterocarpaceae). The species appears to be polyphagous. The egg is
hexagonally pitted.

Drupadia theda. The early stages are markedly different from those of
D. ravindra. The egg is smooth, and the smooth lawa, the colour of which
seems to depend on the colour of the food plant, e.g. often green, but red
when feeding on the flowers of lxora humilis (K. Fiedler, pers. comm.), and
brown pupa are like those of the Polyommatini. It seems unlikely that the
two species are congeneric (Igarashi & Fukuda, 2000).

p. 305 Horaga. The larvae are probably the most extraordinary of all the butterflies.
Those of H. onyx and H. albimacula are green, humped at the thorax and
front of the abdomen, then tapering towards the tail. There are a pair of
short, pointed tubercles onTZ, a long pair of brown subdorsal tentacles on
T3, a single mid-dorsal brown tentacle on A I , a very long green and brown
tipped mid-dorsal tentacle on,A.2 as well as a pair of similar lateral tentacles
extending sideways, a hump onA3, single mid-dorsal brown tentacles onA4,
A.5 and A7 , a long mid-dorsal tentacle similar to those on ,A.2 and, on A8, a
short mid-dorsal tubercle and a pair of very long green and brown-tipped
lateral tentacles extending sideways. Most of the lawa is green but with
some brown dorsal patches. The pupa is green with some dorsal brown
markings and is waisted between thorax and abdomen, and lacks a girdle.
The egg is hexagonally pitted. Neither species has been bred in Malaysia,

33
 but elsewhere the principal food plants are Glochidion species
(Euphorbiaceae), as well as Deutzia pulchra (Saxifragaceae) for
H. albimacula and Coriaria nepalensis (Coriariaceae) and Litsea
rotundifolia (Lauraceae) for H. onyx.

p.312 The egg of Tajuria cippus is smooth (Bascombe et al., 1999) whilst the
eggs of other recorded species of Iolaina are of the usual hexagonally pitted
form.

p. 313 The lawa of Tajuria maculata is quite unlike any other known Tajuria
species. It is brown with numerous yellowish spots and the pupa is similarly
coloured. The larvais not shouldered orwaisted, but more or less onisciform,
and has large single mid-dorsal triangular tubercles extended into long points
on segmentsAl-A7 ,andmuch smaller and blunterpaired subdorsal tubercles
on the thoracic segments. The food plant in Borneo was Scurrula parasitica
(Loranthaceae).

p.31 8 Remelana jangala. Egg flattened, dome-shaped, white and hexagonally

pitted. Lawagreen and onisciform, unlike those of other Iolaina; pupa attached
by a weak girdle. The species has not been bred in Malaysia. In Hong Kong
larval food plants include Litchi chinensis (Sapindaceae), Cleistocalyx
operculatus (Myrtaceae), Eurya japonica (Theaceae) and Kandelia
candel (Rhizophoraceae). The larva is affendedby Polyrhochis ants (Young,
leee).

Ancemo blankn. The white egg is hexagonally pitted. The green larva in
outline resembles the larvae of several Tajuria species. Food plants are
species of Loranthaceae. It is considered that Ancema and Pseudotajuria
should be transferred to Iolaina, leaving in Remelanina only Remelana,which
appears to form a link between Iolaina and Hypolycaenina.

p.324 Artipe eryx.Lawa has the thorax ochreous, segments A3 and A4 whitish,
the remainder brown. Food plants are the flowers and fruits of Gardenia
species (Rubiaceae).

p.325 Bindahara phocides. Lawa of the general shape of Deudorix and Artipe,
blackish with segments A3 and A4 white, bearing a prominent lateral black
spot. Feeds on the fruits of Euonymus and Salacia (both Celastraceae).

p.328 Rapala. The larvae always bear prominent paired subdorsal and lateral
fubercles, and feed on flowers, seed pods or leaves, never boring into fruit.
The larva of R. pheritima is pale yellowish-brown with a dark dorsal line
and large dark green lateral patches outlined with white on each segment.
The food plants are Derris thyrsiJlora and Pithecellobium jiringa (both
Leguminosae). H.S. Barlow (pers. comm.) has bred both R. pheretima and

34
 R. suffusa on the flowers of Bauhinia integrifulia (Leguminosae).

p.333 Bibasis. An additional food plant of B. harisa is Urophyllum arboreum

(Rubiaceae). B. seno has been bred in Malaysia on Pisonia grandis
(Nyctaginaceae).

p.336 Hasora schoenherr has been bred in Malaysia on Fordia ngii

(Leguminosae).

p.337 Choaspes. The black larvae divide into two types: a banded type, and a
spotted type with large dorsal and lateral yellow spots. Both types also have
small subdorsal blue or white spots. The yellow-banded types include
C. benjaminii and C. plateni with yellow bands on each segment, and
C. stigmata with only four white bands. The spotted type includes
C. subcaudatus, C. furcatus and C. xanthopogon. Additional food plants
are species of Tbrminalia, Combretum and Linociera spp. (all Combretaceae)
and Hiptage benghalerzsrs (Malpighiaceae).

p. 338 Celaenorrhinus. Additional food plants for C. qsmoro are Gendarussa

vulgaris (Acanthaceae) and Jasminum anastomosons (Oleaceae). The
foodplants of C. leucocera in India are Strobilanthes species (Acanthaceae).

p.339 Celaenoruhinusficulnea.Thelarva is pale grey with two fine lateral white

lines. The food plant is Didissandra frutescens (Gesneriaceae).

Celaenorrhinus aurivittatus. The larva is very similar to that of

C. leucocera and has been bred in Malaysia on Strobilanthes collinus
(Acanthaceae).

p.340 Darpa. D. pteria has been bred in Malaysia on Desmos dasymaschalus

(Annonaceae). In Laos, D. striata has been bred on Litsea monopetala,
L. lancifolia and Lindera assamica (all Lauraceae).

p.341 Sarangesa dasahara has been bred in Malaysia on Cyathula prostrata

(Acanthaceae).

Coladenia palawano has been bred in Malaysia on Allophylus cobbe

(Sapindaceae), and C. agnioides in Laos on Raphiolepis indica (Rosaceae).

Pseudocoladenia dan has been bred in Malaysia on Cyathala prostrata

(Acanthaceae).

p.342 Satarupa gopala has been bred in Laos on Euodia meliaefolia (Rrfiaceae).
The larva, which has a black head studded with brown bristles, is pale green
with yellow lateral spots on A2-A7 .

35
 Seseria ffinis has been bred on Litsea cubeba (Lauraceae).

p.343 Gerosis limax has been bred in the Peninsula on Abrus precatorius
(Leguminosae).

Mooreana trichoneura has been bred in the Peninsula on Mallotus

paniculatus and Macaranga trichocarpa (both Euphorbiaceae).

p.349 Halpe ormenes and H. flava have been bred on Dendrocalamus giganteus
(Gramineae).

p. 351 Pithauria marsena also breeds on Dendrocalamus giganteus (Gramineae).

p.354 Koruthaialos sindu feeds on Musa sapientum (Musaceae).

p.355 Psolos fuligo. The green larva has a double white spot on ,{6; it has been
bred on Donax canniformls (Marantaceae).

p.356 Ancistroides armatus has been bred on Hedychium coccineum

(Zingiberaceae) in B orneo.

p.357 An additional food plant for Notocrypta clavata is Hedychium coccineum

(Zingiberaceae).

p.361 Suastus. The hemispherical eggs of Suastus gremius are red when recently
laid, and have 14 white ridges (Steiner, 2001a). The final instar lawais whitish
green with a darker dorsal line. The head is grey-white with two broad,
parallel frontal black bands (Igarashi & Fukuda,1997; Steiner, 2001a). The
pupa is brown and covered with wax. Like Plastingia naga and Salanoemia
sala, the larva cuts its shelter off from the food plant prior to pupation and
pupates in the shelter on the ground (Steiner, 2001a). The recorded host
plants of ^S. gremius are Veitchia merillii, Rhapis excelsa, Pholidocorpus
macrocarpus, Chrysalidocarpus lutescens, Cyrtostachys renda, Livistona
sp., Licuala grandis, Daemonorops angustifulia andpossibly also Calamus
polystachys (all Palmae). It is often found on potted plants & planted
ornamental plants grown in towns. In Thailand, it has been found on Cocos
nucifera (Steiner, 2001a). The larvae of ,Saastus everyx and S. minutus are
quite similar to that of S. gremius, and feed on Doemonorops angustifolia
and a species of Calamrls, respectively (both Palmae) (Steiner, 2001a).

p.362 Zographetus. Z. ogygia and Z. doxus occur on Pithecellobium jiringa

(Leguminosae). Z. rama has been bred on Gnetum acuminata (Gnetaceae)
in Palawan.

36
p.363 Hyarotis. The lawae of Hyarotis adrastus and H. iadera are green with
ivory-coloured heads (Steiner, 2001a). In H. adrastus, the head has two
paler frontal lines, while in H. iadera (figured in Davison, t995), the head is
edged with black. The hostplants of H. adrastus are Calamus trachycoleus
and Daemonorops angustifolia (both Palmae). Hyarotis adrastus feeds
on Daemonorops grandis, D. calicarpa and D. verticillaris (all Palmae).

Quedara. The eggs of Quedara monteithi are yellowish, hemispherical

and smooth, and are laid singly (Steiner, 2001a). The larva is whitish green
with two white dorsal longitudinal stripes. In the last two instars, the head is
cream coloured with a wedge-shaped dark frontal stripe and two adjacent
white patches. The larva pupates on the lower surface of a slightly folded
leaflet, secured by a girdle. The pupa is pale green, with a thin orange-brown
dorsal longitudinal line flanked on each side by two widerwhite lines (figured
in Davison, 1995). The known host plants are Calamus manan, C. scipionum,
C. ornatus, C. speciosissimus, Daemonorops grandis, D. angustifolia,
D. didymophylla and Eugeissona tristis (all Palmae).

p.367 Eggs of Plastingia naga are laid in small batches. They are hemispherical
with strong ribs, and are reddish in colour when first laid. The final instar
larva is greenish white; the head is an ivory colour with dark brown to black
edges, and with the mouthpart region whitish (Steiner, 2001a). The host plants
are Caryota mitis (Igarashi & Fukuda, 1997; Steiner, 2001a) and Caryota
maxima (L. Kirton, pers. comm.) (Palmae). The lawabuilds a small pavilion
by folding in a small incised section of the leaflet. On pupation, this pavilion
is excised and the lawa pupates on the ground.

Salanoemia sala has been bred by Steiner and Aminuddin (Steiner &
Aminuddin,1,999; Steiner, 2001a). The yellowish white eggs are hemispherical
with 12-13 radial ribs, and are laid singly. In later stages, the larval head is
light brown with two darker parallel lines running from the top to the
mouthparts. The body is whitish-green with the abdominal contents showing
through the cuticle. The pupa is unmarked yellow, with the sheath of the
proboscis distinctly shorter than the abdomen. The host plants are Calamus
monan, C. scipionum, C. caesius and probably also C. ornotus (Palmae).

p.370 Pyroneura latoia and P. niasanq have been bred by Steiner (2001a) on
Oncosperma horridum and Pinanga scortechinii, respectively (both
Palmae). The final instar lawae are green, reddish in P latoia, and with trvo
paler dorsal lines. The head is ivory-coloured with dark edges and a dark
frontal line that are less pronounced in P. niasana. The mouthparts are
edged with white. ln P latoia, the dark frontal line is also edged with white
and there are two dark frontal eye spots on the head.

37
p.371 The larvae of Lotongus calathus have an association with the ant,
Dolichoderus thoracicus. The ants live and nest in the open-ended shelters
built by the larvae and often co-inhabit these shelters with the gregarious
larvae (Igarashi & Fukuda, 1997; Steiner, 2001a). The adult female seeks
out ant-infested rattans and lays its eggs among the ants (Steiner, 2001a).
The ants benefit from the relationship by the provision of nesting spaces,
while the larvae may benefit from protection provided by the presence of the
ants. All larval stages have an enlargement of the cuticle laterally, which
forms a downward fold over the legs that may protect the larvae from the
ants, although the ants show no signs of aggression towards the larvae. The
lawa is whitish green in later instars, and the head is cream coloured with
conjoined black blotches. The pupa is green with a red mark beneath each
eye. The species has only been found feeding on Calamus monon and
C. scipionum (Palmae) (Steiner, 2001a).

Lotongus avesta. The eggs of Lotongus avesta are dome-shaped with

eleven prominent radial ribs (Steiner, 2001a). The larvae are whitish-green
in later instars. The head is cream coloured, with a wedge-shaped dark
region frontally. The pupa is yellow with short hairs, and is covered with a
waxy substance. The host plant is a species of Salacca (probably Salacca
ffinis) (Palmae).

Zela excellens occurs on Calamus javenszs (Palmae) (Davison, 1995).

The larva is a leaf roller; in the final instar its head is orange and the rest of
the body white, being covered with a waxy secretion (Kirton & Eliot, 2004:
324,p1.1, figs. 13-15).Z.zeushas been bred on the palms Colamus caesius
and C. trachycolezs (Steiner, 2001a, b). The final instar larvahas an orange
head with two black frontal patches (Kirton & Eliot, 2004:325,p1. l,frg.22).
Z. steineri has been bred on Calamus manan (Palmae); the larva is
superficially similar to that of Z. zeus zeus (Kirton & Eliot, 2004: 327).
Z. cowani has been bred on the rattans Korthalsia rostrata and
K. scortechinii (Palmae) (Steiner, 200la,b). In the final instar the larva has
a dull orange head capsule; the pupa is buff and brown (Kirton & Eliot, 2004:
334, pL.3, figs. 6,7). Z. storeyi has been bred by Steiner on the rattan K.
scortechinii (Palmae). The final instar larva is similar to that of Z. cowani
(Kirton & Eliot, 2004:336, pl. 3,frg.I7).

p.373 Additional host plants of Gangara thyrsis are Calamus manan,

C. scipionum, C. caesius, Elaeis guineensis and Chrysalidocarpus
lutescens (all Palmae) (Steiner,200la; Maziah et al., 1992). The larva of
Gangara lebadea is grey-green and covered in white wax, with a dorsal
line of dark dots on the body. It lacks the white filaments found inG thyrsis.
The head has black marks on the frontal region, which form a star-like pattern.
The dark dots on the body and head are formed by the absence ofwhite wax
on these areas (Steiner, 2001a; L. Kirton, pers. comm.). Host plants are

38
 Calamus scipionum, Oncosperma horridum, Daemonorops macrophylla,
Adonidia merillii (Steiner, 2001a) and Arenga undulatifolia (all Palmae)
(Igarashi & Fukuda, 2000).

p . 37 4 Erionota hiraca (8. acroleuca tn C&P4: see Appendix) deposits eggs singly,
mainly on the upper surface of leaves. The larva is greenish and covered
with alayer of powdery white wax, and with the head black (Maziah et al.,
1992 - figured in error as E. torus). The pupa is unmarked yellow, with a
proboscis sheath approximately equal to the length of the abdomen. Host
plants arc Calamus manan, C. ornatuts, C. scipionum, Caryota mitis and
the ornamental palms, Areca triandra and Adonidia merillii (all Palmae)
(Steiner, z}}ta1'lllf:aziah et aL.,1992).Lawae partially cut the leaflet and roll
the severed portion, within which they feed. The lawa of Erionota sybirita
is whitish-green and similar to that of E. hiraca,butlacks a wax layer (Steiner,
2001a). The pupa is brown, with a long proboscis sheath that is up to 40mm
longer than the abdomen. The host plant is the fan palm, Licuala lunstleri
(Palmae).

p.375 Matapa. Lawae of M. druna and M. sasivqrna utilise Dendrocalamus

giganteus (Gramineae).

p.376 Unkana mytheca. The food plant is Pandanus odoratissimus

(Pandanaceae) (Igarashi, 1996).

p.377 Hidari irava also feeds on species of banana, bamboo and other palms,
including the sago palm. The larvae are gregarious (Khoo et al., 1991).

Eetion elia occtrs on Baccaurea sumatrana (Euphorbiaceae).

p. 378 The larva of Acerbas anthea is whitish green; in the final instar the head is
cream coloured and unmarked or with a weakly developed white, wedge-
shaped frontal patch (Steiner, 2001a). The slender pupa is unmarked yellow.
Host plants are Calamus scipionum) C. manan, C. caesius, C. trachycoleus
and probably also Daemonorops angustifolia (all Palmae). In Acerbas
martini,the larva is similar but the anal segment has a black sclerotised spot
flanked by two white semicircles (Steiner, 2001a), and the head has a well
developed white wedge-shaped frontal patch. The pale yellow pupa has a
black anterior spot and a black T:shaped pattern on the thorax. The host
plants are Kortltalsia rigida, K. lanceolata (Steiner, 200Ia) and Arenga
undulatifolia (all Palmae) (Igarashi & Fukuda, 2000).

Pirdana. Additional food plants for P hyela are Cordyline fruticosa

(Agavaceae),. and Peliosanthes viridis and Dracaena species (both
Liliaceae). P distanti occurs on Dracaeno elliptica (Liliaceae).

p.384 Pelopidas group. The larvae feed on Gramineae.

39
 ACKNOWLEDGEMENTS
My thanks go to Henry Barlow for kindly undertaking to see this paper through to publication, and
for his past help, energy and enthusiasm resulting in the publication of the last two revisions of The
Butterflies of the Malay Peninsula. But for him Corbet & Pendlebury would have ended with the 2nd
(1956) edition.
I also wish to thank Suguru Igarashi for his most generous gift of a number of treasured books
with superb coloured plates, which seem to be the monopoly and distinguishing mark of Japanese
authors. I am also grateful for his permission to summarise his breeding records. In addition, I wish
to thank Helmut Steiner for providing translations of relevant parts of his thesis on insects associated
with palms in Malaysia, and for allowing summaries of his findings to be used in this paper.
I am thankful to Dr. Takashi Yokochi for providing information and photographic illustrations
stemming from his work on Tanaecia and Euthqlla. Others to whom I am also grateful for information
and/or illustrations are the late Hugh Storey, C.Y. Chong and their co-workers, Khew Sin Khoon of
Singapore, Laurence Kirton and Henry Barlow. Mr. Teo Thiam Peng of Singapore has kindlypointed
out additional items fortheAppendix. Ms. Kim Chung (London) kindlypreparedthe initial typescript
from my handwritten text, and Ms. S. Lee handled the corrections in Kuala Lumpur.

POSTSCRIPT
At the risk of appearing maudlin I would like to take this opportunity to sing a sort of sentimental
"swan song". I have always felt a great affection for The Natural History Museum, long known to
most and hereafter as the "BIV[INH." Of the curators in charge of the butterflies, the first I was to
know was the distinguished Norman D Riley. When I was a schoolboy of l0 or I I years he told my
father that the Museum would be quite pleased to have a short series of Plebejus (Lysandra) hispana
from the lovely part of Provence where I was lucky enough to have my family home. So the next
Easter holidays I caught and set a short series of the spring brood which was duly handed over. I
remember the thrill when the great man himself showed it to me set out in a drawer of the finest
collection in the world, a thrill compounded soon afterwards by the receipt of a document from the
Trustees thanking me for my gift. The pride of a schoolboy in being associated even in a minimal way
meant a lot to a child, and this kindly and thoughtful action, which may have fostered my life-long
obsession with butterflies, was to be followed later by other kindnesses in both non-entomological
and entomological maffers.
When I retired from the Army in 1958, with an inflated idea of becoming an amateur butterfly
taxonomist, Riley (who had been Keeper of the Department for many years) was still in charge of the
butterflies. I told Riley of my absurd idea of doing an Evans, i.e. of classifuing the whole Lycaenidae
of the world, based on the Museum collection. He kindly advised me to try my hand with something
smaller first! Thus using BMNH material with his blessing I set about trying to solve various
problems in my own collection, resulting in my first publication which Riley kindly ensured was
published in the Museum's Entomology Bulletin (Eliot, 1959). Thereafter he gave me a free hand to
revise the genus Neptis (Eliot, 1969) and even to rearrange the collection, a trust that gave me much
pride and satisfaction.
It has been my great good fortune to receive the same treatment from the successive curators
of the butterfly section, namely Graham Howarth, Dick Vane-Wright and Phil Ackery which allowed
me many hours, probably better numbered in months, of contentment and happiness. How pleasant
it was to arrive mid-morning at the Museum after an early rise and a long and sometimes uncomfortable
journey from Somerset and to enter an atmosphere, which I find diffrcult to describe a sort of
-
amalgam of serenity and total friendliness which was not confined to the first floor, but extended
from the library to the basement. It was delightful to arrive to see an empty table, often bearing a
microscope of a magnificence and ease of use which I could never hope to match at home, and betimes
to see Mrs North advancing down the corridor bearing an unsolicited but much appreciated and
energy-restoring cup of steaming coffee, where I could be trusted to work unsupervised and to be
allowed to select overall some hundreds of butterflies on loan for leisurely dissection and detailed
study at home. I am now rather ashamed that I exploited this friendly trust by irregularly requesting

40
and receiving permission to stay on alone after working hours. I greatly appreciated the extra hour of
working time itris afforded me, in which I often seemed to achieve more than throughout the rest of
the day,before being courteously seen off the premises by some uncomplaining official late worker,
usually from another discipline, just in time to catch the last train home to Somerset.
I think it was Shakespeare, but I cannot put my finger on the quotation, who wrote that
comparisons are odious. I agree, but it is inevitable that among the many friends I made during my
buttirfly years, some names and incidents come to mind first. The first name must be that of Hugh
Storey, since he did so much to help in the production of C&P4. He and I shared an unusual common
misfortune; we both contracted poliomyelitis in fairly early adulthood. [, so much luckier than him,
made a miraculous recovery of two-thirds of my strength and was able to resume a normal life style
and to continue to serve in the Army for another 15 years - where few of my comrades ever
suspected that I suffered from any form of disability. Hugh, so much unluckier, became confined
to
a wheelchair, but nothing daunted, despite a quite heavy load of charitable work to help other
disabled people and other good works (see his obituary in The Malaysian Naturalist 55 part 3)
managed to iravel all over the Peninsula, including many visits to the Langkawi Islands and the
islands off the east coast, in an adapted Land Rover accompanied by Basri, his faithful companion
and assistant and a few friends with butterfly nets who did much successful collecting. This team
was known as Toshforce, and they accounted for many new records and species, Hugh's letters
bringing photographs for identification and an update of the latest news from Malaysia, with a
suitaUte sit of press cuttings. When precise identifications could not be made, the specimens would
usually follow for detailed examination using the Barlow shuttle, together with his generous permission
to give to the BMNH any specimen that I thought should be left there rather than in a private
collection.
Another important figure from the past was the late Brigadier Harry Evans, sitting with his
eye apparently glued to the microscope presented to him by the Bombay Natural History Society in
r..ognltio, of his classic and still widely-used work, The ldentification of Indian Butter/lies. He never
seemed to take a break, which I suppose he could hardly afford to do if he was to be sure of
completing his monolithic self-imposed task of classifying the Hesperiidae of the whole world.
Nevirtheless, he very kindly took time off to give me my first lesson in taxonomy and to teach me
his method of dry dissection of the male genitalia after wetting the abdomen with wood naphtha, in
a method I used on my own collection for a short time but which was hardly necessary in my leisured
home environment. This method, though quick, was rather less than satisfactory as it did not fully
disclose the whole armature and occasionally led to damage which would frustrate later workers.
However, I think more standard and slower methods would not have enabled Evans to complete his
work, so on balance the entomological world has been the beneficiary. As our acquaintance ripened I
grew to become extremely fond of the Brigadier and I think we developed a sort of empathy,
probably because we were both old soldiers who shared the same opinions on most topics. When I
saia gooaUye to him in 1957 on being posted to Malaysia, I could see his health was deteriorating but
he had become as permanent a fixture of the Museum as the building itself and its contents that I
think I expected to find him still beavering away in his accustomed place on my return 3 years later.
So it was with much sadness that I heard a few weeks later, through the kind offices of Graham
Howarth, that he had died.
One of the joys of my life has been the chance to welcome butterfly friends to my home. I
particularly relish the memory of the time spent there by Hugh Storey and Basri.
Another visit I remember with amusement was by the idiosyncratic but distinguished
photographer, Bernard D'Abrera, who came down to photograph the plates in C&P4. I had prepared
Ol bo*.r-of specimens carefully arranged to the correct dimensions for each plate, and supposed that
a long day's work lay ahead of us. In fact, I could hardly hand out the boxes fast enough, and
the
wholi job was compieted in the hour before lunch. I regret that I did not apply a gentle touch of the
brake to make ,u.. ihut any pin had not become loose. Had I done so we could have avoided the few
misplaced specimens that disfigure some of the plates in C&P4. Mea culpa. At lunch I remember we
discussed our mutual love of music, uninformed but genuine on my part, and Bemard told my wife
and me about the choir which he had formed and directed in Australia. We were rather surprised but

4l
entertained when, between the courses, Bernard started to sing excerpts from Grand Opera at the top
of his voice - what a character!
I remember with gratitude the many persons, too many to mention individually, who,
suppose started as "pen friends" but whom I feel became friends in a much wider and truer sense.
especially enjoyed those who sent me entertaining accounts of their adventures, amongst whom
must mention Trig Treadaway, the Philippines expert who introduced me to the fascinating Philippine
butterflies now under such threat. I would have loved to collect there, but only managed to do so just
outside the US Air base at Clarke Field, outside Manila, where I had been sent on duty, and where I
count myself lucky not to have left my bones when the pilot flying me out preferred not to use the
runway but to take off at right angles towards a jungle-covered hill which, to my relief, he cleared by
what seemed a matter of inches. All my life I have been lucky!
Among the many to whom I owe gratitude, my greatest debt is undoubtedly to my darling
wife of 57 years, Betty (nde Thornton; died 18 April 2001). She was unable to work up any interest
in butterflies in general, but did learn to love and recognise the butterflies which would drop down to
settle on the flowers beside her as she worked in her beloved rockery. Without her super-efficient
domestic backup and her tender toleration of all my foibles, the pursuit of my obsession would have
been impossible.

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43
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45
 APPENDIX

THE BUTTERFLIES OF THE MALAY PENINSULA, 4th EDITION

UPDATED ADDENDA AND CORRIGENDA

Page25: Of the Malayan endemics listed in the Table, Nacaduba russelli, Arhopala stubbsi and,Zela
eliotihave recently been found in Borneo.

Page 65: Chilasa paradoxa. The ?-f, leucothoides occurs in two slightly different phenotypes. So
far as known, the typical phenotype, of Bornean origin, has only been found in Singapore
and east of the main range, while to the west of the main range only the Sumatran phenotype,
with larger white submarginal crescents placed a little closer to the hindwing termen, has
been found. The latter, for which the name f. penomima (Martin, 1895) is available if
needed, is shown on Plate 3, figure 9.

Page70,line29, for "l9I0" read "l900".

Page 125,line T,for "Plate 15, figs. 14,16d,l5t"read"Plate 15, figs. 14,166,18 ?"

Page l39,line ll,168 readas 156f

Page 139, line25,15$ readas16S

Page 354: Koruthaialos. Butterflies are found sparingly in the hills above 2,500 feet which are
( 20 mm, t Zt mm; and with a paler,
generally similar to K. sindu, but larger (forewing
yellower and broader forewing band 5-6 mm wide. It is believed that these are a small
race of the Bornean species K.frena Evans, 1949, whose forewing length averages 2mm
gf,eater.

Page 373-74,432,and Plate 58, fig. 2.0: deJong & Treadaway (Igg2,Notizen tiber einige Erionota-
Arten nebst Beschreibung einer neuen Art, Entomol. 2.102:133-142) have shown that
the butterfly called Erionota acroleuca in this book must be known as Erionota hiraca
(Moore, 1881). The latter name was published one month earlier than acroleuca, which
becomes a synonym of hiraca.

Page 379 and 433, Plates 59 and 6 I : de Jong (1992, A note on three species of Taractrocera Bltler,
Zool. Med. 65 257-265) has shown that the butterflies named in this book as T. ziclea
zenia and T. aliena alieno must be known as T. luzonensls (Staudinger, 1889) zenia Evans,
1934, and T. nigrolimbata nigrolimbata (Snellen, 1876) (syn. aliena (Pl6tz, 1883)),
respectively.

Page 411, against P damajanti: for "K.M (2,4)" read "K.M (3,4)".

Page 418, against P achelous malu: for "Penang" read "K. Penang".

Page 426, against H. khoda: for "M 20" read "H 20" .

Page427, against P. dan: for"M 45" read"H45".

Page 433, against T. aliena: for "M" read "M (2)"

46
Page 433: Add apostarc (Snellen, [ 1 880]) after Parnara apostata (Snellen, [ 1 880]). There are two
further subspecies (Chiba & Eliot, 1991, Arevision of the genus Parnara Moore, with
special reference to the Asian species, Ty6 to Ga 42(3): 179-194).

Page437: Note (25), for S read ?.

Page449 The entry for Fiedler & Maschwitz should be dated 1989.

Page 512,line 3: For "Aracaceae'1read "Araceae".

Page 580: 324, delete (right and left valvae).

Legends to coloured plates:

Pl. 1l,fig.2is f .
Pl. 18,fig. 15 Taenaris horsfieldii birchi 6 Ul read as Melanocymafaunulafaunula $.
Pl. 18, fig. 16 Melanocyma faunula faunula $ read as Taenaris horsfieldii birchi d Un.
Pl. 19, fig. 5 is Q.
Pl.zl,frg.24is $.
Pt.24,fig.8 is j.
PL.25,fig. 19 is J.
Pl. 29, frg.20 is P hebe plautus J.
P1.34,fig. 10 is ?.
Pl. 41 , figs. 14 and 18 are $ .

Pl.42,fig.4is ? Un.
Pl.46,frg.27 is ? Un.
4l is A. normani and fig. 43 is A. phanda phanda.
Pl. 51, fig. 35 is S; fig. 36 is ?; fig.
Pl. 53, fig. 16 is C. agni subsp.; C. igna is a distinct Philippine species.
j
Pl. 54, fig. 9 is and fig. l0 is ? Un.
Pl. 56, fig.20 is ?.
Pl. 58, fig. 5 &6 are Unkana ambasa batara.
PI.60, frg.29 is ? Un.
Pl. 62, fie. 2 is ? -f andromima.
Pl. 63, fig.17 is ? Un.

47
 l. Ixias pyrene birdi S 9. Tanaecia violaria 16. Zizeeria maha
syntype t serica S

17. Zizeeria maha

2. Ixias pyrene birdi serica $
bicoloured Q dimorph 10. Tanaecia heringi (underside)
regolis ?
18. Koruthailos frena
j (underside)
3. Ragadia crisilda
critolina $
19. Koruthailos frena
ll. Euthalia agnis hiyamai Q (underside)
4. Ragadia crisilda
holotype j
critolina Q

20. Zela steineri

5. Acraea violae S holotype j

12. Euthalia agnis hiyamai

holotype j
(underside) 21. Zela mars holotype
6. Cethosia cyane ( j (Sumatra)

13. Euthalia agnis hiyamai 22. Zela mars holotype

paratype t 6f (underside)
(Sumaha)
7 . Cethosia cyane $
(underside)

23. Zela zenon $

14. Poritia ibrahimi (Balabak)
$

8. Cethosia cyane Q
24. Zela storeyi
15. Poritia ibrahimi $ holotype 6f
(underside)

Figures are not to a common scale. Figures are reproduced courtesy ofthe following contributors: the
late H.R.M. Storey, C.Y. Chong and co-workers (Figures l-4, 10, l4-I5,18-9), H.S. Barlow (Figure 5),
S.K. Khew (Figures 6-8,16-17), T. Yokochi (Figures 9, ll-13) and L.G. Kirton (Figures 20-24).

48
:15
49
Malayan Nature Journal 2006, 59(1), 51 - 61

Early Stages of Xanthotaenia busiris (Lepidoptera, Nymphalidae), and

the First Report of a LarvalAnal Comb in the Nymphalidae

CARLA PENZ', PHIL DEVRIES Iand LAURENCE KIRTON2

Additional key words: Satyrinae, Amathusiini, Elymnias, Faunis,hostplants, Zingiber,lawa,cervical

gland, anal comb.

Abstract: An illustrated description of the early stages of Xanthotaenia busiris from Peninsular
Malaysia, along with natural history observations is presented for the first time. Although found
widely among the Hesperoidea, our discovery that the larva of Xanthotaenia busiris possesses an
anal comb constitutes the first report of this structure for any member ofthe Nymphalidae, suggesting
it may occur elsewhere within the Papilionoidea. Morphological comparison with Faunis canens
(Amathusiini) and Elymnias hypermneslra (Satyrinae) confirms previous suggestions thatX busiris
should be placed in the Satyrinae.

INTRODUCTION
The butterfly fauna of Peninsular Malaysia is possibly the best known of any tropical
fauna. This stems from a historical interest in butterfly collecting, systematics, and
natural history that was manifested in early treatments (Distant 1882-1886; Seitz
1908-1928). These stimulated a variety of contemporary studies, but the four editions
of Corbet & Pendlebury 0934,1956,1978,1992) and the two books by Igarashi &
Fukuda (1997, 2000) have arguably had the strongest impact on our modern
understanding ofAsian butterflies and their early stages.
The southeast Asian butterfly Xanthotaenia busiris Westwood, 1858 is a
widespread species in lowland rainforests ofPeninsular Malaysia, Thailand, Burma
and Sumatra. Until recently it was placed as a member of the Amathusiini, and field
observations are unanimous in suggesting that the general wing-shape and flight
behaviors ofX busiris resemble members ofthe genus Faunis Hiibner (Morphinae,
Amathusiini) (Fruhstorfer 1913). Nevertheless, as originallypointedto by Fruhstorfer
(1913) in his treatment of the genus, "If Doherfy's statement that the larva of the
only species is entirely naked should be confirmed,thenXanthotaenia comes closest
to the Satyridae." Over 50 years later Col. J. N. Eliot noted from breeding records
of L. Kirton and E.W. Diehl that, unlike amathusiine larvae typically covered in long

1
Department of Biological Sciences, University of New Orleans, New Orleans, Louisiana 70148,
USA.
2 Forest
Research Institute Malaysia, 52109 Kepong, Selangor, Malaysia.

5t
setae, X. busiris lawae were effectively 'naked,' and placedX. busiris as a member
of the Satyrinae in Corbet & Pendlebury 0992). Although Eliot's description ofX
busiris caterpillars (ibid., p.126,'o ...green, smooth, with head horns and tail
points..."), fits the general satyrine gestalt offered in many reference works, none
of these characters are unique to Satyrinae (Freitas & Brown 2004).
Modern systematic methods seek to refine our conceptual ideas ofrelationships
by comparing characters among species and groups, and guide future areas of
investigation. Recently we were able to examine the morphology of early stages of
X. busiris in detail, and noted that its caterpillars possess characters unknown in
the Satyrinae. We therefore felt that a more detailed description of X. busiris
caterpillars was warranted, and such information would be useful to evaluate its
systematic position.
Accordingly, this report describes the early stages of X. busiris from two
sites in Peninsular Malaysia, and includes field observations on larval and adult
behaviors, and food plants. We also provide a comparison ofX busiris early stages
to Elymnias hypermnestra (Linnaeus, 1763) (Satyrinae) and Faunis canens
[Hiibner, 1826) (Morphinae, Amathusiini), plus various other taxa illustrated by
Igarashi & Fukuda (L997). We note several characters of X. busiris that have
never been reported in the Satyrinae or Morphinae, and discuss these in light of
future research directions. A forthcoming phylogenetic study by CMP will provide
additional evidence on the systematic position of Xanthotaenia.

Study sites. Observations and material were gathered from two sites in Malaysia.
(l) Forest Research Institute Malaysia (FRIM), Kepong Selangor (elevation >
200 m): this site was planted approximately 75 years ago with mixed dipterocarp
species on what was tin mining and agricultural land. (2) Genting Sempah, Pahang
(N3" 2l' 32.5",E101" 47' 38.9"; elevation 610 m): this site was once ateaplantation,
now overgrown with secondary vegetation and planted trees. Both study sites
constitute a patchy matrix of forest, plantations, gardens, open areas, and buildings.
Period of field observations: July, 2005.

RESULTS

Host Plant. Zingiber gracile Jack (Zinglberaceae). We found eggs and larvae
most often on isolated plants (n:8), and once on a dense clump of plants (n:1).

Egg.Milky white, round, smooth, 1.9 mm diameter (n:1); laid singly,20-70 cm

above the ground level on nearly mature, but still tender leaves (n:5). The day prior
to hatching, the brown caterpillar head capsule can be seen facing up though the
transparent chorion.

First instar (Fig. 1A). Head width:l.l mm (n:2),brown; vertex with two prominent,
conical protruberances each bearing a short, dorsolaterally directed scolus with a
short, pointed, terminal seta. When head is observed in frontal view the conical
protruberances form a wide V originating at the dorsal epicranial suture. Body

52
cylindrical, smooth, with two long, well separated caudae, each bearing a terminal
seta. Cervical gland present. Rudimentary anal comb present. Total length after
eating egg chorion approximately 7 mm (n:1). Body color of the newly hatched
lawa white, turning green after its first meal of leaf tissue.

Second instar. Head width:l .3-1.4 mm (n:5), green at the frontal region and
brown laterally, including scoli; surfaces of head and scoli covered with small
tubercules; scoli approximately four times longer than in first instar. Body green
dorsally and with a faint tinge or orange below spiracular line; venter white; caudae
white with a faint tinge of orange and slightly longer than in the first instar.

Third instar. Head width:l .7- 1 .8 mm (n:4). Similar to second instar except head
scoli are slightly longer and bear a small white marking attip;minute white tubercles
visible throughout the body; caudae slightly longer and thicker than in the previous
instar.

Fourth instar. Head width:2.6 mm (n:1). Similar to third instar except head scoli
are slightly longer and with outer and inner surfaces pale orange and creamy white
respectively; stronger orange color below spiracular line; caudae green.

Fifth instar (Fig. 1C, D). Head width:3.8-3.9 mm across the stemmata region
(n:2); dark green, changing color gradually to orange or yellow at the oral region; a
distinctive white stripe runs along the entire lateral margin ofthe head, from its base
to the tip ofthe scoli; mandibles yellow with finely toothed dark brown edge. Anterior
region of head covered by small pits, and posterior region covered by tubercles
bearing setae; all setae short. Two thick, blunt or at times slightly bifid scoli (Fig. 1C)
projecting at an angle >90o from each other when head is viewed from front
(3.1-3.5 ffiffi, n:2); green anteriorly, white posteriorly and at tip. Head scoli arise
from conical projections of the gena that give the head a conspicuous medial cleft.
Six stemmata, I-IV contiguous to each other, V positioned ventral to I-IV, and VI
set posterior to I-IV. Body green, with a brown-orange, subspiracular band running
the length ofbody. Cervical gland well developed. Legs andprolegs short, allowing
the body to be held close to the plant surface. Dorsal body cuticle with minute,
sharp, round spines of uneven size and numerous short, bifid setae (all clearly visible
at 30 X). Caudae (2.5-2.'l mm,n:2) green with a slight tinge of red-orange; thick
and densely covered with tubercles bearing sharp setae. Anal comb well developed,
composed of seven curved setae (max length:0.1 mm, n:2) that arise from the
dorsal anal lobe (Fig. 2).

Pupa (Fig. lE). Body length:31 mm (n:1 pupal skin). Thin, elongate, light brown,
thinly mottled with dark brown lines and blotches, and overall resembling a curled,
dead leaf. Headwith two short, wide projections compressed dorsoventrally. Thorax
with a keel at midline. Dorsal edge of wing pads slightly raised, forming small
ridges. Abdomen with lateral dark brown spiracular line that extends from wing pad

53
to cremaster. Abdominal segments immovable. In ventral view, cremaster nearly
as long as A5 + ,{6. Cremaster crown round, perpendicular to long axis of body.
Cremaster hooks long and simple. Silk nearly white.

Caterpillar behavior. We found that first to early third instar larvae made
characteristic feeding damage on the host plant. They ate achannel along one edge
of a leaf blade, leaving a thin pendant strip of leaf tissue, and the caterpillars rested
on this narow hanging peninsula (Fig. 1B). This behavior has not been previously
reported for satyrines or amathusiines, but it occurs in other groups ofNymphalidae
(see DeVries 1987). Generally we found fourth and fifth instars resting along the
dorsal mid-vein of a leaf. One newly molted fourth instar was found resting along
the stem near the terminal leaf of the plant, and one fifth instar was found resting at
the base of a plant with the head pointing downwards. The resting position for all
instars was with the head tilted slightly backward, with the body straight and snug
against the plant surface, and the caudae separated (Fig. lD). We found only a
single caterpillar per occupied plant, although in some cases we found more than
one leaf of the plant with first instar damage.

Adult behavior. Throughout its range all general accounts suggest that adults of
X. busiris are active during the morning hours, they fly close to the ground, often
darting erratically into low vegetation when startled (reminiscent of species of
Faunis), and there is no indication that X. busiris is uncommon (summarized in
Fruhstorfer 1911, Corbet & Pendlebury 1992). Indeed, one author suggests that
X. busiris has recently become so common in some areas as to be a nuisance
(D'Abrera 2001). Our observations at FRIM and Genting Sempah concur with
those of previous authors, excepting that we encountered X. busiris adults only
infrequently roughly three individuals per day. The adults are well known to feed
-
on rotting fruit juices, and at the time of our fieldwork fallen jackfruit, Artocarpus
heteroplryllus (Moraceae), was particularly attractive to X. busiris, along with
otherresident species of fruit-feeding nymphalid genera (e.g., Amathusia,Zeuxidia,
Thaumantis, Thauria, Founis (Morphinae, Amathusiini), Ragadia, Erites, Lethe,
Ypthima, Elymnias, il(ycaleszs (Satyrinae), Tanaecia, Euthalia (Limenitidinae),
Choraxes (Charaxinae)).

Comparison to Elymnias hypermnestra and Faunis canens. To elucidate its

systematic position, we compared 26 characters taken from X. busiris early stages
with the satyrine Elymnias hypermnestra and the amathusiine Faunis canens.
Eggs, larvae and pupae of E. hypermnestro were sampled at FRIM, and a fifth
instar larva of ,E canens was collected at Genting Sempah. Although eggs, first
instars, and pupae of F. canens were unavailable for sfudy, we obtained some
relevant information from the illustrations in Igarashi & Fukuda(1997).In addition
to figures provided here, supplemental color illustrations of the fifth instar and pupa
of X. busiris may be found in Corbet & Pendlebury 0992).

54
We found that X. busiris shares six larval and one pupal character with
E. hypermnestra (Table 1) and other satyrines. The illustrations in Igarashi & Fukuda
(1997) indicate that fifth instar larvae of Lethe and Orsotriaena have an
approximately square-shaped head, while in others (e.g., Ragadia, Mycalesis,
Yphthima, and all amathusiines) the head is rounded. All larval satyrines illustrated
by Igarashi & Fukuda (1997) have short setae in the frontal region of the head, and
this contrasts sharply with the exceptionally long setae of amathusiine larvae. The
small frontal circular pits and posterior tubercles shared by Xanthotaenia and
Elymnias are also clearly present in Lethe, Mycalesis, Yphthima, and Callerebia.
These pits and tubercles are absent in Faunis canens (Table 1) and also in all
amathusiines illustrated by Igarashi & Fukuda (1997). The fifth instar caterpillar of
Xanthotaenia shows the satyrine gestalt in having very short body setae and caudae
with protruding chalazae. Finally, no amathusiine pupae illustrated by Igarashi &
Fukuda (1997) possess the thoracic keel found in Xanthotaenia and Elymnias.
In summary this brief review of 26 characters corroborates the assessments
of Fruhstorfer (1913) and J.N. Eliot (in Corbet & Pendlebury 1992)thatX. busiris
is likely a member of the Satyrinae.

DISCUSSION

Several observations from this study may stimulate future comparative work,
particularly among the satyroid groups. For example, in the fifth instar, the shape of
the head and the presence ofpits in the frontal region clearly associatesXanthotaenia
with basal satyrines, yet neither of these characters were included in the most
comprehensive phylogenetic analysis ofthe Nymphalidae by Freitas & Brown (2004).
The fifth instar lawa of Melanitis leda (Fabricius,l775), the most basal satyrine in
their analysis, lacks filiform setae on the first thoracic segment, yet these setae are
present in X. busiris, E. hypermnestra and F. cqnens (Table l). Furthermore, all
satyrines examined by Freitas & Brown (2003) lack filiform setae on the third
abdominal segment (A3), but these are present in X. busiris, E. hypermnestra and
F. canens (Table 1). It is important to note, however, that the position of the filiform
setae onA3 varies among these species, being posterior to the spiracle inX. busiris,
and anterior to the spiracle in the other two species. Comparison of these and other
characters across a broad range of species will provide an important source of
information for elucidating the systematic position of Xanthotaenia.
Second, caterpillars from many groups of Lepidoptera are known to possess a
cervical gland (Scudder 1889, Mtiller 1886, Stehr Lg87).Available evidence suggests
these glands may function to disseminate defensive chemistry to repel enemies
(Weatherston et al. 1986, DeVries 1987, Attygalle et al. 1993, Hallberg & Poppy
2003), or in one riodinid butterfly, to mediate symbioses with ants (DeVries et al.
2004). Here we showed for the first time that X. busiris caterpillars possess a
cervical gland in all instars. This suggests two areas of comparative and experimental
investigation: what is the function ofthe cervical gland inX. busiris andits relatives,
and how widespread are cervical glands within the context of the Satyrinae and
other nymphalids? The latter question is important because the presence of a cervical

55
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57
 gland has been used to propose that the Southeast Asian Amathusiini is closely
related to the Neotropical Brassolinae (Corbet & Pendlebury lgg2),yet these glands
may be more widespread in satyroid taxa than previously conceived (i.e., Satyrinae
* Brassolinae + Morphinae).
Within Lepidoptera, larval anal combs occur widely in the superfamily
Gelechioidea and the families Tortricidae and Hesperiidae, and arettllized as higher
taxonomic characters (Stehr 1987, Hassenfuss & Kristensen 2003). The anal comb
functions as a mechanical latch mechanism in a frass ejection system that is driven
by a localized increase in larval blood pressure (Caveney et al. 1998). The system
can forcibly eject frass away from the larva considerable distances to eliminate
chemical cues used by potential predators and parasitoids, and thereby increase
larval fitness (Takabayashi & Takahashi 1989, Weiss 2003). The report here is the
first demonstration of a conspicuous anal comb possessed by a caterpillar in the
family Nymphalidae (Fig. 2). The evolution of forcible frass ejection appears to be
prevalent in caterpillars that build shelters, for example the Hesperiidae (Weiss
2003). Our observations indicate that X. busiris caterpillars do not build shelters,
but rather rest and feed in an exposed position on their food plants. The anal comb
of X. busiris is either unique to this butterfly, or perhaps forcible frass ejection
systems may be more widespread in satyrine caterpillars and other groups than
previously thought.
Finally, this study and the host plant checklist ofRobin son et al. (2}}|)indicate
that early stages of X. busiris feed on Zingiber (Zingiberaceae), while Eliot (2006)
reports that larvae also feed on Hedychium coccineum (Zingiberaceae). Together
these observations suggest that X. busiris host plants are members of the
Zingiberaceae, and that Corbet and Pendlebury's (1992) statementthatlarvae feed
on calamus (Arecaceae (: Palmae)) is in error (Eliot, 2006).
Like many others before it, this study showed the importance of early stages
in elucidating butterfly classification, and pointing to new areas of investigation
(e.9., Denis & Schiffermtiller 1775, Miiller 1886, Scudder 1889, DeVries et al.
1985, Penz 1999, DeVries et al. 2004, Freitas & Brown 2004 and references therein).
Despite the availability of modern works like Corb et & Pendlebury ( I 992) and
Igarashi & Fukuda(1997,2000) we find it surprising that so liffle detail is known of
the basic biology of even common tropicalAsian butterflies, particularly in the face
ofwidespread habitat destruction, urbanization and habitat loss. We hope this paper,
in some wdY, stimulates more comparative sfudies on the early stages of Asian
butterflies.

ACKNOWLEDGMENTS
We thank H. S. Barlow for facilitating our fieldwork at Genting Sempah, providing magnificent
hospitality, and for his passionate interest in natural history and conservation. We thank S. K. L.
Hok and Azmi Bin Mahyudin for help rearing caterpillars, Shawn Cheng for help in the field, and
Lillian Chua Swee Lian for identification of the host plant. FRIM facilitated our r...ur.h by providing
housing and laboratory space to CMP and PJD. This work was supported by a grant from the
National Science Foundation (DEB 0527441).

58
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Robinson, G.S., Ackery P.R., Kitching, I.J., Beccaloni, G.W. & Hern6ndez, L.M. 2001. Hostplants of
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60
Figure I - X. busiris natural history. A, First instar head in frontal view; B, caterpillar damage
to two leaves of Z. gracile showing channels and peninsulas; C, Fifth instar head in frontal
view; D, Fifth instar caterpillar in lateral view; E, Exuvia from pupa in lateral view.

everted anal comb

(seven setae)

Frgure 2 - Posterior region of the fifth instar X. busiris showing anal comb. Body setae and
chalazae were omitted from the illustration.
61
 THE MALAYSIAN NATURE SOCIETY

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Moths of Borneo Part 3 by J. D. HOLLOWAY, 1987, 199 pp., 20 plates. 106.25 t25
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103 b & w plates, 10 col. plates. 100 130

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Wayside Trees of Malaya (2 vols) by E. J. H. CORNER, 3rd edn, 1988; Vol. 1,

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An lllustrated Key to Malayan Hard Corals by A. G. SEARLE,2nd edn, 1980,

28 pp., 43 b & w plates. 9 15

Moths of Borneo: with special reference to Mount Kinabalu by J. D. HOLLOWAY,

1976,264 pp.,727 line figs., 31 b & w plates. 25 s0

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Back issues of The Malayan Nature Journal are still in stock. A complete book and price list is available on request at the Society's
Secretariat.

Substantial discounts are available to members. Please write or e-mail for more information on the Society and for membership forms:

THE MALAYSIAN NATURE SOCIETY

P. O. Box 10750
50624 Kuala Lumpur
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Tel: 03-2287 9422
Fax: 03-2287 8773
E-mail: mns @mns.org.my
Website: www.mns.org.my

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