Arhopala Centaurus Java

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Correspondence: Richard I. Vane-Wright, Department ofEntomology, the Natural History Museum, Cromwell Road,London SW7 5BD, UK.Email: [email protected]

Received 7 September 2004; accepted 15 March 2006.

The Arhopala butterflies described by Fabricius: A. centaurus isfrom Java, A. democritus from Phuket (Lepidoptera: Lycaenidae)

Richard I. VANE-WRIGHT and Harish GAONKARDepartment of Entomology, the Natural History Museum, London, UK

Abstract

The origins and identities of two Fabrician butterfly species now included in the genus Arhopala Boisduval,1832, are examined; the species are Papilio centaurus Fabricius, 1775, and Hesperia democritus Fabricius,1793 (Lepidoptera: Lycaenidae). To dispel recent confusion, complete taxonomic histories are presented

for both. The type locality ofArhopala centaurus is established as near Jakarta, Java, and that ofArhopalademocritus as Phuket Island, south-western peninsular Thailand. Amblypodia pseudocentaurus Doubleday,1847, described from Java, is re-established as a subjective synonym ofArhopala centaurus sensu stricto.

The subspecies ofcentaurus that flies in the Malay Peninsula is re-established as A. centaurus nakula (Felder& Felder, 1860). The distinct Australian species misidentified in recent literature as Arhopala centaurusshould be known as Arhopala eupolis (Miskin, 1890). Lectotype designations are made for the nominalspecies Papilio centaurus, Hesperia democritus, Amblypodia pseudocentaurus and Amblypodia nakula.

Key words: Arhopala eupolis, Arhopala nakula, Arhopala pseudocentaurus, Joseph Banks, lectotypedesignation, Niels Tnder Lund, William Hunter.

INTRODUCTION

The oakblues (Arhopala Boisduval, 1832) form themajor genus of the tribe Arhopalini Bingham, 1907, of

the lycaenid subfamily Theclinae (Eliot 1973; Megenset al. 2004a). Alternatively, following Eliot (1990), thismajor group is sometimes demoted to tribal rank, with

the Arhopalina included as a subtribe (Ackery et al.1999). Comprising approximately 200 species, Arho-pala is amongst the largest and most complex of generacurrently recognized among the butterflies (Megens et al.2004b). Distributed widely in the Oriental and Austra-

lian regions, adult Arhopala are generally found in pri-mary forest and second-growth areas, but some species

occur in coastal mangroves and woody savannahs. The

larvae feed mainly on Fagaceae and Euphorbiaceae, but

also utilize a range of other plant families, and always

appear to be associated with ants (Fiedler 1991).

This paper addresses a major nomenclatural problem

that currently affects one of the best-known Oriental

species of the genus. The problem has its origins in poor

scholarship by certain 20th century authors normally

credited with proficiency in such work. Despite thisbeing a very particular case, various arguments pre-

sented here have wide application, because they concern

a name introduced by J.C. Fabricius, Linnaeuss most

able and prolific entomological student. The original

descriptions of certain Fabrician species, belonging to

various insect orders, give very inaccurate information

about the geographic origin of the material on which

they were based. Our example represents a major subset

of these problems relating to critically important but

poorly documented specimens from Java, obtained by

Sir Joseph Banks, naturalist on Captain Cooks HMS

Endeavour. We also take this opportunity to give a briefaccount of the only other species ofArhopala named byFabricius, also misidentified in the past.

Abbreviations for depositories are as follows:

BMNH, 17531881 British Museum, 18811990 Brit-

ish Museum (Natural History), since 1990 the Natural

History Museum, London; EIC, former Honourable

East India Company Museum, London; HM, Hunterian


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Museum, University of Glasgow; LS, Linnean Society of

London; OUM, Oxford University Museum, Oxford;

ZMUC, Natural History Museum of Denmark, Univer-

sity of Copenhagen (formerly the Zoologisk Museum,

University of Copenhagen).

GENUSARHOPALA BOISDUVAL, 1832

(Amblypodia Horsfield, 1829, sensu auctorum: seeHemming (1967): 38, 58.)

Arhopala Boisduval, 1832. Type species by selectionby Scudder (1875): Arhopala phryxus Boisduval, 1832(phryxus is now regarded as a subspecies ofArhopalathamyras (Linnaeus, 1758); for an account of Linnaeanbutterfly types, see Honey & Scoble (2001)).

An Indo-Australasian genus distributed from Sri

Lanka, Pakistan and India to Taiwan, Japan, the Phil-

ippines, Australia and the Solomon Islands. The distinct

genus Amblypodia Horsfield, 1829, was formerly con-fused with Arhopala, due to the false designation ofPapilio apidanus Cramer, 1777, as type species. Thecorrect use ofAmblypodia, with Thecla narada Hors-field, 1829, as type species, and now placed in a separate

tribe (Amblypodiini Doherty, 1886), was determined by

Hemming (1967) (see Pitkin & Jenkins 2003). Papilioapidanus is the type species ofFlos Doherty, 1889, rep-resenting a group of species closely related to Arhopala,but currently separated from it. Evans (1957) gives a

detailed history of Arhopala and associated genera.Eight available generic names are currently included in

the synonymy of Arhopala: Narathura Moore, 1879;

Nilasera Moore, 1881 (of which the type species isPapilio centaurus Fabricius); Panchala Moore, 1882;Acesina Moore, 1884; Darasana Moore, 1884; SatadraMoore, 1884; Iois Doherty, 1889; and Aurea Evans,1957. For original references and type species, see Pitkin

and Jenkins (2003). Other key works on Arhopalainclude de Nicville (1890), Bethune-Baker (1903), Cor-

bet (1941b, 1946), Evans (1957), Eliot (1963, 1972,

1973), Seki et al. (1991), Corbet and Pendlebury(1992), Megens (2002), Huang and Xue (2004), and

Megens et al. (2004a,b).

ARHOPALA CENTAURUSFABRICIUS, 1775

Taxonomic history

Papilio centaurus Fabricius, 1775, has become one ofthe most thoroughly misinterpreted of Fabrician butter-

fly names. As related below, Steven Corbet (1941a) must

bear some responsibility for initiating the recent history

of errors. Usage for centaurus was previously well estab-lished, although not entirely soundly, by Fabricius

(1793), Godart (1824), Horsfield (1829), Hewitson

(1862), Butler (1870), Distant (1885), and de Nicville

(1890). Among the 19th century authors, only Double-

day (1847) made mistakes similar to those introducedby Corbet. Despite Corbets unfortunate intervention,

the correct application of the name centaurus to anOriental insect was otherwise universally maintained

right up to Evans (1957), by most authors until the

1990s, and by some to this day. So far as we have been

able to ascertain, after Doubleday (1847) and Corbet

(1941a), the first recent author to apply centaurus to anAustralian butterfly appears to have been DAbrera

(1977), acting on well-meant but incorrect advice

received from the late John Eliot and Charles Cowan.

Since then, more and more have followed this incorrect

path. It is this confusion that necessitates the historical

review presented here.

Fabricius (1775), who worked in London during the

summer months of 17721775 (Hope 1845: viii), orig-

inally described Papilio centaurus from an unspecifiednumber of specimens from the Banks Collection, and

gave its origin as Habitat in nova Hollandia (=Aus-

tralia). His entire Latin description, in its original lay-

out, is reproduced here:

Centaurus. 329. P[apilio]. P[lebeji]. R[urales]. alis caudatis,coerulescentibus; limbo fusco, subtus cinereis: maculis

baseos ocellaribus.

Habitat in nova Hollandia. Mus. Banks.

Alae anticae supra coerulescentes, margine exteriori et pos-tico fuscis; subtus cinereae, maculis quatuor vel quinque

fuscis, annulo albo cinctis, pone has fascia fusca, albo-

marginata, quae tamen marginem tenuiorem haud attingit.

Posticae coerulescentes, limbo fusco; subtus cinereae, basi

maculis sex vel septem fuscis, annulo albido cinctis; apice

obsolete undatae.

The Banks Collection (BMNH) now includes two

very old specimens over the name centaurus. In additionto this obvious source for original material, we have

searched the Sehested and Tnder Lund collection

(ZMUC) and the HM, where we also found very old

Arhopala specimens associated with the name centau-rus. Banks allowed Fabricius to take certain duplicates,and on occasion Fabricius passed some of these to other

colleagues (Fabricius 1784: 123). As noted in his auto-

biography, Fabricius had close links with Banks and the

Hunter brothers while in London (Fabricius 1805; see

Hope 1845: vi, viii), and with Niels Tnder Lund in

Copenhagen (Hope 1845: xv). However, as detailed


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Rothschild collections, and one in Oxford (ex West-

wood Collection, OUM).

Felder and Felder (1860) described a new nominal

species from peninsular Malaya as Amblypodia nakula.Five years later the Felders transferred nakula to Arho-pala, stating the provenance to be Malacca interior and

Sumatra, and comparing it to Fabriciuss descriptionofcentaurus (Felder & Felder 1865). Hewitson (1862)synonymized both Amblypodia pseudocentaurusDoubleday and A. nakula Felder and Felder (and byimplication, Horsfield and Moores interpretation of

centaurus) with Fabriciuss centaurus, a decision fol-lowed by Kirby (1871) in his influential world catalog.

The next author to study the material in the Banks

and BMNH collections, Joness Icones, and all post-Fabrician taxonomic interpretations, was Butler (1870).

The Banks Collection passed from the LS to the BMNH

in 1863, and Butler would have been able to make direct

comparisons with the rapidly growing BMNH collec-

tion, and consult all the literature. He stated that the

type of centaurus agreed with the Felders nakula. Atfirst sight it seemed surprising to us that he did not

mention any of the BMNH specimens listed by Double-

day (1847), or the three Horsfield specimens from Java

that had arrived only 10 years before (registered as

186015 in the BMNH annual registers). Instead,

Butler notes just one example of centaurus in theBMNH, a Sumatran specimen obtained in 1854 that

formerly belonged to Sir Stamford Raffles. Again,

regrettably, we have been unable to trace this specimen

for comparison (Sumatran centaurus is considered to

belong to the same race as material from the MalayPeninsula, the type-locality ofnakula; see below). How-ever, examination of other entries in Butlers catalog

shows in almost all cases the same pattern: citation of

just a single BMNH specimen. The implication is that

Butler simply considered the particular specimen cited

to be typical, in terms of phenotype, of the Fabrician

taxon in question. Butler is thus (unfortunately) silent

regarding the full range of BMNH material that was

then available to him for comparison.

Distant (1885) summarized the discussions to date

and stated that the true Fabrician centaurus, of which

he had examined the type in the Banksian collectionin the British Museum, was an Oriental species. De

Nicville (1890) reproduced all relevant previous

descriptions and stabilized the nomenclature of the Fab-

rician taxon as Arhopala centaurus. This was followedby virtually all taxonomists and faunistic workers on

the Oriental region, including Bell (1919: 447), who

published the first detailed life history of the species,

from southern India. Thus Moulds (1977), in his bibli-

ography of Australian butterflies, noted centaurus,stated that it was not Australian, and claimed that the

published type locality was an error on Fabriciuss part

(cf. Watkins 1923).

Thus from 1870 until Corbets intervention in 1941

(and even after that), all workers accepted that centau-rus s.s. applied to an Oriental rather than Australiantaxon. However, there was still uncertainty regarding its

origin. This is particularly clear in the work of those

who did not switch to the polytypic species concept, as

introduced by Karl Jordan and others in the mid-1890s.

Amongst butterfly specialists, Frederic Moore was

almost at the opposite extreme to Jordan: Moores gen-

era equated to species groups or even single collective

species in the new taxonomy, whereas his species were

mostly equivalent to what were increasingly regarded as

subspecies.

Moore stayed with this approach throughout his

magnum opus, the remarkable Lepidoptera Indica. Thisgreat series was continued after his death in 1907

by Charles Swinhoe, who evidently did not see fit to

alter Moores taxonomic style. Thus, in dealing with

what we would now consider to be a single polytypic

species, Swinhoe (1911) recognized four separate species-

level taxa: A. pirithous (Sikkim, Assam), A. centaurus(Burma, Singapore, Borneo, Sumatra, Nias, with

nakula as a synonym), A. coruscans (Andamans, SouthIndia, Ceylon), andpseudocentaurus (Java). In thisway the idea that the type material ofcentaurus couldhave come from the Asian mainland, as originally but

incorrectly proposed by Horsfield and Moore (1858),was maintained. This view was reinforced a few years

later by Watkins (1923), who stated that the type of

centaurus in the Banks Collection, though describedfrom New Holland, is certainly non-Australian. Wat-

kins then continued: As pointed out by Butler . . . it

belongs to the race nakula [sic!], Felder, and is probablyfrom Malacca.

As already indicated, Corbet (1941a) was responsible

for introducing several new errors. First, he claimed that

the male Arhopala specimen in the Banks Collection,which was long regarded as the type ofcentaurus, was

not authentic, partly on the grounds that it (and variousother specimens in the Banks Collection) did not have

any original labels. Secondly, that the original descrip-

tion fits a female rather than a male. Corbet further

suggests that neither of the two specimens ofcentaurusin the Banks Collection could have been present at the

time when Fabricius made his original description (prior

to 1775), and that the putative type was probably


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obtained by Koenig in the Malay Peninsula or Peninsu-

lar Siam. As Corbet correctly points out in the intro-

duction to his paper, Johann Gerhard Koenig collected

in those areas during 1778 and 1779 (see also below),

thus implying that the supposed type ofcentaurus wasa later addition to the Banks Collection, and that the

original material that he presumed Banks obtained inNorth Queensland must have been lost or destroyed,

either before or subsequently. A reprint of Watkins

(1923), annotated by Watkins and held in the BMNH,

may have influenced Corbets thinking on possible type

localities.

Despite Corbets views, those writing taxonomic or

faunistic accounts of the Oriental region and of Austra-

lia initially took no notice of these suggestions, and in

this they were supported by Evans (1957), who explic-

itly stated that he saw no reason to upset stability. A

further suggestion of Corbet (1941a) was that Fabri-

ciuss original description ofcentaurus best fits an Aus-tralian taxon, Arhopala eupolis (Miskin 1890). Evans(1957: 136), who revised the whole of this large and

difficult group, dismissed this idea as well, and acted to

maintain the Fabrician name for an Oriental species.

The increasingly influential Eliot (1972) at first followed

Evans but, in the revised third edition ofThe Butterfliesof Malaya Peninsula (Eliot, in Corbet & Pendlebury1978: 468), he unfortunately rejected Evanss plea for

stability and upheld Corbets interpretation. This had

been foreshadowed in print a year earlier by DAbrera

(1977), who noted that Eliot has drawn my attention

to this fact [that the type locality for centaurus was

originally given as Nova Hollandia] and to a ratherextensive correspondence between himself and Col. C.F.

Cowan on the subject, confirming Corbets arguments.

Seemingly Eliot did not do any research on this issue

himself but, in his revision of Corbet and Pendlebury

(1978), added the information supplied by Cowan (in

litt.) on Koenigs collecting activity in Malaya and

Thailand.

Eliot (in Corbet & Pendlebury 1978: 468), closely

echoing the words of Corbet (1941a: 100), confidently

stated there is every reason for supposing that it

[the centaurus specimen] was not in coll. Banks when

Fabricius described centaurus. However, we have beenunable to discover what this every reason might be!Neither Corbet nor Eliot gave explicit grounds for

rejecting the Banks type specimen as part of the type

series. As we demonstrate below by a re-examination of

various specimens and an analysis of Cooks itinerary,

we believe the conclusions of Corbet, as compounded

by Eliot, are erroneous. The changes they proposed have

now been followed uncritically by many taxonomists in

the Indo-Australian region, but not by all, and some

authors have inadvertently used both centaurus andpseudocentaurus as if they applied to two different spe-cies, when in reality they were dealing with one and the

same insect (e.g. Robinson et al. 2001).

Taxonomic conclusions

To arrive at defensible conclusions concerning applica-

tion of the name Papilio centaurus Fabricius, two issuesmust be resolved. First, what is the identity and status

of the centaurus type specimen that has been in theBanks Collection for so long? Apparently it was there

before 1791, the last time Fabricius studied in England,

and at least one specimen seems to have been depicted

by Jones as a Fabrician species during 17831785 and

thus, critically, before Koenigs material reached Lon-

don. Secondly, we must establish the true type locality.

While traveling between Queensland in Australia and

the Cape of Good Hope, HMS Endeavour touched landonly in New Guinea (very briefly), the small island of

Savu (or Sawu) south-west of Timor, and for about 2

months at Batavia (near modern day Jakarta), in Indo-

nesia. Although there is no direct evidence in the Banks,

Solander and Cook journals to show what kinds of

butterflies (for that matter, insects in general) were col-

lected where, there is evidence that they did collect in

Queensland, near Jakarta (Java), and finally at the Cape

of Good Hope in South Africa. There is no evidence

that they collected any material on Savu (H. Gaonkar,

unpubl. data, 2003).

Fabricius generally knew with confidence whichinsects Banks collected in Madeira, Rio de Janeiro, Pat-

agonia, Tahiti and New Zealand, and most of those

obtained from Australia. He did describe a Zygaena

from Madeira, a type of moth that could not possibly

have come from there (O. Karsholt, pers. comm., 2004),

and there may be other individual mistakes. However,

Fabricius was frequently confused about the provenance

of specimens that were collected when Endeavour madelandfall in Java and the Cape. Some of these cases were

correctly interpreted (and some also misinterpreted) in

the late 19th century. The reason for these errors regard-

ing butterflies from Java, the Cape and, in a few cases,Australia, was simply that neither Banks nor Solander

kept (or could keep) any records of their natural history

activities in Java or the Cape. Their serious illnesses, and

how numerous crew members of Cooks HMS Endeav-our died in Java and on the way to the Cape, are nowso well documented that we need not go into detail here

(Banks in Beaglehole 1962; OBrian 1987).


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For example, Papilio astenous Fabricius, 1775, ataxon that Walter Rothschild interpreted as belonging

to Troides helena (Linnaeus, 1758), was undoubtedlycollected in Java, and could only have been collected

there. But Fabricius gave the habitat as in Capite Bonae

Spei Mus. Banks . . ., that is, the Cape of Good Hope

(Rothschild 1895). In this way, Fabricius gave the hab-itat for many insects collected in Java either as Nova

Hollandia (Bankss collecting stop before Java) orCapite Bonae Spei (the Cape of Good Hope, which

was Bankss stop after Java).The original material ofPapilio centaurus must have

reached the Banks Collection before 1772 (Cooks

Endeavour, with Banks and his collections on board,returned to England in 1771; Beaglehole 1962). The

Banks Collection (BMNH) now contains two specimens

over the name centaurus. We consider that the assump-tion of Corbet (1941a), that both of these specimens

came from Malacca, must be wrong. Koenig certainly

collected a number of butterflies in Malacca during

1778 and 1779. Most of these specimens, now in

ZMUC and BMNH, were described by Fabricius in

1781 and 1787, and a few (smaller) species in 1793.

Any Koenig material from Malacca would have come

to Banks in approximately 1786, because all the speci-

mens that Koenig bequeathed to him came to London

after Koenigs death in India in 1785 (Dryander 1800:

vol. 5, 309). Fabricius would then have most certainly

described any new species in these accessions, either in

his Mantissa Insectorum in 1787 or in his EntomologiaSystematica in 1793 (e.g. see the account ofdemocritus

below). But centaurus was described in 1775, longbefore any of Koenigs material had reached Banks. It

seems that because Corbet appreciated this fact, and

accepted the Nova Hollandia source for centauruswithout question, he convinced himself that the material

in the Banks Collection could not have been collected

during the Endeavour voyage, and most likely had beensubstituted by material from the later Koenig accession.

That one or both of the specimens might have been

collected by Banks or other members of the Endeavourcrew in Java does not seem to have occurred to him. As

related further below, it does indeed seem plausible that

the second centaurus specimen in the Banks Collectionis just such a later addition, and could well have beencollected in the Malay/Thai peninsula by Koenig. But

we are confident that this is not the case for the other

Banks specimen that, for so long, was regarded as the

type. We now believe that this example was collected

during the Cook voyage, and it came from Java (see

further below, under type material).

Consequently, the conclusions of the majority of clas-

sical authors are here reinstated. In particular, Double-

dayspseudocentaurus from Java must be regarded as ajunior subjective synonym ofArhopala centaurus (Fab-ricius, 1775), sensu stricto. The name to be applied tothe race ofcentaurus that flies in the Malay Peninsula

is not subspecies centaurus, but centaurus nakula(Felder & Felder, 1860). The name of the separate Aus-

tralian species that has been confused with centaurus(and A. araxes) is Arhopala eupolis. The results of theresearch outlined above are in harmony with the Code

(see ICZN 1999: Articles 23.9 and 23.9.6, but see also

3.2). The overwhelmingly continuous usage of this

name for the Oriental taxon for more than 200 years

(and its continued use in many local checklists) is

enough reason to maintain stability in this case. Finally,

the type locality of the nominotypical subspecies of

Papilio centaurus should be regarded as Java (nearJakarta), not Malaya as proposed by Evans, Nova

Hollandia as given by Fabricius, or North Queen-

sland as suggested by Corbet.

Synonymy of Javanese butterfly to be knownasArhopala centaurus centaurus (currentlyknown asA. pseudocentauruspseudocentaurus)

Papilio (Plebeji Rurales) Centaurus Fabricius, 1775:520, no. 329. Lectotype male, Nova Hollandia

(recte Indonesia, Java, near Jakarta), in Banks Col-lection, BMNH, here designated (examined).

BMNHE# 668233. (Fig. 1.)Papilio centaurus Fabricius, 1775: Fabricius (1781):

117, no. 523; Jones (MS 17831785, volume 6: pl.

22, fig. 1, OUM); Fabricius (1787): 68, no. 646;

Butler (1870): 179; Evans (1957): 135; Zimsen

(1964): no. 118.

Hesperia (Rurales) Centaurus (Fabricius, 1775): Fabri-cius (1793): 275, no. 63.

Polyommatus centaurus (Fabricius, 1775): Godart(1824): 658.

Amblypodia centaurus (Fabricius, 1775): Horsfield(1829): 102; Hewitson (1862): 3,4 (may be a misi-

dentification according to Distant 1885); Butler(1870): 179; Kirby (1871): 419; Norman (1950):

813.

Amblypodia (Arhopala) centaurus (Fabricius, 1775):Watkins (1923): 205.

Arhopala centaurus (Fabricius, 1775): Felder and Felder(1865): 222; de Nicville (1890): 234; de Nicville

and Martin (1896): 464; Swinhoe (1911): 147; Bell


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Figures 14 1 Lectotype ofPapilio centaurus Fabricius, 1775 (male; nr Jakarta, Java 1770, ex J. Banks; forewing length 29.8 mm;BMNH). 2 Arhopala amantes (Hewitson, 1862) in HM (male; India?, collector unknown; forewing length ~28 mm). 3 Arhopalasp. indet., Sehested and Tnder Lund Collection, ZMUC (male; original source unknown; forewing length 25.5 mm). 4 WilliamJones original, unpublished illustration ofPapilio centaurus in the Banks Collection, painting made 17831785 (OUM). Figures13 show upper side on left, underside on right, label below.

1

3

4

2


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(1919): 448450 (biology); Evans (1957): 113;

Eliot (1963): 206; Fleming (1975): 39; Eliot (in

Corbet & Pendlebury 1978): 468; Robinson et al.(2001): 63; Vane-Wright and de Jong (2003): 124.

Amblypodia pseudocentaurus Doubleday, 1847: 24.Lectotype female, Indonesia, Java, BMNH, here des-

ignated (examined). BMNHE# 147889. Formallysynonymized with Arhopala centaurus (Fabricius)by Bethune-Baker (1903): 40. Syn. rev. (Fig. 5)

Amblypodia pseudo-centaurus Doubleday, 1847: West-wood (in Westwood 1852): 478.

Amblypodia pseudocentaurus Doubleday, 1847: Hewit-son (1862): 4, as synonym of centaurus; Corbet(1941a): 100.

Narathura centaurus (Fabricius): Distant (1885): 261.Nilasera centaurus (Fabricius): Moore (1881): 115.Amblypodia amazona Pagenstecher, 1890: 107 (Java),

nomen nudum: Bethune-Baker (1903): 40.Arhopala centaurus pseudocentaurus (Doubleday, 1847):

Fruhstorfer (1914): 158; Corbet (1941b): 163.

Arhopala pseudocentaurus (Doubleday, 1847): Corbetand Pendlebury (1978): 285; (1992): 275; Fleming

(1983): 85; DAbrera (1986): 576; Seki et al.(1991): 53; DAbrera (1998): 162; Bascombe et al.(1999): 209; Osada et al. (1999): 170; Robinsonet al. (2001): 63; Megens (2002): 35; Monastyrskiiand Devyatkin (2003): 37; Huang and Xue (2004):

195; Megens et al. (2004a,b).

Type material ofPapilio centaurus Fabricius

Papilio centaurus was described from an unstated num-

ber of specimens in Mus. Banks, purportedly fromNova Hollandia (Australia). As noted above, there

are two male centaurus in the Banks Collection. On thetestimony of Horsfield (1829), at least one should have

had a ticket in his own (Fabriciuss) hand-writing but,

as Corbet (1941a) pointed out, neither of these speci-

mens has a locality label, nor does either of them have

an original Fabrician ticket. This helped convince

Corbet that neither of them could represent original

material.

One of the specimens (BMNHE# 668232) has no

label at all. It is exceedingly low-set, on a coarse pin

that is different from the pin of the second specimen.The underside of this unlabeled, low-set specimen is

dull, like that ofA. centaurus nakula from the MalayPeninsula and elsewhere on the South-East Asian main-

land (see the underside of a specimen from Laos, beau-

tifully illustrated by Osada et al. 1999: 170). In ourview it is very plausible that this is a Koenig specimen

from Peninsular Thailand, collected during 17781779,

and added by Banks to his collection sometime around

1786. However, there is no independent evidence that

would back such an assertion. The real importance of

this specimen is that it is typical of nakula, and surelyhelped convince Corbet (who in his own mind was

only trying to decide between Malay Peninsula versus

Queensland as the likely origin) that the Banks speci-mens must have been from the former, and therefore

could not be original material for Fabriciuss 1775

description.

The second specimen (BMNHE# 668233), higher

set and on a different type of pin, is clearly the one

referred to by many authors as the Fabrician type. It

has an old label (Papilio P.R. Centaurus, Fab. Entom.

p. 520, n. 329), in addition to a more recent circular

label relating to its accession to the BMNH, a circular

type label, and a drawer label added by Watkins

(Fig. 1). However, the old label offers no proof of

direct connection with Fabricius, and is probably not

the ticket that Horsfield referred to. Lengthy but

superficial (in the sense that we have not undertaken

any biometric or colorimetric analyses) comparison

with a long series of both Malayan and Javanese cen-taurus leaves us a little uncertain regarding its origin,even if we restrict ourselves to a choice between just

Malaya and Java. This insect is more variegated

beneath than the other Banks Collection specimen,

being on the bright end of the range for the Malay

Peninsula population, and at the dull end of the range

for Java. As pointed out by Evans (1957), the mean

difference between the subspecies is clear, but there is a

lot of individual variation. The illustration in Jonessunpublished Icones (MS, volume 6, plate 22, OUM;reproduced here as Fig. 4) is, to our eye, well fitted to

the Javanese phenotype.

However, we also consider that Joness illustration fits

Banks BMNHE# 668233 very well, including wing

shape, setting angle, shape and disposition of underside

spots, and even color (Figs 1,4). A further insight into

the origin of this specimen is afforded by Bethune-Baker

(1903: 40), who evidently considered the Malay Penin-

sula form to be typical centaurus, and usually a dullerblue than the Javanese population. We quote (our

emphasis added):Pseudocentaurus Doubleday (amazona Pagenstecher).

This I take to be the Java form of centaurus, of which I have

a number of specimens before me: the blue of some, not all,

is rather brighter and bluer than ordinary centaurus Fabri-

cius; it is, however, a curious fact that my only specimen,out of a great number, which is exactly typical with the typespecimen in the Banksian collection is one from Java , and


9/17

is not of the dull purple of the common form, but rather

brighter and bluer . . . pseudocentaurus . . . must therefore

be sunk under centaurus Fab.

We imagine, had Bethune-Baker been apprised of

what we consider to be the true origin of the Banks

material, his observation would not have seemed curi-ous to him at all.

The loss of the original Fabrician label from the Banks

Collection type specimen, after so many years and the

vicissitudes that befall collections, is not convincing

evidence of replacement. Taking all the information

together, we are thus quite satisfied that this specimen

(Fig. 1) was originally collected by Banks in Java during

the very difficult phase of the Endeavour voyage, and isnot a later Koenig specimen originating from the Malay

Peninsula, or anywhere else. Accordingly, we designate

the Banks Collection type (BMNHE# 668233; Fig. 1)

as the lectotype of Papilio centaurus Fabricius, 1775.We exclude the second Banks Collection specimen

(BMNHE# 668232) from the type series, as well as

material labeled centaurus that we found in the HM

and ZMUC collections:

As already indicated, based on Fabriciuss known

links with William Hunter and Tnder Lund, we

searched for possible Banks Arhopala material in theHM (Glasgow) and the Sehested and Tnder Lund and

Fabrician collections in the ZMUC (Copenhagen). In

the HM we found a single Arhopala male, labeled Pap.Centaurus Fabr p. 117 No. 523 (Fig. 2). It is quite

similar to A. centaurus, and initially we thought it might

be a syntype, not least because it is set in such a similarfashion to the lectotype (cf. Figs 1,2). However, on the

evidence of the wide black margin to the upperside

wings, and details of the underside pattern, we believe

that this is an example of the related species A. amantes(Hewitson, 1862), a butterfly found from Sri Lanka and

southern India to Myanmar and Laos. It corresponds

well to the image given in DAbrera (1998: 161). Cooks

Endeavour did not make landfall anywhere in thisregion, and thus this specimen could not have been

collected by Banks (prior to 1775). Most likely it is a

contemporary specimen from India, an area from

which, by the mid-late 18th century, English collectorswere receiving much Lepidoptera material (Vane-Wright

& Hughes 2005: 254).

In the Sehested and Tnder Lund Collection

(ZMUC), there are male and female specimens ofArho-pala, both of which carry the name centaurus on oldlabels. The female, which we identify as A. anthelus(Westwood, 1851), is labeled H. Centaurus ex Ind.

Or. More challenging is the male. It bears a label that

appears to have been written at two different times, in

two different hands (Fig. 3). At the bottom, in faded

brown ink, is the single word Centaurus. At the top,

in black ink, it appears to say H. n. sp. ex Ind. orient,

e Sumatra. As pointed out to us by Ole Karsholt (pers.

comm., 2004), the work of Zimsen (1964: 12) suggeststhat most specimens in the Sehested and Tnder Lund

Collection had their original labels replaced by new

ones, written by some unknown person working in the

first half of the 19th century. In this context, inclusion

of Sumatra may be significant. So far as we are aware,

the first people to note in the literature that A. centaurusoccurs on Sumatra were Felder and Felder (1865) and

Butler (1870), the latter documenting a specimen then

in BMNH dating from 1854. Possibly this replacement

Sehested and Tnder Lund label was made rather later

than Zimsen suggests, perhaps even after the appear-

ance of the Butler catalog. However, as Ole Karsholt

(pers. comm., 2006) has further pointed out to us, it is

difficult to imagine anyone working in Copenhagen in

the 1800s having such detailed knowledge of tropical

butterflies.

With respect to the specimen itself (Fig. 3), initially

we thought that this might also be a genuine

A. centaurus, and so conceivably ex Banks. However, ithas wider black borders to the wing uppersides than

true centaurus, and the (damaged) hindwing undersidecannot be matched to any centaurus material that wehave seen. Notably, it has a well-developed tornal lobe.

Frustratingly, the critical forewing underside pattern is

not visible. It has been suggested to us that it could bean example of the Australian species A. madytus Fruh-storfer, 1914, but having compared it with A. madytusmaterial in the BMNH, we do not think it fits this

species either (cf. illustrations in DAbrera 1977: 312;

Braby 2000: 689). It seems to us more comparable to

the Oriental species A. bazalus (Hewitson, 1862). Whilewe remain uncertain as to its identity, it clearly does not

correspond to the lectotype ofcentaurus (Fig. 1), or toJoness painting (Fig. 4). The pin on which it is fixed is

old, short, and of a different gauge from that of the

lectotype. Ole Karsholt (pers. comm., 2006) has pointed

out that Fabricius was not in the habit of giving awayspecimens, unless duplicates, from his own collection,

so it is unlikely that he would have given such a speci-

men to Tnder Lund when no such specimen can be

found in his own collection in ZMUC. We conclude that

this specimen cannot have been part of the original type

series, and there is no evidence to suggest that it was

collected by Banks.


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Type material ofAmblypodiapseudocentaurus Doubleday

Amblypodia pseudocentaurus Doubleday, 1847, wasdescribed from eight syntypes in the collection of the

British Museum: one from Java. Presented by the Hon.

East India Company, one from Moulmein, four

from Silhet, and two from Ceylon, together witheight or nine additional specimens originally in the EIC

that should be included as part of the series by virtue of

the clear reference to Horsfields (1829) account ofcen-taurus. In the BMNH collections we have found onemale and two female specimens ofA. centaurus receivedfrom the EIC in 1860, registered as BMNH 186015

(according to Cowan 1975; the EIC collections were

dispersed to Kew and South Kensington during 1879,

but this is clearly not the case, at least for much of the

insect material). These must be the three specimens

listed by Horsfield and Moore (1858) as remaining in

the EIC collection at that time. In addition, we have alsolocated two specimens apparently presented by Hors-

field to the Felders that probably originated from the

EIC, and one apparently presented to Westwood. The

other three of nine EIC specimens noted by Horsfield

(1829) remain unaccounted for, including the one that

evidently passed to the BMNH sometime between 1829

and the mid-1840s (Doubleday 1847). Of the eight spec-imens listed by Doubleday as being in the BMNH col-

lection in 1847, we have located only the two from Sri

Lanka. From this total of eight putative syntypes that

we have located, we designate the ex-EIC female long

ago placed as the type in the BMNH as the lectotype

ofAmblypodia pseudocentaurus Doubleday, 1847. Thisspecimen (BMNHE# 147889) has been labeled accord-

ingly (Fig. 5), and the other seven specimens have been

labeled as paralectotypes (six in BMNH: BMNHE#

147890147895; one male in OUM labeled W Hors-

field Java/Westw/Arhopala centaurus f. pseudo-centau-rus Doubld. Named by H.H. Druce 19001904.).Horsfields material originated largely or entirely from

East Java, if it is necessary to delimit the type locality

more precisely.

Identity ofArhopala centaurus

Having located the authentic specimen on which Fabr-

iciuss Papilio centaurus is based, and established itsorigin, it remains to confirm the identity of centauruswithin the current framework ofArhopala systematics.In a key with 244 entries, Eliot (1963) divided the

species found in Malaysia into 27 species groups, of

which the centaurus-group was one, noting that the

group included about 15 species mostly from

Papua. Parsons (1998) now recognizes 18 species in

the group. We confirm that the lectotype ofA. centaurus(Fabricius, 1775), as designated here, runs to Eliots

couplet 155 (Eliot 1963: 206) where, based on the false

type locality of Malaysia due to Evans (1957), Eliot

recognized the Malaysian race of this butterfly as cen-taurus centaurus. In the later key to Malaysian Arho-pala revised by Eliot for Corbet and Pendlebury (1978,1992), the lectotype ofcentaurus runs out very early, atentry 4, to A. pseudocentaurus. Following Corbet andPendlebury (1978, 1992), amongst the Malayan fauna,

A. pseudocentaurus is distinguishable from all otherMalaysian species by the silvery green lines edging the

markings in the cell on the forewing beneath. Again,

we confirm that this characteristic is evident in the lec-

totype of centaurus. Corbet and Pendlebury note twoother species in the Indo-Malayan region with similar

venation and silvery markings in the forewing cell

beneath, A. araxes Felder and Felder from Sumatra,Sulawesi, etc., and A. amantes from southern Myanmar.

Distribution ofArhopala centaurus

Because of the confusion over application of the name

centaurus, as well stated by Distant (1885) more than100 years ago, accounts of the distribution of this spe-

cies are also confused in the literature. Following Seki

et al. (1991), we consider A. centaurus to be a polytypicOriental species, the collective taxon extending from Sri

Lanka and Nepal in the west to southern China, and

the western Lesser Sunda Islands and Philippines in the

east. Despite the claim by Bascombe et al. (1999),according to Vane-Wright and de Jong (2003) there is

no reliable record ofA. centaurus from Sulawesi. Thusalthough Detani (1983) recorded A. pseudocentaurus(here regarded as a strict synonym ofA. centaurus) fromthe Banggai Islands, immediately to the east of central

Sulawesi, the identity of the species recorded by Detani

must be regarded as very uncertain (Vane-Wright & de

Jong 2003). Based on Evans (1957), Eliot (in Corbet &

Pendlebury 1978) and Schroeder and Treadaway

(1999), 12 subspecies ofA. centaurus are currently rec-ognized:

Arhopala centaurus pirama (Moore, 1881), from SriLanka and southern India (note: Bethune-Baker(1903): 40, considered this to be a synonym of

coruscans); Arhopala centaurus pirithous (Moore, 1883), from

northern India to southern China and Hainan (Bas-

combe et al. 1999: 210), and recently noted fromVietnam by Monastyrskii and Devyatkin (2003) (but


11/17

Figures 57 5 Lectotype of Amblypodia pseudocentaurus Doubleday, 1847 (female; (East) Java, ex EIC; forewing length

26.3 mm; BMNH). 6 Lectotype ofAmblypodia nakula Felder and Felder, 1860 (male; Malaysia, Malacca, de Castelnau; forewinglength 27.0 mm; BMNH). 7 Lectotype ofHesperia democritus Fabricius, 1793 (male; Phuket 17781779, J.G. Koenig; forewinglength 15.5 mm; ZMUC). All figures show upper side on left, underside on right, label below.

5

7

6


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this record needs to be checked for possible misiden-

tification ofA. c. nakula); Arhopala centaurus coruscans Wood-Mason and de

Nicville, 1880, from the Andaman Islands (India);

Arhopala centaurus nakula (Felder & Felder, 1860),treated as A. c. centaurus by Evans (1957), from

Myanmar, Thailand, Indo-China, Malay Peninsula toSumatra and Borneo, and from Busuanga, Dumaran,

Linapocan and Palawan in the western Philippines

(Schroeder & Treadaway 1999; Treadaway in litt. to

RIVW, 23.xii.2003); also reported from China by

Chou (1994: 635), but needs confirmation regarding

subspecies identity;

Arhopala centaurus dixoni Eliot, 1978, from PulauTioman (Malaysia);

Arhopala centaurus centenitus Fruhstorfer, 1914,from Batu and N. Pagi islands (Indonesia);

Arhopala centaurus centaurus Fabricius, 1775,treated as A. c. pseudocentaurus Doubleday by Evans(1957), from Kangean, Java, Bali, Lombok and Sum-

bawa (Indonesia);

Arhopala centaurus cuyoensis Schroeder and Tread-away, 1999, from Cuyo (Philippines);

Arhopala centaurus babuyana Schroeder and Tread-away, 1999, from Babuyanes (Philippines);

Arhopala centaurus aglais Felder and Felder, 1865,from Leyte, Luzon, Marinduque (type locality of

the synonym A. setsuroi Hayashi, 1981), Masbate,Mindoro, Mindanao, Pollilo, Samar, Sibuyan and

Tawitawi (Philippines; data from Treadaway 1995;

Schroeder & Treadaway 1999);

Arhopala centaurus decimarie Schroeder and Tread-away, 1999, from Homonhon (Philippines); and

Arhopala centaurus dinacola Schroeder and Tread-away, 1999, from Dinagat (Philippines).

Synonymy of Malayan taxon to be known asArhopala centaurus nakula (currently known asA. pseudocentaurus nakula)

Amblypodia nakula Felder and Felder, 1860: 395.Lectotype male, Malaysia, Malacca interior (deCastelnau), BMNH, here designated (examined).BMNHE# 147896. (Fig. 6.)

Amblypodia centaurus centaurus (Fabricius, 1775):Corbet and Pendlebury (1934): 198; Pholboon(1965): 41, 42, 43, 64. Misidentifications.

Arhopala nakula (Felder & Felder, 1860): Felder andFelder (1865): 222, pl. 29, fig. 14; Moore 1879):

835.

Arhopala centaurus (Fabricius, 1775): Swinhoe (1911):147. Misidentification.

Arhopala centaurus centaurus (Fabricius, 1775): Evans(1957): 113; Fleming (1975): 39. Misidentifications.

Arhopala centaurus nakula (Felder & Felder, 1860):Corbet (1941b): fig. 70; Corbet (1946): 87; Corbet

and Pendlebury (1956): 328.

Arhopala pseudocentaurus (Doubleday, 1847): Fiedler

(1991): 167; Huang and Xue (2004): 195.Arhopala pseudocentaurus nakula (Felder & Felder,

1860): Corbet and Pendlebury (1978): 285; Corbet

and Pendlebury (1992): 275; Pinratana (1981): 75;

Seki et al. (1991): 55; Chou (1994): 635.

Type material ofAmblypodia nakulaFelder and Felder

Felder and Felder (1860) originally described Amblypo-dia nakula in a paper listing new butterflies from theMalay Peninsula. No specific distributional data were

given for nakula, or any of the other species describedat the time. Five years later the Felders moved nakulato the genus Arhopala, and gave two localities for itbased on material in their own collection: Malacca Inte-

rior (Com. de Castelnau) and Sumatra (Wallace). Ex

Felder material from both sources is now present in the

BMNH (received as part of the Rothschild Bequest). In

our view only Malayan ex Felder specimens can qualify

as potential syntypic material, as Sumatra was not men-

tioned in the original description, whereas the Malay

Peninsula was. Accordingly, we have selected a specimen

ex Felder Collection (Fig. 6) evidently collected by de

Castelnau as the lectotype ofAmblypodia nakula Felderand Felder, 1860. This specimen (BMNHE# 147896)

bears a label in W.H. Evanss handwriting dated 1955(Fig. 6), stating that the specimen was Incorrectly

placed over type ofvihara, and Is identical with typeofnakula (i.e. the Sumatra specimen now excluded asa valid type).

Identity ofArhopala centaurus nakula

If it is accepted, as established above, that Fabriciuss

centaurus is based on material from Java, then the cor-rect name for the South-East Asian race that includes

Malay Peninsula populations is A. centaurus nakula.

Distribution ofArhopala centaurus nakula

Arhopala centaurus nakula occurs from Indo-Chinathroughout most of Neomalaya (Moulton 1915), from

Myanmar, Indo China, Hainan, Thailand and the Malay

Peninsula to Singkep, Sumatra, Nias, Belitung, Bangka,

the Natuna Islands, Borneo, and Dumaran and the

Calamian group in western Philippines (Evans 1957;

Seki et al. 1991; Schroeder & Treadaway 1999).


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According to Chou (1994), this taxon also occurs in

China but, as noted above, this requires confirmation.

Bascombe et al. (1999) considered the Hong Kong pop-ulation to belong to subspeciespirithous.

ARHOPALA EUPOLIS MISKIN, 1890Synonymy of Australian taxon to be known asArhopala eupolis Miskin, 1890, sp. rev.(currently known asA. centaurus)

Amblypodia eupolis Miskin, 1890: 42. Two male, fourfemale syntypes, Australia: Cooktown and Card-

well, Queensland (in Queensland Museum, Bris-

bane) (Edwards et al. 2001: 200; not examined).Arhopala eupolis (Miskin, 1890): Bethune-Baker

(1903): 41.

Arhopala centaurus eupolis (Miskin, 1890): Waterhouseand Lyell (1914): 124.

Arhopala araxes eupolis (Miskin, 1890): Evans (1957):114; DAbrera (1971): 311.

Narathura araxes eupolis (Miskin, 1890): Quick(1974).

Amblypodia centaurus (Fabricius, 1775): Doubleday(1847): 24; Westwood (1852): 478; Corbet (1934).

Misidentifications.

Arhopala centaurus (Fabricius, 1775): Fiedler (1991):168; Megens (2002): 35; Megens et al. (2004a): 118;Megens et al. (2004b): 1193. Misidentifications.

Arhopala centaurus centaurus (Fabricius, 1775):DAbrera (1977): 311; Common and Waterhouse

(1981): 468; DAbrera (1990): 311; Parsons (1998):386387; Braby (2000): 687; Edwards et al.(2001): 199. Misidentifications.

Type material ofAmblypodia eupolis

Originally described from a syntypic series of both

sexes, collected at Cooktown & Cardwell, Queensland.

See Hancock (1995) for details.

Discussion ofArhopala eupolis

Following Corbet (1941a), DAbrera (1977) and Eliot

(in Corbet & Pendlebury 1978), the name centaurus

Fabricius has in recent years been applied erroneouslyby various authors to an Australian member of the

centaurus group. The species concerned should now beknown as A. eupolis, returned here to full and validspecies status. Corbet (1941a: 100), because of his mis-

taken belief that centaurus had an Australian prove-nance, considered the Australian eupolis (type localitiesCooktown and Cardwell) to be a strict synonym of

centaurus (the most likely type locality, if based onAustralian material, would have been Cooktown: Wat-

kins 1923). Evans (1957), followed by DAbrera (1971)

and Quick (1974), regarded eupolis as a subspecies ofa somewhat more restricted polytypic species, A. araxesFelder and Felder (1865). The latter (type locality

Sulawesi) is now regarded as a separate species thatoccurs from Sumatra and Java to the Lesser Sunda

Islands and the Sulawesi Region (see Vane-Wright & de

Jong 2003).

Distribution ofArhopala eupolis

Taking the works of Evans (1957), Common and

Waterhouse (1981), DAbrera (1990), Parsons (1998),

Braby (2000), Edwards et al. (2001), Tennent (2002)and Vane-Wright and de Jong (2003) together, this spe-

cies apparently occurs as just two weakly separated

races. Since Parsons (1998) proposed that the New

Guinea region populations formerly regarded as sub-

speciesphiltron Fruhstorfer, 1914, cannot be separatedfrom Queensland butterflies, nominate A. eupolis eup-olis is seen to extend along the Queensland coastalbelt, through the islands of the Torres Strait and the

whole of New Guinea, westwards to Aru and Kep.

Kai, and eastwards along the Louisiade Archipelago

to Tagula. Arhopala eupolis asopus Waterhouse andLyell, 1914, is found in north-western Australia and

Northern Territory, including Groote Eylandt. Both

Parsons (1998) and Tennent (2002) regard the

BismarckSolomons taxa previously included within

this collective taxon to comprise a separate species,

A. eurisus Druce, 1891.

ARHOPALA DEMOCRITUS(FABRICIUS, 1793)

Hesperia democritus was misinterpreted throughoutthe 19th century, notably by Butler (1870: 167) and de

Nicville (1890: 157), when this insect was confused

with butterflies such as Papilio bochus Stoll, 1782 (nowin Jamides), and Amblypodia nila Horsfield, 1829.Aurivillius (1898) was the first to indicate the true iden-

tity of this species, which he did by locating an original

Fabrician specimen in ZMUC, collected by Koenig dur-ing his travels to western Malaysia and Thailand in

either 1778 or 1779.

Aurivillius (1898) compared this Copenhagen type

with Arhopala albopunctata (Hewitson, 1869) andA. lycaenaria (Felder & Felder, 1860). Corbet (1934),who examined the ZMUC specimen and noted that

Aurivillius was not certain of the identity, was emphatic


14/17

that Hesperia democritus and Arhopala albopunctatawere the same, a view with which Evans (1957) agreed.

The type specimens of Hewitsons Amblypodiaalbopunctata were captured in Moulmein, about1000 km north of Koenigs presumed locality of Phuket

(see type material below). Evans (1957) recognized

four subspecies ofdemocritus. Among these, accordingto Eliot (in Corbet & Pendlebury 1992), the race found

in the main southern area of the Malay Peninsula, Arho-pala democritus lycaenaria, may represent a distinctspecies from A. democritus, but in the current faunalliterature it is still treated as a subspecies, and the two

might also be confused in the molecular literature (e.g.

Megens 2002; Megens et al. 2004a,b).

Synonymy ofArhopala democritus

Hesperia (Rurales) Democritus Fabricius, 1793: 285,no. 94. Lectotype male (abdomen missing; Corbet

1934), in India Orientali . . . (Phuket, south-

western Thailand), in Sehested and Tnder Lund

Collection, ZMUC, here designated (examined).

(Fig. 7.)

Hesperia democritus Fabricius, 1793: Aurivillius(1898): 147; Corbet (1934); Corbet (1941a): 105;

Zimsen (1964): no. 1042, incorrectly indexed as

Hepialus.Polyommatus democritus (Fabricius, 1793): Godart

(1824): 656.

(Hesperia democritus Fabricius, 1793: de Nicville(1890): 157. Misidentification ofJamides bochus(Cramer (recte Stoll) 1782).)

(Lampides democritus (Fabricius, 1793): Butler (1870):166. Misidentification.)

(Cupido democritus (Fabricius, 1793): Kirby (1871): 352,gives type locality as Ceylon. Misidentification.)

Arhopala democritus (Fabricius, 1793): Aurivillius(1898): 147, 171; Corbet (1934, 1941a): 105;

Evans (1957): 105, synonymy; Fleming (1975): 36;

Eliot (in Corbet & Pendlebury 1978): 291; Corbet

and Pendlebury (1992): 281, index, p. 527, incor-

rectly states originally published as sp. ofPapilio;Pinratana (1981): 84, as nominate subspecies from

Thailand; Seki et al. (1991): 72, 61; Megens (2002):

35; Megens et al. (2004a): 118; Megens et al.(2004b): 1193.Amblypodia albopunctata Hewitson, 1869. Two male,

one female syntype, Myanmar: Moulmein, ex

Atkinson Collection, in BMNH (examined; all

unaccountably discolored, but instantly recogniz-

able as democritus). Synonymized by Corbet(1934); see also Evans (1957: 105).

Amblypodia albopunctata Hewitson, 1869: Kirby(1879): 141.

Type material ofHesperia democritus

Hesperia democritus was described by Fabricius froman unspecified number of specimens (but probably just

one) in Mus. Dom. Lund . . ., collected by Koenig(Corbet 1941a). A single male in the Sehested and

Tnder Lund Collection, ZMUC, illustrated together

with its labels in Fig. 7, is hereby designated the lecto-

type ofHesperia democritus Fabricius, 1793. This selec-tion reflects Corbets (1934) earlier valid restriction, in

which he stated that there appears to be no valid

reason for regarding this specimen as other than the type

ofdemocritus.Taking into account the known distribution of

butterflies with a typical democritus phenotype (seebelow), the original material could only have been col-

lected either in western Malaysia (Kedah), or on the

island of Pulau Salang or Junk Ceylon, now known

as Phuket, the familiar holiday destination in Peninsu-

lar Thailand. Corbet (1941a) concluded that it must

have been the latter, and since there is no evidence

confirming or contradicting this, we accept his infer-

ence. Koenig spent several weeks on Phuket in 1778

and 1779.

Current status and distribution ofArhopala democritus

Arhopala democritus is not found in southern India, butis distributed from Orissa (H. Gaonkar, unpubl. data,

2004) to north-eastern India, Myanmar, Thailand,Malaysia and Neomalaya (Indonesia west of Sulawesi

and Java, including northern Borneo). The following

three subspecies are rather dully marked beneath: A. d.olinda (H. Druce, 1873), endemic to Borneo; A. d. bux-toni (Hewitson, 1878) restricted to Sumatra; and A. d.lycaenaria (Felder & Felder, 1860), the population thatoccurs south of Kedah to Singapore. These three could

collectively constitute a separate species (to which the

name Arhopala lycaenaria would apply) from democri-tus sensu stricto, which flies from Kedawi north to Thai-land and Myanmar, and is brightly marked with silvery-

white beneath (as suggested by the synonym albopunc-tata). Eliot (in Corbet & Pendlebury 1992) notes aspecimen of the bright phenotype collected as far south

as Frasers Hill, suggesting that democritus democritusand democritus lycaenaria overlap without intergrading(see also Fleming 1975, who implies overlap in Kedah).

Material from Orissa and NE India has yet to be deter-

mined to subspecies.


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ACKNOWLEDGMENTS

We are very grateful to our friends and colleagues Phil

Ackery, Kim Goodger and Julie Harvey (BMNH), Ole

Karsholt, Niels Peder Kristensen and Nikolaj Scharff

(ZMUC), Stella Brecknell, Darren Mann, George

McGavin and Stephen Simpson (OUM), C.G. Tread-

away (Limbach, Germany), and Geoff Hancock and

Margaret Reilly (HM) for suggestions, assistance, and

sharing their knowledge. Harry Taylor (BMNH) took

many of the digital photographs, and Campbell Smith

kindly designed the figure layouts. We are grateful to

Geert Brovad (ZMUC) for the images of butterflies in

the Sehested and Tnder Lund collection. The image

from Joness Icones was kindly prepared by RennisonHall (OUM). HG acknowledges the EU-funded Sys-

Resource programme for support while working in Lon-

don. We thank the Department of Entomology (BMNH)

for helping to defray the costs of color reproduction.

The first draft of this paper was written by HG, withsubsequent versions by HG and RIVW. However, the

final manuscript was prepared and submitted by RIVW

alone. Subsequently, one of the referees pointed out

significant errors in the treatment of the HM and

ZMUC material, and the paper was re-drafted by RIVW

to take this into account, again without the benefit of

input from HG. Finally, the author sequence was

reversed, to reflect the fact that RIVW has sole respon-

sibility for the final content. However, it was HG who

first realized that Arhopala centaurus was based onJavanese material, and carried out a large part of the

original work.

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Bell TR (1919) The common butterflies of the plains of India.

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