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Threatened 
Swallowtail Butterflies 
of the World 





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PAPILIO INT. 3 (M), 1986: 207 - 226 ISSN 0109-700B 

date of publication. July 19B7 207 



PAPILIO HIPPONOUS IN THE lUCN 
SWALLOWTAIL RED DATA BOOK, 
A CORRECTION 



J. Haugum Z> N. M. Collins 

Owing to an unfortunate misunderstanding the ranges of P_. 
[Princeps l hipponous Felder & Felder 1862 and P^. [Princeps ] 
pitmani Elwes C de Niceville IBB6 given in the lUCN' Red 
Data Book "Threatened Swallowtail Butterflies of the World" 
(Collins & Morris. 1985] were mixed up. The correct entries 
are as follows: C'p-'^ y<^^ '^t>c:U 'P'OO '0\ 



Digitized by the Internet Archive 
in 2010 with funding from 
UNEP-WCIVIC, Cambridge 



http://www.archive.org/details/threatenedswallo85coll 



Threatened Swallowtail Butterflies 

of the World 

The lUCN Red Data Book 



INTERNATIONAL UNION FOR CONSERVATION OF 
NATURE AND NATURAL RESOURCES 

Threatened 
Swallowtail Butterflies 
of the World 

The lUCN Red Data Book. 



N. MARK COLLINS 

lUCN Conservation Monitoring Centre, Cambridge. U. K. 



MICHAEL G. MORRIS 

Chairman. lUCNISSC Butterfly Specialist Group 
Institute of Terrestrial Ecology. Wareham, U.K. 



With the help and advice of the network of lUCN 
Commissions and other experts throughout the world. 



Photographic plates by 
N. MARK COLLINS 



PUBLISHED BY lUCN, GLAND. SWITZERLAND 
AND CAMBRIDGE, U.K. 1985. 



lUCN 

lUCN (International Union for Conservation of Nature and Natural Resources) is a 
network of governments, non-governmental organisations (NGOs), scientists and 
other conservation experts, joined together to promote the protection and 
sustainable use of living resources. 

Founded in 1948, lUCN has more than 450 member governments and NGOs in over 
100 countries. Its six Commissions consist of more than 700 experts on threatened 
species, protected areas, ecology, environmental planning, environmental policy, 
law and administration, and environmental education. 

lUCN 

monitors the status of ecosystems and species throughout the world; 

plans conservation action, both at the strategic level through the World 

Conservation Strategy and at the programme level through its programme 

of conservation for sustainable development; 

promotes such action by governments, inter-governmental bodies and 

non-governmental organisations; 

provides assistance and advice necessary for the achievement of such action . 

Published by lUCN, Gland, Switzerland and Cambridge, U.K. 
Prepared with the financial assistance of the World Wildlife Fund, the United 
Nations Environment Stamp Conservation Fund, the United Nations Environment 
Programme, Exxon Corporation and Citibank. A contribution to the Global 
Environment Monitoring System. 

^ ^) ^ CITIBANKO 

© 1 985 International Union for Conservation of Nature and Natural Resources 
ISBN No.2 88032-603-6 

Citation: Collins, N.M. and Morris, M.G. (1985). Threatened Swallowtail Butterflies 
of the World. The lUCN Red Data Book. lUCN, Gland and Cambridge. 
viH- 401pp. -I- 8 pis. 

Cover illustration: Papilio homerus by Sarah Anne Hughes. 
Book Design by James Butler. 

Typeset by Text Processing Ltd. , Clonmel, Ireland. 

Printed by Unwin Brothers Limited, The Gresham Press, Old Woking, Surrey, U.K. 



Contents 



Page Number 

1. How to use this book 1 

Acknowledgements 6 

2. The swallowtail butterflies: an introduction to the family 

and its conservation 7 

3. Swallowtails of the world: their nomenclature,distribution 

and conservation status 33 

Swallowtails of the world: an annotated species list 33 

Geographical index to species 124 

4. Analysis of critical faunas 137 

5. Trade in swallowtail butterflies 155 

6. Reviews of threatened species 181 
BARONIINAE 

1. Baronia brevicornis 182 
PARNASSIINAE 

Parnassiini 

2. Parnassius autocrator 185 

3. Parnassius apollo 187 
Zerynthiini 

4. Bhutanitis mansfieldi 192 

5. Bhutanitis thaidina 194 

6. Bhutanitis ludlowi 196 

7. Luehdorfia chinensis 197 

8. Luehdorfia japonica - 198 
PAPILIONINAE 

Leptocercini 

9. Teinopalpus imperialis 200 

10. Teinopalpus aureus 204 

11. Eurytides (Protesilaus) marcellinus 206 

12. Eurytides (Protesilaus) lysithousharrisianus 208 

13. Eurytides (Eurytides) iphitas 211 

14. Graphium (Pathysa) epaminondas 213 

15. Graphium (Pathysa) idaeoides 215 

16. Graphium (Pathysa) megaera 217 

17. Graphium (Arisbe) aurivilliusi 219 

18. Graphium (Arisbe) weberi 220 

19. Graphium (Arisbe) levassori 222 



Contents 



20. Graphium (Graphium) procles 

21 . Graphium (Graphium) meeki 

22. Graphium (Graphium) stresemanni 
Ti. Graphium (Graphium) sandawanum 

24. Graphium (Graphium) mendana 
Troidini 

25. Battus zetides 

26. Parides ascanius 

27. Parides hahneli 

28. Parides pizarro 

29 . Parides stein bachi 

30. Parides coelus 

31. Parides klagesi 

32. Parides burchellanus 

33. Atrophaneura (Atrophaneura) luchti 

34. Atrophaneura (Losaria) palu 

35. Atrophaneura (Pachliopta) jophon 

36. Atrophaneura (Pachliopta) schadenbergi 

37. Atrophaneura (Pachliopta) atropos 

38. Troides (Troides) andromache 

39. Troides (Troides) prattorum 

40. Troides (Troides) dohertyi 

41. Ornithoptera tithonus 

42. Ornithoptera rothschildi 

43. Ornithoptera chimaera 

44. Ornithoptera paradisea 

45 . Ornithoptera meridionalis 

46. Ornithoptera alexandrae 
Al . Ornithoptera aesacus 

48. Ornithoptera Croesus 
Papilionini 

49. Papilio (Pterourus) homerus 

50. Papilio (Heraclides) esperanza 

5 1 . Papilio (Heraclides) aristodemus ponceanus 

52. Papilio (Heraclides) caiguanabus 

53. Papilio (Heraclides) aristor 

54. Papilio (Heraclides) garleppi 

55 . Papilio (Heraclides) himeros 

56. Papilio (Heraclides) maroni 

57. Papilio (Chilasa) maraho 

58. Papilio (Chilasa) osmana 

59. Papilio (Chilasa) carolinensis 

60. Papilio (Chilasa) toboroi 

61 . Papilio (Chilasa) moerneri 

62. Papilio (Papilio) hospiton 

63. Papilio (Princeps) benguetanus 

64. Papilio (Princeps) acheron 

65 . Papilio (Princeps) jordani 

66. Papilio (Princeps) weymeri 

67. Papilio (Princeps) sjoestedti 

68. Papilio (Princeps) manlius 



224 
226 
228 
231 
234 

236 
240 
242 
244 
246 
248 
250 
252 
254 
256 
258 
261 
263 
266 
269 
271 
273 
276 
278 
281 
284 
288 
292 
294 

297 
299 
301 
305 
307 
309 
311 
313 
315 
317 
320 
322 
324 
326 
329 
331 
333 
335 
337 
339 



VI 



Contents 

69. Papilio (Princeps) phorbanta 342 

70. Papilio (Princeps) aristophontes 345 

71. Papilio (Princeps) desmonditeita 2>A1 

72. Papilio (Princeps) antimachus 351 

73. Papilio (Princeps) morondavana 354 

74. Papilio (Princeps) grosesmithi 356 

75. Papilio (Princeps) leucotaenia 358 

76. Papilio (Princeps) mangoura 362 

77. Papilio (Princeps) chikae 364 

78. Papilio (Princeps) neumoegeni 367 



Appendices 



A . List of Papilionidae in threatened categories 369 

B. Papilionidae that require further monitoring and research 371 
Index 375 

Plates 



vu 



1 



How to use this book 



Introduction 

Work on 'Threatened Swallowtail Butterflies of the World' began in 1983. Informa- 
tion has been obtained from a very wide variety of sources including published and 
unpublished material, and from an extensive worldwide correspondence with 
amateur and professional lepidopterists, taxonomists, conservationists and 
government officials. We hope that all have been acknowledged at the end of this 
section. For those not mentioned, please accept our apologies for the omission and 
our thanks for your help. Whilst we have tried to make the book as up-to-date as 
possible and have sought advice worldwide, any mistakes and opinions are, of course, 
the responsibilities of the authors. 

Swallowtail butterflies comprise the family Papilionidae of the insect order 
Lepidoptera. They were chosen for this study because 1) they are relatively well 
known and are familiar to laymen and amateurs as well as professional entomologists, 
2) they are large, often spectacular, insects, relatively easy to see, identify, and 
therefore monitor, and 3) they have a worldwide distribution and have adapted to 
many habitats and foodplant families, particularly in the tropics. 

Layout and content of the book 

The contents of the book have been listed in the preceding pages. The main body of 
the book is divided into a number of sections, each of which will be described and 
explained briefly below. These notes explain the plan of the book, the inter-relation- 
ships of the sections, and the terms and categories utilized. 

The swallowtail butterflies: an introduction to the family and its conser- 
vation This section is a general introduction to the swallowtails. A 
description of their biology is followed by sections summarizing their distribution, 
classification, foodplants and origins. Finally the conservation of swallowtails is 
discussed, including current threats and efforts that are being made , or are needed, to 
combat those threats. 

Swallowtails of the world: their nomenclature, distribution and conservation 
status This is a comprehensive list of the whole family in taxonomic order. 
Each species has been numbered sequentially to permit easy cross-reference with the 
geographical index (see below). Published synonyms have been included where they 
may ease confusion, but no attempt has been made to list all synonyms. All names 
mentioned are indexed at the back of the book. The complete known distribution of 
each species is given, along with notes on taxonomy, conservation status, threats and 
a list of references. Threatened subspecies have been noted in the Hst but, with three 

1 



Threatened Swallowtail Butterflies of the World 

particularly important exceptions, not given a full review. Common names are given 
where known, but no new names have been suggested. To find the species occurring 
in a particular country or geographical region, consult the geographical index to 
species (section 3, page 124). Under each country, or in certain cases each regional 
heading, there is a list of numbers corresponding to the species in the annotated 
taxonomic list which occur in that country or region. 

The annotated taxonomic list was also used as 1) the basis for a critical fauna 
analysis to indicate those countries which have faunas of particular interest 
(section 4, p. 137), 2) the basis for an analysis of trade in swallowtail butterflies 
(section 5, p. 155) and 3) to permit a preliminary assessment of threats to swallowtails 
and the choice of a list for full review (section 6, p. 181). The status of species chosen 
for review in this final section is indicated in the taxonomic list and the name with 
status is also printed in full under the appropriate country or regional heading in the 
geographical index. 

Analysis of critical faunas This is a computer-assisted analysis of the 
distribution of the swallowtails, which has enabled us to pinpoint those faunas that 
are critical to conservation effort. The analysis is based on endemic species within 
political boundaries. It is emphasized that this is a preliminary and coarse-grained 
analysis. It may be used for guidance in international efforts for swallowtail 
conservation but in no way affects national or local considerations. 

Trade in swallowtail butterflies This section describes the extensive 
worldwide trade in butterflies with particular reference to the swallowtails. The main 
kinds of trade are outlined and the areas concerned are pinpointed. An assessment is 
made of the impact on wild populations, with attention given to the various ranching 
projects and their potential as a form of conservation. Appended to the text is a 
complete list of the family Papilionidae showing those species advertised for sale 
during the last five years and the range of prices demanded by dealers. Notes on 
legislation and captive breeding are also included in this table. 

Reviews of threatened species This section contains reviews of the 78 taxa 
which were chosen from the taxonomic list as being threatened or in need of special 
conservation attention. Three important subspecies have been included because they 
are of particular interest for legislative or other reasons. The choice of species for 
review is always problematical and for swallowtails it has been made more difficult by 
the dearth of information for many species (see Appendix B). 

Each review follows a similar format to that of previous lUCN Red Data Books. 
The following sections and headings are used: 

Name and taxonomic position The name comprises generic (and some- 
times subgeneric), specific (and sometimes subspecific) names, authority for the 
description of the taxon and the date when the description was published. The family 
Papilionidae is divided between subfamilies and tribes which are also indicated. 

Red Data Book (RDB) status categories The traditional lUCN RDB status 
categories have been used throughout this volume. It is emphasized that the 
application of these categories to threatened organisms of any kind poses many 
problems and inevitably involves subjective judgements. They should always be 



How to use this book 

regarded as a working tool, subject to change at any time as new information is 
received. The categories are defined as follows: 

Extinct (Ex) 

Species not definitely located in the wild during the past 50 years (criterion as used in 
the Convention on International Trade in Endangered Species of Wild Fauna and 
Flora— CITES). 

Endangered (E) 

Taxa in danger of extinction and whose survival is unlikoly if the causal factors 
continue operating. Included are taxa whose numbers have been reduced to a critical 
level or whose habitats have been so drastically reduced that they are deemed to be in 
immediate danger of extinction. Also included are taxa that are possibly already 
extinct but have definitely been seen in the wild in the past 50 years. 

Vulnerable (V) 

Taxa believed likely to move into the Endangered category in the near future if the 
causal factors continue operating. Included are taxa of which most or all of the 
populations are decreasing because of over-exploitation, extensive destruction of 
habitat or other environmental disturbance; taxa with populations that have been 
seriously depleted and whose ultimate security has not yet been assured; and taxa 
with populations which are still abundant but are under threat from severe adverse 
factors throughout their range. 

Rare (R) 

Taxa with small world populations that are not at present Endangered or 
Vulnerable, but are at risk. These taxa are usually localized within restricted 
geographical areas or habitats or are thinly scattered over a more extensive range. 

Indeterminate (I) 

Taxa known to be Endangered, Vulnerable, or Rare but where there is not enough 

information to say which of the three categories is appropriate. 

Insufficiently Known (K) 

Taxa that are suspected but not definitely known to belong to any of the above 

categories, because of lack of information. 

Out of Danger (O) 

Taxa formerly included in one of the above categories, but which are now considered 
relatively secure because effective conservation measures have been taken or the 
previous threat to their survival has been removed. 

N.B. In practice. Endangered and Vulnerable categories may include, tempora- 
rily, taxa whose populations are beginning to recover as a result of remedial action, 
but whose recovery is insufficient to justify their transfer to another category. 

Threatened is a general term to denote species that are Endangered, Vulnerable, 
Rare , or Indeterminate , and should not be confused with the use of the same word by 
the U.S. Office of Endangered Species. 

The information needed to fulfil the criteria for these categories is often lacking for 
swallowtails, even though they are among the best known of all insects. There is 
generally little difficulty in assessment of the threats to well-known species, 
particularly when they have restricted distributions, but the status of species whose 
habitat requirements are poorly understood is more difficult to assess, particularly 
where their distribution is wide. 



Threatened Swallowtail Butterflies of the World 

The category Insufficiently Known could have been very widely applied in this 
volume, but such action was deemed to be of little conservation purpose. Fourteen 
examples of species reviews in this category are given in section 6, but it will be seen 
that the quality of data is often poor and the review may add relatively little to what 
has already been stated in the species list (section 3). Appendix A (p. 369) is a list of 
those 78 species assigned to RDB categories, followed in Appendix B by a further list 
of almost 100 species for which more data on distribution, ecology and conservation 
status are required before an assignment of RDB category can be made. 

Description As the colour and size of most species of swallowtails will be 
unfamiliar, a brief description is included. In the case of Endangered, most 
Vulnerable and a selection of other interesting threatened species this description is 
supported by a colour plate of a museum specimen. The size of the species is in most 
cases indicated by the length of the forewing. The wings are the main distinguishing 
feature of butterflies and the terms used in describing them are explained in Figure 
1.1. 

Distribution Distributional data are the most recent available and are 
generally only given to the level of region, state or province. Precise localities are 
occasionally mentioned. 

Habitat and Ecology Notes include data on life-cycles and foodplants 
(which are all too often unknown), flight times and seasonal cycles. In some cases the 
biotope is described in a general way, including data on climate, vegetation and 
altitude where appropriate. 

Threats This section includes any known or surmised threats to the survival 
of the species or its populations. In some cases the threats are well defined, in which 
case a higher conservation category may be expected. Often the threats will not be 
known and a species may be known from very few specimens. In such cases holding 
categories such as Indeterminate or Insufficiently Known may be expected. Most of 
the listed species are threatened by loss of habitat. In many reviews where this is so, a 
general account of habitat loss and its causes may be given. Clearly such problems 
affect a wide range of wildlife. 

Conservation Measures Herein are included measures already taken to 
conserve the species, including protection by legislation or gazetting of protected 
areas. Proposed measures are also listed, including legislation, appeals for protective 
measures and further scientific study to establish population sizes, distribution and 
status. 

Index The general index includes all scientific and common names used in 
the book and a selection of key words. 

Plates Finally, colour plates of all but four Endangered and Vulnerable 
swallowtails are presented. A number of species in other threatened categories are 
also included, mainly to demonstrate the range of form to be found within the 
swallowtail family. All photographs have been reduced to a standard size because in 
life many of the species are larger than the page upon which they are illustrated. Their 
actual size is clearly indicated in the legend and in the appropriate review in section 6. 



How to use this book 



Figure 1.1. Explanation of terms used in the description of butterfly wings. In 
general the wings of the two sexes are similar in form but the males of some genera 
(e.g. Parides) have the anal margin of the hindwing folded over to contain secondary 
sexual organs. This fold is variously called the abdominal fold, hair-pouch or 
scent-organ. 




FW 




HW 




UFW: 
UHW: 


upper forewing 
upper hindwing 


LFW: 
LHW: 


lower forewing 
lower hindwing 


1. 
2. 
1&2. 


submarginal 

postdiscal 

distal 


8. 
9. 
10. 


outer margin 

anal or inner margin 

tail 


3. 


discal 


11. 


cell 


4. 

5. 
6. 


basal 
apex 
anal angle or tornus 


12. 
13. 
14. 


apical 

subapical 

subcostal 


7. 


costa or costal margin 


15. 


median 



UFW 




The following terms apply particularly to Ornithoptera males 



16. 


radial band 


17. 


cubital band 


18. 


anal band 



Threatened Swallowtail Butterflies of the World 

Acknowledgements 

We are very grateful to the many institutions and people who have given generous 
assistance in the preparation of this volume and without whom the task could not 
have been attempted. We would particularly like to thank the following: 

Members of the lUCN/SSC Butterfly Specialist Group. 

The staff of the British Museum (Natural History), in particular the Entomology 
librarian Miss P. Gilbert and the staff of the Rhopalocera section of the Entomology 
Department, particularly R.I. Vane-Wright, P.R. Ackery and R.L. Smiles. 

The staff of the Royal Entomological Society of London and particularly the librarian 
Mrs B. Leonard. 

Sally Anne Hughes for the cover drawing. 

All our colleagues at the lUCN Conservation Monitoring Centre and the Institute of 
Terrestrial Ecology, Furzebrook, who gave enthusiastic encouragement and helpful 
advice on many matters. 

Michael Morton of Brunei University and David Webb of Hatfield Polytechnic, 
sandwich students who gave invaluable assistance in data-processing during their 
time at the Centre. 

And the following individuals for providing information and refereeing various 
sections of the book: 



B. d'Abrera 
M.J.C. Barnes 
K.S. Brown Jr. 
M.P. Clifton 
G.A. Dacasin 
M.J. Dourojeanni 
P. Feeny 
A.K. Ghosh 
N. Gunatilleke 
J.D. Holloway 
J.N. Jumalon 
G. Lamas 
D. Lowe 
J.de la Maza E. 
T. Milliken 
A.U. Oboite 
D.F. Owen 
M.en C.H. Perez 
T. Racheli 
R.L. Smiles 
T.W. Turner 
L. Vazquez G. 
D. Wankhar 
M.L. Winston 



P.R. Ackery 

A. Bedford-Russell 

J. P. de Carvalho 

M.J. Coe 

E.W. Diehl 

H. Dove 

T.L. Fenner 

S.K. Ghosh 

D.L. Hancock 

W.Hsu 

J. Kielland 

T.B. Larsen 

R. Macfarlane 

O.H.H. Mielke 

J. Nabhitabahata 

S.N. Okiwelu 

M.J. Parsons 

A. Pinratana 

J. Riley 

D. Spencer-Smith 

A. Tye 

P. Viette 

P.G. Waterman 

A. Youdeowei 



S.A. Ae 
M.J. Braga 
P.B. Clark 
S. Collins 
P. DoUinger 
K. Eltringham 
R.W. Garrison 
H.C.J. Godfray 
J. Haugum 
A.F. Hutton 
J. Kingdon 
R.R.B. Leakey 
D.K. Mandall 
J.S. Miller 
T.R. New 
P. A. Opler 
R. Paulian 
R.M. Pyle 
A.H.B. Rydon 
T.T. Struhsaker 
the late H.Tyler 
M.V. Walker 
J.D. Weintraub 



H.S. Barlow 

F.M. Brown 

Sir C.A. Clarke 

T.C.E. Congdon 

F.M. Dori 

R.D. Estes 

M. Ghilarov 

R. Guilbot 

T. Hidaka 

M. Ishii 

Kim Hon Kyu 

R.E. Lewis 

A.G. Marshall 

L.D. Miller 

D. N'Sosso 

G.W. Otis 

N.D. Penny 

CD. Quickelberge 

T. Saigusa 

B. Turlin 

R.I. Vane- Wright 

B.M. Wankhar 

J. G. Williams 



This book was prepared and published with the support of the World Wildlife Fund, 
the United Nations Environment Stamp Conservation Fund, the United Nations 
Environment Programme, Exxon Corporation and Citibank. 



6 



2 



The swallowtail butterflies: 
an introduction to the family 
and its conservation 



The butterfly family Papilionidae includes some of the most spectacular and 
magnificent of all insects. When compared with other insects, all the swallowtails 
have large wings, ranging from the dainty, 50 mm wingspan of the dragontails of 
India and Malaysia {Lamproptera species), to the giant birdwings {Trogonoptera, 
Troides, Ornithoptera) of Indonesia and New Guinea. The female Queen Alexan- 
dra's Birdwing {Ornithoptera alexandrae) is the largest butterfly in the world, 
attaining a wingspan of over 250 mm. Because of their large wings many swallowtails 
are powerful fliers, the apollos being an exception with their rather feeble flight. The 
colours and patterns of swallowtails are astonishingly rich and diverse. They range 
from sombre black or brown (e.g. Papilio acheron, Troides dohertyi, Plate 3) to the 
lustrous sheen of the gloss swallowtails (e.g. Papilio chikae, Plate 8). Some of the 
birdwings are iridescent and highly changeable in colour; Troides prattorum can 
appear golden-orange, green, pale blue or pink from different angles of view (11). 
In many species the hindwings are extended into tails, superficially resembling the 
long tails of swallows, hence the popular name for these butterflies. The precise 
function of the tails has never been examined and it is not clear whether they assist 
flight or have some other purpose. The term swallowtail is used very loosely at both 
the familial, generic and specific level. Other common names include apollos 
{Parnassius spp), festoons (Allancastria and Zerynthia), gorgons (Meandrnsa), kites 
(Eurytides), dragontails (Lamproptera) swordtails or jays (Graphium), windmills or 
clubtails (Atrophaneura) , birdwings (Trogonoptera, Troides, Ornithoptera) and 
fluted swallowtails (Papilio). Common names are given in the species list (section 3, 
p. 33) and the general index (p. 375). 



Biology of swallowtails 

The adult butterfly is only the last of four distinct phases in the life cycle. The early 
stages comprise the egg, larva or caterpillar, and chrysalis or pupa. The process of 
development, or metamorphosis, takes several weeks unless the species over-winters 
in an immature phase. For example, Papilio machaon eggs hatch after about two 
weeks, the caterpillars feed for 30 days and may then pupate for as little as 14 days or 
may last through the winter. Adult Papilionidae may survive for up to four months, 
but 20-30 days is more usual. Predation and parasitism of butterflies are high at all 
stages of development and only a small proportion will survive to adulthood (47) . The 
eggs of many species, from the primitive Baronia brevicornis of Mexico to the Troides 



Threatened Swallowtail Butterflies of the World 

and Ornithoptera of New Guinea, are heavily attacked by minute parasitic (or, more 
correctly, parasitoid, as they kill their host) chalcid wasps (Hymenoptera: Chalci- 
doidea) (48, 49). Studies on a range of butterflies, including the British race of Papilio 
machaon in the Norfolk Broads, have indicated high predation of the first two larval 
instars by arthropods, particularly spiders (16). Later instars may be too big to be 
taken by arthropods but suffer heavy mortality due to birds. Pupae are severely 
predated by small mammals (16). 

The eggs of swallowtails are spherical, lack any sculpturing and in the birdwings 
may be up to 4 mm in diameter, the largest butterfly eggs of all. Most species can lay 
several hundred eggs, but predators usually prevent such a potential from being 
realized. The genus Ornithoptera is again unusual in generally laying no more than 
30 eggs per brood. Eggs are usually laid on the caterpillar's foodplant, often on the 
underside of leaves where they are inconspicuous and shaded. After hatching, 
caterpillars immediately begin feeding and growing, passing through five moults 
before the quiescent pupal stage. 

The caterpillars of Papilionidae may have fleshy spines or tubercles but they are 
never hairy and always have a curious structure called an osmeterium. This is a forked 
scent gland which the larva can extrude through a slit in the thorax when it is disturbed 
or attacked. The gland secretes a powerful-smelling liquid which consists, in the 
species studied so far, of two aliphatic acids (isobutyric and 2-methyl butyric acids) 
(22,23). When attacked, the larva tries to wipe the gland against its attacker, a 
procedure that is certainly successful against ants (22) and may deter not only other 
predators, but also parasites (47). Mature caterpillars are green in Baronia and the 
Leptocircini, black spotted with red or yellow in the Parnassiinae and Troidini, and 
variable in the Papilionini. Some caterpillars, like those of Papilio polymnestor, 
present a curious 'eyed' appearance and may adopt a threat posture as a deterrent to 
would-be predators (57). Others, such as Papilio clytia, mimic the appearance of 
distasteful species in the caterpillar as well as the adult (13). Species that feed on 
Aristolochiaceae and other poisonous plants are all believed to store the poison and 
themselves become poisonous. This fact may be advertized in a gaudy range of 
patterns and hues that acts as a reminder to predators, particularly birds. Many 
swallowtail larvae change their patterns between instars. Papilio machaon, for 
example, resembles a bird dropping in its first two instars, but thereafter becomes 
conspicuous in a bright pattern of black, orange and green (35). It is fairly 
commonplace for the final instar larva to leave the foodplant before pupating. For 
example, the caterpillar of Ornithoptera paradisea leaves its foodplant, the Aristolo- 
chia vine, and pupates about 1 m above the ground on shrubs and trees (11). 

The pupae of Papilionidae are generally brown, sometimes green, with prominent 
keels or tubercles in the Troidini and a single dorsal protuberance in the Leptocircini 
(30). A characteristic of the Papilionidae is that the fully grown larva spins a silk tail 
pad and girdle prior to pupation. The chrysalis is thus suspended at an angle from its 
substrate, often resembling a broken twig. An exception is the genus Parnassius, in 
which the caterpillar spins a fine silken web and pupates among leaves near the 
ground. Further details of the early stages of Papilionidae are given in Igarashi's 
recent study (34). 

Adult swallowtails have a number of distinctive characteristics, apart from the tails 
already described. In the mouthparts the palps are small, the proboscis well- 
developed. Adults frequently take nectar from flowers whilst still in flight, the 
fluttering stance being quite distinctive. All six legs are fully developed and fit for 
walking (unlike the Nymphalidae), and have a spur-like projection on the tibia of 
each foreleg. The cell is closed on both the fore- and hindwings and only a single anal 



Swallowtail introduction 

vein is present on the hindwing (except in Baronia). In the males of some genera (e.g. 
Parides, Atrophaneura) the anal margin of the hindwing is folded over and contains 
scent brushes used in courtship and mating. 

In the majority of swallowtails the sexes are broadly similar in appearance. 
Nevertheless, there are many species that are sexually dimorphic, often, but not 
invariably, as a result of mimicry. The males and females of birdwings are very 
different (Plates 3,4,5) but are not known to be mimetic except in the case of the 
female Omithoptera croesus lydius (11) (Plate 6). Females of other swallowtails are 
often slightly larger and less colourful than males, as in the birdwings, but again this is 
not always so. For example, the female Papilio antimachus is smaller than the male 
(Plate 8). Females are generally more retiring in their habits and in forests females 
often keep to the tree-tops while males congregate at drinking spots or fly rapidly 
along rides and streams. Males may be highly aggressive, jostling and fighting each 
other for a spot on a hilltop or defending their territory along a stretch of river. 

In some species, such as Graphium idaeoides (Plate 2), Papilio agestor and P. 
slateri, both sexes are mimics. In others, such as Papilio jordani and P. phorbanta 
(Plate 8), only the female is a mimic and extreme sexual dimorphism is apparent. A 
further variation is seen in Eury tides euryleon, a dimorphic species in which each sex 
appears to mimic the same sex of the dimorphic model, Parides iphidamas phalias 
(12). 

Mimicry is a form of protection against predators and is a widespread phenomenon 
amongst butterflies. In classical definitions it consists either of a palatable mimic 
species gaining protection from predators by resembling a distasteful and unpalatable 
model (Batesian mimicry), or of a number of distasteful models joining a mimetic 
ring in which each gains protection from the other (Miillerian mimicry). Recent 
research has shown this to be an over-simplification and many new variations and 
interactions are now recognized (1, 60). 

Table 2.1 lists a small selection of mimics and their models. Models are generally 
unpalatable species from the Danainae, Heliconiinae, Acraeinae or Aristolochia- 
feeding Papilionidae, but other groups may also be used. Papilio laglaizei is unusual 
in being a mimic of a day-flying moth, Alcidis agarthyrsus (Uraniidae). 

In numerous species the females (and less often the males, Papilio paradoxa is an 
example) are polymorphic, i.e. they occur in a number of different forms. The 
African Mocker swallowtail, Papilio dardanus, is yellow and tailed in the male but the 
female is extremely variable and may appear mainly white, yellow or orange, often 
tail-less. The females mimic a variety of distasteful model species (Table 2 A) (4). 

Distribution, classification and origin of swallowtails 

The Papilionidae is a pre-eminently tropical family (ca 20°N-20°S), although some 
species can reach latitudes 6O-70°N and 40-50°S. In an analysis of the distribution of 
Munroe's (42) list of 538 species, Scriber (53, after Slansky, 56) demonstrated the 
great increase in species richness at tropical latitudes. In Figs. 2.1-2.4 we have 
repeated Scriber's analysis using the 573 species in our list (section 3). As expected, 
the pattern is much the same , with the richest areas in the equatorial rain forest zones. 
South East Asia has the highest number of species but interestingly, species richness 
in the eastern Asian region between 20 and 40°N is also high, even exceeding the 
richness of the American and African tropics. This trend is reflected in the analysis of 
critical faunas (section 4, p. 137) where Indonesia, the Philippines and China alone 
are shown to include well over a third of the world's swallowtails. 
The classification of the Papilionidae has been a source of great difficulty to 



Threatened Swallowtail Butterflies of the World 



Table 2.1 A selection of possible models and mimics in the swallowtail family 
Papilionidae (1, 9, 12). Key: m = male, f = female. 



Mimics 



Models 



Indo-Australia 

Graphium idaeoides 
Papilio memnon f. 
Papilio memnon f. 
Papilio memnon f . 
Papilio polytes f . 
Papilio clytia 
Papilio agestor 
Papilio slateri 
Papilio paradoxa 
Papilio clytia clytia 
Papilio paradoxa aegialus 
Papilio jordani f . 
Papilio laglaizei 



Africa and its 
Papilio 
Papilio 
Papilio 
Papilio 
Papilio 
Papilio 
Papilio 
Papilio 
Papilio 



Islands 
phorbanta f. 
dardanus f. 
dardanus f. 
dardanus f. 
dardanus f. 
cynorta f. 
cynorta f. 
echerioides f. 



South America 

Eurytides thymbraeus aconophos 
Eurytides thymbraeus 
Eurytides branchus 
Eurytides belesis 
Eurytides euryleon 
Eurytides euryleon clusoculis 
Eurytides pausanias 



Idea leuconoe 
Atrophaneura varuna f. 
Atrophaneura nox f. 
Atrophaneura coon 
Atrophaneura aristolochiae 
Tirumala septentrionis 
Parantica sita 
Euploea algea 
Euploea mulciber m. 
Euploea klugii 
Euploea diocletianus 
Idea blanchardii 
Alcidis agarthyrsus 

Euploea goudotii 
Amauris albimaculatus 
Danaus chrysippus 
Amauris niavius 
Bematistes poggei m. 
Bematistes epaea f. 
Amauris niavius 
Amauris echerius 
Danaus formosa 

Parides alopius 
Parides photinus 
Parides areas mylotes 
Parides polyzelus 
Parides iphidamas phalias 
Parides lycimenes 
Heliconius wallacei 



taxonomists. The main problem has been that the wide diversity of colouring, life 
cycle and habits is not reflected in clear morphological differences. This has been 
particularly problematical at the generic level where great variations in colour and 
pattern often overlie an essentially similar structural design. In the genus Papilio for 
example, neither Munroe in 1961 (42) nor Hancock in 1983 (30, p. 30) could find 
morphological differences useful in creating generic groupings with a smaller 
number of species. Fortunately however, morphological, biological and chemical 
studies have met with success in developing our understanding of the taxonomy of 
the PapiHonidae at the suprageneric level (26, 42, 43). 

The living Papilionidae are divided into three subfamilies, the Baroniinae, 
Parnassiinae and Papilioninae. Each of these will be discussed only briefly here; the 
reader should refer to the literature for further details (see the bibliographies in 
30, 42) and to section 3 (p. 33) for information on particular species. 

The Baroniinae includes a single species, Baronia brevicornis from Mexico (see 



10 



Swallowtail introduction 

Fig. 2.1. Latitudinal gradients in the species richness of the New World Papilionidae 
(adapted from Scriber (53) and Slansky (56)). 



;^^i.<^Am 



^^ 



Latitude 
70 




J I I I I I l__l 

10 20 30 40 50 60 70 80 

Number of species 



Fig. 2.2. Latitudinal gradients in the species richness of the Eurasian and African 
Papilionidae (adapted from Scriber (53)). 




10 20 30 40 50 60 
Number of species 



11 



Threatened Swallowtail Butterflies of the World 

Fig. 2.3. Latitudinal gradients in the species richness of the Indo-Australian and 
Eastern Asian Papihonidae (adapted from Scriber (53)). 



r>~-^: 




10 20 30 40 50 60 70 80 90 100 110 120 130 

Number of species 



Fig. 2.4. 
world. 



Latitudinal gradients in the species richness of the Papilionidae of the 



Latitude 

70r 



40 



30 



10 



24 



62 



121 
127 



239 



20 



30 



50 



22 



-J I I I ' 



91 



-I 1 \ I I L. 



_l_ 



-I L. 



260 



-I 1 I I I I 



10 20 30 40 50 60 70 80 90 100 110 120 130 140 150 160 170 180 190 200 210 220 230240 250 260 

Number of species 



12 



Swallowtail introduction 

p. 182), considered to be the most primitive living papilionid. Although the adults 
have the fore-tibial spur and osmeterium which are characteristic of the Papilionidae, 
the wing venation is very uncharacteristic in having both anal veins present on the 
hindwing, like Pieridae and Nymphahdae. Baronia is therefore believed to be a 
descendant of primitive papilionid ancestors close to the evolutionary stem shared by 
the Papilionidae and Pieridae (43). Baronia is also most unusual in feeding on Acacia 
(Leguminosae). The position of Praepapilio Colorado, a fossil papilionid from the 
middle Eocene deposits of Colorado, U.S.A., is open to question. It shares some of 
the primitive features found in Baronia, but has recently been elevated to a 
monotypic subfamily of its own, the Praepapilioninae (17). 

The subfamily Parnassiinae is divided into two tribes, Parnassiini and Zerynthiini. 
The former includes the monotypic genera Archon and Hypermnestra and the 
holarctic and species-rich genus Parnassius. Archon and Hypermnestra are confined 
to Asia Minor while Parnassius species are primarily butterflies of the montane areas 
of Europe, northern and central Asia and western North America. Primitively the 
larval foodplant is the lowland vine Aristolochia, as in Archon , but a switch within the 
Aristolochiaceae to the perennial herb Asarum by Luehdorfia, to the family 
Zygophyllaceae by Hypermnestra and to the families Saxifragaceae, Fumariaceae, 
Crassulaceae and Dipsaceae by Parnassius has permitted a great expansion in alpine 
regions. Tectonic upheavals in central and southern Asia have encouraged extensive 
speciation by Parnassius; many species in the Asian mountains and plateaux remain 
very poorly known. The Zerynthiini have remained more conservative and retained 
Aristolochiaceae as larval foodplants. The tribe includes the genera Sericinus, 
Allancastria, Zerynthia, Bhutanitis and Luehdorfia, also found predominantly 
outside the tropics in southern Europe and Asia. Bhutanitis species such as B. 
lidderdalei, the Bhutan Glory, are highly prized and spectacular insects with a 
wingspan of over 10 cm and three tails to the scarlet-spotted hindwing. 

The third subfamily, Papilioninae, is by far the richest (90 per cent of all 
swallowtails) and is divided into three tribes, the Leptocircini, Troidini and 
Papilionini. The Leptocircini includes seven genera which fall into two groups, 
Iphiclides, Teinopalpus and Meandrusa (sometimes called the subtribe Teinopalpiti) 
and Eurytides, Protographium, Lamproptera and Graphium (sometimes called the 
Leptocirciti) (30). The first group is primarily Palaearctic, although Meandrusa 
extends to Indonesia. The larval food plants of Iphiclides and Teinopalpus are 
Rosaceae and Thymeleaceae respectively, but the foodplants of Meandrusa are 
unknown . The second group has a very wide distribution , believed to be the result of a 
spread through Gondwanaland from a North American/Laurasian ancestor (30). 
Eurytides from the Americas is a large genus of over 50 species using Annonaceae, 
Lauraceae and other Magnoliales as larval foodplants. Munroe's subgenera Protesi- 
laus and Eurytides (42) are retained here but were raised to generic level by Hancock 
(30). Protographium is a monotypic genus that feeds on Rauwenhoffta (Annonaceae) 
in Australia. The two species of Lamproptera fly in South East Asia and breed on 
Hernandiaceae and Combretaceae. Finally, the genus Graphium, with over 90 
species, occurs in Africa, Asia and Australia. Munroe (42) adopted a number of 
subgenera that are retained in essence here, but adapted to comply with more recent 
work by Hancock (30) and others. Larval foodplants are variable and include 
Lauraceae for Graphium (Pazala), Aquifoliaceae for Graphium (Pathysa), 
Annonaceae, Malpighiaceae, Apocyanaceae, Annonaceae and Anacardiaceae for 
Graphium (Arisbe), and Annonaceae, Apocyanaceae, Lauraceae, Magnoliaceae, 
Monimiaceae, Hernandiaceae and Piperaceae for Graphium (Graphium) (30). 

The second tribe in the Papilioninae, the Troidini, includes eight genera almost all 

13 



Threatened Swallowtail Butterflies of the World 

species of which feed on Aristolochiaceae. Battus is primitive and occurs in the 
Americas. In South America Euryades is a genus of temperate grasslands while 
Parides is found in tropical forests. Cressida is a monotypic genus from Australia, 
closely related to Euryades. Atrophaneura is mainly eastern Palaearctic and 
Indo-Australian, but with the extraordinary relict species Atrophaneura antenor in 
Madagascar. Finally, the three birdwing genera are Trogonoptera from the forests of 
Sundaland and Palawan, and Troides and Ornithoptera from Indo- Australia. 

The third tribe, Papilionini, includes over 200 species in the single genus Papilio, 
divided into a series of numbered sections by Munroe (42) and into several smaller 
genera by Hancock (30), Miller and Brown (39) and others referred to in those works. 
Here we adopt a conservative nomenclature of subgeneric divisions based on 
Hancock's phylogenetic studies (30) but Munroe's nomenclature (42). The larval 
foodplants of Papilio are extremely variable. The American and Caribbean subgenus 
Papilio (Pterourus) feeds primitively on Rutaceae, but has spread to include at least 
11 other plant families, notably the Lauraceae, Magnoliaceae and Piperaceae (30). 
Papilio (Heraclides), also from the Americas and Caribbean, uses mainly Rutaceae 
and Piperaceae. Papilio (Eleppone) is a monotypic subgenus from Australia that 
feeds on Rutaceae. Papilio (Chilasa) species often mimic distasteful milkweed 
butterflies (Danainae) and they are distributed in eastern and South East Asia, 
mainly on Lauraceae or Magnoliaceae. Papilio (Papilio) includes only the 14 species 
in the machaon group. Some of these butterflies are very familiar to entomologists in 
the northern hemisphere; machaon itself has a vast range stretching from Great 
Britain to Saudi Arabia, North America and Japan. The foodplants are 
Umbelliferae, Compositae or Rutaceae. The final subgenus, Papilio (Princeps), is a 
vast assemblage of over 130 species that feed almost entirely on Rutaceae and inhabit 
Africa, Europe, Asia and Australia. They have evolved a stunning variety of form 
and pattern, from the iridescent blue of Papilio zalmoxis and the giant wings of 
Papilio antimachus in Africa, to the delightful hues of the Gloss or Peacock 
Swallowtails {paris group) and the realistic mimics found in the memnon group of 
Asia. 

In this volume we have adopted a conservative and utilitarian approach to 
nomenclature simply to serve the best interests of our readership. The message of our 
book is conservation rather than taxonomy and our aim is to help the many 
politicians, legislators, scientists, conservancies and individual campaigners whose 
task it is to protect and preserve wild fauna and flora. Those specialist entomologists 
who have a deeper understanding of the inter-relationships and evolution of the 
swallowtails will surely not be misled by such an approach. Readers interested in the 
arguments for and against the alteration of long-established systems of nomenclature 
and classification will not be disappointed by the literature, in which conflicting views 
are regulariy aired (19, 20, 30, 38, 39, 42). 

The present patterns of diversity of the three Papilionidae subfamilies may not 
necessarily reflect their centres of origin. Although Shields and Dvorak (55) 
proposed a Neotropical or Indo-Australian origin for butterflies (including the 
Papilionidae), Hancock has recently argued convincingly against this (30). He 
concludes that in the case of the Papilionidae it is inappropriate to use diversity as an 
indication of centre of origin. The South East Asian fauna contains elements derived 
from the Palaearctic, Australian, Oriental and Afrotropical regions. No doubt it 
owes its richness to speciation patterns on the continental shelves and island arcs that 
comprise the present geography of the area. Similarly, speciation in the neotropics, 
which also has a fairly rich fauna, reflects the great diversity of habitats and perhaps 
the influence of Pleistocene forest refugia. Hancock contends that the centre of origin 

14 



Swallowtail introduction 

of the Papilionidae was Laurasia, one of the two Tertiary super-continents. 
According to the hypothesis, during the Cretaceous period Laurasia divided into 
three sections, each with a subfamily of swallowtails: Baroniinae in the western North 
America block, Papilioninae in the eastern North America-Europe block and 
Parnassiinae in the Asian block (30). Dispersal to Gondwanaland (which gave rise to 
South America, Africa, Australia, Madagascar, India and Antarctica) was 
supposedly facihtated via an island arc between North and South America. The 
theory is supported by the present distribution of the primitive genera from the three 
subfamilies, Baronia (Baroniinae) in Mexico (with the closely related fossil Praepapi- 
lio in Colorado), Archon and SericinuslBhutanitis (Parnassiinae) in western and 
eastern Asia respectively, Eurytides (Protesilaus), Papilio {Pteroiirus) and Battus in 
the Americas, with the most primitive groups in North and Central America (30). 
With Baronia and Praepapilio in the western North America block, this may be the 
most likely area for the origin of the Papilionidae. It must be admitted, however, that 
these ideas are still contentious and there is a need for further research. The fossil 
record for papilionid butterflies is poor and a few new finds could greatly alter the 
present interpretation of patterns. 

Swallowtails and man 

Swallowtails, and indeed all other butterflies, are of value to us in a number of 
different ways. Without doubt the most widely enjoyed aspect is the aesthetic one. 
With their fabulous colours and deHcate flight, the beauty of swallowtails can be 
appreciated by all ages and races of people. 

The enjoyment of swallowtails can take a number of forms, divided broadly 
between appreciation of the living animal and of the dead, dried and set specimen. 
The most natural and rewarding pleasure is to observe them in their native setting, 
flying across a forest glade or sipping water from the edge of a muddy pool. There is 
no need to know the scientific names and life history of swallowtails in order to 
appreciate the extra dimension of richness they can add to our environment. Most 
people around the world may not voice their enjoyment of butterflies, but experience 
in northern industrialized countries is showing that the response is more vociferous 
when the butterflies are suddenly no longer around. 

Swallowtails, being a largely tropical group, are generally not well known to the 
general public in countries with a temperate climate. However, there is a growing 
interest in "butterfly houses", where exotic species fly in large greenhouses amongst 
tropical vegetation. Visitors can walk through to watch, photograph and even handle 
the butterflies. Some of the more common birdwings and swallowtails are often well 
represented in such establishments. The stock is usually imported from the native 
country either as adults or as pupae which emerge in captivity. However, there is 
generally some research going on behind the public galleries and captive breeding is 
possible for an increasingly wide range of species. Established and responsible 
butterfly houses can do much to advance the cause of butterfly conservation, in terms 
of both scientific research and educating the public. So long as wild-caught material is 
harvested in a sustainable manner there will be no threat to the species exhibited. 

The appreciation and use of dead swallowtails and other butterflies is a more 
contentious and complex issue. It can take many forms, from the Papuan farmer who 
decorates his hair with a birdwing, to the Taiwan industrialists that mass produce 
table mats made of butterfly wings set in clear plastic. The collecting method also 
varies widely in approach, from the private collector seeking a specimen or two for 
study, to the disinterested local labourer who is paid by the specimen and employed 

15 



Threatened Swallowtail Butterflies of the World 

by dealers in cities remote from the habitat of the species. The moral issues are 
tortuous and perhaps best guided by a single consideration. Butterflies are living 
beings that fly, breathe and breed; they are not lifeless objects like stamps or 
match-books. They should not be taken merely to satisfy pointless acquisitiveness, 
but should only be collected with restraint and without unnecessary wastage. 
Examples of the types of trade in swallowtail butterflies are given in more detail in 
section 5 (p. 155). As will be seen, small-scale butterfly collecting generally does no 
appreciable harm to the population. The conservation of butterflies is much more 
seriously affected by habitat destruction (see below). Nevertheless, there is an 
unmistakable trend away from collecting and towards monitoring and photographing 
butterflies (51,62). This fashion towards appreciation of the living animal in its 
natural or semi-natural setting is greatly to be applauded and encouraged. 

Swallowtails and science 

Swallowtails have been used as a tool for biological research in a number of different 
fields. In the early 1940s E.B. Ford carried out novel research using chemical analysis 
as a taxonomic tool (26) . He found that pigments could be used to give insight into the 
phylogenetics of swallowtail butterflies. The presence of yellow anthoxanthins was 
believed to be a primitive character shared with certain Pieridae, supporting the 
hypothesis of a common evolutionary stem. The red pigments are of two types, A and 
B. Type A is found in all genera except Battus and Papilio, which have type B, not 
known anywhere else in the Lepidoptera. Type A is widely distributed in butterflies 
and moths and is taken to be the primitive red pigment of the Papilionidae. 

Biochemical analysis has also proved to be central to our understanding of the 
foodplant relationships of the Papilionidae and other phytophagous insects. 
Chemical similarity of larval foodplants is a widespread phenomenon that frequently 
cuts across botanical affinities (21). Correlations have been found between feeding 
habits and a number of natural products, including alkaloids, coumarins, essential 
oils, glucosinolates, acetylenic fatty acids and phenolic glycosides. The role of these 
chemicals is to act as physiological barriers to feeding by animals (27), but they have 
been variously overcome by butterfly larvae in the course of evolution (21). Once an 
animal species has evolved the capacity to metabolize or sequester the poison, any 
plant that uses that particular chemical deterrent becomes a potential foodplant. 

Feeny and his research team have drawn together the foodplant data for the 
Papilionidae and studied the ways in which the butterflies detect the correct plant 
(25). It is now known that host-plant recognition by swallowtails is the result of a 
complex series of tests, including visual searching, contact chemoreception and 
possibly olfaction of plant defence chemicals. It has been demonstrated that the 
families of the order MagnoHales (subclass Magnoliidae, including the Annonaceae, 
Lauraceae and Magnoliaceae) are linked with the Aristolochiaceae and Rutaceae by 
their content of benzylisoquinoline alkaloids. Hydroxycoumarins and furanocouma- 
rins are likewise associated with larval foodplant families, particularly those used by 
Papilio. However, the most consistent chemical link of all is provided by the essential 
oils, which are reported from virtually all families of swallowtail food plants (25). 
Essential oils are volatile organic mixtures, usually containing terpenes and esters, 
that have the odour or flavour of the wide variety of plant families in which they are 
found. The structure of the oil-producing cells in the Rutaceae are similar to those in 
the Aristolochiaceae, the Piperaceae and the MagnoHales families mentioned above. 
Larvae that normally feed on either Rutaceae or Umbelliferae have been shown to be 

16 



Swallowtail introduction 

attracted to the essential oils of both families, explaining the numerous observations 
of atypical egg-laying behaviour (25). 

Research in this field is continually giving rise to fresh challenges and insights that 
are applicable to a wide range of pure and applied biological sciences. In particular, 
there is a growing emphasis on the search for natural plant compounds with 
insect-repellant qualities. The hope is that they will be as effective as the man-made 
chemicals now generally used for plant protection, but less environmentally 
hazardous. A well-known example of a natural product already widely used to 
combat insects is pyrethrum, extracted from the plant Tanacetum cinerariifolium 
(Compositae). 

As has already been mentioned, many species of swallowtails are polymorphic in 
the female, most notably the African Mocker Swallowtail, Papilio dardanus. C.A. 
Clarke and P.M. Sheppard of Liverpool University chose first this species and later 
the South East Asian P. memnon for their examination of the genetic basis of 
mimicry, a study that continued for 30 years and produced a result of unexpected and 
exceptional importance. The story is a complicated one, but will be explained fully in 
order to demonstrate the way in which a seemingly unrelated piece of research may 
finally have a great impact on human well-being. 

The work began with the discovery that Papilio species could be hand-paired quite 
easily, by opening the male's claspers with a fingernail and introducing the male to a 
female. Clarke and Sheppard then set about obtaining specimens of P. dardanus from 
many parts of Africa and specimens of P. memnon from the Far East. By controlled 
cross-breeding of the mimetic and non-mimetic forms in each species they found that 
the genetic coding for the mimetic form was not contained in a single gene . This is best 
explained with an example. Papilio memnon form achates is a mimic of Atrophaneura 
coon, while form agenor is non-mimetic. Breeding experiments showed that mimetic 
achates is dominant to agenor and the various features of the mimic all appear 
together. Such features include yellow body, presence of tails and a small white 
'window' on the hindwing. Very rarely, however, the features were found to be split 
up and although , for example , tails were present and the wing window small , the body 
was black. This suggested that more than one gene locus was involved and that 
random crossing-over of the chromosomes may occasionally split up the several 
closely Hnked loci of what is now known to be a mimicry 'supergene'. Furthermore, 
Clarke and Sheppard found that breeding of hybrids between races produced a great 
increase in phenotypic variability. This indicated that each race had interacting and 
co-adapted gene complexes which insured a low degree of variability in the 
phenotypic expression of the major genes. These interactions were markedly 
disturbed in the race hybrids (6). 

As a result of studies on swallowtail races in which two forms are common but 
intermediates caused by genetic rearrangements are also occasionally found, the 
attention of the scientists was directed to the human Rhesus blood group system, 
where crossovers probably account for the rare genotypes and where there is a 
marked interaction between the Rhesus (Rh) and ABO systems (5). Here, as in the 
mimetic butterflies, the various Rhesus phenotypes are controlled by a series of 
closely linked genes in a supergene. The most important interaction is that ABO 
incompatibility between a mother and her baby often confers protection against 
Rhesus immunization of the mother (5). 

Rhesus immunization is a problem caused by the movement of red blood cells from 
an Rh-positive baby into the blood system of its Rh-negative mother. This generally 
happens at or near to the time of delivery. The mother subsequently develops an 

17 



Threatened Swallowtail Butterflies of the World 

immune response to Rh-positive blood cells and although the first child is safe, having 
already been born, subsequent children may suffer severe rejection problems in the 
womb. The baby may be killed before birth, or die shortly afterwards. Complete 
blood transfusion immediately after birth of the so-called "yellow baby" may save the 
child's life. 

Clearly the protection against Rh-immunization conferred by ABO incompatibi- 
lity of mother and child is greatly to the advantage of second , third and later children . 
The reason for this protection is not clear but Clarke suggested that any Rhesus- 
positive foetal cells which have crossed into the maternal circulation are destroyed by 
the mother's naturally occurring anti-A or anti-B before they have had time to 
stimulate the production of anti-Rh antibodies (5). This protection against Rhesus 
immunization is highly desirable , but it only occurs in the 20 per cent of cases in which 
the foetus is ABO incompatible with the mother. After a great deal of research it 
proved to be possible to imitate the protection by giving suitable antibodies to 
Rh-negative mothers in the 80 per cent of cases where mother and baby are 
compatible on the ABO systems. As a result, nearly all cases of maternal Rh 
immunization can now be prevented (5). The success of the research programme is 
due, in large part, to the usefulness of certain swallowtail butterflies for scientific 
study. 

Threats to swallowtails 

Four main factors threaten swallowtails, as well as other terrestrial forms of wildlife: 
habitat destruction, pollution (which might be considered as a special form of habitat 
destruction), the introduction of exotic species and commercial exploitation. In 
general these threats are either the direct result of increasing human population 
pressure or else are enhanced by it (7). It is now internationally recognized that 
conservation efforts can only reach long-term fruition in partnership with policies for 
population planning and control (32). A full consideration of the global problem of 
human populations outstripping their resources, and of the political, economic and 
social causes, are beyond the scope of this book and other publications should be 
consulted for more detailed studies (3, 24, 33). 

Each of the four types of threat will be briefly reviewed , with examples drawn from 
the swallowtails. 

Habitat alteration and destruction As demonstrated in the companion 
lUCN Red Data Books for plants, mammals, amphibians, reptiles and invertebrates, 
the primary threat to wild species throughout the world is the destruction and 
alteration of the biotopes and habitats in which they live (29, 37, 58, 61). All species 
depend for their survival upon the integrated network of physical and biological 
factors that make up their environment. Most are able to withstand a small degree of 
disturbance and manipulation, but human activities have steadily modified wilder- 
ness areas into man-made landscapes of settlement, agriculture and industry that are 
altered to an extent that precludes coexistence with many wild creatures. 

Habitat alteration has a universal impact on swallowtails and other wildlife, but the 
effects are more noticeable in biomes and ecosystems that are restricted in area. 
Hence the extensive destruction and alteration of African savannas has certainly 
caused great reductions in distribution patterns, but no swallowtail species are 
regarded as threatened there. Even in the more restricted rain forests of the Zaire 
basin only Papilio antimachus is recognized as threatened (category Rare) , and this 
species is still widespread. A similar situation prevails in the massive forest block of 

18 



Swallowtail introduction 

the Amazon basin. Conversely, islands are particularly prone to extinctions because 
they tend to have rich endemic faunas in very restricted areas. Mauritius for 
example, has suffered the extinction of many species of animals and plants, largely 
because of deforestation followed by extensive cultivation of sugar cane. The 
endemic swallowtail Papilio manlius has been fortunate in successfully adapting to 
Citrus for its foodplant, but Libythea cinyras (Libytheinae) is now believed to be 
extinct there and Cyclyrius mandersi (Lycaenidae) may be seriously threatened (14). 
Similarly, Papilio phorbanta has adapted to Citrus on Reunion, but the subspecies 
nana from the Seychelles has not been seen since 1890. A number of endemic taxa in 
Taiwan are under threat, including Papilio maraho (Vulnerable) and Troides aeacus 
kaguya. In Jamaica Eury tides marcellinus is now extremely rare and populations of 
the Homerus Swallowtail {Papilio homerus) have been split up and endangered as a 
result of coffee production and timber extraction. Graphium epaminondas may be 
under similar pressures in the Andaman Islands, as may Graphium levassori on 
Grande Comore. Many other examples of island-living swallowtails at risk will be 
found in section 6, including amongst others the birdwings of South East Asia, 
Papilio hospiton from Corsica and Sardinia and Papilio neumoegeni from Sumba. 

Montane species suffer many of the same problems as those on islands. In the 
tropics there are many swallowtails restricted to a habitat that may have been more 
extensive during the cooler climate of the last Ice Age, but is now found only on 
mountains. In Sabah and Sarawak Papilio acheron and particularly Graphium 
procles have restricted ranges in montane regions. Mt Kinabalu, the only place 
where G. procles flies, is a protected area, but economic considerations have taken 
high priority there and the region has suffered incursions by mining companies and 
recreational developments. In Indonesia the little known species Graphium 
stresemanni is possibly confined to the Manusela range in Seram and Atrophaneura 
luchti has a very restricted montane range in eastern Java. Graphium sandawanum is 
found only on Mt Apo in the Philippine island of Mindanao, and both Papilio chikae 
and P. benguetanus are relict species in the Cordilleras of Luzon. 

In Africa the effect of the last Ice Age on vegetation patterns is still contentious 
(63), but whatever the historical causes, there is no question that the string of relict 
forests running from Mt Kulal in northern Kenya to Mt Mulanje in Malawi now all 
have their own endemic subspecies of butterflies, if not species, and all are under 
threat from deforestation for agriculture and plantation forestry (8). Of particular 
note is the now highly Endangered Taita Blue-banded Swallowtail (Papilio 
desmondi teita) from the Taita Hills in southern Kenya. Other Kenyan montane 
papilios of interest include Papilio dardanus flavicornis from Mt Kulal and P. d. 
ochracea from Mt Marsabit and Mt Nyiru. In Tanzania Papilio sjoestedti flies only in 
montane forests in the regions around Mt Kilimanjaro, Mt Meru and the 
Ngorongoro range, much of which is fortunately protected inside national parks. 
Many more butterflies from East Africa's montane forests could be listed here, 
particularly from the nymphalid genus Charaxes. 

The forms of habitat alteration of most importance to swallowtail butterflies are 
1) deforestation, 2) agricultural conversion and intensification, 3) alteration of 
pastures and 4) industrialization and urbanization. 

Deforestation As a result of the ecology and distribution of swallowtail 
butterflies, forestry practices in temperate latitudes have little impact on their 
populations. An exception is the Japanese Luehdorfia japonica, which is discussed 
below. 

In 1981, following an extensive survey of 76 tropical countries, the Food and 

19 



Threatened Swallowtail Butterflies of the World 

Agriculture Organization (FAO) and the United Nations Environment Programme 
(UNEP) jointly published an assessment of the global tropical forest resources 
(24, 28). These documents have supplied an extremely valuable framework within 
which to discuss the conflicts between conservation and exploitation of forest 
resources, conflicts that must be resolved if sustainable economic growth is to be 
achieved (33). When combined with data on protected areas and the distribution 
patterns of animals, plants and, in the present case, swallowtail butterflies, it becomes 
possible not only to assess current conservation problems, but also to predict which 
regions and species are likely to be of particular concern in the future. 

Tropical forests in the broadest sense are estimated to cover 2970 million hectares, 
i.e. 20 per cent of the land surface of the globe (28). The main forest formations are 
closed broadleaved forest (including rain and monsoon forests, 40 per cent), open 
forest types (including the cerrados of South America and the wooded savannas of 
Africa, 25 per cent), coniferous forests (1 per cent) and shrubland (21 per cent) (28). 
These ecosystems include by far the most important swallowtail habitats, as is clearly 
demonstrated by the distribution patterns in Figs 2.1-2.4. The majority of swallow- 
tail species use larval foodplants that are vines, shrubs or trees, although there are 
important exceptions in the Parnassiinae (see 1.3 below). Exploitation of tropical 
forests by man will inevitably disturb swallowtail populations, but the extent of the 
impact depends on the level of disturbance , the extent of the biotope and its ability to 
recover, and the willingness of people to allow it to do so. 

Tropical forests may be altered in two ways: 1) through complete deforestation 
followed by non-forestry usage such as agriculture, or 2) by selective felling for 
industrial use (mainly rain and monsoon forests) or fuel production (mainly drier 
forest formations). However, it is usual for logged over forests eventually to become 
completely deforested so that for all practical purposes deforestation becomes the 
major factor altering tropical forests (28). 

The most important single cause of deforestation is shifting cultivation , accounting 
for 70 per cent , 50 per cent and 35 per cent of the total areas that have been deforested 
in Africa, Asia and the Americas respectively (24, 28). Thus the division of threats 
between deforestation and agricultural conversion in this discussion is to some extent 
artificial. Some of the conservation problems associated with shifting cultivation will 
be found in section 1.2 below, but might equally well have been discussed here under 
deforestation. Other significant causes of deforestation are clearance to create 
grazing lands (particularly in South and Central America) and settlement of 
agriculturalists along new logging roads with subsequent expansion into the forest 
(particularly in Africa and Asia). 

Logging can damage forests to varying extents . In South East Asia for example , the 
'volume actually commercialized' (VAC), an index of production from the point of 
view of commercial exploitation, ranges from 15 cu. m/hainBurmato90cu. m/hain 
Sabah, i.e. from less than 10 per cent to more than 30 per cent of the corresponding 
standing volume of timber (24). This may mean that whereas Burmese swallowtails 
like the Kaiser-I-Hind {Teinopalpus imperialis) could survive logging, native 
birdwing species in Sabah would be unable to withstand the rapid degradation and 
complete loss of habitat which can result from such intensive logging rates. Another 
variable is the extent to which logging roads are subsequently used as access roads to 
new farms by land-hungry people. This problem is most apparent in Africa and Asia 
where 70 per cent of the deforestation occurs in areas of closed forests that have been 
previously logged. In the Americas the comparable figure is only 44 per cent because 
most logs are transported along rivers or existing roads (28). 

In the drier regions of the tropics, particularly of Africa and Asia, the accelerating 

20 



Swallowtail introduction 

demand for fuelwood and charcoal is causing severe deforestation of open woodland. 
The technology is available to meet these demands through plantation forestry (36) 
and Brazil produces 29 per cent of its fuelwood needs from Eucalyptus plantations 
(28). In Africa and Asia however, the low levels of investment in plantations to meet 
these basic demands of the rural poor must eventually have dire consequences in 
socio-economic and resource conservation terms. 

The global rate of clearance of tropical closed broadleaved forest is now believed to 
be lower than was previously feared. It has been estimated that by the end of the 
century 88 per cent of the present world cover of closed tropical broadleaved forests 
will still remain intact (24, 28). However, this is no cause for complacency, it simply 
means that there is a little more time in which to implement sound conservation and 
management strategies. A large proportion of those remaining forests will be 
disturbed to some degree. 

There are considerable differences between the rates at which countries are 
utilizing and altering their closed forest resources. Whereas Zaire and Brazil will still 
have extensive forest resources in the year 2000, the forests of Peninsular Malaysia, 
Ivory Coast, Nigeria, Sri Lanka and Java will be virtually gone. If present trends 
continue, nine countries will have used practically all their closed broadleaved forests 
within 30 years, and a further 13 will join them within 55 years (28). All 22 of these 
countries between them only account for 11 per cent of the world's tropical closed 
forest, but some, such as Jamaica and Sri Lanka have important swallowtail faunas. 

What impact is tropical deforestation having on swallowtail butterflies? Con- 
sidering in the first place the loss of tropical closed forests, the Asian sector is of 
greatest concern . About one quarter of the world's closed tropical forest area is found 
in Asia, 61 per cent of this being in the islands of South East Asia including New 
Guinea. As will be demonstrated in section 4 (p. 137), this is the world's richest area 
for swallowtail butterflies. Indonesia has 121 species of swallowtails, over 20 per cent 
of the world total. It has also emerged as the world's most important producer and 
exporter of tropical hardwoods (24, vol. 3). Rain forest species from the South East 
Asian islands that have been given threatened status include Graphium idaeoides, G. 
megaera, G. sandawanum, Papilio osmana and P. carolinensis in the Philippines, 
Graphium stresemanni, Ornithoptera aesacus and O. croesus in the Moluccas, O. 
rothschildi, O. chimaera, O. paradisea and particularly the Endangered Queen 
Alexandra's Birdwing, O. alexandrae, in New Guinea and Graphium meeki, G. 
mendana and Papilio toboroi in Bougainville and the Solomon Islands. In continental 
Asia the threats from deforestation are less serious because most swallowtails are 
more widespread in their distribution. However, deforestation throughout its range, 
but particularly in India and Nepal, is placing the Kaiser-I-Hind, Teinopalpus 
imperialism at risk. Numerous species are locally at risk in Peninsular Malaysia, 
including Troides helena, T. aeacus, T. amphrysus, T. cuneifer, Papilio agestor, P. 
mahadeva, P. palinurus, Graphium empedovana, Meandrusa payeni, Lamproptera 
meges and L. curius (2). 

African rain forests are still poorly studied and the two Insufficiently Known 
species, Graphium weberi and G. aurivilliusi from the main Cameroon-Congolese 
block , require further research . There are few endemics in the much-depleted forests 
of the western block, but Graphium maesseni is only known from the Ghanaian type 
series. The Giant African Swallowtail, Papilio antimachus, is now very scarce in this 
western part of its range but although it is confined to primary forests, its future may 
be assured in the forests of the Zaire basin. 

In the South American rain forests the swallowtail fauna also remains poorly 
known. There are few seriously threatened species but the narrow ranges of Parides 

21 



Threatened Swallowtail Butterflies of the World 

hahneli, P. pizarro, P. steinbachi, P. coelus, P. klagesi, P. burchellanus and Papilio 
garleppi are cause for concern . Many other poorly known rain forest species are listed 
in Appendix B. 

Turning to the loss of tropical open woodlands, a number of threats to swallowtail 
butterflies can also be identified. Of the Asian fauna, Papilio jordani is a Rare species 
from possibly open formations of semi-deciduous forest in northern Sulawesi and 
Troides dohertyi is Vulnerable in the nearby Talaud and Sangihe Islands. Two 
Vulnerable species, Papilio neumoegeni from Sumba and Atrophaneura schaden- 
bergi from Luzon, probably fly mainly in open woodlands. Eury tides is a genus of kite 
swallowtails that fly in the open woodlands of the Neotropics. Both the Yellow Kite, 
Eurytides iphitas, and Harris' Mimic Swallowtail, E. lysithous harrisianus, are in 
serious decline in Brazil, and the Jamaican Kite, Eurytides marcellinus is now a great 
rarity in Jamaica. As already noted, there are few threatened species in the extensive 
savanna formations on the African mainland. A number of threatened island species 
have already been mentioned; two others that are confined to open woodland, 
Graphium levassori from the Comores and Graphium pelopidas (possibly a subspe- 
cies of G. leonidas) from Pemba Island and Zanzibar, are at risk. 

Agricultural conversion and intensification The spread of subsistence 
farming has been cited as the biggest single cause of forest clearance , soil degradation 
and loss of wildlife in the world today (18). Landless labourers, sharecroppers and 
marginal farmers now form a majority in rural areas of Latin America and Asia, and 
in Africa the recent and rapid rise in human population levels is imposing severe 
pressure on land unsuitable for settlement (18). Approximately 3.8 million hectares 
of open woodland are being cleared every year, mainly as agriculture extends into 
marginal lands (24). Much of Africa is too infertile and dry for crops or even grazing, 
but farmers are moving further into semi-arid areas in their search for land and food. 
Pastoralists are being squeezed into desert margins, causing overstocking and 
overgrazing which in turn lead to degradation of soils and an increasing proportion of 
plants unpalatable to domestic animals. Inevitably the wildlife suffers too and land in 
national parks throughout Africa, and to a lesser extent other continents, is under 
pressure for grazing and development. In Kenya for example, where the population 
growth rate of about 4 per cent is one of the highest in the world , sections of the Masai 
Mara, Tsavo and Lake Nakuru reserves have already been excised. 

Many examples of threats to swallowtails from deforestation followed by agricul- 
ture have already been cited. A few more that specifically concern agricultural 
pressure may also be mentioned. In the central African countries of Rwanda and 
Burundi agricultural land is so overcrowded that pressures on the few relict forests 
are intense. Encroachment into these forests is depleting forest wildlife such as the 
Vulnerable Cream-banded Swallowtail, Papilio leucotaenia. Papilio morondavana, 
P. grosesmithi and P. mangoura, all endemic to Madagascar, are threatened by 
uncontrolled burning, the spread of shifting agriculture and the failure to sustain 
adequate fallow periods. In Sri Lanka the few remaining forests are under severe 
pressure from agriculturalists and logging companies. The Vulnerable clubtail 
Atrophaneura jophon has its last stronghold in the island's Sinharaja Forest Reserve, 
until recently severely encroached by agriculturalists and fuel-gatherers but now 
under full legal protection. 

In temperate latitudes where agricultural conversion is already extensive, 
emphasis in recent decades has been on intensification of farming practices. Many 
traditional forms of land use are compatible with butterflies, e.g. coppicing and 
moderate grazing, but the demand for higher food production and living standards 

22 



Swallowtail introduction 

has resulted in much more intensive and exhaustive exploitation of the fertile areas. 
In Great Britain, the U.S.A. and many other parts of the northern hemisphere, this 
has caused severe reductions in butterfly populations (41,52,61). Factors detri- 
mental to virtually all forms of wildlife are involved in these processes of 
intensification, including use of insecticides and herbicides, destruction of 
hedgerows, drainage, short fallow periods and heavy use of fertiHzers. With the very 
Hmited swallowtail fauna in the north temperate regions, the family has suffered less 
than many other forms of wildlife. Nevertheless, drainage has severely reduced the 
range of the British Swallowtail {Papilio machaon britannicus) (16), and in Japan 
Luehdorfia japonica is becoming more scarce as its previously lightly managed open 
woodland haunts are destroyed in favour of agriculture or intensive forestry. 

In tropical regions problems associated with intensification of agriculture on the 
main continental blocks are of much less significance to conservation than the 
extension of agriculture into virgin lands. However, in eastern Asia human 
population levels are such that intensive agriculture is already very widespread and 
incompatible with wildlife, notably in the subtropical parts of China, parts of 
Indochina and Thailand, Peninsular Malaysia and Java. In Brazil drainage and 
development of coastal swamps and thickets is threatening the Vulnerable species 
Parides ascanius. Its mimic Eury tides lysithous harrisianus is now seriously Endan- 
gered and has been proposed for Brazil's protected list, an action which will only be 
successful if suitable habitat for the mimic and its model can be set aside and 
maintained. 

Alteration of pastures Permanent grasslands are important habitat for 
many butterflies, particularly in the families Lycaenidae and Hesperiidae, but they 
are of less significance to the Papilionidae. The swallowtail foodplants are generally 
shrubs, vines or trees and most species fly in woodlands or forests. An important 
exception is the north temperate subfamily Parnassiinae, many of which use 
herbaceous species that grow in open situations. Luehdorfia feeds on the aristolo- 
chiaceous herb Asarum, which grows in open woodland, and the foodplant families 
used by Parnassius are all herbs of alpine meadows. In the mountains of the Hindu 
Kush, the Himalaya and western China there are very many species of Parnassius 
that are poorly known and may be at risk from pasture degradation (see appropriate 
countries in section 3 and the discussion of Chinese Parnassius on p. 151). Some are 
in such inaccessible regions that their future may be assured, but others, such as 
Parnassius autocrator, are certainly coming under new pressures. This particular 
species lives in the Hindu Kush of Afghanistan and the Pamir mountains of 
Tadzhikskaya S.S.R. In certain parts of the southern Himalaya, particularly in 
Nepal, there are reports of heavy tourist foot traffic resulting in pollution, trampling, 
erosion and degradation of vegetation through firewood collecting. Such impacts are 
not in the best interests of either the wildlife or the tourist industry. 

Also Endangered is Papilio hospiton from Corsica and Sardinia, primarily as a 
result of alteration and destruction of pastures. The umbelliferous foodplants are 
poisonous to sheep and are destroyed by fires started by local farmers. Pressures 
from recreational developments are reducing the butterfly's range, particularly on 
Corsica. 

Industrialization and urbanization Since the Papilionidae is a mainly 
tropical group, there are relatively few examples of conservation problems resulting 
from these factors. However, the range of Papilio himeros in south-eastern Brazil is 
declining as a result of development in Rio de Janeiro and coastal regions in general. 

23 



Threatened Swallowtail Butterflies of the World 

Schaus' Swallowtail (Papilio aristodemus ponceanus) from the Florida Keys is 
Endangered because of demand for building land and the consequent destruction of 
the hardwood hammocks in which the butterflies live. 

Pollution The effects of atmospheric pollution on butterflies are rather 
poorly understood. An assessment of Europe's threatened butterflies has reported 
that a number of widespread species have suffered severe declines in Fennoscandia, 
northern Germany, Poland, Czechoslovakia, Netherlands, Switzerland, Austria, 
Hungary and the Italian Alps (31). All these areas are to the north and east of the 
principal industrial zones in western Europe from which atmospheric pollution is 
carried on the prevailing south-westerly winds. The decline of Parnassius apollo in 
Norway has, for example, been attributed to 'acid rain' - a weakly acidic solution of 
sulphur and nitrogen oxides in rainwater. It seems likely that acid rain has deleterious 
effects on a wide variety of butterflies and possibly also their foodplants, as well as 
other invertebrates and wildlife. Ozone has been implicated as a particularly toxic 
component of acid rain but further research is needed to demonstrate its effects. In 
Europe there are widespread demands to instigate and increase international efforts 
to improve anti-pollution measures. 

Pesticides are not generally hazardous to butterflies when applied in the recom- 
mended manner directly to the target species. However, there is a significant risk in 
the effects of wind drift, particularly with the growing trend towards aerial 
application of pesticide to rice, wheat and other extensive monocultures. When 
spraying is carried out in a cross-wind pesticide may drift into adjacent natural 
habitats. Although no serious conservation problems have been reported directly 
from this cause, there is still a need for vigilance in the responsible use of pesticides 
(40). 

While in the industrialized countries of the northern hemisphere pollution control 
is showing valuable results, there is little evidence of similar efforts in the developing 
world. Here, serious pollution may result from forest and savanna fires (10, 54, 64) as 
well as industrial effluent and incorrect or excessive use of pesticides. Serious threats 
to all forms of wildlife may be expected from such sources in the future. 

Exotic introductions Introduction of exotic animals and plants inevitably 
upsets the balance of natural communities to some extent. There are no documented 
cases of severe effects on swallowtail butterflies but the spread of Papilio demodocus 
(Orange Dog) from the African mainland to Madagascar, Mauritius and Reunion is a 
matter for concern. Apart from being a minor pest of Citrus, P. demodocus is an 
aggressive species that is capable of ousting less vigorous native species. Research is 
needed concerning the impact P. demodocus has had on the Indeterminate Mauritius 
endemic P. manlius during the three-quarters of a century since demodocus was 
introduced. No effects on the closely related Madagascar endemics Papilio moronda- 
vana (Vulnerable) and P. grosesmithi (Rare) have been noted, nor on the Vulnerable 
Reunion endemic Papilio phorbanta. 

Commercial exploitation The level of commercial exploitation of swallow- 
tails and other butterflies has never been estimated globally. With butterfly trade in 
Taiwan worth about US $20-30 million per year, a speculative figure in the order of 
$100 million per year worldwide may not be unrealistic. As has been noted in the 
above discussion of swallowtails and man, exploitation can be anything from the 
capture, sale or barter of a single specimen for the sake of interest or decoration, to 
large-scale commercial ventures employing hundreds of local collectors. Inverte- 

24 



Swallowtail introduction 

brates can frequently, but not invariably, withstand a considerable level of harvest 
because of their high reproductive capacities. However, heavy exploitation can have 
serious effects under three circumstances: 1) if the population is already critically 
depleted by other factors like habitat destruction, e.g. Papilio hospiton may be 
seriously at risk in Corsica and Sardinia, 2) if the population is small and has a high 
value per individual, e.g. the highly prized species Papilio chikae from Luzon, 3) if 
the species has a low reproductive rate and low juvenile recruitment, e.g. the 
Ornithoptera birdwings of Indonesia, New Guinea and the Solomon Islands. 

The need for self-restraint in the collecting of butterflies and other insects was first 
formally recognized by the Royal Entomological Society of London, which set up a 
Protection Committee in 1923. In 1968, the Society replaced this with the Joint 
Committee for the Conservation of British Insects, which includes representatives 
from all parts of Britain and all the major entomological societies. In 1972 the 
Committee published "A Code for Insect Collecting" which has been reproduced 
and adopted not only in Britain, but also in many parts of Europe and throughout the 
world. The Xerces Society, a U.S. organisation devoted to conservation of insects, 
responded with its own policies in 1975, and the Lepidopterists' Society of America 
pubHshed guidelines in 1982. The only code adopted by commercial entomologists is 
the Entomological Suppliers Association of Great Britain "Code of Conservation 
Responsibility", published in 1974. In this document the trade in a number of British 
and exotic species was restricted to specimens already in circulation (31). 

For a full consideration of the value of rational, sustainable exploitation of 
swallowtails and of the dangers of unsustainable levels of commercialization refer to 
section 5, Trade in swallowtail butterflies (p. 154). 

Conservation of swallowtails 

Documentation and education The documentation of threats to wildlife is 
the first step in any effective conservation programme. This book is an attempt to 
fulfil that requirement for the swallowtails. It is the first worldwide assessment of the 
conservation needs of any invertebrate group and the first Red Data Book (RDB) to 
be based upon a published consideration of every species in the taxon under review, 
in this case the family Papilionidae. 

A number of national RDBs and official or unofficial lists have documented the 
threats to local Papilionidae. The majority of these are for European countries and 
consider only the very limited swallowtail fauna of that region (61 , p. 325). Such lists 
are referenced under individual species in section 3 (p. 33). Although the Papilioni- 
dae is one of the best-known of all insect groups, documentation of the conservation 
and biology of rare swallowtails is relatively sparse because of their mainly tropical 
distribution. Much of the work that has been done is referred to in the reviews of 
threatened species in section 6 (p. 181). 

The lUCN Invertebrate Red Data Book (61) included seven species of swallow- 
tails and a number of other butterflies, but drew attention to the fact that destruction 
of habitat is putting whole invertebrate communities at risk. The category 
Threatened Community, coined in the Invertebrate RDB, has not been utilized here 
because it was considered inappropriate and artificial to describe communities based 
on a single family of butterflies. Nevertheless, the principal remains vahd since 
swallowtails comprise a small, but significant and often highly visible, part of 
innumerable Threatened Communities worldwide. As described above, the most 
severe threat is destruction of entire biotopes and habitats, which destroys plants and 
animals indiscriminately. 

25 



Threatened Swallowtail Butterflies of the World 

The species list in section 3 shows that many swallowtails still remain very poorly 
known. This book is only a preliminary attempt to document what we do know and to 
draw attention to important areas for future research. Hopefully it may act as a 
catalyst for further studies on the biology and conservation of swallowtails. 

Protected areas The protection of swallowtail butterflies in wild and natural 
areas designated as national parks and reserves is a vital priority that is emphasized 
throughout the species reviews in section 6 of this book. Two main needs have 
become evident. 

Firstly, very few existing protected areas have been surveyed for any invertebrate 
groups. The swallowtails and other butterflies provide an opportunity for extending 
the traditional species lists of birds and mammals to include a spectacular and popular 
insect group that is of interest to an increasing number of tourists and visitors to 
protected areas. Once the local species are known, fine opportunities exist for 
tropical countries to emulate the success of the British butterfly houses in more 
natural surroundings. Habitat enrichment to attract butterflies after the fashion of 
the Papua New Guinea ranching programme (see below) could do much to conserve 
swallowtails at the same time as interpreting the value of swallowtails to the general 
public in a pleasantly assimilable fashion. 

Secondly, there are certain countries that have important swallowtail faunas but 
very limited allowances for protected areas. In section 4 of this book. Analysis of 
critical faunas, the distribution of the family Papilionidae is analyzed in detail, 
demonstrating that just five countries, Indonesia, the Philippines, China, Brazil and 
Madagascar, between them contain over half of the world's species of swallowtails . A 
further five countries, India, Mexico, Taiwan, Malaysia and Papua New Guinea, 
brings the total to more than two thirds (p. 141). All of these countries, and many 
others, could use their swallowtail faunas as an extra yardstick in judging the value of 
existing or proposed protected areas. An analysis of the Indonesian fauna, given in 
section 4, demonstrates how this could be done. 

Certain of the world's tropical forest areas are known to be under particularly 
severe threat of total deforestation (44), many of them with valuable endemic 
swallowtails already listed above. An exhaustive account cannot be given here, but a 
few illustrative examples may be cited. Because of economic and population 
pressures most of the Philippines' lowland forests may be gone by the end of the 
century. The national parks system is said to be undergoing review, but there is 
evidence of wholesale abuse of reserve boundaries by logging companies and 
agriculturalists (e.g. see review of Graphium sandawanum) . Similarly, the popula- 
tion profile and extent of logging in Indonesia indicate that the loss of huge areas of 
forest cover is also now inevitable. In recognition of the need for conservation 
planning as part of their development programme, the Indonesian government has 
prepared a thorough conservation plan (59). Malaysia is also drafting a state by state 
conservation strategy that should rationalise the parks system throughout that 
country, but there is a need for haste as the rate of logging in eastern Malaysia is 
growing rapidly, threatening to cause deforestation on the scale seen in Peninsular 
Malaysia before long (24). 

Other priority areas for the protection of swallowtails in reserves include western 
China, the southern foothills of the Himalayas, the western forests of India and the 
forests of Sri Lanka. In the Caribbean, Jamaica's system of reserves is in need of 
revision and there is opportunity for more effective measures in the Dominican 
Republic and possibly Cuba. Brazil's rain forests are still extensive despite increasing 
settlement in Para, Mato Grosso and Rondonia, but the Atlantic seaboard forests are 

26 



Swallowtail introduction 

poorly protected and include many highly endangered species. In Africa there is 
increasing attention being paid to the protection of the relict montane forests, and in 
the unique biotopes of Madagascar more has been done to strengthen the national 
park system in recent years. 

The Tropical Rain Forest and Primates Campaign, launched by the World Wildlife 
Fund in 1982, is currently promoting conservation measures in 14 countries. Many of 
the projects focus on the need to protect forested areas not only as a genetic resource 
for the future , but also as a valuable tool in watershed and soil protection (50) . World 
Wildlife Fund's Plants Campaign, launched in 1984, emphasizes the value of 
protected forests as a source of plants useful to mankind as crops and medicines. 

Where natural forests are already severely depleted, the value of plantation 
forestry as a conservation tool should not be underestimated. Plantations can provide 
timber for fuel, decoration and building, thus alleviating the pressures on the natural 
woodlands and forests that remain. They can also serve as buffer zones around 
natural forest refuges, as has been proposed for the Nyungwe forest in Rwanda (see 
review of Papilio leucotaenia) . However, the aims of resource conservation are 
largely defeated when natural forest areas are cleared to make way for plantations. In 
practically every part of the tropics there are adequate areas of already cleared and 
partially degraded land that could be focal points for afforestation and regeneration. 
International co-operation is needed in a global programme of conservation-orien- 
tated investment in plantation forestry. 

Legislation and international conventions International agreement on 
wildlife trade control is contained in the Convention on International Trade in 
Endangered Species of Wild Fauna and Flora (CITES), which controls and monitors 
import and export of listed species . A total of 87 countries have so far become party to 
CITES, and the European Community has adopted a Regulation (3626/82) to 
enforce the Convention uniformly throughout the Community. Appendix I, which 
contains no insects, is a list of species in which trade is subject to strict regulation and 
commercial trade is virtually prohibited. Appendix II lists species in which trade is 
regulated for the purpose of monitoring. The butterflies listed on Appendix II are the 
birdwings {Ornithoptera, Troides and Trogonoptera) and the Apollo butterfly 
(Parnassius apollo), all in the Papilionidae. These species may be commercially 
traded, but an export permit from the country of export is required before specimens 
may be removed from that country or imported into another state which is party to 
CITES . There are inconsistencies in the present listings of swallowtails on CITES and 
it is perhaps time for a reassessment of their past effectiveness and future value. A 
case could be made for delisting certain birdwings supplied by the ranching trade in 
Papua New Guinea, while possibly raising Queen Alexandra's Birdwing to Appendix 
I (48). Many other threatened butterflies are traded quite heavily and ideally could be 
monitored. However, it must be recognized that there are serious problems in the 
implementation of controls on insects that are difficult to identify (compared with 
many vertebrates) and very easily transportable. 

The European Community Regulation 3626/82 implementing CITES includes an 
Annex of species listed on Appendix II of CITES that the Community treats as 
though they were on Appendix I, i.e. preventing virtually all trade in those species. 
The EC has unexpectedly added the CITES Appendix II Papilionidae to this list, a 
move that was carried out without consultation with the CITES Secretariat or the 
lUCN/SSC Butterfly Specialist Group. The Regulation will severely jeopardize the 
Papua New Guinea birdwing ranching programme and may cause the demise of 
proposed ranching programmes in Indonesia and the Solomon Islands, all of which 

27 



Threatened Swallowtail Butterflies of the World 

rely heavily on European markets. As will be stated on a number of occasions 
throughout this book, butterfly ranching is seen as an important tool for conser- 
vation. International efforts are being made to have the Papilionidae removed from 
Annex C Part I in order to bring the EC Regulation more into line with the 
requirements of CITES.. 

National legislation to protect local swallowtails is now quite commonplace. A full 
consideration of these laws cannot be given here, but protected species are noted in 
section 3; section 5 includes extensive notes on legislation. General restrictions on 
trade in Lepidoptera apply in both East and West Germany (GDR and FRG), 
Kenya, Madagascar, Mexico and Turkey. One or more species of swallowtail are 
protected or banned in trade under national legislation in Austria, Czechoslovakia, 
Finland, France, Germany (GDR), United Kingdom, Greece, Hungary, Luxem- 
bourg, Netherlands, Poland, Switzerland, U.S.S.R., Brazil, India, Indonesia, 
Malaysia, Papua New Guinea and the U.S.A. 

In recognition of the importance of conserving the habitats of threatened animals 
and plants, all northern European countries give greater emphasis to protected areas 
than to legislation concerning individual species. No amount of protective legislation 
will succeed if a species' habitat is permitted to be destroyed. Nevertheless, 
legislation and international conventions do have the added advantage of drawing 
public attention to the plight of particularly rare or threatened species. The U.S. 
Endangered Species Act is probably one of the most effective forms of wildlife 
protection in the world, but is not without its problems. Taxa listed are required to be 
thoroughly studied and a recovery plan drawn up. If so recommended, the Act allows 
for designation of complete protection for habitats critical to the survival of 
threatened taxa, as well as for protection of the taxa themselves. An example of the 
working of the Act is described in the review of Schaus' Swallowtail (p. 301), a 
case-history which also demonstrates that problems still exist in the practical 
application of the Act. 

Management and research Research on the management of threatened 
insects began in Great Britain, where it is now the responsibility of the Institute of 
Terrestrial Ecology. The centres at Furzebrook and Monks Wood have been 
particularly active in management and conservation of rare insects, one of which, the 
British Swallowtail {Papilio machaon britannicus) , has been the subject of long-term 
research and is now one of the world's best-known papilionid taxa. 

The Swallowtail is a very widespread Holarctic species that in Britain has become 
specialized on a single foodplant found only in the fens (marshes) of East Anglia, 
Milk Parsley (Peucedanum palustre). At one time the Swallowtail occurred 
throughout the East Anglian fens and possibly in marshes along the River Thames 
and River Lea (16), but extensive drainage in the early 19th century destroyed most 
suitable habitat. The butterfly survived at Wicken Fen until the early 1950s when that 
population became extinct, leaving the species confined to marshes around the 
Norfolk Broads, notably Hickling Broad (16). In 1975, 228 artificially reared adults 
were released at Wicken Fen in an attempt at re-introduction and it was estimated 
that over 2000 individuals pupated that year, but by 1980 the population was once 
again extinct (15). The failure was attributed to a gradual lowering of water levels in 
Wicken Fen, which had a deleterious impact on the foodplants. The Swallowtail is 
one of about 15 target species in a new campaign of research and conservation of 
British butterflies currently being run by World Wildlife Fund (U.K.). 

In the U.S.A. Schaus' Swallowtail {Papilio aristodemus ponceanus) is now 

28 



Swallowtail introduction 

confined to the Florida Keys. It is listed as Threatened but has been proposed for 
re-Usting as Endangered under the Endangered Species Act. Schaus' Swallowtail has 
been the subj ect of intensive biological research but despite these efforts the species is 
still declining, mainly as a result of habitat destruction. For a full account of research 
on this swallowtail refer to section 6, p. 301. 

These attempts at swallowtail management have perhaps not met with the success 
that was hoped for, but important lessons have been learnt. It has been recognized 
that recovery programmes are expensive on resources and may be very risky. 
Long-term planning and adequate provision for protected areas are undoubtedly 
preferable. Nevertheless, as the threats to swallowtails and other butterflies become 
more intense, there will be a growing need for careful management studies, 
particularly in tropical regions. Research on the birdwing butterflies has demon- 
strated that an intimate knowledge of breeding biology and general ecology can pay 
dividends in terms of both conservation and rational exploitation (45, 48). Further 
carefully directed research could be of great benefit to the conservation of the family 
as a whole. 

In 1984 the Lepidoptera Specialist Group of the Species Survival Commission of 
lUCN was divided into a Butterfly SpeciaHst Group and a Moth Specialist Group. 
The Butterfly Group inherited certain long-standing priorities such as the conser- 
vation of the Monarch Butterfly over-wintering grounds in Mexico and the protection 
of Queen Alexandra's Birdwing in Papua New Guinea (61), but now has the 
opportunity to re-examine butterfly conservation problems, particularly those of the 
swallowtails. With their worldwide connections the Specialist Groups are of great 
value in centralizing conservation data and in advising on priorities for conservation 
attention. Attracting funds for conservation projects is more difficult for inverte- 
brates than vertebrates , but the recommendations of the Group should be recognized 
as being of international priority and importance. 

Ranching and farming of swallowtails The term farming' in this context 
means that young are reared in captivity from parents that are also held in captivity. 
'Ranched' swallowtails are captured as young stages of wild parentage and reared to 
the adult stage in captivity. Farming of swallowtails on a large scale is relatively rare 
but perfectly possible, particularly since the technique of hand-mating has been 
perfected for Papilio. As noted above, the hybrids used in the study of the genetics of 
mimicry were farmed, as were the Swallowtails that were released at Wicken Fen. 
Farming has also been used as a conservation measure for the Apollo, Parnassius 
apollo. Normally this species breeds one generation per year, over-wintering in the 
egg stage . By artificially rearing two generations per year, material for recolonization 
of depleted areas can be rapidly accumulated (46). 

The finest example of successful butterfly ranching is without question the 
development of the birdwing industry in Papua New Guinea. The programme is 
described in the trade review in section 5 so only a brief summary is needed here. In 
Papua New Guinea the Insect Farming and Trading Agency (I.F.T.A.) of the 
Department of Primary Industry sells high quality specimens of birdwings ranched 
locally, returning three quarters of all profits to the ranchers (45). The I.F.T.A. 
demonstrates ways to enrich the habitat of the birdwings by planting foodplants and 
nectar plants around the gardens of the ranchers, and provides basic equipment for 
rearing the pupae in cages, killing the adults and storing them safely for later setting 
and sale. Imperfect and unwanted specimens are returned to the wild in order to 
keep up the stock of individuals visiting the gardens. The main species ranched are 

29 



Threatened Swallowtail Butterflies of the World 

Troides oblongomaculatus and Ornithoptera priamus, but recent research into 
foodplant requirements and conservation status has suggested that O. goliath, O. 
victoriae and 0. chimaera could also be ranched and traded (48). The butterfly 
ranching project in Papua New Guinea has demonstrated that trade and conser- 
vation can be of mutual benefit. Careful biological studies of other large and 
spectacular swallowtails, with a view to replacing the trade in wild-caught specimens 
with ranched specimens, is greatly to be encouraged. Although a thorough analysis 
of the potential for butterfly ranches around the world has never been made, there 
are certainly innumerable opportunities throughout tropical Africa, South America 
and Asia (45). 

References 

1. Ackery, P.R. and Vane- Wright, R.I. (1984). Milkweed Butterflies: Their Cladistics and 
Biology. British Museum (Natural History), London, and Cornell University Press. 
420 pp. 

2. Barlow, H.S. (1983). Butterfly protection in Peninsular Malaysia. Manuscript /n//tt., 4 pp. 

3. Brandt Commission (1980). North-South: A Programme for Survival. Pan, London. 
304 pp. 

4. Carcasson, R.H. (1981). Collins Handguide to the Butterflies of Africa. Collins, London. 
109 pp. 

5. Clarke, C.A. (1967). Prevention of Rh-haemolytic disease. British Medical Journal ^•. 
7-12. 

6. Clarke, C.A. and Sheppard, P.M. (1963). Interactions between major genes and 
polygenes in the determination of the mimetic patterns of Papilio dardanus. Evolution 17: 
404-413. 

7. Collins, N.M. (1984). The impact of population pressure on conservation and develop- 
ment. Research in Reproduction 16: 1-2. 

8. Collins, N.M. and Clifton, M.P. (1984). Threatened wildlife in the Taita Hills. Swara 7(5): 
10-14. 

9. Corbet, A.S. and Pendlebury, H.M. (1978). The Butterflies of the Malay Peninsula. (Third 
edition, revised by J.N. Eliot). Malayan Nature Society, Kuala Lumpur. 578 pp. 

10. Crutzen, P.J., Heidt, L.E., Krasnec, J. P., Pollock, W.H. and Seller, W. (1979). Biomass 
burning as a source of atmospheric gases CO, H,, N2O, NO, CH3CI and COS. Nature 282: 
253-256. 

11. D'Abrera,B. (1975). Birdwing Butterflies of the World. Landsdowne Press, Melbourne. 
260 pp. 

12. D'Abrera, B. (1981). Butterflies of the Neotropical Region. Part 1. Papilionidae and 
Pieridae. Lansdowne Editions, Melbourne, in association with E.W. Classey, Faringdon. 
172 + xvi pp. 

13. D'Abrera, B. (1982). Butterflies of the Oriental Region. Parti. Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi -I- 244 pp. 

14. Davis, P.M.H. and Barnes, M.J.C. (in press). The butterflies of Mauritius. Journal of 
Research on the Lepidoptera. 

15. Dempster, J. P. and Hall, M.L. (1980). An attempt at re-establishing the swallowtail 
butterfly at Wicken Fen. Ecological Entomology 5: 327-334. 

16. Dempster, J. P., King, M.L. and Lakhani, K.H. (1976). The status of the swallowtail 
butterfly in Britain. Ecological Entomology 1: 71-84. 

17. Durden, C.J. and Rose, H. (1978). Butterflies from the Middle Eocene: the earliest 
occurrence of fossil Papilionidae (Lepidoptera). Pearce-Sellards Serial of the Texas 
Memorial Museum 29: 1-25. 

18. Eckholm, E. (1982). Down to Earth. Pluto Press, London. 238 pp. 

19. Ehrlich, P.R. and Murphy, D.D. (1981 (82)). Butterfly nomenclature: a critique. Journal 
of Research on the Lepidoptera 20:1-11. 

20. Ehrlich, P.R. and Murphy, D.D. (1981 (83)). Nomenclature, taxonomy and evolution. 
Journal of Research on the Lepidoptera 20:199-204. 

30 



Swallowtail introduction 

21. Ehrlich, P.R. and Raven, P.H. (1964). Butterflies and plants: A study in coevolution. 
Evolution 18: 586-608. 

22. Eisner, T. and Meinwald, Y.C. (1965). Defensive secretion of a caterpillar (Papilio). 
Science 150: 1733-1735. 

23. Eisner, T., Pliske, T.E., Ikeda, M., Owen, D.F., Vazquez, L., Perez, H., Franclemont, 
J.G. and Meinwald, J. (1970). Defense mechanisms of arthropods. XXVII. Osmeterial 
secretions of papilionid caterpillars {Baronia, Papilio, Eurytides). Annals of the Entomolo- 
gical Society of America 63: 914-915. 

24. FAO/UNEP (1981). Tropical Forest Resources Assessment Project. (3 volumes). FAO, 
Rome. 

25. Feeny, P., Rosenberg, L. and Carter, M. (1983). Chemical aspects of oviposition 
behaviour in butterflies. In: Herbivorous Insects: Host-seeking Behaviour and Mechan- 
isms. Ed. S. Ahmad. Pp. 27-76. 

26. Ford,E.B. (1944). Studies on the chemistry of pigments in the Lepidoptera, with reference 
to their bearing on systematics. 4. The classification of the Papilionidae. Transactions of the 
Royal Entomological Society 94: 201-223. 

27. Fraenkel, G.S. (1959). The raison d'etre of secondary plant substances. Science 129: 
1466-1470. 

28. Global Environment Monitoring System, UNEP (1982). The global assessment of tropical 
forest resources. GEMSPAC Information Series No. 3, 14 pp. 

29. Groombridge, B. and Wright, L. (1982). The lUCN Amphibia-Reptilia Red Data Book. 
Part 1. lUCN, Gland, xliii + 426 pp. 

30. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

31. Heath, J. (1981). Threatened Rhopalocera (Butterflies) in Europe. Nature and Environ- 
ment Series No. 23. Council of Europe, Strasbourg. 

32. lUCN Commission on Ecology (1984). Population and Natural Resources . Commission on 
Ecology Occasional Paper Number 3, 12 pp. 

33. lUCN/UNEP/WWF. (1980). World Conservation Strategy. lUCN, Gland. 

34. Igarashi, S. (1979). Papilionidae and Their Early Stages. Vol. 1 219 pp., Vol. 2 102 pp. of 
plates. Kodansha, Tokyo. (In Japanese). 

35. Larsen, T. (1984). Butterflies of Saudi Arabia and its Neighbours. Stacey International, 
London. 160 pp. 

36. Leakey, R.R.B. and Last, F.T. (in press). Deforestation in the tropics: how to mount a 
counter attack. Spectrum. Central Office of Information. 

37. Lucas, G. and Synge, H. (1978). The lUCN Plant Red Data Book. lUCN, Gland. 540 pp. 

38. Miller, L.D. and Brown, F.M. (1981 (83)). Butterfly taxonomy: a reply. Journal of 
Research on the Lepidoptera 20: 193-198. 

39. Miller, L.D. and Brown, F.M. (1981). A catalogue/checklist of the butterflies of America 
north of Mexico. Memoirs of the Lepidopterists' Society 2: vii + 280 pp. 

40. Moriarty,F. (1975). Pollutants and Animals. Chapter6: Where have all the butterflies gone? 
Pp. 81-97. Allen and Unwin, London. 

41. Morris, M.G. (1981). Conservation of butterflies in the United Kingdom. Beiheft 
Veroffentlichung Naturschutz Landschaftspflege Baden-WUrttemberg 21: 35-47. 

42. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supplement 17: 1-51. 

43. Munroe, E. and Ehrlich, P.R. (1960). Harmonization ofconcepts of higher classification of 
Papilionidae. Journal of the Lepidopterists' Society 14: 169-175. 

44. Myers, N. (1979). Conversion Rates in Tropical Moist Forests. Report to National 
Academy of Sciences. National Research Council, Washington D.C. 205 pp. 

45. National Research Council (1983). Butterfly Farming in Papua New Guinea. Managing 
Tropical Animal Resources Series. National Academy Press, Washington D.C. 30 pp. 

46. Nikusch, I. (1981). Die Zucht von Parnassius apollo Linnaeus mit jahrlich zwei 
Generationen als Moglichkeit zur Erhaltung bedrohter Populationen. Beiheft Veroffentli- 
chung Naturschutz Landschaftspflege Baden-Wiirttemberg 21: 175-176. 

47. Owen, D.F. (1971). Tropical Butterflies . Clarendon Press, Oxford. 214 pp. 

31 



Threatened Swallowtail Butterflies of the World 

Troides (Lepidoptera: Papilionidae) in Papua New Guinea. Final report to the Depart- 
ment of Primary Industry, Papua New Guinea. Ill pp. 

49. Perez R., H. (1971). Algunas consideraciones sobre la poblacion de Baronia brevicornis 
Salv. (Lepidoptera, Papilionidae, Baroniinae) en la region de Mezcala, Guerrero. Anales 
del Instituto de Biologia Universidad de Mexico 42, Ser. Zool. (1): 63-72. 

50 . Poore , D . ( 1 976) . Ecological Guidelines for Development in Tropical Rain Forests . lUCN , 
Gland. 39 pp. 

51. Pyle, R.M. (1984). The Audubon Society Handbook for Butterfly Watchers. Charles 
Scribner's Sons, New York. 274 pp. 

52. Pyle, R.M., Bentzien, M. and Opler, P. (1981). Insect conservation. Annual Review of 
Entomology 26: 233-258. 

53. Scriber, J.M. (1973). Latitudinal gradients in larval feeding specialization of the world 
Papilionidae (Lepidoptera). Psyche 80: 355-373. 

54. Seller, W., and Crutzen, P.J. (1980). Estimates of gross and net fluxes of carbon between 
the biosphere and the atmosphere from biomass burning. Climatic Change 2: 207-247. 

55. Shields, O. and Dvorak, S.K. (1979). Butterfly distribution and continental drift between 
the Americas, the Caribbean and Africa. Journal of Natural History 13: 221-250 

56. Slansky, F. (1972). Latitudinal gradients in species diversity of the New World swallowtail 
butterflies. Journal of Research on the Lepidoptera 11(4): 201-218. 

57. Smart, P. (1975). The Illustrated Encyclopedia of the Butterfly World. Hamlyn, London. 
275 pp. 

58. Thornback, J. and Jenkins, M. (1982). 77ie/C/CAf Mamma/ /?edDafflfioo*.Pflrt7.IUCN, 
Gland, xl + 516 pp. 

59. UNDP/FAO National Parks Development Project (1981/1982). National Conservation 
Plan for Indonesia. Vols 1-8. FAO, Bogor. 

60. Vane-Wright, R.I. (1976). A unified classification of mimetic resemblances. Biological 
Journal of the Linnean Society 8: 25-56. 

61. Wells, S.M., Pyle, R.M. and Collins, N.M. (1983). r/ie/{/CN/MvmeZ)rate/?eJZ)afaBoo^. 
lUCN, Cambridge and Gland. L + 632 pp. 

62. Whalley, P. (1980). Butterfly watching. Severn House, London. 

63. White, F. (1981). The history of the Afromontane archipelago and the scientific need for its 
conservation. African Journal of Ecology 19: 33-54. 

64. Woodwell, G.M., Hobbie, J.E., Houghton, R.A., Melillo, J.M., Moore, B., Peterson, 
B.J. and Shaver, G.R. (1983). Global deforestation: contribution to atmospheric carbon 
dioxide. Science 222: 1081-1086. 



32 



Swallowtails of the world 



3 



Swallowtails of the world: their 
nomenclature, distribution and 
conservation status 



This section is divided into two parts. The first is a complete list of swallowtail species 
arranged in taxonomic order. Details of distribution, conservation status and 
coinmon names are given. The second part is a geographical analysis, listing 
swallowtail species by region and country (p. 124). 

Swallowtails of the world: an annotated species list 

The arrangement of this list is a slightly amended version of Hancock's recent 
taxonomic re-assessment of the Papilionidae (109), which itself relies heavily on the 
earlier work of Munroe (188), Rothschild and Jordan (144, 145, 233, 234) and many 
others. These references give basic background information on all groups of species 
in the list that follows. Important regional works include those of Berger (16), 
Pennington (208), Van Son (273) and Carcasson (34) for Africa , D'Abrera for 
Australia (48), the Afrotropics (50), the Neotropics (51) and the Orient (52), Corbet 
and Pendlebury for Malaysia (45), Common and Waterhouse for Australia (40), 
Tsukada and Nishiyama (262) for the South East Asian islands, Rothschild and 
Jordan (234) and D' Almeida (55) for South America and Chou (35) for China. Many 
other valuable volumes are cited in the references, which follow the species list. 
Important systematic works include Ackery (1) and Eisner (70-74 and numerous 
others) for the Parnassiinae, Jordan (144, 145) for the Papilionidae and Haugum and 
Low (119) and D'Abrera (49) for the birdwings. Ford (88) gives a useful account of 
general systematics in the Papilionidae. Useful illustrations are given by Lewis (169) 
and Smart (250) , who also gives a species list . Authorities and dates of description are 
taken mainly from Bryk (28-31) and other authors, and have not necessarily been 
checked back to the original publication. 

The use of this list has been described in section 1 . It is essentially a vehicle for notes 
on the distribution and conservation of all swallowtail butterflies. Our main aim has 
been to produce a document of utilitarian value to conservation planners in wildlife 
and national park authorities, most of whom may not be professional entomologists. 
Common names have been given where known. These have been taken from dealers' 
lists and the literature. 

Occasionally it has been necessary to note conflicting opinions where species 
taxonomy and nomenclature may require further study or revision. However, we 
must emphasize that the Hst is not an account of primary taxonomic research. 

We have not attempted to examine the anatomy and morphology of the species 

33 



Threatened Swallowtail Butterflies of the World 

listed, such a work being beyond our brief and capacity. Instead we have been guided 
by an extensive literature search combined with correspondence from about 100 
expert lepidopterists worldwide. With 573 species listed, differences of opinion were 
inevitable and in such cases we have given all aspects of the argument in an attempt to 
present a balanced document. 

This section of the book will need to be constantly up-dated. All errors and 
omissions remain the responsibility of the authors, but in our attempts to reduce these 
to a minimum we would particularly like to acknowledge the advice and help of four 
people well-versed in the taxonomy and nomenclature of the swallowtails, David L. 
Hancock, Jan Haugum, Tomaso Racheli and Bernard Turlin. 



34 



Swallowtails of the world 

Order: LEPIDOPTERA (Butterflies and moths) 
Family: PAPILIONIDAE (Swallowtail butterflies) 

Subfamily: BARONIINAE 

Genus: Baronia Salvin 

1 Baronia brevicornis Salvin, 1893 
Rare — refer to section 6, p. 182. 

Restricted to a very small area of Mexico and particularly important 
/- jO because of its 'relict' nature as the sole member of the primitive subfamily 

/^ Baroniinae. Common name: Baronia. Refs: 10,51,62,69, 75, 128, 

129, 141, 189, 209, 210, 211, 212, 240, 269, 276, 277, 278, 290. 



Subfamily: PARNASSIINAE 
Tribe: Parnassiini 

The evidently sedentary nature of parnassiine populations and consequent 
description of many subspecies, particularly of European species, has 
tended to obscure the conservation status of species in this tribe (1,31, 141). 
We are grateful to B. Turlin for allowing us to quote from his unpublished 
list of Parnassius subspecies (265). 

Genus: Archon Hiibner 

2 Archon apollinus (Herbst, 1798) 
/ ^j-}^ A narrow range including Greece, Bulgaria, Romania, Turkey, Syria, Iran, 

Iraq, Lebanon, Jordan, Israel and the U.S.S.R. (Armenia and Turkmenis- 
tan). Fairly rare in collections, although this may be because it flies very 
early in the season. Not known to be threatened as a species. Listed as 
Vulnerable in Europe (123). Vulnerable and protected in Greece. Six 
subspecies. Common name: False Apollo (125). Other refs: 1, 141, 
169, 264. 

Genus: Hypermnestra Menetries 

^ 3 Hypermnestra helios (Nickerl, 1846) 

'yi A narrow range in Afghanistan, Pakistan (Baluchistan), Iran and U.S.S.R. 
L,^^ (Uzbekistan, Kirghizia and possibly Turkmenistan). Rather poorly known 

and often considered to be a rarity, but it is not known to be threatened and 
is apparently rather common in the Maimana Province of Afghanistan 
(118). The taxon Hypermnestra was transferred to an independent tribe by 
Hiura in 1980 (126) but this was not followed by Hancock (109). Seven 
subspecies. Common name: Desert Apollo (80). Other refs: 1,71, 
141,169,259. 

. ,i^ , Genus: Parnassius Latreille 
V Species-group: szechenyii 



\y/* 4 Parnassius szechenyii Frivaldszky, 1886 
"^J^ Western China (north-eastern Xizang Zizhiqu (Tibet), Qinghai, Gansu, 



35 



Threatened Swallowtail Butterflies of the World 

Sichuan (Szechwan) and Yunnan). No threats known but apparently rather 
rare (264). Possibly eleven distinguishable subspecies (265). Other refs: 
1,31,169. 

5 Parnassius cephalus Grum-Grshimailo, 1891 

Western China (Xizang Zizhiqu (Tibet), Gansu, Sichuan (Szechwan) and 
Qinghai). Not known to be threatened but apparently rather rare (264). 
Eleven distinguishable subspecies, not including maharaja (265). Other 
refs: 1,31. 

6 Parnassius maharaja Avinoff, 1916 

North-western India (Jammu and Kashmir: Ladakh Range). Not known to 
be threatened. Treated only as a subspecies of P. cephalus by Ackery (1), 
Eisner (73) and Hancock (109). A full species in Bryk (31), Munroe (188), 
Smart (250), Talbot (259) and Turlin (265). 

Species-group: delphius Eversmann 

7 Parnassius delphius (Eversmann, 1843) 

Afghanistan, U.S.S.R. (Tadzhikistan, Kirghizia and Uzbekistan), 
northern Pakistan, northern India (including Jammu and Kashmir, Hima- 
chal Pradesh and Uttar Pradesh), Bhutan and western China (Xizang 
Zizhiqu (Tibet), Xinjiang Uygur (Sinkiang) and Qinghai). Widely distri- 
buted, generally rare but locally common, not known to be threatened, but 
requiring further research. Protected by law in India (182) and included in 
the U.S.S.R. Red Data Book, category Vulnerable (8). Up to forty-four 
subspecies (265). Common name: Banded Apollo (80). Also Astor Banded 
Apollo (ssp. nicevillei), Chitral Banded Apollo (ssp. chitralica), Hunza 
Banded Apollo (ssp. hunza), Kafir Banded Apollo (ssp. kafir), Pir Panjal 
Banded Apollo (ssp. atkinsoni), Sikkim Banded Apollo (ssp. lampidius) 
and Tibet Banded Apollo (ssp. latonius) (80). Other refs: 1,4,31, 
71,74,169,259,264. 

8 Parnassius stoliczkanus C. & R. Felder, 1864 

Afghanistan (Badakhshan and possibly Nuristan) (118), northern India, 
(Jammu and Kashmir, Himachal Pradesh and Uttar Pradesh), Pakistan, 
and China (Xizang Zizhiqu (Tibet)). Stated by Evans (80) to be very rare 
and given only subspecific status by him, by Eisner (73) and by Ackery (1), 
but listed as a full species by Munroe (188) with nine subspecies by Bryk 
(31), with seven subspecies by Talbot (259) and with fifteen subspecies by 
Turlin (265). Not mentioned by Hancock (109). It is protected by law in 
India (182). Further information needed on this species. Common name: 
Ladak Banded Apollo (80). 

9 Parnassius patricius "Niepeh, 1911 

U.S.S.R. (Kirghizia); a rather narrow range but not known to be threat- 
ened. Five subspecies (265). Other refs: 1, 31. 

10 Parnassius acdestis Grum-Grshimailo, 1891 

U.S.S.R. (Kirghizia), Nepal, northern India (Jammu and Kashmir, and 
Sikkim), Bhutan and western China (Xizang Zizhiqu (Tibet), Xinjiang 
Uygur (Sinkiang) and Sichuan (Szechwan)). No threats known but appar- 
ently very local and rather rare (264). Nineteen subspecies (265). Other 
refs: 1,31. 

36 



Swallowtails of the world 

Species-group: imperator Oberthiir 

11 Parnassius imperator OhQvXhm , 1883 

North-eastern India (Sikkim) (80), western China (Xizang Zizhiqu (Tibet), 
Qinghai, Gansu, Sichuan (Szechwan) and Yunnan). Not known to be 
threatened as a species, but P. i. aiigustus is protected by law in India (182). 
It seems to be a common subalpine species in China (264). Twenty 
subspecies (265). Common name: Imperial Apollo (80). Other refs: 
1,4,31,207. 

Species-group: charltonius Gray 

12 Parnassius charltonius Gray, 1853 

Afghanistan, U.S.S.R. (Kirghizia, eastern Uzbekistan and Tadzhikistan), 
Pakistan, northern India (including Jammu and Kashmir, Uttar Pradesh 
and Himachal Pradesh) and China (Xizang Zizhiqu (Tibet)). Not known to 
be threatened as a species and is common in Kashmir, but the nominate 
subspecies is protected by law in India (182). Twenty subspecies (265). 
Common name: Regal Apollo (80, 282). Other refs: 1, 4, 31, 72, 74, 259. 

13 Parnassius inopinatus Kotzsch, 1940 

Found only in mountain ranges in north-western Afghanistan where it is 
very localised (264). Little is known about it , but it has recently appeared for 
sale on dealers' lists. No threats recognized, but possibly a candidate for 
Rare status. More information is required. Two subspecies. Other refs: 
1,74,265. 

14 Parnassius loxias Piingeler, 1901 

U.S.S.R. (Kirghizia and Tadzhikistan) and China (Xinjiang Uygur 
(Sinkiang)). A narrow range, possibly extremely rare but little information 
available from these inaccessible regions (264). Not known to be threatened 
but more data required. Two subspecies (265). Other refs: 1, 288. 

15 Parnassius autocrator Avinoff, 1913 
Rare — refer to section 6, p. 185. 

Afghanistan and U.S.S.R. (Tadzhikistan). Originally described as a 
subspecies of P. charltonius Gray. Bryk (31) agrees with this and it is 
omitted by Munroe (188) but thought to be a full species by recent authors 
(1, 109, 250). Narrow range in the Pamir mountains and Hindu Kush and a 
rarity according to Smart (250) . An extremely Rare species in the U.S.S.R., 
where it is threatened by the degradation of high mountain pastures (260) . It 
is included in the U.S.S.R. Red Data Book, category Rare (8). Careful 
monitoring and further study are required. Only one subspecies has been 
described (221) but up to four have been listed. Other ref: 74. 

Species-group: tenedius Eversmann 

16 Parnassius tenedius Eversmann, 1851 

Eastern U.S.S.R. (Tuvinskaya, Chitinskaya and Yakutskaya), Mongolia 
and China (Nei Monggol (Inner Mongolia)) . Not known to be threatened as 
a species, but included in the U.S.S.R. Red Data Book, category 
Vulnerable (8). Very wide range in tundra and mountains (264). Five 
subspecies (265). Other refs: 1, 31, 169. 

Species-group: acco Gray 

17 Parnassius acco Gray, 1853 

37 



Threatened Swallowtail Butterflies of the World 

Pakistan, northern India (Jammu and Kashmir and further east in Sikkim), 
Nepal and China (Xizang Zizhiqu (Tibet)). Possibly also Bhutan if P. 
hannyngtoni Avinoff (below) is regarded as a subspecies. Stated by Antram 
(4) , Evans (80) and Turlin (264) to be exceedingly, or very rare (in India (?)) 
but present status not known. Flies at high altitudes (5500-6200m) so it is 
very rarely captured (264). The subspecies P. a. geminifer is protected by 
law in India (182). More information is needed, particularly from the Indian 
highlands and Tibet. Ten subspecies. Common name: Varnished Apollo 
(80). Other refs: 1, 31, 169, 259. 

18 Parnassius przewalskii AXphexdiky , 1887 

Western China (including Xizang Zizhiqu (Tibet), Sichuan (Szechwan), 
and Yunnan) . Four subspecies (265) . Although recognized by some authors 
(31, 188, 191) as a good species, it is not mentioned by Ackery (1) or 
Hancock (109) and was placed as a subspecies of P. acco by Eisner (73) . Not 
known to be threatened but apparently very rare (264). 

19 Parnassius hannyngtoni Avinoff, 1916 

North-eastern India (Sikkim), China (Xizang Zizhiqu (Tibet)) and Bhutan 
(?). Very rare, according to Evans (80) and Talbot (259) and protected by 
law in India (182), but specific status uncertain. Flies at a very high level 
(6000m) (264). Ackery (1) and Eisner (73) treat it as a subspecies of P. acco 
as, more doubtfully, does Smart (250). Hancock (109) does not mention it, 
but Turlin accepts it as a good species with two subspecies (265). More 
information is required on this species. The spelling of the trivial name 
varies greatly with the authors quoted but Avinoff's original 'hunnyngtoni' 
was arhended to the above form by Bryk (31), Munroe (188) and Talbot 
(259) and is presumably correct since the species was dedicated to 
Hannyngton (259). Common name: Hannyngton's Apollo (80). 

Species-group: simo Gray 

20 Parnassius simo Gray, 1853 

U .S . S . R . (Kirghizia and Tadzhikistan) , Pakistan, northern India (including 
Jammu and Kashmir, and Sikkim), Nepal, western China (Xizang Zizhiqu 
(Tibet), Xinjiang Uygur (Sinkiang) and Gansu) and Mongolia. Not known 
to be threatened. Thirty-four subspecies (265). Common name: Black- 
edged Apollo (80). Other refs: 1, 4, 31, 71. 

Species-group: hardwickii Gray 

21 Parnassius hardwickii Gray, 1831 

Himalayas: northern India (including Jammu and Kashmir, and Sikkim), 
Pakistan, Nepal, Bhutan and China (Xizang Zizhiqu (Tibet)). Not known 
to be threatened. Five subspecies (265). Common name: Common Blue 
Apollo (80, 251). Other refs: 1, 4, 31, 169, 251, 264. 

Species-group: mnemosyne Linnaeus 

22 Parnassius Orleans Oberthiir, 1890 

Southern and western China (Xizang Zizhiqu (Tibet), Xinjiang Uygur 
(Sinkiang), Qinghai, Gansu, Shaanxi (Shensi), Sichuan (Szechwan) and 
Yunnan) and Mongolia. Not known to be threatened. Sixteen subspecies 
(265). Other refs: 1, 31, 169, 207, 264. 

38 



Swallowtails of the world 

23 Parnassius clodius Menetries, 1855 

Western U.S.A. (Alaska, Washington, Idaho, Utah, Montana, Wyoming, 
Oregon, Nevada and California) and south-western Canada (British 
Columbia). Usually abundant and not known to be threatened as a species. 
However, two of the twelve subspecies are in decline. The Californian 
subspecies P. c. strohbeeni formerly occurred in the Santa Cruz Mountains, 
California, but is now believed to be extinct. P. c. shepardii has a restricted 
distribution in the north-western states. It is now absent from former haunts 
in the Snake River Canyon on the border of Oregon and Idaho (216) and is 
Hsted as a Special Species by the Washington State Department of Game. 
Many conservationists blame logging and dam-building for the decline of 
these two subspecies, but others blame drought (216). Common name: 
Clodius Parnassian (216). Other refs: 1, 31, 55, 137, 169, 264, 269. 

24 Parnassius eversmanni Menetries, 1849 

Eastern U.S.S.R, Mongolia to northern China, North Korea, Japan 
(Hokkaido), Alaska and Canada (Yukon Territory, Northwest Territories 
and British Columbia). Included in the U.S.S.R. Red Data Book, category 
Vulnerable (8). Inhabits lowland and tundra. Populations difficult to assess 
but widespread. Not known to be threatened. Twelve subspecies (265). 
Common name: Eversmann's Parnassian (216). Other refs: 1, 31, 137, 141, 
149, 169, 248, 264, 269. 

25 Parnassius felderi Bremer , 1861 

Eastern U.S.S.R. (Khabarovsk Kray). The species is not mentioned by 
Hancock (109), but is listed by Turlin with three subspecies (265). No 
threats known, but little information available. 

26 Parnassius ariadne Lederer, 1853 (= clarius Eversmann, 1843) 
U.S.S.R (Altay mountains and Tadzhikistan) and western MongoHa. Not 
known to be threatened. Localized in distribution (264) and rare in 
collections. Two subspecies (265). Other refs: 1, 31, 169. 

27 Parnassius nordmanni (Nordmann, 1849) 

U.S.S.R. (eastern Armenia and the Bol'shoy Kavkaz (Caucasus) moun- 
tains; Azerbaydzhan and Georgia) and eastern Turkey. Locally distributed 
(264), sometimes abundant (193), but very rare in collections. Inhabits 
inaccessible areas above 2000m and has a short flight season (90,245). 
Included in the U.S.S.R. Red Data Book (8, 260), but known to be present 
in at least seven reserves and probably several more (193). Four subspecies 
described (239, 265), but their distinction is somewhat speculative (193). 
Other refs: 1,31,227. 

28 Parnassius mnemosyne (Linnaeus, 1758) 

Hilly or mountainous regions in Spain (Pyrenees), France, Norway, 
Sweden, Finland, Switzerland, F.R.G., Liechtenstein, Austria (Endan- 
gered/Vulnerable (93)), Italy, Sicily, Albania, Yugoslavia, Greece, Bulga- 
ria, Turkey, Romania, Poland, G.D.R. (?), Czechoslovakia, Hungary, 
U.S.S.R (Latvia, Lithuania, Estonia and Ukraine to Armenia, Bol'shoy 
Kavkaz (Caucasus) mountains, Uzbekistan, Tadzhikistan, Kirghizia and 
the Ural mountains), Syria, Lebanon, Iran, Iraq and Afghanistan. Not 
known to be threatened at the specific level, but now Rare over the whole of 
itsrangeintheU.S.S.R.andlistedintheU.S.S.R. RedDataBook(8,260). 

39 



Threatened Swallowtail Butterflies of the World 

Protected by law in Czechoslovakia, Finland, G.D.R., Greece, Hungary, 
Poland and Lithuania (123). Another species which has been excessively 
subdivided, 125 subspecies being listed by Bryk (31). Common name: 
Clouded Apollo (125). Other refs: 1, 71, 72, 74, 169, 264. 

29 Parnassius stubbendorftMenethes, 1849 

Eastern U.S.S.R., China (Xizang Zizhiqu (Tibet), Heilongjiang, Gansu, 
Sichuan (Szechwan) and Qinghai), Mongolia, North Korea, South Korea 
and Japan (Hokkaido). Twenty-five subspecies listed by Igarashi (141), 28 
by Turlin (265). Not known to be threatened and seemingly quite common 
(264). Other refs: 1, 31, 149, 169, 247. 

30 Parnassius glacialis Butler, 1886 

Eastern China (Hubei, Shandong, Jiangsu, Anhui and Zhejiang), North 
Korea, South Korea and Japan (Hokkaido, Honshu and Shikoku). 
Accepted as a good species by recent authors (1 , 141 , 169, 247), though not 
by Bryk (31) (subspecies of stubbendorfi.) or Munroe (188). The larval 
stages are quite distinct from those of clearly related species. Apparently 
common (264). Fifteen subspecies (265). Other ref: 149. 

Species-group: apollo Linnaeus 

31 Parnassius apollonius (Eversmann, 1847) 

U.S.S.R. (Uzbekistan, Tadzhikistan, Kirghizia) and China (western Xin- 
jiang Uygur (Sinkiang)). Twelve subspecies (265), most of which are 
difficult to distinguish. No evidence of being under threat. Other refs: 
1,31,169,264. 

32 Parnassius honrathi Staudinger, 1882 

Central Asia: U.S.S.R. (western Uzbekistan (?), Tadzhikistan and possibly 
southern Kirghizia) and north-eastern Afghanistan (Pamir Mountains). 
Five subspecies (265). No evidence of being in danger. Other refs: 
1,31,74,169,264. 

33 Parnassius bremeri Bremer, 1864 

Eastern U.S.S.R., northern China (Heilongjiang, Hebei and Shanxi), 
North Korea, South Koreawhereit is threatened (152), Japan (Hokkaido) 
and possibly eastern Mongolia. The Japanese population is either extinct or 
the record is erroneous since no recent specimens are known (264). No 
information available about the status but declining in Korea due to 
overcoUection and loss of foodplant (Dicentra perigrina) on skiing slopes. 
Conservation measures proposed in Korea include legislation prohibiting 
collection of the species and a captive breeding programme (153). More 
information is needed on this species, which has up to sixteen subspecies 
(265). Other refs: 1, 31, 71, 141, 149, 169. 

34 Parnassius jacquemontii^oi?,d\iva\, 1836 

North-eastern Afghanistan (including Badakhshan), northern Pakistan, 
north-western India (Jammu and Kashmir), U.S.S.R. (Tadzhikistan 
(Pamirs) and Uzbekistan) and south-western China (Xizang Zizhiqu 
(Tibet), Xinjiang Uygur (Sinkiang), Gansu and Sichuan (Szechwan)). 
Twenty-five subspecies (265); the nominate subspecies is protected in India 
(182). Not known to be in danger. Common name: Keeled Apollo (80). 
Other refs: 1,31, 74, 169, 259. 

40 



Swallowtails of the world 

35 Parnassius epaphus Oberthiir, 1879 

Afghanistan, Pakistan, northern India (including Jammu and Kashmir, and 
Sikkim), Nepal, Bhutan, China (Xizang Zizhiqu (Tibet), Xinjiang Uygur 
(Sinkiang), Sichuan (Szechwan), Gansu and Qinghai) and possibly 
U.S.S.R. (Tadzhikistan). A fairly broad distribution and not known to be 
threatened. Up to 37 subspecies (265); P. e. hillensis is protected by law in 
India (182). P. beresowskyi Staudinger from China is not mentioned by 
Munroe (188) or Ackery (1) (even as a subspecies) and is treated as a 
subspecies of epaphus by Bryk (31) and Smart (250). Common name: 
Common Red Apollo (80). Other refs: 1, 169, 259. 

36 Parnassius actius {^\ersmdLX\n, 1843) 

U.S.S.R. (Turkmenistan, Uzbekistan (?), Tadzhikistan, Kirghizia, 
Kazakhstan), north-eastern Afghanistan, northern Pakistan, north- 
western India (Jammu and Kashmir) and south-western China (Xinjiang 
Uygur (Sinkiang) and Gansu). Very rare in the U.S.S.R. and declining due 
to changes in high mountain meadows (260). Included in the U.S.S.R. Red 
Data Book, category Vulnerable (8). More information is needed on this 
species, which has up to nineteen subspecies (265). Other refs: 
1,31,74,169,259. 

37 Parnassius phoebus (¥ahx'ic\\xs, 1793) 

Palearctic: Alps (France, southern F.R.G., Switzerland, Liechtenstein (?) 
and Austria), U.S.S.R. (Ural mountains, Kazakhstan, Altay, Siberia and 
Kamchatka), Mongolia, China (Xinjiang Uygur (Sinkiang)), western 
U.S.A. (Alaska, Washington, Idaho, Montana, South Dakota, Wyoming, 
California, Nevada, Utah, Colorado and New Mexico) and Canada (British 
Columbia and Alberta). Not under threat as a species, although some of the 
45 subspecies (265) may be threatened. Included in the U.S.S.R. Red Data 
Book, category Vulnerable (8) and listed as Vulnerable throughout Europe 
(123). Protected in France. Common names: Small Apollo (125), Phoebus 
Parnassian (216). Other refs: 1, 31, 62, 72, 118, 137, 169, 269. 

38 Parnassius tianschanicus Oberthiir, 1879 

U.S.S.R. (Uzbekistan, Tadzhikistan and Kirghizia), Afghanistan, Pakis- 
tan, India (Jammu and Kashmir) and western China (Xinjiang, Uygur 
(Sinkiang)). Locally common but generally very rare in the U.S.S.R. where 
it is declining and is included in the U.S.S.R. Red Data Book, category 
Vulnerable (8, 118, 260). More information is required. Up to eighteen 
subspecies (265). Common name: Large Keeled Apollo (80). Other refs: 
1,31,74,259. 

39 Parnassius nomion Fischer de Waldheim, 1823 

Eastern U.S.S.R. (Irkutsk, Buryatskaya, Amurskaya, Khabarovsk and 
Altay), Mongolia, China (Gansu, Qinghai, Shaanxi (Shensi), Heilong- 
jiang, Liaoning and Nei Monggol (Inner MongoHa) (?)), North Korea and 
South Korea. A doubtful record from North America (Alaska) (141) and 
another from California (55). Authority wrongly given as Hiibner by 
Munroe (188) and Smart (250). Up to 31 subspecies (265) including P. 
nomius Grum-Grshimailo from China, which is not listed by Munroe (188), 
Hancock (109) or Ackery (1) (even as a subspecies) and is treated as a 
subspecies oi nomion by Bryk (31) and Eisner (73). Other refs: 149, 169. 

41 



Threatened Swallowtail Butterflies of the World 

40 Parnassius apollo Linnaeus, 1758 
Rare — refer to section 6, p. 187. 

Among mountains at subalpine levels in France, Andorra, Spain (59), 
Netherlands (only old and questionable records), Norway (Vulnerable, 
possibly Endangered (123)), Sweden, Finland, F.R.G., Switzerland, 
Liechtenstein, Austria (Endangered/Vulnerable (93)), Italy, Sicily, 
G.D.R. (Extinct), Poland, Czechoslovakia, Hungary (migratory), Yugo- 
slavia, Greece, Albania, Bulgaria, Turkey (including the border area with 
Iran and Iraq), Romania, Syria, U.S.S.R. (Latvia, Lithuania, Ukraine, to 
Armenia, Bol'shoy Kavkaz mountains (Caucasus), Ural mountains and 
Siberia), Mongolia and China (Xinjiang Uygur (Sinkiang)). Concern for 
this species seems to be based on local threats to populations, particularly 
those in 'developed' areas of western Europe. In other parts of Europe P. 
apollo can be very numerous. The inclusion of P. apollo on Appendix 2 of 
CITES is questionable when so many threatened species are excluded. The 
unnecessary division of the species (over 160 'subspecies' are Hsted by Bryk 
(31)) has led to exaggerated fears for particular populations. However, 
numbers of 'Apollos' are taken every year in western Europe for trade, 
populations are often very isolated and it is declining, threatened, or rare in 
many countries (64, 123). It is decHning sharply in all parts of the U.S.S.R. 
and is included in the U.S.S.R. Red Data Book (category Vulnerable) and 
the Red Book of the Ukrainian S.S.R. (8, 260). There is no information 
from the eastern section of the Apollo's range. The 87 countries that are 
party to CITES are obliged to invoke national legislation implementing the 
convention and all countries in the European Economic Community have 
already done so. The Apollo is also protected by law in several other 
countries including Austria, Czechoslovakia, Finland, G.D.R. , Greece and 
Poland. The species should be closely monitored. Common name: Apollo. 
Other refs: 1, 30, 100, 101, 118, 125, 141. 



Subfamily: PARNASSIINAE 
Tribe: Zerynthiini 

Genus: Sericinus Westwood 

41 Sericinus montela Gray, 1843 (= telamon Donovan, 1798) 

Eastern U.S.S.R. (PrimorskiyKray), China (Heilongjiang,JiHn,Liaoning, 
Hebei, Shandong, Anhui, Jiangsu, Hubei, Hunan, Jiangxi and Gansu), 
North Korea and South Korea. Threatened by changes in flood plain 
vegetation in the U.S.S.R., where it is included in the U.S.S.R. Red Data 
Book, category Rare (8,260). More information is required on this 
primitive and important zerynthiine. Eleven subspecies. Other refs: 
1,31,141. 

N.B. The following six species have seen several changes of genus. First 
described as Thais by Fabricius in 1807, this invalid name (homonym) was 
replaced with Parnalius by Rafinesque in 1815, and almost simultaneously 
with Zerynthia by Ochsenheimer in 1816. The latter name achieved popular 
usage and a recent attempt to revive the validity of the name Parnalius was 
suppressed by the Commission on Zoological Nomenclature (118, 194). 
The name Allancastria was raised by Bryk in 1934 and applied to those 

42 



Swallowtails of the world 

species of Zerynthia found in Asia Minor. This course has been followed by 
Hancock (109). 

Genus: Allancastria Bryk 

42 Allancastria cerisy (Godart, 1824) 

Albania, Cyprus, Crete, Greece, Yugoslavia, Bulgaria, Romania, Turkey, 
U.S.S.R. (Armenia and Bol'shoy Kavkaz (Caucasus) mountains), Iran, 
Iraq, Syria, Israel and Lebanon. The species is quite local and often rare 
(160). Declining rapidly over the whole of its range in the U.S.S.R. 
Included in the Red Book of the Ukraine S.S.R. and the U.S.S.R. Red 
Data Book (8, 260). Localised but common in Turkey (264). Protected by 
law in Greece . Not known to be threatened in the rest of its extensive range , 
but clearly requires monitoring. Eleven subspecies. Common name: 
Eastern Festoon (125). Other refs: 1, 31, 141, 157, 158, 169, 194. 

43 Allancastria deyrollei Oberthiir, 1872 

Turkey, Syria, Lebanon, Israel, Jordan, Iraq and possibly Iran (158). 
Common and widespread in the Lebanon (158). Treated as a subspecies of 
A. cerisy by Bryk (28) and Igarashi (141) and not listed by Ackery (1), but 
Larsen (157, 158) has shown it to be distinct. Other ref: 264. 

44 Allancastria caucasica Lederer, 1864 

U.S.S.R. (Armenia and Bol'shoy Kavkaz (Caucasus) mountains) and 
northern Turkey (160). Specific status uncertain. Regarded as a subspecies 
oi A. cerisy by most authors (31, 141). Larsen considers it to be distinct 
(160) as does Turlin (264). Conservation status needs clarification. 

45 Allancastria louristana (Le Cerf, 1908) 

West and south-west Iran, mountains of Louristan. Originally described as 
a subspecies of A. cerisy, but recently raised to full species status. Not 
known to be threatened, but conservation status requires confirmation. 
Ref: 162. 

Genus: Zerynthia Ochsenheimer 

46 Zerynthia poly xena {Hqvus &.Schititxm\i\\ex, 111^) 

Southern France, Italy, Sicily, Austria, Hungary, Czechoslovakia, Yugo- 
slavia, Albania, Greece, Bulgaria, Romania and south-western U.S.S.R. 
Widely distributed though local and not under threat as a species. 
Nevertheless, it is rare and declining rapidly in the U.S.S.R.; in need of 
habitat protection to ensure its survival. Included in the U.S.S.R. Red Data 
Book, category Vulnerable (8,260); listed as Vulnerable throughout 
Europe and protected by law in Czechoslovakia, Greece, Hungary and 
some of the Austrian provinces (123). Continuous monitoring of this 
species is necessary. Nineteen subspecies. Z. hypermnestra Scopoli, 1763 is 
an invalid homonym (125). Common names: Southern Festoon (125), 
Birthwort Butterfly (253). Other refs: 1, 31, 141, 169, 194. 

47 Zerynthia rumina (Linnaeus, 1758) 

Southern France, Italy, Spain, Portugal, Algeria, Morocco and Tunisia. 
Often fairly common within its somewhat restricted range but has been 

43 







Threatened Swallowtail Butterflies of the World 

listed as Vulnerable throughout Europe. Threatened to varying degrees in 
France (where it is more local than Z. polyxena (264) and the distinctive 
form honoratii is protected by law), Spain and Italy (123). Five subspecies. 
Common name: Spanish Festoon (125). Other refs: 1, 31, 141, 169. 

Genus: Bhutanitis Atkinson 

48 Bhutanitis mansfieldi (Riley, 1939) 
Rare — refer to section 6, p. 192 

Known to Ackery (1) only from the female holotype from Yunnan, China, 
but a contemporary male and a second female have since been found. 
Recent specimens have also come from Sichuan (Szechwan) and a second 
subspecies described (238). B. mansfieldi was separated as the type species 
of a monobasic genus Yunnanopapilio by Hiura in 1980 (126) but this was 
reduced to a subgenus by Saigusa and Lee (238). The name is not used by 
Hancock (109). Other refs: 118, 199, 229, 264. 

49 Bhutanitis thaidina (Blanchard, 1871) 
Rare — refer to section 6, p. 194 

A narrow range in China (Yunnan, Sichuan (Szechwan) and Shaanxi 
(Shensi) provinces). Rare status is justified, at least until more is known 
about the species. No subspecies described. Refs: 1, 19, 31, 207. 

50 Bhutanitis lidderdalii Atkinson, 1873 

Bhutan, northern India (Assam, Sikkim, Manipur and Nagaland), 
northern Burma, Thailand and China (Sichuan (Szechwan) and Yunnan 
provinces) (31). Not so restricted as B. thaidina and probably not in danger 
at the moment, but needs monitoring to ascertain its actual status. Up to 
three subspecies, the nominate subspecies protected by law in India (182), 
the third recently described from Thailand. The Thailand population is 
apparently confined to northern areas around Chiang Mai (141) and is 
considered to be a relict (7). Hundreds of individuals are exported from 
Thailand annually to collectors (7). Deforestation may also be a threat and 
more data are needed. Common name: Bhutan Glory (80). Other refs: 
1,259. 

51 Bhutanitis ludlowi Gabriel, 1942 
Insufficiently Known — refer to section 6, p. 196 

Bhutan: Trashiyangsi Valley only. The type-series is unique (1). Other ref: 
91. 



\ Genus: Luehdorfia Criiger 

r * N.B. The genus Luehdorfia has a very confusing taxonomy. There is much 

\ variation between authors regarding species and the distribution of the 

individual species is also inconsistently reported. 

52 Luehdorfia chinensis Leech, 1893 

Insufficiently Known — refer to section 6, p. 197 

Eastern China (Anhui, Hubei, Jiangsu and Jiangxi provinces). Status not 
entirely certain: variously treated as a separate species (109, 141), as a 
subspecies of L. japonica (1) or L. puziloi (30, 31) or ignored (188). Lee 
(163) and Igarashi (141) are followed as authors who have studied the early 

44 



y" 



Swallowtails of the world 

stages. The above status seems appropriate until its specific identity and 
possible threats can be assessed. Two subspecies (141). 

53 Luehdorfia japonica Leech, 1889 
Indeterminate — refer to section 6, p. 198 

Restricted to the island of Honshu, Japan (141). Ackery (1) includes 
chinensis within this species (although its distribution is in southern China 
disjunct from the range oi japonica) while Bryk includes chinensis in L. 
puziloi (30, 31). The subspecies supposed to occur in Taiwan appears to be 
either a doubtful record or now extinct (141,246,264). One or two 
subspecies. Other refs: 149, 166, 247. 

54 Luehdorfia puziloi (Erschoff, 1872) 

Extreme south-eastern U.S.S.R. (Primorskiy Kray), north-eastern China 
(Manchuria), North Korea, South Korea and Japan (Honshu and Hok- 
kaido). Appears to be declining in Korea due to overcollecting and 
pollution (153), and in Japan due to overcollecting and habitat destruction 
(124). Conservation measures proposed in Korea include legislation 
prohibiting collecting and a programme of captive breeding (153). It has 
been included in the U.S.S.R. Red Data Book, category Rare (8). Not 
known to be threatened over the rest of its range, but clearly requires 
monitoring. Five subspecies. Other refs: 1, 141, 149, 247. 

Subfamily: PAPILIONINAE 
Tribe: Leptocircini 

Genus: Iphiclides Hiibner 

Iphiclides podalirius (Linnaeus, 1758) 

Europe (excluding the British Isles, Norway, Sweden (?) and Finland), 
North Africa and the Middle East (possibly excluding Saudi Arabia, 
Yemen, South Yemen, Oman, United Arab Emirates and Qatar), Afg- 
hanistan, Pakistan, India and China. Generally common and not seriously 
threatened although it is protected by law in Czechoslovakia, G.D.R., 
Hungary, Luxembourg and Poland, included in the U.S.S.R. Red Data 
Book (8), listed as Endangered-Rare in the provinces of Austria (93), and 
protectsd in seven of them, and Indeterminate throughout Europe (123). 
Six subspecies including /. p. feisthamelii Duponchel from Morocco, 
Algeria, Tunisia, Spain, Portugal and France, which is variously treated as a 
doubtfully good species (250) or a subspecies of /. podalirius (125). 
Common names: Scarce Swallowtail (125, 253, 282), Sail Swallowtail (253), 
Pear-tree Swallowtail (158). Other refs: 141, 202. 

56 Iphiclides podalirinus (OherthuT, 1890) 

China (Xizang Zizhiqu (Tibet) and Yunnan). A good species in Munroe 
(188) and Hancock (109) but stated by Smart (250) to be possibly a 
subspecies of /. podalirius. Not known to be threatened, but more data 
needed. 



Genus: Teinopalpus Hope 

Teinopalpus imperialis Hope, 1843 
Rare — refer to section 6, p. 200 

45 





Threatened Swallowtail Butterflies of the World 

Nepal, northern India (West Bengal, Meghalaya, Manipur, Sikkim and 
Assam), Bhutan, northern Burma and China (Hubei and Sichuan). A 
prized species that commands high prices on dealers lists and is hunted 
mercilessly in the Himalayas. It is described as rare by Talbot (259) and is 
protected by law in India (182). The habitat is in mountainous regions, it 
keeps to the tops of trees , has a strong flight and is difficult to capture (4) . T. 
behludinii (Pen, 1937) from Sichuan is almost certainly referable to T. 
imperialis (185,207). Two subspecies. Common name: Kaiserihind (80), 
Kaiser-I-Hind (251). Other refs: 52, 135, 141, 179, 197, 286. 

58 Teinopalpus aureus Mell, 1923 

Insufficiently Known — refer to section 6, p. 204. 

Despite its unique appearance , its specific status is uncertain ; it is an isolated 
taxon of Teinopalpus from south-eastern China (Guangdong Province). If 
distinct it is presumably Rare, but more information is required. Common 
name: Golden Kaiser-I-Hind. Refs: 52, 178. 



Genus: Meandrusa Moore 

59 Meandrusa sciron (Leech, 1890) 

China, India (Sikkim and Assam), Bhutan, south-western Thailand and 
Burma. The name hercules Blanchard is a synonym of sciron. The name 
gyas Westwood is a synonym of lachinus Fruhstorfer, which is currently 
regarded as conspecific with sciron. Status and distribution of the species 
not well known but not known to be threatened. Not uncommon in India, 
where it is protected by law under the name gyas (182). More information 
needed. Two subspecies. Common name: Brown Gorgon (80). Other refs: 
4,141,207,259. 

60 Meandrusa pay eni {^o\i,d\x\a\, 1836) 

Northern India (Assam and Sikkim), Bhutan, southern Burma, northern 
Thailand, northern Vietnam, Laos, China (Hainan (Guangdong prov.)). 
Peninsular and Eastern Malaysia, Brunei, Indonesia (Sumatra, Java and 
Kalimantan (?)) (52, 262). Not thought to be threatened across most of its 
range but considered to be Vulnerable and in need of protection in 
Peninsular Malaysia ( 10). Six or seven subspecies. Common names: Yellow 
Gorgon (80), Outlet Sword (3), the Sickle (3). Other refs: 4,45, 
87,141,259. 



0^ 



\>^' 




Genus: Eurytides Hiibner 

N.B. The genus name Eurytides is usually applied to new world forms 
whereas Graphium is applied to old world forms. There is controversy over 
the morphological distinction of the two genera and the traditional division 
is upheld here. 

Species-group: marcellus Cramer 

61 Eurytides (Protesilaus) marcellus (Cramer, 1777) 

Canada (southern Ontario) and eastern U.S.A. Local, but not threatened. 
Several named seasonal forms, but subspecies doubtful. Common name: 
Zebra Swallowtail (154, 216). Other refs: 55, 62, 137, 234, 269. 



46 



Swallowtails of the world 

62 Eury tides (Protesilaus) epidaus (Doubleday, 1846) 

Mexico, Guatemala, Honduras, Belize, Nicaragua (?) and Costa Rica. 
Seasonally abundant in Costa Rica, flying in open areas associated with 
deciduous forest (61). Generally common and not threatened. Three 
subspecies. Other refs: 51, 55, 57, 62, 232, 234, 269. 

63 Eurytides (Protesilaus) zonaria (Butler, 1869) 

Confined to Hispaniola. Said to be widespread and not uncommon (243) 
though it is not well known. No threatened category has been given but the 
status of the species should be better determined if possible. Common 
name: Haitian Kite (230). Other refs: 51, 55, 234. 

64 Eurytides (Protesilaus) marcellinus (Doubleday, 1845) 
Vulnerable — refer to section 6, p. 206. 

Restricted to Jamaica. The butterfly is not common and because of the 
relatively small size of Jamaica, the species' status needs to be carefully 
monitored. Common name: Jamaican Kite (21,230). Other refs: 
51,55,234,266,267. 

65 Eurytides (Protesilaus) celadon (Lucas, 1852) 

Restricted to Cuba but the butterfly is 'apparently widespread' in that 
country (230). Common name: Cuban Kite (230). Other refs: 
21,51,55,234. 

66 Eurytides (Protesilaus) philolaus (Boisduval, 1835) 

Mexico, Belize, Guatemala, El Salvador (?), Honduras, Nicaragua and 
Costa Rica. Common and sometimes very abundant (57, 269); no known 
threats. Common name: Dark Zebra Swallowtail (216). Other refs: 
51,55,62,175,234. 

67 Eurytides (Protesilaus) anaxilaus (C. & R. Felder, 1864) 

{=arcesilaus 'LMcas, 1852) 
Northern Venezuela and Colombia. Not known to be threatened (234). 
Other ref: 51. 

68 Eurytides (Protesilaus) xanticles (Bates, 1863) 

Northern Colombia and Panama, apparently with a restricted range. 
Generally uncommon although it is fairly common in the Panama Canal 
Zone where there is a national park (221). No threats known, but further 
information is needed for this narrowly distributed species. Other refs: 
51,55,234. 

69 Eurytides (Protesilaus) oberthueri (Rothschild & Jordan, 1906) 
Honduras and Mexico (disjunct range) (269). Specific status questionable, 
possibly only a form of E. (P.) philolaus, but accepted by Munroe (188) and 
Hancock (109). Only three specimens appear to have been recorded (221). 
Other refs: 51, 55, 62, 234, 275. 

Species-group: bellerophon Dalman 

70 Eurytides (Protesilaus) bellerophon (Dalman, 1823) 

Northern Argentina, south-eastern Brazil and possibly Paraguay. Appar- 
ently not common, but not known to be threatened. More information 
needed. Refs: 51, 55, 122, 234. 

47 



Threatened Swallowtail Butterflies of the World 

Species-group: protesilaus Linnaeus 

71 Eurytides (Protesilaus) agesilaus (Guerin and Percheron, 1835) 
Mexico, Central America and South America (excluding Chile and 
Uruguay). Rare in Costa Rica, common in Panama (61). Generally quite 
abundant and not threatened. Five subspecies (also variously treated as 
forms or good species including E. (P.) a. autosilaus (Bates, 1861)). Other 
refs: 26, 51, 55, 62, 122, 234, 269. 

72 Eurytides (Protesilaus) orthosilaus (Weymer, 1889) 

Paraguay and Brazil. Formerly believed to be quite rare, but now known to 
occur over a very wide area of cerrado in the Mato Grosso of central Brazil 
(180), where it can be seen almost any day in the year. Males frequent 
favoured sandy areas and females seek nectar at cerrado flowers (23) . Other 
refs: 26, 51, 55, 234. 

73 Eurytides (Protesilaus) helios (Rothschild and Jordan, 1906) 
Southern Brazil, northern Argentina and possibly Paraguay. Not uncom- 
mon, but not known to be threatened. Refs: 26, 51, 55, 122, 234. 

74 Eurytides (Protesilaus) stenodesmus (Rothschild and Jordan, 1906) 
Paraguay, Brazil and northern Argentina. Common and not threatened. 
Sometimes confused with E. (P.) helios. Refs: 51, 55, 122, 234. 

75 Eurytides (Protesilaus) earis (Rothschild and Jordan, 1906) 

Ecuador (55) and Brazil (26). An uncommon and little known species, but 
no threats are recognized. More information required. Other ref: 234. 

76 Eurytides (Protesilaus) telesilaus (Felder, 1864) 

Panama, Colombia, Venezuela, Guyana, Surinam, French Guiana, Trini- 
dad, Ecuador, Brazil, Peru, Bolivia, Paraguay and possibly Argentina. Not 
known to be threatened. Ranched in Brazil (24). Four subspecies. Common 
name: Southern White Page (9, 282). Other refs: 26, 51, 55, 234, 269. 

77 Eurytides (Protesilaus) agidari (D'Almeida, 1937) 

Brazil, from Belem (Para) to Benjamin Constant (Amazonas). No known 
threats but more data required. Refs: 51, 55. 

78 Eurytides (Protesilaus) embrikstrandi (D'Almeida, 1936) 

Brazil. A little known and fairly recently described species. No threats 
known. Refs: 51,55. 

79 Eurytides (Protesilaus) travassosi (D'Almeida, 1938) 

Brazil. Another little known and relatively recently described species. Not 
known to be threatened Refs: 51, 55. 

80 Eurytides (Protesilaus) molops (Rothschild and Jordan, 1906) 
Colombia, Venezuela (?), Guyana, Surinam, French Guiana (?), Brazil, 
Ecuador, Peru and Bolivia. Not recognized as threatened. Three subspecies 
(all treated as full species by D'Almeida (55)) but E. (P.) m. hetaerius was 
transferred to E. (P.) macrosilaus by Hancock (109) and replaced in this 
species by E. (P.) m. leucosilaus (Zikan) (115). Other refs: 51, 234. 

81 Eurytides (Protesilaus) macrosilaus (Gray, 1852) 

Mexico, Guatemala, Honduras, Belize and Nicaragua. Conspecific with E. 
(P.) protesilaus, according to most authors (188, 234), but listed as a species 
by D'Almeida (55) and Hancock (109). E. (P.) penthesilaus (Felder, 1864) 

48 



Swallowtails of the world 

from Mexico is regarded as a subspecies of E. (P.) protesilaus by most 
authors (188, 234, 269), as a subspecies of E. (P.) macrosilaus by Hancock 
(109), and only by D'Almeida (55) as a good species. It should be included 
under E. (P.) macrosilaus if this is to be accepted as a good species (115). 
Two further subspecies were placed here by Hancock (109). 

82 Eurytides (Protesilaus) nigricornis (Staudinger, 1884) 

Eastern Paraguay and southern Brazil. No known threats but more data 
needed. Ranched in Brazil (24). Listed by D'Abrera (51) as a subspecies of 
E. (P.) protesilaus, but given full species rank by Hancock (109). The larvae 
apparently feed on Lauraceae, whilst E. (P.) protesilaus utilizes Magno- 
Haceae (115). 

83 Eurytides (Protesilaus) protesilaus (Linnaeus, 1758) 

Central America, Trinidad and South America (excluding Chile and 
Uruguay). Apparently common and not threatened. Three subspecies 
according to Hancock (109). Common name: Northern White Page (9), 
Swordtail (282). Other refs: 26, 51, 55, 61, 62, 115, 122, 188, 234, 269. 

84 Eurytides (Protesilaus) glaucolaus (Bates, 1864) 

Panama, South America (excluding Chile, Uruguay and Argentina) and 
possibly Costa Rica (51). Not known to be threatened. Sometimes difficult 
to separate from E. (P.) molops (51). Four subspecies. Other refs: 55, 234. 

Species-group: asius Fabricius 

85 Eurytides (Protesilaus) asius (Fabricius, 1781) 

South-western Brazil and eastern Paraguay. Not known to be threatened. 
Ranched in Brazil (24). Other refs: 51, 55, 234. 

86 Eurytides (Protesilaus) microdamas (Burmeister, 1878) 

Paraguay and adjacent areas of Argentina and Brazil. Little known but no 
threats recognized. Refs: 51, 55, 122, 234. 

87 Eurytides (Protesilaus) thymbraeus (Boisduval, 1836) 

Mexico, Guatemala, Belize, Honduras and El Salvador. Apparently fairly 
common and widespread within its range (234, 269) and not known to be 
threatened. Two subspecies. Other refs: 51, 55, 62. 

88 Eurytides (Protesilaus) belesis (Bates, 1864) 

Mexico, Guatemala, Honduras and Nicaragua. Uncommon in some areas 
(57), but not rare and not known to be threatened. Dimorphic, but no 
accepted subspecies. Other refs: 51, 55, 62, 232, 234, 269. 

89 Eurytides (Protesilaus) branchus (Doubleday, 1846) 

Mexico, Guatemala, Honduras, Belize (?), El Salvador (?), Nicaragua and 
Costa Rica. Apparently common north of Costa Rica, but rare within Costa 
Rica (61). Not believed to be threatened. Two forms, analogous to those of 
E. (P.) belesis. Other refs: 51, 55, 57, 62, 234, 269. 

90 Eurytides (Protesilaus) ilus Fabricius, 1793 

Colombia, Panama and northern Venezuela. Rare and Httle known but not 
recognized as threatened. Refs: 51, 55, 234. 

91 Eurytides (Protesilaus) lysithous (Hiibner, 1821) 

Brazil, Argentina and eastern Paraguay. Apparently not uncommon and 
not threatened as a species. Ranched in Brazil (24). E. (P.) lysithous mimics 

49 



Threatened Swallowtail Butterflies of the World 

various species of Parides and seven taxa are treated variously as subspe- 
cies, species, or merely forms (234). E. (P.) kumbachi from Salto Grande is 
regarded as an aberration of E. (P.) lysithous by D'Almeida (55) and 
Hancock (109). 

Eury tides (Protesilaus) lysithous harrisianus (Swainson, 1822) 
Endangered — refer to section 6, p. 208. 

This Brazilian sub-species mimics the Vulnerable Parides ascanius and is 
itself seriously Endangered (283). Nearly all known colonies have been 
destroyed by development and only a single known locality remains (283). 
The subspecies is now on the official list of Brazilian animals threatened 
with extinction (23). Common name: Harris' Mimic Swallowtail. Other 
refs:26,55,122. 

92 Eurytides (Protesilaus) ariarathes (Esper, 1788) 

Colombia, Venezuela, Guyana, Surinam, French Guiana, Brazil, Ecuador, 
Peru and Bolivia. Females resemble the females of Aristolochia feeding 
Papilios and exhibit geographical variation (51). Not uncommon and no 
known threats. Up to nine subspecies. Other refs: 26, 55, 234. 

93 Eurytides (Protesilaus) harmodius (Doubleday, 1846) 

Colombia, Ecuador, Peru and Bolivia. Common and not threatened. Five 
or six subspecies. The female of the nominate form is known from only one 
specimen. Refs: 51, 55, 234. 

94 Eurytides (Protesilaus) trapeza (Rothschild and Jordan, 1906) 

Known only from Ecuador and north-eastern Peru (restricted range), but 
not uncommon. Common in the Napo province of Ecuador but declining in 
Pastaza province (221). Two subspecies. Other refs: 51, 55, 234. 

95 Eurytides (Protesilaus) xynias (Hewitson, 1875) 

Ecuador, Bolivia and Peru. Not uncommon and not threatened. Two 
subspecies. Refs: 51, 55, 234. 

96 Eurytides (Protesilaus) phaon (Boisduval,1836) 

Mexico, Guatemala, Honduras, Belize, Nicaragua, Costa Rica (Hkely, but 
no records yet, 61), Panama (?), Colombia, Venezuela and western 
Ecuador. Not known to be threatened. Two forms, not thought to be of 
subspecific status (234). Other refs: 51, 55, 57, 62, 269. 

97 Eurytides (Protesilaus) euryleon (Hewitson, 1855) 

Costa Rica, Panama, western Colombia and Ecuador. Present in most 
habitats throughout the year, at least in Costa Rica (61). Not known to be 
threatened. Five subspecies. Other refs: 51, 55, 234. 

98 Eurytides (Protesilaus) pausanias (Hewitson, 1852) 

Costa Rica, Panama, Colombia, Venezuela, Trinidad, Guyana, Surinam, 
French Guiana, northern Brazil, Ecuador, Peru and Bolivia. Very rare in 
Costa Rica and Panama (61,274), generally uncommon and possibly 
threatened. Little known of its biology and ecology. Mimics the unpalatable 
Heliconius wallacei (Heliconiinae) (51). More data required on conser- 
vation status in the main part of its range. Up to four subspecies. Other refs: 
9,55,234. 

99 Eurytides (Protesilaus) protodamas (Godart, 1819) 

Southern Brazil, Paraguay and Argentina. Not known to be threatened. 

50 



Swallowtails of the world 

Two forms, not regarded as subspecies. Ranched in Brazil (24). Other refs: 
51,55,122,234. 

Species-group: thyastes Drury 

100 Eury tides (Eury tides) marchandi (Boisduval, 1836) 

Mexico, Central America, Colombia and western Ecuador. Inhabits rain 
forest up to 1000 m in Costa Rica (61). Less common than E. (E.) thyastes 
but not rare or threatened. Two subspecies. Other refs: 51,55, 57, 
62, 234, 269. 

101 Eurytides (Eurytides) thyastes (Drury, 1782) 

Eastern Ecuador, Peru, BoHvia and Brazil. Not uncommon and not threat- 
ened. Ranched in Brazil (24). Three subspecies. Other refs: 51, 55, 234. 

102 Eurytides (Eurytides) calliste (Bates, 1864) 

Mexico, Guatemala, Belize, Honduras, El Salvador (?), Nicaragua (?), 
Costa Rica and Panama (261). Apparently common in Mexico, but rarely 
seen in Costa Rica (61). Not known to be threatened. Two subspecies. 
Other refs: 51, 55, 62, 234, 269. 

103 Eurytides (Eurytides) leucaspis (Godart, 1819) 

Ecuador, Colombia, Peru and Bolivia. Common and not threatened. Two 
subspecies. (222). Other refs: 51, 55, 234. 

104 Eurytides (Eurytides) lacandones (Bates, 1864) 

Mexico, Central America, Colombia, Ecuador, Peru and Bolivia. Possibly 
conspecificwith£. (E.)dioxippus (222). Rare in Costa Rica (61). Generally 
poorly known, but wide-ranging and not believed to be threatened. Two 
subspecies. Other refs: 51, 55, 62, 234, 269. 

105 Eurytides (Eurytides) dioxippus (Hewitson, 1855) 

Colombia. Possibly conspecific with E. (E.) lacandones (222). Restricted 
range, but not uncommon and not known to be threatened. Other refs: 
51,234. 

Species-group: dolicaon Cramer 

106 Eurytides (Eurytides) serville (Godart, 1824) 

Colombia, Venezuela, Brazil, Ecuador, Peru and Bolivia. Possibly ccnspe- 
cific with E. (E.) columbus (222). Common and not threatened. Two 
subspecies. Other refs: 51, 55, 234. 

107 Eurytides (Eurytides) columbus (Kollar, 1850) 

Colombia, north-western Ecuador and possibly Venezuela. Possibly con- 
specific with E. (E.) serville (222). No known threats. Other refs: 
51,55,234. 

108 Eurytides (Eurytides) orabilis (Butler, 1872) 

Guatemala, Costa Rica, Panama and Colombia. In Costa Rica always 
associated with primary forest; always uncommon and solitary but present 
throughout the year (61). Not recognized as threatened. Two subspecies. 
Other refs: 51,55,234. 

109 Eurytides (Eurytides) salvini (Bates, 1864) 

Mexico, Guatemala and Belize. Not particularly common but not known to 
be threatened. Refs: 51, 55, 57, 62, 234, 269. 

51 



Threatened Swallowtail Butterflies of the World 

110 Eury tides (Eury tides) callias (Rothschild and Jordan, 1906) 
Eastern Ecuador and Peru. Not known to be threatened and not uncom- 
mon. Refs: 51,55,234. 

111 Eury tides (Eury tides) dolicaon (Cramer, 1775) 
South America (excluding Chile and Uruguay). Ranched in Brazil (24). 
Seven, possibly eight subspecies. Other refs: 26, 51, 54, 55, 122, 234. 

112 Eurytides (Eury tides) iphitas (Hiibner, 1821) 
Vulnerable — refer to section 6, p. 211. 

Brazil . Extremely rare and not seen for several decades (23) . Believed to be 
seriously threatened but more details needed. Common name: Yellow 
Kite. Other refs: 51, 55, 139, 234. 

Genus: Protographium Munroe 

113 Protographium leosthenes (Doubleday, 1846) 
Australia; restricted to the east coast from Cape York southwards to 
Sydney and (as a separate subspecies) to a small area of the Northern 
Territory. Apparently not uncommon and not threatened, but status needs 
to be monitored. Two subspecies. Refs: 14, 48, 141. 

Genus: Lamproptera Gray 

114 Lamproptera meges (Zinken-Sommer, 1831) 
North-eastern India (Assam), Burma, Thailand, Laos, Vietnam, southern 
China (including Hainan (Guangdong prov. )), Kampuchea, Peninsular and 
Eastern Malaysia, Philippines, Brunei and Indonesia (Sumatra, Babi Is, 
Nias, Bangka, Java, Sulawesi and Kalimantan (52, 262)). Not known to be 
threatened in most of its range, but considered to be Vulnerable and in need 
of some protection in Peninsular Malaysia (10). Ten subspecies (262). Com- 
mon name: Green Dragontail (45, 80, 282). Other refs: 87, 131, 141, 259. 

115 Lamproptera curius (Fabricius, 1787) 
North-eastern India (Assam), Burma, Thailand, southern China (including 
Hainan (Guangdong prov.)). Hong Kong, Laos, Vietnam, Kampuchea, 
Peninsular and Eastern Malaysia, Philippines (Palawan), Brunei and 
Indonesia (Sumatra, Bangka, Nias, Bunguran, Java and Kalimantan 
(52, 262)). Not known to be threatened in most of its range, but considered 
to be Vulnerable and in need of some protection in Peninsular Malaysia 
(10). Four subspecies. Common name: White Dragontail (3,80). Other 

._ refs: 45,87, 131,141,259. 

Genus: Graphium Scopoli 
Subgenus: Pazala Moore 

,A^^ Species-group: alebion Gray 

116 Graphium (Pazala) eurous (Leech, 1892-94) 

3^ Northern India (Kashmir and Jammu, Himachal Pradesh, Uttar Pradesh, 

Assam, Sikkim and Manipur), Nepal, northern Burma, south-western and 
central China and Taiwan. Common and not threatened. Five subspecies 
including caschmirensis Rothschild from Northern India (Jammu and 

52 



Swallowtails of the world 

Kashmir, and the Himalayas). Common name: Six-bar Swordtail (80, 251). 
Otherrefs:4, 52, 141,259. 

117 Graphium (Pazala) mandariniis (Oberthur, 1879) (= glycerion Gray) 
China, Burma and Nepal. Not common, but status and distribution not well 
known and further information required. Common name: Spectacle 
Swordtail (80, 251). Other refs: 52, 207. 

118 Graphium (Pazala) alebion (Gray, 1853) 

China and Taiwan. Status, exact distribution and nature of any threats not 
known. Two or more subspecies. Refs: 52, 141. 

119 Graphium (Pazala) tamerlanus (Oberthur, 1876) 

China (including Xizang Zizhiqu (Tibet)). Apparently not common, but 
species little known and more information required. Refs: 188, 250. 

Subgenus: Pathysa Reakirt 
Species-group: antiphates Cramer 

120 Graphium (Pathysa) aristeus (Cramer, 1775) 

Northern India (Assam and Sikkim), Burma, south-eastern China (includ- 
ing Hainan (Guangdong prov.)), Thailand, Laos, Vietnam, Kampuchea, 
Peninsular and Eastern Malaysia, Philippines, (possibly not Panay, Negros 
and Leyte), Indonesia (excluding Sulawesi, Sula,Bangka and Lombok(?)), 
Brunei, Papua New Guinea, Bismarck Archipelago (including New 
Britain) and Australia (northern Queensland) (52, 262). Collected for 
trade in Papua New Guinea (36). G. (P.) a. anticrates is protected by law in 
India (182). Nine subspecies (262). Common name: Five-bar Swordtail 
(282), Chain Swordtail (80). Other refs: 4, 40, 45, 48, 87, 131, 176. 

121 Graphium (Pathysa) nomius (Esper, 1798) 

Southern and eastern India (including Assam and Sikkim), Nepal, Sri 
Lanka, Bangladesh (?), Burma, Thailand, Laos (?), Vietnam (?) and 
Kampuchea (?) (52). Fairly common and not known to be threatened. Two 
subspecies. Common name: Spot Swordtail (80,251,282). Other refs: 
4, 259, 287. 

122 Graphium (Pathysa) rhesus (Boisduval, 1836) 

Indonesia (Sulawesi, Butung, Tanahjampea, Tukangbesi and Sula 
(52, 262)). Regarded as not uncommon and not threatened. This species 
appears to be the replacement species for G. (P.) aristeus in Sulawesi (52). 
Four subspecies. Other ref: 121. 

123 Graphium (Pathysa) dorcus (de Haan, 1840) 

Sulawesi (Indonesia). Not a well-known species, possibly confined mainly 
to mountainous areas. Although no threats are known, this species is a 
rarity and more information is required. Two subspecies. Refs: 

52,121,262. 

124 Graphium (Pathysa) androdes (Boisduval, 1836) 

Indonesia (Sulawesi and Sula (262)). Apparently not uncommon and not 
known to be threatened. Two subspecies. Other ref: 121. 

125 Graphium (Pathysa) epaminondas Oberthur, 1879 
Insufficiently Known — refer to section 6, p. 213. 

53 



Threatened Swallowtail Butterflies of the World 

Confined to the Andaman Is; distribution within the group poorly known. 
Ahhough not particularly rare in its chosen haunts, the butterfly has a very 
restricted range and requires further study before its status can be verified. 
It is sometimes treated as a subspecies of G. (P.)antiphates(5, 80, 188, 259) 
but seems to be distinct (250, 262). 

126 Graphium (Pathysa) euphrates (C. & R. Felder, 1862) 

Philippines (Luzon, Mindoro, Palawan and Balabac, possibly other islands 
too), Malaysia (only Banggi I., off Sabah) and Indonesia (Sulawesi, 
Halmahera and Obi). A rather disjunct distribution, but not uncommon. 
Not known to be threatened. Three or perhaps more subspecies 
(48,141,262). 

127 Graphium (Pathysa) decolor Staudinger, 1888 

Sabah (Eastern Malaysia) and the Philippines, including Palawan but 
excluding Panay , Cebu and Masbate (262) . Usually rare and local, except in 
Mindoro, where it is common in season. Not known to be threatened. Five 
subspecies (262). Often confused with G. (P.) euphrates, as by D'Abrera 
(52). 

128 Graphium (Pathysa) antiphates (Cramer, 1775) 

North-eastern and southern India, Nepal, Sri Lanka, Burma, south-eastern 
China (including Hainan (Guangdong prov.)), Thailand, Laos, Vietnam, 
Kampuchea, Peninsular and Eastern Malaysia, Brunei and Indonesia 
(Sumatra, Mentawai Is, Nias, Bangka, Bali, Lesser Sunda Is (except 
Sumba), Sulawesi, Bunguran (Natuna Is) and Kalimantan) (52, 191, 262). 
Not threatened as a species but the subspecies in Sri Lanka is apparently 
very rare (259). Twelve subspecies. Common name: Five-bar Swordtail 
(45, 80, 251). Other refs: 4, 87, 131, 141, 143, 251, 287. 

129 Graphium (Pathysa) agetes (Westwood, 1841) 

Northern India (Assam and Sikkim), Burma, Thailand, Laos, Vietnam, 
south-eastern China (including Guangdong prov.), Kampuchea (?), Pen- 
insular and Eastern Malaysia (Sarawak and Sabah) and Indonesia (Suma- 
tra, Kalimantan (?)) (52, 262). Widespread, often common and not 
threatened. Uncommon in Peninsular Malaysia; found on open hilltops 
above 1000 m. Five subspecies (262). Common name: Four-bar Swordtail 
(80). Other refs: 45, 87, 131, 259. 

130 Graphium (Pathysa) stratiotes (Grose-Smith, 1897) 

Eastern Malaysia (Sabah and Sarawak), possibly also Brunei and Kaliman- 
tan (Indonesia) (52, 262). Not particularly common but not recognized as 
threatened. Other refs: 131, 259. 

Species-group: macareus Godart 

131 Graphium (Pathysa) phidias (Oberthiir, 1896) 

Vietnam, Laos (?). Recorded from only a small area and possibly 
Vulnerable but very Httle information available. Refs: 52, 262. 

132 Graphium (Pathysa) encelades (Boisduval, 1836) 

Indonesia (Sulawesi). Not uncommon and not recognized as being threat- 
ened. Refs: 52, 262. 

133 Graphium (Pathysa) idaeoides (Hewitson, 1853) 
Rare — refer to section 6, p. 215. 

54 



Swallowtails of the world 

Philippines (Luzon, Samar, Leyte and Mindanao) (146). In view of the 
widespread habitat destruction in the Philippines, this species is a probable 
candidate for Vulnerable status in the future . A mimic of the sympatric but 
more widespread danaine Idea leuconoe. Much prized by Japanese and 
other collectors (274). Other refs: 52, 147, 148, 262. 

134 Graphium (Pathysa) delesserti (Guerin, 1839) 

Thailand, Peninsular and Eastern Malaysia, Philippines (Palawan), Brunei 
and Indonesia (Sumatra, Nias, Bangka, Bunguran, Java (probably extinct) 
and Kalimantan) (52, 262). Often common and no threats known although 
it appears to have become extinct in Java (45). If this species is extinct in 
Java it is likely to be due to loss of habitat ; deforestation is very extensive on 
this densely populated island. The extremely scarce female resembles 
Ideopsis gaura (Danainae) (45). Four subspecies. Common name: Zebra 
(251), Malayan Zebra (45). Other refs: 87, 131. 

135 Graphium (Pathysa) xenocles (Doubleday, 1842)(= leucothoe Westwood) 
Northern India (Assam and Sikkim), Nepal, Bhutan, Burma, Thailand, 
Laos, Vietnam and China (Hainan (Guangdong prov.)) (52). Apparently 
common and not threatened. Four subspecies. Common name: Great 
Zebra (80), Greater Zebra (251). Other refs: 4, 259. 

136 Graphium (Pathysa) macareus (Godart, 1819) 

Northern India (Assam, Sikkim and Manipur), Nepal, Burma, Thailand, 
Laos, Vietnam, Kampuchea, China (Hainan (Guangdong Prov.)), Pen- 
insular and Eastern Malaysia, Brunei, Philippines (Palawan) and Indonesia 
(Sumatra, Java, Bali and Kalimantan) (52, 262). Relatively common and 
not threatened. Eleven subspecies (52, 262). Common name: Lesser Zebra 
(80, 251). Other refs: 45, 87, 131, 259. 

137 Graphium (Pathysa) rumaceus (Westwood, 1872) 

Thailand, Peninsular and Eastern Malaysia, Brunei and Indonesia (Suma- 
tra and Kalimantan) (52,262). Uncommon (45), but not rare and not 
thought to be threatened. Three subspecies. Common name for ssp. G. (P.) 
r. pendleburyi: Pendlebury's Zebra (45). Other ref: 87. 

138 Graphium (Pathysa) megarus (Westwood, 1841) 

Northern India (Assam and Sikkim), Burma, Thailand, Laos, Vietnam, 
south-eastern China (including Hainan (Guangdong prov.)), Kampuchea, 
Peninsular and Eastern Malaysia, Brunei and Indonesia (Sumatra, Java, 
Bah and Kalimantan) (52, 262). Rather common and not considered to be 
threatened, although the nominate subspecies is protected by law in India 
(182). Eight subspecies. Common name: Spotted Zebra (80,282). Other 
refs:45, 87, 131,259. 

139 Graphium (Pathysa) megaera (Staudinger, 1888) 
Indeterminate — refer to section 6, p. 217. 

Philippines (Palawan). This species merits Indeterminate status because of 
its restricted range in a habitat that is being increasingly disrupted. Refs: 
52, 262. 

140 Graphium (Pathysa) stratocles (C. & R. Felder, 1862) 

Phihppines (Luzon, Mindoro, Marinduque, Bohol, Mindanao, Calamians 
and Palawan) (52, 262). Not uncommon and not known to be threatened. 
Three subspecies. 

55 



(pyUy^' 



Threatened Swallowtail Butterflies of the World 

141 Graphium (Pathysa) deucalion (Boisduval, 1836) 
Indonesia (Sulawesi, Moluccas (Halmahera, Bacan and Ternate) and Irian 
Jaya (Biak)(115)). Not known to be threatened. Two (262) or three (104) 
subspecies. Other refs: 48, 52. 

142 Graphium (Pathysa) thule (Wallace, 1865) 
Indonesia (Irian Jaya) and Papua New Guinea. Certainly not common, but 
not known to be threatened. Rarely collected but possibly overlooked as it 
mimics the common danaines Ideopsis juventa and Tirumala hamata. One 
subspecies with three forms (104). Other ref: 48. 

Subgenus: Arisbe Hiibner 
Species-group: angolanus Goeze 

143 Graphium (Arisbe) endochus (Boisduval, 1836) 
Madagascar only. Apparently well distributed and not currently threat- 
ened, but it is a forest species and its status in a country which is being 
rapidly deforested should be carefully monitored. Refs: 33, 34, 50, 
204,206,285. 

144 Graphium (Arisbe) angolanus (Goeze, 1779) (= pylades F., 1793) 
^f Southern and tropical Africa: distributed in woodland over most of the 

(^SdtA^ continent. Common, often abundant and not thieatened. Three subspe- 
^^ ':i'le- <^i^s- Common names: Angola White Lady Swallowtail (282, 285), White 

^ Lady Swallowtail (34, 44, 214). Other refs: 20, 33 50, 63, 97 165. See note 

after References. «'^ U^^t. *, t^ cr u^^ St^ tt^O 1=1^. f^(^^k 

145 Graphium (Arisbe) taboranus (Oberthiir, 1886) 
Zaire, Tanzania, Angola, Zambia, Malawi and Namibia. Not uncommon 
and not known to be threatened. Two subspecies. Common name: Tabora 
Swallowtail (215,285). Other refs: 33,34,50,63,97, 165. See note after 
References. 

146 Graphium (Arisbe) morania (Angas, 1849) 
South Africa (Cape Province, Natal and Transvaal), Namibia, Zimbabwe, 
southern Malawi and Mozambique. Not uncommon and not threatened. 
Common name: White Lady Swallowtail (285), Lesser White Lady 
Swallowtail (214). Other refs: 34, 50, 63, 115. 

Species-group: ridleyanus White 

147 Graphium (Arisbe) ridleyanus (White, 1843) 
Lowland forest in West Africa: Sierra Leone, Liberia, Ivory Coast, Ghana, 
Togo, Benin, Nigeria, Cameroon, Central African Republic (?), Gabon, 
Congo, Zaire, Uganda, western Tanzania, Zambia and Angola. Rather 
widely distributed and not known to be threatened. Common names: 
Acraea Swallowtail (282, 285), Red Graphium (34), Ridley's Swallowtail 
(215). Other refs: 20, 33, 50, 115. 

Species-group: philonoe Ward 

148 Graphium (Arisbe) philonoe (Ward, 1873) 
Forest and woodland in East Africa: Kenya, Tanzania, southern Sudan, 
northern Uganda, south-west Ethiopia, Mozambique and Malawi. 

56 



Swallowtails of the world 

Common in much of its range and not threatened. Two subspecies. 
Common names: White-dappled Swallowtail (285), Eastern Graphium 
(34). Other refs: 33,50,97. 

Species-group: adamastor Boisduval 

149 Graphium (Arisbe) almansor (Honrath, 1884) 

Cameroon, Central African Republic, Sudan, Ethiopia, Equatorial Guinea 
(?), Gabon, Congo, Zaire, Uganda, western Kenya, Rwanda (?), Burundi 
(?), north-west Tanzania, Angola and Zambia. Uncommon and local, 
though widely distributed. Not threatened. Four subspecies. Refs: 
33,34,50,151,285,291. 

150 Graphium (Arisbe) poggianus (Honrath, 1884) 

Angola, Zaire and Zambia (115). Given as a form of G. (A.) almansor by 
Berger (16) but accepted as a full species by Hancock (109), under the name 
carchedonius Karsch. The name poggianus follows Hancock (115). Uncom- 
mon but no evidence of being threatened. See note after References. 

151 Graphium (Arisbe) adamastor (Boisd. 1836) (= carchedonius Karsch) 
Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin and 
Nigeria; also a subspecies in Central African Republic and northern Zaire 
(? disjunct range). Uncommon and local but not threatened. Two subspe- 
cies. Refs: 20,33,34,50,285. 

152 Graphium (Arisbe) agamedes (Westwood, 1842) 

Lowland forest in Ghana, Togo, Benin, Nigeria, Cameroon, Central 
African Republic, Equatorial Guinea (?), Gabon (?), northern Congo and 
northern Zaire. Local and uncommon but not regarded as threatened. 
Common name: Glassy Graphium (34). Other refs: 20, 33, 50, 285. 

153 Graphium (Arisbe) aurivilliusi (Seeldrayers, 1896) 
Insufficiently Known — refer to section 6, p. 219. 

West Africa: Type specimen labelled "Congo" but probably present-day 
Zaire (34). Known only from the type series and possibly not specifically 
distinct (form of agamedes ? (250)), but recognized by Carcasson (34). 
Other refs: 33,50,285. 

154 Graphium (Arisbe) olbrechtsi (Berger, 1950) 

Zaire. Uncommon and scarce. This recently-described species may deserve 
Rare status when more information becomes available. Two subspecies. 
Refs: 33, 34, 50, 115, 285. See note after References. 

155 Graphium (Arisbe) odin (Strand, 1910) 

Cameroon, Congo, Central African Republic and Zaire. Not common but 
not threatened. Three (possibly only two) subspecies. Refs: 34, 50, 285. See 
note after References. 

156 Graphium (Arisbe) auriger (Butler, 1877) 

West Africa: Gabon. Uncommon but not known to be threatened. More 
information needed. Refs: 34, 50, 285. See note after References. 

157 Graphium (Arisbe) hachei (Dewitz, 1881) 

Lowland forest in Cameroon, Equatorial Guinea (?), Central African 
Republic, Gabon, Congo, western Zaire and Angola. Uncommon, but not 

57 



Threatened Swallowtail Butterflies of the World 

threatened. Two or three subspecies. Common name: Milky Graphium 
(34). Other refs: 33, 50, 285. 

158 Graphium (Arisbe) weberi (Holland, 1917) 
Insufficiently Known — refer to section 6, p. 220. 

Cameroon. Known only from the type specimen in Pittsburgh, U.S.A. (50). 
More information required. Common name: Weber's Swallowtail (285). 
Other refs: 33, 34. See note after References. 

159 Graphium (Arisbe) fulleri (Grose-Smith, 1883) 

West Africa: Cameroon, Gabon and the Congo. Possibly a form of G. (A.) 
ucalegonides, but regarded as a full species by Carcasson (34). Uncommon, 
but not known to be threatened. Common name: Fuller's Swallowtail (270). 
Other refs: 33, 50, 285. See note after References. 

160 Graphium (Arisbe) ucalegonides (Staudinger, 1884) 

Ghana, Togo, Benin, Nigeria, Cameroon, Central African Republic, 
Equatorial Guinea (?), Gabon, Congo, Zaire and Angola. Not uncommon 
and not threatened. Two subspecies. Refs: 34, 50, 285. See note after 
References. 

161 Graphium (Arisbe) ucalegon (Hewitson, 1865) 

Lowland forest in Nigeria, Cameroon, Central African Republic (?), 
Gabon, Congo, Zaire, Uganda, Rwanda (?), Burundi (?), Tanzania and 
Angola. Not uncommon and not considered to be threatened. Two 
subspecies. Common name: Creamy Graphium (34). Other refs: 
20,33,50,151,285. 

162 Graphium (Arisbe) simoni (Aurivillius, 1899) 

Cameroon, Gabon, Congo and Zaire. Possibly a form of G. (A.) ucalegon 
(250), ahhough regarded as a full species by Carcasson (34). Uncommon 
but not known to be threatened. Common name: Simon's Swallowtail 
(285). Other refs: 33, 50. 

Species-group: leonidas Fabricius 

163 Graphium (Arisbe) cyrnus (Boisduval, 1836) 

Madagascar. Apparently common throughout the island. Not known to be 
threatened. Two subspecies (206), not recognized by D'Abrera (50). Other 
refs: 33, 34. 

164 Graphium (Arisbe) leonidas (Fabricius, 1793) 

Throughout tropical and southern Africa, common in forest, woodland and 
gardens but absent from dry and montane areas. Not threatened. Three 
subspecies, two on small islands and so very restricted in range. Larva now 
generally found on the introduced Custard Apple (Annona reticulata 
(Annonaceae)), but this is presumably not the original foodplant. Sexes 
similar. Common names: Veined Swallowtail (44,285), Common 
Graphium (34). Other refs: 20, 33, 50, 63, 97, 214, 215. 

165 Graphium (Arisbe) pelopidas (Oberthiir, 1879) 

Tanzania (Pemba Island and possibly Zanzibar). Recognized as a species by 
D'Abrera (50) but not by other authors, including Munroe (188), Carcasson 
(34), Smart (250) and Hancock (109, 113). Should possibly be regarded as a 
subspecies of G. (A.) leonidas as are the populations on Sao Tome and 

58 



Swallowtails of the world 

Principe in the Gulf of Guinea. Status of Zanzibar population particularly 
uncertain. Should the Pemba Island population be considered as a full 
species it would certainly be given Rare or even Vulnerable status. Pemba 
Island is largely covered with clove plantations but natural habitat may still 
be found in the Ngezi forest, north of Pemba (264). 

166 Graphium (Arisbe) levassori (Oberthiir, 1890) 
Vulnerable — refer to section 6, p. 222. 

Comoros (Grande Comoro) only (205). Range very circumscribed and 
shrinking. Other refs: 33, 34, 50, 206. 

Species-group: tynderaeus Fabricius 

167 Graphium (Arisbe) tynderaeus (Fabricius, 1793) 

Lowland forest in West Africa: Sierra Leone, Liberia, Ivory Coast, Ghana, 
Togo, Benin, Nigeria, Cameroon, Central African Republic (?), Gabon 
(?), Congo and Zaire. Widely distributed, not uncommon and no threats 
recognized. Common name: Green-spotted Swallowtail (285). Other refs: 
20, 34, 50. 

168 Graphium (Arisbe) latreillanus (Godart, 1819) 

Lowland forest in West Africa: Sierra Leone, Liberia, Ivory Coast, Ghana, 
Togo, Benin, Nigeria, Cameroon, Central African Republic (?), Gabon 
(?), Congo, Zaire, Uganda and Angola. Not threatened. Two subspecies. 
Common names: Coppery Swallowtail (285), Olive Graphium (34). Other 
refs: 20, 33, 50. 



Subgenus: Graphium Scopoli 
Species-group: policenes Cramer 

169 Graphium (Graphium) junodi (Trimen, 1893) 

Restricted to Mozambique. There has also been a single record from 
eastern Zimbabwe but this was probably a vagrant (115). Uncommon or 
scarce, but not known to be threatened. Common names: Mozambique 
Swallowtail (285), Junod's Swordtail (44), Scarce Swordtail (44). Other 
refs: 33,34,50.214. 

170 Graphium (Graphium) nigrescens (Eimer, 1889) 

Cameroon , Gabon and Zaire. Species of uncertain status (50) but said to be 
rare (285). More information required. Common name: Dusky Swordtail 
(285). Other refs: 33, 34. 

171 Graphium (Graphium) polistratus Grose-Smith, 1889 

East Africa, largely coastal from southern Somalia southwards to Delagoa 
Bay, i.e. Somalia, Kenya, Tanzania, Malawi and Mozambique. Not 
particularly common but no threats known. Graphium sisenna (Mabille, 
1890) was listed by Munroe (188) but recognised as a synonym of G. (G.) 
polistratus by Hancock (109). Common name: Dancing Swordtail (285), 
Dusky Swordtail (214). Other refs: 33, 34, 50, 63. 

172 Graphium (Graphium) policenes (Cramer, 1775) 

Tropical and southern Africa, widely distributed. Common in woodland 
and forest and not threatened. Most authors do not accept described 
subspecies to be valid (33, 50, 97). Larvae feed on Artobotrys 

59 



Threatened Swallowtail Butterflies of the World 

(Annonaceae). G. (G.) boolae was shown to be an aberration of G. (G.) 
policenes (41). Common names: Small-striped Swordtail (285), Small 
Striped Swordtail (214), Common Swordtail (34). Other refs: 20, 63. 

Species-group: porthaon Hewitson 

173 Graphium (Graphium) porthaon (Hewitson, 1865) 

East Africa, fairly widely distributed in woodland in western Transvaal 
(South Africa), Mozambique, Botswana, Zimbabwe, Malawi, Zambia, 
Zaire, Tanzania and eastern Kenya. Apparently not uncommon and no 
threats known. Two subspecies of which G. (G.) p. tanganyikae is rare in 
Tanzania (34, 151). Common names: Cream-striped Swordtail (285), 
Cream Swordtail (34), Pale Spotted Swordtail (214). Other refs: 
33,50,63,97,165. 

Species-group: illyris Hewitson 

174 Graphium (Graphium) illyris (Hewitson, 1873) 

Lowland forest in Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, 
Togo, Benin, Nigeria, Cameroon, Gabon (?), Congo and Zaire (34). Not 
uncommon but not known to be threatened. Common names: Yellow- 
banded Swordtail (285), Cream-banded Swordtail (34). Other refs: 
20,33,50,103. 

175 Graphium (Graphium) gudenusi (Rebel, 1911) 

Eastern Zaire, south-western Uganda, Rwanda and Burundi, a rather 
restricted range in highland forests (263). Scarce and local, but not 
threatened at present. However, human population pressure on some parts 
of its range, e.g. the highlands of Rwanda, is increasing and the species 
needs further study and monitoring. Common name: Kigezi Swordtail 
(285). Other refs: 33,34,50. 

176 Graphium (Graphium) kirbyi (Hewitson, 1872) 

East Africa: Kenya and Tanzania. Local but sometimes common in coastal 
forests and not threatened. Common name: Kirby's Swordtail (34, 285). 
Other refs: 33,50. 

Species-group: colonna Ward 

177 Graphium (Graphium) colonna (Ward, 1873) 

East Africa, coastal woodlands and forests in South Africa (Natal, 
Transvaal (?)), Swaziland (?), Mozambique, Malawi, Tanzania, Kenya, 
Somalia and southern Ethiopia (34) . Not known to be threatened. Common 
names: Mamba Swordtail (285), Black Swordtail (34). Other refs: 
33,50,63,214. 

Species-group: antheus Cramer 

178 Graphium (Graphium) antheus (Cramer, 1779) 

Most of continental Africa. Common and not in danger. G. (G.) mercutius 
Grose-Smith and Kirby, is not listed by Munroe (188), D'Abrera (50), 
Carcasson (34) and most other authors and is only a form of G. (G.) antheus 
(115). Common name: Large Striped Swordtail (44,285). Other refs: 
20,33,63,97,165,214,215. 

60 



Swallowtails of the world 



179 Graphium (Graphium) evombar (Boisduval, 1836) 

Madagascar, distributed over the whole island. Common and not threat- 

ened. Refs: 33, 34, 50, 206. ''"^ Co^>^-<^ '^ -- - L^y^^^ ^^c <.?(.) o ( 

Species-group: eurypylus Linnaeus 

180 Graphium (Graphium) arycles (Boisduval, 1836) 

Southern Burma, Thailand, Kampuchea (?), Peninsular and Eastern 
Malaysia, Philippines (Palawan), Brunei and Indonesia (Sumatra, Bangka, 
Java and Kalimantan) (52,262). Scarce but not known to be threatened 
although the nominate subspecies is protected in India (182). Three (262) or 
four (52) subspecies. Common name: Spotted Jay (80). Other refs: 
45,87,231. 

181 Graphium (Graphium) bathycles (Zinken-Sommer, 1831) 

Southern Thailand (?), Peninsular and Eastern Malaysia, Philippines 
(Palawan), Brunei and Indonesia (Sumatra, Java and Kalimantan) 
(52, 262). Not rare and not threatened. Two subspecies, a third, chiron, 
having recently been raised to the specific level (52,262) (see below). 
Common name: Veined Jay (111). Other refs: 45, 80, 87, 131, 231, 259. 

182 Graphium (Graphium) chiron (Wallace, 1865) 

Nepal, northern India (Assam), Burma, southern China, Thailand, Laos, 
Vietnam, Kampuchea and Peninsular Malaysia (76). Formerly regarded as 
a subspecies of G. (G.) bathycles but raised to specific level by Saigusa et at. 
(237). There is doubt concerning the validity of the name chiron. Hancock 
(115) considers it to be a homonym requiring replacement. Both chironides 
Honrath, 1884 and clanis (Jordan, 1909) are available and Hancock 
suggests the latter as being more appropriate, chironides having been used 
as an infrasubspecific title. We do not presume to alter the nomenclature 
here, but adopt the most recent published judgement. Three subspecies. 
Common name: Veined Jay (80). Other refs: 52, 109, 259, 262. 

183 Graphium (Graphium) leechi (Rothschild, 1895) 

China. Little information available and not known to be threatened. Ref: 

188. 

184 Graphium (Graphium) procles (Grose-Smith, 1887) 
Indeterminate — refer to section 6, p. 224. 

Montane forest in Eastern Malaysia (Sabah). Considered to be a good 
species by D'Abrera (52), Tsukada and Nishiyama (262) and Robinson 
(231) (who notes its similarity to G. doson) Other ref: 131. 

185 Graphium (Graphium) meyeri (Hopffer, 1874) 

Indonesia (Sulawesi and Sula) (262). Not a well-known species, but not 
considered to be threatened. Two subspecies (52). 

186 Graphium (Graphium) eurypylus (Linnaeus, 1758) 

North-eastern India, Bangladesh, Burma, Thailand, Andamans, Nicobar 
Is, southern China (including Hainan (Guangdong prov.)), Vietnam, Laos, 
Kampuchea, South East Asia to Papua New Guinea and north-eastern 
Australia (52, 262). Collected for trade in Papua New Guinea (36). G. (G.) 
e. macronius is protected by law in India (182). Not threatened. About 
twenty subspecies (262). Common name: Great Jay (80), Pale Green 
Triangle (282). Other refs: 5, 14, 40, 45, 48, 87, 176, 259. 

61 



Threatened Swallowtail Butterflies of the World 

187 Graphium (Graphium) doson (C. & R. Felder, 1864) 

Northern and southern India, Nepal, Sri Lanka, Bangladesh, Burma, 
Thailand, south-east China (including Hainan (Guangdong prov.)), Tai- 
wan, Japan, Vietnam, Laos, Kampuchea, Malaysia, Brunei, Indonesia 
(Sumatra, Nias, Mentawai Is, Bangka, Java, Bali, Lombok, Sumbawa, 
Bawean and Kalimantan) and the Philippines (including Palawan) 
(52, 191,262)). Widespread and often common, but scarce in Honshu, 
Japan, where it only occurs in the southern part (82). Fourteen subspecies 
(262). Common name: Common Jay (45,80,251). Other refs: 87,141, 
149, 246, 259. 

188 Graphium (Graphium) evemon (Boisduval, 1836) 

North-eastern India (Assam), Burma, Thailand, southern China, Laos, 
Vietnam, Kampuchea, Malaysia, Brunei and Indonesia (Sumatra, Simeu- 
lue, Nias, Mentawai Is, Java and Kalimantan) (52,191,262). Fairly 
common and not threatened but one subspecies, G. (G.) e. albociliates is 
protected by law in India (182). Seven subspecies (262). Common name: 
Lesser Jay (3, 80). Other refs: 45,87,131,231,259. 

Species-group: agamemnon Linnaeus 

189 Graphium (Graphium) agamemnon (Linnaeus, 1758) 

Southern and northern India, Nepal, Sri Lanka, Andamans, Nicobar, 
Bangladesh, Burma, Thailand, Laos, Kampuchea, southern China (includ- 
ing Hainan (Guangdong prov.)), Taiwan, South East Asia to Papua New 
Guinea, Bougainville, Solomon Is and Australia (northern Queensland) 
(52, 262). Farmed and collected for trade in Papua New Guinea (36, 68). 
Range similar to that of G. (G. ) sarpedon but not reaching Japan. Common 
and not threatened. Twenty or twenty-one subspecies. Common names: 
Tailed Jay (45, 80, 251), Tailed Green Jay, Green-spotted Triangle (282). 
Other refs: 5, 14, 40, 48, 87, 141, 176, 191, 220, 231, 246, 287. 

190 Graphium (Graphium) macfarlanei (Butler, 1877) 

Indonesia (Buru, Obi, Halmahera, Batjan, Seram (?) and Irian Jaya), 
Papua New Guinea, Bismarck Archipelago (including New Britain) and 
north-eastern Australia. Farmed and collected for trade in Papua New 
Guinea (36,68). Not threatened. Three subspecies. Other refs: 
14,40,48, 141,176. 

191 Graphium (Graphium) meeki (Rothschild and Jordan, 1901) 
Rare — refer to section 6, p. 226. 

Solomon Is (Santa Isabel (171) and Choiseul) and Papua New Guinea 
(Bougainville)(219). Despite D'Abrera's statement that it is probably 
extinct (48), examples are sent in from time to time to the Bulolo Agency 
(36). The species is little known and may be classed as Rare until more 
details are available. A second subspecies G. (G.) meeki inexpectatum was 
described from Bougainville in 1981 (181). Other ref: 142. 

Species-group: sarpedon Linnaeus 

192 Graphium (Graphium) gelon Boisduval, 1859 

New Caledonia (including Loyalty Is). Restricted range, but stated to be 
'not rare' (48) and not recognizably threatened. Other ref: 133. 

62 



Swallowtails of the world 

193 Graphium (Graphium) macleayanum (Leach, 1814) 

Australia including Tasmania, and Lord Howe and Norfolk Is. Recently 
recorded from mainland New Guinea (48). Not threatened as a species. 
Four subspecies. Common name: Macleay's Swallowtail or Swordtail 
(282). Other refs: 14, 95, 141, 176, 289. 

194 Graphium (Graphium) weiskei (Ribbe, 1900) 

All of New Guinea (Indonesia (Irian Jaya) and Papua New Guinea) (48) in 
suitable habitat. Very common in some areas although the foodplant is still 
unknown. It is collected for trade but this is occasionally halted for periods 
of up to one year (36). It is not known to be threatened. Common name: 
Purple-spotted Swallowtail (282). Other ref: 120. 

195 Graphium (Graphium) stresemanni Rothschild, 1916 
Rare — refer to section 6, p. 228. 

Known only from the island of Seram in the Moluccas (Indonesia) , where it 
is Rare (48) . Not listed by Munroe (188) as a species but recognized by most 
other authors. Other refs: 48, 196. 

196 Graphium (Graphium) codrus (Cramer, 1776) 

Phlippines (not Palawan, Calamian Is, or Sulu Is), Indonesia (Sulawesi, 
Moluccas (including Sula, Buru, Ambon and Seram but not Obi) and Irian 
Jaya (including Waigeo, Biak, Kai and Aru), Papua New Guinea, Bismarck 
Archipelago (including New Britain and the Admiralty Is), Bougainville 
and the Solomon Is (not Rennell) (262). It is collected for trade in Papua 
New Guinea and G. (G.) c. auratus is farmed on Manus I. (36). Some local 
populations may be threatened but the species is not. Fourteen subspecies. 
Other refs: 48, 52, 68, 127, 141, 181, 218. 

197 Graphium (Graphium) empedovana (Corbet, 1941)(= empedocles F., 
1787) 

Peninsular and Eastern Malaysia, Philippines (Palawan), Indonesia (Suma- 
tra, Bangka, Java and Kalimantan) and Brunei (262). Sometimes regarded 
as a subspecies of the Papuan G. (G.jco(/n« (45, 52). Uncommon generally 
and possibly Vulnerable in Malaysia (10). G. (G.) empedocles is regarded 
by some as a senior synonym of G. (G. ) empedovana (262) . Not known to be 
threatened over most of its range. Other ref: 87. 

198 Graphium (Graphium) cloanthus Westwood, 1841 

Disjunct range: northern India, Nepal (251), Bhutan, northern Burma, 
southern and central China, Taiwan, northern Thailand and Indonesia 
(Sumatra)(52. 262). Common and not threatened. Three subspecies includ- 
ing G. (G.) clymemts Leech, a little known form from central and western 
China, but not including G. (G.) sumatranum (see below) (52, 141, 259). 
Common name: Glassy Bluebottle (80, 251). 

199 Graphium (Graphium) sumatranum (Hagen, 1894) 

Indonesia (Sumatra). Recognized as a subspecies of G. (G) cloanthus by 
most authors (52, 262) but given species rank by Hancock (109) on the 
strength of its isolated distribution and morphological differences. No 
known threats, but more information needed. 

200 Graphium (Graphium) monticolum (Fruhstorfer, 1897) 

Indonesia (southern Sulawesi). Regarded by some as a subspecies of G. 
(G.) sarpedon {12\ , 262) , but by others as a full species (52, 109). Graphium 

63 



Threatened Swallowtail Butterflies of the World 

sarpedon textrix Tsukada and Nishyama, 1980, belongs to this species. No 
threats are known, although more information is needed. 

201 Graphium (Graphium) milon C. & R. Felder, 1864 

Indonesia (Sulawesi (including Banggai) and Moluccas: Sula, Buru, 
Ambon and Seram but not Morotai) (262). Status revised by Murayama in 
1978 (190) but not accepted as a good species by Hancock (109). Not known 
to be threatened. Six subspecies (262). 

202 Graphium (Graphium) sandawanum Yamamoto, 1977 
Vulnerable — refer to section 6, p. 231. 

A recent discovery (1977) , known only from the Philippines (Mindanao: Mt 
Apo) (52, 262). Other ref: 289. 

203 Graphium (Graphium) sarpedon Linnaeus, 1758 

Southern India, Sri Lanka, northern India, Nepal, Bangladesh (?), Burma, 
south-eastern and western China (including Hainan (Guangdong prov.)), 
Taiwan, South Korea (?), Japan, Thailand, Laos, Kampuchea, Malaysia, 
Philippines, Brunei, Indonesia (not Moluccas) , Papua New Guinea (includ- 
ing New Britain), Bismarck Archipelago, Solomon Is, Vanuatu (New 
Hebrides) (102) and north-eastern Australia (52, 262). Very common and 
not threatened. Widely collected for trade. Sixteen recognized subspecies 
(262) not including milon and monticolum which are regarded as separate 
species (52, 109, 262) or textrix which is a synonym, of monticolum (52). G. 
(G.) protensor Gistel, is regarded as a synonym of G. (G.) sarpedon (109). 
Common names: Blue Triangle (282), Common Bluebottle (45, 80, 251). 
Other refs: 4, 14, 36, 40, 48, 87, 131, 141, 149, 176, 207, 220, 259. 

Species-group: mendana Godman and Salvin 

204 Graphium (Graphium) mendana (Godman and Salvin, 1888) 
Rare — refer to section 6, p. 234. 

Papua New Guinea (Bougainville) and Solomon Is. A little-known species 
though specimens are occasionally sent to the Bulolo Insect Trading 
Agency from Bougainville (36). Four subspecies. Other refs: 48, 99, 
181,218. 

Species-group: wallacei Hewitson 

205 Graphium (Graphium) wallacei (Hewitson, 1859) 

Indonesia (Moluccas (possibly not Buru or Seram), Aru and Irian Jaya), 
and Papua New Guinea where it is collected for trade (36). Not threatened. 
Two subspecies. Other ref: 48. 

206 Graphium (Graphium) hicetaon (Mathew, 1886) 

Solomon Is. Not rare (48) and not known to be threatened. Other refs: 
219, 220. 

207 Graphium (Graphium) browni (Godman and Salvin, 1879) 

Bismarck Archipelago (including New Britain) . Fairly restricted range , but 
not rare (48) and not known to be threatened. 

Subfamily: PAPILIONINAE 
Tribe: Troidini 

Genus: Battus Scopoli 
64 



u 



1/ 



Swallowtails of the world 

Species-group: philenor Linnaeus 

208 Battus philenor {VAnmieus, 1771) 
Southern Canada, U.S.A. and Mexico. Not uncommon and not known to 
be threatened. Two subspecies (55). Common names: Pipevine Swallowtail 
(154, 216), Blue Swallowtail (216). Other refs: 62, 137, 141, 234, 269. 

209 Battus zetidesy[\ix\xoe,\91\ 
Vulnerable — refer to section 6, p. 236. 

Known only from high elevations in Haiti and the Dominican Republic. 
"Apparently very rare" (230), but "locally common under the right 
conditions and at the right times of year" (243). Further information 
needed. Originally described as zetes (Westwood, 1847) (230,234). 
Common name: Zetides Swallowtail (230). Other refs: 51, 55, 139. 

210 Battus devilliers (Godan, 1823) 
Cuba and the Bahamas: Andros I. (230, 234). Not known to be threatened 
in Cuba, but the population on Andros is rare and precarious (268). Status 
needs clarification and careful monitoring. Very high prices demanded by 
dealers for this species. Common name: Devilliers Swallowtail (230). Other 
refs: 51,55. 

Species-group: polydamas Linnaeus 

211 Battus polydamas (hmnaexx^, 1758) 
Southern U.S.A., Mexico, Central America, South America, Antigua, 
Bahamas, Cuba, Dominica, Dominican Republic, Grenada, Guadeloupe, 
Haiti, Jamaica, Martinique, Puerto Rico, St. Kitts, St. Lucia, Trinidad and 
Tobago. Very common and not threatened; often associated with disturbed 
habitats, rarely entering the forest (61 ). Well differentiated local races (51). 
Thirteen or fourteen subspecies. Common names: Polydamas Swallowtail 
(154,216,230), Gold Rim (9,216), Black Page (9,282). Other refs: 
21,26,55,62,137,234,269. 

212 Battus streckerianiis {WonraXh, 1884) 

Peru; uncommon, but no threats known. Refs: 51, 55, 234. 

< M3 fifl««5 fl/-c/j/rfflmfl5 (Boisduval, 1836) 

^p^ Coastal areas of central Chile. Appears to be common and not threatened. 
A male labelled Mendoza, Argentina, was described in 1925 by Ehrmann as 
the type specimen of Papilio lindeni (51). Other refs: 55, 234. 

214 Battus polystictus (Butler, 1874) 
Brazil, Paraguay and Argentina. Fairly widely distributed, not uncommon 
and not thought to be threatened. Ranched in Brazil (24). Two subspecies. 
Other refs: 51,55,122,234. 

215 Battus philetas (Hewitson, 1869) 
Northern Peru and Ecuador. Not uncommon and not recognized as 
threatened. Two subspecies. Refs: 51, 55, 234. 

216 Battus madyes (Doubleday, 1846) 
Peru, Bolivia and Argentina. Not uncommon and not known to be 
threatened. Up to six subspecies. Refs: 51, 55, 234. 

65 



r 





Threatened Swallowtail Butterflies of the World 

111 Battus eracon (Godman and Salvin, 1897) 

Western Mexico. Local and uncommon, but not known to be threatened. A 
possible candidate for listing because of its restricted range. Further study 
and reserarch are necessary. Refs: 51, 55, 234, 269. 

218 Battus belus (Cramer, 1777) 

Mexico, Central America, Trinidad, Colombia, Venezuela, Guyana, 
Surinam, French Guiana, Ecuador, Peru, Brazil and Bolivia. Apparently 
not threatened, but is encountered as solitary individuals in humid forest 
and is rarely abundant (at least in Costa Rica) (61). Five or six subspecies; 
B. b. chalceus is only known from the type collection (174). Common name: 
the Belus (9). Other refs: 51, 55, 234, 269. 

219 Battus crassus (Cramer, 1777) 
Costa Rica, Panama (?), Colombia, Venezuela, Guyana, Surinam, French 
Guiana (?), Ecuador, Peru, Bolivia, Brazil, northern Argentina and 
possibly Paraguay. Widespread and not threatened, but solitary and rarely 
collected (61). Other refs: 26, 51, 55, 234. 

220 Battus laodamas (Felder, 1859) 
Mexico, Central America and Colombia. Little known but not threatened. 
Extremely local, but fairly common in places (61). Four subspecies. Other 
refs: 51, 55, 57, 62, 234, 269. 

221 Battus lycidas (Cramer, 1777) 
Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Trinidad, 
Colombia, Venezuela, Ecuador, Surinam, Peru, Bolivia and Brazil. Little 
known but not threatened. Very rare in Costa Rica (61). Common name: 
Lycidas Swallowtail (9, 282). Other refs: 51, 55, 150, 234. 

Genus: Euryades C. & R. Felder 

222 Euryades duponchelii (Lucas, 1839) 
Woodlands in southern Brazil, Paraguay, Argentina and Uruguay (51). 
Less common than E. corethrus but not known to be threatened. Very high 
prices demanded by dealers for this species. The two species belonging to 
the genus Euryades are closer to the Australasian Cressida than to other 
Neotropical Aristolochia feeders (51). Other ref: 55. 

Euryades corethrus (Boisduval, 1836) 

Pampas in southern Brazil, Paraguay, Argentina and Uruguay. Not 
uncommon but not known to be threatened. Very high prices demanded by 
dealers for this species. Refs: 51, 55, 141. 

Genus: Cressida Swainson 

224 Cressida cressida (F., 1775) (= heliconides Swainson, 1832) 

Indonesia (Lesser Sunda Is: Timor, Leti, Moa, Babar, Wetar and Tanimbar 
Is (?)) , eastern Papua New Guinea and Australia (north Northern Territory 
and north and eastern Queensland) (262). Generally common and not 
threatened. Five subspecies. Common name: Big Greasy Butterfly (282). 
Other refs: 14, 40, 48, 141, 176. 



66 



x^(^r\\^ ^^^ 5wfl//owrai75 of the world 

y^enus: Parides Hiibner 
HyV^^ Species-group: ascanius Cramer 

225 Parides gundlachianus (C. & R. Felder, 1864) 

/-^^ Restricted to forests in Cuba. Not exceptionally rare but requires moni- 
L^' toring. Parides columbus (Herrich-Schaeffer, 1862) is a homonym. 

Common name: Gundlach's Swallowtail (230). Other refs: 51, 55, 234. 

226 Parides alopiiis (Godman and Salvin, 1890) 

According to standard works this species is found in western Mexico, 
Guatemala, Honduras and Nicaragua (51,55,61,62,234), but recent 
unpublished work by Racheli shows it to be a Mexican endemic (221). The 
position should be clarified as soon as possible. It is certainly uncommon 
(269) and known from very few localities, but is not known to be threatened. 



^ 



111 Parides photinus (Douhleday, 1844) 

Mexico, Guatemala, Honduras, Nicaragua and Costa Rica. In many places 
abundant (269); generally common and not threatened. In Costa Rica 
moderately common in remnant patches of forest, 300-800 m (61). Life 
cycle known and described (61). Parides dares (Hewitson, 1869) from 
Nicaragua may be a hybrid, form (250), aberration (109), or synonym (268) 
of Parides phodnus. Other refs: 51, 55, 62, 234. 



228 Parides montezuma (Westwood, 1842) 

Mexico, Guatemala, Honduras, Nicaragua and Costa Rica. Occurs from 
sea level to 700 m in dry forest habitats (61) . Generally fairly common north 
of Costa Rica, but rare or locally common within Costa Rica (61). Not 
threatened. Other refs: 51, 55, 62, 234, 269. 

229 Parides phalaeciis (Hewitson, 1869) 

Eastern Ecuador and possibly Peru. Very scarce and perhaps meriting Rare 
status, but more information required. Refs: 51, 55, 234. 

230 Parides agaviis (Drury, 1782) 

Brazil, Paraguay and Argentina. Not uncommon and not thought to be 
threatened. Ranched in Brazil (24). Other refs: 51, 55, 234. 

231 Parides proneus {Hiibncx , \%25) 

South-eastern Brazil. Not uncommon and not known to be threatened. 
Refs: 26,51,55,234. 

232 Parides ascanius (Cramer, 1775) (= orophobus D'Almeida, 1942) 
Vulnerable — refer to section 6, p. 240. 

Only known from scattered points on the coast of Rio de Janeiro state, 
Brazil where it is declining rapidly due to habitat destruction. Protected by 
law in Brazil. Common names: Fluminense Swallowtail, Ascanius Swallow- 
tail (282). Other refs: 51, 55, 234. 

233 Parides bunichus (Hubner, 1821) (= chamissonia Eshscholtz, 1821) 
Brazil and northern Argentina. Not uncommon and in no known danger. 
Two subspecies including Parides bunichus diodorus (Hopffer) which is 
treated by D'Almeida (55) as a separate species. Other refs: 26,51, 
122, 234. 

67 



Threatened Swallowtail Butterflies of the World 

234 Parides diodorus (Hopffer, 1866) 

Brazil (Piavi, Goias and Minas Gerais states). Given as a subspecies of 
bunichus (= chamissonia) by Rothschild & Jordan (234); not listed by 
D'Abrera (51). Not known to be threatened. 

Species-group: aeneas Linnaeus 

235 Parides perrhebus {^o\sd\x\a\, 1836) 

South-eastern Brazil, northern Argentina, Paraguay and Uruguay. Gen- 
erally common and not threatened. Two subspecies, but probably better 
placed with P. bunichus. Refs: 51, 55, 122, 234. 

236 Parides hahneli (Staudinger, 1882) 
Rare — refer to section 6, p. 242. 

Brazil. Although a few specimens have featured in dealers' lists in the last 
few years, the species must be regarded as very scarce. Occasionally 
ranched. Common name: Hahnel's Amazonian Swallowtail. Refs: 27, 51, 
55,234,283. 

237 Parides mithras (Grose-Smith, 1902) (= triopas Godart, 1819) 
Guyana, Surinam, French Guiana and northern Brazil (Obidos). Not 
uncommon but not known to be threatened . Further data needed in order to 
assess conservation status. Two subspecies. Refs: 51, 55, 234. 

238 Parides chabrias (Hewitson, 1852) 

Peru and Ecuador (Upper Amazon). An uncommon species, probably 
conspecific with P. mithras. Further data needed in order to assess 
conservation status. Refs: 51, 55, 234. 

239 Parides quadratus (Staudinger, 1890) 

Brazil (Amazonas) and Peru (Upper Amazon). Apparently a rare species 
(221). More information required. Two subspecies. Other refs: 51, 55, 234. 

240 Parides pizarro {SidiUdmgex , 1884) 
Insufficiently Known — refer to section 6, p. 244. 

Peru (Upper Amazon) and Brazil (Acre State). Again, a fairly scarce and 
little known species. Two subspecies including Parides pizarro kuhlmanni 
May (25, 156), with Parides steinbachi from Bolivia possibly a third (156). 
Other refs: 51,55,234. 

241 Parides steinbachi (Rothschild, 1905) 
Insufficiently Known — refer to section 6, p. 246. 

Eastern Bolivia. Again a very scarce and uncommon species. It has been 
suggested that it may be a subspecies of Parides pizarro (25, 156). D'Abrera 
regards Parides pizarro kuhlmanni from Peru and western Brazil as a 
subspecies of Parides steinbachi (51). Other refs: 55, 139, 234. 

242 Parides coelus (Boisduval, 1836) 
Insufficiently Known — refer to section 6, p. 248. 

Known only from French Guiana. Little is known about this scarce species, 
but it may be under threat. More details needed. Refs: 51, 55, 139, 234. 

243 Parides tros (Fabricius, 1793) (= dardanus Fabricius) 

Brazil (Rio de Janeiro) and northern Argentina. Generally common (55) 
and not threatened. Other refs: 26, 51, 54, 122, 234. 

68 




Swallowtails of the world 

244 Parides aeneas (Linnaeus, 1758) 

Colombia, Guyana, Surinam, French Guiana. Brazil, Ecuador, Peru and 
Bolivia. Not uncommon and not known to be threatened. Parides schiippi 
(Rober, 1927) appears to be a form of Parides aeneas (109). Six subspecies. 
Other refs: 51,55,234. 

245 Parides klagesi (Ehrmann, 1904) 
Insufficently Known — refer to section 6, p. 250. 

Venezuela only. Another scarce and little-known species; the male, taken 
only in 1983, has not been described. More data are required. Refs: 
51,55,139,234. 

246 Parides orellana (Hewitson, 1852) 

Peru and Brazil (Upper Amazon). Not at all common, but not known to be 
threatened. More information required. Refs: 51, 55, 234. 

247 Parides childrenae (Gray, 1832) 

Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Colom- 
bia and Ecuador. Extremely rare in Mexico (62). Moderately common in 
primary rain forest (at least in Costa Rica), but intolerant of areas without 
substantial forest cover (61). Not believed to be threatened, but poor 
forest management might cause a decline. Three subspecies. Other refs: 
51,55,234,269. 

248 Parides sesostris (Cramer , 1779) 

Mexico, Central America (possibly excluding El Salvador), Trinidad and 
South America (excluding Chile, Uruguay and possibly Paraguay); widely 
distributed and not threatened. Four subspecies. Common name: 
Southern Cattle Heart (9,282). Other refs: 51,55,57,61,62,122, 
234, 269. 

249 Parides burchellanus (Westwood, 1872) 
Vulnerable — refer to section 6, p. 252. 

Central Brazil. A little-known and very scarce species; more information is 
needed. Refs: 26, 51, 55, 145, 234. 

250 Parides polyzelus (C. & R. Felder, 1865) 

Mexico, Guatemala, Belize and Honduras. Generally common and not 
threatened. Two subspecies. Refs: 51, 55, 57, 62, 234, 269. 

251 Parides iphidamas (Fahricius, 1793) 

Mexico, Central America, Colombia, Venezuela, Ecuador and northern 
Peru. Generally common and not threatened. Occurs from sea level to 
1200 m in Costa Rica, tolerant of many habitats (61). Five or six subspe- 
cies. Other refs: 51, 55, 57, 62, 234, 269. 

252 Parides vertumnus (Cramer, 1779) 

Colombia, Venezuela, Guyana, Surinam, French Guiana, Ecuador, Bra- 
zil, Peru and Bolivia. Generally common and not threatened. Five to seven 
subspecies. Refs: 51, 55, 234. 

253 Parides cutorina (Staudinger, 1898) 

Eastern Peru and Ecuador (Upper Amazon). Scarce and localized but not 
known to be threatened. More information required. Refs: 51, 55, 234. 

69 



Threatened Swallowtail Butterflies of the World 

254 Parides lycimenes {BohduMsA, 1870) 

Mexico, Central America, Colombia, Venezuela and Ecuador. Fairly 
common and not threatened. Three subspecies. Refs: 55,57, 
61,62,234,269. 

255 Parides phosphorus (Bates, 1861) 

Colombia, Venezuela (?), Guyana, Brazil, Ecuador and eastern Peru. Not 
uncommon but not known to be threatened. Status uncertain, further data 
required. Three subspecies. Refs: 51, 55, 234. 

256 Parides drucei (Butler, 1874) 

Ecuador, Peru, Brazil and Bolivia. Not uncommon and not threatened. 
May be a subspecies of P. anchises. Refs: 55, 234. 

257 Parides erlaces (Gray, 1852) 

Ecuador, Peru, Brazil, Bolivia and northern Argentina. Not uncommon 
nor threatened. Five subspecies. Refs: 51, 55, 122, 234. 

258 Parides nephalion (Godart, 1819) 

South-eastern Brazil, northern Argentina and Paraguay. Generally 
common and not threatened. May be a subspecies of P. anchises. Ranched 
in Brazil (24). Other refs: 26, 51, 55, 122, 234. 

259 Parides anchises (Linnaeus, 1758) 

Colombia, Venezuela, Ecuador, Peru, Trinidad, Guyana, Surinam, French 
Guiana, Brazil, eastern Bolivia, Paraguay and northern Argentina. Widely 
distributed and not uncommon. Ten subspecies. Parides hedae (Foetterle, 
1902) from Brazil is not included as a species by Munroe (188) or Hancock 
(109) and considered to be possibly a form (234, 250) or subspecies (55) of 
P. anchises. P. eversmanni (Ehrmann, 1925) is an invalid name, the male of 
this supposed species was in fact P. anchises while the female was P. 
eurimedes (55, 109). Common name: the Cattle Heart (9). Other 
refs: 26, 122. 

260 Parides erithalion (Boisduval, 1836) 

Costa Rica, Panama, Colombia and northern Venezuela. Generally 
common and not threatened, sea level to 700 m (61). Four subspecies. 
Other refs: 55, 234. 

Species-group: lysander Cramer 

261 Parides panthonus {Cxamex , 1780) 

Brazil, Guyana, French Guiana and Surinam. Not common but not 
regarded as threatened. However, P. p numa has not been recorded since 
1920 (23). Two subspecies. Other refs: 26, 51, 55, 234. 

262 Parides aglaope (Gray, 1852) 

Brazil, southern Peru and eastern Bolivia. Not common, but far from rare 
and not threatened. Refs: 26, 51, 55, 234. 

263 Parides castilhoi (D'Almeida, 1967) 

Brazil (Castilho, Rio Parana, Sao Paulo) (51). More information is required 
on this little known species. 

264 Parides lysander (Cramer, 1775) 

Colombia, Brazil, Guyana, Surinam, French Guiana, Ecuador and Peru. 
Often common (55) and not threatened. Two subspecies. Other ref: 51. 

70 



Swallowtails of the world 

265 Parides echemon (Hiibner, 1813) (= echelus Hubner, 1815) 

Guyana, Surinam, French Guiana and Brazil (lower Amazon). Common 
and not threatened. Two subspecies. Refs: 51, 55, 234. 

266 Parides zacynthus (Fabricius, 1793) 

Eastern Brazil. Fairly common and not threatened. P. z. polymetus is 
ranched in Brazil (24). Two subspecies. Other refs: 51, 55, 234. 

267 Parides neophiliis (Geyer, 1837) 

Colombia, Venezuela, Guyana, Surinam, French Guiana, Trinidad, 
southern and western Brazil, Ecuador, Peru, Bolivia, Paraguay and 
north-eastern Argentina. Common and not threatened. Six subspecies. 
Common name: Spear-winged Cattle Heart (9,282). Other refs: 
234,55,51. 

268 Parides eurimedes (Stoll, 1780) (= areas Cramer, 1777) 

Mexico, Guatemala, Belize (?), El Salvador, Honduras, Nicaragua, Costa 
Rica, Panama, Colombia, Venezuela, Ecuador, Guyana, Surinam and 
French Guiana. Generally common and not threatened. Adoption of the 
name eurimedes follows Hancock (115), areas being preoccupied. Parides 
eversmanni (Ehrmann, 1925) is an invalid name, the male of this supposed 
species was in fact P. anchises while the female was P. eurimedes (55, 109). 
Five, possibly six subspecies. Other refs: 51, 57, 61, 62, 234, 269. 

269 Parides timias (Gray, 1852) 

Ecuador. Fairly common, despite the restricted range. Two subspecies, 
possibly conspecific with P. eurimedes. Refs: 51, 55, 234. 



270 



271 




Genus: Atrophaneura Reakirt 
Subgenus: Pharmacophagus Haase 

Species-group: anterior Drury 

Atrophaneura (Pharmacophagus) antenor (Drury, 1773) 
Madagascar. This species is the only Afrotropical representative of the 
Troidini. Well distributed and apparently not uncommon but the status of 
the species needs to be continually monitored. Collected commercially. 
Refs: 33,34,50, 141,204,206. 

Subgenus: Atrophaneura Reakirt 

Species-group: latreillei Donovan 

Atrophaneura (Atrophaneura) daemonius (Alpheraky, 1895) 

China. Very little information available and none on current status. Further 

research required. Ref: 109. 

Atrophaneura (Atrophaneura) plutonius (Oberthiir, 1907) 
South-western China, Bhutan, Nepal and north-eastern India (Sikkim, 
Manipur, Nagaland, Mizoram and Assam (?)) (52, 94). A rare species of 
high elevations, not known to be threatened but requires monitoring. The 
subspecies pembertoni (Pemberton's Chinese Windmill (80)) and tytleri 
(Tytler's Chinese Windmill (80)) are protected by law in India (182). Three 
subspecies. Common name: Chinese Windmill (94,251), Pemberton's 
Chinese Windmill (80), Tytler's Chinese Windmill (80). Other ref: 259. 



71 



Threatened Swallowtail Butterflies of the World 

273 Atrophaneura (Atrophaneura) alcinous (Klug, 1836) 

Southern China, northern Vietnam (?), northern Laos (?), Taiwan, North 
Korea, South Korea, Japan (to northern Honshu) and U.S.S.R. (Pri- 
morskiy Kray). Apparently common and not in danger as a species but rare 
in the U.S.S.R. and hsted in the U.S.S.R. Red Data Book, category Rare 
(8, 260). Seven subspecies. Common name: Chinese Windmill (80). Other 
refs: 52,141, 149,246,247. 

274 Atrophaneura (Atrophaneura) latreillei (Donovan, 1826) 
Afghanistan (136), northern India (including Uttar Pradesh, Sikkim, 
Assam, Manipur, Nagaland and Meghalaya (?)), Nepal, Bhutan (?), 
northern Burma, southern China and northern Vietnam (?) (52). Not rare 
and not thought to be threatened although the subspecies, ^4. (A.) I. kabrua 
isprotectedby law in India (182). Common name: Rose Windmill (80, 251). 
Three subspecies. Other refs: 4, 141, 259. 

275 Atrophaneura (Atrophaneura) polla (de Niceville, 1897) 
North-eastern India (Assam, Manipur, Chin Hills of Nagaland, Arunachal 
Pradesh), Burma (eastern Bhamo and Bernandmyo of the Shan states), 
northern Thailand (?), south-western China (?) and northern Laos (?) 
(52, 284). Recorded as being very rare by Evans (80) and Talbot (259). 
Protected by law in India (182). More information needed on distribution 
and status. Common name: De Niceville's Windmill (80, 94). Other ref: 
109. 

276 Atrophaneura (Atrophaneura) crassipes (Oberthur, 1879) 
North-eastern India (Manipur), Burma (southern Shan states), northern 
Thailand, northern Laos, northern Vietnam (Tonkin) (94) and possibly 
southern China. Described as very rare (259) and perhaps deserving some 
conservation status, but more information required. It is protected by law in 
India (182). Common name: Black Windmill (80, 94). Other ref: 52. 

277 Atrophaneura (Atrophaneura) adamsoni (Grose-Smith, 1896) 

Burma and Thailand. Locally common in the latter country (168) but said to 
be generally rare (259). Not known to be threatened, but further research 
needed. Common name: Adamson's Rose (80). Other ref: 52. 

278 Atrophaneura (Atrophaneura) nevilli (Wood-Mason, 1896) 
North-eastern India (Assam), Burma (Shan states) and western China 
(52, 94) . Said to be very common in western China though very rare in India 
(259) where it is protected (182) but not known to be threatened. Common 
name: Nevill's Windmill (80, 94). Other ref: 5. 

279 Atrophaneura (Atrophaneura) laos (Riley and Godfrey, 1921) 
Northern Thailand and Laos. Said to be very rare (168), but status not 
known with any certainty. More data required for conservation assessment. 
Other ref: 52. 

280 Atrophaneura (Atrophaneura) mencius (Felder, 1862) 

Sichuan (Szechwan), central and south-eastern China (52,207). Two 
subspecies, although D' Abrera notes that A.(A.)m. rhadinus is closer to A . 
(A.) nevilh and may be a distinct species (52). No information on threats; 
more research needed. 

72 



Swallowtails of the world 

281 Atrophaneura (Atrophaneura) impediens (Rothschild, 1895) 

China including Sichuan (Szechwan) (207) and Taiwan. No data on 
conservation status; more research needed. Two subspecies (52). 

282 Atrophaneura (Atrophaneura) febanus (Fruhstorfer, 1908) 

Restricted to Taiwan. Apparently common, but status should be monitored 
in view of the limited distribution. D' Abrera \\^X?, febanus as a subspecies of 
impediens but this has not been demonstrated adequately (52). Not listed by 
Hancock (109). Other refs: 141 , 246. 

283 Atrophaneura (Atrophaneura) hedistus (Jordan, 1928) 

Southern China (Yunnan). Also little known, with no information available 
on status. Regarded by D'Abrera (52) as a subspecies oiA. (A.) dasarada. 

284 Atrophaneura (Atrophaneura) dasarada (Moore, 1857) 

Northern India, Nepal, Bhutan, Burma, south-eastern China (including 
Hainan (Guangdong prov.)) (52). Not rare and not threatened. Five 
subspecies. Common name: Great Windmill (80,251). Other refs: 
4,141, 168,259. 

285 Atrophaneura (Atrophaneura) polyeuctes (Doubleday, 1842) 
Northern India, Nepal, Bhutan (?), Burma, northern Thailand, Laos, 
Vietnam, southern China (including Yunnan) and Taiwan (52). Up to five 
subspecies. The species lama (Oberthiir) described from western China is 
generally regarded as a subspecies of polyeuctes. A. (A.) philoxenus Gray is 
a synonym. Common name: Common Windmill (80, 251). Other refs: 
4, 141, 168,246,259. 

Species-group: nox Swainson 

286 Atrophaneura (Atrophaneura) semperi (C. & R. Felder, 1861) 
Philippines (not Cebu). Not common but not known to be threatened. 
Seven subspecies (52). A. (A.) erythrosoma Reakirt is a synonym. Other 
refs: 141, 147,262. 

287 Atrophaneura (Atrophaneura) kuehni (Honrath, 1886) 

Indonesia (eastern and northern Sulawesi). Little known about status, 
possibly very rare ( 121 ). It is certainly very rare in collections. More data are 
required on its conservation status. Two subspecies (121). A. (A.) 
mesolamprus Rothschild is believed to be a synonym (52). Other ref: 262. 

288 Atrophaneura (Atrophaneura) luchti (Roepke, 1935) 
Rare — refer to section 6, p. 254. 

Indonesia (Java) . Restricted to the mountains in the far east of Java. Little is 
known about this species and the specific status has been brought in 
question due to its similarity with /I. (A.) priapus(\\%). Otherrefs: 52, 262. 

289 Atrophaneura (Atrophaneura) hageni (Rogenhofer, 1889) 

Indonesia: restricted to high plateau of Sumatra. Uncommon, but not 
known to be threatened. Refs: 52, 141. 

290 Atrophaneura (Atrophaneura) priapus (Boisduval, 1836) 

Indonesia (Java and southern Sumatra) (262). The three 'species' sycorax, 
priapus and hageni are often confused and may represent one, two or three 
good species. Not known to be threatened, though status uncertain. More 

73 





Threatened Swallowtail Butterflies of the World 

information and close monitoring are needed. Two (52) or three (262) 
subspecies. Common name: White-head Batwing (3). 

291 Atrophaneura (Atrophaneura) sycorax (Grose-Smith, 1885) 

Southern Burma, Thailand, Peninsular Malaysia and Indonesia (Sumatra 
and western Java) (52, 262). Rather rare (45) but not known to be 
threatened. Three subspecies (52, 262). Common name: White-head 
Batwing (80). Other refs: 87, 168, 259 (as ^. (A.) priapus). 

292 Atrophaneura (Atrophaneura) horishanus (Matsumura, 1910) 
Restricted to Taiwan. Apparently common but status should be kept under 
review. A. (A.) sauteri Heyne is a synonym. Refs: 52, 141, 246. 

293 Atrophaneura (Atrophaneura) aidoneus (Doubleday, 1845) 

Northern India (including Uttar Pradesh, Sikkim, Assam, Meghalaya (?), 
Manipur (?) and Nagaland (?)), Bhutan, Burma, northern Laos, northern 
Vietnam, southern China (including Hainan (Guangdong prov.)) (52). Not 
common but not regarded as threatened. Common name: Lesser Batwing 
(80). Other refs: 4, 259. 

Atrophaneura (Atrophaneura) varuna (White, 1868) 
North-eastern India (including Sikkim), eastern Nepal (?), Burma, Thai- 
land, northern Laos, northern Vietnam and Peninsular Malaysia (52). Not 
rare across most of its range and not known to be threatened. In Peninsular 
Malaysia it is uncommon and flies at high elevations (45). Believed to be 
extinct in Singapore (45). Two subspecies. Common name: Common 
Batwing (3, 80). Other refs: 4, 87, 141, 168, 259. 

295 Atrophaneura (Atrophaneura) zaleucus (Hewitson, 1865) 
Burma, Thailand, northern Laos, northern Vietnam and possibly south- 
western China (52). Not generally common, but not known to be threat- 
ened. Sometimes regarded as a subspecies of A. (A.) varuna (259). 
Common name: Burmese Batwing (80). Other refs: 141, 168. 

296 Atrophaneura (Atrophaneura) nox (Swainson, 1822-23) 
Peninsular and Eastern Malaysia, southern Thailand (?), Brunei (?) and 
Indonesia (Sumatra, Nias, Mentawai Is, Java, Bali and Kalimantan) 
(52, 262). A widespread but often local species of well-wooded localities; 
extinct in Singapore (45). Nine subspecies. Common name: Malayan 
Batwing (45). Other refs: 87, 131, 141. 

297 Atrophaneura (Atrophaneura) dixoni (Grose-Smith, 1901) 

Indonesia (northern and central Sulawesi). Status uncertain, said to be 
uncommon to rare, presumably localized (121) More data are required. 
Other refs: 52, 262. 

Subgenus: Losaria 

Species-group: coon Fabricius 

N.B. The next four species represent a distinctive facies in the genus 
Atrophaneura (called the coon group by Hancock (109)) and are here 
placed in the subgenus Losaria. Tsukada and Nishiyama treat Losaria as a 
full genus (262). 

298 Atrophaneura (Losaria) neptunus (Guerin-Meneville, 1840) 
Southern Burma, southern Thailand, Peninsular and Eastern Malaysia, 
Philippines (Palawan), Brunei and Western Indonesia (Sumatra, Nias, 

74 



Swallowtails of the world 

Simeulue and Kalimantan). Less common than A. (L.) coon but widely 
distributed and not known to be threatened. Eight subspecies. Common 
name: Yellow-bodied Clubtail (80), Yellow-red Clubtail (3). Other refs: 
45,52,87, 131, 168,259,262. 

299 Atrophaneura (Losaria) pain (Martin, 1912) 
Insufficiently Known — refer to section 6, p. 256. 

Indonesia (western Sulawesi (Palu)). Taxonomic status no longer in doubt 
(114). Said to be a subspecies of A. (L.) coon by Tsukada and Nishiyama 
(262) but it is apparently distinct and isolated from other members of the 
coon group (1 14, 121). Believed to be restricted to the west coast where it is 
rare and localized (172). 

300 Atrophaneura (Losaria) coon (Fabricius, 1793) 

Northern India (Assam), southern Burma, Thailand, Laos, Vietnam, 
Kampuchea, China (Hainan (Guangdong prov.)). Peninsular Malaysia, 
Nicobar Is, Indonesia, (Sumatra, western Java and Bawean Is) (52, 262). 
The record from Assam (80) seems to be erroneous, as do some attribu- 
tions of subspecies (52). A. (L.) c. sambilanga from the Nicobar Is is 
protected under Indian law (182) and is said to be very rare (5). A. (L.) c. 
doubledayi is extinct in Singapore (45). Eight subspecies excluding pa/w. 
Common name: Common Clubtail (45, 80, 282). Other refs: 4, 87, 107, 
141, 168.259. 

301 Atrophaneura (Losaria) rhodifer (Butler, 1876) 

Restricted to the Andaman Is. Possibly to be classified when its status is 
clearer but described as not rare (5, 80, 259) and so not listed as threat- 
ened. Common name: Andaman Clubtail (80). Other ref: 52. 

Subgenus: Pachliopta 

Species-group: polydorus Boisduval 

N.B. The genus Pachliopta has been synonymized with Atrophaneura 
(107) but we retain Pachliopta at the subgeneric level as representing a 
distinct fades within the genus. Both genera have in the past been called 
Polydorus. 

302 Atrophaneura (Pachliopta) hector (Linnaeus, 1758) 

Southern and eastern India, Sri Lanka, Andamans and possibly the coast 
of western Burma (52). Generally common and not threatened but said to 
be very rare in the Andamans (5, 259). Protected by law in India (182). 
Common names: Common Rose (3, 282), Crimson Rose (80). Other refs: 
4, 287. 

303 Atrophaneura (Pachliopta) jophon (Gray, 1852) 
Vulnerable — refer to section 6, p. 258. 

Sri Lanka. Declining due to loss of habitat (52, 5?)). A. (P.) pandiyana from 
southern India is sometimes regarded as a subspecies oi A. (P.) jophon. 
Common name: Ceylon Rose (80). Other refs: 259, 287. 

304 Atrophaneura (Pachliopta) pandiyana (Moore, 1881) 

Southern India. Uncommon but not considered to be threatened. (This 
species and A. (P.) jophon are sometimes regarded as being conspecific). 
Common name: Malabar Rose (80). Other refs: 4, 52. 

75 



Threatened Swallowtail Butterflies of the World 

305 Atrophaneura (Pachliopta) oreon (Doherty, 1891) 

Indonesia: Lesser Sunda Is (Sumba, Alor, Flores and Solor) (52, 262). 
Possibly threatened in view of the very narrow range, but insufficient 
information available on status. Two subspecies (262). Other ref: 109. 

306 Atrophaneura (Pachliopta) liris (Godart, 1819) 

Indonesia: Lesser Sunda Is (Timor, Sawu, Wetar, Moa, Leti, Kisar, Babar, 
Damar, Romang and Tanimbar) (262). Apparently common and not 
threatened. Six subspecies. Other refs: 48, 130. 

307 Atrophaneura (Pachliopta) polyphontes (Boisduval, 1836) 

Indonesia (Sulawesi, Salayar, Talaud, Sangihe (?) and the Moluccas (Sula, 
Ternate , Bacan , Halmahera and Morotai)) (52, 262) . Five subspecies (262) . 
Other refs: 48, 141. 



,;^ 



« 308 Atrophaneura (Pachliopta) schadenbergi (Semper, 1886) 
^^ Vulnerable — refer to section 6, p. 261. 



Philippines: Luzon and the Babuyan Is. A locally distributed species with 
J^'- seasonally limited occurrence in lowland and medium elevation wooded 

areas. Threatened by habitat destruction. Two subspecies. Refs: 52, 177, 
262. 

309 Atrophaneura (Pachliopta) mariae (Semper, 1874) 
Philippines (Luzon, Samar, Leyte, Bohol, Mindanao, Sibuyan and Polillo). 
Neither rare nor threatened. Three subspecies. Refs: 52, 141, 147, 177,262. 

310 Atrophaneura (Pachliopta) phegeus (Hopffer, 1886) 
Southern Philippines (Mindanao, Samar, Leyte and Cebu). Not rare and 
not considered to be threatened. A. (P.) leytensis (Murayama) is believed 
to be a synonym. Refs: 52, 141, 147, 177, 262. 

311 Atrophaneura (Pachliopta) phlegon (C. & R. Felder, 1864) 
Philippines (Mindoro, Panay, Guimaras, Marinduque and Mindanao 
(262)). Not uncommon and not known to be threatened. Two subspecies 
including Bryk's {192>Q) strandi irom the Philippines (262). A. (P.) phlegon 
is regarded as a subspecies of annae by D'Abrera (52) but annae is normally 
regarded as a synonym of this species as issabinae (Seyer) (107). Other refs: 
52, 177. 

312 Atrophaneura (Pachliopta) atropos (Staudinger, 1888) 
Indeterminate — refer to section 6, p. 263. 

Restricted to Philippines (Palawan). Not much is known about its status, 
but it seems to be uncommon and is listed as Indeterminate until more 
information is available. Refs: 52, 141, 262. 

313 Atrophaneura (Pachliopta) kotzebuea (Eschscholtz, 1821) 
Widely distributed in the Philippine Is south of Babuyan Is, but absent from 
Palawan (262). Flies on edges of rain forest (262). A. (P.) kotzebuea is 
recognized by Tsukada and Nishiyama (262), but not by Munroe (188) or 
Hancock (109). Not specifically known to be threatened, although any 
Philippine endemic dependent upon rain forest must be considered to be at 
some risk. Five subspecies (262). 

314 Atrophaneura (Pachliopta) aristolochiae (Fabricius, 1775) 
Afghanistan (118), India, Nepal, Sri Lanka, Burma, Thailand, southern 
and eastern China (including Hainan (Guangdong prov.)). Hong Kong, 

76 





Swallowtails of the world 

Taiwan, Japan (south-western Okinawa only), Laos, Vietnam, Kampu- 
chea, Andamans, Nicobar Is, Peninsular and Eastern Malaysia, Brunei, 
Philippines (Palawan and Leyte). Indonesia (Sumatra, Nias. Enggano, 
Bangka, Java. Bali, Kangean, Lombok, Sumbawa, Sumba, Flores, Tanah- 
jampea and Kalimantan) (262). Generally common and not threatened. 
Up to twenty subspecies. Common name: Common Rose (45, 80, 251). 
Other refs: 4. 5, 52, 87, 131, 141, 143, 147, 168, 246, 259. 287. 

315 Atrophaneiira (Pachliopta) polydonis {Linnaeus, 1758) 

Indonesia (Moluccas (not Morotai, but including Seram and Ambon), 
Tanimbar, Irian Jaya, Kai, Aru, Waigeu and Skouten Is), Papua New 
Guinea, Bismarck Archipelago (including New Britain), Bougainville, 
Solomons (including San Cristobal and Ulawa I.), Trobriand Is, D'Entre- 
casteaux Is, Louisiade Archipelago and Australia (northern Queen- 
sland)(262). Generally common and not threatened as a species, though 
some subspecies may be threatened (138). About four subspecies are 
farmed in Papua New Guinea (36). Up to thirty-one subspecies are known 
(262). Common name: Red-bodied Swallowtail (282). Other refs: 14. 40, 
48,68, 141, 176. 

Genus: Trogonoptera Rippon 

N.B. Both species of Trogonoptera are included on Appendix II of the 
Convention on International Trade in Endangered Species of Wild Fauna 
and Flora (CITES) . The 87 countries that are party to CITES are obliged to 
invoke national legislation implementing the Convention. In line with this 
requirement import and export restrictions have been implemented by 
national legislation in the Federal Republic of Germany, the Netherlands 
and the United Kingdom, and by Regulation 3626/82 of the Commission of 
the European Community. 

316 Trogonoptera brookiana (Wallace, 1855) 

Peninsular and Eastern Malaysia, Indonesia, (Sumatra, Mentawai Is (284) 
and Kalimantan) and Brunei (52, 262). Locally common (especially males) 
and not thought to be threatened, despite the large numbers taken for the 
decorative trade. Protected by law in Malaysia and Indonesia but still 
collected commercially, possibly as many as 125 000 specimens are expor- 
ted per year (10). In Malaysia the species is 'protected" but not "totally 
protected". This means that licences and permits may be issued for the 
taking, import , export or possession of specimens. A permit is also required 
to accompany the possession of a specimen. Five subspecies. Common 
name: Raja(h) Brooke's Birdwing (3, 45, 49, 282). Other refs: 87, 131,141, 
201. 

317 Trogonoptera trojana (Honrath, 1886) 

Philippines (Palawan, Balabac) (52, 262, 281). Less common and with a 
much more restricted range than T. brookiana but not presently regarded 
as being threatened. Nevertheless males are collected in great numbers, 
particularly in the rain forests of northern Palawan, and the forests 
themselves are under growing pressure from timber interests (56). In 
common with D"Abrera (49) most authors consider T. trojana to be a 
distinct species, but Igarashi (141) regards it as a subspecies of T. 
brookiana. 

11 




Threatened Swallowtail Butterflies of the World 

Genus: Troides Hiibner 

N.B. All species of Troides are included on Appendix II of the Convention 
on International Trade in Endangered Species of Wild Fauna and Flora 
(CITES). The 87 countries that are party to CITES are obliged to invoke 
national legislation implementing the Convention. In line with this require- 
ment import and export restrictions have been implemented by national 
legislation in the Federal Republic of Germany, the Netherlands and the 
United Kingdom, and by Regulation 3626/82 of the Commission of the 
European Community. 

Subgenus: Ripponia Haugum & Low 

Species-group: hypolitus Cramer 

318 Troides (Ripponia) hypolitus (Cramer, 1775) 

Indonesia (Sulawesi (including Sangihe and Talaud) and the Moluccas: 
Sula, Morotai, Halmahera, Ternate, Buru, Seram, Saparua, Ambon (52, 
262) and Obi (252)). Not rare and not regarded as being threatened, though 
protected in Indonesia. Four subspecies. Other refs: 48, 141, 188. 

Subgenus: Troides Hiibner 

Species-group: amphrysus Cramer 

319 Troides (Troides) cimeifer (Oberthur, 1879) 

Southern Thailand, Peninsular Malaysia and Indonesia (Sumatra and Java) 
(52, 262). Not rare and not known to be threatened across most of it range, 
but considered to be Vulnerable and in need of some protection in 
Peninsular Malaysia (10). Three subspecies. Common name: Golden 
Birdwing (3). Other refs: 45, 49, 87, 141. 

320 Troides (Troides) amphrysus (Cramer, 1782) 

Southern Burma (Mergui Archipelago), southern Thailand, Peninsular and 
Eastern Malaysia, Brunei, Indonesia (Kalimantan, Sumatra, Simeulue (?), 
Nias, Batu, Mentawai Is, Enggano and Java) (49, 52, 191, 262). Not rare 
and not known to be threatened except in Malaysia where said to be 
Vulnerable (10). Protected in Indonesia. Six subspecies. Common names: 
Golden Birdwing (3), Malay Birdwing (80). Other refs: 45, 87, 131, 141, 
168, 231. 

321 Troides (Troides) miranda (Butler, 1869) 

Brunei, Eastern Malaysia (Sabah and Sarawak), Labuan I. and Indonesia 
(northern Kalimantan and western Sumatra) (52, 262). Igarashi's map (141) 
erroneously shows most of the Malaysian peninsula, as well. Not rare and 
not known to be threatened, though protected in Indonesia. Two subspe- 
cies. Other refs: 49, 131,231. 

322 Troides (Troides) andromache (Staudinger, 1892) 
Indeterminate — refer to section 6, p. 266. 

Eastern Malaysia (Sabah and Sarawak) (49, 132, 231) but not recorded 
from Indonesia (Kalimantan) although it is protected there. Very localised. 
Two subspecies. Other refs: 11, 52, 131. 

78 



Swallowtails of the world 



323 



324 



325 



326 



327 



328 



329 






Species-group: Helena Linnaeus 

Troides (Troides) magellanus (C. & R. Felder, 1862) 
Philippines (Batan Is, Babuyan Is, Luzon, Polillo, Marinduque, Cuyo I., 
Samar, Cebu, Leyte, Bohol and Mindanao) and Taiwan (Hung-t'ou Hsii Is 
only) (262). Not uncommon and not thought to be threatened. Three 
subspecies (262) Other refs: 49, 52, 141, 147, 246. 

Troides (Troides) prattorum (Joicey and Talbot, 1922) 

Indeterminate — refer to section 6, p. 269. 

Indonesia (Buru) only; a very restricted range. The species is rare and little 

known as Buru was previously almost impossible to visit due to its use as a 

prison camp (49). It is now apparently accessible to collectors and 

specimens are beginning to appear on dealers' lists. Other ref: 48. 

Troides (Troides) minos (Cramer, 1779) 

Western and south-western India (49, 52). Despite its restricted range the 
species is not known to be threatened, but continuous monitoring is 
recommended. Some authorities have regarded it as a subspecies of T. (T.) 
Helena. Other refs: 4, 15, 80, 259. 

Troides (Troides) aeaciis (C. & R. Felder, 1860) 

Northern India, Nepal, Burma, China, Thailand, Laos (?), Vietnam (?), 
Taiwan, Kampuchea, Peninsular Malaysia (49, 52) and Indonesia (Suma- 
tra) (118). Generally common and not threatened though classified as 
Vulnerable and possibly in need of protection in Peninsular Malaysia (10). 
It is very uncommon in Sumatra ( 1 18) . In 1974 the Entomological Suppliers 
Association of Great Britain agreed not to trade in the endemic Taiwan 
subspecies T. a. kagiiya on the grounds that such trade could threaten the 
well-being of the taxon (123). Three subspecies. Common name: Golden 
Birdwing (80, 251). Other refs: 4, 45, 87, 141, 168, 207, 246, 259. 

Troides (Troides) plateni Staudinger, 1888 

Philippines (only known positively in Palawan (281), but possibly occurs in 
Calamian Is) (262). A common species easily raised in captivity and not 
thought to be at risk (56). Igarashi (141) and, by implication, Munroe (188), 
Smart (250) and Hancock (109) regard it as a subspecies of T. (T.) 
rhadamantus , but D'Abrera emphasises its specific distinction (49, 52). 
Other ref: 119. 

Troides (Troides) rhadamantus (Lucas, 1835) 

Philippines (except Palawan, Dumaran, Bugsuk and Calamian Is) (49, 52, 
262). Present in Luzon, Polillo, Mindoro, Marinduque, Romblon, Mas- 
bate, Samar, Panay, Guimaras, Negros, Cebu, Leyte, Panaon, Dinagat, 
Bohol, Camiguin, Mindanao, Basilan, Jolo, Tawitawi (262). Not uncom- 
mon and not known to be threatened. Listing of this species under 
Indonesian legislation refers to T. (T.) dohertyi, sometimes considered to 
be a subspecies of r. ('r.)r;iadfl/«a«fi«( 118, 262). Other refs: 141, 147,281. 

ides) dohertyi Rippon, 1893 
Vulnerable— TTefer to section 6, p. 271. 

THaorTesiatTalaud Is and Sangihe) (262); listed as threatened on account of 
its very restricted range. However, the remarks on the specific distinctness 
of T. (T.) plateni apply in exactly the same way to T. (T.) dohertyi. 

79 



Threatened Swallowtail Butterflies of the World 

Protected in Indonesia under the name T. (T.) rhadamantus, of which \t has 
been considered to be a subspecies (118, 246). Common name: Talaud 
Black Birdwing. Other refs: 49, 52, 119, 141. 

330 Troides (Troides) helena (Linnaeus, 1758) 

From north-eastern India through Indo-China and southern China to 
Indonesia i.e. north-eastern India (including Orissa and Sikkim), Nepal, 
Bangladesh, Burma, southern China (including Hainan (Guangdong 
prov.)), Hong Kong, Thailand, Vietnam, Laos, Kampuchea (?), Andaman 
Is, Nicobar Is, Peninsular and Eastern Malaysia, Indonesia (Sumatra, Nias, 
Enggano, Java, Bawean, Kangean Is, Bali, Lombok, Sumbawa, Bunguran 
(Natuna), Sulawesi, Butung, Tukangbesi and Kalimantan) and Brunei (49, 
52 , 262) . Widely distributed and often common , but classified as Vulnerable 
and possibly requiring protection in Peninsular Malaysia (10). Protected in 
Indonesia. Seventeen subspecies. Common name: Common Birdwing (45, 
80, 251). Other refs: 4, 5, 14, 87, 131, 141, 143, 168, 231, 259. 

331 Troides (Troides) oblongomaculatus (Goeze, 1779) 

Indonesia (southern Sulawesi, Salayar (Kabia), Tanahjampea, Sula, Buru, 
Seram, Ambon, Seramlaut, Watubela, Banda Is, Salawati, Skouten Is 
(Yapen) and Irian Jaya) and Papua New Guinea (49, 52, 262). Generally 
common and certainly not threatened. It is easily reared and commonly sold 
on the butterfly market in Papua New Guinea (17). Either six (49) or nine 
(262) subspecies. The Sulawesi records may refer to hybrids (115). Other 
refs: 14, 36, 48, 217. 

332 Troides (Troides) darsius (Gray, 1852) 

Sri Lanka only. Not rare and not threatened. Regarded as a subspecies of T. 
(T.) helena by earlier authors (80, 259, 287) but now generally accepted as a 
full species. Other refs: 49, 52. 

333 Troides (Troides) criton (C. & R. Felder, 1860) 

Indonesia, confined to the Moluccas (Morotoi, Ternate, Halmahera 
(Jailolo Gilolo), Bacan and Obi) (49). May be confused with the hybrid 
celebensis from Sulawesi which, though criton-Wke., has nothing whatever to 
do with this species (118). Reasonably common (48) and not thought to be 
threatened, though protected in Indonesia. Possibly two subspecies. 

334 Troides (Troides) riedeli (Kirsch, 1885) 

Indonesia (Tanimbar Is). The species has a restricted range and might be 
considered for threatened status. However, it is said not to be rare (48) and 
it is protected by law in Indonesia. The species should be carefully 
monitored and extensive suitable habitat protected. Any serious habitat 
disruption would necessitate immediate listing as Vulnerable and appro- 
priate conservation measures. Other refs: 49, 262. 

335 Troides (Troides) plato Wallace, 1865 

Indonesia (Timor). Apparently very rare (48) but locally common (118). 
More information is required on its status and monitoring is recommended. 
It is protected in Indonesia. Earlier regarded as a subspecies of haliphron 
and not listed as a good species by several authors (109, 141, 188). 
Reinstated as a species by Haugum and Low who had studied its genitalia 
(118, 119). Other refs: 49, 262. 

80 



Swallowtails of the world 

336 Troides (Troides) vandepolli (Snellen, 1890) 

Indonesia (north-west Java and western Sumatra) (49, 52, 262). Uncom- 
mon, but not known to be threatened though it is protected in Indonesia. 
Two subspecies (262), which are possibly full species (49). 

337 Troides (Troides) haliphron (Boisduval, 1836) 

Southern Indonesia: Sulawesi, Salayar (Kabia), Tanahjampea, Kalao, 
Bonerate, Sumbawa, Sumba, Komodo (?), Rinja (?), Flores, Solor Is, 
Alor, Wetar, Romang, Damar, Leti, Moa, Babar and the Tanimbar Is (49, 
262). Not rare and not known to be threatened but it is protected in Indone- 
sia. Either nine (49) or eleven (262) subspecies. Other refs: 48, 52, 141. 

338 Troides (Troides) staudingeri (Rober, 1888) 

Confined to the Babar (Babber) group in Indonesia east of Timor (49). 
Raised to a full species by Haugum and Low (118, 1 19) who had studied the 
genitalia , but previously considered to be a subspecies of haliphron by other 
authors. Protected in Indonesia under T. (T.) haliphron. Not known to be 
threatened. Other ref: 109. 

Genus: Ornithoptera Boisduval 

N.B. All species of Ornithoptera are included on Appendix II of the Con- 
vention on International Trade in Endangered Species of Wild Fauna and 
Flora (CITES). The 87 countries that are party to CITES are obliged to 
invoke national legislation implementing the Convention. In Hne with this 
requirement import and export restrictions have been implemented by 
national legislation in the Federal Republic of Germany, the Netherlands 
and the United Kingdom, and by Regulation 3626/82 of the Commission of 
the European Community. Hancock (109) reduced Ornithoptera to a 
subgenus of Troides but we choose to retain Ornithoptera as a full genus for 
practical reasons of usage. 

Species-group: tithonus de Haan 

339 Ornithoptera goliath Oberthiir, 1888 

Indonesia (Seram, Waigeo and Irian Jaya) , Papua New Guinea and Goode- 
nough I. The species is protected in Indonesia; the Papua New Guinea 
subspecies is also protected but has been recommended as a good candidate 
for farming (36). Adults have been noted as scarce in nature (119), Out a 
recent report by M.J. Parsons (in litt.) records that O. goliath may also be 
locally common and does not require protected status. In view of this 
reassessment, O. goliath is not listed as threatened. Ten subspecies have 
been described but this has been reduced to two (49) or five (119). Common 
name: Goliath Birdwing (282). Other refs: 48, 183, 217. 

340 Ornithoptera tithonus de Haan, 1840 
Insufficiently Known — refer to section 6, p. 273. 

Indonesia (Irian Jaya, Waigeo, Salawati and Mysol (Misool) Is). A rela- 
tively rare species (119) with a restricted range (117) but one subspecies has 
been repeatedly collected in recent years (119). Protected in Indonesia. 
Three subspecies, one as yet undescribed. Other refs: 48, 49. 

341 Ornithoptera rothschildi Kenrick, 1911 
Indeterminate — refer to section 6, p. 276. 

Indonesia (endemic to the Arfak Mountains, Irian Jaya). Although the 

81 



Threatened Swallowtail Butterflies of the World 

species has an extremely limited distribution, it is 'not rare, localized, and 
occurs in some abundance at certain sites' (119). It is under heavy 
commercial collecting pressure (117), despite being protected in Indonesia. 
Or^/f/zopferflfl/cfl/cefle-KobayashiandKoiwaya, 1978from Irian Jaya(Arfak 
Mountains) is considered by Haugum and Low (119) to be a natural hybrid 
of O. rothschildi and O. priamus poseidon. Common name: Rothschild's 
Birdwing (49). Other refs: 48, 155. 

342 Ornithoptera chimaera (Rothschild, 1904) 
Indeterminate — refer to section 6, p. 278. 

Papua New Guinea and Indonesia (Irian Jaya). A montane species, scarce 
but widely distributed in the east (119); protected in Papua New Guinea and 
Irian Jaya but recommended as a good candidate for farming in the former 
(36). Two subspecies are recognized in the most recent study. Common 
name: Chimaera Birdwing. Other refs: 48, 49, 117, 183. 

343 Ornithoptera paradisea Si&xxdmgQX , 1893 
Indeterminate — refer to section 6, p. 281. 

Papua New Guinea and Indonesia (Irian Jaya): widespread but localised in 
seven main areas (119). The species is protected in both Papua New Guinea 
and Irian Jaya. Six named subspecies, of which the nominate subspecies has 
been extinct for many years. Common names: Paradise Birdwing (49, 282), 
Tailed Birdwing (282). Other refs: 14, 48, 141, 183, 217. 

344 Ornithoptera meridionalis (Rothschild, 1897) 
Vulnerable — refer to section 6, p. 284. 

Extreme south-eastern Papua New Guinea (where it is protected) and 
Indonesia (Irian Jaya). Very rare and localized but not uncommon where it 
occurs (119). It must be considered Vulnerable due to the danger from 
habitat destruction, sedentary behaviour of adults and the extremely low 
number of eggs laid by each female (118). Dealers' prices are exceptionally 
high (3). One or two subspecies. Other refs: 48, 49, 108, 141, 183. 

Species-group: priamus Linnaeus 

345 Ornithoptera alexandrae (Rothschild, 1907) 
Endangered — refer to section 6, p. 288. 

Restricted to a very small part of south-eastern Papua New Guinea where it 
is protected from collectors (183) but under threat from habitat destruction. 
Conservation of this species is a priority for any programme on swallowtails. 
Common name: Queen Alexandra's Birdwing (49). Other refs: 14, 48, 1 19, 
141. 

346 Ornithoptera victoriae (Gray, 1856) 

Widely distributed throughout the Solomons Is and Bougainville (Papua 
New Guinea) (48, 49, 119, 141, 188, 219, 250) and not rare (171), but in 
demand by collectors so that Vulnerable status could be applied to some 
subspecies. May be threatened by agriculture and forestry in localized sites 
(171). Declining rapidly on Malaita due to intense deforestation (220). 
Protected on Bougainville (Papua New Guinea), but may be struck off the 
protected list in the near future to allow farming, since it occurs in large 
numbers and the larvae feed on the common and easily grown Aristolochia 
tagala (36). Seven subspecies (32). O. allottei Rothschild (Abbe Allotte's 
Birdwing) from the Solomon Archipelago (Bougainville and the south of 

82 



Swallowtails of the world 

Malaita) is generally regarded as a hybrid of O. victoriae x O. priamiis 
urvillianus; Racheli (219) gives references to the different views. The 
hybrid is protected on Bougainville (Papua New Guinea). Common name: 
Queen Victoria's Birdwing (49). Other ref: 183. 

347 Ornithoptera aesacus (Ney, 1903) 
Indeterminate — refer to section 6, p. 292. 

Indonesia; known only from the the island of Obi in the Moluccas (48, 49, 
119). Very rare and with a restricted distribution on this heavily logged 
island. When more information is available Vulnerable or Endangered 
status may prove appropriate. 

348 Ornithoptera croesus Wallace, 1859 
Vulnerable — refer to section 6, p. 294. 

Indonesia: Moluccas (Halmahera, Ternate, Tidore, Bacan, Morotai and 
Sanana (Sula Is)). Not common, conservation status in some doubt, but 
possibly threatened by insecticide sprays (119) and certainly in demand by 
collectors. Two subspecies are distinct whilst there is some doubt about a 
third which has recently been described from Sanana (262). O. c. lydiiis is 
unusual in that the female mimics unpalatable species of Danainae. Other 
refs: 49, 141. 

349 Ornithoptera priamiis {Linnaeus, 1758) 

Indonesia (Ambon, Seram, Waigeo, Misool, Kai Is, Aru Is, Tanimbar Is 
and Irian Jaya), Papua New Guinea and offshore islands, Bismarck Archi- 
pelago (including New Britain), Trobriand Is, D'Entrecasteaux Is, 
Woodlark I., Louisiade Archipelago, Bougainville and the Solomon Is 
(excluding San Cristobal) (119). Protected in Indonesia. Following Han- 
cock (109), the Australian forms euphorion and richmondia (north-eastern 
coastal regions southwards to New South Wales) are here given full species 
status, but other authors may include these as subspecies of priamus (115). 
While many races are generally common, some of the eleven to fifteen 
subspecies merit special consideration. Some of the Papua New Guinea 
subspecies are farmed (36). O. p. caelestis (Rothschild), generally 
regarded as a subspecies of O. priamus (119, 141), from the Louisiade 
Archipelago suffers from a very restricted range, pressures on its habitat 
and overcollecting, although it is now being farmed (36, 83, 138). O. p. 
miokensis (Ribbe) from the Duke of York Is, O. p. boisduvali (Montrou- 
zier) from Woodlark I. and O. p. demophanes (Friihstorfer) (possibly only 
a form of O. p. poseidon (Doubleday); the New Guinea Birdwing) from 
the Trobriand Is are all threatened by pressures on their habitat, a 
restricted range and to a much lesser extent overcollecting (138). O. p. 
urvillianus (Guerin-Meneville), D'Urville's Birdwing, from the Bismarck 
Archipelago (New Ireland), Bougainville and the Solomon Is (excluding 
San Cristobal and Rennell) (119, 141, 219, 220) is farmed in the first two 
areas (36). Common name: Priam's Birdwing (49). Other refs: 14, 40, 48, 
68, 176, 217, 255. 

350 Ornithoptera euphorion (Gray, 1853) 

Cooktown to Mackay, northern Queensland, AustraHa. Listed as a 
subspecies oi priamus by D'Abrera (49) and Haugum and Low (119), but 
accepted as a full species by Hancock (109). Protected in Queensland 
under the name O. priamus, the Cairns Birdwing (258). This species 

83 



Threatened Swallowtail Butterflies of the World 

should be monitored because of its restricted distribution in an area of 
extensive habitat destruction. Common name: Cairn's Birdwing (49). 

351 Ornithoptera richmondia (Gray, 1852) 

Eastern coastal region of Australia: southern Queensland and northern 
New South Wales, a restricted range. Specific identity doubtful and 
generally regarded as a subspecies of O. priamus (40, 119, 141, 176), 
although accepted as a full species by Hancock (109) and D'Abrera (49). 
Protected in Queensland (115). As for O. euphorion, monitoring of this 
species is highly recommended. Common name: Richmond Birdwing (49). 
Other ref: 48. 

Subfamily: PAPILIONINAE 
Tribe: Papilionini 

Genus: Papilio Linnaeus 

N.B. The genus Papilio sensu lato is retained throughout. The generic 
names proposed by Hancock (109) have been used at the subgeneric level. 

Subgenus: Pterourus Scopoli 

Species-group: glaucus Linnaeus 

352 Papilio (Pterourus) glaucus L., 1758 (= turnus L., 1771) 

Canada, eastern and south-eastern U.S.A. (including Alaska and Texas) 
and Mexico. Common and not generally threatened, but listed as a Special 
Species by Washington State Department of Game. Three subspecies. 
Common name: Tiger Swallowtail (154, 216). Other refs: 55, 62, 137, 141, 
234, 269. 

353 Papilio (Pterourus) alexiares Hopffer, 1886 

Restricted to eastern Mexico, but not known to be threatened. Two 
subspecies. Refs: 51, 55, 62, 175, 234, 269. 

354 Papilio (Pterourus) multicaudatus Kirby, 1884 (= daunus Bois., 1836) 
South-western Canada (British Columbia and Alberta), western and 
mid-western U.S.A. (Washington, Montana, North Dakota, South 
Dakota, Oregon, Idaho, Wyoming, Nebraska, California, Nevada, Utah, 
Colorado, Kansas, Arizona, New Mexico and Texas), Mexico and Guate- 
mala. Fairly widely distributed and not threatened. Common names: 
Two-tailed Tiger Swallowtail (216), Three-tailed Swallowtail (282). Other 
refs: 55, 62, 137, 169, 234, 269. 

355 Papilio (Pterourus) rutulus Lucas, 1852 

Western and mid-western U.S.A. (Washington, Idaho, Montana, South 
Dakota (Black Hills), Wyoming, Oregon, California, Nevada, Utah, 
Colorado, Arizona and New Mexico), south-western Canada (British 
Columbia) and northern Mexico (Baja California). Common and not 
threatened but rare east of the Rockies. Three subspecies (269) of doubtful 
vahdity (137). Common name: Western Tiger Swallowtail (216). Other 
refs: 55, 62, 141,234. 

356 Papilio (Pterourus) eurymedon Lucas, 1852 

South-western Canada (British Columbia), western U.S.A. (Washington, 
Idaho, Montana, Oregon, Wyoming (?), California, Nevada, Utah, 

84 



Swallowtails of the world 

Colorado, Arizona and New Mexico) and northern Mexico (Baja Califor- 
nia). According to D'Almeida (55) it is also found further south in Mexico, 
Belize and possibly Guatemala but this is questionable . Not uncommon and 
not known to be threatened. Common name: Pale Tiger Swallowtail (216). 
Other refs: 62, 137, 234, 269. 

Species-group: troilus Linnaeus 

357 Papilio (Pterourus) pilumnus Boisduval, 1836 

Mexico and Guatemala; occasional records from adjacent U.S.A. (Texas 
and southern Arizona), Honduras and El Salvador. Not threatened. Refs: 

51, 55, 62, 137, 169, 216, 234, 269. 

358 Papilio (Pterourus) palamedes Drury, 1773 

South-eastern U.S.A. (Pennsylvania, Maryland, West Virginia, North 
Carolina, South Carolina, Georgia, Florida, Tennessee (?), Alabama, 
Mississippi, Missouri, Arkansas, Louisiana, Texas and Nebraska (?)) and 
Mexico. Not uncommon and not known to be threatened. Two subspecies. 
Common name: Palamedes Swallowtail (154, 216, 230). Other refs: 55, 137, 
169, 269. 

359 Papilio (Pterourus) troilus Linnaeus, 1758 

Eastern Canada and eastern and south-eastern U.S.A. (to Texas). Gen- 
erally common and not threatened. Two subspecies. Common names: 
Spicebush Swallowtail (154, 216, 230), Green-clouded Swallowtail (216). 
Other refs: 55, 137, 141, 234, 269. 

360 Papilio (Pterourus) ascolius C. & R. Felder, 1864 

Nicaragua (?), Costa Rica, Panama, Colombia and western Ecuador, a 
subspecies is also found in French Guiana and possibly Guyana and 
Surinam (55). Unusual in being an ithomiine mimic. Rare in Costa Rica 
(61). Scarcer than P. (P.) zagreus but not thought to be threatened. More 
data required. Five subspecies. Other ref: 234. 

361 Papilio (Pterourus) neyi Niepeh, 1909 

Ecuador. A little-known species, probably only a form of P. (P.) zagreus 
(250). Further information is needed on both the taxonomic and conser- 
vation status of this species. Other refs: 51, 55. 

Species-group: zagreus Doubleday 

362 Papilio (Pterourus) zagreus Doubleday, 1847 

Colombia, Venezuela, Ecuador, Peru and Bolivia. Not uncommon and not 
known to be threatened. Two subspecies. P. (P.) chrysoxanthus Fruhstor- 
fer, 1915 is a synonym. Refs: 51, 55, 169, 234. 

363 Papilio (Pterourus) bachus C. & R. Felder, 1865 

Colombia, Ecuador, Peru and Bolivia. Not uncommon and not known to be 
threatened. Two subspecies. Refs: 51, 55, 234. 

Species-group: scamander Boisduval 

364 Papilio (Pterourus) hellanichus Hewitson, 1868 

Uruguay, Brazil and Argentina. Not common, but not recognized as 
threatened. Refs: 51, 55, 122, 234. 

85 



Threatened Swallowtail Butterflies of the World 

365 Papilio (Pterourus) scamander Boisduval, 1836 

Argentina, Brazil and possibly Paraguay. Not uncommon and not known to 
be threatened. Ranched in Brazil (24). Three (possibly four) subspecies. 
Considered synonymous with P. (P.) grayi Boisduval, 1836, P. (P.) 
eurymander Rothschild and Jordan, 1906 and P. (P.) joergenseni Rober, 
1925 (51). Other refs: 26, 55, 122, 234. 

366 Papilio (Pterourus) xanthopleura Godman and Salvin, 1868 

Peru (Upper Amazon) and Ecuador. It may also occur in eastern Bolivia. 
Uncommon and could possibly be designated Rare, but more information 
needed. Refs: 51, 55, 234. 

367 Papilio (Pterourus) birchalli Hewitson, 1863 

Costa Rica, Panama and Colombia. Widely distributed in Costa Rica, but 
seldom collected (61). Uncommon generally, though possibly less so than 
P. (P.) xanthopleura. Not known to be threatened. Two subspecies. Other 

refs: 51, 55, 234. 

Species-group: homerus Fabricius 

368 Papilio (Pterourus) cleotas Gray, 1832 

Costa Rica, Panama, Colombia, Venezuela, Brazil and northern Argentina 
(51, 55, 234). Possibly also in Paraguay and Bolivia. Rare in Costa Rica 
(61). Ranched in Brazil (24). Not known to be threatened. 

369 Papilio (Pterourus) menatius Hiibner, 1819 

Panama, Colombia, Guyana, Surinam, French Guiana, Brazil, Ecuador, 
Peru and Bolivia. Generally common and not known to be threatened. Up 
to ten subspecies. P. (P.) judicael Oberthiir, 1888 is considered a hybrid 
between P. (P.) menatius and P. (P.) warscewiczi by Hancock (109) but this 
has been questioned (221). P. (P.) aristeus Cramer, 1781 is a homonym. 
Other refs: 51, 55, 234. 

370 Papilio (Pterourus) phaeton (Lucas, 1857) 

Costa Rica, Panama, Colombia, Venezuela. Usually regarded as a subspe- 
cies of cleotas, but accepted by Hancock as a good species (109). Little 
information available, but not known to be threatened. Three subspecies. 
Other ref: 51. 

371 Papilio (Pterourus) victorinus Doubleday, 1844 

Mexico and Central America (excluding Panama). Rare in Costa Rica. 

r\ ,,, Flies at canopy level (61). Not believed to be threatened. Three subspecies. 

\ A' P- (P-) diazi Racheli and Sbordoni, 1975 was recently discovered in Mexico 

V^ (Morelos) (225). However according to Hancock (109, 112) and Tyler (268) 

it is of hybrid origin, probably between P. (P.) victorinus and P. (P.) 

garamas. Other refs: 51, 55, 62, 234, 269. 

372 Papilio (Pterourus) garamas Geyer, 1829 

Mexico. Not uncommon but not regarded as threatened. Up to five 
subspecies have been described from Central America but most of these 
probably belong to P. (P. ) abderus (115, 234) . Subspecies 5_yerfra is placed in 
garamas by DeVries (61), but this may be better placed under abderus. For 
the present purpose we retain garamas as a Mexican endemic, but a 
thorough taxonomic clarification is required. Other refs: 51, 62, 175, 261, 
269. 

86 



Af 



Swallowtails of the world 

373 Papilio (Pterourus) abderus Hopffer, 1856 
Mexico and Central America. Often regarded as a subspecies of P. (P.) 
garamas (see above), this taxon requires further study. Not known to be 
threatened. Four subspecies. Refs: 55, 62, 234, 269. 

374 Papilio (Pterourus) homerus Fabricius, 1793 
Endangered — refer to section 6, p. 297. 
Jamaica. This much-prized species was formerly classified as Vulnerable 

V^ but has now been raised to Endangered. Its mountain habitats seem to be 

threatened and specimens are still in demand (130). Its status should be 
closely monitored. Common name: Homerus Swallowtail (21, 230). Other 
refs:51,55, 234, 267, 283. 

375 Papilio (Pterourus) warscewiczi Hopffer, 1886 
Ecuador, Peru and Bolivia. Not uncommon and not threatened. Three 
subspecies. Refs: 51, 55, 234. 

376 Papilio (Pterourus) cacicus Lucas, 1852 
Colombia, Ecuador, Venezuela and Peru. Not common but not known to 
be threatened. Up to four subspecies. Refs: 51, 55, 234. 

377 Papilio (Pterourus) euterpinus Godman and Salvin, 1868 
Colombia, Ecuador and northern Peru. Scarce, possibly deserving Rare 
status, but not known to be threatened. More information needed. Two 
subspecies. Refs: 51, 55, 234. 

Subgenus: Heraclides Hiibner 

Species-group: thoas Linnaeus 

378 Papilio (Heraclides) esperanza Beutelspacher, 1975 
Vulnerable — refer to section 6, p. 299. 

Mexico (Oaxaca state). Very rare and with a restricted range. Little 
information available as its locality is kept secret. Refs: 18,51,62, 137, 175, 
268, 276. 



,,^ 






379 Papilio (Heraclides) andraemon Hiibner, 1823 




Cuba, Bahamas, Cayman Is and sporadically Florida (270). Introduced into 
Jamaica where it is a pest of Citrus. Otherwise not common but not 
threatened as a species. Three subspecies. Common names: Bahaman 



^\/^ Swallowtail (230), Andraemon Swallowtail (21). Other refs: 51, 55, 169, 

234, 269. 

380 Papilio (Heraclides) machaonides Esper, 1796 
Confined to Hispaniola (Haiti and Dominican Republic) and the Cayman Is 

' (Grand Cayman) (51). A report of its occurrence in Puerto Rico (230) 

seems doubtful (92). A common butterfly throughout Hispaniola (243) 
despite reports to the contrary (230), occurring from sea-level to high 
elevation. Common name: Machaonides Swallowtail (230). Other refs: 55, 
234. 

381 Papilio (Heraclides) thersites Fabricius, 1775 
Confined to Jamaica, where it is 'local and rather rare but widespread' (230) 
but is not threatened (266). Its status should be carefully monitored. 
Common names: False Androgeus Swallowtail (230), Thersites Swallowtail 
(21). Other refs: 55,234. 

87 







Threatened Swallowtail Butterflies of the World 

382 Papilio (Heraclides) astyalus Godart, 1819 

U.S.A. (southern Texas), Mexico, Central America and South America 
(except for Chile). At least the nominate subspecies is common and 
abundant. Encountered as solitary individuals along forest edges (61). 
Ranched in Brazil (24). Five subspecies (22). P. (H.) lycophron Hiibner is a 
synonym. Common name: the Lycophron (9). Other refs: 26, 51, 55, 62, 
137, 234, 269. 

^^ ^ 383 Papilio (Heraclides) androgens Cramer, 1775 

Mexico, Central America, Cuba, Hispaniola, Trinidad, Tobago, St. Lucia 

„. ,^ and South America (except Chile and Uruguay). Generally common and 

^ ^^ not threatened. Tolerant of open areas and secondary growth (61). Three 

TiY subspecies. Common names: the Androgens (9), Androgeus Swallowtail 

^ (230), Queen Page (9, 282). Other refs: 26, 51, 55, 62, 234, 269. 

384 Papilio (Heraclides) ornythion Boisduval, 1836 
U.S.A. (southern Texas), Mexico and Guatemala. Not uncommon, but not 
known to be threatened. Common name: Ornythion Swallowtail (154). 
Other refs: 51, 55, 57, 62, 137, 234, 269. 

385 Papilio (Heraclides) aristodemus Esper, 1794 
U.S.A. (Southern Florida), Cuba, Cayman Is, Hispaniola, Bahamas and 
possibly Puerto Rico. Not common anywhere, though probably not 
threatened as a species. Five subspecies (including two recently described 
from the Bahamas) (38). Common name: Dusky Swallowtail (230). Other 
refs: 51,55,216,234,269. 

Papilio (Heraclides) aristodemus ponceanus Schaus, 1911 

Endangered — refer to section 6, p. 301. 

The Floridan subspecies, poAJcefl/?j« Schaus, is an Endangered taxon and is 

protected under the U.S. Endangered Species Act (47, 137, 283). It has 

recently been relisted as Endangered (270). Common name: Schaus' 

Swallowtail (154, 216). 

386 Papilio (Heraclides) caiguanabus Poey, 1851 
Indeterminate— refer to section 6, p. 305. 

Restricted to Cuba, where it is rare (230): more information on its status and 
prospects is required. Common name: Poey's Black Swallowtail (230). 
Other refs: 51, 55, 234. 

387 Papilio (Heraclides) aristor Godart, 1819 
Indeterminate — refer to section 6, p. 307. 

Restricted to Haiti and the Dominican Republic (Hispaniola) where it is 
said to be very rare (230), but may be locally common in xeric areas (243). 
More data on status are required as there is little information available. 
Common name: Scarce Haitian Swallowtail (230). Other refs: 51, 55, 234. 

388 Papilio (Heraclides) thoas Linnaeus, 1771 
U.S.A. (Texas, Colorado and Kansas), Mexico, Central America, Cuba, 
Jamaica, Antigua, Trinidad and South America (possibly not Chile). Often 
common and not threatened. Ranched in Brazil (24). Seven, or eight 
subspecies. Common names: Thoas Swallowtail (154, 216, 230), King Page 
(9, 282), Citrus or Orange Swallowtail (282). Other refs: 21, 26, 51, 55, 61, 
62, 137, 150, 175, 234, 269. 

88 



Swallowtails of the world 

389 Papilio (Heraclides) cinyras Menetries, 1857 

Peru and eastern Ecuador to Brazil (central Amazonas) and Bolivia to 
Brazil (Mato Grosso) (51). Often regarded as a subspecies of thoas , but with 
a very distinct appearance. Little information, but not believed to be 
threatened. 

390 Papilio (Heraclides) homothoas Rothschild and Jordan, 1906 
Trinidad, Colombia and Venezuela. Although the range is fairly restricted, 
the species is not threatened. Common name: Small King Page (9). Other 
refs: 51,55,234. 

391 Papilio (Heraclides) cresphontes Cramer, 1777 

Southern Canada, U.S.A. (east of the Rockies and including the southern 
states), Mexico, Central America, Colombia (?), Cuba, possibly Bermuda 
and Bahamas. Widely distributed and not threatened. Common names: 
Giant Swallowtail (60, 154, 216, 230), Orange Dog (216). Other refs: 51, 55, 
57,61,62, 137,234,269. 

392 Papilio (Heraclides) paeon Boisduval, 1836 

Costa Rica, El Salvador, Colombia, Venezuela, Ecuador, Peru and 
Bolivia. Not uncommon and no threats recognized. Three, possibly four, 
subspecies. Refs: 51, 55, 57, 234. 

Species-group: torqiiatus Cramer 

393 Papilio (Heraclides) garleppi Staudinger, 1892 
Insufficiently Known — refer to section 6, p. 309. 

Guyana, Surinam, French Guiana, Bolivia, Brazil and Peru. Not known to 
be in danger but reported to be a very rare species (268). Other refs: 51 , 55, 
234. 

394 Papilio (Heraclides) torquatus Cramer, 1777 

Mexico, Central America, Trinidad and South America (except Chile and 
Uruguay). Generally common and not threatened. Six subspecies including 
P. (H.) t. tolas Godman and Salvin, 1890 from Mexico, possibly Belize and 
Guatemala. P. (H.) tolus is generally regarded as a subspecies of P. (H.) 
torquatus (51, 234) and by only a few authorities as a good species (175). P. 
(H.) tasso (Staudinger, 1884) is an aberration of P. (H.) torquatus (109). 
Common name: the Torquatus (9). Other refs: 26, 55, 61, 62, 169, 269. 

395 Papilio (Heraclides) hectorides Esper, 1794 

South-eastern Brazil, Paraguay, northern Argentina and Uruguay (51). 
Not uncommon and not considered to be threatened. Ranched in Brazil 
(24). Other refs: 55, 122, 169, 234. 

396 Papilio (Heraclides) lamarchei Staudinger, 1892 

Brazil (near the Argentine border), Bolivia and northern Argentina. 
Uncommon but not known to be threatened. Refs: 51, 55, 122, 234. 

397 Papilio (Heraclides) himeros Hopffer, 1866 
Vulnerable — refer to section 6, p. 311. 

Brazil and according to D' Almeida (55) also from Corrientes in northern 
Argentina. Two subspecies, both of which are exceedingly rarely 
encountered and decHning (23). Other refs: 26, 51, 122, 139. 234. 



Threatened Swallowtail Butterflies of the World 

Species-group: anchisiades Esper 

398 Papilio (Heraclides) hyppason Cramer, 1775 
Venezuela (?), Guyana, Surinam, French Guiana, Brazil, Peru and Bolivia. 
Not uncommon and apparently not threatened. Refs: 51, 55, 169, 234. 

399 Papilio (Heraclides) pelaus Fabricius, 1775 
Cuba, Jamaica, Hispaniola and Puerto Rico. Apparently not uncommon 
but widely distributed and not threatened. Three subspecies. Common 
names: Prickly Ash Swallowtail (230), Pelaus Swallowtail (21). Other refs: 
51,55,234. 

400 Papilio (Heraclides) oxynias (Geyer, 1827) 
Cuba. Widespread in that country, according to Riley (230), but status 
requiring clarification. Common name: Cuban Black Swallowtail (230). 
Other refs: 51,55, 169,234. 

401 Papilio (Heraclides) epenetus Hewitson, 1861 
Known only from western Ecuador. Certainly uncommon but vulnerability 
not known and more information needed. Larvae feed on Citrus. Refs: 51, 
55, 169, 234. 

402 Papilio (Heraclides) erostratus Westwood, 1847 
Eastern Mexico (not western as given by D'AImeida (55)), Guatemala and 
Belize. Not common, but not known to be threatened. Other refs: 51, 62, 
234, 269. 

403 Papilio (Heraclides) erostratinus Vazquez, 1947 
Mexico. Described fairly recently but known from several localities and not 
thought to be threatened. Tyler tentatively suggests it might be a northern 
subspecies of P. (H.) erostratus (269). Other refs: 51, 55, 62. 

404 Papilio (Heraclides) pharnaces Doubleday, 1846 
Southern Mexico and possibly Guatemala (51). Not uncommon and, 
despite the narrow range, apparently not threatened. Other refs: 55, 62, 
234, 269. 

405 Papilio (Heraclides) chiansiades Westwood, 1872 
Ecuador and Peru (Upper Amazon). Generally common and not threat- 
ened. Refs: 51,55, 169,234. 

406 Papilio (Heraclides) dospassosi Rutimeyer, 1969 
South-east Colombia, Putumayo Valley (236). Listed by Hancock (109) but 
not by D'Abrera (51). No further information on taxonomic status or 
threats. This species is apparently only known from the type specimen. 
Possibly only a subspecies of Eury tides trapeza (221). It could be a candidate 
for conservation concern if shown to be a good species. 

J^Ay 407 Papilio (Heraclides) anchisiades Esper, 1788 

Southern U.S.A. (Texas), Mexico, Central America, South America 
Ss^ .. (excluding Chile and Uruguay) and Trinidad. Widespread and generally 

1^ ., ,>y ' very abundant; not threatened. Tolerant of a very wide range of habitats. 

Ranched in Brazil (24). Three or four subspecies (all apparently common). 

Common names: Ruby-spotted Swallowtail (216), Orange Dog (9, 282). 

Other refs: 26, 51, 55, 57, 62, 141, 154, 169, 234, 269. 

90 



,5 



Swallowtails of the world 

408 Papilio (Heraclides) maroni Moreau, 1923 
Insufficiently Known — refer to section 6, p. 313. 

Known only from French Guiana (male and female) (186). but very little 
information available. Smart (250) notes its similarity to the common 
species P. (H.) isidorus. Other refs: 51, 55, 139, 187. 

409 Papilio (Heraclides) rogeri Boisduval, 1836 

Mexico (Yucatan), Belize and possibly Guatemala (51), a restricted range. 
The species seems to be fairly common, at least in Yucatan, and not 
threatened. Other refs: 55, 62, 234, 269. 

410 Papilio (Heraclides) isidorus Doubleday, 1846 

-O Costa Rica (61), Panama, Colombia, Ecuador, Bolivia and Peru. Generally 

common and not threatened. Apparently very rare, montane at its northern 
limits in Costa Rica, not collected in recent years (61). Four subspecies. 
Other refs: 51,55,234. 

411 Papilio (Heraclides) rhodostictus Butler & Druce. 1874 

Panama, Costa Rica, Colombia and western Ecuador. Appently not 
uncommon , at least in the southern part of its range , but rarely recorded in 
Costa Rica (61). Not known to be threatened. Three subspecies. Other refs: 
51,55,234. 

Subgenus: Eleppone Hancock 

Species-group: anactus Macleay 

412 Papilio (Eleppone) anactus Macleay, 1826 

Eastern Australia and possibly New Caledonia (vagrants?) (133). Common 
in at least parts of its range and not threatened. Common name: Dingy 
Swallowtail (282). Other refs: 40, 48, 106, 141, 176. 

Subgenus: Chilasa Moore 

Species-group: elwesi Leech 

413 Papilio (Chilasa) elwesi Leech, 1889 

Eastern and central China. Scarce and not well known, but not thought to 
be threatened. The subspecies P. (C.) e. maraho (see below) has been 
raised to a full species. Refs: 52, 141. 

414 Papilio (Chilasa) maraho (Shiraki and Sonan, 1934) 
Vulnerable — refer to section 6, p. 315. 

Taiwan. Sometimes regarded as a subspecies of P. (C.) elwesi (52), but 
Igarashi (141), Hancock (109) and Haugum (118) believe that it is quite 
distinct. It certainly merits conservation concern due to habitat destruction 
within its restricted distribution and the high prices paid for it by collectors. 
It has been suggested that this species is rarely collected because of its 
montane distribution (118), but Taiwan is such a small and crowded island 
that all populations of the butterfly are considered to be at risk. Other ref: 
246. 

Species-group: clytia Linnaeus 

415 Papilio (Chilasa) clytia Linnaeus, 1758 
India, Sri Lanka, Nepal, Bhutan (?), Bangladesh, Burma, Thailand, 
southern China (including Hainan (Guangdong prov.)), Hong Kong, 

91 




Threatened Swallowtail Butterflies of the World 

Vietnam, Laos, Kampuchea, Peninsular Malaysia, Philippines, Andamans 
and Indonesia (Flores, Alor, Timor and Moa) (52, 262). Generally common 
and not threatened, but the nominate subspecies is protected by law in India 
(182). Up to seven subspecies. Common name: Common Mime (45, 80, 
251). Other refs: 4, 5, 48, 87, 141, 143, 259, 287. 

416 Papilio (Chilasa) paradoxa Zinken-Sommer, 1831 

Northern India (including Assam), Bangladesh, Burma, southern China 
(?), Thailand, Vietnam, Laos, Kampuchea, Peninsular and Eastern Malay- 
sia, Philippines (Palawan), Brunei and Indonesia (Sumatra, Bangka, Nias, 
Batu and Kalimantan) (52, 262). Never common (269) but not known to be 
threatened. However, P. (C.) paradoxa telearchus is protected by law in 
India (182). A number of forms, mimicking different Danainae. Seven 
subspecies. Common name: Great Blue Mime (3, 45, 80, 282). Other refs: 
4, 87, 131. 

Species-group: veiovis Hewitson 

417 Papilio (Chilasa) veiovis Hewitson, 1853 

Indonesia (Sulawesi). Not uncommon, but not known to be threatened. 
Refs: 52, 262. 

418 Papilio (Chilasa) osmana Jumalon, 1967 
Vulnerable — refer to section 6, p. 317. 

Philippines (southern Leyte and northern Mindanao). Scarce everywhere 
in its restricted range (262). Other refs: 52, 148. 

419 Papilio (Chilasa) carolinensis Jumalon, 1967 
Vulnerable — refer to section 6, p. 320. 

Philippines (north-eastern Mindanao). Restricted range, claimed by Tsu- 
kada and Nishiyama (262) to be the rarest Philippine swallowtail. Other 
refs: 52, 148. 

Species-group: agestor Gray 

420 Papilio (Chilasa) agestor (Gray, 1832) 

Northern India, Nepal, Bhutan (?), Bangladesh (?), Burma, Thailand, 
central and southern China, Taiwan, Vietnam, Laos, Kampuchea (?) and 
Peninsular Malaysia (52). Not threatened across most of its range but 
considered to be Vulnerable in Peninsular Malaysia (10). Six subspecies. 
Common name: Tawny Mime (80, 251). Other refs: 45, 87, 141, 246, 259. 

421 Papilio (Chilasa) epycides (Hewitson, 1862) 

Northern India (Assam, Sikkim and Manipur), Bhutan, Burma, Thailand, 
southern China, Taiwan, Vietnam (?), Laos (?) and Peninsular Malaysia 
(?). No known threats to the species, although the nominate subspecies is 
protected by law in India (182). Six subspecies. Common name: Lesser 
Mime (80). Other refs: 4, 45, 87, 141, 246, 259. 

422 Papilio (Chilasa) slateri (Hewitson, 1857) 

Northern India (Sikkim and Assam), Bhutan (?), Burma, Thailand, 
southern China (?), northern Vietnam, Laos, Peninsular and Eastern 
Malaysia, Brunei and Indonesia (Kalimantan (?) and northern Sumatra) 
(52, 262). Not known to be threatened as a species but some subspecies may 
be rare and the nominate subspecies is protected by law in India (182) . Four 

92 



Swallowtails of the world 

(262) to six (52) subspecies. Common names: Blue-striped Mime (80), 
Brown Mime (3). Other refs: 4, 45, 87, 131. 141. 

Species-group: laglaizei Depuiset 

423 Papilio (Chilasa) laglaizei Depuiset, 1877 
Indonesia (Irian Jaya, Waigeo and Aru) and Papua New Guinea. Local but 
not threatened. Refs: 14, 48, 141, 256. 

424 Papilio (Chilasa) toboroi Ribbe, 1907 
Rare — refer to section 6, p. 322. 

Papua New Guinea (Bougainville (203)) and Solomons (Santa Isabel and 
Malaita). Said to be common (48), but the price of specimens is relatively 
high (142). It is rarely collected for trade in Papua New Guinea (36). More 
information is needed. Two subspecies. Other refs: 132, 141 , 218, 219, 228, 
256. 

425 Papilio (Chilasa) moerneri Aurivillius, 1919 
Vulnerable — refer to section 6, p. 324. 

Restricted to the Bismarck Archipelago (New Ireland and possibly New 
Britain and New Hanover). Known from very few specimens. It was 
regarded as extinct by D'Abrera (48) but this is not certain as large areas of 
potential habitat remain unexplored (83). Two subspecies have been 
described. Other refs: 6, 217, 235. 

Subgenus: Papilio Linnaeus 

Species-group: machaon Linnaeus 

426 Papilio (Papilio) alexanor Esper, 1799 
Southern France, southern Italy, Sicily, Yugoslavia, Albania, Greece, 
Turkey, Syria, Lebanon, Israel, Jordan, Iraq, Iran, western Pakistan, 
Afghanistan (including Nangrahar province) and U.S.S.R. (Turkmenis- 
tan). Extremely rare in Pakistan (Baluchistan) (259). Uncommon in the 
western part of its range and listed as Vulnerable in Europe (123), but 
apparently not threatened as a species. Declining in the U.S.S.R. due to 
degradation of alpine meadows and included in the U.S.S.R. Red Data 
Book, category Vulnerable (8, 260). Extinct in Yugoslavia, Endangered in 
the Balkan Peninsula and Italy (123), protected by law in Greece. Six 
subspecies. Common names: Southern Swallowtail (125, 282), Tiger 
Swallowtail (158), Baluchi Yellow Swallowtail (80). Other refs: 141, 169. 

427 Papilio (Papilio) hospiton Guenee, 1839 
Endangered — refer to section 6, p. 326. 

Endemic to Corsica and Sardinia; status not thoroughly known, but giving 
rise to concern and listed as Endangered (123). Trade is restricted by law in 
France and the Netherlands. Common name: Corsican Swallowtail (125). 
Other ref: 202. 

428 Papilio (Papilio) machaon Linnaeus, 1758 
, Entire Palearctic region through the U.S.S.R. to China (164, 207) and 

(, J^ Japan (including Nepal, Bhutan and Taiwan) as far south as some oases in 

r<f^ .Xj/ Saudi Arabia and Oman and the high mountains of Yemen (161); Pakistan, 

. ■ , V northern India, northern Burma, Canada and Alaska. Often common and 

-^K ^ not threatened as a species. Thirty seven subspecies (141). P. (P.) m. 

93 




^ 



Threatened Swallowtail Butterflies of the World 

saharae flies in Yemen and parts of the Sahara ( 161 ) . Larsen (161) lists it as a 
full species, noting that its larva is very different to that of machaon in other 
parts of Arabia. Papilio (P.) machaon is protected by law in six of the 
Austrian provinces, Czechoslovakia, Hungary, the U.K. (subspecies 
britannicus) and India (subspecies verityi) (182). It is listed as Vulnerable in 
South Korea (152), the U.S.S.R. Red Data Book (8) and the Austrian Red 
Data Book (93). Common names: Swallowtail (3, 125, 282), Old World 
Swallowtail (154, 216), Common Yellow Swallowtail (80, 251). Other refs: 
52, 55, 137, 149,, 169, 202, 246, 259, 269. 

429 Papilio (Papilio) hippocrates C. & R. Felder, 1864 

Japan. Generally regarded as a subspecies of P. (P.) machaon (141, 149, 
279), but accepted as a full species by Hancock (109). 

430 Papilio (Papilio) zelicaon Lucas, 1852 

Mexico, western U.S.A. including Alaska, and Canada (Yukon Territory 
(?), British Columbia, Alberta, Saskatchewan and Manitoba). Common 
and not threatened. P. (P.) gothica Remington from Canada (British 
Columbia and Alberta) , western U.S.A. and Mexico is widely regarded as a 
biological race or subspecies of P. (P.) zelicaon or as the yellow form of P. 
(P.) nitra (109). Not threatened. Common name: Anise Swallowtail (216). 
Other refs: 55, 62, 137, 141, 169, 226, 234, 244, 269. 

431 Papilio (Papilio) indra Reakirt, 1866 

Western U.S.A. (Washington, Oregon, California, Nevada, Utah, Colo- 
rado and possibly Arizona). Uncommon but not generally threatened. 
Listed as a Special Species by Washington State Department of Game. 
Seven subspecies. Common names: Indra Swallowtail (216), Short-tailed 
Black Swallowtail (175). Other refs: 62, 77, 78, 79, 137, 169, 234, 269. 

432 Papilio (Papilio) polyxenes Fabricius, 1775 (= ajax L., 1758) 
Southern Canada, U.S.A., Cuba, Mexico, Central America, Colombia, 
Venezuela, Ecuador and northern Peru. Widely distributed, often common 
and not threatened. Populations may fluctuate widely in numbers (61). 
Four subspecies. Common names: Eastern Black Swallowtail (216), 
American Black Swallowtail (230), Parsnip Swallowtail (154). Other refs: 
55, 62, 137, 141, 169, 234, 269. 

433 Papilio (Papilio) brevicauda Saunders, 1869 

Eastern Canada (southern Labrador, Newfoundland, Quebec, New Bruns- 
wick, Nova Scotia), mainly maritime. It has been described as a subspecies 
of P. (P.) polyxenes (234) or of P. (P.) ajax (55) the latter a synonym of the 
former. Not particularly common but not known to be threatened. Three 
subspecies (not well defined) (137). Common names: Short-tailed Swallow- 
tail (154, 216), Maritime Swallowtail (216). Other refs; 84, 85, 269. 

434 Papilio (Papilio) oregonia Edwards, 1876 

North-western U.S.A. (eastern Washington, Idaho, western Montana, 
North Dakota (?) and eastern Oregon) and south-western Canada (British 
Columbia and Alberta (?)). Possibly a form of P. (P.) bairdi (55, 234, 250). 
Not common but not recognized as being widely threatened. Listed as a 
Special Species by Washington State Department of Game. Two subspe- 
cies. Common name: Oregon Swallowtail (216). Other refs: 65, 137, 141, 
213, 269. 

94 



Swallowtails of the world 




Vv 



435 Papilio (Papilio) kahli F. & R. Chermock, 1937 

Canada (Manitoba and Saskatchewan). Specific status doubtful (250, 269). 
' Regarded as a subspecies of P. (P.) nitra by D'Almeida (55). Restricted 
range, but not recognized as threatened. Other refs: 137, 216. 

436 Papilio (Papilio) nitra Edwards, 1883 

Canada (Alberta) and eastern side of Rocky Mountains in U.S.A. 
(Montana, Wyoming and Colorado). Another taxon of doubtful specific 
status, probably a form or subspecies of P. (P.) zelicaon . Occurs early in the 
year in high mountains and so rarely taken, but not believed to be 
threatened. Refs: 55, 137, 216, 234, 269. 

437 Papilio (Papilio) coloro Wright, 1905 {=nidkini Chermock, 1977) 
U.S.A. (southern Nevada, south-eastern Utah, southern California and 
western Arizona). Specific status doubtful (250, 269), possibly a form of P. 
(P.) bairdi (55), P. (P.) polyxenes (118) or P. (P.) zelicaon. Restricted 
range, but not recognized as threatened. Common name: Desert Swallow- 
tail (216). Other refs: 86, 137. 

Papilio (Papilio) bairdi Edwards, 1869 
Mid-western U.S.A. (Montana (?), Idaho, Wyoming, South Dakota, 
Nebraska, Utah, Colorado, Arizona and New Mexico). Not known to be 
threatened. Common names: Western Black Swallowtail (216), Baird's 
Swallowtail (216). Other refs: 55, 62, 137, 141, 234, 269. 

439 Papilio (Papilio) joanae Heitzman, 1974 

U.S.A. (Missouri only). Specific status not certain, possibly a form of P. 
(P.) polyxenes (250); it is not included in Howe (137). The range is 
restricted, but the butterfly is not regarded as a threatened species. Other 
refs: 216, 269. 

Subgenus: Princeps Hiibner 

Species-group: xiithus Linnaeus 

440 Papilio (Princeps) xuthiis Linnaeus, 1760 

Southern China, northern Burma, Japan, Taiwan, Guam, Ogasawara- 
shoto (Bonin I.), Wake (?), Midway Is (?) and Hawaii. Common and not 
threatened. Two subspecies. Common name: Chinese Yellow Swallowtail 
(80). Other refs: 52, 92, 141, 149, 207. 

441 Papilio (Princeps) benguetaniis Joicey and Talbot, 1923 
Vulnerable — see section 6, p. 329. 

Philippines (Luzon only). Occasionally regarded as a subspecies of P. (P.) 
xuthus, but very different in appearance. Refs: 52, 141, 148, 262. 

Species-group: demolion Cramer 

442 Papilio (Princeps) euchenor Guerin-Meneville, 1829 

Indonesia (Irian Jaya) , Papua New Guinea, Bismarck Archipelago (includ- 
ing New Britain), D'Entrecasteaux Is, Trobriand Is (?) and the Louisiade 
Archipelago (?). Not uncommon and not threatened as a species though 
some subspecies may be threatened ( 138). Collected for trade in Papua New 
Guinea. This species was separated as the type of a monobasic genus, 
Sugurua, by K. Okano in 1983 (198) but this has not been generally 
accepted. Thirteen subspecies. Other refs: 14, 48, 52, 141, 217, 262. 



95 



Threatened Swallowtail Butterflies of the World 

443 Papilio (Princeps) gigon C. & R. Felder, 1864 

Indonesia (Sulawesi, Banggai Is (?), Sula, Talaud and Sangihe). Not 
uncommon and not known to be threatened. Three subspecies. Refs: 52, 
141,262. 

444 Papilio (Princeps) demolion Cramer, 1779 

Burma, Thailand, Peninsular and Eastern Malaysia, Brunei, Philippines 
(Palawan) and Indonesia (Sumatra, Mentawai Is, Nias, Bangka, Java, 
Lombok, Sumbawa and KaHmantan) (52, 191, 262). Fairly common and 
not threatened. Three subspecies. Common names: Banded Swallowtail 
(3, 45), Burmese Banded Swallowtail (80). Other ref: 141. 

445 Papilio (Princeps) liomedon Moore, 1874 

Southern India. Uncommon, but protected by law (182) and not known to 
be threatened. It has been considered to be only a subspecies of P. (P.) 
demolion (80), but is apparently distinct (144). Common name: Malabar 
Banded Swallowtail (80). Other refs: 4, 52. 

Species-group: protenor Cramer 

446 Papilio (Princeps) protenor Cramer, 1775 

Northern India, (including Kashmir and Jammu, Sikkim and Assam), 
Bangladesh, Burma, southern China (including Hainan (Guangdong 
prov.)), northern Vietnam (?), northern Laos (?), Taiwan, North Korea, 
South Korea and Japan. Common and not threatened. Five subspecies. 
Common name: the Spangle (80, 251). Other refs: 4, 52, 141, 149, 153, 207, 
246, 259. 

447 Papilio (Princeps) demetrius Cramer, 1782 

Japan, North Korea, South Korea and China. Not uncommon except in 
Korea where it is considered to be Extinct or Endangered (152, 153). 
Losses in Korea suggest that monitoring throughout its range might be 
advisable. Sometimes considered to be only a subspecies of P. (P.) protenor 
(141). Other refs: 149,247. 

448 Papilio (Princeps) macilentus Janson, 1877 

Japan, North Korea, South Korea where it is Vulnerable (152) and eastern 
and central China. Common and not threatened. Refs: 141, 149, 247. 

Species-group: bootes Westwood 

449 Papilio (Princeps) bootes Westwood, 1842 

North-eastern India (Uttar Pradesh, Sikkim, Assam, Meghalaya, Manipur 
and Nagaland), northern Burma, northern Laos and south-western China, 
a restricted range. Not common but not regarded as threatened. It is 
protected by law in India (182). Three subspecies (not including P. (P.) 
janaka — see below). Common name: Tailed Redbreast (80). Other refs: 4, 
52, 141, 259. 

450 Papilio (Princeps) janaka Moore, 1857 

Northern India (including Sikkim) and Nepal. Often regarded as only a 
subspecies of P. (P.) bootes (see above) (80, 141). Not known to be 
threatened. Common name: Tailed Redbreast (80, 251). Other refs: 4, 52. 

96 



Swallowtails of the world 

Species-group: memnon Linnaeus 

451 Papilio (Princeps) lampsacus Boisduval, 1836 

Indonesia (western Java), only known from the localities of Mt Gede and 
Mt Mas. Apparently rather scarce although said to be abundant on Mt Gede 
by Tsukada and Nishiyama (262) who say "It is regrettable that collecting is 
not permitted on Mt Gede now as it rises within the National Park area". 
Further information and monitoring of this restricted species are needed. 
Signs of habitat disruption would necessitate immediate listing as threat- 
ened. Other refs: 45, 52. 

452 Papilio (Princeps) forbesi Grose-Smith, 1883 

Indonesia (northern Sumatra). Despite the restricted range, abundant in 
some localities, not uncommon and not known to be threatened. Female 
mimics Atrophaneura {Atrophaneura) hageni (262). Other refs: 45, 52. 

453 Papilio (Princeps) acheron Grose-Smith, 1887 
Rare — refer to section 6, p. 331. 

Northern Borneo: Sabah, Sarawak (Eastern Malaysia), Brunei and 
possibly Kalimantan (Indonesia). Conservation concern is the result of its 
restricted known range and recent evidence of deforestation on Mt 
Kinabalu. The taxonomic status of this species in relation to the two 
preceding needs clarification (45). Other refs: 11, 52, 131, 231, 262. 

454 Papilio (Princeps) oenomaus Godart, 1879 '^ ' ^ :^\ , .- .' ' " •**: ^^^5 f«^ '^^ 
Indonesia (Lesser Sunda Is: Timor, Wetar, Moa, Leti, Kisar and Romang) 

(262). Common (48) and presumably not threatened. Two subspecies. 

455 Papilio (Princeps) ascalaphus Boisduval, 1836 

Indonesia (Sulawesi, Salayar and Sula Is). Restricted range, but not 
uncommon and not known to be threatened. Two subspecies. Refs: 141, 
262. 

456 Papilio (Princeps) rumanzovia Eschscholtz, 1821 

Philippines (except Palawan), Batu Is and Indonesia (Talaud and Sangihe). 
Also known from stray butterflies in southern Taiwan but not established 
(52, 262). Not uncommon and not thought to be threatened. Other ref: 141. 

457 Papilio (Princeps) deiphobus Linnaeus, 1758 

Indonesia (Moluccas). Not common but not thought to be threatened. At 
least two subspecies. Ref: 109. 

458 Papilio (Princeps) alcmenor C. & R. Felder, 1864 

North-eastern India, Nepal, Bhutan, Burma and China. Generally 
common and not threatened. This species was formerly known as P. (P.) 
rhetenorWestwood, 1841. Common name; Redbreast (80, 251). Other refs: 
4, 52, 207, 259. 

459 Papilio (Princeps) thaiwanus Rothschild, 1898 

Taiwan only. Common, despite its restricted range and not therefore Hsted, 
though its status should clearly be monitored. Refs: 52, 141, 246. 

460 Papilio (Princeps) polymnestor Cramer, 1775 

Southern India and the east coast, and Sri Lanka. Not uncommon and not 
thought to be threatened. At least two subspecies. Common name: Blue 
Mormon (80). Other refs: 4, 52, 141, 259, 287. 

97 




Threatened Swallowtail Butterflies of the World 

461 Papilio (Princeps) lowi Druce, 1873 

Philippines (Palawan and Balabac) (52, 262) . D' Abrera claims that it is also 
found in northern Borneo (52). Despite restricted range apparently not 
uncommon and not threatened. Males are seen widely but the females are 
more secretive; the young can be raised on Citrus (56). 

462 Papilio (Princeps) memnon Linnaeus, 1758 

North-eastern India (Sikkim, Assam and Nagaiand), Nepal, Bhutan (?), 
Bangladesh, Nicobar Is, Andaman Is (stragglers only, 5), Burma, western, 
southern and eastern China (including Hainan), Taiwan, southern Japan, 
Ryukyu Is, Thailand, Laos, Vietnam, Kampuchea, Eastern and Peninsular 
Malaysia, Brunei and Indonesia (Sumatra, Mentawai Is, Nias, Batu, 
Simeulue, Bangka, Java, Kalimantan and the Lesser Sunda Is (except 
Timor, Wetar, Babar and Tanimbar)) (52, 191, 262). Common (especially 
since the cultivation of Citrus (45)) and not threatened. Up to 13 subspecies, 
highly mimetic and polymorphic in the female. Common name: Great 
Mormon (45, 80, 251). Other refs: 87, 131, 141, 149, 207, 231, 246, 259. 

463 Papilio (Princeps) mayo Atkinson, 1873 

Endemic to the Andamans. A little known butterfly for which more 
information is needed. Not rare according to Evans (80), though Tsukada 
and Nishiyama (262) suggest that it may be uncommon. Arora and Nandi 
state that males are common, females rare (5), and it is protected by Indian 
law (182). The female mimics Atrophaneura (Losaria) rhodifer Atkinson. 
Common name: Andaman Mormon (80). Other refs: 52, 259. 

Species-group: helenus Linnaeus 

464 Papilio (Princeps) noblei de Niceville, 1889 

Burma, Thailand, Laos and Vietnam (52). Said to be rare (80, 259), but 
current status not known. Two subspecies. Common name: Noble's Helen 
(80). Other refs: 144, 168. 

465 Papilio (Princeps) antonio Hewitson, 1875 

Philippines (Mindanao, Leyte and possibly Negros) (52, 262). Not known 
to be threatened but possibly could be classed as Rare. Poorly known and 
locally distributed but not always rare in its locality when found (262) . Other 
ref: 144. 

466 Papilio (Princeps) biseriatus Rothschild, 1895 

Indonesia (restricted to Timor) . Usually regarded as a subspecies of P. (P.) 
helenus, but recognized as a full species by Hancock ( 109). Further data may 
reveal threats to this very restricted taxon. 

467 Papilio (Princeps) iswara White, 1842 

Southern Burma, southern Thailand, Eastern and Peninsular Malaysia, 
Brunei and Indonesia (Sumatra, Bunguran, Bangka and Kalimantan) (52, 
262). Not uncommon and not threatened. Two subspecies. Common 
names: Great Helen (80), Large Helen (3). Other refs: 45, 87, 259. 

468 Papilio (Princeps) hystaspes (C. & R. Felder, 1862) 

Philippines from Luzon south to Mindanao, excluding Palawan. Sometimes 
regarded as a subspecies of helenus (52) but recognized as a full species by 
Hancock (109) and Tsukada and Nishiyama (262). Widespread and not 
known to be threatened. 

98 



Swallowtails of the world 

469 Papilio (Princeps) sataspes C. & R. Felder, 1864 

Indonesia (Sulawesi, Banggai and Sula). There is no evidence that it is 
threatened. Three subspecies (262). Other ref: 109. 

470 Papilio (Princeps) helenus Linnaeus, 1758 

Southern and north-eastern India, Nepal, Sri Lanka, Burma, Thailand, 
Bangladesh, Bhutan, southern China (including Hainan (Guangdong 
prov.)), Vietnam, Laos, Taiwan, southern Japan, South Korea, Ryukyu Is, 
Kampuchea, Peninsular and Eastern Malaysia, Brunei, Philippines and 
Indonesia (Sumatra, Java, Bangka, Kalimantan and the Lesser Sunda Is 
(not Tanimbar)) (52, 262). Common and not threatened. Up to thirteen 
subspecies. Common name: Red Helen (45, 80, 251). Other refs: 4, 87, 131, 
141, 149,246,259. 

471 Papilio (Princeps) iswaroides Fruhstorfer, 1897 

Peninsular Malaysia, Indonesia (northern Sumatra) and possibly the 
extreme south of Thailand. It has been suggested that it may also occur in 
northern Borneo (262), but there has been no record so far. Two 
subspecies. Other refs: 45, 52. 

Species-group: polytes Linnaeus 

472 Papilio (Princeps) jordani Fruhstorfer, 1902 
Rare — refer to section 6, p. 333. 

Indonesia (endemic to northern Sulawesi) where it is reported to be 
extremely rare (262). The female mimics Idea blanchardii (Danainae). 
Other refs: 52, 121, 144. 

473 Papilio (Princeps) polytes Linnaeus, 1758 

India, Nepal, Sri Lanka, Burma, Thailand, southern and western China 
(including Hainan (Guangdong prov . ) ) , Taiwan , Ryukyu Is (Japan) , Guam 
(221), Vietnam, Laos, Kampuchea, Nicobar Is, Andamans, Eastern and 
Peninsular Malaysia, Brunei and Indonesia (except Moluccas and Irian 
Jaya) (52, 262). Common and not threatened. Here we follow Hiura and 
Alagar (127) and Tsukada and Nishiyama (262) who regard the subspecies 
from the Philippines and Moluccas as a separate species, P. (P.) alphenor 
Cramer (see below). Seventeen subspecies (23 if P. (P.) alphenor is 
included)(262). P. (P.) waZ/cenJanson from southern India is probably only 
a form of P. (P.) polytes (4, 80, 259). It is very uncommon and would be 
classed as Rare if a good species. P. (P.) sakontala Hewitson, 1864 from 
northern India (including Sikkim and Assam) is another taxon thought by 
some authors to be a form of P. (P.) polytes rather than a good species (4, 
80). It also is uncommon. Common name: Common Mormon (45, 80). 
Other refs: 5, 48, 87, 131, 141, 143, 144, 149, 207, 246, 287. 

474 Papilio (Princeps) alphenor (Cramer, 1776) 

Indonesia (Moluccas, Sangihe and Talaud Is) and the Philippines (including 
Palawan). Earlier regarded as a subspecies of P. (P.) polytes, but separated 
as a full species in the revisionary work by Hiura and Alagar (127). 
Widespread and not known to be threatened. Other ref: 262. 

475 Papilio (Princeps) ambrax Boisduval, 1832 

Papua New Guinea (including Woodlark I. (184), and Fergusson I. in the 
Entrecasteaux Is), Indonesia (Irian Jaya and Aru) and north-eastern 

99 



Threatened Swallowtail Butterflies of the World 

Australia. Collected for trade in Papua New Guinea (36). Not rare and not 
threatened. Five subspecies. Other refs: 14, 40, 48, 141, 176. 

476 Papilio (Princeps) phestus Guerin-Meneville, 1830 

Papua New Guinea (Bismarck Archipelago (including New Britain and 
Admiralty Is) and Bougainville) and the Solomons (except Malaita, 
Guadalcanal and San Cristobal) (219). Common and not known to be 
threatened. Four subspecies. Other refs: 48, 68. 

Species-group: nephelus Boisduval 

477 Papilio (Princeps) diophantus Grose-Smith, 1882 

Indonesia (northern Sumatra). Uncommon. Not thought to be threatened, 
though more information needed. The female is very scarce in collections. 
Refs: 45, 52, 144, 262. 

478 Papilio (Princeps) nephelus Boisduval, 1836 

Nepal, north-eastern India (including Assam and Sikkim), Bhutan (?), 
Burma, Thailand, Vietnam, Laos, southern China (including Hainan 
(Guangdong prov.), Taiwan, Kampuchea, Eastern and Peninsular Malay- 
sia, Brunei and Indonesia (Sumatra, Mentawai Is, Nias, Batu, Java and 
Kalimantan) (52, 191, 262). Common and not threatened. Eleven subspe- 
cies including chaon Westwood (115). P. (P.) nubilus Staudinger from 
Sumatra and northern Borneo was placed as a possible aberration of P. (P) 
nephelus, or a hybrid , by Hancock (109) and is regarded as a hybrid between 
P. (P. ) nephelus and P. (P. ) polytes by Tsukada and Nishiyama (262) . More 
information required. Common names: Yellow Helen (80, as P. chaon). 
Black and White Helen (3, 45), Banded Helen (3). Other refs: 87, 131 , 141, 
231,246. 

479 Papilio (Princeps) castor Westwood, 1842 

North-eastern India (Assam, Sikkim), Bhutan, Thailand, southern China, 
Taiwan, Vietnam, Laos and possibly Kampuchea (52). Common and not 
threatened. Deceptively similar to members of the subgenus Chilasa (52). 
Seven subspecies. Common name: Common Raven (80). Other refs: 4, 
141,246. 

480 Papilio (Princeps) mahadeva Moore, 1878 

Burma, Thailand, Laos (?), Vietnam (?), Kampuchea (?) and northern 
Peninsular Malaysia. Definitely uncommon but probably not meriting 
conservation status, though more information is needed. May be Vulnera- 
ble and in need of protection in Peninsular Malaysia (10). Deceptively 
similar to members of the subgenus Chilasa (52). Possibly two subspecies. 
Common name: Burmese Raven (45, 80). Other ref: 87. 

481 Papilio (Princeps) dravidarum Wood-Mason, 1880 

Southern India. Uncommon, but not known to be threatened. Deceptively 
similar to members of the subgenus Chilasa (52). Common name: Malabar 
Raven (80, 282). Other ref: 4. 

Species-group: fuscus Goeze 



T-^ -1 J__ 1 OiC^ 



■»B2. PgEilio [Prin ceps 1 hipponoua C. & R. Felder. 1862 ^^ 

Philippines: Palawan. Banggi. Baeilan. Luzon. Mindoro.' Panay *' 
Negros. Cebu. Bohol. Leyte. Samar. Mindanao (Hancock 19B5). 

Indonesia: Sangihe and Talaut To =„^ 

^ "° laiaut Is.. and possibly northern 

Sulawesi as vagrants. Two subspecies. 



Swallowtails of the world 

species is urgently needed. Up to six subspecies. Other refs: 52, 110, 118, 
121, 262. 

%B3. Papillo [Princepa l pitwani Elwes & de Niceville. IBB6. 
South Burma: Tenasserim, Tavoy. Thailand. N. Vietnam [110). 
? Laos. China: Hainan. Possibly three subspecies. 

262). A report of its occurrence in Assam is very doubtful (259). Often 
regarded as a subspecies of hipponous (52, 262), but given as a full species 
by Hancock (109, 110). Not known to be threatened. Common name: 
Pitman's Helen (80). Other refs: 118, 121. 

484 Papilio (Princeps) fusciis Goeze, 1779 

Andamans (5), Nicobar Is, Peninsular and Eastern Malaysia, Brunei, 
Indonesia (except Sumatra (?), Java and the Lesser Sunda Is), Papua New 
Guinea, Bismarck Archipelago (except Admiralty Is), Bougainville, Solo- 
mon Is (except Santa Cruz), Vanuatu (= New Hebrides) (Torres Is), 
Trobriand Is (?), D'Entrecasteaux Is (?), Louisiade Archipelago (?) and 
Australia (Queensland and New South Wales) (262). Common generally 
and not threatened although the subspecies P. (P.) f. andamaniciis 
(Andaman Helen (80))from the Andamans and Nicobar Is is protected by 
Indian law (182). Two subspecies are farmed in Papua New Guinea (36). Up 
to twenty-two subspecies. It has been surmised that P. (P.) heringi is a rare 
natural hybrid, P. (P.) fiiscus x P. (P.) tydeiis (111, 254) but this has been 
disputed (224). Common name: Blue Helen (3). Other refs: 4, 14, 40, 45, 
48, 52, 87, 110, 131, 141, 176, 219, 220. 

485 Papilio (Princeps) canopus Westwood, 1842 

Indonesia (Lesser Sunda Is, including Tanimbar), Vanuatu (New 
Hebrides) (242) and Australia (Northern Territory and northern Queen- 
sland). P. (P.) c. hypsicles has recently been revised to a full species by 
Hancock (110) who notes its close relationship to P. (P. ) woodfordi and not 
to P. (P.) canopus, as was previously thought. Common and not known to 
be threatened. Twelve subspecies if P. (P.) c. hypsicles is included (262). 
Other refs: 40, 48, 176. 

486 Papilio (Princeps) albinus Wallace, 1865 

Papua New Guinea and Indonesia (Irian Jaya and Moluccas). Not rare, 
except in the Moluccas, and not considered to be threatened. Collected for 
trade in Papua New Guinea (36). Two subspecies. Other refs: 48, 118. 

487 Papilio (Princeps) hypsicles Hewitson, 1868 

Vanuatu (New Hebrides). Status recently revived to a full species by 
Hancock (110) who noted it to be close to P. (P.) woodfordi and not to P. 
(P.) canopus which it superfically resembles and of which it was thought to 
be a subspecies. Restricted range but not known to be threatened. Other 
ref: 262. 

488 Papilio (Princeps) woodfordi Godman and Salvin, 1888 

Solomons, (except San Cristobal and Santa Cruz) and Bougainville where it 
is farmed (36). Not rare (48) and not known to be threatened. Four 
subspecies including P. (P.) w. ptolychus Godman and Salvin, 1888 (219). 
Other ref: 220. 

101 



l^tf.yJ'^ 



Threatened Swallowtail Butterflies of the World 

489 Papilio (Princeps) heringi Niepelt, 1924 
Indonesia (endemic to Halmahera, Moluccas). A little-known species that 
was accepted as a full species and placed in the aegeus group by Munroe 
(188), but regarded as a rare natural hybrid of P. (P.) fuscus and P. (P.) 

\ tydeus by Hancock (109, 111). In a recent reassessment P. (P.) heringi is 

"r'J placed as a full species near \\\q fuscus group (224). It has rarely been 

\ collected and the rapid acceleration of deforestation on Halmahera is a 

matter of concern. More information is required on this species. 

Species-group: amynthor Boisduval 

490 Papilio (Princeps) amynthor Boisduval, 1859 
New Caledonia and Norfolk I. Apparently not uncommon, feeding on 
cultivated Citrus (133) and not threatened. Formerly known as P. (P.) 
ilioneus Donovan. Other refs: 48, 95, 111. 

,^^2^ u^cL.s-''^ 491 Papilio (Princeps) schmeltzii HemchSchaiier, 1^69 

Fiji. Despite its restricted range, the species is apparently common (48) and 
not threatened. Other ref: 111. 

492 Papilio (Princeps) godeffroyi Semper, 1866 
Western Samoa. Could possibly be classed as Rare, but more information 
on status required. Refs: 111, 144. 

Species-group: gambrisius Cramer 

493 Papilio (Princeps) inopinatus Butler, 1883 
Indonesia (Lesser Sunda Is: Romang, Babar, Damar and Tanimbar). Not 
rare and not threatened. Specific status slightly doubtful (48) Two subspe- 
cies. Other refs: 111,262. 

494 Papilio (Princeps) bridgei Mathew, 1886 
Throughout the Solomons (except for Santa Cruz) and Bougainville (219) 
where it is collected and farmed for trade (36). Common (48) and not 
threatened. Seven subspecies including P. (P.) b. erskinei Mathew, 1886 
(219) and the recently described P. (P.) b. michae from Malaita (220). 
Other refs: 111, 144. 

495 Papilio (Princeps) wejwerz Niepelt , 1914 
Rare — refer to section 6, p. 335. 

Bismarck Archipelago (Admiralty Is). A rare species according to 
D'Abrera (48) but its rain forest habitat is still fairly extensive (217). The 
rutaceous foodplant is common on Manus I. (217). Sometimes available in 
tradeasexpupae(36).P. (F.) carrerefz Oberthiir, 1914 is a synonym, but the 
order of priority of the two names has been difficult to establish, both being 
published in the first six months of 1914. Ebner (68), assisted by R.I. 
Vane-Wright, decided on cartereti as the senior name, but weymeri has 
been adopted by Hancock (109). Other ref: 111. 

496 Papilio (Princeps) gambrisius Cramer, 1777 
Indonesia (Seram, Ambon and Buru). A good species according to 
D'Abrera (48), but specific status slightly in doubt (144). More information 
needed before conservation status can be properly assessed. Two subspe- 
cies. Other ref: 111. 

102 




v< 




Swallowtails of the world 

497 Papi,lio (Princeps) tydeus C. & R. Felder, 1860 

Indonesia (Moluccas: Bacan, Ternate, Halmahera, Morotai and Obi). Said 
to be common (48) and presumably not threatened. It has been surmised 
that P. (P.) heringi Niepelt is a rare natural hybrid P. (P.) tydeus x P. (P.) 
fuscus (111, 254), but this has been disputed (224). Two subspecies. 

498 Papilio (Princeps) aegeus Donovan, 1805 

Eastern Australia (including Lord Howe I. (95)), Indonesia (Irian Jaya), 
Papua New Guinea, Bismarck Archipelago, D'Entrecasteaux Is, Louisiade 
Archipelago (?) and Trobriand Is (?). Common and not threatened. 
Farmed and some collected for trade in Papua New Guinea (36). Seven 
(111) or eleven subspecies. P. (P.) oberon Grose-Smith, 1897 from the 
Solomons (Santa Cruz) is now thought to be a local form of P. (P.) aegeus 
rather than a good species (111, 241). Common name: Orchard Butterfly 
(282). Other refs: 14, 40, 48, 141, 176, 188, 219. 

Species-group: cynorta Fabricius 

499 Papilio (Princeps) cynorta Fabricius, 1793 

Lowland forest in Sierra Leone, Guinea (?), Liberia, Ivory Coast, Ghana, 
Togo, Benin, Nigeria, Cameroon, Central African Republic (?), Gabon 
(?), Congo, Zaire, western Kenya, Uganda, Angola and an isolated 
population in Ethiopia. Generally common and not threatened. Two 
subspecies. Common name: Common White-banded Papilio (34). Other 
refs: 20, 33, 50, 285. 

500 Papilio (Princeps) plagiatiis Aurivillius, 1898 

Forests in Nigeria, Cameroon, Central African Republic, southern Sudan, 
Congo and northern Zaire. Rather uncommon but not thought to be 
threatened. Refs: 20, 34, 50, 285. 

501 Papilio (Princeps) zoroastres Druce,1878 

Cameroon, Equatorial Guinea (Bioko, formerly known as Fernando Poo 
I.), Gabon (?), Congo, Zaire, south-eastern Sudan, Uganda, western 
Kenya, Rwanda, Burundi (?), north-western Tanzania, Zambia and 
Angola. Generally common and not threatened. Five subspecies. Refs: 20, 
33, 34, 50, 214, 285. 

502 Papilio (Princeps) echerioides Trimen, 1868 

Forests in East Africa: southern Ethiopia, Kenya, Tanzania, Malawi, 
Zambia, Mozambique , Zimbabwe and South Africa (Natal and Transvaal) . 
Not uncommon and not threatened. Six subspecies. Common names: 
White-banded Swallowtail (44, 285), Southern White-banded Papilio (34). 
Other refs: 33, 50, 63, 89, 97, 165, 214. 

503 Papilio (Princeps) jacksoni Sharps, 1891 

Kenya, Uganda, Zaire, eastern Congo (?), Burundi (?), Rwanda, western 
Tanzania, Zambia and Malawi . Common in some places and not threatened 
as a species, but one subspecies is rare in Tanzania (151). Five subspecies. 
Common name: Jackson's Swallowtail (285). Other refs: 33, 34, 46, 50, 97. 

504 Papilio (Princeps) fuelleborni Karsch, 1900 

Tanzania and Malawi. Restricted range, but not known to be threatened. 
Common name: Fuelleborn's Swallowtail (285). Other refs: 33, 34, 50. 

103 



Jt 



Threatened Swallowtail Butterflies of the World 

505 Papilio (Princeps) sjoestedti Aurivillius,1908 
p Rare — refer to section 6, p. 337. 

Known from montane forest in only three localities, all in northern 

V^ t Tanzania: Mt Meru, Ngorongoro Crater and, as a different subspecies, Mt 

NJ^^r\ r Kilimanjaro, where it is abundant. Mt Meru and Mt Kilimanjaro are both 

j rvT well protected national parks indicating that this species, although very 

V narrowly distributed, is not threatened at the moment. Common name: 

Kilimanjaro Swallowtail (285). Other refs: 33, 34, 43, 50, 151. 

Species-group: rex Oberthiir 

\\ 506 Papilio (Princeps) rex Oberthiir, 1886 

\ _/^ Highland forest in Nigeria, Cameroon, Sudan, Ethiopia, eastern Congo, 

.j\i Zaire, Uganda, western Kenya, Rwanda, Burundi and Tanzania. Not 

' i uncommon (though difficult to capture) and not threatened. It has been 

suggested that this species mimics the danaid Danaus formosa Godman 

(50). Seven subspecies. Common names: Regal Swallowtail (285), King 

Papilio (34). Other refs: 33, 105. 

Species-group: nireus Linnaeus ^ \.^.AjoP- ^ 

-507 Papilio (Princeps) epiphorbas Boisduval, 1833 /^ '^'^^^ f^\K~i 

C^v^A^r*^^ -Sv.'^^^ Madagascar. Well distributed over the whole island (and Comorojs^ '^^«- 
, I \^^Z ^ 6^ ' according to D'Abrera (50)) and not threatened. Other refs: 34, 204, 206. 

C^j(ju- C^^"^"^ ' j08 Papilio (Princeps) manlius Fabricius, 1798 
Indeterminate — refer to section 6, p. 339. 

Mauritius. Despite the restricted range, widespread loss of vegetation and 
reports of nearing extinction (165), this species is currently quite common 
and widespread due to its ability to utilize cultivated Citrus (13). On the 
other hand there is concern that it may be unable to compete with the 
introduced and spreading P. (P.) demodocus (12). Other refs: 34, 50, 206. 

509 Papilio (Princeps) phorbanta Linnaeus, 1771 
Vulnerable — refer to section 6, p. 342. 

Endemic to Reunion. Two specimens of a dwarf form, subspecies nana 
Oberthiir, also included in D'Abrera (50), and allegedly confined to the 
Seychelles possibly represent wind-blown vagrants from Reunion and were 
not established on these islands (118, 167, 206). An alternative theory 
suggests that the taxon was artificially introduced to the Seychelles but did 
not become established (116). The restricted range and rarity of P. (P.) 
phorbanta is indicative of Vulnerable status. Protected since 1979. 
Common name: Papillon La Pature. Other ref: 34. 

510 Papilio (Princeps) charopus Westwood, 1843 
Highland forests in Cameroon, Equatorial Guinea (Bioko, formerly known 
as Fernando Poo L), Zaire, Rwanda, Burundi and Uganda. The type 
locality of Ashanti, Ghana is apparently erroneous as the type almost 
certainly came from Cameroon (116). Fairly common and not threatened. 
Two subspecies (115, 116). Common names: Westwood's Swallowtail 
(285), Blue-banded Swallowtail (34). Other refs: 20, 33, 50. 

511 Papilio (Princeps) hornimani Distant, 1879 
Highland forest in East Africa: northern Tanzania and south-eastern 
Kenya. Not uncommon and not thought to be threatened although a 

104 



Swallowtails of the world 

subspecies is reported to be rare in Tanzania (151). Common name: 
Horniman's Swallowtail (285). Other refs: 33, 34, 50. 

512 Papilio (Princeps) mackinnoni Sharpe, 1891 
Highland forest in Sudan, Kenya, Uganda, Zaire, Rwanda, Burundi, 
Zambia, Malawi, Tanzania and Angola. Common in places, (e.g. Kenya) 
and not threatened as a species, but one subspecies is rare in Tanzania. Four 
named subspecies (116). Common name: MacKinnon's Swallowtail (34, 
285). Other refs: 33, 50. 

513 Papilio (Princeps) sosia Rothschild and Jordan, 1903 
Senegal, Gambia, Sierra Leone, Guinea, Liberia, Ivory Coast, Ghana, 
Togo, Benin, Nigeria, Cameroon, Gabon, Congo, Central African 
Republic, Zaire, Uganda and northern Angola. Less common than P. (P.) 
nireus but fairly frequent and not threatened. Two subspecies, of which P. 
(P.) s. debilis Storace appears to be of hybrid origin (116). Common name: 
Straight-banded Swallowtail (285). Other refs: 20, 33, 34, 50. 

514 Papilio (Princeps) aethiopsis (Hancock, 1983) 
Adopted by Hancock (109) as a replacement name for aethiops Rothschild 
and Jordan, 1905. Ethiopia and north-western Somalia. A restricted range 
and the butterfly is 'local and generally uncommon' (285). It is not known to 
be threatened though more information is desirable. Common name: 
Abyssinian Blue-banded Swallowtail (285). Other refs: 34, 50, 116. 

515 Papilio (Princeps) nireiis Linnaeus, 1758 
Forests throughout southern and tropical Africa. Very common and not 
threatened. Three subspecies (four including P. (P.) n. aristophontes) 
(116). Common names: Green-banded Swallowtail (44), Narrow Blue- 
banded Swallowtail (34, 285). Other refs: 20, 33, 50, 63, 97, 156, 165, 214, 
215. 

516 Papilio (Princeps) aristophontes Oberthiir, 1897 
Indeterminate — refer to section 6, p. 345. 

r j^ fipN^ Comoro Is. Taxonomic status as a good species recently revived (50), but 
still open to doubt. Regarded as a subspecies of P. (P.) nireus by most 
authors including Carcasson (34) and Hancock (116). Information on 
conservation status required. Other refs: 206, 279. 



\h 



517 Papilio (Princeps) oribazus Boisduval, 1836 

Madagascar. Well distributed, except in the west, and apparently quite 
common and not threatened. Refs: 50, 204, 206. 

518 Papilio (Princeps) thuraui Karsch, 1900 

East Africa: highland forest in southern Tanzania, Malawi and northern 

J Zambia (115). Considered to be scarce and very local (43, 50, 285), perhaps 

j'^ deserving conservation status and certainly requiring monitoring. It has 

\i y been reported to be rare in Tanzania (43, 151) and possibly threatened by 

rJ^ deforestation if this proceeds unchecked, although it is quite firmly 

\n established at present (42). P. (P.) t. ufipa iS' apparently Vulnerable in 

Tanzania where it is restricted to the Ufipa plateau (151). Five subspecies 

according to Hancock (116) including P. (P.) t. occidua previously placed 

under P. (P.) desmondi, and P. (P.) t. cyclopsis and P. (P.) t. ufipa 

105 



Threatened Swallowtail Butterflies of the World 

previously placed as subspecies of P. (P.) bromiiis. Common name: 
Blue-spotted Black Swallowtail (285). Other refs: 34. 97. 

519 Papilio (Princeps) desmondi Van Someren, 1939 

East Africa: Kenya, Tanzania, Malawi, and Zambia. Fairly common and 
not threatened. The taxonomy and nomenclature are confused. P. (P.) 
magdae G\{{ord, 1961 was a replacement name for F. (P.) brontes Godm^n, 
1885 and, therefore, has the same type locality, Mt Kilimanjaro (115). P. 
(P.) d. magdae is reported by Kielland to be rare in Tanzania (151). Four 
subspecies including P. (P.) d. teita. P. (P.) d. occidiia Storace, described 
from Zaire, and also occuring in northern Zambia and Malawi, does not 
belong here but appears to be a subspecies of P. (P.) thuraui (115, 116). 
Common name: Godman's Swallowtail (285). Other refs: 33,34,50,96,97. 

Papilio (Princeps) desmondi teita Van Someren, 1960 

Endangered — refer to section 6, p. 347. 

South-eastern Kenya (Taita Hills) only. A recently-described taxon (271), 

treated as a subspecies of P. (P. ) desmondi by D' Abrera (50) and Carcasson 

(34). Common name: Taita Blue-banded Swallowtail. Other refs: 39, 116, 

272. 

520 Papilio (Princeps) inter jecta Van Someren, 1960 

East Africa: Uganda and western Kenya. Recently described (271) and 
accepted as a good species by D' Abrera (50), Carcasson (34) and Hancock 
« /, 1 c (109, 116). Conservation status not known; further research required. 

T Lw Ajp ^<v ^^5^^)' ^^^ Papilio (Princeps) bromius Doubleday, 1845 

f<=\?i -o '^i {'■'^^(^ Forests in Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, 

^\^ Benin, Nigeria, Cameroon, Central African Republic, Sudan, Ethiopia (?), 

\i ^^ ^ Equatorial Guinea (?), Sao Tome, Gabon, Congo, Zaire, Uganda, Kenya, 

\ vT^ Rwanda, Burundi, Tanzania and Angola. Common and not threatened as a 

*^ species. P. (P.) b. cyclopsis from northern Malawi and Zambia, and P. (P.) 

b. ufipa from south-western Tanzania do not belong here but appear to be 

subspecies of P. (P.) thuraui (116). Four subspecies (116). Common name: 

Broad Blue-banded Swallowtail (34, 285). Other refs: 20, 33, 50. 

522 Papilio (Princeps) chrapkowskii Suffert, 1904 
Highland forest in Kenya and eastern and central Uganda. Often regarded 
as a subspecies of bromius. D' Abrera follows this course, although the 
name is missing from his index (50). 

Species-group: zalmoxis Hewitson 

523 Papilio (Princeps) zalmoxis Hewitson, 1864 
Lowland forest in Liberia, Ivory Coast, Ghana, Togo (?), Benin (?), 
Nigeria, Cameroon, Central African Republic, Equatorial Guinea (?), 

\^ Gabon, Congo and Zaire. Males commonly collected (though females very 

^P scarce) and species not known to be threatened. Often placed in the genus 

0^ Iterus. Common names: Giant Blue Swallowtail (285), Great Blue Papilio 

(34). Other refs: 20, 50, 105. 

Species-group: zenobia Fabricius 

\ 524 Papilio (Princeps) gallienus Distant, 1879 

Lowland forest in Nigeria, Cameroon, Gabon, Equatorial Guinea (?), 
Congo and Zaire. Uncommon (285), but not markedly so and not 

106 



N. 




n 



Swallowtails of the world 

threatened. Common name: Large White-banded Papiho (34). Other ref: 
20. 

525 Papilio (Princeps) mechowi Dewitz, 1881 

Cameroon, Central African Republic, southern Sudan, Gabon (?), Congo, 
Zaire, Uganda and Angola. Common in places and not threatened. Two 
subspecies. Common name: Mechow's Swallowtail (285). Other refs: 33, 
34, 50. 



\r^ 



v> 



V 





526 Papilio (Princeps) zenobius Godart, 1819 
Lowland forest in Sierra Leone, Guinea (?), Liberia, Ivory Coast, Ghana, 
Togo, Benin, Nigeria, Cameroon, Central African Republic, southern 
Sudan, Uganda (?), Equatorial Guinea (Bioko, formerly known as 
Fernando Poo L), Gabon (?), Congo, Zaire and Angola. Carcasson (34) 
and D'Abrera (50) recognize cypraeofila but do not cite zenobius. In view of 
the possibility of confusion between zenobius and zenobia, this seems a 
good idea. Generally common and not threatened. P. (P.) cypraeofila 
Butler is a synonym. Three subspecies. Other refs: 20, 285. 

527 Papilio (Princeps) mechowianus Dewitz, 1885 

Angola, Zaire, Central African Republic and the Congo according to 
Carcasson (33, 34) but Williams (285) and Lewis (169) state that it also 
occurs in West Africa from Liberia (i.e. also Ivory Coast, Ghana, Togo, 
Benin, Nigeria and Cameroon). Fairly common and not threatened. Other 
ref: 50. 

528 Papilio (Princeps) androniciis Ward, 1871 

Cameroon and possibly adjacent areas. Uncommon but not known to be 
threatened. Refs: 50, 133. 

529 Papilio (Princeps) zenobia Fabricius, 1775 

Sierra Leone, Guinea (?), Liberia, Ivory Coast, Ghana, Togo, Nigeria, 
Cameroon, Central African Republic, Congo, Zaire and western Uganda. 
Generally common and not known to be threatened. Common name: 
Zenobia PapiHo (34). Other refs: 20, 33, 50, 285. 

530 Papilio (Princeps) maesseni Berger, 1974 

Ghana (recorded only from Likpe in the northern Volta Region (222)) and 
Togo (115). Not known to D' Abrera (50) or Carcasson (34) , nor Hsted in the 
Zoological Record but listed without comment by Hancock (109) who 
believes it to be a valid species (115). Further information is needed on the 
distribution and conservation of this obscure butterfly. 

Species-group: antimachiis Drury 

531 Papilio (Princeps) antimachus Drury, 1782 
Rare — refer to section 6, p. 351. 

Lowland forest in southern Guinea , Sierra Leone (200) , Liberia (98) , Ivory 
Coast, Ghana, Togo (?), Benin (?), Nigeria (159), Cameroon, Central 
African Republic, Equatorial Guinea (including Bioko, fo'rmerly known as 
Fernando Poo I.), Gabon, Congo, Zaire, Uganda, Rwanda and Angola 
(58). Males not uncommonly collected, females very scarce. Often placed in 
the genus Drury ia. Common names: African Giant Swallowtail (285), 
Giant Papilio (34). Other refs: 20, 33, 50, 66, 67, 81, 89, 105, 170, 257. 

107 



Threatened Swallowtail Butterflies of the World 

Species-group: menestheus Drury 

532 Papilio (Princeps) ophidicephalus Oberthiir, 1878 

Eastern Kenya, Tanzania, south-eastern Zaire, Malawi, Zambia, Mozam- 
bique, Zimbabwe and South Africa (Natal, Transvaal and Cape Province). 
Not uncommon and not threatened. Ten subspecies, several of them with 
restricted ranges (274). Common names: Emperor Swallowtail (44, 249, 
285), Snake's Head Swallowtail (249). Other refs: 33, 34, 50, 63, 97, 165, 
214,215. 

533 Papilio (Princeps) menestheus Drury, 1773 

Senegal, Guinea (?), Sierra Leone, Liberia (?), Ivory Coast, Ghana, Togo, 
Benin, Nigeria and Cameroon. Common and not threatened. Common 
name: Drury's Emperor Swallowtail (285). Other refs: 20, 33, 34, 50. 

534 Papilio (Princeps) lormieri Distant, 1874 

'irv^.e-ivv*, tc^sjie^ Forests up to 2000 m in Nigeria, Cameroon, Central African Republic, 

( P /v^J^'<1*-T^t-«of south-western Sudan, Equatorial Guinea (?), Gabon (?), Congo, Zaire, 

f ' Uganda, western Kenya and Angola. Not uncommon and not threatened. 

<144£> e^ MMAaetuiior Replaced by P. (P.) menestheus west of the Niger and by P. (P.) 

ite.^ ^>^»^^^p ."^W-, '^'^^' ' ophidicephalus from the Kenya coast to South Africa. Sexes similar, three 

\{ Cj- te(f. subspecies. Common names: Western Emperor Swallowtail (285), 

l*\«il f r ■ Emperor Swallowtail (34). Other refs: 33, 50. 

Species-group: demoleus Linnaeus 

535 Papilio (Princeps) morondavana Grose-Smith, 1891 
Vulnerable — refer to section 6, p. 354. 

Madagascar only. The rarest of the Malagasy endemics, though stated to be 
less rare than supposed by Paulian and Viette (206). Threatened by loss of 
habitat (280) and possibly its popularity with commercial collectors (205). 
Common name: Madagascan Emperor Swallowtail. Other refs: 34, 50, 81. 

536 Papilio (Princeps) grosesmithi Rothschild, 1926 
Rare — refer to section 6, p. 356. 

Madagascar only, particularly the west. Probably less rare than P. (P.) 
morondavana, but commercial collecting and habitat destruction need to be 
monitored. Refs: 34, 50, 206. 

537 Papilio (Princeps) erithonioides Grose-Smith, 1891 

Madagascar only, mainly in the west of the island. Not uncommon and 
apparently not threatened. Refs: 34, 50. 

538 Papilio (Princeps) demodocus Esper, 1798 

(jo^^K'b^oS T'' Tropical and southern Africa, Saudi Arabia, Yemen, South Yemen, Oman; 
. , . introduced to Madagascar, Mauritius and Reunion. Abundant in open 

- S<-t v.fr*A \^>-»' habitats and not threatened. A minor pest on Citrus. Aggressive towards 
other butterflies flying in its vicinity (50). Sexes ahke, two subspecies. 
Common names: Citrus Swallowtail (34, 44, 249, 285), Christmas Butterfly 
(44, 249), Orange Dog (34), African Lime Butterfly (161). Other refs: 20, 
33, 63, 97, 204, 214, 215. 

539 Papilio (Princeps) demoleus Linnaeus, 1758 

Oman, United Arab Emirates, Saudi Arabia, Kuwait, Bahrain, Qatar, 
western and possibly also eastern Afghanistan, Sri Lanka, India, Nepal, 

108 



^ 






^0 




,vS^ 



:v 




^ ' Swallowtails of the world 

Vietnam, Laos, Andamans (5), Burma, Thailand, Kampuchea, southern 
China (including Hainan (Guangdong prov.)), Taiwan, Japan (rare strays). 
Peninsular Malaysia, Singapore, Philippines, Indonesia (Sumatra, Sula, 
Talaud, Flores, AlorandSumba), Papua New Guinea, Australia (including 
Lord Howe L (95)), apparently Hawaii and possibly other Pacific Ocean 
islands. Another aggressive and very common species. Six subspecies. 
Common names: Lime or Lemon Butterfly (45, 80, 251), Chequered 
Swallowtail (282). Other refs: 4, 33, 40, 48, 52, 87, 1 18, 141, 148, 149, 165, 
176, 246, 262. 

Species-group: leucotaenia Rothschild 

540 Papilio (Princeps) leucotaenia Rothschild, 1908 
Vulnerable — refer to section 6, p. 358. 

Montane forest in East Africa: south-western Uganda, Rwanda, Burundi 

^ and north-eastern Zaire. A rare species threatened by deforestation. 

Common name: Cream-banded Swallowtail (285). Other refs: 33, 34, 50. 

Species-group: delalandei Godart 

Papilio (Princeps) delalandei Godart, 1824 

Madagascar. Well distributed in the forests of Madagascar, especially in the 

east; not known to be threatened. Refs: 34, 50, 206. 

542 Papilio (Princeps) mangoura Hewitson, 1875 
Rare — refer to section 6, p. 362. 

Madagascar. Distributed in the eastern rain forests and usually regarded as 
rare (263). However, Paulian & Viette (206) treat it as less rare than is 
usually thought. At present it may not be threatened, but with deforestation 
proceeding very quickly, its status needs to be carefully monitored. Local 
catchers decoy male P. (P.) wflrtgoMra with female P. (P.) delalandei, which 
has similar yellow-barred wings (263). Other refs: 34, 50, 195. 

Species-group: phorcas Cramer 

543 Papilio (Princeps) constantiniis Ward, 1871 
Woodland and forested rivers in Ethiopia, Kenya, Somalia, Tanzania, 
Congo, south-east Zaire, Malawi, Mozambique, Zambia, Zimbabwe and 
South Africa (Natal and northern Transvaal). Not uncommon and not 
threatened. Three subspecies. Common name: Constantine's Swallowtail 
(34, 44, 285). Other refs: 33, 50, 63, 97, 214. 

544 Papilio (Princeps) phorcas Cramer, 1775 

Forests throughout tropical Africa excluding Ethiopia. Very common in 

\\ places and not threatened. Six subspecies. Common names: Green-patch 

Swallowtail (285), Green Swallowtail (34). Other refs: 20, 33, 50, 97, 215. 

545 Papilio (Princeps) dardanus Brown, 1776 

Wooded and forested areas throughout tropical and southern Africa. This 
species is a famous mimetic and polymorphic swallowtail . Not threatened as 
a species but three subspecies are Rare or Vulnerable in Tanzania (151). 
There are twelve subspecies including P. (P.) d. meriones from Madagascar 
and P. (P.) d. humbloti restricted to the Comoros (263). P. (P.) d. 
flavicornis is restricted to Mt Kulal and P. (P.) d. ochracea to Mt Marsabit 
and Mt Nyiru; all in northern Kenya (50). P. (P.) nandina Rothschild from 



vV 



109 



J 



Threatened Swallowtail Butterflies of the World 

Kenya was recently demonstrated to be a rare natural hybrid between P. 
(P.) phorcas and P. (P.) dardanus (37). Common name: Mocker Swallow- 
tail (34, 44, 249, 285). Other refs: 20, 33, 63, 97, 165, 206, 214, 215. 

Species-group: hesperus Westwood 



V 



. 546 Papilio (Princeps) euphranor Trimen, 1868 

\ V^ Restricted to highland forest in South Africa (Cape Province, Natal and 

\ . iv" \ , Transvaal). Uncommon (34); but no threats known. Has been proposed for 

^ \\ inclusion in a Red Data Book for South Africa (192), ahhough no such 

work has been started. Common names: Forest Kite, Bush Kite (285), 

Natal Swallowtail (34). Other refs: 33, 50, 63, 214. 

547 Papilio (Princeps) pelodurus Butler, 1895 
Highland forest (800-1900m) in East Africa: Malawi, Zambia (115) and 
eastern Tanzania. Restricted range, limited to the distribution of its 
foodplant, Cryptocarya liebertiana (Lauraceae), in evergreen forests. It is 
apparently not uncommon, although it will be under threat in Tanzania if 
deforestation proceeds unchecked (42, 43). Two subspecies. Common 
name: Eastern Black and Yellow Swallowtail (285). Other refs: 33, 34, 50. 

548 Papilio (Princeps) hesperus Westwood, 1843 
Lowland forest in Ivory Coast, Ghana, Togo, Benin, Nigeria, Cameroon, 
Central African Republic, southern Sudan, Equatorial Guinea (Bioko, 

\ VT^ ""N^ formerly known as Fernando Poo I.), Congo, Zaire, Uganda, Rwanda (?), 

^^ J^ Burundi (?), western Tanzania and north-western Zambia. Replaced by P. 

\ J (P.) pelodorus in Malawi and Tanzania and by P. (P.) horribilis from the 

x!'^ Ivory Coast to Sierra Leone (34). Not uncommon, though difficult to 

capture and not threatened. Three subspecies. Common names: Black and 
Yellow Swallowtail (285), Hesperus Swallowtail (34). Other refs: 33, 50. 

549 Papilio (Princeps) horribilis Butler, 1874 
West Africa; Sierra Leone, Liberia, Ivory Coast and Ghana. Given as a full 
species by D'Abrera (50) and Carcasson (34), but not mentioned by other 
authors (188, 250, 285); Gifford (97) treats it as synonymous with P. (P.) 
pelodurus. Not uncommon and probably not threatened. Other ref: 41. 

550 Papilio (Princeps) nobilis Rogenhoffer, 1891 
. jv Highland forest in southern Sudan, Kenya, Uganda, Burundi, Rwanda, 
^ r^^'- Tanzania and Kivu Province, Zaire (34, 285). Not uncommon and not 

^ r^ recognized as being threatened. Three subspecies. Common name: Noble 

>/ Swallowtail (34, 285). Other refs: 33, 50. 

Species-group: palinurus Fabricius 

\^ 551 Papilio (Princeps) crino Fabricius, 1792 

Southern and western coastal India and Sri Lanka (52). Not threatened. 
Common name: Common Banded Peacock (80). Other refs: 4, 259, 287. 

552 Papilio (Princeps) blumei Boisduval, 1836 
Indonesia (Sulawesi). Much in demand for the decorative trade but prices 
have fallen recently and it is not known to be threatened. Monitoring of 
status is required. Two, possibly four subspecies (262). Other refs: 52, 141. 

553 Papilio (Princeps) buddha Westwood, 1872 
Southern India. Locally common and not rare (118). Protected by law 

110 



c 



^ 



!>' 



Swallowtails of the world 

(182), but not known to be threatened. Common name: Malabar Banded 
Peacock (80). Other refs: 4, 52, 259. 

554 Papilio (Princeps) palinurus Fabricius, 1787 

Burma, western Thailand, Peninsular and Eastern Malaysia, Brunei and 
Indonesia (Sumatra, Nias and Kalimantan) (52, 262). Generally common, 
not threatened except in Malaysia where possibly Vulnerable and in need 
of protection (10). Two subspecies (262). The five subspecies listed by 
D'Abrera (52) include P. (P.) daedalus subspecies (see below) and a 
synonym. Common names: Banded Peacock (3, 45), Burmese Banded 
Peacock (80), Moss Peacock (3). Other refs: 87, 131, 141, 168, 259. 

555 Papilio (Princeps) daedalus C. & R. Felder, 1861 

Philippines, including Palawan but excluding Sulu Archipelago (52, 262). 
Formerly regarded as the eastern subspecies of P. (P.) palinurus (52), 
which is distributed further to the west, but raised to full species rank by 
Hiura and Alagar (127). Abundant and not threatened. Two subspecies. 

Species-group: paris Linnaeus 

556 Papilio (Princeps) chikae Igarashi, 1965 
Endangered — refer to section 6, p. 364. 

Philippines: Luzon; only known from the Baguio and Bontoc areas in the 
north of the island (262). The recent discovery of this species has created a 
high demand, particularly from Japanese collectors. This, along with its 
limited distribution, danger of habitat destruction and apparent ease of 
capture all indicate Endangered status. Other refs: 2, 52, 53, 140, 148. 

557 Papilio (Princeps) maackii Menetries, 1859 

Japan, southern China, North Korea, South Korea, Taiwan and extreme 
south-eastern U.S.S.R. (Sikhote-Alin, etc. and Kuril Is); a similar, but 
slightly more northerly, distribution to P. (P.) Manor. Two subspecies. 
Generally common but rapidly declining in the U.S.S.R. and listed in the 
U.S.S.R. Red Data Book (8, 260). Other refs: 52, 141, 149, 169. 

558 Papilio (Princeps) Manor Cramer, 1776 

North-eastern Burma, Sichuan (Szechwan) and southern China, Japan, 
South Korea, North Korea, northern Laos and Vietnam. Generally 
common and not threatened. Seven subspecies; okinawaensis is now rare 
on Okinawa (2). Common name: Chinese Peacock (80). Other refs: 141, 
143, 149, 246, 259. 

559 Papilio (Princeps) syfanius Oberthiir, 1886 

Western China (Yunnan and Sichuan). Usually considered to be a 
subspecies of bianor but treated as a full species by Hancock (109). Not 
known to be threatened. 

560 Papilio (Princeps) polyctorBoisduva] , 1836 

Eastern Afghanistan, Pakistan, northern India (including Sikkim and 
Assam), Nepal, Bhutan (?), Burma, Thailand, northern Vietnam and Laos 
(?). Generally common and not threatened. Six subspecies, including the 
recently described P. (P.) p. pinratanai (223). Common name: Common 
Peacock (80, 251). Other refs: 4, 52, 136, 141, 168, 259. 

561 Papilio (Princeps) dialis (Leech,, 1893) 

Southern China (including Hainan (Guangdong prov.)), north-eastern 

111 



y^ 



Threatened Swallowtail Butterflies of the World 

Burma, northern Vietnam, Laos and Taiwan. Generally common and not 
threatened. Five subspecies. Common name: Black-crested Spangle (94). 
Other refs: 52, 141,246,259. 

562 Papilio (Princeps) elephenor Doubleday, 1845 

North-eastern India (Jorehat, Cachir and Sadiya in Assam, Khasia in 
Meghalaya, Nagaland and Manipur), Burma and northern Thailand. 
Although status given variously as rare or very rare, it seems marginal for 
threatened status. It is protected by law in India (182). More information 
from the eastern part of its range is desirable. Two subspecies. Common 
names: Yellow-crested Spangle (80, 94), Black-crested Spangle (P. (P.) e. 
schanus) (80). Other refs: 4, 52, 168, 259. 

563 Papilio (Princeps) paris Linnaeus, 1758 

Southern and north-western India, Bangladesh, Bhutan (?), Burma, 
northern Thailand, Laos, Vietnam, Sichuan (Szechwan) and southern 
China (including Hainan (Guangdong prov.)). Peninsular Malaysia (76), 
Taiwan and Indonesia (Sumatra and Java) (52, 262). Common in many 
areas and not threatened. Ten subspecies. Common name: Paris Peacock 
(P. (P.) p. paris) (80, 282), Tamil Peacock (P. (P.) p. tamilana) (80). Other 
refs: 4, 45, 141, 143, 168, 207, 246, 259. 

564 Papilio (Princeps) karna C. & R. Felder, 1864 

Philippines (Palawan), Eastern Malaysia, Brunei and Indonesia (Kaliman- 
tan, Java and Sumatra). Generally uncommon but not known to be 
threatened. On Palawan females remain near the foodplant, said to be a 
species of large forest tree; males congregate at drinking spots; numbers 
stable (56). More information required from other areas. Possibly three or 
more subspecies. Other refs: 45, 52, 131, 148, 262. 

565 Papilio (Princeps) arcturiis Westwood, 1842 

North-western India, Nepal, Bhutan, Bangladesh (?), Burma, western and 
central China and Thailand. Not uncommon and not threatened. Two 
subspecies. Common name: Blue Peacock (80, 282). Other refs: 4, 52, 141, 

207, 259. 

566 Papilio (Princeps) hoppo Matsumara, 1908 

Taiwan. Not uncommon and not thought to be threatened, despite its 
restricted range. Requires monitoring because of the butterfly trade in 
Taiwan. Refs: 52, 141,246. 

567 Papilio (Princeps) krishna Moore, 1857 

North-eastern India (including Manipur, Sikkim and Assam), Nepal and 
western China (52). A narrow range. Apparently uncommon and com- 
manding high prices in the trade, but not thought to be threatened. Two 
subspecies. Common name: Krishna Peacock (80, 251). Other refs: 141, 
259. 

Species-group: peranthus Fabricius 

568 Papilio (Princeps) neumoegeni Honrath, 1890 
Vulnerable — refer to section 6, p. 367. 

Known only from Sumba in the Lesser Sunda Is in Indonesia, whence some 
numbers have recently been distributed by the trade; very restricted range, 
current status uncertain. Refs: 52, 134, 262. 

112 



f Swallowtails of the world 

569 Papilio (Princeps) peranthus Fabricius, 1787 

Indonesia (Java, Bali, Lombok, Bawean, Kangean, Sumbawa, Flores, 
Adonara, Solor, Pantar, Alor, Tanahjampea, Kalao, Bonerate, Salayar 
and Sulawesi) (52, 262). The Sulawesi form is probably specifically distinct 
(118) but further assessment is needed to confirm this. Not uncommon but 
not thought to be threatened. Six to eight subspecies. Other ref: 141. 

570 Papilio (Princeps) pericles Wallace, 1865 

Indonesia (Lesser Sunda Is (Timor, Wetar, Moa, Leti, Babar, Damar, 
Romang and Tanimbar)) (262). Not rare (48) or threatened, but not 
common. 

571 Papilio (Princeps) lorquinianus C. & R. Felder, 1865 

Indonesia (Moluccas (Morotai, Ternate, Halmahera, Bacan and Seram) 
and western Irian Jaya). Scarce throughout its range but not known to be 
threatened. Five subspecies. Refs: 48, 141. 

Species-group: ulysses Linnaeus 

572 Papilio (Princeps) ulysses Linnaeus, 1758 

Indonesia (Moluccas and Irian Jaya), Papua New Guinea, Bismarck 
Archipelago (including New Britain), Trobriand Is, D'Entrecasteaux Is 
(?), Louisiade Archipelago (?), north-eastern Australia and the Solomons 
(except Malaita, San Cristobal and Santa Cruz). Much prized for its beauty, 
but not threatened as a species. Collected for trade in Papua New Guinea 
(36, 68). Protected by law in Queensland, Australia (258). Sixteen 
subspecies. Common names: Ulysses Butterfly (282), Blue Mountain 
Butterfly (282), Mountain Emperor. Other refs: 14, 40, 48, 141, 142, 176, 
217,219. 

573 Papilio (Princeps) montrouzieri^ohdxivaX. 1859 

New Caledonia (including the Loyalty Is) . Despite this restricted range , it is 
not rare (48) but neither is it ubiquitous (133). Its status should be closely 
monitored. Other ref: 219. 

The following species were listed by Munroe (188) but not by Hancock (109): 

1. Parnassius pythiaKoXhschiXd, 1932 

2. Parnassius rothschildianus Bryk, 1932 

3. Parnassius stenosemus WonraXh , 1890 

4. Eurytides chibcha(¥2LSs\,\9l2) 

5. Eurytides hipparchus (Staudinger, 1884) 

6. Graphium sisenna (Mabille, 1890) 

7. Papilio peleides Esper , 1793 

The three parnassians were listed as subspecies of other Parnassius species in a 
revision by Bryk (31), the Eurytides species were regarded as "aberrations" by 
Hancock (109), Graphium sisenna from Mozambique is synonymous with Graphium 
(Graphium) polistratus, and Papilio peleides is listed in the British Museum (Natural 
History) records as possibly an artefact of South American origin. 

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113 



Threatened Swallowtail Butterflies of the World 

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Moscow-Leningrad, 365-366 (In Russian). 

228. Ribbe, C. (1907). Zwei neuen Papilioformen von der Salomon-Insel Bougainville. 
Deutsche Enlomologische Zeitschrift "Iris" 20: 59-63. 

229. Riley, N.D. (1939). A new species of Armandia (Lep. Papilionidae). The Entomologist. 
72: 206-208, 267. 

230. Riley, N.D. (1975). A Field Guide to the Butterflies of the West Indies. Collins, London. 
244 pp. 

231. Robinson, J. C. (1975-6). Swallowtail butterflies of Sabah. Sabah Society Journal 6: 5-22. 

232. Ross, G. (1964). Life history studies on Mexican butterflies I. Journal of Research on the 
Lepidoptera. 3: 9-17. 

233. Rothschild, W. (1895). A revision of the papilios of the eastern hemisphere, exclusive of 

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234. Rothschild, W. and Jordan, K. (1906). A revision of the American Papilios. Novitates 
Zoologicae 13: 411-752. (Facsimile edition ed. P.H. Arnaud, 1967). 

121 



Threatened Swallowtail Butterflies of the World 

235. Rothschild, W. and Jordan, K. (1901). On some Lepidoptera. Novitates Zoologicae 1901: 
401-407. 

236. Riitimeyer, E. (1969). A new Papilio from Colombia and a new sphingid from New 
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237. Saigusa, S., Nakanishi, A., Shima,H. and Yata, O. (1977). Phytogeny and biogeography 
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238. Saigusa, T. and Lee, C.-L. (1982). A rare papilionid butterfly Bhutanitis mansfieldi 
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239. Salk, P. (1983). Neue beobachtungen zur Biologie von Parnassius nordmanni Men., 
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240. Salvia, O. (1893). XIX. Description of a new genus and species of Papilionidae from 
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241. Samson, C. (1979). Butterflies (Lepidoptera: Rhopalocera) of the Santa Cruz group of 
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242. Samson, C. (1983). Butterflies (Lepidopera: Rhopalocera) of Vanuatu. Naika (Journal of 
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243. Schwartz, A. (1984). In litl., 16 May. 

244. Shapiro, A.M. (1975). Papilio "gothica" and the phenotypic plasticity of P. zelicaon 
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245. Sheljuzhko, L. (1961). Parnassius nordmanni Men in Kleinasien. Entomologische 
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246. Shirozu, T. (1960). Butterflies of Formosa in Colour. Hoikusha. Osaka, Japan (in 
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247. Shirozu, T. andHara, A. (1960). Early Stages of Japanese Butterflies in Colour. Hoikusha, 
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248. Shirozu, T. and Hara, A. (1962). Early Stages of Japanese Butterflies in Colour. Hoikusha, 
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249. Skaife, S.H. (1953). African Insect Life. Longmans Green and Co., Cape Town. 387 pp. 

250. Smart, P. (1975). The Illustrated Encyclopedia of the Butterfly 'World. Hamlyn, London. 
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251. Smith, C. (1975). Commoner Butterflies of Nepal. Tribhuvan University, Kathmandu, 
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252. Stahl, G. (1979). Ripponia hypolitus Cramer. A description of the form 'antiope' from 
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135-139. 

253. Stanek, V.J. (1969). The Pictorial Encyclopedia of Insects. Hamlyn, London. 544 pp. 

254. Straatman, R. (1963). A hybrid between Papilio aegeus aegeus and Papilio fuscus 
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255. Straatman, R. (1969). Notes on the biology and host plant associations of Ornithoptera 
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256. Straatman, R. (1975). Notes on the biologies of Papilio laglaizei and P. toboroi 
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257. Struhsaker, T.T. (1983). In litt., 30 April. 

258. Sullivan, Hon. V.B., Minister for Primary Industries (1974). Fauna Conservation 
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259. Talbot, G. (1939). The Fauna of British India, Including Ceylon and Burma. Butterflies 
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260. Tanasiychuk, V.N. (1981). Data for the "Red Book" of Insects of the U.S.S.R. 
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261. Thomas, C. and Cheverton, M. (1982). Cambridge butterfly expedition to Central 
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262. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands. 'Vol. I 

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Swallowtails of the world 

Papilionidae. (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 

263. Turlin, B. (1983). In lilt., 15 September. 

264. Turlin, B. (1983). In litt., 1 July. 

265. Turlin, B. (1983). Systematique des Parnassius. Manuscript in lilt.. 10 October. 

266. Turner, T.W. (1983). In lilt.. 27 July. 

267. Turner, T.W. (1983). The status of the Papilionidae, Lepidoptera of Jamaica with 
evidence to support the need for conservation of Papilio homerus Fabricius and Eurylides 
marcellinus Doubleday. Unpublished report. 14 pp. 

268. Tyler, H.A. (1983). In litt., 13 March. 

269. Tyler, H.A. (1975). The Swallowtail Butterflies of North America. Naturegraph 
Publishers, viii + 192 pp. 

270. U.S. Fish and Wildlife Service (1983). Proposed delisting of Bahama Swallowtail 
Butterfly and reclassification of Schaus Swallowtail Butterfly from Threatened to 
Endangered. Federal Register 48(\68): 39096-39099. 

271. Van Someren, V.G.L. (1960). Systematic notes on the associated blue-banded black 
Papilios of the bromius-bronles-sosia complex of Kenya and Uganda, with descriptions of 
two new species. Boletim do Sociedade de Esludos da Provincia de Mocambique 123: 
65-78. 

272. Van Someren, V.G.L. (1974). Listof foodplants of some East African Rhopalocera, with 
some notes on the early stages of some Lycaenidae. Journal of the Lepidopterists' Society 
28:315-331. 

273. Van Son G. (1949). The Butterflies of Southern Africa. Part I. Papilionidae and Pieridae 
Transvaal Museum, Pretoria. 237 pp + 41 pis. 

274. Vane-Wright, R.I. (1984). In lilt., 3 July. 

275. Vazquez G., L. (1953). Observaciones sobre Papilios de Mexico, con descripciones de 
algunas formas nuevas; una especie nueva para Mexico y localidades nuevas de algunos 
otros. III. Anales del Instituto de Biologid Universidad Nacional Autonoma de Mexico. 24: 
170-175. 

276. Vazquez G., L. (1983). In litt., 15 March. 

277. Vazquez G., L. and Perez R., H. (1961). Observaciones sobre la biologia de Baronia 
brevicornis Salv. (Lepidoptera: Papilionidae — Baroniidae). Anales del Instituto de 
Biologid Universidad Nacional Autonoma de Mexico. 32: 295-311. 

278. Vazquez G., L. and Perez, H. (1967). Nuevas observaciones sobre le biologia de Baran/a 
brevicornis SdL\\. Lepidoptera: Papilionidae — Baroniinae. Anales del Instituto de Biologia 
Universidad Nacional Autonoma de Mexico. 37: 195-204. 

279. Viette, P. (1980). Mission lepidopterologique a la Grande Comore (Ocean Indien 
occidental). Bulletin de la Societe Entomologique de France 85: 226-235. 

280. Viette, P. (1983). In litt., 22 March. 

281. Walker, M.V. (1983). In litt. to S.M. Wells, 11 April. 

282. Watson, A. and Whalley, P.E.S. (1975). The Dictionary of Butterflies and Moths in 
Colour. McGraw-Hill Book Company. 296 pp. 

283. Wells, S.M., Pyle, R.M. and ColUns, N.M. (1983). The lUCN Invertebrate Red Data 
Book. lUCN, Cambridge and Gland. L + 632 pp. 

284. Whitten, A. (undated). In litt. 

285. Williams, J.G. (1969). A Field Guide to the Butterflies of Africa. Collins, London. 238 pp. 

286. Wood, G.L. (1982). The Guinness Book of Animal Facts and Feats. Third Edition. 
Guinness Superlatives Ltd., Enfield. 250 pp. 

287. Woodhouse, L.G.O. (1950). The Butterfly Fauna of Ceylon. Second Edition. Colombo 
Apothecaries' Co. Ltd., Colombo. 231 + xxxii pp. 

288. Wyatt, C. and Omoto, K. (1963). Auf der Jagd nach Parnassius autocrator Avin. 
Zeitschrift Wiener Entomologische Gesellschaft 48: 163-170. 

289. Yamamoto, A. (1977). A new species of Graphium from Mindanao (Lepidoptera: 
Papilionidae). Tyo to Ga. (Transactions of the Lepidopterists' Society of Japan). 28: 
87-88. 

290. Yuste, F., Perez, R., H. and Walls, F. (1972). Compounds of papilionid caterpillars 
{Baronia brevicornis Salv). Experientia 28: 1149. 

123 



Threatened Swallowtail Butterflies of the World 

Note added in proof: In a recent publication some Graphium species-groups have been revised 
(Hancock, 1985). Conclusions affecting this text may be summarized as follows: 

1 . G. schaffgolschi , formerly regarded as a second subspecies of G. taboranus (species 145), is 
raised to a full species distributed in Angola, Namibia, Zaire and Zambia. 

2. G. poggianus (species 150) is now recorded from Tanzania. 

3. G. oibrechtsi and G. odin are now regarded as conspecific with G. auriger (species 154-156). 

4. G. weberi and G. ucalegonides are now regarded as conspecific with G. fulleri (species 
158-160). 

291. Hancock, D.L. (1985). Systematic notes on some African species of Graphium Scopoli. 
Papilio International 2: 97-103 (plus correction in subsequent issue). 

Geographical index to species 

This index lists the species of swallowtails found in countries and regions of the world. 
It is as complete and accurate as possible at the time of going to press, but new data are 
always emerging and conservation status categories are constantly reviewed and al- 
tered. Any new information, amendments or comments on the list will be welcomed. 

Under each country or region a list of numbers will be found. These numbers refer 
to the 573 species recognized in this book and discussed in the main taxonomic list. In 
addition the number, name and status of any threatened species is printed in full. Full 
reviews of these threatened species will be found in section 6 of this book. To find the 
appropriate page number, first refer back to the species list. 

Many countries have been listed individually, but to save space and avoid 
unnecessary repetition some are grouped into regions. These regions usually contain 
either relatively few species (e.g. North Africa), or the constituent countries have 
similar swallowtail faunas (e.g. Europe, Middle East), or the distributional informa- 
tion for individual countries within the region is poor (e.g. West Africa, Indochina, 
Central America). Readers interested in the fauna of one particular country which is 
not individually listed should note the regional species numbers and refer to the 
distributions given in the full taxonomic list. Because of the exceptionally large size of 
the U.S.S.R. and the variation within its swallowtail fauna it has been divided into 
three separate regions. 

EUROPE 

Europe has a poor swallowtail fauna which is concentrated in the south and east of the 
continent. To save repetition it has not been divided into its constituent countries, i.e. 
Albania, Andorra, Austria, Belgium, Bulgaria, Czechoslovakia, Denmark, Finland, 
France (including Corsica), Federal Republic of Germany, German Democratic 
Republic, Gibraltar, Greece, Hungary, Iceland, Ireland, Italy (including Sardinia 
and Sicily), Liechtenstein, Luxembourg, Malta, Monaco, Netherlands, Norway, 
Poland, Portugal, Romania, Spain, Sweden, Switzerland, United Kingdom and 
Yugoslavia. The U.S.S.R. is considered separately. 

2, 28, 37, 40. 42, 46, 47, 55, 426-428 (11 species) 

40. Parnassiiis apollo RARE (Albania, Andorra, Austria. Bulgaria, 

Czechoslovakia, Finland, France, F.R.G., G.D.R., Greece, Hungary, 

Italy (including Sicily), Liechtenstein, Norway, Poland, Romania, Spain. 

Sweden. Switzerland and Yugoslavia) 

427. Papilio {Papilio) hospiton ENDANGERED (Corsica and Sardinia) 

124 



Swallowtails of the world 
NORTH AFRICA AND THE MIDDLE EAST 

This region has a relatively poor swallowtail fauna and is, therefore, treated as three 
large subregions. 

North Africa 

(Algeria, Canary Is, Egypt, Libya, Morocco and Tunisia) 

47, 55, 428 (3 species) 

Arabian Peninsula 

(Bahrain, Kuwait, Oman, Qatar, Saudi Arabia, United Arab Emirates, 

Yemen and South Yemen) 

55(?), 428, 538, 539 (3 or 4 species) 

Middle East 

(Cyprus, Israel, Jordan, Lebanon, Syria and Turkey) 

2, 27, 28, 40, 42, 43, 426, 428 (8 species) 

40. Parnassiiis apollo RARE (Syria and Turkey) 

U.S.S.R. 

The U.S.S.R. has been divided into three regions, (total for all regions 35 species). 

European U.S.S.R. 

(including Armenia, Azerbaydzhan, Byelorussia, Estonia, Georgia, Lat- 
via, Lithuania, Moldavia and Ukraine) 
2, 27, 28, 40, 42, 44, 46, 55, 428 (9 species) 
40. Parnassius apollo RARE 

Soviet Central Asia 

(Kazakhstan, Kirghizia, Tadzhikistan, Turkmenistan and Uzbekistan) 
2, 3, 7, 9, 10, 12, 14, 15, 20, 26, 28, 31, 32, 34, 35(?), 3^3 8, 55(?), 426, 428 
(19-21 species) 

15. Parnassius autocrator RARE (Tadzhikistan) 

Siberia 

(Asiatic U.S.S.R. excluding Soviet Central Asia) 

16, 24-26, 29, 33, 37, 39-41, 54, 273, 428, 557 (14 species) 
40. Parnassius apollo RARE 

MIDDLE ASIA TO INDOCHINA AND JAPAN 



Afghanistan 

3, 7, 8, 12, 13, 15, 28, 32, 3^36, 38, 55, 274, 314, 426, 428, 539, 560 (19 

species) 

15. Parnassius autocrator RARE 

Bangladesh 

121(?), 186, 187, 189, 203(?), 330, 415, 416, 420(?), 428(?), 446, 462, 470, 
563, 565(?) (10-15 species) 

125 



Threatened Swallowtail Butterflies of the World 



Bhutan 



Burma 



China 



India 



7, 10, 17(?), 19, 21, 35, 50, 51, 57, 59, 60, 135, 198, 272, 274(?), 284, 285(?), 
293, 415(?), 420(?), 421, 428, 458, 462, 470, 478(?), 479, 560(?), 563(?), 
565 (22-30 species) 

51 . Bhutanitis ludlowi INSUFFICIENTLY KNOWN 
57. Teinopalpus imperialis RARE 

50,57,59,60, 114-117, 120, 121, 128, 129, 135, 136, 138, 180, 182, 186-189, 
198, 203, 274-278, 284, 285, 291, 293-295, 298, 300, 302, 314, 320, 326, 330, 
415, 416, 420-422, 428, 440, 444, 446, 449, 458, 462, 464, 467, 470, 473, 478, 
480, 483, 539, 554, 558, 560-563, 565 (68 species) 
57. Teinopalpus imperialis RARE 

(including Hong Kong and Xizang Zizhiqu (Tibet)) 

4, 5, 7, 8, 10-12, 14, 16-22, 24, 29-31, 33-41, 48-50, 52, 54-60, 114-120, 

128, 129, 135, 136, 138, 182, 183, 186-189, 198, 203, 271-274, 275(?), 

276(?), 278, 280, 281, 283-285, 293, 295(?), 300, 314, 326, 330, 413, 415, 

416(?), 420, 421, 422(?), 428, 440, 446-449, 458, 462, 470, 473, 478, 479, 

483, 539, 557-559, 561, 563, 565, 567 (99-104 species) 

40. Parnassius apollo RARE (Xinjiang Uygur (Sinkiang)) 

48. Bhutanitis mansfieldi RARE (Yunnan) 

49. Bhutanitis thaidina RARE (Yunnan, Sichuan and Shaanxi) 

52. Luehdorfia chinensis INSUFFICIENTLY KNOWN (Anhui, Hubei, 
Jiangsu and Jiangxi) 

57. Teinopalpus imperialis RARE (Hubei, Sichuan) 

58. Teinopalpus aureus INSUFFICIENTLY KNOWN (Guangdong) 



(mainland only; for Andaman and Nicobar Islands see Indian Ocean) 
6, 7, 8, 10-12, 17, 19-21, 34-36, 38, 50, 55, 57, 59, 60, 114-116, 120, 121, 
128, 129, 135, 136, 138, 182, 186-189, 198, 203, 272, 274-276, 278, 284, 285, 
293, 294, 300, 302, 304, 314, 325, 326, 330, 415, 416, 420-422, 428, 445, 446, 
449, 450, 458, 460, 462, 470, 473, 478, 479, 481, 551, 553, 560, 562, 563, 565, 
567 (77 species) 
57. Teinopalpus imperialis RARE 

Indochina 

(Kampuchea, Laos, Thailand and Vietnam) 

50, 59, 60, 114, 115, 120, 121, 128, 129, 131, 134-138, 180-182, 186-189, 
198, 203, 273, 274, 275(?), 276, 277, 279, 285, 291, 293-295, 296(?), 298, 
300, 314, 319, 320, 326, 330, 415,416, 420-422, 444, 446, 449, 462, 464, 467, 
470, 471(?), 473, 478-480, 483, 539, 554, 558, 560-563, 565 (66-69 species) 



Iran 
Iraq 
Japan 



2, 3, 28, 42, 43(?), 45, 55(?), 426, 428 (7-9 species) 

2, 28, 42, 43, 55(?), 426, 428 (6 or 7 species) 

(including Ryukyu Is) 

24, 29, 30, 33, 53, 54, 187, 203, 273, 314, 428, 429, 440, 446-448, 462, 470, 

473, 539, 557, 558 (22 species) 



126 



Swallowtails of the world 

53. Luehdorfia japonica VULNERABLE 

North Korea 

24, 29, 30, 33, 39, 4L 54, 273, 428, 446-448, 557, 558 (14 species) 

South Korea 

29, 30, 33, 39, 41, 54, 203(?), 273, 428, 44^448, 470, 557, 558 (14 or 15 
species) 

Mongolia 

16, 20, 22, 24, 26, 29, 33, 37, 39, 40, 428 (11 species) 
40. Parnassius apollo RARE 

Nepal 

10, 17, 20, 21, 35, 57, 116, 117, 121, 128, 135, 136, 182, 187, 189, 198, 203, 
272, 274, 284, 285, 294(?), 314, 326, 415, 420. 428, 446, 450, 458, 462, 470, 
473, 478, 539, 560, 565, 567 (37 or 38 species) 
57. Teinopalpus imperialis RARE 

Pakistan 

3, 7, 8, 17, 20, 21, 34-36, 38, 55, 426, 428, 560 (14 species) 

Sri Lanka 

121, 128, 187, 189, 203, 302, 303, 314, 332, 415, 460, 470, 473, 539, 551 (15 

species) 

303. Atrophaneura (Pachliopta) jophon VULNERABLE 

Taiwan 

116,118,187,189,198,203,273,281,282,285,292,314,323,326,414,420, 
421 , 428, 440, 446, 456, 459, 462, 470, 473, 478, 479, 539, 557, 561 , 563, 566 
(32 species) 
414. Papilio (Chilasa) maraho VULNERABLE 

SOUTHEAST ASIA TO AUSTRALIA 

N.B. No swallowtails have been recorded from New Zealand. 

Australia 

113, 120, 186, 189, 190, 193, 203, 224, 315, 349-351, 412, 475, 484, 485, 498, 
539, 572 (19 species) 

Brunei 

60, 114-116, 120, 128, 130(?), 134, 136-138, 180, 181, 186-189, 197, 203, 
296(?), 298, 314, 316, 320, 321, 330, 416, 422, 444, 453, 467, 470, 473, 478, 
484, 554, 564 (35-37 species) 
453. Papilio (Princeps) acheron RARE 

Indonesia ^ ^^ Ot^^--^ 

60, 114, 115, 120, 122-124) 126, 128, 129,® 134, 136-138, lB, ul 180;* /~^\? 
j^^i^^f^r^ 181,qi5-190, 194-^,^, 205, 224,@^291, 296-300,1305-307^314-316, ^ /*V5j^ 
3 1 8^32 1 , 324 , 326 , 329-3^ , 333-34'4 , 347-349 , 4 1 5-417 , 422 , 423 , 442^44 , ^-"""^ 
451 ,452,^5-457, 462, 466, 467, 469-475 ,>477, 478, 482, 484-486, 483, 493, 
i496-498, 539,(552, 554,T63, 564,^^6§i^572 (121 species) v 
195. Graphium (Graphium) stresemanni RARE (Moluccas) _^ 



Threatened Swallowtail Butterflies of the World 

288. Atrophaneura (Atrophaneura) luchti RARE (Java) 

299. Atrophaneura (Losaria) palu INSUFFICIENTLY KNOWN 

(Sulawesi) 
324. Troides (Troides) prattorum INDETERMINATE (Buru) 
329. Troides (Troides) dohertyi VULNERABLE (Talaud Is and 

Sangihe) 
340. Ornithoptera tithonus INSUFFICIENTLY KNOWN (Irian Jaya) 
34L Ornithoptera rothschildi INDETERMINATE (Irian Jaya) 

342. Ornithoptera chimaera INDETERMINATE (Irian Jaya) 

343. Ornithoptera paradisea INDETERMINATE (Irian Jaya) 

344. Ornithoptera meridionalis VULNERABLE (Irian Jaya) 

347. Ornithoptera aesacus INDETERMINATE (Moluccas) 

348. Ornithoptera croesus VULNERABLE (Moluccas) 

472. Papilio (Princeps) jordani RARE (Sulawesi) 

568. Papilio (Princeps) neumoegeni VULNERABLE (Lesser Sunda Is) 

Eastern Malaysia "^r^ 

(Sabah and Sarawak) ' ^-^ 

60, i;4, 115, 120, 126-130, 134, 136-138, 180, 181, 184', 18^189, 197, 203, 

296, 298, 314, 316, 320-322, 330, 416, 422, 444, 453, 461(?), 4^2, 467, 470, 

473, 47$, 484, 554, 564 (42-43 species) ~^ ,-i.^ 6- l^xJ^ ^'-- -^n^^? 
184. Graphium (Graphium) procles INDETERMINATE (Sabah) 

322. Troides (Troides) andromache INDETERMINATE 
453. Papilio (Princeps) acheron RARE 

Peninsular Malaysia 

(including Singapore) / i 

60, 114, 115, 120, 128, 129, 134, 136-138, 180-182, 186-189, 197, 203, 291, 
294, 296, 298, 300, 314, 316, 319, 320, 326, 330, 415, 416, 420, 421(?), 422, 
444, 462, 467, 470, 471, 473, 478, 480, 4^4, 539, 554, 563 (46-47 species) 
(Malaysia as a whole has 54-56 species) 

Papua New Guinea 

(including Bismarck Archipelago, Bougainville and associated islands) 

120, 142, 186, 189-191, 193, 194, 196, 203-205, 207, 224, 315, 331, 339, 

342-346, 349, 423-^25, 442, 475, 476, 484, 486, 488, 494, 495, 498, 539, 572 

(37 species) 

191. Graphium (Graphium) meeki RARE (Bougainville) 

204. Graphium (Graphium) mendana RARE (Bougainville) 

342. Ornithoptera chimaera INDETERMINATE 

343. Ornithoptera paradisea INDETERMINATE 

344. Ornithoptera meridionalis VULNERABLE 

345. Ornithoptera alexandrae ENDANGERED 

424. Papilio (Chilasa) toboroi RARE (Bougainville) 

425. Papilio (Chilasa) moerneri VULNERABLE (Bismarck Archipel- 
ago) 

495. Papilio (Princeps) weymeri RARE (Bismarck Archipel- 
ago) 

Philippines 

114, 115, 120, 126, 127, 133, 134, 136, 139, 140, 180, 181, 186, 187, 189, 196, 
197, 202, 203, 286, 298, 308-314, 317, 323, 327, 328, 415,416, 418, 419, 440, 
441, 444, 456, 461, 465, 468, 470, 474, 483, 539, 555, 556, 564 (49 species) 

128 



v^ 



Swallowtails of the world 

133. Graphium (Pathysa) idaeoides RARE 

139. Graphium (Pathysa) megaera INDETERMINATE (Palawan) 

202. Graphium (Graphium) sandawamtm VULNERABLE (Mindanao) 

308. Atrophaneura (Pachliopta) schadenbergi VULNERABLE (Luzon, 

Babuyan) 

312. Atrophaneura (Pachliopta) atropos INDETERMINATE (Palawan) 

418. Papilio (Chilasa) osmana VULNERABLE (Leyte and Mindanao) 

419. Papilio (Chilasa) carolinensis VULNERABLE (Mindanao) 
441. Papilio (Princeps) benguetamis VULNERABLE (Luzon) 
556. Papilio (Princeps) chikae ENDANGERED (Luzon) 

Solomon Islands 

186, 191, 196, 203, 204, 206, 315, 346, 349, 424, 476, 484, 488, 494, 572 (15 

species) 

191. Graphium (Graphium) meeki RARE 

204. Graphium (Graphium) mendana RARE 

424. Papilio (Chilasa) toboroi RARE 

TROPICAL AND SOUTHERN AFRICA 



This region has been subdivided into individual countries except for western and 
southern Africa. Madagascar and other islands are listed under Oceanic Islands. 

Southern Africa 

(Botswana, Lesotho, Namibia, South Africa, Swaziland and Zimbabwe) 
144-146, 164, 172, 173, 177, 178, 502, 515, 532, 538, 543, 545, 546 (15 
species) 

Western Africa 

(Benin, Gambia, Ghana, Guinea, Guinea-Bissau, Ivory Coast, Liberia, 
Mali, Mauritania, Niger, Nigeria, Senegal, Sierra Leone, Togo and 
Burkina Paso (formerly Upper Volta)) 

144, 147, 151, 152, 160, 161, 164, 167, 168, 172, 174, 178, 499, 500, 506, 513, 
515, 521, 523, 524, 526, 527(?), 529-531, 533, 534, 538, 544, 545, 548, 549 
(31-32 species) 

531. Papilio (Princeps) antimachus RARE (Benin (?), Ghana, Guinea, 
Ivory Coast, Liberia, Sierra Leone and Togo (?)) 

Angola 

144, 145, 147, 149, 150, 157, 160, 161, 164, 168, 172, 178,499-501,512,513, 
515, 521, 525-527, 531, 534, 538, 544, 545 (27 species) 
531. Papilio (Princeps) antimachus RARE 

Burundi 

144, 149(?), 161(?), 164, 172, 175, 178, 501(?), 503(?), 506, 510, 512, 515, 
521, 538, 540, 544, 545, 548(?), 550 (15-20 species) 
540. Papilio (Princeps) leucotaenia VULNERABLE 

Cameroon 

144, 147, 149, 152, 155, 157-162, 164, 167, 168, 170, 172, 174, 178,499-501, 
506, 510, 513, 515, 521, 523-526, 528, 529, 531, 533, 534, 538, 544, 545, 548 
(39 species) 

129 



Threatened Swallowtail Butterflies of the World 

158. Graphium (Arisbe) weberi INSUFFICIENTLY KNOWN 
531. Papilio (Princeps) antimachus RARE 

Central African Republic I 

144, 147(?), 149, 151, 152, 155, 157, 160, 161(?), 167(?), 168(?), 172, 178, 
y\jX 499(?), 500, 513, 515, 521, 523, 525-527, 529, 531, 534, 538, 544, 545, 548 
(24-29 species) 
531. Papilio (Princeps) antimachus RARE 



\ 



Chad 
Congo 



144, 168, 172, 178, 515, 538, 544(?), 545 (7-8 species) 

144, 147, 149, 152, 155, 157, 159-162, 164, 167, 168, 172, 174, 178, 499-501, 
503(?), 506, 513, 515, 521, 523-527, 529, 531, 534, 538, 543-545, 548, 550 
(37-38 species) 
531. Papilio (Princeps) antimachus RARE 

144(?), 164, 172, 178, 515, 538, 545 (6-7 species) ^ Ae " 

Equatorial Guinea , ^^^ /^ ^'^ 

(including Bioko (formerly Fernando Poo)) U ^^\\^'^'^ 

144, 149(?), 152(?), 157(?), 160(?), 164, 172, 178, 501, 510, 515, 521(?), «'^J.^> 
523(?), 524(?), 526, 531, 534(?), 538, 544, 545, 548 (13-21 species) ^ 

531. Papilio (Princeps) antimachus RARE 

Ethiopia 



Djibouti 



Gabon 



144, 148, 149, 164, 172, 177, 178, 499, 502, 506, 514, 515, 521(?), 538, 543, 
545 (15 or 16 species) 

144, 147, 149, 152(?), 156, 157, 159-62, 164, 167, 168, 170, 172, 174, 178, 
499(?), 501(?), 513, 515, 521, 523, 524, 525(?), 526(?), 531, 534(?), 538, 
544, 545 (25-31 species) 
531. Papilio (Princeps) antimachus RARE 



Kenya 



144, 148, 149, 164, 171-173, 176-178, 499, 501, 503, 506, 511, 512, 515, 

519-522, 532, 534, 538, 543-545, 550 (30 species) 

519. Papilio (Princeps) desmondi teita ENDANGERED 

Malawi 

144, 145, 146, 148, 164, 171-173, 177, 178, 502-504, 512, 515, 518, 519, 532, 
543-545, 547 (22 species) 

Mozambique 

144, 146, 148, 164, 169, 171-173, 177, 178, 502, 532, 538, 543-545 (16 
species) 

Rwanda 

144, 149(?), 161(?), 164, 172, 175, 178, 501, 503, 506, 510, 512, 515, 521, 

531, 538, 540, 544, 545, 548(?), 550 (18-21 species) 

531. Papilio (Princeps) antimachus RARE 

540. Papilio (Princeps) /eMCOfaen/a' VULNERABLE 

Somalia 

144, 164, 171, 172, 177, 178, 514, 515, 538, 543-545 (12 species) 

130 



Swallowtails of the world 

Sudan 

144, 148, 149, 164, 172, 178, 500, 501, 506, 512, 515, 521, 525, 526, 534, 538, 
544, 545, 548, 550 (20 species) 

Tanzania 

(including Zanzibar and Pemba Island) 

144, 145, 147-149, 161, 164, 165, 171-173, 17^178, 501-506, 511, 512, 515, 

518, 519, 521, 532, 538, 543-545, 547, 548, 550 (34 species) 

505. Papilio (Princeps) sjoestedti RARE 

Uganda 

144, 147-149, 161, 164, 168, 172, 175, 178, 499, 501, 503, 506, 510, 512, 513, 

515, 520-522, 525, 526(?), 529, 531, 534, 538, 540, 544, 545, 548, 550 (31-32 

species) 

531. Papilio (Princeps) antimachiis RARE 

540. Papilio (Princeps) leucotaenia VULNERABLE 

Zaire 

144, 145, 147, 149-155, 157, 160-162, 164, 167, 168, 170, 172-175, 178, 
499-501, 503, 506, 510, 512, 513, 515, 521, 523-527, 529, 531, 532, 
534, 538, 540, 543-545, 548 (48 species) 
153. Graphium (Arisbe) aurivilliusi INSUFFICIENTLY KNOWN 

531. Papilio (Princeps) antimachus RARE 

540. Papilio (Princeps) leucotaenia VULNERABLE 

Zambia 

144, 145, 147, 149, 150, 164, 172, 173, 178, 501-503, 512, 513, 515,518, 519, 

532, 538, 543-545, 547, 548 (23 species) 

NORTH AMERICA 

Canada 

23, 24, 37, 61, 208, 352, 334^356, 359, 391, 428, 430, 432-436 
(18 species) 

Mexico 

1, 62, 66, 69, 71, 81, 87-89, 96, 100, 102, 104, 109, 208, 211, 217, 218, 220, 
221, 226-228, 247, 248, 250, 251 , 254, 268, 352-358, 371-373, 378, 382-384, 
391, 394, 402-404, 407, 409, 430, 432 (52 species) 

1. Baronia brevicornis RARE 
378. Papilio (Heraclides) esperanza VULNERABLE 

U.S.A. 

23, 24, 37, 39(?), 61, 208, 211, 352, 354-359, 382, 384, 385, 388, 391, 407, 
428, 430-432, 434, 436-439 (28-29 species) 

385. Papilio (Heraclides) aristodemus ponceanus ENDANGERED 
(Florida) 

ISLANDS OF THE CARIBBEAN 

Bahamas 

210, 211, 379, 385, 391 (5 species) 

131 



Threatened Swallowtail Butterflies of the World 

Bermuda 

391 (1 species) 

Cayman Islands 

379, 380, 385 (3 species) 

Cuba 

65, 210, 211, 225, 379, 383, 385, 386, 388, 391, 399, 400, 432 (13 species) 

386. Papilio (Heraclides) caiguanabus INDETERMINATE 

Hispaniola (Dominical Republic and Haiti) 

63, 209, 211, 380, 383, 385, 387, 399 (8 species) 
209. Battus zetides VULNERABLE 

387. Papilio (Heraclides) aristor INDETERMINATE 

Jamaica 

64, 211, 374, 379, 381, 388, 399 (7 species) 

64. Eury tides (Protesilaus) marcellimis VULNERABLE 

374. Papilio (Pterourus) homerus ENDANGERED 

Lesser Antilles 

211,383, 388 (3 species) 

Puerto Rico 

211, 385(?), 399 (2-3 species) 

Trinidad and Tobago 

76, 83, 98, 211, 218, 221, 248, 259, 267, 383, 388, 390(?), 394, 407 (13-14 
species) 

CENTRAL AND SOUTH AMERICA 

Due to the relatively small size of its constituent countries, Central America has been 
treated regionally. 

Central America 

(Belize, Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua and 

Panama) 

62, 66, 68, 69, 71, 76, 81, 83, 87-90, 96-98, 100, 102, 104, 108, 109, 211, 

218-221, 227, 228, 247, 248, 250, 251, 254, 260, 268, 354, 356, 357, 360, 

367-371, 373, 382-384, 388, 391, 392, 394, 402, 404(?), 407, 409-411, 432 

(57-58 species) 

Argentina 

70, 71, 73, 74, 76(?), 83, 86, 91, 99, 111, 211, 213, 214, 216, 219, 222, 223, 
230, 233, 235, 243, 248, 257-259, 267, 364, 365, 368, 382, 383, 388, 394-397, 
407 (36-37 species) 

397. Papilio (Heraclides) himeros VULNERABLE 

Bolivia 

71, 76, 80, 83, 84, 92, 93, 95, 98, 101, 103, 104, 106, 111, 211, 216, 218, 219, 
221, 241, 244, 248, 252, 256, 257, 259, 262, 267, 362, 363, 365, 368(?), 369, 

375, 382, 383, 388, 389, 392-394, 396, 398, 410 (43-44 species) 
241 . Parides steinbachi INSUFFICIENTLY KNOWN 

393. Papilio (Heraclides) garleppi INSUFFICIENTLY KNOWN 

132 



Swallowtails of the world 

Brazil 

70-80, 82-86, 91, 92, 98, 99, 101, 106, 111, 112, 211, 214, 218, 219, 221-223, 
230, 231-237, 239, 240, 243, 244, 246, 248, 249, 252, 255-259, 261-267, 364, 
368, 369, 382, 383, 388, 389, 393-398, 407 (74 species) 
91. Eury tides (Protesilaus) lysithous harrisianus ENDANGERED 
112. Eury tides (Eury tides) iphitas VULNERABLE 
232. Parides ascanius VULNERABLE 
236. Parides hahneli RARE 

240. Parides pizarro INSUFFICIENTLY KNOWN 
249. Parides burchellanus VULNERABLE 
393. Papilio (Heraclides) garleppi INSUFFICIENTLY KNOWN 
397. Papilio (Heraclides) himeros VULNERABLE 

Chile 

211,213, 388(?) (2-3 species) 

Colombia 

67, 68, 71, 76, 80, 83, 84, 90, 92, 93, 96-98, 100, 103-108, 111, 21 1, 218-221, 
244, 247, 248, 251, 252, 254, 255, 259, 260, 264, 267, 268, 360, 362, 363, 
367-370, 376, 377, 382, 383, 388, 390-392, 394, 406, 407, 410, 41 1 , 432 (59 
species) 

Ecuador 

71,75,76,80,83,84,92-98,100,101,103,104,106,107,110,111,211,215, 
2 1 8 , 2 1 9 , 22 1 , 229 , 238 , 244 , 247 , 248 , 25 1-257 , 259 , 264 , 267-269 , 360-363 , 
366, 369, il5-?>ll, 382, 383, 388, 389, 392, 394, 401 , 405, 407, 410, 41 1 , 432 
(64 species) 

French Guiana 

71, 76, 80(?), 83, 84, 92, 98, 111, 211, 218, 219(?), 237, 242, 244, 248, 252, 

259, 261, 264, 265, 267, 268, 360, 382, 383, 388, 393, 394, 398, 407, 408 

(29-31 species) 

242. Parides coelus INSUFFICIENTLY KNOWN 

393. Papilio (Heraclides) garleppi INSUFFICIENTLY KNOWN 

408. Papilio (Heraclides) maroni INSUFFICIENTLY KNOWN 

Guyana 

71, 76, 80, 83, 84, 92, 98, 111, 211, 218, 219, 237, 244, 248, 252, 255, 259, 

26 1 , 264 , 265 , 267 , 268 , 360( ? ) , 369 , 382 , 383 , 388 , 393 , 394 , 398 , 407 (30-3 1 

species) 

393. Papilio (Heraclides) garleppi INSUFFICIENTLY KNOWN 

Paraguay 

70(?), 71, 72, 73(?), 74, 76, 82-86, 91, 99, 111, 211, 214, 219(?), 222, 223, 
230, 248(?), 258, 259, 267, 365(?), 368(?), 382, 383, 388, 394, 395, 407 
(26-32 species) 

Peru 

71, 76, 80, 83, 84, 92-95, 98, 101, 103, 104, 106, 110, 111, 211, 212, 215, 216, 
218, 219, 221, 229(?), 238-240, 244, 246, 248, 251-253, 255-257, 259, 262, 
264, 267, 362, 363, 366, 369, 375-377, 382, 383, 388, 389, 392, 394, 398, 405, 
407, 410, 432 (58-59 species) 
240. Parides pizarro INSUFFICIENTLY KNOWN 

133 



Threatened Swallowtail Butterflies of the World 

393. Papilio (Heradides) garleppi INSUFFICIENTLY KNOWN 

Surinam 

71, 76, 80, 83, 84, 92, 98, 111, 211, 218, 219, 221, 237, 244, 248, 252, 259, 

261, 264, 265, 267, 268, 360(?), 369, 382, 383, 388, 393, 394, 398, 407 (30-31 

species) 

393. Papilio (Heradides) garleppi INSUFFICIENTLY KNOWN 

Uruguay 

211, 222, 223, 364, 368(?), 382, 388, 395 (7 or 8 species) 

Venezuela 

67, 71, 76, 80(?), 83, 84, 90, 92, 96, 98, 106, 107(?), Ill, 211, 218, 219, 221, 
245, 248, 251, 252, 254, 255(?), 259, 260, 267, 268, 362, 370, 376, 382, 383, 
388, 390, 392, 394, 398(?), 407, 432 (35-39 species) 
245. Parides klagesi INSUFFICIENTLY KNOWN 

INDIAN OCEAN ISLANDS 

Including Madagascar, Andaman and Nicobar Islands. For Sri Lanka, see section on 
Middle Asia to Indochina and Japan. 

Andaman and Nicobar Islands 

125, 186, 189, 300-302, 314, 330, 415, 462, 463, 473, 484, 539 (14 species) 
125. Graphium (Pathysa) epaminondas INSUFFICIENTLY KNOWN 
(Andamans) 



-, Comoros 



P^-y^^.^"' ' 166, 507(?), 516, 545 (3-4 species) 

\-^' '^ 166. Graphium (Arisbe) levassori VULNERABLE 

^ ' s<^^ V 516. Papilio (Princeos) aristoohontes INDETERMl 



''S.- 



^ \ 516. Papilio (Princeps) aristophontes INDETERMINATE 

car 

143, 163, 179, 270, 507, 517, 535-538, 541, 542, 545 (13 species) 



C^o\ Madagascar 

! V^ 1 

535. Papilio (Princeps) morondavana VULNERABLE 

536. Papilio (Princeps) grosesmithi RARE 
542. Papilio (Princeps) mangoura RARE 

Mauritius 

508, 538 (2 species) 

508. Papilio (Princeps) manlius INDETERMINATE 

Reunion 

509, 538 (2 species) 

509. Papilio (Princeps) phorbanta VULNERABLE 
PACIFIC OCEAN ISLANDS 



All islands east of the Solomons, north to Hawaii. 

Fiji 

491 (1 species) 

134 



Swallowtails of the world 



Hawaiian Islands 

440, 539 (2 species) 

New Caledonia (including the Loyalty Islands) 
192, 412(?), 490, 573 (3-4 species) 

Vanuatu {New Hebrides) 

203, 484, 485, 487 (4 species) 

Western Samoa 

492 (1 species) 

Other Pacific Islands (including Lord Howe, Norfolk, Guam etc.) 
440, 473, 490, 498, 539 (5 species) 






135 



4 



Analysis of critical faunas 



The aim of this section is to analyse the world distribution of swallowtails so as: 

1) to identify countries with particularly rich, valuable or threatened swallow- 
tail faunas; 

2) to present a detailed analysis of the distribution and points of interest in the 
swallowtail fauna of a single country, Indonesia; 

3) to present selective notes on interesting aspects of a few other countries 
recognized as having important swallowtail faunas. 

Similar methods may be applied to any country and to other groups of animals or 
plants. Baseline data for an analysis may be obtainable from the lUCN Conservation 
Monitoring Centre. 

Introduction 

In their work of defining geographical units for conservation Dasmann (7, 8, 9) and 
Udvardy (28) recognized the need to consider the distribution of species as well as the 
distribution of ecosystems and biomes. Ideally, all species and all ecosystems should 
be catalogued , but of course such a task is impossible . Many groups of species remain 
too poorly known to be catalogued, and under man's influence ecosystems are 
changing too quickly for their boundaries to be identified. Udvardy finally proposed a 
system of biogeographical realms, provinces and biomes that included some 
consideration of the distribution of species, but relied heavily on an analysis of 
angiosperms, mammals and birds. Molluscs were the only invertebrates that received 
some attention (28). In terms of biomass, species richness, energy and nutrient flux 
arthropods dominate the animal life of terrestrial ecosystems from tropical rain 
forests to antarctic regions, yet they received no study whatsoever. Udvardy was 
aware of the anomaly, and implied that groups such as arthropods might reveal 
zoogeographical entities markedly different from those of land vertebrates (28). 
Certainly they would be expected to reveal patterns in finer detail. To some extent 
this has been borne out. For example, in a study of the Indo-AustraUan area it has 
been shown that although the faunal elements of butterflies and birds are well- 
matched, those of bats are somewhat simpler because of their more recent evolution 
and spread (14). In this book we cannot hope to redress the imbalance, but we shall 
demonstrate that simple analyses of insect groups can give information of value to 
conservation planners and administrators. 
The phylum Arthropoda is a vast assemblage of invertebrates comprising about 

137 



Threatened Swallowtail Butterflies of the World 

one million described species from a living total generally estimated at 2-5 million 
(18). Higher estimates of up to 30 million have been made (11), but have little 
credence so far. At least three quarters of these species are insects and it is perhaps 
because of this high diversity, rather than in spite of it, that they have so far received 
little attention in biogeographical analyses for conservation purposes. The diversity 
of insects certainly requires that survey work for conservation assessments needs to 
be highly selective, but there are no grounds for the view that insects (or other 
invertebrates) are too complex for their use to be contemplated. Certain groups are 
sufficiently well known ecologically and taxonomically, sufficiently visible and 
obtainable , to be of great value both for conservation planning from biogeographical 
analyses and for environmental monitoring on a more local scale (13). There is a vast 
literature on the zoogeography of insects, from the early observations on patterns by 
the father of zoogeography, A.R. Wallace (29), to modern numerical and cladistic 
analyses. Regrettably little of this work concerns itself with conservation, although 
some is certainly of considerable significance to the subject. It is hkely that different 
groups would be appropriate 'indicators' for the various biogeographical realms as 
well as for different ecological habitats. For example , the phytophagous Lepidoptera 
might prove valuable for terrestrial habitats but whereas the milkweed butterflies 
(Danainae) might be chosen for a study in South East Asia, they would be of little use 
in South America, where the related glasswings (Ithomiinae) are more diverse. For a 
number of reasons swallowtail butterflies have great potential for inclusion in 
faunistic analyses and environmental monitoring: 1) they are large, often brightly 
patterned and recognizable on the wing, 2) the males can laometimes be baited or 
decoyed, 3) they inhabit a wide range of latitudes, altitudes and biotopes, 4) there are 
numerous local, as well as widespread, species, subspecies and forms, 5) they often 
have specific requirements for foodplants, but the variety of plant families used is 
wide, particularly in temperate regions. 

In Europe and North America in recent years there have been numerous ad hoc 
efforts to protect important insect localities. The butterflies of San Bruno Mountain 
in San Francisco and the Large Blue (Maculinea arion) sites in Britain are good 
examples (30). In these cases the sites were identified because butterflies were known 
to be rare and threatened. If conservation is to advance as a practical but scientific 
discipline methods must be adopted that will identify conservation objectives in 
advance, not just in response to crises. As the World Conservation Strategy 
recommends, conservation objectives must be combined with social and economic 
objectives in formulating development policies (15). 

There are very few general surveys or biogeographical studies of insects that have 
played a major role in identifying key conservation requirements. One of the most 
detailed is the study by Pyle (22, 23) of the butterflies of Washington State, U.S.A. 
By superimposing the range limits of 116 species Pyle identified 17 biogeographic 
provinces. These did not compare closely with the major North American provinces 
of Dice (10), which were among the first faunistic divisions to be named for the state. 
A resemblance was found with Dalquest's (6) mammalian provinces, but his western 
Washington province included all or part of nine of Pyle's butterfly provinces. 
Different groups of organisms will never reveal identical faunal provinces, but there 
are clear indications that butterflies offer an excellent level of detail. Pyle's work 
showed that the proportion of protected areas varied very widely in the butterfly 
provinces, from 0.2 per cent to 57 per cent. From this information it was possible to 
recommend focal points for conservation action. 

Pioneering work by K.S. Brown Jr. on the distribution of neotropical forest 
butterflies has played a large part in locating those areas of South American rain 

138 



Critical faunas 

forest that survived the arid Pleistocene era, served as refugia for rain forest animals 
and plants, and are still rich in species (2, 3, 4, 5). The identification of the Peruvian 
rain forest refugia was based on the distribution of glasswing butterflies (Nymphali- 
dae: Ithomiinae) (4). Lamas (16) extended the study to cover the whole of Peru and a 
large number of species of Nymphalidae, Papilionidae and Pieridae. As in Pyle's 
study, the distribution ranges of the butterflies were mapped and biogeographical 
provinces identified. Twelve provinces were delimited in lowland, lower montane 
and upper montane forests, comparing favourably with Brown's presumed tropical 
forest refuge areas for Peru (4) . A further 36 provinces were identified in the lowland, 
lower montane, upper montane and paramo-puna non-forest zones (16). All 48 
provinces were clearly distinguishable on the basis of their butterfly faunas, many of 
which included endemic species or subspecies. A logical extension of the analysis 
would be to examine the protected areas within the 48 provinces and plan a 
conservation strategy for the future. 

A larger scale but complementary approach has been developed by Ackery and 
Vane-Wright in their worldwide study of the milkweed butterflies ( 1 ) . This group of 
insects is centred on South East Asia and as a consequence of a thorough 
distributional analysis it was possible to identify "critical faunas" for conservation, 
based on the whereabouts of narrowly distributed species. The technique is 
particularly valuable in identifying larger areas of responsibility, such as nations, 
islands or states. 

In the ensuing pages we shall try to combine these two methods in an effort to 
demonstrate how they may be used to identify priorities in a conservation strategy 
based on swallowtails. The method is intended to be applicable to any other group of 
animals or plants, indeed it may prove to be most effective when a range of life forms 
is analysed together. We shall identify countries with critical swallowtail faunas and 
try to examine the distribution and vulnerability of swallowtails within a few of those 
countries. It will not be possible to analyse the distributions as finely as Pyle did for 
Washington (22, 23) or Lamas did for Peru (16), nor would it be particularly valuable 
to do so without the identification of a precise need . What we wish to demonstrate and 
emphasize is that swallowtail and other butterflies can help in identifying conser- 
vation priorities, and that the methodology to do so is available. 

Methods 

This analysis is essentially an assessment of the distribution of endemic species . It is a 
form of biogeographical analysis but lacks the temporal element, seeking only to 
describe the distribution of present faunas, not their origins. Ackery and Vane- 
Wright used physical boundaries to delimit their units of endemism ( 1 ) but we choose 
to adopt political boundaries at the first level of analysis. Hence , whereas Ackery and 
Vane- Wright have analysed, for example, the faunas of Borneo, Sulawesi and 
Luzon, our analysis begins with Malaysia, Indonesia and the Philippines. In the 
second level of the analysis we examine the distribution of swallowtails in regions, 
provinces or other administrative blocks that comprise those countries with 
important faunas. The reason for choosing national and provincial boundaries is that 
these are the units within which conservation is administered. Two exceptions have 
been made in cases where countries are under the control ofa remote government. 
The French Overseas Departments of French Guiana, Reunion and New Caledonia 
have been considered individually, as have the Andaman and Nicobar Islands, which 
are administered by India. 
In following the method of analysis it will be useful to refer to Table 4.1. The initial 

139 



Threatened Swallowtail Butterflies of the World 

object is to find the smallest number of countries that includes all the 573 species of 
swallowtails recognized in this work. To do this, the distribution of endemics is 
analysed and the countries are listed in descending order of richness (Table 4.1, 
column A). Where more than one country has the same number of endemics the 
countries are listed in descending order of their total number of species (column F). 
Countries with endemic species clearly must accept ultimate responsibility for the 
conservation of those species, but if their endemic faunas could be positively 
conserved then their non-endemic species might also be protected by the same action. 
Hence in Indonesia (country number 1) 53 endemic swallowtails are found together 
with 68 non-endemic species (column B). Thus a total of 121 species have already 
been included (column E). In the Philippines (country number 2) 21 endemics are 
found, plus 28 non-endemics. However, 24 of those non-endemics are also found in 
Indonesia (column C), leaving only four non-endemics and 21 endemics, a total of 25 
species (column D) newly accounted for and to be added to the cumulative total 
(column E). In all, 43 countries contain one or more endemic swallowtails, and by 
repeating the analysis for each country in turn 561 species may be accumulated. The 
remaining 12 species are distributed between countries with no endemic swallowtails 
and are to be found in a minimum of eight countries, making 51 countries in all. 

It should be noted that the listing is only one of many possible forms of analysis and 
should not detract from other considerations. For example, Burma (country number 
44) has no endemics but has the fifth largest swallowtail fauna in the world and could 
clearly play a very important role in conservation. By contrast, Italy (country number 
51) has no endemics and a small total fauna of eight species, but two of these are 
threatened, Parnassius apollo (Rare) and Papilio hospiton (Endangered), the latter 
species requiring priority conservation action. Papilio hospiton is an example of a 
butterfly with a very narrow distribution that crosses national borders, in this case 
Corsica (France) and Sardinia (Italy); another good example is Papilio leucotaenia 
(Vulnerable) which has a narrow range spanning the borders of four central African 
countries. Clearly this analysis of critical faunas should wherever possible be used in 
addition to careful and independent national assessments. 

We cannot emphasize too strongly that this analysis provides no excuse for 
disregarding the conservation needs of swallowtail butterflies in unlisted countries. 

Results 

As Table 4.1 shows, the five countries with the highest swallowtail species endemism 
are Indonesia (53), Philippines (21), China (15), Brazil (11) and Madagascar (10). If 
their national conservation plans could ensure the protection of swallowtails these 
five countries alone could between them account for 309 swallowtail species, almost 
54 per cent of the world total. A further five countries, India, Mexico, Taiwan, 
Malaysia and Papua New Guinea, bring the total to 68 per cent. Increments decrease 
as further blocks of five countries are added, with 15, 20, 25, 30, 35, 40, and 45 
countries respectively including 77, 90, 93, 95, 96, 97 and 99 per cent of the world's 
swallowtails. 

Having identified these 51 countries of particular importance we would ideally 
have presented an analysis of each, examining the present conservation status of 
swallowtails within those countries and making recommendations for action. 
However, this would be a major undertaking beyond our present resources. Instead 
we offer below a single moderately detailed study , followed by brief notes on selected 
countries. Should this prove to be of interest to other countries they might choose to 
include a more detailed analysis in the formulation of their own 

140 













Critical faunas 


Table 4.1. Analysis 


of critical swallowtail faunas 










A 


B 


C 


D 


E 


F 




Endemic 


Non-ende 


mic species 


Newly 


Cumulative Total 




species 


Not 


Also 


account- 


species- 


species 






found in 
previous 
countries 


found in 
previous 
countries 


able 
species 
(A + B) 


list 

(Dn + 

D„.,) 


per 

country 

(A-l-B-l-C) 


1. Indonesia 


53 


68 





121 


121 


121 


2. Philippines 


21 


4 


24 


25 


146 


49 


3. China 


15 


61 


28 


76 


222 


104 


4. Brazil 


11 


63 





74 


296 


74 


5. Madagascar 


10 


3 





13 


309 


13 


6. India 


6 


8 


63 


14 


323 


77 


7. Mexico 


5 


37 


10 


42 


365 


52 


8. Taiwan 


5 





27 


5 


370 


32 


9. Malaysia 


4 


1 


51 


5 


375 


56 


10. P.N.G. 


4 


8 


25 


12 


387 


37 


11. U.S.A. 


4 


6 


19 


10 


397 


29 


12. Cuba 


4 


3 


6 


7 


404 


13 


13. Ecuador 


3 


19 


42 


22 


426 


64 


14. Colombia 


3 


10 


46 


13 


439 


59 


15. Australia 


3 


2 


14 


5 


444 


19 


16. Andaman & Nicobar 3 





11 


3 


447 


14 


17. Jamaica 


3 





4 


3 


450 


7 


18. Zaire 


2 


44 


2 


46 


496 


48 


19. Cameroon 


2 


2 


35 


4 


500 


39 


20. U.S.S.R. 


2 


11 


22 


13 


513 


35 


21. Tanzania 


2 


11 


21 


13 


526 


34 


22. French Guiana 


2 





29 


2 


528 


31 


23. Japan 


2 





20 


2 


530 


22 


24. Canada 


2 





16 


2 


532 


18 


25. Sri Lanka 


2 





13 


2 


534 


15 


26. Haiti 


2 


2 


4 


4 


538 


8 


27. New Caledonia 


2 





2 


2 


540 


4 


28. Comoro Is 


2 





2 


2 


542 


4 


29. Peru 




1 


57 


2 


544 


59 


30. Bolivia 







43 


1 


545 


44 


31. Venezuela 







38 


1 


546 


39 


32. Gabon 







30 


1 


547 


31 


33. Bhutan 







29 


1 


548 


30 


34. Ghana 




1 


22 


2 


550 


24 


35. Solomon Is 







14 


1 


551 


15 


36. Afghanistan 







18 


1 


552 


19 


37. South Africa 




1 


10 


2 


554 


12 ■ 


38. Iran 




1 


7 


2 


556 


9 


39. Vanuatu 







3 


1 


557 


4 


40. Mauritius 







1 


1 


558 


2 


41. Reunion 







1 


1 


559 


2 


42. Fiji 










1 


560 


1 


43. Western Samoa 










1 


561 


1 


44. Burma 





2 


66 


2 


563 


68 


45. Laos 





2 


46 


2 


565 


48 


46. Honduras 





1 


33 


1 


566 


34 


47. Argentina 





1 


36 


1 


567 


37 



141 



Threatened Swallowtail Butterflies of the World 



Table 4.1. (cont.) 







A 


B 


C 


D 


E 


F 






Endemic 


Non-endei 


Tiic species 


Newly 


Cumulative Total 






species 


Not 


Also 


account- 


species- 


species 








found in 


found in 


able 


list 


per 








previous 


previous 


species 


(D„ + 


country 








countries 


countries 


(A + B) 


D,.,) 


(A-t-B-t-C) 


48. 


Uganda 





2 


30 


2 


569 


32 


49. 


Ethiopia 





1 


15 


1 


570 


16 


50. 


Mozambique 





1 


15 


1 


571 


16 


51. 


Italy 





2 


6 


2 


573 


8 



conservation strategies, calling upon lUCN's information resources when needed. 
Much of the information needed to formulate such an analysis is already contained 
within this book. Lists of endemics and other species of interest can easily be 
generated by referring to section 3. 

Indonesia The swallowtails of Indonesia and their distribution between the 
islands are given in Table 4.2. Further information on these species will be found in 
the main species list in section 3. Indonesia as a whole has 44 per cent endemism in 
the swallowtails, a very high level which compares closely with the levels for 
mammals and birds (17). 

In Table 4.3 the relationships between the swallowtail faunas of seven Indonesian 
provinces are summarized. The geography of Indonesia has made the demarcation of 
these provinces relatively simple. The range of each province is defined at the bottom 
of Table 4.2. There are a few departures from Indonesian administrative provinces, 
notably in the Moluccas (Maluku) and Lesser Sundas, but we have followed the 
divisions made in the National Conservation Plan for Indonesia (27). 

Sumatra shares 42 of its 49 species with Malaysia, 36 of them with Peninsular 
Malaysia and 34 with Eastern Malaysia. Only two of these species, Troides aeacus 
and Papilio iswarioides, are not found on other Indonesian islands. Of the remaining 
seven, Graphium cloanthiis has a strange disjunct range mainly in India, Indo-China 
and China and the other six are Indonesian endemics. Four are found only on 
Sumatra, namely Graphium sumatranum, Atrophaneura hageni, Papilio forbesi and 
Papilio diophantus . Two species, Atrophaneura priapus and Troides vandepolli, are 
shared with Java. Both Papilio forbesi and P. diophantus are from northern Sumatra, 
which includes the Gunung Leuser National Park, over 8000 sq. km in extent. 
Papilio forbesi females mimic A . hageni which is restricted to the high plateau and 
probably also occurs in Gunung Leuser as well as other highland areas. P. forbesi 
may be present in the Sumatra Selatan Proposed National Park in the southwest 
corner of the island. This reserve is over 3500 sq. km in extent and covers the 
southern end of the Barisan range, which reaches lower altitudes than in the north. 
Unfortunately there has been extensive damage in certain parts of the reserve, 
mainly from agriculturalists (23, vol. 2). In general Sumatra's butterfly fauna is likely 
to be well protected. However, much of its fauna is shared with Peninsular Malaysia 
and with the increasing extent of logging and the imminent disappearance of virtually 
all the lowland forest in that state, Sumatra's forests will, in a world context, become 
increasingly important as conservation zones. 

Kalimantan has no known endemic swallowtails, sharing all its species with the 
Eastern Malaysian states of Sarawak and Sabah. Thirty-five of its 36 species are 



142 



Critical faunas 
Table 4.2. Distribution of swallowtail butterflies within Indonesia 





IN 


SM 


KA 


JA 


LS 


sw 


MO 


IJ 


Order LEPIDOPTERA 


















Family Papilionidae 


















Meandrusa payeni 


+ 


+ 


+ 


+ 










Lamproptera meges 


+ 


+ 


+ 


+ 




+ 






Lamproptera curias 


+ 


+ 


+ 


+ 










Graphium aristeus 


+ 


+ 


+ 


+ 


+ 




+ 


+ 


Graphium rhesus* 


+ 










+ 


+ 




Graphium dorcus* 


+ 










+ 






Graphium androcles* 


+ 










+ 


+ 




Graphium euphrates 


+ 










+ 


+ 




Graphium antiphales 


+ 


+ 


+ 


+ 


+ 


+ 






Graphium agetes 


+ 


+ 


+ 












Graphium encelades* 


+ 










+ 






Graphium delesserti 


+ 


+ 


+ 


+ 










Graphium macareus 


+ 


+ 


+ 


+ 










Graphium ramaceus 


+ 


+ 


+ 












Graphium megarus 


+ 


+ 


+ 


+ 










Graphium deucalion ^ — 


+ 










+ 


+ 


+ 


Graphium thule 


+ 














+ 


Graphium arycles 


+ 


+ 


+ 


+ 










Graphium bathycles 


+ 


+ 


+ 


+ 










Graphium meyeri* 


+ 










+ 


+ 




Graphium eurypylus 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Graphium doson 


+ 


+ 


+ 


+ 


+ 








Graphium evemon 


+ 


+ 


+ 


+ 










Graphium agamemnon 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Graphium macfarlanei 


+ 












+ 


+ 


Graphium weiskei 


+ 














+ 


Graphium stresemanni* 


R 












R 




Graphium codrus 


+ 










+ 


+ 


+ 


Graphium empedovana 


+ 


+ 


+ 


+ 










Graphium cloanlhus 


+ 


+ 














Graphium sumatranum* 


+ 


+ 














Graphium monticolum* 


+ 










+ 






Graphium milon* 


+ 










+ 


+ 




Graphium sarpedon 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Graphium wallacei 


+ 








+■ 


+ 






Cressida cressida 


+ 








+ 








Atrophaneura kuehni* 


+ 










+ 






Atrophaneura luchti* 


R 






R 










Atrophaneura hageni* 


+ 


+ 














Atrophaneura priapus* 


+ 


+ 




+ 










Atrophaneura sycorax 


+ 


+ 




+ 










Atrophaneura nox 


+ 


+ 


+ 


+ 










Atrophaneura dixoni* 


+ 










+ 






Atrophaneura neptunus 


+ 


+ 


+ 












Atrophaneura palu* 


K 










K 






Atrophaneura coon 


+ 


+ 




+ 










Atrophaneura oreon* 


+ 








+ 








Atrophaneura liris* 


+ 








+ 








Atrophaneura polyphontes* 


+ 










+ 


+ 




Atrophaneura aristolochiae 


+ 


+ 


+ 


+ 


+ 


+ 







143 



Threatened Swallowtail Butterflies of the World 



Table 4.2. (cont.) 



< 





IN 


SM 


KA 


JA 


LS 


sw 


MO 


IJ 


Atrophaneura polydorus 


+ 






+ 


+ 




+ 


+ 


Trogonoptera brookiana 


+ 


+ 


+ 












Troides hypolitus* 


+ 










+ 


+ 




Troides cuneifera 


+ 


+ 




+ 






' 




Troides amphrysus 


+ 


+ 


+ 


+ 










Troides miranda 


+ 


+ 


+ 












Troides prattorum* 


I 












I 




Troides aeacus 


+ 


+ 














Troides dohertyi* 


V 










V 






Troides Helena 


+ 


+ 


+ 


+ 


+ 


+ 






Troides oblongomaculatus 


+ 










+ 


+ 


+ 


Troides criton* 


+ 










+ 






Troides riedeli* 


+ 








+ 








Troides plato* 


+ 








+ 








Troides vandepolli* 


+ 


+ 




+ 










Troides haliphron* 


+ 








+ 




+ 




Troides staudingeri* ^^ 
Ornithoptera goliatK^ 


+ 








+ 








+ 


" 










+ 


+ 


Ornithoptera tithonus* 


K 














K 


Ornithoptera rothschildi* 


I 














I 


Ornithoptera chimaera 


I 














I 


Ornithoptera paradisea 


I 














I 


Ornithoptera meridionalis 


V 














V 


Ornithoptera aesacus* 


I 












I 




Ornithoptera croesus* 


V 












V 




Ornithoptera priamus 


+ 








+ 




+ 


+ 


Papilio clytia 


+ 








+ 








Papilio paradoxa 


+ 


+ 


+ 












Papilio veiovis* 


+ 












+ 




Papilio slateri 


+ 


+ 


+ 












Papilio laglaizei 


+ 














+ 


Papilio euchenor 


+ 














+ 


Papilio gigon* 


+ 










+ 


+ 




Papilio demolion 


+ 


+ 


+ 


+ 


+ 








Papilio lampsacus* 


+ 






+ 










Papilio forbesi* 


+ 


+ 














Papilio oenomaus* 


+ 








+ 








Papilio ascalaphiis* 


+ 










+ 


+ 




Papilio rumanzovia 


+ 












4- 




Papilio deiphobus* 


+ 












+ 




Papilio memnon 


+ 


+ 


+ 


+ 


+ 








Papilio biseriatus* 


+ 








+ 








Papilio iswara 


+ 


+ 


+ 












Papilio sataspes* 


+ 










+ 


+ 




Papilio helenus 


+ 


+ 


+ 


+ 


+ 








Papilio iswaroides 


+ 


+ 














Papilio jordani* 


R 












R 




Papilio polytes 


+ 


+ 


+ 


+ 


+ 




+ 




Papilio alphenor 


+ 










+ 


+ 




Papilio ambrax 


+ 














+ 


Papilio diophantus* 


+ 


+ 














Papilio nephelus 


+ 


+ 


+ 


+ 











144 



V' 



Critical faunas 
Table 4.2. (cont.) 





IN 


SM 


KA 


JA 


LS 


sw 


MO 


IJ 


Papilio hipponous* 


+ 






+ 










Papilio fuscus 


+ 




+ 






+ 


+ 


+ 


Papilio canopus 


+ 








+ 








Papilio albinus 


+ 












+ 


+ 


Papilio heringi* 


+ 












+ 




Papilio inopinatus* 


+ 








+ 








Papilio gambrisius* 


+ 












+ 




Papilio tydeus* 


+ 












+ 




Papilio aegeus 


+ 














+ 


Papilio demoleus 


+ 


+ 






+ 


+ 


+ 




Papilio blumei* 


+ 










+ 






Papilio palinurus 


+ 


+ 


+ 












Papilio paris 


+ 


+ 




+ 










Papilio karna 


+ 


+ 


+ 


+ 










Papilio neumoegeni* 


V 








V 








Papilio peranthus* 


+ 






+ 


+ 


+ 






Papilio pericles* 


+ 








+ 








Papilio lorquinianus* 


+ 












+ 


+ 


Papilio ulysses 


+ 












+ 


+ 



Key: 

IN: Indonesia 

SM: Sumatra, including Nias, Simeuiue, Siberut, Lingga, Bangka, Belitung, Riau, 

Batu, Enggano, Mentawai 
KA: Kalimantan, including Bunguran (Natuna Is) 
JA: Java and Bali, including Madura, Kangean and Bawean 
LS: Lesser Sundas, including Lombok, Sumbawa, Sumba, Tenggara, Timor, Roti, 

Wetar, Leti, Moa — Babar, Tanimbar, Flores 
SW: Sulawesi, including Talaud, Sangihe, Salayar, Banggai, Tanahjampea, Butung, 

Tukangbesi I. 
MO: Moluccas, including Halmahera, Bacan, Sula, Buru, Ambon, Seram, Obi, 

Morotai 
IJ: Irian Jaya, including Kai, Aru, Misool, Waigeo, Biak, Saiawati 

Note: Records indicate presence over part or all of the areas listed. Detailed records are 
available from our database on request. Graphium procles (Indeterminate), Graphium 
stratiotes, Troides andromache (Indeterminate) and Papilio acheron (Rare) are all endemic to 
Borneo but have not as yet been recorded on the Kalimantan side of the border and are, 
therefore, not listed above. 
*: Endemics to Indonesia 
+ : Records exist for this region 
V: Vulnerable in these regions 
R: Rare in these regions 
I: Indeterminate in these regions 

K: Insufficiently Known in these regions 



also shared with Sumatra (Table 4.3). Borneo as a whole has four endemics, 
Graphium stratiotes, Graphium procles (Indeterminate) Troides andromache 
(Indeterminate) and Papilio acheron (Rare). With the possible exception of G. 

145 



Threatened Swallowtail Butterflies of the World 

procles, which has a restricted distribution in the Crocker range of Sabah, all of these 
might well be expected to occur in northern Kalimantan, an area well served by the 
giant Sungai Kayan-Mentarang Reserve (4000 sq. km) (27, vol. 5). The borders of 
this reserve are undergoing reassessment and possible alteration as a result of the 
large human populations inside. A survey of the butterflies could add extra valuable 
criteria towards the final decisions. 

The Javanese fauna also has close links with Sumatra and the mainland, sharing 30 
of its 35 species with Malaysia. Atrophaneura priapus is found in southern Sumatra as 
well as Java and its relationship with the Sumatran species A . hageni is not at all clear. 
The Rare Javan endemic y4. luchti from the mountains of eastern Java differs only 
slightly from A . priapus and there are doubts as to its taxonomic status. Regardless of 
this, the Ijen, Merapi and Moeleng reserves may be important to the survival of the 
butterfly. Troides vandepolli from north-western Java and western Sumatra is 
uncommon and protected by Indonesian law. Papilio lampsacus is endemic to 
western Java, occurring only on Gunung (Mt) Gede and Gunung Mas. The Ujung 
Kulon and Gunung Gede-Pangrango National Parks may effectively protect the 
habitat of both these species. Their status should be carefully monitored and 
immediate action taken against habitat disruption, particularly on Gunung Gede. 

As might be expected from the biogeographical considerations, the regions lying 
on the Asian continental plate (the 'Sundaic' regions of Malaysia, Sumatra, Borneo, 
Palawan, Java and Bah) show strong faunistic similarities in their swallowtails, but 
quite marked differences begin to develop in the 'Wallacean' regions eastwards from 
the Lesser Sundas and Sulawesi. Of the 30 species in the Lesser Sundas only 14 are 
shared with Malaysia. Some species with a mainly Australasian distribution begin to 
appear, such as Cressida cressida and Papilio canopus. Of the nine endemics, Troides 
riedeli (Tanimbar Is), T. plato (Timor) and T. staudingeri (Babar group) are 
protected by law. Widespread Lesser Sundas endemics include the common 
Atrophaneura liris and Papilio oenomaus and the scarcer P. perides. Atrophaneura 
oreon is found only on Sumba, Alor, Flores and Solor, Papilio biseriatus only on 

Table 4.3. Patterns of species richness and distribution in Indonesian swallowtails 



Province''' 


No. of 


Endemics 


Non-endemics 




No. 1 


of species shared with 






Species 


No. % 












































1 


2 


3 


4 


5 


6 


7 


1. Sumatra 


49 


4 8 


45 




32 


13 


9 


5 


4 




2. Kalimantan 


36(2' 





36<2> 


35 


— 


26 


12 


9 


5 


5 


3. Java(+ Bali) 


35 


2 6 


33 


32 


26 


— 


13 


9 


4 


4 


4. Lesser Sundas 


30 


10 33 


20 


13 


12 


13 


— 


10 


7 


6 


5. Sulawesi 


38 


11 29 


27 


9 


9 


9 


10 


— 


19 


6 


6. Moluccas 


37 


9 24 


29 


5 


5 


4 


6 


19 


— 


15 


7. Irian Jaya 


26 


2 8 


24 


4 


5 


4 


5 


6 


15 


— 


8. Indonesia 


121<2' 


53'^' 44 


68<2' 

















'" Limits of provinces are defined in Table 4.2 

*2' Not including Graphium stratiotes, Graphium procles, Troides andromache or Papilio 

acheron, all of which are endemic to Borneo but have not been recorded on the Kalimantan 

side of the border. 
*^' 37 species are endemic to only one of areas 1-7 ; a further 16 species are endemic to Indonesia 

but occur in more than one of areas 1-7. 

146 



Critical faunas 

Timor, P. inopinatus on Romang, Babar, Damar and Tanimbar and P. neumoegeni 
only on Sumba. The last of these species is category Vulnerable and the other three 
are certainly in need of closer study and assessment. In general the Lesser Sundas are 
under-represented in terms of protected areas. The Indonesian Conservation Plan 
(27, vol. 4) includes many recommendations for new reserves and parks, including 
acquisition of new reserves on Sumbawa, Flores, Timor and Sumba. The sites should 
be surveyed for local swallowtails. 

Sulawesi and its associated islands (see Table 4.2) contain 1 1 endemic swallowtails, 
second only to New Guinea and its associated islands, which have 14. The rich 
endemic fauna of Sulawesi is possibly the result of a very complex geological history 
which has included the fusion of island arcs from both the Sunda and Australasian 
continental shelves. In the swallowtails there is clear evidence of a stronger rela- 
tionship with the Australasian region, 19 species being shared with the Moluccas 
(including Sula) but only 9 with Kalimantan (Table 4.3). Unfortunately many 
Sulawesi biotopes have been and continue to be destroyed by deforestation (25). 
Large areas stretching between Ujung Pandang (Makassar) and the Northern and 
Central Districts are covered by the unpalatable and useless grass known as lalang 
(Imperata cylindrica) (25), implying severe overexploitation and deterioration of 
soils. 

Two endemic species of Graphium, G. dorcus and G. monticolum are mainly 
restricted to the high mountains of Sulawesi , dorcus in the northern andcentral areas, 
rarely southern, monticolum mainly central and southern above 1000m (12). G. 
dorcus is rare and its habitat is not properly known, but G. monticolum is quite 
common (12). Two endemics are widespread in Sulawesi, Papilio veiovis and Papilio 
blumei. They are likely to be recorded in a number of parks and reserves. Southern 
Sulawesi is believed to be the main area for Graphium encelades, but the female is 
unknown and males are very rare. More data are needed to ascertain its conservation 
status. Two Atrophaneura species, A. dixoni and A. palu (Insufficiently Known), 
have a mainly central Sulawesi distribution. Both are uncommon, the latter particu- 
larly so. The Lore-Lindu Proposed National Park in central Sulawesi may be an 
important locality for these species. Northern Sulawesi has a particularly vulnerable 
swallowtail fauna, including the endemics Atrophaneura kuehni (very rare, also 
eastern), A. dixoni (also central) and Papilio jordani (Rare). The Dumoga-Bone 
National Park in northern Sulawesi, set up in a joint venture between the Indonesian 
Government, WWF/IUCN and the World Bank, is likely to be of particular value to 
conservation of the region's wildlife. In eastern Sulawesi the Morowali Reserve may 
prove to be of equal value. Sulawesi is the main centre for the widespread species 
Papilio ascalaphus, an Indonesian endemic also found on the Salayar and Sula 
islands. Just north of Sulawesi lie the Talaud and Sangihe Islands, which have two 
important endemic swallowtails, Troides dohertyi (Vulnerable) and Papilio hippo- 
nous. Papilio alphenor is also found here, as well as in the Philippines and the 
Moluccas, but not on the Sulawesi mainland. The need for a reserve on Sangihe island 
is recognized in the Indonesian Conservation Plan and Gunung Sahendaruman has 
been suggested (27, vol. 6). 

The Moluccas and Sulawesi share a number of important Indonesian endemics 
between them . Some of them are only on Sulawesi and the Sula Islands, not reaching 
further east, namely Graphium rhesus, G. androcles, G. meyeri, Papilio gigon, P. 
ascalaphus and P. sataspes. These species would need to be considered in a conser- 
vation plan for either the Moluccas or Sulawesi. Human populations are low in the 
Sula Islands and much of the area is thickly forested. A reserve is urgently needed and 

147 



Threatened Swallowtail Butterflies of the World 

has been proposed for Pulau Taliabu. Other Indonesian endemics extend from 
Sulawesi across to the main Moluccan islands, these including G. milon, Atro- 
phaneura polyphonies and Troides hypolitus. 

In addition, there are nine swallowtail species restricted to the Moluccas, out of a 
total of at least 25 endemic butterflies. Six of these are found on Halmahera and its 
associated islands, namely Troides criton (fairly common, protected by law), 
Ornithoptera aesacus (Obi only. Indeterminate, also protected), O. croesus (Vul- 
nerable), Papilio deiphobus, P. heringi (very poorly known and rare) and P. tydeus 
(common). These islands have the richest fauna and flora in the Moluccas, including 
35 (nearly half) of the province's endemic birds. Despite several proposals for 
establishing reserves in the group, none yet exist or have been approved. Defores- 
tation through commercial timber exploitation is having an increasing impact on 
Halmahera, and the lack of protected areas is a matter for concern (24). In the 
southern part of the Moluccas three further endemic species are found. Graphium 
stresemanni (Rare) and Papilio gambrisius both occur on Seram and are probably 
present in the 1800 sq. km Manusela Reserve. The Indonesia Conservation Plan 
suggests another reserve in the mountains of the Sahuai Peninsula (27, vol. 7). 
Finally, Troides prattorum (Indeterminate) is endemic to Buru, a steeply forested 
mountainous island once used as a prison . There are no reserves on the island , but the 
proposed Gunung Kelapat Muda Reserve (27) would almost certainly protect this 
endemic birdwing. In general the superbly rich butterfly fauna of the Moluccas is very 
poorly protected. The existing reserve system is quite inadequate but the proposals in 
the Indonesian Conservation Plan, if accepted, would remedy this regrettable 
situation. 

Irian Jaya has 26 of the 42 species found in New Guinea and its associated islands. 
Endemics to Irian Jaya and its islands include Ornithoptera tithonus (Irian Jaya, 
Waigeo, Salawati and Misool, Vulnerable) and O. rothschildi, (Arfak mountains 
only, western Irian Jaya, Indeterminate) both listed as threatened and considered in 
detail in section 6. A further three birdwing species, O. chimaera (Indeterminate), O. 
paradisea (Indeterminate) and O. meridionalis (Vulnerable) are New Guinea 
endemics also discussed in section 6. There are two endemic Graphium species in 
New Guinea, G. thule and G. weiskei, the latter being common and widespread. The 
former is poorly known and rarely collected, but may have been overlooked as it is a 
mimic of common species of Danainae. Finally, the two endemic Papilio species, P. 
laglaizei and P. euchenor are widespread and common, although the former, an 
interesting mimic of the day-flying moth Alcidis agarthyrsus (Uraniidae), is quite 
localized. 

Irian Jaya already has a fine network of reserves, including the giant Lorentz 
Reserve (16 750 sq. km) on the south coast and reserves in western Waigeo, but there 
are no fully designated national parks. As the Indonesian National Conservation 
Plan says, Irian Jaya is the richest and most important province in terms of faunal and 
floral wealth (27, vol. 7). The magnificent birdwing butterflies are a very significant 
part of the fauna and it is vital to ensure that these are thoroughly protected. There 
are great opportunities for conservation in the province, but land use conflicts must 
be quickly resolved. The potential for wildlife industries is very great, to include 
birdwing ranching as well as crocodile farms and deer ranches. 

To summarize, Indonesia has the finest swallowtail fauna in the world, certainly 
worthy of consideration in the country's conservation planning. On the Sunda shelf 
the Sumatran fauna is rich, the Kalimantan fauna poorly studied. The Javan 
endemics are as safe as the island's reserved areas, but those in the Lesser Sundas are 
poorly protected. Sulawesi has a very rich swallowtail fauna, probably well 
represented in parks and reserves, but poorly known. The northern Moluccas include 

148 



Critical faunas 

important endemic swallowtails that are very poorly protected and require priority 
action. Irian Jaya has a spectacular fauna that appears to be quite well protected and 
has great potential for interpretation to local and foreign visitors and entomologists. 
Indonesia has at least 17 rare and threatened species of swallowtails. The main threat 
is loss of habitat as a result of agriculture, commercial logging and other forms of 
human impact. 

Philippines With a total of 49 species and 21 endemics, nine of which are 
threatened (one Endangered, five Vulnerable, one Rare, two Indeterminate), the 
Philippines has one of the most important swallowtail faunas in the world (Table 4.4). 
An important factor contributing to this is that the Philippines span a wide latitudinal 
and biogeographical belt, from Palawan which is the only Philippine island on the 
Sunda shelf, to Luzon with its Asian mainland connections and Mindanao with 
submerged island arc connections to Sulawesi. This variety is evidenced by the 
frequent occurrence of pairs of closely related but distinct species that replace each 
other in Palawan (and Borneo) and the rest of the Philippines. Examples of such pairs 
are Atrophaneura aristolochiae and A . kotzebuea , Troides plateni{'Pa\2iVjan endemic) 
and T. rhadamantus , Papilio helenus and P. hytaspes. Parts of the fauna are still very 
poorly known, and the fact that new species have been discovered on various islands 
during the past 20 years implies that still more might be expected. Recently described 
species include Graphium sandawanum (\911 , Vulnerable), Papilio osmana (1967, 
Vulnerable) and P. carolinensis (1967, Vulnerable), P. chikae (1965, Endangered). 
Three threatened species are considered to be relicts of the Ice Age climate, and they 
now survive only on high mountains. These are Graphium sandawanum (Vulner- 
able) on Mt Apo, and Papilio benguetanus (Vulnerable) and P. chikae (Endangered) 
in the Cordillera Central of Luzon. Such localities are particularly vulnerable to 
disturbance and slow to recover. Many of the central Philippine islands are almost 
completely turned over to agriculture: Cebu, for example, has no forest left. In many 
places inappropriate agricultural methods have degraded the soils to such an extent 
that only the unpalatable grass Imperata cylindrica will grow. Deforestation on 
Mindanao is of particular international concern and has been aggravated by extensive 
fires following the drought of 1982-3. Samar still has extensive forests with great 
potential for conservation. 

Table 4.4. Distribution of swallowtail butterflies within the Philippines (after 
Tsukada and Nishiyama) (22) 

PH LU MR SA LE PN NE CE BO MN PL 

Order LEPIDOPTERA 

Family Papilionidae 
Lamproptera meges 
Lamproptera curias 
Graphium aristeus 
Graphium euphrates 
Graphium decolor 
Graphium idaeoides* 
Graphium delesserti 
Graphium macareus 
Graphium megaera* 
Graphium stratocles* 
Graphium arycles 
Graphium bathycles 
Graphium eurypylus 

149 



+ 


+ 


+ 


+ 


+ 


+ 










+ 


+ 








+ 


+ 


+ 






+ 


+ 


+ 


+ 


+ 


R 


R 




R 


R 


+ 










+ 

T 










+ 


+ 


+ 






+ 










+ 










+ 


+ 


+ 


+ 


+ 



+ 


+ 




+ 


+ 


+ 




+ 


+ 


+ 




+ 




+ 




I 


+ 


+ 




+ 




+ 


+ 


+ 



Threatened Swallowtail Butterflies of the World 
Table 4.4. (cont.) 





PH 


LU 


MR 


SA 


LE 


PN 


NE 


CE 


BO 


MN 


PL 


Graph ium doson 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Graphiuin agamemnon 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Graphium codrus 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 




Graphium empedovana 


+ 




















+ 


Graphium sandawanum* 


V 


















V 




Graphium sarpedon 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Atrophaneura semperi* 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Atrophaneura neptunus 


+ 




















+ 


Atrophaneura schadenbergi* 


V 


V 




















Atrophaneura mariae* 


+ 


+ 




+ 


+ 








+ 


+ 




Atrophaneura phegeus* 


+ 






+ 


+ 






+ 




+ 




Atrophaneura phlegon* 


+ 




+ 






+ 








+ 




Atrophaneura atropos* 


I 




















I 


Atrophaneura kotzebuea* 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 




Atrophaneura aristolochiae 


+ 




















+ 


Trogonoptera trojana* 


+ 




















+ 


Troides magellanus 


+ 


+ 




+ 


+ 






+ 


+ 


+ 




Troides plateni* 


+ 




















+ 


Troides rhadamantus* 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 




Papilio clytia 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Papilio paradoxa 


+ 




















+ 


Papilio osmana* 


V 








V 










V 




Papilio carolinensis* 


V 


















V 




Papilio benguetanus* 


V 


V 




















Papilio demotion 


+ 




















+ 


Papilio rumanzovia 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 




Papilio lowi 


+ 




















+ 


Papilio antonio* 


+ 








+ 










+ 




Papilio hystaspes* 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 




Papilio helemis 


+ 




















+ 


Papilio alphenor 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Papilio pitmani 


+ 


+ 














+ 


+ 


+ 


Papilio demoleus 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Papilio daedalus* 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


Papilio chikae* 


E 


E 




















Papilio karna 


+ 




















+ 



Key: 

PH: Philippines 

LU: Luzon, including Batan Islands, Babuyan islands and Polillo Islands 

MR: Mindoro 

SA: Samar 

LE: Leyte 

PN: Panay 

NE: Negros 

CE: Cebu 

BO: Bohol 

MN: Mindanao 

PL: Palawan, including Balabac and Calamian Islands 

Note: Records indicate presence over part or all of the areas listed. Detailed records are 
available from our database on request. 

150 



Critical faunas 



Table 4.4. (cont.) 

*: Endemics to Philippines 

+ ; Records exist for this region 

E: Endangered in these regions 

V: Vulnerable in these regions 

R: Rare in these regions 

I: Indeterminate in these regions 



The national park system in the Philippines was analysed in a 1982 report that drew 
attention to the deficiencies in the existing state of affairs (21). Considerable 
confusion existed on the purposes, objectives and management of national parks and 
there was evidence of substantial violation of protected area boundaries. Since that 
report the situation has apparently not improved. Sections of Mt Apo National Park 
(which has an endemic swallowtail) have been excised and other protected areas are 
reported to be neglected. It is difficult to obtain precise data on the protected areas of 
the Philippines, but we understand that the national system is under review. For the 
sake of the exquisite fauna and flora of the Philippines and of the natural resources of 
the Filipino people this review should be completed and its recommendations 
consolidated and enforced at the earliest opportunity. Conflicts between the 
exploitation and conservation of the natural resources of the Philippines are 
increasing day by day. 

China China is a very large country with a wide range of habitats and has a 
huge swallowtail fauna of 104 species including 15 endemics (see section 3). Five 
species are threatened or Insufficiently Known, i.e. Parnassius apollo (Rare), 
Bhiitanitis mansfieldi (Rare), Bhiitanitis thaidina (Rare), Luehdorfia chinensis 
(Insufficiently Known) and Teinopalpus aureus (Insufficiently Known) (see section 
6). China is particularly rich in Parnassius and has a fauna of 26 species including the 
three endemics P. szechenyii, P. cephalus, P. przewalskii and several rare and poorly 
known species, including P. acdestis, P. loxias, P. acco, P. hannyngtoni, P. actius and 
P. tianschanicus. All these species have a montane distribution in western and/or 
northern China, but there is too little information available to make specific 
conservation recommendations. In the genus Graphium three species are very poorly 
known, G. mandarinus, G. alebion and G. leechi, the last two being endemic. Five 
species of Atrophaneura. A. daemonius,A. plutonius,A. mencius,A. impediens and 
A. hedistus, are poorly known and/or rare. Some of these are also restricted to higher 
altitudes. Of the 25-27 species in the genus Papilio. P. elwesi (endemic), P. syfanius 
and P. krishna are rare and require further study, but most of the others have a wide 
distribution. Information on the ecology and precise distribution of Chinese 
swallowtails is very scarce. As a result it is difficult for an outsider to relate China's 
environmental problems to the conservation of her swallowtails. Suffice it to say that 
the Chinese swallowtail fauna is of international value and concern, stretching as it 
does from tropical climates to the snows of the Himalayas. The faunas of the western 
and northern mountainous regions and plateaux are of particular significance. 

In recent years Japanese entomologists have forged links with Chinese in Sichuarl 
and Yunnan, resulting in important collections of Bhutanitis mansfieldi and the 
slightly more common B. thaidina. In years to come the accessibility of these 
important regions will hopefully improve and opportunities for joint programmes of 
environmental assessment will be possible. It is important that commercial collecting 
is not encouraged in the absence of impact assessment and monitoring. There is 
potential for development of seasonal butterfly ranches in western China. 

151 



Threatened Swallowtail Butterflies of the World 

Brazil Brazil has a very rich swallowtail fauna of 74 species, including 11 
endemics. Seven species and one subspecies have been given threatened status, three 
of the species also being found in other countries. Threats directly affecting these taxa 
are described in section 6. A preliminary analysis of the Brazilian fauna suggests that 
southern Brazil (from Rio Grande do Sul to Sao Paulo) has a surprisingly rich fauna of 
about 39 species; it is not only the Amazon basin that needs consideration. At one 
time the Atlantic seaboard of eastern Brazil was covered in a strip of semi-deciduous 
forest, but only 1-2 per cent remains intact. Many species of animals and plants are 
threatened with extinction, including primates like the Golden-headed Lion Tamarin 
{Leontopithecus chrysomelas) and the Woolly Spider Monkey {Brachyteles arach- 
noides). The beautiful butterfly Parides ascanius (Vulnerable) is in the same 
position. The forests of the Mato Grosso are also rich in species (about 37 in all), 
while still more records might be expected from Amazonas (about 43 species so far). 

Madagascar Madagascar has only 13 swallowtails, but ten of these, (77 per 
cent) are endemic. The fauna has developed mainly from African stock with one 
notable exception, Atrophaneura anterior, a troidine species with relatives in Asia 
and South America, but not in Africa. A chain of mountains runs from north to south 
down the spine of Madagascar, the eastern side is wet and forested, the western side 
relatively dry with a variety of woodland and savanna ecosystems. Four endemics are 
mainly distributed in the dry west and central regions, namely Papilio erithonioides, 
P. grosesmithi (Rare), P. morondavana (Vulnerable) and Atrophaneura antenor. 
The last three species are traded commercially and require monitoring. Rain forest 
endemics of the eastern side include Graphium endochus (uncommon), Papilio 
oribazus, P. delalandei and P. mangoura (Rare). The other two endemics, 
Graphium evombar and G. cyrnus are widespread and common (19, 20). A number 
of reserves and other protected areas may ensure the perpetuation of these and other 
Malagasy endemics (see species accounts in section 6), but there is international 
concern at the alarming rate of degradation of Madagascar's vegetation and soils. 

Conclusion 

These notes on the swallowtail faunas of the world are necessarily brief. It is 
recognized that far more detail would be needed precisely to identify centres of 
species richness and importance within a country, and to plan a system of protected 
areas around those centres. These few examples must serve to indicate the potential 
conservation value of investigating swallowtails and other insect groups. Swallow- 
tails not only require species-oriented conservation action, but can also act as 
indicators of general threats, and as pointers for delimitation of biogeographical 
boundaries for planning the conservation of whole biotopes, their faunas and floras. 
Addition of other groups of butterflies, insects or invertebrates (e.g. molluscs) will 
strengthen and diversify the analysis, particularly when the data are synthesized with 
a consideration of vertebrates and angiosperms. Critical fauna analyses will serve to 
direct national and international attention to faunistically important countries, states 
and provinces, but local knowledge is the basis for more detailed conservation 
planning. It is axiomatic that baseline data for conservation can only be obtained by 
the encouragement and completion of field research. 

References 

1. Ackery, P.R. and Vane-Wright, R.I. (1984). Milkweed Butterflies: Their Cladistics and 
Biology. British Museum (Natural History), London, and Cornell University Press. 
448 pp. 

152 



Threatened Swallowtail Butterflies of the World 

2. Brown, K.S., Jr. (1975). Geographical patterns of evolution in Neotropical Lepidoptera. 
Systematics and derivation of known and new Heliconiini (Nymphalidae: Nymphalinae). 
Journal of Entomology 44: 201-242. 

3. Brown, K.S., Jr. (1982). Historical and ecological factors in the biogeography of 
aposematic neotropical butterflies. American Zoologist 22: 453-471. / 

4. Brown, K.S., Jr. (1977). Geographical patterns of evolution in neotropical Lepidoptera: 
Differentiation of the species of Melinaea and Mechanitis (Nymphalidae, Ithomiinae). 
Systematic Entomology 2: 161-197. 

5. Brown, K.S., Jr. and Ab'Saber N.A. (1979). Ice-age forest refuges and evolution in the 
neotropics: Correlation of paleoclimatological, geomorphological and pedological data 
with modern biological endemism. Paleoclimas 5: 1-30. 

6. Dalquest, W.W. (1948). Mammals of Washington. University of Kansas Natural History 
Museum, Lawrence. 444 pp. 

7. Dasmann, R.F. (1972). Towards a system for classifying natural regions of the world and 
their representation by national parks and reserves. Biological Conservation 4: 247-255. 

8. Dasmann, R.F. (1973). A system for defining and classifying natural regions for purposes 
of conservation. lUCN Occasional Paper No. 7: 47 pp. 

9. Dasmann, R.F. (1974). Biotic provinces of the world. lUCN Occasional Paper No. 9: 57 
pp. 

10. Dice, L.R. (1943). The Biotic Provinces of North America. University of Michigan Press, 
Ann Arbor. 78 pp. 

11. Erwin, T.L. (1982). Tropical forests: their richness in Coleoptera and other arthropod 
species. The Coleopterisis' Bulletin 36: 74-75. 

12. Haugum, J., Ebner, J. and Racheli, T. (1980). The Papilionidae of Celebes (Sulawesi). 
Lepidoptera Group of 1968 Supplement 9: 21 pp, 1 map, 2 pi. 

13. Holloway, J.D. (1980). Insect surveys — an approach to environmental monitoring. Atti 
XII Congresso Naz. Ital. Ent. 12. 

14. Holloway, J.D. and Jardine, N. (1968). Two approaches to zoogeography: a study based 
on the distributions of butterflies, birds and bats in the Indo-Australian area. Proceedings 
of the Linnean Society of London 179: 153-188. 

15. lUCN/UNEP/WWF. (1980). World Conservation Strategy. lUCN, Gland. 

16. Lamas, G. (1982). A preliminary zoogeographical division of Peru, based on butterfly 
distributions (Lepidoptera, Papilionoidea). In Prance, G.T. (ed.) Biological Diversifi- 
cation in the Tropics. Proceedings of the Fifth International Symposium of the Association 
for Tropical Biology. Columbia University Press, New York. 714 pp. 

17. Mackinnon, J. and Wind, J. (1980). Birds of Indonesia. FAO, Bogor. xii + 55 pp. 

18. Parker, S.P. (1982). Synopsis and Classification of Living Organisms. McGraw Hill, New 
York. 2 vols, 1232 pp. 

19. Paulian, R. (1951). Papillons Communs de Madagascar. L'Institut de Recherche 
Scientifique, Tananarive-Tsimbazaza. 90 pp. 

20. Paulian, R. and Viette, P. (1968). Insectes Lepidopteres Papilionidae. Faune de 
Madagascar 27, 97 pp, 19 pi. ORSTOM, CNRS, Paris. 

21. Pollisco, F.S. (1982). An analysis of the national park system in the Philippines. 
Likas-Yaman 3(12): 56 pp. 

22. Pyle, R.M. (1976). The eco-geographic basis for Lepidoptera conservation. Unpublished 
Ph.D. thesis, Yale University, New Haven, Connecticut 369 pp. 

23. Pyle, R.M. (1980(82)). Butterfly eco-geography and biological conservation in Washing- 
ton. Atala 8(1): 1-26. 

24. Smiet, F. (1982). Threats to the Spice Islands. Oryx 16: 323-328. 

25. Straatman, R. (1968). On the biology of some species of Papihonidae from the island of 
Celebes (East-Indonesia). Entomologische Berichten 28: 229-233. 

26. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands. Vol. I 
Papihonidae. (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 

27. UNDP/FAO National Parks Development Project (1981/1982). National Conservation 
Plan for Indonesia. Vols. 1-8. FAO, Bogor. 

28. Udvardy, M.D.F. (1975). A classification of the biogeographical provinces of the world. 
lUCN Occasional Paper No. 18: 48 pp. 

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Threatened Swallowtail Butterflies of the World 

29. Wallace, A.R. (1876). The Geographical Distribution of Animals. Macmillan, London. 2 
vols. 

30. Wells, S.M., Pyle, R.M. and Collins, N.M. (1983). The lUCN Invertebrate Red Data Book. 
lUCN, Gland and Cambridge, 632 pp. 



154 



5 



Trade in swallowtail butterflies 



The worldwide trade in butterflies is big business, running into tens of millions of 
dollars annually. The precise amount is controversial; a recent estimate of US $10-20 
million per year (21) must be conservative since there have been reports that the 
export trade from Taiwan alone is $20-30 million (16, 17). Butterflies can be bought 
from hundreds of dealers throughout the world but particularly in Hong Kong, 
Japan, Korea, Taiwan, Malaysia, western Europe, the U.S.A., Brazil and Peru. In 
this section we examine the main areas of commercial interest and ask whether the 
trade is likely to damage wild populations in a severe or permanent way. All prices 
quoted are in U.S. dollars. 

Introduction 

Butterfly collecting began in the 16th and 17th centuries when large numbers of new 
and exotic species were brought back by explorers to be described, classified and 
studied by the scientists of the day. It evolved into a popular hobby during the 19th 
century, pursued by educated aristocrats, middle-class citizens, doctors and cler- 
gymen who had the necessary money and time to spare. Some of these became 
leading authorities on the subject. Large collections were built up, especially by 
entomologists such as Lord Rothschild who were wealthy enough to send expeditions 
to remote areas. 

The current trade supplies dead specimens to scientists, museums and private 
collectors, eliminating their need to travel long distances. The number of commercial 
dealers has risen dramatically over the last three decades (20) and dealers' lists 
advertise a huge range of butterflies and other insects. During the past five years over 
80 per cent of the world's swallowtail species have been made available through 
international trading organizations, but less than 10 per cent are known to be farmed 
or ranched (see Table 5.1). Prices range very widely; cheaper species include the 
common Oriental and Australasian Graphium sarpedon (Blue Triangle or Common 
Bluebottle butterfly) and the Brazilian Eurytides stenodesmus at $0.30 or less. At the 
other end of the scale are rare birdwings like Omithoptera alexandrae (Queen 
Alexandra's Birdwing) an imperfect male specimen of which was recently advertised 
for $2850, and Omithoptera meridionalis and Omithoptera paradisea (Paradise 
Birdwing), specimens of which have been advertised for $7000! All three are 
restricted to New Guinea where they are protected by law. O. paradisea is far more 
plentiful than O. meridionalis and O. alexandrae, but certain rare subspecies are 
highly valued. It is usual for prices to be correlated with rarity, but this is not always 
clear. Surprisingly, some specimens of Zerynthia polyxena (Southern Festoon) from 
the Alps of Haute-Provence, France, were offered between 1978 and 1984 for the 
equivalent of about $2500 -$4000 (1, 11). The high price may indicate that the 

155 



Threatened Swallowtail Butterflies of the World 

specimens were of a very rare form since this species is widely distributed and rarely 
advertised for more than $2. Up until the 1970s the highest price paid was for a male 
Ornithoptera allottei (Abbe Allottes Birdwing) from Bougainville which was sold for 
the equivalent of $1500 at an auction in Paris in 1966 (33). O. allottei is only known 
from about twelve specimens and is thought to be a natural hybrid between O. 
victoriae regis (Queen Victoria's Birdwing) and O. priamus urvillianus (Priam's 
Birdwing) (8). 

High prices may also be paid for species which are fairly widely distributed but rare 
and difficult to obtain. In the foothills of the Himalaya mountains Teinopalpus 
imperialis (Kaiser-I-Hind) is difficult to capture since it is a strong flyer which keeps to 
the tree tops. In 1955 a collector paid the equivalent of about $250 for a single 
specimen (33) and the species now costs up to $75 a pair. Species from inaccessible 
countries rarely appear on the market and when they do they tend to have high price 
tags. For example, a pair of Bhutanitis thaidina from China was advertised by a West 
German dealer in 1983 for the equivalent of about $225. 

The table at the end of this section lists all swallowtail species and shows which of 
these have legislation protecting them, which are reared in captivity and which are 
currently in trade . The range of prices demanded for a particular species is shown by a 
series of price categories. In addition to species the list also includes several rare 
subspecies that command much higher prices than the nominate form. The 
information used in the table has been obtained from worldwide lists and correspond- 
ence between 1980 and 1985. Higher prices are often asked for female specimens 
because they are less easily attracted to baits and decoys. For example female 
specimens of Graphium latreillanus are offered at prices much greater than those for 
males, accounting for the wide range of prices asked for this species. The same 
consideration applies particularly to females of Papilio antimachus and P. zalmoxis 
which are seldom collected. Space in the table does not allow a full consideration of 
the prices of males and females separately. Where threatened species appear in trade 
they normally command very high prices, as do most rare species or subspecies. 

Some collectors will go to great lengths to obtain rarities, either for their own 
collections or for sale at a profit. The high prices paid for butterflies in recent years 
have attracted financiers interested only in investment, with little thought for the 
insects in the wild (5). The largest and one of the rarest butterflies in the world, 
Ornithoptera alexandrae of Papua New Guinea, is mainly threatened by defores- 
tation but collecting was at one time also a problem. Since 1968 it has been illegal to 
collect this species but until recently one smuggler was reputedly earning as much as 
$800 per specimen (16). With Ornithoptera allottei now believed to be a hybrid, 
Ornithoptera alexandrae has probably replaced it as the world's most valuable species 
(although specimens of Ornithoptera meridionalis and Ornithoptera paradisea have 
been advertised at higher prices). In 1979 it was estimated that fine specimens of 
Ornithoptera alexandrae were worth up to $2000 (7, 31), but this has since been 
exceeded. Commercial collecting by expatriates has now been stopped completely in 
Papua New Guinea and several people have been deported for smuggHng (16). 
Ornithoptera alexandrae and some other rare, valuable and coveted species are 
seriously endangered, mainly by habitat loss rather than trade alone (see section 6). 
They include Papilio hospiton (Corsican Swallowtail) from Corsica and Sardinia, 
Papilio homerus (Homerus Swallowtail) from Jamaica, a female of which was 
advertised in the U.S.A. in 1984 for $2800, and the uncommon Malagasy species 
Papilio morondavana, females of which are frequently advertised by French dealers 
for about $120 -$150. 

An important development in the last decade, particularly in the United Kingdom, 

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Trade 

has been the rise of the "butterfly house". These enterprises have concentrated in 
providing a display of butterflies, particularly exotic species, in "artificial jungles", 
that is, tropical conditions with appropriate vegetation and climate. Butterfly houses 
need a constant supply of living specimens for display. Although some butterflies are 
reared in the houses, most are obtained from their countries of origin as pupae or 
adults, using the fast, reliable airfreight services now available from many parts of the 
tropics. Good prices are paid for butterflies used in living displays especially when the 
supply is regular and reliable. Swallowtails are probably in demand because of their 
size, bright colours and exotic appeal. The main exporting countries are in the tropics 
where there is a diverse range of beautiful species. Farmed (i.e. bred from parental 
stock held in captivity) or ranched (i.e. conceived by wild parents but reared from 
young stages under controlled conditions) specimens of certain species are available , 
but most material appears to be caught in the wild. 
To summarize, there are essentially three different sorts of trade :- 

(i) Low value , high volume . Large numbers of common species , often of low 
quality are used in a range of ornaments and decorations, less as 
specimens. The trade of Taiwan is typical of this category, with annual 
sales estimated at 15-500 million butterflies. 

(ii) High value, low volume. High quality specimens, sometimes with 
scientific data including date and place of capture, are sold to museums, 
students and collectors. Many dealers in Europe, North America and 
Japan, produce catalogues of specimens in this category. Trade from the 
insect ranching programme in Papua New Guinea is at the top end of this 
market (15, 24). Linking (i) and (ii) is the trade in high value, high quality 
ornamental items. This includes wall mounts and glass domes containing 
mounted butterflies in life-like settings, or jewellery such as that based on 
the iridescent Morpho menelaus (Blue Morpho) of Brazil. 

(iii) The live trade. Fairly low but continuous volume, medium value. Living 
butterflies, usually pupae or adults, are despatched to butterfly houses 
for display to the paying public. This trade is probably the fastest growing 
of the three types. 

Countries of origin 

The Orient The Oriental region contains an extremely diverse butterfly 
fauna (see sections 3 and 4). The major threat in the region is loss of habitat resulting 
from deforestation, agriculture and urbanization. In most areas collecting is 
small-scale, often by Japanese collectors, but there are large-scale international 
trading centres in Taiwan and Malaysia, and significant commercial enterprises in 
Hong Kong and Korea (26). 

Taiwan has a butterfly trade which operates on a vast scale involving about 20 000 
people including 10 000 collectors (11, 16, 30). Estimates of the number of butterflies 
traded vary from 15 to 500 million (26). Virtually all butterflies are caught in the wild 
and sold to the 30 or so factories which process them (19, 30). Many species are 
imported in bulk from overseas, particularly Morpho species from Brazil, Charaxes 
from Africa and birdwings from South East Asia. In most cases the bodies are 
discarded for pig feed and the wings are glued to paper bodies with bristle antennae 
and laminated between sheets of clear plastic. They are then used to make 
bookmarkers, coasters, table mats, wall decorations and even plastic toilet seats! 
Others are made into pictures or used to decorate hand-bags or purses. Taiwan's 

157 



Threatened Swallowtail Butterflies of the World 

export trade was valued at $30 million in 1969 (16) and about $21 million in 1976 (17). 

Individual specimens are also sold by a number of dealers in Taiwan, where prices 
vary according to rarity, size and beauty, from about $0.02 for a specimen of the 
endemic Papilio machaon sylvina to $50 for the endemic Papilio maraho. Males of P. 
maraho, now recognized as a Vulnerable species, were advertised in 1983 by a 
German dealer for the equivalent of about $75. Although an island with an area of 
only 35 960 sq. km, Taiwan has about 400 butterfly species of which 40 are endemic 
(19). About 100 species, including 20 crop pests, need no protection, while the 
remainder may need some form of management ( 19) . Between 50 and 60 of these are 
very rare (19) and the Entomological Suppliers Association of Great Britain has 
banned trade in the rare Taiwan endemic Troides aeacus kaguya (14). There are no 
captive-breeding programmes though a few people rear rare papilionids and Attacus 
atlas (Atlas moth) to obtain quality specimens for collectors (19). There is no 
conclusive evidence that the butterfly industry poses a threat to most butterfly 
species, but there is now serious concern for Papilio maraho. The major threats to 
Taiwan's butterflies undoubtedly come from habitat destruction for agriculture and 
forestry. The lowland forests are virtually gone and human population pressure is 
extending deforestation into more mountainous areas. In densely populated areas 
urbanization, air pollution and pesticides are taking their toll. There is apparently no 
environmental legislation in Taiwan, although there is growing conservation 
concern. Four national parks have been designated, but there are so far no laws to 
ensure rational exploitation of butterflies and the long-term survival of the industry 
(11, 16, 26, 28, 30). The import ofspecimens to Taiwan from overseas has never been 
measured and it is not known what effect the market has in the countries of origin. 
More research is needed to assess what proportion of the Taiwan trade is in foreign 
species, and the impact caused by the demand. 

Large numbers of Trogonoptera brookiana (Rajah Brooke's Birdwing) are 
exported from Malaysia (23). There is evidence that the trade exceeds 125 000 
specimens annually, all of which are apparently collected in the wild. Under 
Malaysian law the species is "protected" but not "totally protected" , and collecting is 
permissible with a permit which is easily obtained. The species is also listed on 
Appendix II of CITES (Convention on International Trade in Endangered Species of 
Wild Fauna and Flora). This Appendix implies that commercial trading is allowed 
providing a permit from the country of export is obtained . This can provide a method 
of monitoring trade levels but there is evidence that most exports of T. brookiana are 
unlicensed. Many dealers list Trogonoptera brookiana for sale in bulk quantities of up 
to 1000 specimens and large numbers have recently been used for artwork (20). In 
1983 batches of 100 were on offer by a West German dealer for about $20 and in the 
U.S.A. a dealer was selling 100 for $30. There have been recent suggestions that the 
species is being ranched or farmed, but the evidence is circumstantial. Certain dealers 
in Peninsular Malaysia sell a large proportion of specimens that are in perfect 
condition, suggesting that they are reared from pupae. The breeding localities, 
foodplants and habits of the young stages are being kept a closely-guarded secret (9) . 
The main threat to Trogonoptera brookiana and other Malaysian butterflies is habitat 
destruction rather than trade. Nevertheless, the trade in this species should be 
carefully monitored since the population may be brought to such a low level by loss of 
habitat that a continuation of heavy collecting could be a serious threat (23). 

There is little information on trade in the Philippines and Indonesia although 
collecting by tourists, private collectors and local people on behalf of dealers is said to 
be affecting some species, including the Endangered Papilio chikaem Luzon (29, also 
see p. 364). Indonesia has legislation protecting certain species, mainly birdwings, 

158 



Trade 

but it is not known how effective this has been. Some of the species are often 
advertised in dealers' lists. 

Butterfly trading based in India and Indochina is now quite extensive and occurs at 
all levels, from personal collectors to substantial businesses. In South Korea the 
decline in populations of species such as Parnassius bremeri, Papilio demetrius, P. 
machaon and P. macilentus has been partly attributed to over-collection (18). There 
is little information from Thailand, but at least one Bangkok business uses local 
collectors to provide material for the international market. In the mountains around 
Kathmandu in Nepal collecting at times reaches absurd proportions. There are 
reports from Phulchoki Mountain that encampments of foreign entomologists wait 
for hill-topping males of Teinopalpus imperialis and kill every specimen that flies in. 
The Nepal government requires a permit for collecting, but the legislation seems to 
be openly abused. There is also a substantial market in dead specimens offered by 
street hawkers in Kathmandu (12). In 1980, the Government of India passed an 
amendment to the 1972 Wildlife Protection Act listing a large number of butterflies 
as fully protected. This certainly helps to draw attention to the need for restraint in 
collecting certain species, but the law is very difficult to enforce. Some foreign 
tourists visit northern India specifically to collect butterflies known to be fully 
protected. Capturing material for personal study would no doubt be acceptable, but 
there are fears that large numbers of specimens find their way to the market-place. 
There are two large commercial "farms" in the country which have apparently been 
affected by this legislation but it is thought that neither had a captive breeding 
programme. The Indian Government is interested in setting up a farming project for 
common species and two trial farms are planned for north-western and eastern 
India. 

China has one of the world's most important swallowtail faunas, but because of its 
relative inaccessibility in recent years many species still remain poorly known. 
However, during the 1980s Japanese entomologists have travelled in the rich 
collecting areas of Yunnan and Sichuan, bringing back little-known species such as 
Bhutanitis mansfieldi (see p. 192). There is also evidence that local Chinese have 
been trained to collect specimens and are regularly sending material to Japan. Large 
numbers of Bhutanitis thaidina have been traded in this way (see p. 194). The export 
of material from China for academic study is greatly to be encouraged, but the 
build-up of a large commercial operation should be viewed with caution. Many high 
altitude species are very local and have short flight seasons; they may be vulnerable 
to over-exploitation. Monitoring of the trade would be advisable. There are oppor- 
tunities for summer butterfly ranching programmes in western China, worthy of 
investigation as a source of butterflies for dealers, a seasonal income for local people 
and a means of protection for the butterflies. 

Australasia There is relatively little commercial trade in butterflies in 
Australia, partly because of the low numbers of amateur enthusiasts in that country. 
The few commercial enterprises in Australia have perhaps up to 80 per cent of their 
trade in butterflies originating from outside the country. Endemic swallowtails of 
particular interest, such as Graphium macleayanum and Protographium leosthenes, 
have been intermittently in trade but have not been offered for some years. The 
endemic birdwings, Ornithoptera richmondia, O. euphorion and O. priamus prono- 
miis, are in demand and also in trade, but not in large numbers. These species, 
together with the endemic subspecies of the Blue Mountain Butterfly, Papilio idysses 
joesa, are protected in Queensland; however, the main threat to all these butterflies 
is destruction of the rain forest habitat rather than commercial collecting. Other 

159 



Threatened Swallowtail Butterflies of the World 

Australian swallowtails are occasionally offered, including Cressida cressida, 
commonly known by the extraordinary name Big Greasy Butterfly because of the 
way it rapidly loses its wing-scales, producing a greasy effect. The Australian 
papilionid fauna is a small one and the commercial trade is generally low in volume 
and high in quality and value. 

Papua New Guinea probably runs the best example of a high value, low volume 
trade, exporting high quality specimens of native butterflies. The spectacular 
birdwings of the genus Ornithoptera are the greatest attraction for collectors and an 
important natural resource for the country. Mainly because of habitat destruction, 
but possibly aided by collecting, some species have become very rare, in particular 
Ornithoptera alexandrae. In 1968, the seven rarest Ornithoptera species, i.e. 
alexandrae, allottei, chimaera (Chimaera Birdwing), go/Za//!, meridionalis , paradisea 
and victoriae were protected by a national law to prevent collecting (21). 

Before Papua New Guinea's independence in 1975 all collecting was carried out for 
expatriate dealers by native collectors. They often received as little as $0.20 for 
butterflies which could fetch up to $250 overseas (7). A few years ago a butterfly 
dealer visited the island of Nimoa south-east of Papua New Guinea and persuaded 
villagers to collect hundreds of adults and chrysalids of the spectacular birdwing 
Ornithoptera priamus caelestis, endemic to the Louisiade Archipelago (8). The 
villagers were paid trifling sums for a haul of several hundred butterflies worth about 
$5000 (26) . Not only were the villagers exploited , but no more birdwings were seen in 
the vicinity for many years. Eventually they were reintroduced by the Insect Farming 
and Trading Agency (I.F.T.A.) from a population on the other side of the island 
and are now in trade (26, 27). Nowadays, only nationals are permitted to profit 
from the trade, which is co-ordinated by I.F.T.A. at Bulolo. Villagers are encouraged 
to ranch or, to a lesser extent, collect the commoner unprotected species such as 
Ornithoptera priamus and Troides oblongomaculatus . By 1978 over 500 people were 
involved in this rapidly expanding village industry. All business is handled by 
I.F.T.A. which pays the villagers all profits less 25 per cent. There has been an 
increase in the trade in quality specimens from Papua New Guinea. In 1979 villagers 
were being paid $37 for a box of butterflies but by 1980 this had risen to $50 (21). A 
hard-working butterfly rancher may earn up to $1200 annually (21). By 1983 the 
annual trade in insects, mainly butterflies was estimated to be $110 000 and 
increasing. Approximately 30 per cent of all butterflies reaching the Agency are 
ranched and the rest are collected as adults in the wild, but 50 per cent of the revenue 
comes from the usually better quality ranched specimens (21). Swallowtails are more 
commonly ranched than butterflies of other groups and so the proportion of ranched 
specimens is higher than 30 per cent of the total. 

Ranching is widely encouraged by the I.F.T.A. because it produces fresh 
undamaged specimens and the habitat enrichment is also a good conservation 
measure. Under this practice usually only the pupae are kept in captivity. Wild 
butterflies are free to come and go and the rancher ensures that a proportion of those 
reared in captivity is released to continue the cycle. Children and potential ranchers 
are shown which species can easily be ranched, and which plants are attractive as food 
or for nectar. An average ranch will have a garden of about 0.2 ha surrounded by a 
thick hedge of Hibiscus, Bougainvillea or similar nectar-bearing plants. These attract 
the butterflies but also keep out livestock. Foodplants cultivated inside the fence 
include Aristolochia tagala (Dutchman's Pipe Vine) for the common birdwings, and 
Evodia spp. for Papilio ulysses (27). The pupae are removed and usually placed into 
an emergence cage just before the adults emerge. The ranchers are taught how to 
judge the correct time and also how to kill and pack into boxes the newly emerged 
butterflies (12, 15, 24, 27, 31). 

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Trade 

A large number of other Papuan butterflies are collected as adults but these tend to 
be mainly common species such as Graphium sarpedon and G. agamemnon (Tailed 
Jay or Green-spotted Triangle butterfly). Adult Graphium weiskei (Purple-spotted 
swallowtails) are collected only as adults because the food plant and larval biology are 
unknown. This species may be common in some areas but occasionally collection has 
to be halted for anything up to a year whilst the numbers increase (27). Ornithoptera 
victoriae is common on Bougainville and the I.F.T.A. may in future recommend its 
removal from the protected list in order to allow ranching. O. chimaera and O. goliath 
have also been suggested as good candidates for future ranching projects although 
their foodplants are more difficult to cultivate. As well as being a valuable village 
industry in Papua New Guinea, the butterfly ranching project demonstrates that 
butterfly trade and conservation can be of mutual benefit (27) and the future could be 
promising for both the trade and the ranching programme. 

Africa and Madagascar Large numbers of African Charaxes (Nymphali- 
dae), Papilio and other genera appear on dealers' lists. Many are only listed as being 
from Africa, West Africa or East Africa, but the Central African Republic, 
Madagascar, Malawi and to a lesser extent Congo and Gabon seem to be regular 
suppliers. Malawi is chiefly a source of Charaxes species, whereas both the Central 
African Republic and Madagascar appear to be suppliers of a wide range of 
butterflies. 

Reports from the Central African Republic indicate that every year for the past 
fifteen years hundreds of thousands of butterflies have been indiscriminately caught 
by hundreds of local collectors for sale overseas. Collecting of this sort could have an 
effect on some species or populations. More information is needed from this and 
other African countries. Some nations would benefit from a farming or ranching 
scheme, particularly for Papilio and Charaxes species. 

Madagascar is well known for its unique and endemic fauna, including many 
butterflies. There is no commercial ranching and specimens are collected by local 
collectors for expatriate dealers. The collectors are usually untrained and are only 
interested in large numbers of different species. The resulting indiscriminate 
collecting may be highly destructive and wasteful since many specimens are damaged 
and later discarded. Female Papilio morondavana, a large black and yellow 
Vulnerable swallowtail, can command prices up to $150 (see p. 000). Other valuable 
Malagasy species are the two Rare species Papilio grosesmithi (see p. 000) and P. 
mangoura (see p. 000), and the more common Graphium evombar. Specimens from 
Madagascar are subject to a heavy export tax but the fact that some species are 
common and cheap on many dealers' lists may indicate large-scale smuggling. 

The African species commanding the highest prices are often large and spectacu- 
lar, such as Papilio antimachus (African Giant Swallowtail, $20 for a male in 1983 in 
the USA) and Papilio zalmoxis (Great Blue Papilio, $10-15 in 1983 in the USA). 
Others are very scarce, such as Papilio sjoestedti (Kilimanjaro Swallowtail, Rare, see 
p. 000) from Tanzania, which has been commanding particularly high prices ($212 for 
a female on one American list in 1984), or Papilio leucotaenia, (Cream-banded 
Swallowtail, Vulnerable, see p. 000) which is restricted to relict forests in Central 
Africa. Collecting is normally done by local people for overseas dealers. In Africa 
there is no legislation preventing the collecting of butterflies except in Kenya where a 
permit is required to collect Lepidoptera and Coleoptera, and the Province of the 
Cape of Good Hope, South Africa, where 17 Lepidoptera are protected, none of 
them swallowtails. 

South and Central America Commercial trade in the Caribbean Islands is 

161 



Threatened Swallowtail Butterflies of the World 

very variable in extent. There are many endemic taxa of swallowtails in the 
Caribbean, particularly endemic subspecies, but on the whole these have been little 
exploited. Some of the commercial collecting in Jamaica has caused unease, 
particularly the exploitation of the endemic, and very valuable, Papilio homerus 
(p. 297). One commercial enterprise is established in the Dominican Republic, but 
most trading activity in the Caribbean Islands seems to be by visiting dealers from 
elsewhere, particularly the U.S.A. Apart from the trade in swallowtails from Jamaica 
and Hispaniola, highly-priced specimens of Battus devilliers (from the Bahamas) 
have recently been offered and other species from various parts of the Caribbean are 
advertised from time to time. On the whole, however, the volume of trade is low, is 
concentrated on the high value end of the market, and is not to be compared with the 
volume of trade in butterflies from Peru, Brazil or even Mexico. 

Butterfly "farms" exist in Costa Rica and some species of Morpho (Morphidae) 
and some swallowtails are ranched in Brazil (21, 25, 32). The beautiful iridescent 
blue Morpho butterflies are much in demand for high quahty jewellery and other 
decorative purposes and 50 million a year are used in Brazil alone (6, 26). Although 
the law requires that morphos in trade must have been farmed, there is evidence that 
farms or ranches only meet part of the trade and many specimens continue to be 
caught in the wild (21). Survey work is needed to assess the impact of this trade. In 
Brazil one butterfly, Parides ascanius, is completely protected by law and another, 
Eurytides lysithous harrisianus , has been proposed for listing. In Honduras there is a 
low value, high volume trade similar to that in Taiwan, but on a smaller scale (26). In 
Peru there is a considerable insect industry, with large numbers of swallowtails 
coming out of the classic Tingo Maria and Rio Satipo regions as well as other areas. A 
major butterfly collecting company reportedly operates in Colombia (20). Mexico 
has a large export trade in butterflies, all of which is required to be licensed under 
Mexican legislation. Butterflies originating in Bolivia, Colombia, Costa Rica, 
Ecuador and Venezuela are also not uncommon on dealers' Hsts. 

North temperate regions In general the trade in temperate species is for 
serious collectors rather than the ornamental market. Butterflies of the Palearctic 
region are often in demand and although some farms exist, most species are collected 
as young stages or adults in the field. Eastern Palearctic species are particularly 
difficult to obtain and China has species of Papilio and Parnassius that would be 
suitable for farming or ranching. There is extensive European legislation to prevent 
and monitor butterfly collecting and trade. The EEC is party to CITES and thus 
monitors trade in birdwings and Parnassius apollo in all EEC countries. Some or all of 
the native Papilionidae are protected in Austria, Czechoslovakia, Finland, France, 
East Germany (DDR), Luxembourg, Netherlands, Poland, Switzerland, Great 
Britain and the U.S.S.R. (Lithuania). Turkey has a complete ban on collecting and 
export of butterflies (14). In the U.S.A. the Floridan subspecies Papilio aristodemus 
ponceanus (Schaus' Swallowtail) is currently proposed as endangered under the 1972 
Endangered Species Act, but Papilio andraemon bonhotei (Bahama Swallowtail) is 
proposed for delisting as a result of its improved circumstances (see p. 301). 

Butterfly trade and conservation 

The impact of private collectors The major threat to butterflies is loss of 
habitat caused by the actions of humans (2). Urbanization, deforestation, overgraz- 
ing, agricultural intensification, expansion of subsistence agriculture and atmos- 

162 



Trade 

pheric pollution all take their toll. Nevertheless, there are circumstances in which 
private collectors may have a serious impact, notably when populations are already 
severely reduced by other factors (22). 

Such was probably the fate of Lycaena dispar (the Large Copper, Lycaenidae) 
which became extinct in Britain, on the edge of its range, in 1847/8. Habitat 
destruction by drainage was the main cause of the demise, but collectors may have 
dealt the final blow. More recently, Canadian naturalists expressed concern over 
excessive collecting of butterflies in the Great Lakes area. Species that were already 
threatened by habitat loss and pollution became the target of collectors interested by 
their increased rarity value. As a result, Artogeia virginiensis (the West Virginia 
White, Pieridae) has been listed as endangered and protected in Ontario (2), but 
subsequent investigations indicate that the species may be more widespread than was 
at first realized (4). 

Even butterflies with a seemingly wide range may have vulnerable isolated 
populations with slight pattern variations coveted by collectors. In 1926 a collector 
tried to wipe out an entire local population of Parnassius apollo (Apollo butterfly) in 
the Italian Alps to increase the value of specimens already possessed (3). 

Recently some other examples of irresponsible exploitation of butterflies by 
collectors have come to light. In the Far East, Japanese collectors are particularly 
implicated. Package tours to collect butterflies have been accused of causing serious 
reductions in the butterfly populations of some areas of the Himalayas (10). In 
Luzon, Japanese collectors have been known to offer cameras for specimens of 
Papilio chikae, a beautiful iridescent swallowtail (9), and a female specimen on a 
West German dealer's list in 1983 was offered for the equivalent of about $150. It is 
now quite common for lepidopterists to travel to the tropics on collecting tours. 
Collecting a few specimens for a private collection can do no harm, but it seems to be 
common practice to collect large numbers of specimens for sale to dealers. The 
money obtained may be offset against the cost of the travel. Reports indicate that 
Japanese collectors frequently travel to Indonesia, the Philippines or other parts of 
Asia with the sole purpose of collecting large numbers of butterflies. The recent 
rediscovery of Atrophaneura palu in western Sulawesi has resulted in a number of 
highly-priced specimens appearing in the European trade and yet no data on the 
habitat or ecology of the species have been published for the benefit of the scientific 
community. 

There is a danger that collectors may be unable to recognize when they are 
depleting butterfly stocks below the threshold of recovery, particularly when they 
only visit the breeding areas for short periods of time or employ unsupervised 
assistants. Many species of butterfly can reproduce exponentially if conditions are 
right, but they may also decrease exponentially when conditions are bad. This means 
that a seemingly common species may very suddenly plummet into rarity or even 
extinction. 

The impact of commercial collectors Given suitable conditions and 
resources most butterflies are able to reproduce sufficiently to prevent permanent 
reduction of their populations by collectors. However, the evidence that millions of 
specimens are now involved in the international trade in ornaments is disturbing. In 
Taiwan the butterfly populations are reputedly remaining constant despite the 
enormous pressure on them (26), but there is also evidence that more material is 
imported to Taiwan from Africa and South America, and that the Taiwan trade is in 
decline. There is clearly a need for an independent assessment. Similarly the main 
collecting areas in South America and Africa require further research, not only to 

163 



Threatened Swallowtail Butterflies of the World 

protect the butterflies, but also to ensure long-term rational utilization of the 
resources that are the livelihood of many people. 

As a result of their limited size , island populations are often particularly vulnerable 
to excessive collecting as well as habitat destruction. For the giant birdwings of New 
Guinea, Indonesia and the Solomon Islands the problem is exacerbated because they 
generally lay no more than 30 eggs per generation. In such cases trade does need to be 
controlled and in Papua New Guinea the butterfly ranching programme is a good 
example of how this can be done successfully. 

There is some controversy over the effect of removing large numbers of males from 
butterfly populations. Males tend to be collected more than females because they are 
often brighter and more easily captured on hilltops or at baits of urine or rotten fruit. 
One male may be capable of fertilizing several females, but there is little scientific 
information on the degree of male depletion a population can withstand. In Brazil 
where 50 million male morpho butterflies enter the trade annually, a survey was 
undertaken to determine the proportion of males and females in nature (6). Results 
showed a ratio of ten or even several hundred males to every female in most species 
and it was concluded that the number of specimens taken by commercial collecting 
would not threaten populations of the species taken. These sex ratios have been 
widely disputed since the habits of females make them much more difficult to capture 
(13) and the sex ratio of captive-bred butterflies has not been shown to depart 
significantly from 1:1. More research is needed on this important aspect of butterfly 
reproduction and its implications to management. 

Collecting and legislation CITES lists the birdwing butterflies (Ornithop- 
tera, Trogonoptera, and Troides) and Parnassius apollo on Appendix II. Since these 
listings there has been a significant reduction in advertised trade of some species, 
although CITES Appendix II seeks only to regulate trade , not to prevent it. A total of 
87 countries are now party to CITES. 

Many examples of legislation have been cited in the above sections. National 
legislation or voluntary restrictive codes are increasingly necessary in regions where 
habitat destruction is so extensive that the relatively minor effects of collecting may 
have a serious impact (13). Such laws or codes should encourage scientific study for 
management purposes and can be used to encourage public awareness of butterfly 
conservation needs. In the U.S.A., the Endangered Species Act not only prevents 
collecting but also specifies a recovery programme and allows for the protection of 
habitat which is critical to the survival of the species. Legislation against collectors 
can only be successful if habitat is set aside to conserve wild populations. Emphasis 
should be put on protected areas as well as protected species. This course of action 
requires thorough survey work in order to delineate critical habitat, and expert 
ecological studies to ensure adequate long-term management. 

In countries with large areas of natural habitat, private collectors are most unlikely 
to have a significant impact on butterfly populations. Even when butterflies are 
protected within national parks there would generally be no harm in permitting 
amateur collecting on a small scale. It is particularly important that future 
generations of entomologists are not dissuaded from their studies by unnecessarily 
severe restrictions. 

Commercial collecting has not been independently assessed in any of the main 
tropical centres of origin. There are insufficient data to judge whether monitoring or 
regulation of the trade is warranted. Research into the origin and extent of world 
trade in ornamental butterflies is urgently required. 

Conclusions Private collecting of butterflies can be an instructive hobby 

164 



Trade 

and is important for research into ecology, population dynamics, genetics and 
taxonomy. It brings a great deal of pleasure to many people and does not usually 
threaten butterfly species. However, in a small number of cases irresponsible 
over-collection can cause a permanent decline. The latter is particularly true if the 
species has a very small range, has naturally low populations and low reproductive 
rate, or has already been severely reduced by other impacts. 

Commercial collecting can be an important source of income and should not be 
dismissed as necessarily harmful. If populations are harvested in a sustainable 
manner, then both conservation and commercial interests can be satisfied. Habitat 
destruction is the main cause of decHne in butterfly populations, but there is a danger 
that commercial collecting levels that were sustainable in the past may become 
damaging in the ever-decreasing areas of suitable habitat. Such may soon be the case 
for Trogonoptera brookiana, which lives only in the primary lowland forests of 
Malaysia and Indonesia. Responsible commercial collectors should take all possible 
steps to conserve the habitats in which they collect. This is clearly essential for both 
the trade and the species. Where possible, captive breeding should be encouraged, to 
provide high quality specimens for the trade, to provide local employment and to 
ease the pressure on wild populations. At present , less than 10 per cent of swallowtail 
species are known to be farmed or ranched (see Table 5.1). Commercial organiz- 
ations should invest in research to raise this figure. 

Legislation against collecting and trade is unlikely to preserve a species unless 
parallel measures to protect its habitat are also enforced. Preliminary assessments 
are needed in order to decide whether extensive monitoring of international butterfly 
commerce is advisable. 



References 

1. Anon. (1978). Extract from Nice Matin in litt. 

2. Anon. (1982). Save the butterflies. Nature Canada 11(4): 46. 

3. Bourgogne, J. (1971). Un temoignage de plus sur la destruction de la nature (Papilioni- ■ 
dae). Alexanor 7: 50. 

4. Brownell, V.R. (1981). The West Virginia white butterfly (Artogeia virginiensis Edwards) in 
Canada: a status report. Nongame Program, Wildlife Branch, Ontario Ministry of Natural 
Resources. 55 pp + appendices. 

5. Campbell, G. (1976). Why not try a flutter on butterflies? Daily Telegraph 13 March. 

6. Carvalho, J. CM., and Mielke, O.H.M. (1972). The trade of butterfly wings in Brazil and 
its effects upon survival of the species. Proceedings 19th International Congress of 
Entomology (Moscow) 1 : 486-488. 

7. Cherfas, J. (1979). How to raise protection money. New Scientist 6 December. 

8. D'Abrera, B. (1975). Birdwing Butterflies of the World. Lansdowne Press, Melbourne. 260 
pp. 

9. D'Abrera, B. (1982). Butterflies of the Oriental Region. Part L Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi + 244 pp. 

10. Dasgupta, M.A. (1979). Save our butterflies. Letter to the Editor. Illustrated Weekly of 
India 23 September. 

11. Donahue, J. P. (1984). In litt. ,24 May. 

12. Feeny, P. (1984). Pers. comm., 1 August. 

13. Heath, J. (1981). Insect conservation in Great Britain (including "A code for insect 
collecting". Beiheft Veroffentlichung Naturschutz Landschaftspflege Baden-Wiirttemberg 
21: 219-223. 

14. Heath, J. (1981). Threatened Rhopalocera (Butterflies) in Europe. Nature and Environ- 
ment series (Council of Europe) no. 23, 157 pp. 

15. Hutton, A.F. (1978). Conservation and utilization of the insect resources of Papua New 
Guinea. Unpublished report. 

165 



Threatened Swallowtail Butterflies of the World 

16. Inskipp, T. and Wells, S. (1979). International Trade in Wildlife. Earthscan, London. 104 
pp. 

17. Jackman, B. (1976). Bye-bye birdwing. Sunday Times 12 September. 

18. Kyu, K.H. (1982). Statussurvey of Alpine butterfly fauna. South Korea. WWF Yearbook 
1982. World Wildlife Fund, Gland, Switzerland, 492 pp. 

19. Marshall, A.G. (1982). The butterfly industry of Taiwan. Antenna 6: 203-4. 

20. Nagano, CD. (1984). The International Trade in Butterflies. Unpublished manuscript. 27 
pp. 

21. National Research Council (1983). Butterfly Farming in Papua New Guinea. Managing 
Tropical Animal Resources Series. National Academy Press, Washington, D.C. 

22. Owen, D.F. (1974). Trade threats to butterflies. Oryx June. 

23. Owen, D.F. (1976). Rajah Brooke's birdwing. Oryx 13: 259-261. 

24. Parsons, M. (undated). Insect Farming and Trading Agency Farming Manual. Insect 
Farming and Trading Agency, Bulolo, P.N.G. 

25. Pyle, R.M. (1979). How to conserve insects — for fun and necessity. Terra 17(4): 18-22. 

26. Pyle, R.M. (1981). Butterflies: now you see them. International Wildlife 11(1): 4-11. 

27. Pyle, R.M. and Hughes, S.A. (1978). Conservation and utilisation of the insect resources 
of Papua New Guinea. Unpublished report to the Wildlife Branch, Dept. of Nature 
Resources, Independent State of Papua New Guinea. 157 pp. 

28. Severinghaus, S.R. (1977). The butterfly industry and butterfly conservation in Taiwan. 
Atala 5(2): 20-23. 

29. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands. Vol. I 
Papilionidae . (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 

30. Unno, K. (1974). Taiwan's butterfly industry. Wildlife 16: 356-359. 

31. Vietmeyer, N.D. (1979). Butterfly ranching is taking wing in Papua New Guinea. 
Smithsonian 10(2): 119-135. 

32. Wiltshire, E. P. (1959). First impressions of the tropical forests of southeastern Brazil and 
their Lepidoptera. Journal of the Lepidopterists' Society 13(2): 79-88. 

33. Wood, G.L. (1982). The Guinness Book of Animal Facts and Feats. Third Edition. 
Guinness Superlatives Ltd., Enfield. 250 pp. 



166 



Trade 



Table 5.1: Swallowtail butterflies in trade, 1980-1985 



Data in this table are drawn from a selection of trade literature acquired from dealers 
worldwide. We would like to be informed of other species known to be traded and of 
species already listed here but found to be traded at prices markedly different from 
those given. The numbers next to each species are the same as those used in the 
species list (section 3) and may be used for cross-reference. 



Key: 



lUCN 



conservation status categories (column CAT) 

E Endangered 

V Vulnerable 

R Rare 

I Indeterminate 

K Insufficiently Known 



Legislation (column LAW) 

Lists all species which are protected by law in part of their range. In addition, 
Parnassius apollo and all species of the three birdwing genera Trogonoptera, Troides 
and Ornithoptera are listed on appendix 2 of the Convention on International Trade 
in Endangered Species of Wild Fauna and Flora (CITES). 

Ranching or farming (column R/F) 

Shows traded specimens which are ranched, reared or farmed in captivity. Most will 

be ranched. 

Price categories (columns A-H) 

Exchange rates have fluctuated widely during the preparation of this table. The price 
equivalents in the various ranks are therefore very approximate and the table can 
only be used as a guide. Source data are held at the Conservation Monitoring Centre 
and are available to interested parties. 





UK 


USA 


GERMANY 


FRANCE 


JAPAN 


A. 


Up to £0.60 


Up to$l 


Up to DM3 


Up to Fr6 


Up to Y240 


B. 


£0.60-1.20 


$1-2 


DM3-6 


Fr6-12 


Y24O-480 


C. 


£1.20-5 


$2-7.50 


DM6-20 


Fr 12-50 


Y480-1800 


D. 


£5-20 


$7.50-30 


DM20-80 


Fr50-200 


Y 1800-7200 


E. 


£20-50 


$30-75 


DM80-200 


Fr200-500 


Y7200-18000 


F. 


£50-100 


$75-150 


DM200-400 


Fr500-1000 


Y 18000-36000 


G. 


£100-1000 


$150-1500 


DM400-4000 


FrlOOO- 10000 


Y36000-360000 


H. 


£1000+ 


$1500+ 


DM4000+ 


FrlOOOO+ 


Y360000+ 



167 



Threatened Swallowtail Butterflies of the World 
Table 5.1 



CAT LAW R/F 



ABCDEFGH 



Order: LEPIDOPTERA 
Family: PAPILIONIDAE 
Subfamily: BARONIINAE 

1 Baronia brevicornis 
Subfamily: PARNASSINAE 
Tribe: PARNASSIINI 

2 Archon apollinus 
Hypermnestra helios 



9 
10 
11 



ParnasS: 
Parnass, 
Parnass, 
Parnass, 
Parnass, 
Parnass, 
Parnass, 
Parnass, 

12 Parnass, 

13 Parnass, 

14 Parnass 

15 Parnass 

16 Parnass, 

17 Parnass 

18 Parnass 
Parnass 
Parnass 
Parnass 
Parnass 

23 Parnass 

24 Parnass, 

25 Parnass, 

26 Parnass, 

27 Parnass, 

28 Parnass, 

29 Parnass, 

30 Parnass, 

31 Parnass, 

32 Parnass, 

33 Parnass, 

34 Parnass, 

35 Parnass, 

36 Parnass, 

37 Parnass, 

38 Parnass, 

39 Parnass, 

40 Parnass, 



19 
20 
21 

22 



MS szechenyii 
us cephalus 
us maharaja 
us delphius 
us stoliczkanus 
us patricius 
us acdestis 
us imperator 
us charltonius 
us inopinatus 
us loxias 
us autocrator 
us tenedius 
us acco 
us przewalskii 
us hannyngtoni 
us simo 
us hardwickii 
us Orleans 
us clodius 
us eversmanni 
us felderi 
us ariadne 
us nordmanni 
us mnemosyne 
us stubbendorfi 
us glacialis 
us apollonius 
us honrathi 
us bremeri 
us jacquemontii 
us epaphus 
us actius 
us phoebus 
us tianschanicus 
us nomion 
us apollo 



Tribe: ZERYNTHIINI 

41 Sericinus montela 

42 Allancastria cerisy 

43 Allancastria deyrollei 

44 Allancastria caucasica 

45 Allancastria louristana 



R 



+ + 



R 



+ 



R 



+ + 



+ + + 
+ 
+ 



+ 
+ 



+ + 



+ + + 



+ + 


+ 


+ 




+ 


+ 


+ + 


+ 
+ 




+ 




+ 
+ 


+ 




+ 
+ 
+ 


+ 


+ 


+ 




+ 


+ 
+ + 


+ 


+ 


+ 


+ 


+ 






+ 


+ 
+ 



168 



Trade 



Table 5.1 (cont.) 





CAT 


LAW 


R/F 


A 


B 


C 


D 


E F G H 


46 Zerynthia polyxena 




* 


o 


+ 


+ 


+ 




+ 


47 Zerynthia rumina 




* 


o 




+ 


+ 


+ 




48 Bhutanitis mansfieldi 


R 
















49 Bhutanitis thaidina 


R 














+ + + 


50 Bhutanitis lidderdalii 




* 










+ 




51 Bhutanitis ludlowi 


K 
















52 Luehdorfia chinensis 


K 
















53 Luehdorfia japonica 


V 






+ 


+ 


+ 






54 Luehdorfia puziloi 










+ 


+ 







Subfamily: PAPILIONINAE 
Tribe: LEPTOCIRCINI 

55 Iphiclides podalirius 

56 Iphiclides podalirinus 

57 Teinopalpus imperialis 

58 Teinopalpus aureus 

59 Meandrusa sciron 

60 Meandrusa payeni 

61 Eurylides marcellus 

62 Eurytides epidaus 

63 Eurytides zonaria 

64 Eurytides marcellinus 

65 Eurytides celadon 

66 Eurytides philolaus 

67 Eurytides anaxilaus 

68 Eurytides xanticles 

69 Eurytides oberthueri 

70 Eurytides bellerophon 

71 Eurytides agesilaus 

72 Eurytides orthosilaus 

73 Eurytides helios 

74 Eurytides stenodesmus 

75 Eurytides earis 

76 Eurytides telesilaus 

77 Eurytides aguiari 

78 Eurytides embrikstrandi 

79 Eurytides travassosi 

80 Eurytides molops 

81 Eurytides macrosilaus 

82 Eurytides nigricornis 

83 Eurytides protesilaus 

84 Eurylides glaucolaus 

85 Eurytides asius 

86 Eurytides microdamas 

87 Eurytides thymbraeus 

88 Eurytides belesis 

89 Eurytides branchus 

90 Eurytides ilus 

91 Eurytides lysithous 
ssp. harrisianus 

92 Eurytides ariarathes 

93 Eurytides harmodius 



R 

K 



V 



+ 



+ + 

+ + + + 

+ + + 
+ + 



+ 
+ + + 



+ 



+ + 



+ + 



+ + 
+ 

-f + 

+ + 

+ + 

+ 
+ + 
+ + 

+ + 

+ 

+ + 



+ 



169 



Threatened Swallowtail Butterflies of the World 
Table 5.1 (cont.) 



CAT LAW R/F 



ABCDEFGH 



94 Eurytides trapeza 

95 Eurytides xynias 

96 Eurytides phaon 

97 Eurytides euryleon 

98 Eurytides pausanias 

99 Eurytides protodamas 

100 Eurytides marchandi 

101 Eurytides thyastes 

102 Eurytides calliste 

103 Eurytides leucaspis 

104 Eurytides lacandones 

105 Eurytides dioxippus 

1 06 Eurytides serville 

107 Eurytides columbus 

108 Eurytides orabilis 

109 Eurytides salvini 

110 Eurytides callias 

1 1 1 Eurytides dolicaon 

112 Eurytides iphitas 

113 Protographium leosthenes 

114 Lamproptera meges 

115 Lamproptera curius 

116 Graphium eurous 

117 Graphium mandarinus 

118 Graphium alebion 

1 19 Graphium tamerlanus 

120 Graphium aristeus 

121 Graphium nomius 

122 Graphium rhesus 

123 Graphium dorcus 

124 Graphium androcles 

125 Graphium epaminondas 

126 Graphium euphrates 

127 Graphium decolor 

128 Graphium antiphates 

129 Graphium agetes 

130 Graphium stratiotes 

131 Graphium phidias 

132 Graphium encelades 

133 Graphium idaeoides 

134 Graphium delesserti 

135 Graphium xenocles 

136 Graphium macareus 

137 Graphium ramaceus 

138 Graphium megarus 

139 Graphium megaera 

140 Graphium stratocles 

141 Graphium deucalion 

142 Graphium thule 

143 Graphium endochus 

144 Graphium angolanus 

145 Graphium taboranus 



V 



K 



R 





+ 


+ 






+ 


+ 








+ 


+ 


+ 






+ 




+ 


+ 




+ 


+ 








+ 


+ 


+ 






+ 


+ 








+ 




+ 






+ 










+ 




+ 
+ 






+ 


+ 








+ 


+ 




+ 




+ 










+ 


+ 








+ 






+ 








+ 


+ 


+ 






+ 


+ 
+ 




+ 


+ 
+ 


+ 






+ 




+ 

+ 
+ 
+ 


+ 


+ 
+ 


+ 










+ 










+ 











+ 



+ 



+ + + 



+ 




+ 




+ 






+ 


+ 




+ 




+ 








+ 




+ 
+ 




+ 




+ 






+ 


+ 




+ 


+ 




+ 
+ 



170 



Table 5.1 (cont.) 



Trade 



CAT LAW R/F 



ABCDEFGH 



146 Graphium morania 

147 Graphium ridleyanus 

148 Graphium philonoe 

149 Graphium almansor 
ssp carchedonius 
ssp kigoma 

150 Graphium poggianus 

151 Graphium adamastor 

152 Graphium agamedes 

153 Graphium aurivilliusi 

154 Graphium olbrechtsi 

155 Graphium odin 

156 Graphium auriger 

157 Graphium hachei 

158 Graphium weberi 

159 Graphium fulleri 

160 Graphium ucalegonides 

161 Graphium ucalegon 

162 Graphium simoni 

163 Graphium cyrnus 

164 Graphium leonidas 

165 Graphium pelopidas 

166 Graphium levassori 

167 Graphium tynderaeus 

168 Graphium latreillanus 

169 Graphium junodi 

170 Graphium nigrescens 

171 Graphium polistratus 

172 Graphium policenes 

173 Graphium porthaon 

174 Graphium illyris 

175 Graphium gudenusi 

176 Graphium kirbyi 
111 Graphium colonna 

178 Graphium antheus 

179 Graphium evombar 

180 Graphium arycles 

181 Graphium bathycles 

182 Graphium chiron 

183 Graphium leechi 

184 Graphium procles 

185 Graphium meyeri 

186 Graphium eurypylus 

187 Graphium doson 

188 Graphium evemon 

189 Graphium agamemnon 

190 Graphium macfarlanei 

191 Graphium meeki 

192 Graphium gelon 

193 Graphium macleayanum 

194 Graphium weiskei 

195 Graphium stresemanni 



K 



K 



R 



R 



+ 




+ 


+ 






+ 


+ 


+ 


+ 




+ 
+ + 


+ 


+ 




+ 


+ 


+ 




+ 


+ 


+ 
+ 




+ 




+ 


+ 




+ 


+ 






+ 


-1- 


+ 




+ 


+ 


+ 




+ 


+ 






+ 




+ 





+ + 



+ + 









+ 












+ 


+ 










+ 












+ 












+ 








+ 


+ 






+ 




+ 


+ 






+ 




+ 






+ 






+ 


+ 










+ 












+ 




+ 








+ 


+ 


+ 








+ 


+ 


+ 








+ 










o 


+ 


+ 


+ 






o 




+ 


+ 
+ 


+ 
+ 








+ 


+ 


+ 





+ + + 



171 



Threatened Swallowtail Butterflies of the World 



Table 5.1 (cont.) 





CAT LAW R/F 


A B C D 


E F G H 


196 Graphium codrus 




o 


+ + 




197 Graphium empedovana 






+ + 


+ 


198 Graphium cloanthus 






+ + + 




199 Graphium sumatranum 






+ 


, 


200 Graphium monticolum 






+ 




201 Graphium milon 






+ + + 




202 Graphium sandawanum 


V 






+ + 


203 Graphium sarpedon 






+ + 




204 Graphium mendana 


R 


o 






205 Graphium wallacei 




o 


+ 




206 Graphium hicetaon 




o 


+ 




207 Graphium browni 






+ 




Tribe: TROIDINI 










208 Battus philenor 






+ 




209 BaWMi zetides 


V 








210 Battus devilliers 








+ + 


211 Battus polydamas 






+ + 




212 Ba«MS streckerianus 










213 Battus archidamas 






+ 




214 Battus polystictus 




o 


+ + 




215 Battus philetas 






+ 




216 Battus madyes 






+ + + 




217 Battus eracon 








+ 


218 Ba«Ms fte/(« 






+ + 




219 Battus crassus 






+ + 




220 Battus laodamas 






+ + 




221 Sa«(/s lycidas 






+ + 




222 Euryades duponchelii 






+ 


+ 


223 Euryades corethrus 






+ + 


+ 


224 Cressida cressida 






+ + + 




225 Parides gundlachianus 










226 Parides alopius 






+ 




227 Parides photinus 






+ + 




228 Parides montezuma 






+ + + 




229 Parides phalaecus 










230 Parides agavus 




o 


+ 




231 Parides proneus 






+ 




21)1 Parides ascanius 


V 






+ 


233 Parides bunichus 






+ 




234 Parides diodorus 










235 Parides perrhebus 






+ + + 




236 Parides hahneU 


R 


o 




+ + 


111 Parides mithras 






+ 




238 Parides chabrias 






+ 




239 Parides quadratus 










240 Parides pizarro 


K 








241 Parides steinbachi 


K 








242 Parides coelus 


K 








243 Parides tros 






+ 


+ 


244 Parides aeneas 






+ + + 





172 



Table 5.1 (cont.) 



Trade 



CAT LAW R/F 



ABCDEFGH 



245 Parides klagesi 

246 Parides orellana 

247 Parides childrenae 

248 Parides sesostris 

249 Parides burchellanus 

250 Parides polyzeliis 

251 Parides iphidamas 

252 Parides vertumnus 

253 Parides cutorina 

254 Parides lycimenes 

255 Parides phosphorus 

256 Parides drucei 

257 Parides erlaces 

258 Parides nephalion 

259 Parides anchises 

260 Parides erithalion 

261 Parides panthonus 

262 Parides aglaope 

263 Parides castilhoi 

264 Parides lysander 

265 Parides echemon 

266 Parides zacynthus 

267 Parides neophilus 

268 Parides eurimedes 

269 Parides timias 

270 Atrophaneura anterior 

271 Atrophaneura daemonius 

272 Atrophaneura phitonius 
TTi Atrophaneura alcinous 
21 A Atrophaneura latreillei 

275 Atrophaneura polla 

276 Atrophaneura crassipes 

277 Atrophaneura adamsoni 

278 Atrophaneura nevilli 

279 Atrophaneura laos 

280 Atrophaneura mencius 

281 Atrophaneura impediens 

282 Atrophaneura febanus 

283 Atrophaneura hedistus 

284 Atrophaneura dasarada 

285 Atrophaneura polyeuctes 

286 Atrophaneura semperi 

287 Atrophaneura kuehni 

288 Atrophaneura luchti 

289 Atrophaneura hageni 

290 Atrophaneura priapus 

291 Atrophaneura sycorax 

292 Atrophaneura horishanus 

293 Atrophaneura aidoneus 

294 Atrophaneura varuna 

295 Atrophaneura zaleucus 

296 Atrophaneura nox 



K 



R 







+ 




+ 


+ 


+ 




+ 




+ 




+ 


+ 






+ 


+ 


+ 


+ 




+ 


+ 






+ 


+ 


+ 


+ 


+ 


+ 




+ 


+ 


+ 






+ 


+ 




+ 




+ 




+ 




+ 
+ 




+ 


+ 


+ 




+ 






+ 


+ 


+ 




+ 


+ 




+ 




+ 


+ 
+ 










+ 


+ 


+ 




+ 


+ 


+ 


+ 


+ 


+ 



+ 



+ + 







+ 


+ 






+ 




+ 
+ 
+ 


+ 
+ 

+ 
+ 


+ 


+ 






+ 


+ 


+ 




+ 


+ 




+ 






+ 


+ 




+ 


+ 




+ 


+ 











173 



Threatened Swallowtail Butterflies of the World 
Table 5.1 (cont.) 



CAT LAW R/F 



ABCDEFGH 



297 Atrophaneura dixoni 




298 Atrophaneura neptunus 




299 Atrophaneura palu 


K 


300 Atrophaneura coon 




301 Atrophaneura rhodifer 




302 Atrophaneura hector 




303 Atrophaneura jophon 


V 


304 Atrophaneura pandiyana 




305 Atrophaneura oreon 




306 Atrophaneura liris 




307 Atrophaneura polyphonies 




308 Atrophaneura schadenbergi 


V 


309 Atrophaneura mariae 




310 Atrophaneura phegeus 




311 Atrophaneura phlegon 




312 Atrophaneura atropos 


I 


313 Atrophaneura kotzebuea 




314 Atrophaneura aristolochiae 




315/4 trophaneura polydorus 




316 Trogonoptera brookiana 




317 Trogonoptera trojana 




318 Troides hypolitus 




319 Troides cuneifer 




320 Troides amphrysus 




321 Troides miranda 




322 Troides andromache 


I 


323 Troides magellanus 




324 Troides prattorum 


I 


325 Troides minos 




326 Troides aeacus 




327 Troides plateni 




328 Troides rhadamantus 




329 Troides dohertyi 


V 


330 Troides helena 




331 Troides oblongomaculatus 




332 Troides darsius 




333 Troides criton 




334 Troides riedeU 




335 Troides plato 




336 Troides vandepolli 




337 Troides haliphron 




338 Troides staudingeri 




339 Ornithoptera goliath 




340 Ornithoptera tithonus 


K 


341 Ornithoptera rothschildi 


I 


342 Ornithoptera chimaera 


I 


ssp. chary bdis 




343 Ornithoptera paradisea 


I 


344 Ornithoptera meridionalis 


V 


345 Ornithoptera alexandrae 


E 


346 Ornithoptera victoriae 




ssp. regw 











+ 


+ 














+ 


+ 


+ 


+ 












+ 


+ 


+ 


+ 
+ 










+ 


+ 
+ 


+ 

+ 
+ 
+ 




+ 










+ 


+ 
+ 


+ 
+ 
+ 


+ 










o 


+ 


+ 


+ 


+ 










o 


+ 


+ 


+ 


+ 










o 


+ 


+ 

+ 
+ 

+ 


+ 
+ 
+ 
+ 
+ 

+ 

+ 
+ 
+ 
+ 
+ 


+ 
+ 
+ 
+ 
+ 
+ 
+ 

+ 

+ 


+ 
+ 

+ 
+ 

+ 


+ 
+ 
+ 

+ 


+ 

+ 

+ 




o 




+ 


+ 
+ 
+ 

+ 
+ 


+ 

+ 
+ 

+ 

+ 
+ 
+ 


+ 

+ 

+ 
+ 
+ 
+ 

+ 


+ 
+ 

+ 

+ 

+ 


+ 

+ 
+ 
+ 

+ 
+ 
+ 


+ 
+ 



+ 



174 



Trade 



Table 5.1 (cont.) 





CAT 


LAW 


R/F 


ABC 


D 


E 


F 


G 


H 


347 Ornithoptera aesacus 


I 














+ 


+ 


348 Ornithoptera croesus 


V 






+ 


+ 


+ 




+ 




349 Ornithoptera priamus 




* 





+ 


+ 


+ 


+ 






ssp. admiralitatis 




* 


? 


+ 


+ 










ssp. arruana 




* 




+ 


+ 










ssp. boisduvaU 




* 






+ 










ssp. bornemanni 




* 




+ 


+ 










ssp. caelestis 




* 


o 


+ 


+ 


+ 


+ 






ssp. demophanes 




* 




+ 


+ 










ssp. hecuba 




* 
















ssp. miokensis 




* 


? 




+ 










ssp. poseidon 




* 


o 


+ 












ssp. priamus 




* 




+ 




+ 








ssp. pronomus 












+ 








ssp. teucnis 




* 




+ 












ssp. urvilliana 




* 


o 


+ 


+ 










350 Ornithoptera euphorion 




* 






+ 


+ 








351 Ornithoptera richmondia 










+ 











Tribe: PAPILIONINI 

352 Papilio glaucus 

353 Papilio alexiares 

354 Papilio multicaudatus 

355 Papilio rutulus 

356 Papilio eurymedon 

357 Papilio pilumnus 

358 Papilio palamedes 

359 Papilio troilus 

360 Papilio ascolius 

361 Papilio neyi 

362 Papilio zagreus 

363 Papilio bachus 

364 Papilio hellanichus 

365 Papilio scamander 

366 Papilio xanthopleura 

367 Papilio birchalli 

368 Papilio cleotas 

369 Papilio menatius 

370 Papilio phaeton 

371 Papilio victorinus 

372 Papilio garamas 

373 Papilio abderus 

374 Papilio homerus 

375 Papilio warscewiczi 
yit Papilio cacicus 

377 Papilio euterpinus 

378 Papilio esperanza 

379 Papilio andraemon 

380 Papilio machaonides 

381 Papilio thersites 

382 Papilio astyalus 

383 Papilio androgens 



+ + 



+ 


+ 




+ 


+ 




+ 


+ 






+ 


+ 


+ 




+ 


+ 


+ 


+ 


+ 


+ 




+ 


+ 





+ + 

+ 
+ + + + 

+ + 

+ 
+ 

+ 
+ 

+ 



+ + 
+ + + 



+ + 



175 



Threatened Swallowtail Butterflies of the World 
Table 5.1 (cont.) 



CAT LAW R/F ABCDEFGH 



384 Papilio ornythion + + 

385 Papilio aristodemus + 
ssp. ponceanus E * 

386 Papilio caiguanabus I 

387 Papilio aristor I 

388 Papilio thoas o + + + 

389 Papilio cinyras + 

390 Papilio homothoas + 

391 Papilio cresphontes + 

392 Papilio paeon + + + 

393 Papilio garleppi K + 

394 Papilio torquatus + + 

395 Papilio hectorides o + + 

396 Papilio lamarchei 

397 Papilio himeros V 

398 Papilio hyppason + + 

399 Papilio pelaus 

400 Papilio oxyuias 

401 Papilio epenetus 

402 Papilio erostratus + 

403 Papilio erostratinus 

404 Papilio pharnaces + 

405 Papilio chiansiades + + 

406 Papilio dospassosi 

407 Papilio anchisiades o + + 

408 Papilio maroni K 

409 Papilio rogeri 

410 Papilio isidorus 

411 Papilio rhodostictus 
All Papilio anactus 

413 Papilio elwesi 

414 Papilio maraho V 

415 Papilio clytia * 

416 Papilio paradoxa * 

417 Papilio veiovis 

418 Papilio osmana V 

419 Papilio carolinensis V 

420 Papilio agestor 

421 Papilio epycides * 

422 Papilio slateri * 

423 Papilio laglaizei 

424 Papilio toboroi R 

425 Papilio moerneri V 

426 Papilio alexanor + 

427 Papilio hospiton E * 

428 Papilio machaon * o + + + 

429 Papilio hippocrates + 

430 Papilio zelicaon + 

431 Papilio indra + + 

432 Papilio polyxenes + + + 

433 Papilio brevicauda 

434 Papilio oregonia + 

176 



+ 




+ 






+ 


+ 


+ 


+ 


+ 
+ 


+ 


+ 


+ 






+ 


+ 


+ 
+ 


+ 




+ 


+ 




+ 




+ 




+ 


+ 




+ 




+ 










+ 


+ 


+ 






+ 







Table 5.1 (cont.) 



Trade 



CAT LAW R/F ABCDEFGH 



435 Papilio 

436 Papilio 

437 Papilio 

438 Papilio 

439 Papilio 

440 Pap/7;o 

441 Papilio 

442 Papilio 

443 Papilio 

444 Papilio 

445 Papilio 

446 Papilio 

447 Papilio 

448 Pap(7(o 

449 PapfY/o 

450 Pap/7/o 

451 Pa/7(7/o 

452 Pflp///o 

453 Papilio 

454 Papilio 

455 Papilio 

456 Papilio 

457 Papilio 

458 Pap/7/o 

459 Pfl/7(7;'o 

460 Pa/?(7/o 

461 Papilio 

462 Papilio 

463 Papilio 

464 Papilio 

465 Papilio 

466 Papilio 

467 Papilio 

468 Pap(7;o 

469 Pa/7/7(o 

470 Pfl/7(7(0 

471 Papilio 
All Papilio 

473 Pap/7/o 

474 Papilio 

475 Pap(7/o 

476 Papilio 
All Papilio 

478 Pap/7/o 

479 Pap/7/o 

480 Pap/7/o 

481 Pa/7j7/o 

482 Pap/7/o 

483 Pflp/7;o 

484 Pap(7/o 

485 Pap(7(o 

486 Pflp(7/o 



kahli 

nitra 

coloro 

bairdi 

joanae 

xuthus 

benguetanus 

euchenor 

gigon 

demolion 

liomedon 

protenor 

demetrius 

macilentus 

bootes 

janaka 

lampsacus 

forbesi 

acheron 

oenomaus 

ascalaphus 

rumanzovia 

deiphobus 

alcmenor 

thaiwanus 

polymnestor 

lowi 

memnon 

mayo 

noblei 

anionio 

biseriatus 

iswara 

hystaspes 

sataspes 

helenus 

iswaroides 

jordani 

polytes 

alphenor 

am b rax 

phestus 

diophantus 

nephelus 

castor 

mahadeva 

dravidarum 

hipponous 

pitmani 

fuscus 

canopus 

albinus 



R 



R 











+ 












+ 




o 


+ 


+ 


+ 
+ 


+ 
+ 






+ 


+ 


+ 






o 


+ 


+ 


+ 






o 


+ 
+ 


+ 


+ 
+ 
+ 


+ 


+ 






+ 


+ 

+ 


+ 
+ 


+ 






+ 


+ 










+ 


+ 


+ 






+ 


+ 


+ 








+ 


+ 
+ 


+ 
+ 






o 


+ 
+ 


+ 
+ 


+ 

+ 
+ 


+ 
+ 


+ 


o 


+ 


+ 
+ 


+ 






o 


+ 
+ 


+ 


+ 






o 




+ 


+ 
+ 












+ 


+ 


+ 




+ 


+ 


+ 








+ 












+ 


+ 


+ 
+ 


+ 
+ 


+ 


o 


+ 


+ 


+ 







+ + 



177 



Threatened Swallowtail Butterflies of the World 
Table 5.1 (cont.) 



CAT LAW R/F 



ABCDEFGH 



487 Papilio hypsicles 

488 Papilio woodfordi 

489 Papilio heringi 

490 Papilio amynthor 

491 Papilio schmeltzii 

492 Papilio godeffroyi 

493 Papilio inopinatus 

494 Papilio bridgei 

495 Papilio weymeri 

496 Papilio gambrisius 

497 Papilio tydeus 

498 Papilio aegeus 

499 Papilio cynorta 

500 Papilio plagiatus 

501 Papilio zoroastres 

502 Papilio echerioides 

503 Papilio jacksoni 

504 Papilio fuelleborni 

505 Papilio sjoestedti 

506 Papilio rex 

507 Papilio epiphorbas 

508 Papilio manlius 

509 Papilio phorbanta 

510 Papilio charopus 
ssp. juvenilis 

511 Papilio hornimani 

512 Papilio mackinnoni 

513 Papilio sosia 

514 Papilio aethiopsis 

515 Papilio nireus 

516 Papilio arisiophonies 

517 Papilio oribazus 

518 Papilio ihuraui 

519 Papilio desmondi 
ssp. /e/Ya 

520 Papilio interjecta 

521 Papilio bromius 

522 Papilio chrapkowskii 

523 Papilio zalmoxis 

524 Papilio gal lien us 

525 Papilio mechowi 

526 Papilio zenobius 

527 Papilio mechowianus 

528 Papilio andronicus 

529 Papilio zenobia 

530 Papilio maesseni 

531 Papilio antimachus 

532 Papilio ophidicephalus 

533 Papilio menesiheus 

534 Papilio lormieri 

535 Papilio morondavana 

536 Papilio grosesmithi 



R 



R 



I 
V 



R 



V 
R 



+ 
+ + 



+ 











+ 


+ 


o 






+ 


+ 


+ 


o 






+ 
+ 


+ 

+ 
+ 




o 


+ 


+ 


+ 








+ 




+ 


+ 










+ 


+ 


+ 




+ 


+ 


+ 
+ 







+ + 



+ + 



-I- + + 



+ + 
+ + 



+ + 



+ 



+ 


+ 




+ + 


+ 




+ + 


+ 


+ 




+ 


+ 



+ + 



+ 



+ 

+ + + 

+ + + 
+ 
+ + 
+ + 

+ + + + 
+ + + + 



178 



Table 5.1 (cont.) 



Trade 



CAT LAW R/F 



ABCDEFGH 



537 Papi 

538 Papi 

539 Papi 

540 Papi 

541 Papi 

542 Papf 

543 Papi 

544 Pap/ 

545 Pap« 

546 Papi 

547 Papi 

548 Papi 

549 Papj 

550 Pap; 
ssp 

551 Papi 

552 Pap/ 

553 Papi 

554 Papj, 

555 Papi 

556 Papj 

557 Pap/, 

558 Papi 

559 Pap/ 

560 Papi 

561 Pap/ 

562 Pap/- 

563 Papi 

564 Pap/ 

565 Papi 

566 Pap/ 

567 Pap/ 

568 Papi 

569 Pap/ 

570 Papi 

571 Pap/ 

572 Pap/, 

573 Pap 



'io erithonioides 
'io demodocus 
'io demoleus 
io leucotaenia 
io delalandei 
io mangoura 
io constantinus 
io phorcas 
io dardanus 
io euphranor 
io pelodorus 
'io Hesperus 
'io horribilis 
'io nobilis 
crippsianus 
'io crino 
'io blumei 
'io buddha 
'io palinurus 
'io daedalus 
'io chikae 
'io maackii 
'io bianor 
'io syfanius 
'io polyctor 
'io dialis 
io elephenor 
io paris 
io karna 
io arcturus 
io hoppo 
io krishna 
io neumoegeni 
io peranthus 
io pericles 
io lorquinianus 
io ulysses 
io montrouzieri 



V 
R 



V 





+ 


+ 


+ 




+ 






+ 


+ 


+ 








o 


+ 
+ 

+ 


+ 
+ 

+ 


+ 
+ 
+ 
+ 


+ 

+ 


+ 






+ 


+ 


+ 


+ 
+ 


+ 






+ 


+ 

+ 

+ 
+ 


+ 

+ 

+ 
+ 

+ 


+ 
+ 
+ 
+ 


+ 
+ 


+ 
+ + 


o 




+ 


+ 








o 


+ 
+ 


+ 

+ 
+ 


+ 
+ 


+ 


+ 




o 


+ 
+ 


+ 
+ 


+ 
+ 
+ 
+ 

+ 
+ 
+ 


+ 
+ 
+ 

+ 
+ 
+ 
+ 


+ 

+ 
+ 


+ 
+ 


o 


+ 


+ 


+ 


+ 
+ 


+ 





179 



6 



Reviews of threatened species 



The following 78 reviews summarize our present knowledge of those swallowtail 
species for which some degree of threat has been established or is suspected. The 
order of appearance is given on the Contents pages. A list of the 78 species sorted into 
their threatened categories is given in Appendix A at the end of this section (p. 369). 

The criteria used in selecting species for full review have been outlined in section 1 , 
How to use this book. There it was emphasised that Red Data Book categories are 
constantly being updated and changed. There are always species that could arguably 
be in a higher category, while others could be in a lower one. To be categorized as 
threatened (i.e. Endangered, Vulnerable, Rare or Indeterminate) a species must be 
at risk on a world scale, hence the absence of an RDB category for a particular species 
implies nothing about localized threats. Species not given a full review may 
nevertheless be threatened over part of their range, and the full species list in section 3 
(p. 33) should be consulted for details. 

As noted in section 1 , there are very many species that have not been given an RDB 
category but for which monitoring and further information are still required. Those 
species, of which there are almost 100, are listed in Appendix B (p. 371). 

We would be grateful if readers could bring new information and shortcomings in 
these reports to our attention. 



181 



Threatened Swallowtail Butterflies of the World 

Baronia brevicornisSalvin, 1893 RARE 

Subfamily BARONIINAE 



Summary Baronia brevicornis (the Baronia) is the most primitive living 
swallowtail, a 'living fossil' of great scientific interest. The species is confined to 
Mexico and although it is well distributed there, its populations should continue to be 
monitored, studied and protected. 

Description Baronia brevicornis is a dark brown tailless butterfly with 
variable ochreous markings (Plate 1.1). The wingspan is 55-65 mm, and the female is 
larger than the male but otherwise similar in appearance (2). Both sexes are rather 
variable in colour and some individuals, particularly females, are much darker than 
others (3). The antennae are very short (11). Although the adults have the fore-tibial 
spur which is characteristic of the Papilionidae, the wing venation is very uncharac- 
teristic in having both anal veins on the hindwing, like Pieridae and Nymphalidae 
(8, 12). It must therefore be assumed that Baronia brevicornis is primitive and 
diverged before the time when the vein was lost by other members of the Papilionidae 
(12). 

Male: UFW ground colour brown with yellow-ochre oval spots forming three 
arches on the wing. 

UHW with yellow-ochre markings which include two large patches, one covering 
two-thirds of the cell, the other a triangle below the cell, and a discal row of spots. 

LFW/LHW paler than the upper surface. The subapical spots of the forewing, and 
all of the hindwing spots are silver. 

The female is larger than the male, with more extensive yellow-ochre markings. 

The mature larvahas a black, yellow or green (15) head and a green body with small 
tubercles which may be either black or yellow (6, 8, 12). The body has a continuous 
yellow dorsal stripe and white transverse lines on each segment (6,8,12). In common 
with all other Papilionidae, the caterpillar possesses a defensive osmeterium 
(4,6,8,11,12). 

Distribution This species is restricted to areas where its foodplant occurs in 
the Sierra Madre del Sur in south-west Mexico. It has been recorded from localities in 
the States of Jalisco, Oaxaca, Chiapas, Guerrero, Morelos, Puebla, Michoacan, and 
Colima (6, 9, 10, 12, 14). Its occurrence is very local. 

This distribution is somewhat enigmatic for such a primitive species, being far 
removed from South-East Asia, the region which, on grounds of having the greatest 
diversity of swallowtails, has often been presumed to be their centre of origin (12). 
However, in a recent assessment Hancock has succinctly re-evaluated the data and 
proposed from fossil and geological evidence that North America holds the key to the 
ancestry of the Papilionidae (5). Praepapilio Colorado, a fossil papilionid from middle 
Eocene deposits in Colorado , U . S . A . , shares some of the primitive features found in 
Baronia (5) and might easily be included within that genus (1). At the time when the 
family was differentiating, Mexico was separated from South America by a 
considerable body of water and the western Sierra Madre was a peninsula, 
presumably with Baronia brevicornis inhabiting the tip (12). The impact of 
physiographic, climatic and vegetational factors on the distribution of Baronia 
brevicornis has been assessed, and the presence of disjunct populations as far south as 
Oaxaca and Chiapas is believed to be the result of southward movements of the 

182 



*' Baronia brevicornis 

lepidopteran fauna during the Pleistocene Ice Ages (10). The present extent of the 
range of Baronia brevicornis is limited to alititudes of 505-1335 m (10), where its 
foodplant grows. 

Habitat and Ecology Baronia brevicornis was described in 1893 (11) but it 
was not until 1961 that Vazquez and Perez elucidated some of its fascinating life 
history (10, 14). The habitat is thickets of deciduous scrub composed mainly of the 
very unusual foodplant Acacia cochliacantha (Leguminosae) (not A . cymbispina, as 
is often quoted) (10). The Acacia is known locally as 'cubata' and the thickets are 
called 'cubatera' (14). Other plants in the thickets are Bursera spp. (Torchwood), 
Ipomoea spp. (Morning Glory), Mimosa polyantha and Neobuxbaumia mezcalaensis 
(12). The Acacia trees are bare until the first rains come and, as the leaves grow out, 
the butterflies emerge. Females lay their eggs singly, few on each tree (14). The 
butterfly is said to be single-brooded, and the larvae dig themselves into loose ground 
for pupation (10, 14). The adults have a short flight range and the males are 
territorial, defending their areas from intrusion by assailing intruders from a vantage 
point high in the branches of an Acacia tree (7). At Morelos during July, both males 
and females come down to drink on the sandy banks of small streams (7). 

At suitable localities and in the right season Baronia may be quite numerous 
(7, 10). The population in the Mezcala region of Guerrero State has been studied for 
ten years (9). Questions elucidated in that study include the influence of rainfall on 
adult emergence and the duration of egg-laying periods, larval stages and adult 
populations (9). Population parameters have been described, notably the drop in 
populations during the 1960s and the impact of natural predators on the species (9). 
The eggs of Baronia brevicornis are parasitized by chalcid wasps of the family 
Trichogrammatidae. The caterpillars suffer little predation, possibly because of the 
osmeterium (see below), but some are taken by bugs of the family Pentatomidae. 
Adults are devoured by spiders in the Argiopidae and by robber flies ( Asilidae) which 
catch them in flight (9). 

As noted above, Baronia brevicornis is the 'living fossil' of the Papilionidae. 
Because of its primitive characteristics and unusual distribution and feeding habits, it 
is of very great scientific interest. Studies of the osmeterial secretions of Baronia 
showed them to be mainly composed of two aliphatic acids, isobutyric acid and 
2-methyl butyric acid (4). When disturbed, the larvae extrude the osmeterial gland 
and try to wipe it against the attacker. The secretion is strongly odorous and visably 
coats the erected gland (4). In Papilio machaon at least, the same acids are strongly 
repellent to ants (4). Other studies have demonstrated the presence of cholesterol, 
three saturated hydrocarbons and two monounsaturated acids in the caterpillar 
bodies, although the siginificance of these discoveries is not clear (16). 

Threats Baronia brevicornis is under no immediate threat, but with such a 
limited distribution it must be considered to be rare on a world basis. Being of such 
great evolutionary interest, the populations attract considerable attention from 
collectors and scientists worldwide . The human population of Mexico is growing at an 
alarming rate and this may eventually lead to degradation of the Acacia thickets in 
which the butterflies breed. 

Conservation Measures The Mexican Government should recognize the 
international importance of Baronia brevicornis and, whilst ensuring that popula- 
tions are protected from excessive exploitation, should encourage further research 
and study by local and expatriate scientists. The extent of Acacia thickets within 

183 



Threatened Swallowtail Butterflies of the World 

national park boundaries is unknown . An assessment of these , and their utilization by 
Baronia would be a necessity for its long-term protection from habitat destruction . A 
colony of Baronia does occur within the Canon del Sumidero National Park near 
Tuxta Gutierrez, Chiapas (7). 

References 

1. Brown, K.S., Jr. (1982). Historical and ecological factors in the biogeography of 
aposematic neotropical butterflies. American Zoologist 22: 453-471. 

2. D'Abrera, B. (1981). Butterflies of tiie Neotropical Region. Parti. Papilionidae and 
Pieridae. Lansdowne Editions, Melbourne, xvi + 172 pp. 

3. Diaz Frances, A. and Maza E. J. de la, (1978). Gui'a illustrada de las mariposas Mexicanas. 
Parte 1. Familia Papilionidae. Sociedad Mexicana de Lepidopterologia A.C. Publicaciones 
Espec tales 3: 15 pp. 

4. Eisner, T., Pliske, T.E., Ikeda, M., Owen, D.F., Vazquez, L., Perez, H., Franclemont, 
J.G. and Meinwald, J. (1970). Defense mechanisms of arthropods. XXVII. Osmeterial 
secretions of papilionid caterpillars (Baronia, Papilio, Eurytides). Annals of the Entomolo- 
gical Society of America 63: 914-915. 

5. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

6. Igarashi, S. (1979). Papilionidae and Their Early Stages. Vol 1: 219 pp.. Vol 2: 102 pp. of 
plates. Kodansho, Tokyo. (In Japanese). 

7. Maza E. J. de la (1984). In litt. , 3 January. 

8. Perez R., H. (1969). Quetotaxia y morfologia de la oruga de Baronia brevicornis Salv. 
(Lepidoptera Papilionidae Baroniinae). Anales del Insiituto de Biologia Universidad de 
Mexico. 40, Ser. Zool. (2): 227-244. 

9. Perez R., H. (1971). Algunas consideraciones sobre la poblacion de Baronia brevicornis 
Salv. (Lepidoptera, Papilionidae, Baroniinae) en la region de Mezcala, Guerrero. Anales 
del Institute de Biologia Universidad de Mexico 42, Ser. Zool. (1): 63-72. 

10. Perez R., H. (1977). Distribucion geografica y estructura poblacional de Baronia 
brevicornis Salv. (Lepidoptera, Papilionidae, Baroniinae) en la Repiiblica Mexicana. 
Anales del Insiituto de Biologia Universidad de Mexico 48, Ser. Zool. (5): 151-164. 

11. Salvin, O. (1893). Description of a new genus and species of Papilionidae from Mexico. 
Transactions of the Entomological Society of London 1893 (4): 331-332. 

12. Tyler, H.A. (1975). The Swallowtail Butterflies of North America Naturegraph, California, 
viii + 192 pp., 16 pi. 

13. Tyler, H.A. (1983). In litt., 13 March. 

14. Vazquez G., L. and Perez R., H. (1961). Observaciones sobre la biologia de Baronia 
brevicornis Salv. (Lepidoptera: Papilionidae — Baroniinae). Anales del Institute de Biolo- 
gia Universidad de Mexico. 32: 295-311. 

15. Vazquez G., L. and Perez R., H. (1967). Nuevas observaciones sobre le biologia de 
Baronia brevicornis Salv. Lepidoptera: Papilionidae — Baroniinae. Anales del Institute de 
Biolegia Universidad de Mexico. 37: 195-204. 

16. Yuste, F., Perez, H. and Walls, F. (1972). Compounds of papilionid caterpillars (Baronia 
brevicornis S.). Experientia 28: 1149. 



184 



Parnassius autocrator 
Parnassius autocrator Avinoff, 1913 RARE 

Subfamily PARNASSIINAE Tribe PARNASSIINI 



Summary Parnassius autocrator flies at high altitudes in the Hindu Kush of 
Afghanistan and the Pamir mountains of Tadzhikskaya S.S.R. It is extremely local in 
its distribution and possibly threatened by degradation of high mountain pastures in 
the U. S.S.R. The status of the species in Afghanistan is presently unknown. 

Description Parnassius autocrator was originally described as a subspecies 
of P. charltonius Gray. Bryk agreed with this (3) and Munroe omitted the species 
from his list (6), but recent authors consider it to be a full species (4, 5, 8, 9, 11). 
Parnassius autocrator is somewhat unusual in appearance, lacking the red spots so 
characteristic of many other Parnassius species, but with yellow marks on the hind 
wings (8). The sexes differ in that the female has a wide orange patch on the UHW, 
while the male has only a wavy yellow-orange line (1) (Plates 1.3 and 1.4). 

UFW/UHW chalky white with shadowy black markings and transparent apices. 
Hind wings with a row of black submarginal lunules and a bright yellow postdiscal 
bar. The undersides are similar. 

Distribution Restricted to a narrow range in the Pamir Mountains of 
south-eastern Tadzhikistan (Tadzhikskaya) S.S.R. in the U. S.S.R. and the Hindu 
Kush in north-eastern Afghanistan (1,4, 11). Very few specimens have been taken in 
the U. S.S.R. and it is not known to complete its life cycle there (7). 

Habitat and Ecology Parnassius autocrator is a member of the charltonius 
species group (5), whose young stages feed on Fumariaceae. The foodplant of P. 
autocrator is Corydalis adiantifolia (and possibly C. hindukushensis) , a very local 
plant that grows on steep, rocky slopes and cliffs at altitudes around 3000 m and 
above (4, 11). In north-eastern Afghanistan the butterflies are found in the narrow 
steep-sided valleys where the foodplant occurs (11). They fly at altitudes between 
2800 and 4000 m, but are most numerous between 3200 and 3500 m (11), sometimes 
in the company of P. charltonius (A, 11). The adults emerge around the second week 
of July and are on the wing for four weeks. Females flutter around the foodplant or 
settle on stones or thistle flowers, taking off at the slightest disturbance (11). The 
males are more active, soaring about a metre from the ground up and down the 
slopes, mainly between 09.00 and 15.00 hrs (11). 

The climate at these altitudes in the Hindu Kush is dry and hot in summer with deep 
snow in winter. The landscape in summer is bare and dry with thorny, succulent and 
aromatic shrubs growing on rocky bare earth and scree (11). This is in great contast to 
the meadows in which alpine parnassians such as P. phoebus fly. The U. S.S.R. 
populations of P. autocrator are said to fly in pastures (9) but the composition and 
extent of these habitats are unknown. 

Threats Listed in the U. S.S.R. Red Data Book (2) and stated by 
Tanasiychuk (9) to be an extremely rare species in the U. S.S.R., threatened with 
extinction because of degradation of high mountain pastures. No further detail is 
given and the statement is taken to be rather speculative. However, the habitat and 
foodplants of Parnassius autocrator may be damaged by heavy grazing of sheep and 
goats. The species is known to be extremely local in north-eastern Afghanistan, but 

185 



Threatened Swallowtail Butterflies of the World 

no threats are known (11). The region is at present inaccesible, but there are reasons 
to fear that environmental degradation may be an increasing problem. 

Conservation Measures No protected areas are known in this region and 
there is very little information on the extent of human impact on the fauna and flora of 
the high pastures where the butterfly lives. The main part of the butterfly's range is 
clearly in Afghanistan and conservation efforts should be directed to that country. 
More data are required on the precise distribution, biology and habitat requirements 
of Parnassius autocrator. When the region is once again open to scientists it will be 
necessary to develop a conservation plan. 



References 

1. Ackery, P.R. (1975). A guide to the genera and species of Parnassiinae (Lepidoptera: 
Papilionidae). Bulletin of the British Museum (Natural History). Entomology 31: 71-105. 

2. Bannikov.A.G. and Sokolov, V.I. (eds) (1984) r/ie/?erf Date BooA:o/f/ie[/55/?. Rare and 
Threatened Species of Animals and Plants. Lesnaya Prom. Press, Moscow. (In Russian). 

3. Bryk, F. (1935). Lepidoptera, Parnassiidae. Part 2. (Subfam. Parnassiinae). Tierreich 65: 
51 + 790 pp., 698 figs. 

4. Eisner, C. andNaumann,C.M. (1980). ParnassianaNovaLVII. BeitragzurOkologieund 
Taxonomie der Afghanischen Parnassiidae (Lepidoptera). Zoologische Verhandelingen 
178: 35 pp. + 9 pi. 

5. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersial: 1-48. 

6. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
£«/owc>/og«r Supplement 17: 1-51. 

7. Nekrutenko, Y.P. (1984). In litt.,20 June. 

8. Smart, P. (1975). The Illustrated Encyclopedia of the Butterfly World. Hamlyn, London. 275 
pp. 

9. Tanasiychuk, V.N. (1981). Data for the "Red Book" of Insects of the U.S. S.R. 
Entomologicheskoye Obozreniye 60 (3): 168-186. 

10. Turlin, B. (1984). In litt., 1 June. 

11. Wyatt,C. andOmoto, K. (1963). Aui der iagd nach Parnassius autocrator Awin. Zeitschrift 
der Wiener Entomologischen Gesellschaft 48: 163-170. 



186 



*" Parnassius apollo 

Parnassius apollo (Linnaeus, 1758) RARE 

Subfamily PARNASSIINAE Tribe PARNASSIINI 



Summary Parnassius apollo, the Apollo butterfly, was the first insect to be 
included under the Convention on International Trade in Endangered Species of 
Wild Fauna and Flora (CITES). Prone to local subspeciation in montane and 
northern Eurasia, the Apollo has many named regional populations. A number of 
these are Extinct or Endangered although others are still numerous. 

Description The Apollo is a medium-sized (50-80 mm wingspan) butterfly 
with rounded, chalky white wings, with black spots, grey markings and transparent 
areas lacking scales. The pattern, density and intensity of marking varies according to 
locality, but the hindwings always have striking scarlet spots. The antennal shaft is 
grey with darker rings (1, 23). 

Distribution Parnassius apollo was formerly widely distributed in Europe 
and Asia, although its range is now somewhat depleted. In northern Europe it has 
been recorded from Norway, Denmark, Sweden and Finland across to Latvia and 
central Siberia in the U.S.S.R. In central and southern Europe it has been recorded in 
Spain, France, Switzerland, Italy, Liechtenstein, Austria, West Germany, Nether- 
lands (vagrants?). East Germany, Poland, Czechoslovakia, Hungary, Yugoslavia, 
Greece, Bulgaria, Turkey, Romania, and Syria, eastwards into the Ukraine, 
BoFshoy Kavkaz (Caucasus) and Siberia in the U.S.S.R., China (Sinkiang) and 
Mongolia (3, 4, 5, 8, 18, 19, 21, 22, 23, 35). Populations are often isolated, disjunct 
and local. 

Habitat and Ecology This butterfly, a relict of the glacial epoch, occurs in 
subalpine situations between 750 and 2000 m in the Alps and associated ranges, but 
near sea level in the northern parts of its range. The larvae feed exclusively on 
stonecrops {Sedum spp.) (24). There is one generation per year, with over-wintering 
in the egg stage. Its population biology differs from one colony to another and 
individuals may be numerous or extremely rare. The Apollo seems to be able to 
withstand a certain degree of grazing of its habitats, which are commonly rocky and 
relatively xeric (5). Major alterations affecting the host plant are not tolerated 
(21,23). Additional details for specific races are available in the literature 
(12,26,28,30,37,39,40). 

Threats Parnassius apollo is reported to have declined and become rare or 
endangered in all or part of the following countries: Bulgaria, Czechoslovakia, 
Finland (26,27), France, West Germany (2,5,6,35), East Germany (extinct), 
Greece, Italy, Liechtenstein (4), Netherlands, Norway, Poland (11), Romania, 
Spain (38), Sweden (25), Switzerland (9, 17) and the U.S.S.R. (Ukraine, Carpa- 
thians and Crimea) (21 , 22). The species has long been prized by collectors, who aim 
to possess as many of the different variants as possible . The trade therefore comprises 
a substantial value and volume (34). Rare or very limited subspecies may command 
impressive prices. There is disagreement over the importance of over-collecting in 
bringing about the observed declines. Collecting has been considered a potential 
threat in Finland given the current depleted condition of populations (26, 27). 
Over-collecting is reported to have occurred in Spain (34), Silesia (26, 27) and Italy, 

187 



Threatened Swallowtail Butterflies of the World 

where an effort was made to extirpate purposefully a rare, local race of Apollo in 
order to enhance the value of specimens already in hand (7). Market collecting has 
also been held partly responsible for the Polish decline of the species (12). None of 
these assertions contain numerical data. One lepidopterist, finding seven Apollos 
killed along 1 km of road, suggested that vehicles were a greater mortality factor than 
collectors (36), and vehicles on a motorway system near Bozen, South Tyrol (Italy) 
nearly wiped out a local race of Apollo (34) . However, most observers agree that it is 
not collectors or cars, but 'Tannensoldaten' (ranks of conifers in plantations) that 
threaten Europe's parnassians. Examples of afforestation interfering with Apollo 
survival have been cited in Bavaria (10, 33), Poland (12), Spain (20) and Switzerland 
(21). Succession of suitable habitat to scrubland is another recurring threat, for 
instance in Poland and Switzerland (21). Climatic change (Finland) and acid rain 
(Norway) are among purported causes of the widespread withdrawal of P. apollo 
from much of Fennoscandia (21), although agriculture has also been blamed in 
Finland (27) and the causes in Sweden are not properly understood (25). Urbani- 
zation has been given as another reason for decline in Finland and tourist 
development is imphcated in Bulgaria (21). On the whole, habitat change seems to be 
a far more important threat to Apollo than collecting. Whatever the major cause or 
causes, the effects are dramatic, at least locally: in France, two subspecies are 
Extinct, one Endangered and seven Vulnerable (16, 21); in East Germany, the insect 
is Extinct (21) and in Poland the decline is widespread and continuing: between 1950 
and 1965 three colonies disappeared from Pieniny National Park and many other 
Polish populations are considered Endangered (12-16, 30, 31, 32, 40). 

Conservation Measures Parnassius apollo was the first invertebrate to be 
included in Appendix II of the Convention on International Trade in Endangered 
Species of Wild Fauna and Flora (CITES), requiring monitoring of imports and 
exports to and from signatory states. Laws exist (21) at the national, regional or local 
level to protect P. apollo in the following countries: Austria, Czechoslovakia, 
Finland, France, Greece, Netherlands, Switzerland, West Germany (since the 19th 
century (34)) and Poland. However, these laws and regulations usually address 
protection of individuals (i.e. collecting restrictions) instead of habitats, and may 
have little effect. For example, despite legal protection accorded in 1952, the Apollo 
has continued to decline dramatically in Poland (11, 12, 30, 32). P. apollo does exist 
in a number of national parks and other protected areas, such as Tatra and Pieniny 
National Parks in Poland, but this does not always lead to its conservation (15) since 
specific management measures have seldom been taken. An attempt has been made 
to reintroduce P. apo/Zo into the Pieniny Mountains in Poland (16, 31); the results are 
not yet known. Publicity items have been produced stressing the need for conser- 
vation of P. apollo, such as a poster issued by the Federation Frangaise des Societes 
de Sciences Naturelles, depicting four of the threatened French races of the butterfly. 
Many authors have called for specific measures on behalf of P. apollo. In Poland 
the status of the butterfly inside and outside the national parks should be assessed in 
order to identify the factors detrimental to its survival, and the government should 
implement the aims of the 1952 law protecting the butterfly (15, 31). Reserves for the 
Apollo are needed in the mountains of Crimea (39). The Spanish Lepidoptera Red 
Data Book (38) offers suggestions for conserving the butterfly in the Pyrenees, and 
strong measures (including a new national park for this and other wildlife) are 
recommended in a later Spanish appraisal (20) . Artificial breeding of two generations 
per year (instead of the usual one) has been suggested as a means of reinforcing 
populations or enhancing reintroduction attempts in Scandinavia (28, 29). Reintro- 

188 



Parnassius apollo 

ductions should be attempted only with larvae or pupae, to avoid emigration of the 
adults (29). Past attempts to release butterflies have not succeeded. All countries 
where the species occurs should sponsor detailed surveys of Apollo colonies, and take 
steps to establish reserves and implement management measures where possible. 
Reforestation with conifers should be avoided and scrub succession should be 
arrested in important Apollo habitats. The provisions of CITES should be enforced, 
to determine levels of commercial trade in the butterfly and its possible impact on 
populations. Attention should also be given to Parnassius phoebus and P. mnemo- 
syne, which may also be threatened in Europe (21, 26, 27, 34). 

As a large , conspicuous and very attractive butterfly, the Apollo has a certain value 
as a tourist attraction in some alpine areas. 



References 

1. Ackery, P.R. (1975). A guide to the genera and species of Parnassiinae (Lepidoptera: 
Papilionidae). Bulletin of the British Museum (Natural History). Entomology 3\: 71-105. 

2. Anon. (1976). Rote Liste bedrohterTiere in Bayern (Wirbeltiere und Insekten). Schriften 
der Natur. Lans. Bayern 3: 1-12. 

3. Bernardi.G., Nguyen, T. and Nguyen, T.H. (1981). Inventaire, cartographic et protection 
des Lepidopteres en France. Beiheft Veroffenilichungen Naturschutz Landschaftspflege 
Baden-Wtirttemberg 21 : 59-66. 

4. Biedermann, J. (1982). Lebensraum fur Insekten. Liechtensteiner Umweltbericht June 
1982, 4-5. 

5. Blab, J. and Kudrna, O. (1982). Naturschutz Aktuell, HilfsprogrammfurSchmetterlinge. 
Kilda-Verlag, Greven. 135 pp. 

6. Blab, J., Nowak, E. and Trautmann, W. (1977). Rote Liste der Gefahrdeten Tiere und 
Pflanzen in der Bundesrepublik Deutschland. Naturschutz Aktuell 1: 1-67. 

7. Bourgogne, J . ( 1971 ). Un temoignage de plus sur la destruction de la nature (Papilionidae). 
Alexanor 7: 50. 

8. Bryk, F. (1935). Lepidoptera. Parnassiidae pars II (Subfam. Parnassiinae). Tierreich 65: Li 
+ 790 pp., 698 figs. 

9. Burckhardt, D., Gfeller, W. and Muller, H.U. (1980). Geschutzte Tiere der Schweiz. 
Schweiz. Bund, fiir Naturschutz, Basel. 223 pp. 

10. Christensen, G. (1975). Wer rottet aus? Entomologische Zeitschrift 85: 246-48. 

1 1 . Dabrowski, J.S. (1975). Some problems in the preservation of butterflies in Poland. Atala3: 
4-5. 

12. Dabrowski, J.S. (1980). The protection of the Lepidopterofauna - the latest trends and 
problems. Nota Lepidopterorum 3: 114-118. 

13. Dabrowski, J.S. (1980). O stanie zagrozenia lepidopterofauny w niektorych parkach 
narodowych Poiski. Wiadomosci Entomol. 1: 143-149. 

14. Dabrowski, J.S. (1980). Mizeni biotopu jasone cervenookeho — Parnassius apollo (L.) v 
Polsku a nutnost jeho aktivni ochrany (Lepidoptera, Papilionidae). Casopis Slezskeho 
Muzea Opava (A) 29: 181-185. 

15. Dabrowski, J.S. (1981). Remarks on the state of menacing the lepidopteran fauna in 
National Parks. Part 11 (general): the National Park in Tatra. Zeszyty Naukowe 
Uniwersytetu Jagiellonskiego, Prace Zool 27: 77-100. 

16. Dabrowski, J.S. and Palik, E. (1979j. Uwagi o stanie zagrozenia w parkach narodowych, 
Czesc 1 : Zmiany zachodzace we wspolczesnej lepidopterofaunie Pieninskiego Parku 
Narodowego, ze szczegolnym uwzglednieniem zanikania gatunku Parnassius apollo (L.), 
(Lepidoptera: Papilionidae), Dokumentacja n/t na zlec. Kom. Nauk.: 'Czlowiek i 
Srodowisko' PAN, 1-38 (maszynopis). 

17. Gfeller, W. (1975). Geschutzte Insekten in der Schweiz. Mitteilungen der Schweizerischen 
Entomologischen Gesellschaft 48: 217-213. 

18. Gomez Bustillo, M.R. andFernandez-Rubio, F. (1974). Mariposas de la Peninsula Iberica 
(tomo 1). Servicio de Publicaciones del Ministerio de Agricultura, Madrid. 198 pp. 

189 



Threatened Swallowtail Butterflies of the World 

19. Gomez Bustillo, M.R. and Fernandez-Rubio, F. ( 1974). Mariposas de la Peninsula Iberica 
(tomo 2). Servicio de Publicaciones del Ministerio de Agricultura, Madrid. 258 pp. 

20. Gomez-Bustillo, M.R. (1981). Protection of Lepidoptera in Spain. Beiheft Veroffentli- 
chungen Naturschutz Landschaftsflege Baden-Wiirttemberg 21: 67-72. 

21. Heath, J. (1981). Threatened rhopalocera (butterflies) in Europe. Council of Europe, 
Nature and Environment Series No. 23, 157 pp. 

22. Heath, J. and Leclerq, J. (Eds)(1981). European Invertebrate Survey. Provisional Atlas of 
the Invertebrates of Europe, Maps 1-27. Institute of Terrestrial Ecology, Monks Wood and 
Faculte des Sciences Agronomiques, Gembloux. 

23. Higgins, L.G. and Riley, N.D. (1980). A Field Guide to the Butterflies of Britain and 
Europe. 4th ed. Collins, London. 384 pp. 

24. Igarashi, S. ( 1979). Papilionidae and their early stages. 2 vols. , 219 pp. , 102 pi. Kodansho, 
Tokyo. (In Japanese). 

25. Janzon, L-A. and Bignert, A. (1979). Apollofjarilen i Sverige, Fauna Flora, Uppsala 74: 
57-66. 

26. Mikkola, K. (1979). Vanishing and declining species of Finnish Lepidoptera Notulae 
Entomologicae 59: 1-9. 

27. Mikkola, K. (1981). Extinct and vanishing Lepidoptera in Finland. Beiheft Veroffentli- 
chungen Naturschutz. Landschaftspflege Baden-Wiirttemberg 21: 19-22. 

28. Nikusch, I. (1981). Die Zucht von Parnassius apollo Linnaeus mit jahrlich zwei 
Generationen als Moglichkeit zur Erhaltung bedrohter Populationen. Beiheft Veroffentli- 
chungen Naturschutz Landschaftspflege Baden-Wiirttemberg 2\: 175-176. 

29. Nikusch, I. (1982). First trials to save threatened populations of Parnassius apollo by 
transplantation to new suitable biotypes. Third European Congress of Lepidopterology, 
Cambridge. In press. 

30. Palik, E. (1966). On the process of dying out of Parnassius apollo Linne. Tohoku Koncho 
Kenkyu 2: 45-47. 

31. Palik, E. (1980). The protection and reintroduction in Poland of Parnassius apollo 
(Linnaeus) (Papilionidae). Nota Lepidopterorum 2: 163-164. 

32. Palik, E. (1981). The conditions of increasing menace for the existence of certain 
Lepidoptera in Poland. Beiheft Veroffentlichungen Naturschutz Landschaftspflege Baden- 
Wiirttemberg 21: 31-33. 

33. Pfaff, G. (1935). Wer rottet aus? Entomologische Zeitschrift 49: 105-107. 

34. Pyle, R.M. (1976). The eco-geographic basis for Lepidoptera conservation. Part iii. A 
review ofworld Lepidoptera conservation. Yale Univ. PhD. Thesis, Publ. Univ. Microfilm 
Int. 369 pp. 

35. Rowland-Brown, H. (1913). Parnassius apollo in Germany. Entomologist 46: 289-290. 

36. Schmiedel, R. ( 1934). Tragt der sammler die schuld am Ruckgang unserer Insektenfauna? 
Entomologische Rundschau 51: 174—177. 

37. Svenson, I. (1981). Changes in the Lepidoptera fauna of Sweden after Linnaeus. Beiheft 
Veroffentlichungen Naturschutz. Landschaftspflege Baden-Wiirttemberg 21: 23-30. 

38. Viedma, M.G. de and Gomez-Bustillo, M.R. (1976). Libro Rojo de los Lepidopteros 
Ibericos. Instituto Nacional para la Conservacion de la Naturaleza, Madrid. 120 pp. 

39. Yermolenko, V.M. (1973). On protection of useful, relict and endemic insects of the 
Ukrainian Carpathian mountains and mountains of Crimea. In: On Insect Protection. 
Armenian Academy of Sciences Symposium, Yerevan. Pp. 29-35. 

40. Zukowski, R. (1959). Extinction and decrease of the butterfly Parnassius apollo on Polish 
territories. Sylwan 103: 15-30. 

This review has been adapted from The lUCN Invertebrate Red Data Book, whose 
authors and contributors are gratefully acknowledged. 



190 



Bhutanitis mansfieldi 
Bhutanitis mansfieldi ( Riley, 1939) RARE 

Subfamily PARNASSIINAE Tribe ZERYNTHIINI 



Summary Until recently Bhutanitis mansfieldi was known from only two 
female specimens and one male collected in 1918 from Yunnan Province, south- 
western China. In spring 1981 a team of Japanese mountaineers rediscovered this 
enigmatic and beautiful rarity on Mt Gonga in Sichuan. More information on its 
conservation status is needed. 

Description Bhutanitis mansfieldi was originally described from a single 
female specimen (4) (Plate 1.5). A second female was known to have been collected 
(2), but its whereabouts remained a mystery until March 1983, when it was 
rediscovered in the British Museum (Natural History) (3). In 1982 a Japanese insect 
dealer obtained a very battered male specimen, illustrated in (3). Also in 1982 
Saigusa and Lee described a second subspecies, pulchristata. from 1 1 males and three 
females taken in Sichuan (5). The sexes are similar, with a forewing length of about 
40 mm. 

UFW ground colour pale yellow with eight black, transverse bands (which extend 
into the UHW) including a broad outer margin. 

UHW elongated, with a scalloped edge, a deep notch at the anal angle and three 
tails, of which the outermost is long and club-shaped and the others shorter. There is a 
broad, black distal band, a large bright red patch extending from the anal margin, two 
blue eye-spots, and four pale yellow submarginai lunules (1,3, 4). 

LFW/LHW differs only slightly from the upperside. One of the discal spots is red, 
and the submarginai lunule between the first two tails is orange. 

Distribution The type specimen of this species was found by M.J. Mansfield 
in a collection of butterflies made by a British botanist, G. Forrest, who collected 
extensively in Yunnan, China. The specimen lacked data, but was assumed to have 
originated in that province. No further material was collected until spring 1981 when 
two members of a mountaineering party sent to Mt Gonga in Sichuan by the 
Hokkaido Alpine Association unexpectedly collected 14 specimens near Xinxing 
(2200 m a.s.l.) (5). A further party of Chinese collectors sponsored by a pool of 
Japanese entomologists was then sent to Xinxing in 1983, where they caught about 30 
specimens early in the season (6). Snow was still present in patches. In view of these 
records from Sichuan, there may be cause to question the assumption that Yunnan 
was the provenance of the type material . There are no records from Laos, Vietnam or 
Burma, which border onto Yunnan. 

Habitat and Ecology In common with Bhutanitis thaidina, B. mansfieldi 
may feed on species of Aristolochia (Aristolochiaceae), a genus of climbing vines. If 
this is so, the habitat may be in valley forests suitable for supporting the vine. 
However, there would seem to be every possibility that Bhutanitis may also have 
adapted to a more temperate alpine genus or family of plants, as have most of the 
Parnassiini. Luehdorfia, which is believed to be derived from Bhutanitis, uses 
Asarum as a foodplant, another genus within the Aristolochiaceae but with a more 
temperate distribution. There is no information on breeding biology or young stages, 
although it is known that the adults fly early in the season, before B. thaidina (6). 
There is a great need for further study of both these beautiful species. 

191 



Threatened Swallowtail Butterflies of the World 

Much of Yunnan and Sichuan is rugged, highland country. Yunnan is mainly a 
plateau region at an altitude of about 2000 m in south-western China while Sichuan, 
lying north of Yunnan, includes some lower-lying regions. The cHmate is temperate 
alpine, dry and bright in winter, wet in summer. The terrain is mountainous with 
fertile valleys. Despite being rich in minerals (particularly Yunnan), the region is so 
remote from administrative centres that it has remained under-developed and very 
poorly known to both local and foreign entomologists. 

Threats Now that a precise locality has been discovered there is clearly a 
need to plan and execute conservation measures. Any collecting in the region should 
be moderated and sustainable. There is no precise information on environmental 
threats in the region of Xinxing, nor on the provision for protected areas nearby. 
More information is needed. 

Conservation Measures No measures have been taken so far, although the 
rediscovery of the butterfly by Japanese entomologists is a great step forward. The 
Japanese have shown that they have the resources and the lines of communication to 
find the butterfly, they should also be encouraged to allocate further resources to 
ensuring the long-term survival and protection of this species. 

References 

1. Ackery, P.R. (1975). A guide to the genera and species of Parnassiinae (Lepidoptera: 
Papilionidae). Bulletin of the British Museum (Natural History). Entomology 31: 71-105. 

2. Mansfield, M.J. (1941). Bhutanitis mansfteldi Riley not unique. Entomologist 74: 44. 

3. Okano, K. (1984). Color illustration of Bhutanitis mansfieldi (Riley, 1940) (Papilionidae): 
with some notes on the same species. Tokurana (Acta Rhopalocera) 6/7: 61-65. 

4. Riley, N.D. (1939). A new species oiArmandia (Lep. Papilionidae). The Entomologist 72: 
206-208, 267. 

5. Saigusa, T. and Lee, C. (1982). A rare papilionid butterfly Bhutanitis mansfieldi (Riley), its 
rediscovery, new subspecies and phylogenetic position. Tyo to Ga 33: 1-24. 

6. Turlin, B. (1984). In litt., 1 June. 



192 



Bhutanitis thaidina 
Bhutanitis thaidina Blanchard, 1871 RARE 

Subfamily PARNASSIINAE Tribe ZERYNTHIINI 



Summary Bhutanitis thaidina has a fairly wide distribution in south- 
western China and Tibet, but was until recently known from very few specimens. 
Further documentation of the precise localities of the known populations, their 
exploitation by local collectors, and the life history of this beautiful species is needed. 

Description Bhutanitis thaidina is a medium-sized three-tailed butterfly 
with a forewing length of 42-48 mm. The sexes are similar in appearance, the female 
slightly larger than the male (1, 3). 

UFW yellow ground colour reduced to eight transverse lines separating broad 
bronze-black bands from the costal margin to the inner margin (1). 

UHW elongated with a scalloped outer margin and three tails of which the outer is 
long and club-shaped and the others shorter. The dark forewing bands continue into 
the discal region of the hindwing as streaks and spots. There is a broad, black distal 
band with a large red patch, four orange-yellow submarginal lunules and three Hght 
blue postdiscal spots (1). 

LFW/LHW paler than the upperside, with broader yellow lines and a smaller, 
paler red patch on the LHW. 

Distribution Bhutanitis thaidini occurs in south-western China where it has 
been recorded from the upper Jinsha Jiang (Yangtzekiang) River in Yunnan and 
Sichuan (Szechwan) provinces, from Kangding (Tatsienlu) in Sichuan, and localities 
in Shaanxi (Shensi) province (1,3,4). Records also exist for Mou-pin in Tibet 
(Xizang Zizhiqu) (2). 

Habitat and Ecology According to Bryk (3) the foodplant of Bhutanitis 
thaidina is Aristolochia sp. (Aristolochiaceae), a genus of scrambling vines. This 
requires confirmation from field studies. The genus Luehdorfia, derived from 
Bhutanitis, has changed its feeding habits to the more temperate aristolochiaceous 
genus Asarum. No further details are available on the breeding biology or habitat of 
the species. 

The range of this species is mainly high altitude plateaux (over 2000 m) with a 
rugged and highly dissected terrain. The natural vegetation would be variable, but 
the foodplants are likely to be found in forested country. 

Threats As in the case of Bhutanitis mansfieldi, B. thaidina is too poorly 
known for environmental threats to be properly documented. Highly priced 
specimens occasionally appear on dealers' lists, but no information is given on their 
precise sources. It has been reported that teams of Chinese collectors trained and 
sponsored by Japanese entomologists and dealers travelled to various parts of 
Sichuan during the collecting seasons of 1981, 1982 and 1983. Numerous specimens 
were taken near the town of Kangding, where the species is very local but common 
during the flight season (5). Collecting on this scale is most unlikely to pose an 
immediate threat to the butterfly, but there is a need to monitor the trade in order to 
ensure sustainable exploitation of the populations. 

Conservation Measures Unfortunately Tibet and south-western China are 
very difficult of access, but there is a great need for more research into the 

193 



Threatened Swallowtail Butterflies of the World 

Lepidoptera of these regions. It is now clear that the specimens rarely appearing in 
the market are freshly caught, probably through the arrangements of Japanese 
entomologists. The entomologists who have developed lines of access and communi- 
cation to these remote parts of China should be encouraged to ensure assessments of 
the impact of their collecting. It is apparent that Bhutanitis thaidina is fairly 
widespread and there is perhaps potential for a local industry in butterfly ranching 
along the Hnes of that in Papua New Guinea. Research into the breeding biology and 
management of Bhutanitis would be an essential preliminary step. There would 
certainly be a valuable market for this and many other Chinese species (see section 5 
of this book). It would be essential to monitor and regulate any trade in a sustainable 
manner, both to ensure the welfare of the butterfly and to safeguard the ranching 
programme itself. Now that populations are beginning to be located, the Chinese 
authorities might also ensure that adequate habitat is protected in national parks and 
reserves. There is a need for information on protected areas on this region. 

References 

1. Ackery, P.R. (1975). A guide to the genera and species of Parnassiinae (Lepidoptera: 
Papilionidae). Bulletin of the British Museum (Natural History). Entomology 31: 71-105. 

2. Bouchard, M.E. (1871). Remarques sur la faune de la principaute thibetaine du Mou-pin. 
Comptes Rendus Hebdomadaires des Seances de L'Academie de Science. Paris 72: 807-813. 

3. Bryk, F. (1935). Lepidoptera. Parnassiidae pars II (Subfam. Parnassiinae). Tierreich 65: Li 
+ 790 pp., 698 figs. 

4. Pen, D. (1936). The Papilionidae of south-western Szechwan. Journal of the West China 
Border Research Society 8: 153-165. 

5. Turlin, B. (1984). In litt., 1 June. 



194 



Bhutanitis ludlowi 
Bhutanitisludlowi Gabriel, 1942 INSUFFICIENTLY KNOWN 

Subfamily PARNASSIINAE Tribe ZERYNTHIINI 



Summary Bhutanitis ludlowi is known only from the type series, three 
males and one female collected in the Trashiyangsi Valley in Bhutan in 1933 and 1934. 
Its presence and status there need to be ascertained. 

Description Bhutanitis ludlowi is as beautiful as B. mansfieldi and B. 
thaidina, as well as being rather larger, with a forewing length of about 59 mm (male) 
or 63 mm (female). The sexes are similar (1, 3) (Plate 1.6). 

UFW ground colour dull black crossed by eight transverse, off-white lines. 

UHW elongated with scalloped outer margin and three tails, a long, spatulate, 
outermost tail, and two smaller ones. The transverse lines continue from the UFW, 
and a large velvety black and red patch, four dark grey lunules, and three large, 
blue-grey spots are present. 

LFW/LHW with more pronounced transverse lines and yellow-ochre submarginal 
lunules (1, 3). 

Distribution Bhutanitis ludlowi has only been recorded from the 
Trashiyangsi Valley in north-eastern Bhutan ( 1 ). The type series of three males and 
two females is unique. No other specimens have been found since the original 
collections in 1933 and 1934 (3). 

Habitat and Ecology There is no published information on the habitat or 
ecology of Bhutanitis ludlowi. It may use Aristolochia as a foodplant, as does 
Bhutanitis thaidina (I), but its habitat may be too high for these vines. The specimens 
were collected at an altitude of 2300-2500 m (3). Information on the vegetation at this 
altitude is given in reference (2) and summarized in the review of Teinopalpus 
imperialis. 

Threats No information is available specifically concerning the butterfly. 
Bhutan is certainly the best forested area in the Himalaya, but even here the resource 
is being eroded by unregulated grazing, forest fires and high consumption of wood for 
fuel and construction. Such degradation will ultimately affect all wildlife, including 
butterflies , and is extremely difficult to reverse in the Himalayan conditions of climate 
and topography. 

Conservation Measures The presence of Bhutanitis ludlowi in the 
Trashiyangsi Valley needs to be confirmed. Appropriate measures for its conser- 
vation and protection may then be developed. Bhutan has a good conservation 
record, with about 20 per cent (9213 sq. km) of its area protected as wildlife 
sanctuaries and reserved forests. The giant Jigme Dorji wildlife sanctuary 
(7900 sq. km) extends along the northern border of Bhutan and may well include B. 
ludlowi habitat within its boundaries. A butterfly survey of the sanctuary would be 
extremely valuable. 

References 

1. Ackery, P.R. (1975). A guide to the genera and species of Parnassiinae (Lepidoptera: 
Papilionidae). Bulletin of the British Museum (Natural History). Entomology 31: 71-105. 

195 



Threatened Swallowtail Butterflies of the World 

2. FAO/UNEP (1981). Tropical Forest Resources Assessment Project. Forest Resources of 
Tropical Asia. FAO, Rome. 475 pp. 

3. Gabriel, A.G. (1942). A new species of Bhutanitis (Lep. Papilionidae). Entomologist 75: 
189. 



196 



Luehdorfia chinensis 
Luehdorfia chinensis Leech, 1893 INSUFFICIENTLY KNOWN 

Subfamily PARNASSIINAE Tribe ZERYNTHIINI 



Summary Luehdorfia chinensis is a poorly known species restricted to 
certain eastern provinces in China. Its taxonomic status is uncertain and its 
conservation status is obscured by the inaccessibility of its habitat and the lack of 
information on its biology. 

Description The taxonomic status of Luehdorfia chinensis is not entirely 
certain. It was not listed by Munroe (6), but has since been variously treated as a 
separate species (3,4), and as a subspecies of L. japonica (1) or L. puziloi (2). 
Recently however, the young stages have been described and the species is now 
generally considered to be distinct (5). L. chinensis is similar to L. japonica in 
appearance (see next review), but differs in having a red submarginal band and 
yellow marginal spots on the UHW. Igarashi (1) gives two subspecies, chinensis and 
lenzeni. 

Distribution Luehdorfia chinensis has a restricted distribution in the 
provinces of Anhui (Anhwei), Hubei (Hupeh), Jiangsu (Kiangsu), and Jiangxi 
(Kiangsi) in eastern China (1). 

Habitat and Ecology The foodplant of Luehdorfia is Asarum (Aristolo- 
chiaceae), a genus of about 100 species of perennial herbs with vestigial or no petals to 
the flowers. Further details of the young stages and breeding biology are given by Lee 
(5) and Igarashi (4) (in Chinese and Japanese respectively). 

Threats There is no information on threats to this poorly known species. 
However, the eastern lowlands of China are highly developed and support high 
human populations . If the habitat is open deciduous woodland , as it is for L. japonica 
(see next review), there is cause for concern because the level of destruction of such 
formations is likely to have been high. Listing as Insufficiently Known is appropriate 
until further details emerge. 

Conservation Measures More details are required concerning the habitat, 
biology and conservation status of Luehdorfia chinensis. 

References 

1. Ackery, P.R. (1975). A guide to the genera and species of Parnassiinae (Lepidoptera: 
Papilionidae). Bulletin of ihe British Museum (Natural History). Entomology 31: 71-105. 

2. Bryk, F. (1935). Lepidoptera. Parnassiidae pars II (Subfam. Parnassiinae). Tierreich 65: Li 
+ 790 pp., 698 figs. 

3. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

4. Igarashi, S. (1979). Papilionidae and Their Early Stages. Vol 1: 219 pp.. Vol 2: 102 pp. of 
plates. Kodansho, Tokyo. (In Japanese). 

5. Lee, C.-L. ( 1978). The early stages of Chinese Rhopalocera — Luehdorfia chinensis Leech 
(Parnassiidae: Zerynthiinae). Acta Entomologica Sinensis 21: 161-163. (In Chinese). 

6. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supp\ement 17: 1-51. 

197 



Threatened Swallowtail Butterflies of the World 

LuebdorRa japonica Leech, 1889 VULNERABLE 

Subfamily PARNASSIINAE Tribe ZERYNTHIINI 



Summary Luehdorfia japonica is found in open, deciduous woodland in 
western Honshu, Japan. Suitable habitat is disappearing through changes in 
management and deforestation for development. Research on the management of 
the butterfly is needed, together with identification of appropriate reserves and 
protected areas. 

Description Luehdorfia japonica is a relatively small butterfly with a 
forewing length of 30-35 mm, the female slightly larger than the male (8, 9) (Plate 
1.2). The sexes are otherwise alike. 

UFW pale yellow with a black base and outer margin, and black bands across the 
wing (1,5, 8, 9). 

UHW also pale yellow with black bands, a medium length tail, a deep notch at the 
anal angle, and a scalloped outer margin. The outer third of the wing is black with 
orange submarginal patches and blue postdiscal spots. There is a small, blue anal 
eye-spot with a large, crimson patch above it (1, 5, 8, 9). 

LFW/LHW virtually the same as the upper, differing only on the hindwing. The 
crimson patch near the anal angle is extended as a band to the costal margin, and the 
orange submarginal patches are also merged into a continuous band (1, 5, 8). 

The young stages have been described by Igarashi (5) and Shirozu and Hara (11) 
(in Japanese). 

Distribution Luehdorfia japonica is restricted to the western part of 
Honshu (3), the main island of Japan. The northern limit is Yuri in Akita Prefecture, 
the eastern limit is the Tama Hills in the vicinity of Tokyo and the western limit is 
around Hagi city in Yamaguchi Prefecture (8). The subspecies, L. j. formosana 
Rothschild, 1918, listed by Shirozu (10), appears to be either a doubtful record or is 
now extinct. L. puziloi also flies in Japan, but the two species are almost entirely 
separated. Where they do occur together, in Akita, Yamagata, Nagano, Niigata and 
Yamanashi Prefectures, hybrids are occasionally found (8). These areas are known as 
the ''Luehdorfia line". 

Habitat and Ecology In Japan at least, the habitat oi Luehdorfia japonica is 
open formations of deciduous broad-leaved forest at middle altitudes (2). Such forest 
is a succession stage towards dense, climax forest with a closed canopy and is 
therefore naturally unstable (2). The climax forest is unsuitable for the butterfly. 

The foodplants are small perennial herbs from the family Aristolochiaceae, 
Asarum nipponicum, A. tamaense, A. blumei, A. caulescens and /I. sieboldi (1,5). 
The aduh butterflies emerge early in the spring (March in the west, April generally 
and May/June in mountainous or northern areas) and seek nectar in the flowers of 
Erythronium japonicum and Viola. The eggs hatch within several days of laying (4). 
The larvae feed on Asarum and grow rapidly, changing into pupae at the end of May 
or beginning of June (4). The pupae may be found attached to the stems of shrubs or 
herbs near the ground (8). They pass about ten months in diapause (4). 

The pupal diapause in Luehdorfia japonica has been the subject of research 
programmes by Japanese teams and is now known to be complicated. Two types of 
diapause seem to be experienced in one generation. One is summer diapause 

198 



*" Luehdorfia japonica 

maintained by the long day and high temperatures in summer and broken by the short 
day (below 15 hours) and moderate temperatures (14-25°C) in autumn; the other is 
winter diapause terminated by the winter cold (4, 6, 7). Consequently, the differenti- 
ation of the adult resumes in autumn and continues at a low rate during the winter, 
reaching completion in spring. 

Threats The deciduous broadleaved forests of Japan were formerly man- 
aged as a source of timber and charcoal and the constant thinning encouraged 
Luehdorfia japonica (2). In recent times however, the forests are not used as a 
resource . They are either left to reach dense climax woodland or else they are cleared 
for residential developments and amenities such as golf courses. The resulting 
depletion of suitable habitat for L. japonica is causing continuous reductions in its 
range. 

Conservation Measures There are still many good localities for Luehdorfia 
japonica in western Honshu, but the constant and accelerating loss of its habitat 
requires conservation action. There are no permanent reserves for the butterfly and 
suitable localities should be identified and surveyed. Once protected, habitats will 
need to be carefully managed in order to maintain suitable conditions. A research 
programme into the management of the butterfly is desirable. The butterfly is not 
difficult to rear artificially and suitable localities could be kept well stocked. Pupae 
are already occasionally advertised for sale. More data on the habitat and biology of 
Luehdorfia japonica are needed, but the programme of research into its pupal 
diapause has given a good basis for future management studies. 

References 

1. Ackery, P.R. (1975). A guide to the genera and species of Parnassiinae (Lepidoptera: 
Papilionidae). Bulletin of the British Museum (Natural History). Entomology 31: 71-105. 

2. Ae, S.A (1983). In lilt., March 18. 

3. Bryk.F. (1935). Lepidoptera. ParnassiidaeparsII (Subfam. Parnassiinae). TierreichbS: Li 
+ 790 pp. 

4. Hidaka, T., Ishizuka, Y. and Sakagami, Y. (1971). Control of pupal diapause and adult 
differentiation in a univoltine papilionid butterfly, Luehdorfia japonica. Journal of Insect 
Physiology 17: 197-203. 

5. Igarashi. S. (1979). Papilionidae and Their Early Stages. Vol 1: 219 pp., Vol 2: 102 pp. of 
plates. Kodansho, Tokyo. (In Japanese). 

6. Ishii, M. and Hidaka, T. (1982). Characteristics of pupal diapause in the univoltine 
papilionid Luehdorfia japonica (Lepidoptera, Papilionidae). Kontyu 50: 610-620. 

7. Ishii, M. and Hidaka, T. (1983). The second pupal diapause in the univoltine papilionid, 
Luehdorfia japonica (Lepidoptera: Papilionidae) and its terminating factor. Applied 
Entomology and Zoology 18: 456-463. 

8. Kawazoe, A. and Wakabayashi, M. (1976). Colored Illustrations of the Butterflies of Japan. 
Hoikusha, Osaka, Japan. 422 pp. 

9. Leech, J.H. (1889). Description of a new Luehdorfia from Japan. The Entomologist 
22(309): 25-27. 

10. Shirozu, T. (1960). Butterflies of Formosa in Colour. Hoikusha. Osaka, Japan (In 
Japanese). 483 pp. 

1 1 . Shirozu, T. and Hara, A. ( 1960). Early Stages of Japanese Butterflies in Colour. Hoikusha, 
Osaka. Vol. 1, 142 pp. (In Japanese). 



199 



Threatened Swallowtail Butterflies of the World 

Teinopalpus imperialis Hope , 1843 RARE 

Subfamily PAPILIONINAE Tribe LEPTOCIRCINI 



Summary Teinopalpus imperialis, the Kaiser-I-Hind, is a superb green, 
black and orange, tailed butterfly from the forests of the Himalaya, Burma and 
China. Although widely distributed, it is very local and rare. Some of the forests still 
retain their original character, but human population pressure is increasing their 
degradation through fire, deforestation, grazing and fuelwood collecting. As well as 
supporting unique wildlife , the forests play a vital role in controlling the flow of water 
to the lowlands. 

Description Teinopalpus imperialis, the Kaiser-I-Hind, is a large and 
beautiful, tailed butterfly with sexes that differ in size and coloration and were 
originally described as two separate species (Plate 1.7). There are two (or perhaps 
three) subspecies. The wingspan of the nominate subspecies is 90-120 mm, (male 
forewing length 52 mm, female 60-65 mm); the female T. i. imperatrix de Niceville, 
is even larger with a wingspan of 130 mm , a forewing length of 70 mm and five distinct 
tails (2, 4, 5, 17). Teinopalpus behludinii (15) from Sichuan is almost certainly 
referable to T. imperialis, but the type specimen is apparently untraceable (12, 15) 
and no other specimens are known. New material is needed to establish behludinii as 
a good species. Until this is forthcoming it may be assumed that behludinii is a 
subspecies or population of imperialis. 

Male: UFW with a black ground colour densely suffused with bands of green 
scales. UHW scalloped, with a long golden-tipped tail. The green bands of the UFW 
continue, but the distal region has a bright orange patch and a series of submarginal 
lunules in orange and bright green. 

LFW basal area densely covered with green scales, the distal two-thirds yellow- 
brown with narrow black bands. LHW similar to UHW but with broader yellow 
markings (2,4,5, 17). 

Female: UFW basal area resembling the male but the outer two-thirds of the wing 
has broad, indistinct alternate grey and green bands. UHW deeply scalloped with two 
long black tails and a shorter one between them . The yellow discal patch of the male is 
replaced by a much larger dark grey patch, with yellow markings beneath. 

LFW/LHW similar to the male but with the yellow-brown colour replaced by grey 
and with a broad ochreous submarginal band on the LFW (2, 4, 5, 17). 

Distribution Teinopalpus imperialis is found at high altitudes from Nepal to 
southern Burma and in the Chinese provinces of Sichuan (as T. behludinii) and Hubei 
(11). As Mell (11) suggested, records from Yunnan are to be expected. The records 
from Guangdong are referable to T. aureus (see next review). T.i. imperialis occurs 
between 2000 and 3500 m in Nepal, across north-eastern India and Bhutan to 
northern Burma and at lower altitudes (over 1000 m) in China (11). In India it has 
been recorded from Sikkim, the Darjeeling region of northern West Bengal, the 
Khasia Hills, north of Cherrapunji in Meghalaya, and the states of Assam and 
Manipur in the north-east. It may also prove to be present in Arunachal Pradesh and 
Nagaland, completing the circle around the headwaters of the Brahmaputra River. 
T.i. imperatrix occurs only in Burma, where its distribution records indicate an 
extension to lower altitudes than the nominate subspecies. It is known from the Shan 
States in the north, southwards to the Ataran River in Karen State, and Mergui in 

200 



Teinopalpus imperialis 

Upper Tenasserim State (2,4,5,17). Although there have been no pubhshed 
records of Teinopalpus imperialis in the highlands of western Thailand, its presence 
there is a possibility. In China Teinopalpus imperialis is possibly quite widespread, 
perhaps occurring in the montane plateau of Yunnan and Sichuan, eastward to the 
highland parts of Hubei (11). Unfortunately there have been no reports from China 
for almost 50 years (11, 15). 

Habitat and Ecology The Kaiser-I-Hind is restricted to mountainous 
wooded districts, where it flies high up in the canopy ( 17) . It is said to descend nearer 
the ground from 8.00 to 11.00 a.m. on sunny days, when it can be attracted by 
baiting (17). Its flight is very rapid, particularly in the male, and the best localities for 
observation are open places and mountain tops surrounded by forests (10). The 
butterflies fly up through the forest to the hilltops in search of a mate. Often the 
males will defend mating territories on the summits, fighting off other suitors. In 
cloudy or misty weather the butterflies rest on low bushes, concealed by the ventral 
camouflage of the wings (8). 

Adults have not been seen to visit either flowers or water (8). Okano (14) gives 
details of the foodplants and young stages of Teinopalpus (in Japanese). The 
caterpillar is green with a large head and the pupa is smooth, green and horned 
(13, 17). Larvae pupate at the end of September and adults emerge in April-May in 
Sikkim, May-July in Manipur (8). There are reports that the species is double- 
brooded, at least in India (18). The foodplants are Daphne spp., laurel-like small 
trees or shrubs in the Thymeleaceae (8, 13, 14). The habitat in which Teinopalpus 
imperialis flies is mainly broad-leaved evergreen forest, sometimes mixed with 
conifers and often well-stocked with various species of oak {Quercus) (3, 16). The 
foodplants are generally a component of the understorey. 

Threats The butterfly is very local and is never reported to be abundant, 
although it is fairly widespread (17). For its survival it appears to be dependent upon 
the natural forests that harbour its foodplant Daphne in the Himalaya, Burma and 
China. There is no evidence of its spread into degraded forest, the regenerative 
stages of shifting cultivation, or agricultural land. Nothing has been published 
concerning threats specific to the butterfly, but there is a growing literature describ- 
ing the disturbing extent of deforestation over the entire area of the butterfly's 
range. 

In Nepal deforestation has largely resulted from rising human population and 
migration into the Terai, or southern lowland strip, where broad-leaved forests 
grow. Loss of these lowland forests is running at an estimated 80 000 ha per year, 
almost 5 per cent of the 1980 total forest cover (6). The migrants come from the hill 
areas, which are more difficult to cultivate. Virtually all of central Nepal is now 
deforested although certain spots known to contain Teinopalpus, such as Phulchoki 
mountain, still remain wooded. 

In India the Himalayan forests are possibly being deforested more slowly than in 
Nepal, although the available data are acknowledged as being unreliable (6). Few 
data are available for specific regions, but the isolated montane area of Meghalaya 
seems to be particularly seriously deforested. These hills have a very high rainfall 
and Cherrapunji, with an average rainfall of 1150 cm per year, is one of the wettest 
places on earth (1). The traditional form of agriculture was shifting cultivation, or 
'jhumming', but over-exploitation has led to severe soil erosion (1). Such degrada- 
tion in the lower altitude regions has undoubtedly led to more severe environmental 
pressures and forest degradation in the high altitude central core of Meghalaya. In 

201 



Threatened Swallowtail Butterflies of the World 

Darjeeling the forests are reported to be virtually all gone and in Sikkim also large 
areas are denuded and eroded (7). 

In Bhutan the forests are said to be subjected to heavy biotic interference, 
particularly grazing and fires (6). In Burma deforestation is relatively limited, but 
again the main pressure on forests is rising human population and the increased need 
for fuelwood, grazing and land for shifting cultivation (6). Few data are available for 
the Chinese part of the butterfly's range , but specimens have been recorded as low as 
1000-1400 m (11), where habitat destruction is likely to have been severe. 

Information on the degradation of the Himalayan forests is scarce and often out of 
date. It is thus difficult to obtain an accurate impression of human impact on the 
butterfly's habitat, but the indications are that its range must be shrinking rapidly. As 
well as supporting important flora and fauna, not to mention human communities, 
these forests have important cHmatic and environmental influences. Most of the 
important rivers of northern India originate in the Himalaya. The huge River 
Brahmaputra has its headwaters in the habitat of Teinopalpiis imperialis. Biotic 
interference is often serious in the lower zones of hardwood forest, declining higher 
up. Fires may be devastating at any altitude, and grazing and fuelwood collection are 
on the increase. In general most of the high altitude coniferous forests seem to have 
retained their original character, particularly in the Himalaya, but increasing 
population pressure may eventually lead to intensified use of the land (6). In this 
eventuality great skill will be needed if environmental degradation is to be avoided. 

Despite the wide range of T. imperialis, some of the reports of over-collecting must 
be a matter for concern. For example, in July 1984 on Phulchoki (Pulchoki) mountain 
near Kathmandu in Nepal, every specimen flying in was being caught by foreign 
collectors (7). Although such an isolated event is unlikely to have a serious impact on 
the butterfly species, other hills may be receiving the same treatment. Moreover, in 
both Nepal and India collecting is illegal without a licence. Excessive collecting in this 
way reduces the natural beauty of the area. 

Conservation Measures Specific measures for the Kaiser-I-Hind are diffi- 
cult to propose at this stage. Collecting could pose a problem in certain localized areas 
and the legislation already enacted in India and Nepal should be enforced more 
effectively. The best way to preserve the Kaiser-I-Hind is rational utilization and 
protection of the forests in which it lives. The Government of Nepal is preparing a 
National Conservation Strategy that will be of benefit to all wildlife (9) . In India such 
strategies would have to be on a state-wide basis, since each state is virtually 
autonomous in its forestry practice (6). The presence of Teinopalpus imperialis in 
protected areas should be noted and research towards management measures 
encouraged. 



References 

1. Agarwal, A., Chopra, R. and Sharma, K., Eds. (1982). The State of India's Environment 
1982. A Citizens' Report. Centre for Science and Environment, New Delhi. 192 pp. 

2. Antram, C.B. (1924). Butterflies of India. Thacker, Spink & Co., Calcutta and Simla. 226 
pp. 

3. Champion, H.G. and Seth, S.K. (1968). A Revised Study of the Forest Types of India. 
Publication Division, Government of India, Delhi. 

4. D Abrera, B. (1982). Butterflies of the Oriental Region. Part 1. Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi + 244 pp. 

5. Evans, W.H. (1932). The Identification of Indian Butterflies. Bombay Natural History 
Society. 2nd ed., revised. 454 pp. 

202 



*" Teinopalpus imperialis 

6. FAO/UNEP (1981). Tropical Forest Resources Assessment Project. Forest Resources of 
Tropical Asia. FAO, Rome. 475 pp. 

7. Feeny, P. (1984). Pers. comm.. 1 August. 

8. Ghosh, S.K. and Mandal, D.K. (1983). Review of Kaiser-I-Hind butterfly, Teinopalpus 
imperialis Hope. Unpublished manuscript. 3 pp. 

9. His Majesty's Government of Nepal/IUCN (1983). National Conservation Strategy for 
Nepal. A Prospectus. lUCN, Gland. 36 pp. 

10. Jordan, K. (1909). In Seitz, A. Macrolepidoptera Fauna Indo-Australasia 9: 17-109. 

11. Mell, R. (1938). Beitrage zur Fauna Sinica. Deutsche Entomologische Zeiischrift 17: 
197-345. 

12. Moonen, J.J.M. (1984). Notes on eastern Papilionidae. Papilio International 1(3); 47-50. 

13. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supp\emem 17: 1-51. 

14. Okano, K. (1983). Some ecological notes on Teinopalpus. Tokurana 5: 94-100. (In 
Japanese). 

15. Pen, D. (1936). The Papilionidae of south-western Szechwan. Journal of the West China 
Border Research Society 8: 153-165. 

16. Stainton, J.D.A. (1972). Forests of Nepal. John Murray, London. 181 pp. 

17. Talbot, G. (1939). The fauna of British India, including Ceylon and Burma. Butterflies voU. 
Taylor and Francis Ltd., London, reprint New Delhi 1975. 600 pp. 

18. Wankhar, D.M. (1984). In litl., 11 April. 



203 



Threatened Swallowtail Butterflies of the World 

Teinopalpus aureus Mell, 1923 INSUFFICIENTLY KNOWN 

Subfamily PAFILIONINAE Tribe LEPTOCIRCINI 



Summary Teinopalpus aureus, sometimes known as the Golden Kaiser-I- 
Hind, is a superb green and golden tailed butterfly from China. No specimens have 
been seen for many years and there is a great need for more information on the 
distribution, ecology and conservation status of the species. 

Description Teinopalpus aureus is a large and beautiful butterfly with a 
forewing length of 45-55 mm . Its taxonomic status is uncertain because the butterfly is 
so poorly known. It was described as a subspecies of T. imperialis (3), but Munroe (4) 
and Hancock (2) list it as a full species. The female is unknown. 

UFW black with dense, bright green scaling and a black band with a bright 
yellow-green distal border. UHW with a large, golden-yellow discal patch, concen- 
tric bands of black-blue, orange and green, and a similar long, yellow-tipped tail. The 
underside is similar, but somewhat paler (1). 

Distribution There is very little information on the distribution of T. aureus. 
The type locality is in south-east China where it has been recorded from montane 
forests near Lianping in the north of Guangdong (Kwangtung) province (3). In 
addition, there is a colour photograph of this species in the British Museum (Natural 
History) labelled "Haut-Donnai, Annam" , suggesting it may also be found in Vietnam 

Habitat and Ecology Teinopalpus aureus is a montane species, probably 
flying in habitats above 1000 m, in a region that would once have had broad-leaved 
and coniferous evergreen forests at such altitudes. The species is so poorly known that 
there are no published details of its present habitat, breeding biology or foodplant 
requirements. However, it is known to fly in late March and April (3). The closely- 
related Teinopalpus imperialis (see previous review) feeds on Daphne, a laurel-like 
small tree or shrub in the Thymeleaceae. 

Threats The extent of montane land in the vicinity of Lianping, the locality 
for the specimens in the British Museum (Natural History), appears from maps to be 
very limited. In all probability the species is very rare but fairly widespread, although 
it seems strange that such a striking butterfly has not been recorded more often. If, Hke 
Teinopalpus imperialis, T. aureus flies in montane evergreen forest, then its survival in 
southern China is likely to be severely imperilled by the extent of deforestation there. 

Conservation Measures Teinopalpus aureus is extremely poorly known , but 
is likely to be threatened by deforestation of its montane habitats. It may already be 
extinct and searches are needed to establish its present distribution. Once found, 
protected areas and ecological study will be needed to ensure its future survival. 

References 

1. D'Abrera, B. (1982). Butterflies of the Oriental Region. Pari 1. Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi + 244 pp. 

2. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

204 



Teinopalpiis aureus 

3. Mell, R. (1938). Beitrage zur Fauna Sinica. Deutsche Entomologische Zeitschrift 17: 
197-345. 

4. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supplement 17: 1-51. 



205 



Threatened Swallowtail Butterflies of the World 

Eurytides (Protesilaus) marcellinus {Douhleday, 1845) VULNERABLE 

Subfamily PAPILIONINAE Tribe LEPTOCIRCINI 



Summary Eiiry tides marcellinus, the Jamaican Kite Swallowtail, is ende- 
mic to Jamaica and participates in spectacular migrations across the island. Its 
populations fluctuate widely and its main stronghold is threatened by loss of the 
foodplant Oxandra lanceolata as a result of cultivation. 

Description This species is easily distinguished from other Jamaican species 
by its small size (forewing length 30-35 mm), long slender tails and pattern of 
longitudinal bands of black and delicate pale blue-green (1,5) (Plate 1.8). Its 
nomenclature has been somewhat changeable at both the specific (1) and generic 
levels (3, 4). The most recent revision places it in the revived genus Protesilaus, as 
part of the marcellus species group (1,3,4) but here we retain the name Eury tides. 
The sexes are beHeved to be similar (5). 

UFW/UHW black with bold pale blue-green stripes and submarginal spots and a 
bright red anal spot. 

LFW/LHW similar but with brown ground colour and a wide red stripe on the 
hindwings. Tails about 10 mm long (5, 6, 2). 

The eggs are spherical, pale brown when laid, becoming black as they approach 
hatching (7). The mature larva is blue-green dorsally and grey ventrally with an 
orange head; the pupa dark brown with no anterior prominences (7). 

Distribution Endemic to Jamaica, with a former range which includes the 
parishes of St Elisabeth in the south-west, St Andrew and Kingston, and St Thomas 
in the south-east (1). The species is still locally common at Roselle (probably 
synonymous with Rozelle Falls) in the south-east but is rare elsewhere (7). 

Habitat and Ecology Prior to 1968 the species was found only sporadically. 
In 1968 T.W. Turner found the main breeding colony at Roselle and raised specimens 
from eggs (1). The preferred habitat is a wooded area with the calciphile foodplant 
Oxandra lanceolata, a 10-15 m straight-trunked tree in the family Annonaceae, 
known locally as the Black lancewood or Okra. At least two broods of the buttefly 
occur each year, in May-July and September. Adults are abundant for only one week 
in May or June and are otherwise rare. Pupal diapause may occur. According to 
D'Abrera (2), Eury tides marcellinus is known to maintain very low populations for 
long periods, occasionally undergoing a population explosion and appearing all over 
the island. The males are strongly migratory and participate in spectacular westerly 
migrations in some years; the purpose of these movements is not clear. In the absence 
of the foodplant from most of the island, such mass movements are doomed to 
reproductive failure. 

Threats The breeding habitat is very restricted and is being severely 
deforested for cultivation. The larval foodplants are unable to survive the disturb- 
ance (7). The breeding ground is not officially protected. 

Conservation Measures No long-term conservation measures have been 
taken so far, but the scientific community in Jamaica has shown considerable concern 
and started a dialogue with the owners of the 2 sq. km breeding ground. About a 

206 



Eurytides marcellimis 

quarter of a hectare has recently been set aside as a reserve where it is hoped the larval 
foodplant can be preserved. An adjacent landowner is prepared to provide land for 
the foodplant to be planted and funds are urgently required for this project. Since the 
tree is straight-trunked, it may even prove to be a valuable plantation species. 

Further measures required include a survey of the distribution of Oxandra, 
identification of Oxandra to local landowners and a plea for help in protecting the 
trees, establishment of Oxandra seedlings in nearby properties in order to extend the 
range of the plant and the butterfly, and an investigation of predators of the 
caterpillars and pupae with a view to preventing heavy losses (7) . The butterfly can be 
raised in captivity but the adults are virtually impossible to contain (7). 

The Natural Resources Conservation Department of Jamaica should be encou- 
raged to take some responsibility for the protection of Eurytides marcellimis, and to 
take steps to create permanently protected breeding grounds. 

References 

1. Brown, F.M. and Heinemann, B. (1972). Jamaica and its Butterflies. Classey, London. 
478 pp. 

2. D"Abrera,B. (1981). Butterflies of the Neotropical Region. Parti. Papilionidae and Pieridae . 
Lansdowne Editions, Melbourne, xvi + 172 pp. 

3. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersial: 1-48. 

4. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supp\cmQn\ 17: 1-51. 

5. Riley, N.D. ( 1975). A Field Guide to the Butterflies of the West Indies. Collins, London. 244 
pp. 

6. Rothschild, W. and Jordan, K. (1906). A revision of the American Papilios. Novitates 
Zoologicae 13: 411-752. (Facsimile edition ed. P.H. Arnaud, 1967). 

7. Turner, T.W, (1983). The status of the Papilionidae, Lepidopteraof Jamaica with evidence 
to support the need for conservation of Papilio homerus Fabricius and Eurytides marcellinus 
Doubleday. Unpublished report. 14 pp. 



207 



Threatened Swallowtail Butterflies of the World 

Eurytides (Protesilaus) lysithous harrisianus (Swainson, 1822) ENDANGERED 

Subfamily PAPILIONINAE Tribe LEPTOCIRCINI 

Summary Eurytides lysithous is a kite swallowtail which mimics various 
species of Parides swallowtails in different parts of its range . The strikingly patterned 
subspecies E. I. harrisianus, Harris' Mimic Swallowtail, resembles the Vulnerable 
species P. ascanius (also in this volume). Common in the nineteenth century, it was 
still found regularly around Rio de Janeiro, Brazil, until the 1940s. Nearly all known 
colonies have been destroyed by development and only a single known locality 
remains, in which the unpalatable model P. ascanius may already be extinct. E. I. 
harrisianus has recently been proposed to be added to Brazil's list of animals 
threatened with extinction. 

Description All members of Munroe's lysithous species-group are mimetic 
of Troidini or Heliconiinae (9). Recently, Hancock (7) (following D'Almeida (6)) 
has taken a new look at the group and reduced it to 15 species, renaming it after the 
most primitive member Eurytides asius. New data resulting from biological and 
morphological studies of the species indicate that still further taxonomic analysis at 
the subgeneric level is needed (2). Eurytides lysithous is a highly variable species (5) 
and E. I. harrisianus looks and behaves very much like Parides ascanius. It is a 
medium-sized black swallowtail with narrow tails and, in its commonest form 
(platydesma), both wings are crossed by a broad white band (Plate 2.1). The 
hindwing is variably marked with large red spots which appear similar to the rose-red 
patch on P. ascanius. Unlike the model, the mimic possesses a red streak at the base 
of the wings underneath. The antennae, tongue and tails are short for a swallowtail. 
The immature stages are identical to those of other subspecies of G. lysithous (3, 6). 
The taxonomic position of the genus is much disputed. Munroe (9) placed the 
lysithous species-group in the genus Eurytides subgenus Protesilaus. In a recent 
revision, Hancock has raised Protesilaus to generic status and transferred several 
Eurytides species-groups into this new genus (8). Other authorities consider 
Eurytides to be a synonym of Graphium, a name normally reserved for Old World 
species. Hancock has also treated E. chibcha, hipparchus and kumbachi as 
aberrations of other species and removed them from Munroe's lysithous species- 
group, renaming it the asius species-group. In addition, harrisianus has been 
considered to be a form (5) and even a full species (7), as well as a subspecies. These 
arguments do not affect the main matter under discussion here, i.e. the conservation 
of this highly endangered taxon. 

Distribution The species Eurytides lysithous occurs in Brazil and eastern 
Paraguay. The subspecies E. I. harrisianus formerly occurred in southern Espirito 
Santo and along the whole coast of the State of Rio de Janeiro, Brazil, but is now 
known only from Barra de Sao Joao , on the eastern coast of Rio de Janeiro. Potential 
habitat and strong colonies of the model species (P. ascanius) occur widely in 
lowland Rio de Janeiro, but no E. I. harrisianus have been seen in these sites since 
1945 (4, 6). In the single known locality, four specimens were collected between 1977 
and 1982 (4); the colony was strong and permitted extensive mark-recapture and 
biological work in 1984 (2). 

Habitat and Ecology Much is now known of the biology of the taxon. It flies 
in habitats adjacent to the lowland swamps occupied by its model, rarely in the 

208 



*" Eurytides lysithous harrisianus 

swamps themselves, and sometimes on adjacent hillsides. Males do not obviously 
frequent hilltops in search of mates, as many other Eurytides do, but are attracted to 
damp earth, where they were formerly often captured in Rio de Janeiro. Females are 
commonly encountered near their hostplants, which are various genera in the 
Annonaceae, especially Xylopia and Rollinia (2). The flight period is from 
September to February, with the pupal diapause lasting up to nine months (4, 6, 7). 
The adults fly rapidly and erratically along corridors in their dense, scrubby habitat, 
preferring areas with alternating large blocks of strong sun and deep shade. In the 
only known colony, two principal morphs are present, platydesina (mimicking 
Parides ascanius, about 60 per cent at present), and oedippus (mimicking P. 
zacynthus and P. neophilus, about 30 per cent at present) (2). The remaining 10 per 
cent are genetic recombinants, including typical harrisianus. These two morphs both 
fly from early morning (07.00 a.m. ) throughout the day. However, the latter tend to 
be more active when their models are out, in the early morning and very late 
afternoon, while the former fly with P. ascanius in the warmer hours. Flight activity is 
strongly correlated with sun and high humidity. Flowers commonly visited include 
Lantana, Inga and Eupatorium . The colony numbered 50-200 individuals in late 1984 
(2). 

Threats The site of the only known colony is undergoing development as a 
recreational area. There is to be some emphasis on wildlife conservation, but without 
special care the butterfly is likely to be lost in the general alteration of habitat (4, 7). 
Unusual climatic conditions in 1983 greatly reduced the only known colony (1). At 
the current level of disruption, extinction of this colony is likely by 1990 ( 10). With the 
disappearance of P. ascanius from the site due to development of subcoastal marshes, 
the platydesma morph may be negatively selected and become quite reduced in 
numbers. 

Conservation Measures In June 1982 G. /. harrisianus was proposed to the 
Brazilian Government agency responsible for inclusion on the official list of species 
threatened with extinction. Intensive searching is needed for any additional colonies 
which might occur in remote regions or in the Reserva Biologica P090 das Antas, Rio 
de Janeiro, where P. ascanius is still present to serve as a model. Officials responsible 
for the planning and management of the recreation area under construction at Barra 
de Sao Joao should be informed of the species' presence and petitioned to include 
measures for its conservation in plans for the area. If further populations can be 
found, urgent studies on the swallowtail's management ecology should be 
undertaken. 

As the only clear Batesian mimic of the vulnerable Parides ascanius, this 
population occupies a unique position for mimicry research. Possible further studies 
of the model/mimic pair include abundance ratios, association in the field (especially 
in light of the unusual situation of the model and mimic occupying slightly displaced 
microhabitats) and synchrony of generations (10). 

References 

1. Brown, K.S., Jr. (1983). In lift. 6 April. 

2. Brown, K.S., Fr. (1984). In litt. 20 November. 

3. Burmeister, A.M. (1879). Description Physique de la Republique Argentine Vol. V. 
(Lepidopteres), Atlas p. 9, No. 23. 

4. Callaghan, G., Laranja, J., Otero, L., Brown, K.S., Jr. (1982). Observations in 1940-1982 



I 



in Rio de Janeiro coastal lowlands. 



209 



Threatened Swallowtail Butterflies of the World 

5. D'Abrera, B. (1981). Butterflies of the Neotropical Region. Part 1. Papilionidae and 
Pieridae. Lansdowne Editions, Melbourne, xvi + 172 pp. 

6. D'Almeida, R.F. (1922). Melanges Lepidopterologiques, I. Eludes sur les Lepidopteres du 
Bresil. R. Friedlander und Sohn, Berlin. 

7. D'Almeida, R.F. (1966). Caidlogo dos Papilionidae Americanos . Sociedade Brasileira de 
Entomologia. Sao Paulo, Brazil. 

8. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersial: 1-48. 

9. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist 17: 1-51. 

10. Otero, L.S. and Brown, K.S., Jr. (in press). Biology and ecology of Parides ascanius 
(Cramer, 1775) (Lep., Papilionidae), a primitive butterfly threatened with extinction. 
Atala. 

This review has been extensively adapted from an entry in The lUCN Invertebrate 
Red Data Book, whose authors and contributors are gratefully acknowledged. 



210 



9(LprR B'y-~n-'^ err" '>^ Eurytides iphitas 

Eury tides (Eurytides) iphitas HiibneT, 1821 VULNERABLE 

Subfamily PAPILIONINAE Tribe LEPTOCIRCINI 



Summary Eurytides iphitas, the Yellow Kite, is a poorly known swallowtail 
from south-eastern and east-central Brazil. It has not been seen for many years and is 
believed to be in serious decline in the states of Rio de Janeiro and Espirito Santo. 

Description Eurytides iphitas is a relatively large, tailed butterfly (Plate 
2.3). The female is apparently unknown, but may be expected to resemble the male 
since the sexes of the close relative E. dolicaon are very similar. E. iphitas closely 
resembles E. dolicaon, but whereas in the former the long, narrow tails are only 
tipped with yellow, in the latter the tails are mainly yellow (2, 8, 9). 

UFW/UHW creamy-buff with a black submarginal border, outer margin and apex, 
oblique black bands on the forewing and a row of small white submarginal spots on 
the hindwing (2). 

LFW/LHW similar to the upperside, but with less black on the wing borders, more 
black scaling on the veins and more prominent white submarginal spots on the 
hindwing (2). 

The immature stages of E. iphitas have not been described, but are probably 
similar to those of E. dolicaon, in which the final instar caterpillar is yellow-green 
with a black band and black spotting (3). 

Distribution E. iphitas is endemic to south-eastern and east-central Brazil. 
It has not been seen for many years, even in a known colony locality at Boca do Mato 
near Rio de Janeiro (1). It has previously been recorded from further north in the 
state of Espirito Santo, but no localities are known there at present (1, 2, 4). 

Habitat and Ecology Very little information is available on this or other 
members of the dolicaon species-group (5,7). According to Hancock (5) the larvae of 
the group are smooth and found on Annonaceae or Lauraceae; the pupa is slender 
with a long dorsal protuberance. Eurytides iphitas occurs at medium elevations (2) 
probably in seasonally moist, open deciduous woodland. Eurytides dolicaon, a close 
relative of E. iphitas which also occurs in this part of Brazil, lays its eggs on the 
underside edges of leaves of Mespilodaphne indecora (Lauraceae) (3). li is a 
multivoltine species, with generations renewed every month during summer. After 
April the development of caterpillars is slow and over-wintering in the pupal stage 
usually occurs until mid-August (3). E. iphitas might be expected to have similar 
habits. 

Threats Habitat destruction in the south-eastern coastal states of Brazil is 
an increasingly serious threat to the wildlife. Urbanization, drainage and develop- 
ment will undoubtedly cause the decline of many species, as has been found with 
Eurytides lysithous harrisianus. There is insufficient information on the fauna of the 
several protected areas in the region (6) to know whether these measures have been 
effective. Eurytides iphitas is believed to be a relict species with a naturally small 
range being further reduced by destruction of suitable habitat. Its present rarity in 
regions close to human population centres is a matter for great concern. As the 
type-species of the genus, E. iphitas is of great importance to taxonomists. 

211 



Threatened Swallowtail Butterflies of the World 

Conservation Measures Searches for extant colonies are urgently needed in 
protected and unprotected areas, in order that data on habitat, ecology and present 
distribution may be gathered. Only then may its conservation status be properly 
determined and appropriate action taken. The situation of this butterfly could be 
extremely grave and no time should be lost in locating and protecting its present 
haunts. 



References 

1. Brown, K.S. Jr. (1983). In ////., 6 April. 

2. D'Abrera, B. (1981). Butterflies of the Neotropical Region. Parti. Papilionidae and Pieridae . 
Lansdowne Editions, Melbourne, xvi + 172pp. 

3. D'Aimeida,R.F. (1924). Lespapiiionidesde Rio de Janeiro. Description de deux chenilles. 
Annates de la Societe Entomologique de France 93: 23-30. 

4. D'Almeida, R.F. (1966). Catdlogo das Papilionidae Americanos. Sociedade Brasileira de 
Entomologia. Sao Paulo, Brazil. 

5. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

6. lUCN (1982). lUCN Directory of Neotropical Protected Areas. Tycooly International, 
Dublin. 436 pp. 

7. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
En/owo/ogwf Supplement 17: 1-51. 

8. Rothschild, W. and Jordan, K. (1906). A revision of the American Papilios. Novitates 
Zoologicae 13: 411-752. (Facsimile edition ed. P.H. Arnaud, 1967). 

9. Tyler, H.A. (1983). In /;//., 30 September. 



212 



Graphium epaminondas 
Graphium (Pathysa) epaminondas Oberthur, 1879 INSUFFICIENTLY KNOWN 
Subfamily PAPILIONINAE Tribe LEPTOCIRCINI 



Summary Graphium epaminondas is one of three swallowtails confined to 
the Andaman Islands. Intensification and expansion of cultivation in former forest 
and clear felling for timber are severely reducing the area of natural habitat. A 
conservation plan for the islands is needed, coupled with a survey of important 
butterfly habitats. 

Description Graphium epaminondas is a relatively large black, white and 
orange swordtail butterfly with a forewing length of 43 mm, and similar sexes (3). 

UFW with a brown ground colour and broad transverse white bands; LFW may be 
lighter but is otherwise similar in appearance (2,6). 

UHW with a slightly scalloped outer edge and a long, tapering dark brown tail with 
a white tip. The discal area is white with brown shadows from the LHW pattern (2,5). 
Postdiscal and submarginal areas are patterened with brown (2,6). There is an 
orange and black tornal spot. 

LHW with narrow black bands in the discal region and a postdiscal band of large, 
orange spots, small black lunules and a dark outer margin. 

Distribution Graphium epaminondas is confined to the Andaman Islands 
(India) where it may only occur on South Andaman . The islands are situated between 
India and Burma in the south-east of the Bay of Bengal. 

Habitat and Ecology Details of the life-history and foodplants of this 
species are unknown. Graphium epaminondas is in the antiphates species group, of 
which the larvae generally feed on Annonaceae or Lauraceae (4). G. epaminondas 
has sometimes been treated as a subspecies of antiphates (6), but is now generally 
accepted as a full species (2, 4). There are 13 species of Papilionidae recorded from 
the Andamans (1). In addition to G. epaminondas , Papilio mayo and Atrophaneura 
rhodifer are endemic to the islands. The natural vegetation of the islands is rain forest 
and all three butterflies are probably forest inhabitants. 

Threats The forest flora of the Andaman Islands is of Malesian origin, but 
has its own unique character and a high level of endemicity . A number of articles and 
reports describe the increasing rate of forest destruction in the Andamans and, in the 
absence of protected areas, there is good reason for conservation concern. The main 
threats are the intensification and expansion of cultivation, clear-felling for timber 
and planting of forest monocultures. One cause is the growing human population of 
the Andamans, which is being boosted at a rate of up to 1000 per month (in 1980) by 
new settlers from Bihar in India and by Karens from Burma (7). The heavy monsoon 
rains are apparently causing severe erosion of the thin soils in deforested areas, 
ultimately leading to degradation and loss of natural habitats. 

All three endemic papilionids are poorly known and very little has been published 
on their habitat and behaviour. A 1980 report states that Graphium epaminondas and 
Atrophaneura rhodifer are not rare, and that PapiUo mayo males are common but 
females rare (1,5). These assessments seem to be based mainly on numbers of 
museum specimens and an objective field assessment of conservation needs is 
required. 

213 



Threatened Swallowtail Butterflies of the World 

Conservation Measures At the moment there is too little information on 
Graphium epaminondas and the other endemic papilionids of the Andamans to 
propose conservation measures. The Botanical Survey of India has drafted proposals 
for a land use and conservation study of the Andamans, hopefully this will result in 
the development of protected areas. An entomological and zoological input into 
these studies might be encouraged. 

References 

1. Arori, G.S. and Nandi, D.N. (1980). On the butterfly fauna of Andaman and Nicobar 
Islands (India). I. Papilionidae. Records of the Zoological Survey of India 77: 141-151. 

2. D'Abrera, B. (1982). Butterflies of the Oriental Region. Part 1. Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi + 244 pp. 

3. Evans, W.H. (1932). The identification of Indian butterflies. Diocesan Press, Madras. 2nd 
ed., revised. 454 pp. + 31 plates. 

4. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

5. Haugum, J. (1983). In litt.,2 June 1983. 

6. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands. Vol. I 
Papilionidae. (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 

7. Wright, A. (1980). Destruction in the Andamans. Oryx 15: 315-316. 



214 



*" Graphium idaeoides 

Graphium (Pathysa) idaeoides Hewitson, 1853 RARE 

Subfamily PAPILIONINAE Tribe LEPTOCIRCINI 



Summary Graphium idaeoides is rare throughout its range in the Phihp- 
pine islands of Luzon, Samar, Leyte and Mindanao. It is a remarkably realistic mimic 
of Idea leuconoe (Danainae), which has a much wider distribution in South East 
Asia. The main stronghold is eastern Mindanao, where rain forests are under 
increasingly severe threat from exploitation and recently from forest fires, even 
within national park boundaries. 

Description Graphium idaeoides is a large white butterfly with reticulated 
black markings and a forewing length of 70-80 mm (Plate 2.4). It closely mimics the 
sympatric species Idea leuconoe (Nymphalidae: Danainae), which has a forewing 
length of around 73 mm (3). The sexes are alike (1,3,9). 

UFW/LFW white with heavy black scaling over the veins, especially in the apical 
and submarginal regions and a row of large , black postdiscal spots. The basal part has 
a creamy or yellowish tinge to the ground colour, varying between island races (9). 

UHW/LHW similar to upper surface. 

Distribution Graphium idaeoides occurs in the eastern Philippines, from 
Luzon to Mindanao. It has been recorded from localities in the north-east, east and 
south of Luzon, from Leyte, Samar, and various localities throughout eastern 
Mindanao (3,5). In addition, there is apparently a disjunct population in the 
Zamboanga Peninsula in the south-west of Mindanao (1,3, 9). 

Habitat and Ecology Graphium idaeoides is a rare and local species 
preferring the lowland rain forests (less than 330 m a.s.l.) on the eastern side of the 
Philippines, where rainfall is continuously high. Jumalon notes that Graphium 
idaeoides flies in forests 100-200 m above sea level in an area north-west of the Mt 
Diwaba range in Mindanao (3, 4). Adults are reported to fly along sheltered and 
shady streams at a height of 3-10 m, much in the manner of the model. Idea 
leuconoe, but with a slightly heavier flight (3). The main flight season is possibly 
February to April after the period of heavy rains, although specimens have also been 
taken in virtually every other month of the year (3, 8, 9). Males defend definite 
territories believed to extend up to 100 m along streams. They may adopt one or 
more perches, from which sorties are made to repel intruders of the same or different 
species. Neither the fast-flowing, upper reaches of the streams on hillsides nor the 
coastal 3 km or so of rivers are used by G. idaeoides, although its model may be seen 
in such places. Females are more elusive than males; Jumalon notes that of 45 
specimens taken between 1958 and 1962, only three were females (3). Unlike some 
other papilionids, G. idaeoides does not fly through the tangled undergrowth of the 
forest (3). Many specimens have been taken along water-courses in secondary 
growth forest, indicating a certain tolerance of disturbance (3). Other species with 
which it shares its habitat include the birdwings Troides magellanus and T. 
rhadamantus (3). The species may have several broods per year, but the foodplants 
and young stages are completely unknown (9). Graphium idaeoides is in the 
macareus group of species (2,7), some of which feed on Aquifoliaceae (2). 

Threats Although widely distributed, Graphium idaeoides seems to be 
rare and local throughout its range (8,9), often very difficult to locate (3). The 

215 



Threatened Swallowtail Butterflies of the World 

hazards to this species are not unique, but common to many other PhiHppine 
endemics. The primary threat is habitat destruction caused by the accelerating rate of 
deforestation and soil degradation throughout the Philippines. Much of the habitat of 
Graphium idaeoides in eastern Luzon, Leyte and northern Mindanao has already 
been cleared for arable crops and coconut plantations, although there are still 
considerable areas of forest on Samar. Its main stronghold seems to be the rainforests 
in the Agusan Valley of north-eastern Mindanao and forested areas in the extreme 
south-east of the island (8), regions which are now under increasingly heavy threat 
from commercial logging and the 'kaingin' farmers (shifting cultivators) who follow 
in their wake. One of the finest national parks in the Philippines, Mt Apo National 
Park, certainly includes this butterfly in its lower reaches, but there have been reports 
that large areas of the Park have been leased for agricultural development. For 
further details of this, refer to the review of the Mt Apo endemic Graphium 
sandawanum. It is now feared that the drought of 1982-3 and the fire that followed in 
its wake have destroyed large areas of Mindanao's eastern forests. The full extent of 
defoliation will only be known when aerial or satellite surveys have been undertaken 
(6). General data on deforestation in Mindanao are given in the reviews of two 
Mindanao endemics, Papilio osinana and Papilio carolinensis . Within the next two 
decades the conservation status of all these butterflies will become increasingly 
serious. 

Conservation Measures The reported threat to Mt Apo National Park is a 
serious matter not only for this butterfly, but for many other species of rain forest 
wildlife, including the Philippine national bird, the Monkey-eating Eagle {Pitheco- 
phaga jefferyi). Maintenance of the integrity of Mt Apo National Park is therefore a 
vital priority. The lowland forests of the Agusan watershed may also prove to be 
important for the conservation of this and many other lowland species since it is one of 
few extensive areas in Mindanao below 200 m a.s.l. The apparently small and very 
local nature of the populations of Graphium idaeoides is a matter for concern. 
Although the reason may be related to its status as a mimic ot Idea leuconoe, further 
study is necessary in order to ascertain its foodplant and breeding ecology. 

References 

1. D'Abrera, B. ( 1982). Butterflies of the Oriental Region. Part 1. Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi -I- 244 pp. 

2. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

3. Jumalon, J.N. (1964). Haunt and habits of Graphium idaeoides (Hewitson) (Papilionidae; 
Lepidoptera). The Philippine Journal of Science 93(2): 207-216. 

4. Jumalon, J.N. (1967). Two new papilionids. Philippine Scientist 1(4): 114-118. 

5. Jumalon, J.N. ( 1969). Notes on the new range of some Asiatic papilionids in the Philippines. 
The Philippine Entomologist 1(3): 251-257. 

6. Lewis, R.E. (1984). In lilt., 17 April. 

7. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supplement 17: 1-51. 

8. Treadaway, C.G. (1984). In //«., 25 May. 

9. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands. Vol. 1 
Papilionidae. (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 



216 



^ Graphium megaera 

Graphium (Pathysa) megaera Staudinger, 1888 INDETERMINATE 

Subfamily PAPILIONINAE Tribe LEPTOCIRCINI 



Summary Graphium megaera is a small swallowtail confined to the forests 
of Palawan in the Philippines. The status of the species is uncertain but, along with 
many other endemic species of wildlife , it is likely to be declining in the face of rapidly 
accelerating deforestation. The national parks system in the Philippines is undergo- 
ing redevelopment. The Province of Palawan needs the protection of significant areas 
of wilderness to ensure the long term survival of its unique fauna and flora. 

Description This small tailless papilionid has a forewing length of about 
35-40 mm. It has been variously stated that the female is unknown (4,7,1 1) and that 
the sexes are similar (3). 

UFW with a dark brown ground colour, submarginal and postdiscal rows of white 
spots. 

UHW similar but with a scalloped outer margin. White markings include 
horseshoe-shaped submarginal spots, postdiscal spots and small discal spots. 

LFW/LHW. The lower surface has a lighter ground colour, but is otherwise 
similar. 

Distribution Graphium megaera is found only on the Philippine island of 
Palawan, where it is fairly widespread, particularly in central and northern regions 
(4, 10). Atrophaneura atropos is also endemic to Palawan and the review of that 
species later in this volume should also be consulted. 

Habitat and Ecology Palawan is a fairly large island ( 1 1 500 sq . km) , 53 per 
cent of which is still covered with tropical lowland dipterocarp and montane forest 
(8). The greater part is in the southern and south-western regions (1). The western 
coast-line tends to have a more seasonal climate and the known areas of distribution 
of Graphium megaera are in the central highlands and eastern lowlands (11). The 
main habitat is lowland, but records indicate a potential distribution up to about 
800 m (10). The butterflies are to be seen flying along the rivers, the males coming 
down to moist spots with a slow, gliding flight (11). Actively on the wing on fine 
mornings, the main flight season is March to April (11), sometimes going through into 
August (4, 10). Females are extremely shy and have been rarely, if ever seen. The 
early stages and foodplants are unknown, but some other species in the macareus 
group feed on Aquifoliaceae (6,7). 

Threats The main threat to the fauna of Palawan is deforestation by timber 
companies and agricuhuraHsts. Much of the island is under concession to logging 
companies (44 per cent of total land area in 1976), and the rate of deforestation is 
increasing rapidly (1,5). The same applies to the rest of the Philippines and nearby 
Sabah (5). Long-term plans include reforestation of logged sites but the experience in 
South East Asia is that this promise is rarely kept. Incursions by shifting cultivators 
and illegal squatters often follow in the wake of the logging road graders (5,8). 
Mining for silica sand, copper, chrome, nickel and mercury may also have a local 
impact on forests, but these threats are more serious to freshwaters and along 
coastlines (2). The precise degree of threat to Graphium megaera is unlikely to be 
critical at this time, but with deforestation accelerating and the Philippine national 

217 



Threatened Swallowtail Butterflies of the World 

parks system uncertain, there is concern for all Palawan's native and endemic 
wildlife. 

Conservation Measures The whole of Palawan was at one time designated 
as a wildlife sanctuary and national park, but this was to no practical purpose (1,2). 
Now it seems that the Philippine Government is restructuring its entire national park 
system in favour of creating fewer parks, but of a higher, international, standard 
(8, 9). It is not clear whether any new parks will be created on Palawan, although St 
Paul's Underground River National Park will presumably remain. There is certainly 
potential for siting more parks in Palawan, where large areas of wilderness are still 
available in the northern highlands around Cleopatra's Needle, in the central 
highlands around an area called The Teeth, and in the southern regions south of Mt 
Mantalingajan (1). It has been suggested that both Cleopatra's Needle and Mt 
Mantalingajan should be designated as Wildlife Sanctuaries, with access only for 
scientific research (9). The status of Graphium megaera and other butterflies needs 
urgent assessment in Palawan, together with rapid recommendations and action in 
siting well-protected national parks on the island. 

References 

1. Bruce, M.D. (Ed.) (1980). The Palawan Expedition Stage I. Traditional Explorations, 
Sydney. 47 pp. 

2. Bruce, M.D. (Ed.) (1981). The Palawan Expedition Stage II. Associated Research, 
Exploration and Aid Pty Ltd., Sydney. 139 pp. 

3. D'Abrera, B. (1982). Butterflies of the Oriental Region. Part 1. Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi + 244 pp. 

4. Dacasin, G.A. (1984). In litt., 25 April. 

5. FAO/UNEP (1981). Tropical Forest Resou-ces Assessment Project. Forest Resources of 
Tropical Asia. FAO, Rome. 475 pp. 

6. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

7. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
£rttowo/og/sf Supplement 17: 1-51. 

8. National Environment Protection Council (1979). Philippine Environment 1979. NEPC 
Third Annual Report. NEPC, Ministry of Human Settlements, Republicof the Philippines. 
158 pp. 

9. Pollisco, F.S. (1982). An analysis of the national park system in the Philippines. 
Likas-Yaman, Journal of the Natural Resources Management Forum 3(12): 56 pp. 

10. Treadaway, C.G. (1984). In litt., 25 May. 

11. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands Vol. 1 
Papilionidae. (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 



218 



'' Graphium aurivilliusi 

Graphium (Arisbe) aurivilliusi Seeldrayers, 1896 INSUFFICIENTLY KNOWN 
Subfamily PAPILIONINAE Tribe LEPTOCIRCINI 



Summary Graphium aurivilliusi is known only from the type series from 
present-day Zaire. More information on its status and distribution is needed. 

Description The male Graphium aurivilliusi is a small blackish-brown 
tailless butterfly with green-tinged white markings and a wingspan of about 75 mm 
(4, 8). The female is unknown (8). This species is in the ucalegon species-groups of 
Berger (2) and Munroe (7), or the adamastor species-group of Hancock (6). 

UFW/LFW blackish-brown with green-tinged white markings, these including 
three elongated cell spots, two large subapical spots and a broad discal band (8). 

UHW/LHW with a broad, green-tinged white discal band, abdominal area, and 
many small postdiscal and submarginal spots (8). 

Distribution This species is known only from the type series from the 
"Congo" in West Africa (4, 7, 8). There are presumably at least two specimens since 
the two illustrations given by Berger (1,2) are different. The precise collecting 
locality is unknown (8) but is almost certainly in present-day Zaire (1, 2, 3). 

Habitat and Ecology Unknown. Other members of the adamastor group 
feed on Anacardiaceae (6). 

Threats This butterfly is so rare and poorly known that there are insuffi- 
cient data to assess current threats. The forests of Zaire and Congo are better 
protected than in any other African countries. 

Conservation Measures No proposals are possible without habitat informa- 
tion. Congo has 1300 sq. km of rain forest legally protected from logging, and Zaire 
has 56 900 sq. km(5). Logging in these countries is currently less than 0.2 per cent per 
year. 

References 

1. Berger, L.A. (1950). Catalogues raisonnees de la faune entomologique du Congo Beige. 
Lepidopteres-Rhopaloceres, 1, Family Papilionidae. Annates du Musee Royale du Congo 
Beige Ser. 4to (C: Zool.) 8: 1-104. 

2. Berger, L.A. (1951). Systematique des Papilionidae de la faune ethiopienne. 3me Congres 
National des Sciences, Bruxelles 8: 47-50. 

3. Carcasson, R.H. (1980). Collins Handguide to the Butterflies of Africa. Collins, London. 
Hardback edition, 188 pp. 

4. DAbrera, B. (1980). Butterflies of the Afrotropical Region. Lansdowne Press, xx -I- 593 pp. 

5. FAO/UNEP (1981). Tropical Forest Resources Assessment Project. Forest Resources of 
Tropical Africa. Part 1: Regional Synthesis. FAO, Rome, 108 pp. 

6. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

7. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Ewtowo/og/i? Supplement 17: 1-51. 

8. Williams, J. G. (1969). A Field Guide to the Butterflies of Africa. Collins, London. 238 pp. , 24 
col. pi. 

219 



Threatened Swallowtail Butterflies of the World 

Graphium (Arisbe) weberi Holland, 1917 INSUFFICIENTLY KNOWN 

Subfamily PAPILIONINAE Tribe LEPTOCIRCINI 



Summary Graphium weberi comes from Cameroon whence it is known 
only from a single specimen. Assessments of the taxonomic status and distribution of 
the species are required. 

Description Graphium weberi is a small brown and ochreous tailless 
butterfly in the ucalegon group of Munroe (7), and the adamastor group of Hancock 
(4). It has a wingspan of about 80 mm (8). Since the species has been so rarely 
collected, doubts may be raised about its taxonomic status. The closely related and 
sympatric G. ucalegonides and G.fulleri are similar in appearance and known to be 
variable (5,6). 

UFW blackish-brown with a broad, pale ochreous median band separated into 
large spots, and a line of large subapical spots (8). 

UHW also blackish-brown with a pale ochreous band and pale ochreous 
submarginal spots (8). 

LFW/LHW pale brown with reddish wing bases (8). 

Distribution This species from Cameroon, West Africa, is known only from 
the type specimen, held at the Carnegie Museum, Pittsburgh, Pennsylvania (2). It is 
not clear on what basis Williams noted that the sexes are alike (8). 

Habitat and Ecology Believed to be confined to tropical rain forest in 
Cameroon (1, 5). Feeding requirements unknown, but other members of the group 
feed on Anacardiaceae (4). 

Threats This butterfly is so rare and poorly known that there are insuffi- 
cient data to assess current threats. However, it may be noted that Cameroon 
currently contains no forest which is legally protected from logging (3). 

Conservation Measures Three forest sites in Cameroon are scheduled for 
full protection: Korup (c. 90 000 ha), Dja (c. 540 000 ha) and Pangar-Djerem (c. 
480 000 ha). Legislation is expected during the current five-year plan. The World 
Wildlife Fund is assisting the newly-formed Department of Wildlife and National 
Parks in various ways. It is not known whether Weber's swallowtail occurs in these 
proposed parks. 

References 

1. Carcasson, R.H. (1980). Collins Handguide to the Butterflies of Africa. Collins, London. 
106 pp. 

2. D'Abrera, B. (1980). Butterflies of the Afrotropical Region. Lansdowne Press, xx + 593 pp. 

3. FAO/UNEP (1981). Tropical Forest Resources Assessment Project. Forest Resources of 
Tropical Africa. Pari 1: Regional Synthesis. FAO, Rome, 108 pp. 

4. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

5. Hancock, D.L. (1983). In to., 21 June. 

6. Hancock, D.L. (1984). In litt., 25 June. 

220 



Graphium weberi 

7. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
£/?/owc»/og/i/ Supplement 17: 1-51. 

8. Williams, J.G. (1969). A field guide to the butterflies of Africa. Collins, London. 238 pp. 

Note added in proof: In a very recent paper, Hancock (9) placed Graphium weberi: as 
a new synonym of G. fulleri, thereby invalidating the above assessment. G. fulleri is 
widespread in the forests of Cameroon, Gabon, Congo Republic, Ivory Coast, 
Ghana, Angola and Zaire. 

9. Hancock, O.L. (1985). Systematic notes on some African species of Graphium Scopoli. 
Papilio International 2(1-2); 97-103. 



221 



Threatened Swallowtail Butterflies of the World 

Graphium (Arisbe) levassoh Oberthiir, 1890 VULNERABLE 

Subfamily PAPILIONINAE Tribe LEPTOCIRCINI 



Summary Graphium levassori is confined to Grande Comore in the 
Comoro Islands, where it is extremely scarce. There is no information on its biology 
or ecology and surveys and studies are urgently needed. 

Description This pale yellow butterfly with black markings is in the 
leonidas group (4) and has similar sexes, though the females are slightly larger (Plate 
2.5). The forewing length is 43-55 mm (3, 7). 

UFW pale yellow with a black apex, outer and costal margins; wing base reddish 
with a small black cell spot, two yellow apical spots and a narrow yellow streak 
between the cell and the outer margin (3, 6). 

UHW also pale yellow, with a black border to the scalloped outer margin. There is 
some brown scaling at the apex with two yellow lunules and a silvery-white costal 
margin (3, 6, 7). 

LFW/LHW differs from the uppersides in having fawn instead of black scaling and 
brick-red wing base (3, 6, 7). 

Distribution This species is endemic to Grande Comore in the Comoro 
Islands, between Mozambique, on the African mainland, and Madagascar in the 
Indian Ocean (3,6). 

Habitat and Ecology The four islands of the Comoros are volcanic in 
origin, Grande Comore being the most recent. On Grand Comore there are two 
upHfted areas, the volcanic massif of La Grille (1087 m) in the north, and the still 
active volcano of Karthala (2560 m) in the south. All the islands are characterized by 
moderate rainfall (1000-5000 mm per year) and soils with a very poor capacity for 
water retention. There are no streams or rivers on Grande Comoro, although 
subterranean hollows in the lava may hold water and encourage growth of forest (9). 
Patches of forest may still be found at low and middle altitudes e.g. near 
Nioumbadjou (550 m), but the main forests occur on the flanks of La Grille and 
Karthala up to a maximum of 1900 m ( 1 ). The forests on La Grille have been severely 
depleted by banana cultivation, but they are more extensive on Karthala, on whose 
western and south-western sides forest may grow at altitudes as low as 550-800 m ( 1 ) . 
Graphium levassori inhabits forested areas and is believed to be very local in its 
distribution (5). In 1980 it was reported from forests near M'Lima Manda, north of 
Karthala, where it was said to be not rare but difficult to capture (9). However, a 
further visit to search for the butterfly in 1983 was unsuccessful (8). Little is known of 
the habits or biology of this species, but other members of the leonidas group feed on 
Annonaceae, the Custard Apple family (4). G. levassori has been found at sea level 
in the north of the island, where the cultivated Custard Apple {Annona reticulata) is 
common (2), but this seems to be an exceptional record and has not been recently 
confirmed. The adults may be on the wing all year round since dry periods are not 
usually prolonged, but they are certainly on the wing at the beginning of the heaviest 
rains in April (9). 

Threats Graphium levassori is one of many species of insects and other 
organisms, including Papilio aristophontes (see separate review), that are confined to 

222 



Graphium levassori 

the forests of Grande Comore and the other Comoro Islands. Even within these 
forests G. levassori seems to be restricted to medium altitudes and very patchily 
distributed. There is considerable evidence that the forests are under increasingly 
severe pressure and that if forest destruction continues to accelerate, extinctions are 
inevitable. 

The estimated human population of Grande Comore in 1976 was 150 000, and 
growing rapidly. The island is mainly farmed at the subsistence level and income from 
the main cash crops such as vanilla, copra, cacao, sisal, coffee, cloves and essential 
oils is very low. The main subsistence crop is bananas, plantations and gardens of 
which clothe the island. Even forests on La Grille and Karthala, from a distance 
apparently dense and healthy, may be found on closer inspection to include few large 
trees and a dense understorey of banana trees (9). The wild trees are unable to 
reproduce and regenerate, and they are gradually retreating higher and higher (9). 
Bananas are grown at altitudes up to 1200 m; the destruction is particularly serious on 
La Grille, but also extensive on Karthala (1, 9). The gradual erosion of the forests by 
the planting of bananas has been continuing for at least the past 25 years (9). 

Conservation Measures Even though Graphium levassori may be able to 
eke out an existence on cultivated Custard Apple, the only way to ensure its 
long-term survival is protection of its forest home from the constant incursion of 
banana growers. At present there are no protected areas on the islands, a situation 
which should be remedied as soon as possible. It is clearly necessary to create a series 
of reserves in representative biotopes. In the middle altitudes of Karthala it is almost 
too late to find undisturbed forest, but some areas should be protected and allowed to 
regenerate. At higher altitudes of 1200 m and above, the government could consider 
making the whole of Karthala a protected area. This would not affect many people 
and would ensure the survival of at least the higher altitude species for posterity. 

Conservation measures specifically for Graphium levassori are not possible at this 
stage. More data are needed on its distribution and ecology in order that measures for 
its protection may be incorporated into efforts to conserve representative biotopes. 
Being large and spectacular as well as rare, Graphium levassori is prized by collectors 
(5) . A farming or ranching programme on Custard Apple bushes could be developed 
into a useful local industry as well as a valuable measure for conservation purposes. 

References 

1. Benson, C. W. ( 1960). The birds of the Comoro Islands: results of the British Ornithologists' 
Union Centenary Expedition 1958. Ibis 103b: 5-106. 

2. Collins, S. (1983). In liti., 12 July. 

3. D'Abrera, B. (1980). Butterflies of the A frotropical Region. Lansdowne Press, xx + 593 pp. 

4. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
£«/owo/og/sf Supplement 17: 1-51. 

5. Paulian, R. (1983). In litt., 10 May. 

6. Paulian, R. and Viette, P. (1968). Faiine de Madagascar. XXVII Insectes Lepidopteres 
Papilionidae. O.R.S.T.O.M. and C.N.R.S., Paris. 97 pp., 19 pi. (2 col.), 34 figs. 

7. Smart, P. (1975). The Illustrated Encyclopedia of the Butterfly World. Hamlyn, London. 
275 pp. 

8. Tudin, B. (1983). In litt., 15 September. 

9. Viette, P. (1980). Mission lepidopterologique a la Grande Comore (Ocean Indien 
occidental). Bulletin de la Societe Entomologique de France 85: 226-235. 



223 



Threatened Swallowtail Butterflies of the World 

Graphium (Graphium) procles Grose-Smith, 1887 INDETERMINATE 

Subfamily PAPILIONINAE Tribe LEPTOCIRCINI 



Summary Graphium procles is restricted to the Crocker Range in the East 
Malaysian state of Sabah in northern Borneo . It is best known from Mt Kinabalu , the 
highest mountain between the Himalayas and New Guinea, where it occurs in 
montane rain forest. Mt Kinabalu is designated as a national park, but economic 
interests such as copper mining, logging, farming and recreation are eroding the park 
boundaries. 

Description Graphium procles is a tailless butterfly with a forewing length 
of about 39^2 mm. The female is very rarely seen, and has not been described in the 
literature. 

UFW black with a broad median band tapering from the inner margin towards the 
apex, a row of small submarginal spots, and some irregular subcostal spots and 
streaks mainly in the subapical region. Markings are pale blue-green, becoming 
yellower towards the apex (7). 

UHW black with a scalloped outer margin, and a continuation of the UFW median 
band which crosses the cell and tapers sharply towards the anal angle. The subcostal 
region is white and other markings are pale blue-green, including the submarginal 
spots (7). 

LFW/LHW with paler markings and a slightly browner ground colour. The LHW 
has irregular dark orange or buff discal markings (6) . The subcostal part of the LHW 
band is cream-coloured (6). 

Distribution Graphium procles is endemic to Sabah (a state of Malaysia) in 
northern Borneo. It occurs in the west, only on Gunung (Mt) Kinabalu and the 
Crocker range (6, 7). 

Habitat and Ecology Graphium procles is restricted to lower montane 
forest (4) at elevations over 1000 m (6). It is best known from Gunung Kinabalu, 
where the species as a whole is not uncommon, possibly the most frequently 
encountered species in the subgenus (1). Females, however, are extremely rare (7). 
The early stages and foodplants are unknown (7) but other members of the eurypylus 
group feed on Annonaceae, Lauraceae or Magnoliaceae (3). Adults have been 
recorded at all times of year, flying along streams or visiting moist spots (7). 
Graphium procles shares its habitat with the widespread Graphium bathycles, but 
their main flight seasons tend to be separate. 

Threats The main threat to Graphium procles is destruction of its restricted 
montane habitat. Mt Kinabalu is the only part of the Crocker Range that is 
designated as a national park, and even this is under pressure from a number of 
quarters. Increased tourism in recent years has brought necessary developments such 
as access roads and a visitor centre , but plans for a golf course and other recreational 
facilities in the virgin forests of the Pinosuk plateau threaten to destroy habitats and 
are of questionable conservation value. Parts of the Pinosuk plateau have already 
been lost to forest clearance for cattle farming, timber processing and hydro-electric 
projects (2) . Many of the lower slopes of western Kinabalu have long been cleared by 
shifting cultivators (2). 

224 



Graphium procles 

On eastern Kinabalu the discovery of porphyry copper has resulted in the lease of 
2556 ha of national park land to a joint Japanese-Malaysian mining company. The 
lease runs from 1973 for 30 years and the Mamu Copper Mine has completely 
changed the eastern face of Kinabalu (2). These alterations to Mt Kinabalu are a 
serious threat to its butterflies and the great wealth of other endemic animals and 
plants that occur there. Two other threatened papilionids, Papilio acheron and 
Troides andromache, are also found on Mt Kinabalu (see separate reviews). 

Conservation Measures Kinabalu is the highest mountain in South East 
Asia and the highest point between the Himalayas and New Guinea (5). Its value as 
an important biogeographical site and as a water catchment area led to its designation 
as a national park in 1964 (5). It now acts as a vital conservation area and a major 
tourist attraction. Short-term economic developments may threaten the long-term 
value of the region as a biological storehouse and source of tourist revenues. Sensitive 
planning is needed if further losses of these rare habitats and their fauna and flora are 
to be avoided. 

References 

1. Barlow, H.S., Banks, H.J. and Holloway, J.D. (1971). A collection of Rhopalocera 
(Lepidoptera) from Mt Kinabalu, Sabah, Malaysia. Oriental Insects 5: 269-296. 

2. Davis, S. (1983). Draft review for lUCN Plant Sites Red Data Book. 

3. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

4. Holloway, J.D. (1978). Butterflies and Moths. In Kinabalu Summit of Borneo. Sabah 
Society Monograph, 25-278. 

5. Luping, D.M., Wen., C. and Dingley, E.R. (Eds) (1978). Kinabalu. Summit of Borneo. 
Sabah Society Monograph, Malaysia. 

6. Robinson, J.C. (1975-6). Swallowtail butterflies of Sabah. Sahah Society Journal 6: 5-22. 

7. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands. Vol. I 
Papilionidae. (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 



225 



Threatened Swallowtail Butterflies of the World 

Graphium (Graphium) meeki (Rothschild & Jordan, 1901) RARE 

Subfamily PAPILIONINAE Tribe LEPTOCIRCINI 



Summary Graphium meeki is a very rare and locally distributed butterfly 
from the mountainous districts of Bougainville, Choiseul and Santa Isabel. With 
extensive mining, logging and agricultural activities within its range, there are fears 
that local population extinctions may be occurring even before the distribution of the 
butterfly is properly known. With more data on its distribution and biology, it could 
be a good candidate for small-scale farming. 

Description The authors have not seen a specimen or photograph of a male 
Graphium meeki, but the sexes are probably similar, as in the closely related G. 
agamemnon . 

Female : UFW ground colour black with blue-green markings consisting of rows of 
spots in the subcostal, subapical, submarginal and median areas (1,5). Forewing 
length about 53 mm (7). 

UHW black with a slightly scalloped outer margin, no tail, some white spots and a 
row of large blue-green discal-submarginal spots. 

LFW/LHW similar to the upper sides but scaled with white and with a sub-basal red 
bar (1,7). 

Distribution This butterfly is endemic to the Solomon Archipelago, east of 
Papua New Guinea. It has only been recorded from Bougainville (Papua New 
Guinea), Choiseul, and Santa Isabel (Solomon Is) (4, 5). 

Habitat and Ecology Graphium meeki is confined to mountainous, inland 
areas (5). Much of its range remains little explored and although generally rare it may 
be locally common. The foodplants are unknown, but other species in the 
agamemnon group feed on Annonaceae or Magnoliaceae (2). In northern Bougain- 
ville G. meeki has been found at 600-800 m in cleared areas of rain forest (6). The egg 
is large, pale yellow and laid on young leaves of Litsea sp. (Lauraceae). Caterpillars 
of the second or third instar are completely black with no markings (6). 

Threats No specific threats have been identified so far (3), but the species is 
so rarely collected and poorly known that populations could be disappearing before 
they are recorded. The Solomon Islands Government requires an export permit for 
all butterfly captures, but this is difficult to enforce. In any case, habitat destruction is 
likely to be a more significant threat. There are major logging interests throughout 
the Solomons archipelago. Habitat destruction is extensive on Bougainville, where 
open-cast copper mining is a major industry. 

Conservation Measures Further study of the distribution and biology of 
Graphium meeki is urgently needed. Forest exploitation could already be reducing 
the range of Graphium meeki, possibly causing extinctions of some of its apparently 
highly localized populations. The status of this species should be reviewed as soon as 
more data are forthcoming. There is a possibility that it could be Vulnerable, if not 
Endangered. If a small farming operation could be initiated it would be a good 
conservation measure as well as providing local employment. There would certainly 
be some demand for such a little-known species. 

226 



Graphium meeki 



References 



1. D'Abrera, B. (1971). Butterflies of the Australian Region. Lansdowne Press, Melbourne. 
415 pp. 

2. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

3. Macfarlane. R. (1983). In litt., 15 March. 

4. Parsons, M.J. (1980). In liti.,1 February. 

5. Racheli, T. ( 1980). A list of the Papihonidae (Lepidoptera) of the Solomon Islands, with 
notes on their geographical distribution. Australian Entomologists' Magazine. 7: 45-59. 

6. Racheli, T. (1984). In litt., 18 June, incorporating observations from R. Straatman. 

7. Rothschild, W. and Jordan, K. (1901). On some Lepidoptera. Novitates Zoologicae 1901; 
401-407. 



227 



Threatened Swallowtail Butterflies of the World 

Graphium (Graphium) stresemanni (Rothschild, 1916) RARE 

Subfamily PAPILIONINAE Tribe LEPTOCIRCINI 



Summary Graphium stresemanni is a little-known butterfly from Seram in 
Maluku Province (Moluccas), Indonesia. Although Seram has not yet attracted 
large-scale logging operations comparable to other areas of the province, the threat 
remains. The Manusela Reserve on Seram promises to protect the island's wildlife 
but has yet to be designated as a national park. There is a substantial trade in 
butterflies on Seram, involving whole villages. Documentation and rationalization of 
this trade is needed. 

Description Graphium stresemanni is a brown, tailed butterfly with white, 
pale blue, pale green and yellow markings and a forewing length of about 40 mm 
(2,8). The species was not listed by Munroe (4) because it was originally described as 
a subspecies of G. weiskei, but it is now generally accepted as a full species of some 
rarity (2, 6, 7, 8). The sexes are broadly similar. 

UFW dark brown ground colour with pale blue patches below the cell and in the 
postdiscal area, yellow subapical spots and a row of small white, submarginal spots 
(2,4). 

UHW with a similar brown ground colour, pale green markings in the basal and 
discal regions, and a row of larger, white, submarginal spots. 

LFW/LHW similar to the upperside (2). 

Distribution Graphium stresemanni is only found on Seram (otherwise 
known as Serang or Ceram) , one of the larger and more southerly islands of Maluku 
Province (Moluccas), Indonesia. Several of the specimens listed by Okano (6) came 
from around the 1000 m level in the Manusela range of central Seram. 

Habitat and Ecology Very little is known about this butterfly and its 
ecological requirements have not been researched. Its young stages and foodplants 
have not been recorded, but the sarpedon group, of which stresemanni is a member, 
feed on Winteraceae, Monimiaceae, Lauraceae or Hernandiaceae, rarely other 
groups (3). The main range of the group is the eastern Palaearctic, but a number of 
species reach into Indo-Australia. G. sarpedon extends throughout South East Asia 
and New Gmnea to Queensland, Australia, but other species are more restricted, in 
some cases as a result of the fluctuations in sea level resulting from the Ice Ages. G. 
empedovana is found from Palawan to Sumatra while G. codrus extends eastwards, 
from Sulawesi and the Philippines across to the Solomons. A few species occur only 
over a very small range, either because they have evolved there or because they are 
now restricted to a small fragment of a biome that may once have been more 
extensive. G. sandawanum (see next review) is restricted to montane forest on 
Mt Apo, Mindanao, and G. monticolum is only found in similar habitat on Sulawesi 
(3). Seram rises to over 3000 m in its central mountain range and G. stresemanni is 
probably another isolated montane relict species, perhaps with its main populations 
in the Manusela range. 

Seram is the largest of the Maluku islands, very mountainous with central peaks 
rising to over 3000 m and with a rainfall of around 2000 mm falling mainly between 
May and October (1,9). The natural vegetation includes lowland and montane 
forests. The lowland forests are mainly semi-deciduous and some of the common tree 

228 



Graphiiim stresemanni 

genera found there are Canarium, Eucalyptus, Intsia, Agathis, Diospyros and 
Palaquium. 

Threats Maluku Province has a unique and scientifically extremely impor- 
tant fauna. At least 25 species of butterfly (1), 14 birds (5) and five mammals as well as 
many snails and reptiles are endemic to the islands. Seventy per cent of the islands' 
area is still untouched, but large-scale commercial logging is affecting the forests of 
the northern and central islands (9) and may in future spread south to Seram. Much of 
Seram consists of steep hills and deep valleys that are essential not only to the wildlife, 
but also as watersheds supplying the coastal towns and villages. These regions need to 
be permanently protected from logging operations. 

Many of Seram's indigenous people traditionally trade in wildlife. Wild sago and 
damar resin are important forest products, and plants, cockatoos and lories have long 
been taken for barter. This formerly local trade has recently become lucrative and 
international, and butterflies have become increasingly in demand (9). According to 
a recent report, there are several whole villages on Seram whose inhabitants depend 
totally on butterfly catching (9). About 90 species of butterfly occur in Maluku, 
including two threatened birdwings(Or/i/7/zopreraa^5fla« (Obi only) and O. croesus) 
as well as Graphium stresemanni and a number of other endemic butterflies, including 
the swallowtails Troides criton, Papilio deiphobus and Papilio tydeus. Graphium 
stresemanni has been seen on sale in Hatumetan on the southern coast of Seram (10) 
as well as in a number of trade catalogues. 

Conservation Measures Manusela Reserve (proposed as a national park) 
covers a large area of Seram (180 000 ha (5, 9)), and extends from sea level to the 
peak of Seram's (and the Province's) highest mountain, Gunung Pinaia (3027 m). All 
Seram's ecosystems are represented and the island's 14 endemic birds and five 
endemic mammals have also been recorded there (5,9). Graphium stresemanni and 
many other endemic butterflies are certainly to be found in the area. There is a great 
need for more information from Manusela. 

Feasibility studies and a management plan for the proposed Manusela National 
Park have been prepared (9) but as yet there is no legislation to give the reserve full 
national park status. At present the reserve has no headquarters, no facilities and 
presumably no guards (1). Steps to ensure the future of the reserve are urgently 
needed since the pressure of economic development in the forests of north and 
central Maluku must eventually spread southwards into the relatively untouched 
island of Seram. Part of the northern lowlands of the reserve have already been 
logged over, prior to reserve status being gazetted (5). 

Seram is too large to be adequately served by one national park, even one as large 
as Manusela, and the Indonesian Conservation Plan recommends another in the 
mountainous interior of the Sahuai peninsula (5). Gunung Sahuai in west Seram is 
covered in primary rain forest and is an important water catchment area and resource 
for wildlife. Logging concessions have already been requested (5) and gazetting as a 
protected area is urgently needed. 

With such a superb butterfly fauna, Maluku Province could do more to increase 
awareness of this important heritage. Local people involved in the butterfly trade are 
already aware of the international demand for specimens. Studies towards the 
development of butterfly farms or ranches would help to meet this demand and at the 
same time encourage local interest and conservation. The local tourist board could 
encourage appreciation of the butterfly fauna through its brochures and adver- 
tisements. 

229 



Threatened Swallowtail Butterflies of the World 

References 

1 . Anon, (undated). Indonesia National Parks and Nature Reserves. Directorate General of 
Tourism, Jakarta. 150 pp. 

2. D'Abrera, B. (1971). Butterflies of the Australian Region. Lansdowne Press, Melbourne. 
415 pp. 

3. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

4. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
£Mtomo/og(5f Supplement 17: 1-51. 

5. National Park Development Project, UNDP/FAO (1981). National Conservation Plan for 
Indonesia Vol 7: Maluku and Irian Jaya. FAO, Bogor. 

6. Okano, K. (1983). On the data of the type-specimen Graphium stresmanni (sic) 
(Papilionidae) Tokurana (Acta Rhopalocera) 5: 88. 

7. Saigusa,S.,Nakanishi, A.,Shima,H.andYata,0.(1977).Phylogenyandbiogeographyof 
the subgenus Graphium Scopoli. Acta Rhopalocera 1: 2-32. 

8. Smart, P. (1975). The Illustrated Encyclopediaof the Butterfly World. Hamlyn, London. 275 
pp. 

9. Smiet, F. (1982). Threats to the Spice Islands. Oryx 16(4): 323-328. 
10. Walker, M.V. (1983). In litt., 11 April. 



230 



Graphium sandawanum 
Graphium (Graphium) sandawanum Yamamoto, 1977 VULNERABLE 

Subfamily PAPILIONINAE Tribe LEPTOCIRCINI 



Summary Graphium sandawanum is confined to Mt Apo in Mindanao, the 
Philippines. The mountain has been a national park since 1936 but has suffered 
constant encroachment by squatters. In a recent proclamation the government has 
agreed to lease 56 per cent of the park's area to the squatters ; much of this area is still 
under virgin forest. Drought followed by fires in 1983 caused a further loss of forest on 
Mt Apo. There is international concern for the fate of the park, which is a stronghold 
for the Endangered Philippine Monkey-eating Eagle as well as this rare and beautiful 
butterfly. 

Description This recently discovered, black and pale yellow, short-tailed 
butterfly has a forewing length of about 38 mm in the male, and 41 mm in the female. 
The sexes are similar, the female with a slightly paler ground colour (3, 7, 11). 

UFW with a dark brown or black ground colour and a very broad yellow-green 
discal band running from the anal margin and tapering into three subapical spots 
(3,7,11). 

UHW similar, with the yellow-green discal band tapering towards the anal angle. 
There is also a row of pale green submarginal lunules (3, 7, 11). 

LFW/LHW. The lower surface is blackish-brown with a paler distal border. The 
discal band is yellow green as on the upper surface, but there are a number of small 
red or yellow markings on the distal edge of the band on the hindwing. The ratio of the 
red-spotted form to the yellow-spotted form has been estimated at 7:3 in the male 
(11). 

Distribution This remarkable species was discovered as recently as 1977 on 
Mt Apo (2954 m) in central Mmdanao, Philippines (12). So far it has been found 
nowhere else. After some 50 years of collecting by lepidopterists in the Mt Apo 
region it is quite extraordinary to find that in its chosen locations both males and 
females of this species are relatively abundant (10). No-one knows why it was not 
discovered earlier. 

Habitat and Ecology Graphium sandawanum is considered to be an Ice Age 
relict of Palaearctic continental stock (11). According to Tsukada and Nishiyama, it 
usually flies in the mossy, montane forest of Mt Apo (11), the broad green bands on 
the wings making it very conspicuous. The foodplants and young stages are unknown, 
but other members of the sarpedon group feed on Winteraceae, Monimiaceae, 
Lauraceae or Hernandiaceae, rarely other groups (4) . Graphium sarpedon itself also 
flies on Mt Apo ( 12) , but is less common at altitudes over 1000 m, where sandawanum 
flies (11). Graphium sandawanum has been captured from January to June , October 
and December, and is probably present all year round, possibly with reduced 
populations during the heavy rains of February to May (7, 10, 11). Mt Apo may be 
approached from Davao or Cotabato. The butterflies apparently fly in greater 
abundance on the western, Cotabato, side (11). 

Threats Mt Apo is the highest mountain in the Philippines (2909 m) and 
was designated as a national park as long ago as 1936, with a total area of 76 900 ha 
(9). Since then it has undergone a number of amendments in terms of classification 

231 



Threatened Swallowtail Butterflies of the World 

and land use (9), but none so alarming as the recent newspaper and other reports that 
56 percent of the area, almost 42 000 ha, is to be reclassified as 'disposable land' and 
handed over to illegal squatters (2, 7). Squatters have settled in the park since 1945 
and current records indicate that 4034 famiHes are living within the park boundaries 
(2). Attempts have been made in the past to reforest the lower slopes of Mt Apo and 
restore the natural beauty and integrity of the park (1). The latest proclamation 
indicates that these measures have been largely unsuccessful. The 42 000 ha will be 
leased to the squatters for 25 years, but only 35 percent of this is currently suitable for 
commercial agriculture. The other 65 percent is still under forest but is likely to be 
exploited very quickly since the new proclamation will allow squatters to take up 
loans to expand their farms (2). 

A further threat only recently being fully realized is the serious drought of 1982-3 
that has affected eastern Mindanao, eastern Borneo and perhaps other islands. In the 
wake of the drought, huge fires have destroyed vast areas of forest. In Mindanao 
whole mountains have been burnt through and both Mt Apo and Mt Katanglad have 
suffered badly (8). The full extent of defoliation is not yet known and will only be 
discovered from aerial or satellite survey. 

Public concern has been mainly for the Endangered Phillippine Monkey-eating 
Eagle (Pithecophaga jefferyi) (6) but there is undoubtedly a wide range of other 
species, including Graphiiim sandawanum, whose livelihood is threatened by these 
recent developments. 

Recent reports confirm that quantities of G. sandawanum specimens are reaching 
butterfly dealers in Manila and are being sold on the international market (5). The 
most likely source of the material is local youths employed by dealers in remote cities 
(possibly even overseas) to collect the butterflies on Mt Apo. Such arrangements are 
not only irresponsible but are also quite illegal since the butterfly only occurs within 
national park boundaries. The Philippine authorities should take steps to prevent 
such blatent abuse of their national legislation. 

Conservation Measures There is evidence that the proclamation to lease 
such a huge area to squatters on Mt Apo was not supported by a thorough survey of 
the forests and agricultural land within the boundaries adopted (2,7). Those areas 
irretrievably occupied by squatters are of no value for the conservation of Mt Apo's 
wildlife , but to expand squatters' privileges into virgin forested land of a national park 
would appear to be a short-sighted move. Much basic scientific work remains to be 
done on Mt Apo. Undoubtedly many species await discovery and only thorough 
survey work wil! reveal the full international importance of the park. Meanwhile, the 
national park authorities should take immediate steps to prevent the constant erosion 
of the park's boundaries and wildlife . Conflicting land-use needs will not be solved by 
ignoring the problem. 



References 

1. Anon. (1980). Logging firms tapped for Mt Apo reforestation. Evergreen (Publ. Bureau of 
Forest Development) 5(4); 18. 

2. Anon. (1983). BFD moves to save Philippine Eagle. Bulletin Today, 1 December: 8. 

3. D'Abrera, B. (1982). Butterflies of the Oriental Region. Part 1. Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi + 244 pp. 

4. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1 48. 

5. Jumalon, J.N. (1984). In litt., 10 July. 

232 



Graphium sandawanum 

6. King, W.B. (1981). Endangered Birds of the World. The ICBP Bird Red Data Book. 
Smithsonian Institution Press, Washington D.C. 

7. Lewis, R.E. (1984). In lilt., 5 and 9 February. 

8. Lewis, R.E. (1984). In litt.. 17 April. 

9. Pollisco, F.S. (1982). An analysis of the national park system in the Phillippines. 
Likas Yaman, Journal of the Natural Resources Management Forum 3(12): 56 pp. 

10. Treadaway, C.G. (1984). In litt., 25 May. 

11. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands Vol. I 
Papilionidae. (transl. K. Morishita). Plapac Co. Plapac Co. Ltd., Tokyo. 457 pp. 

12. Yamamoto, A. (1977). A new species of Graphium from Mindanao (Lepidoptera: 
Papilionidae). Tyo to Ga. ( Transactions of the Lepidopterists' Society of Japan). 28: 87-88. 



233 



Threatened Swallowtail Butterflies of the World 

Graphium (Graphium) mendana (Godman and Salvin, 1888) RARE 

Subfamily PAPILIONINAE Tribe LEPTOCIRCINI 



Summary Graphium mendana has spread throughout the Solomon archi- 
pelago with the exception of San Cristobal, but is apparently rare everywhere. 
Logging and agriculture are probably the main threats to the species, but 
considerably more biological and ecological data are needed before a rational 
conservation programme can be proposed. 

Description Graphium mendana is a relatively large butterfly with a 
forewing length of up to 52 mm. The sexes are similar. Four subspecies have been 
described (1,2,4,6). 

UFW with a black upper surface and a median band of spots from the middle of the 
inner margin to the wing apex. The colour, originally described as 'cyan' (blue- 
green) (2) seems to vary between green and yellow (1, 2, 6, 7). Submarginal spots 
may be present in some subspecies (1,2,6,7). In the original description G. 
mendana was believed to be closely related to G. sarpedon (2), but Munroe placed it 
in the eurypylus group (5). In a more recent revision, G. mendana has become the 
nominate species in a new group (3). 

UHW with a scalloped outer margin and a long, broad tail with a rounded tip. The 
UHW is black with a large, pale blue-green basal patch extending along the 
abdominal part of the wing and a white anal margin (1,2). 

LFW brown-black with smaller round spots forming the median band; LHW dull 
black with no basal patch. Markings on the LHW vary according to subspecies; G.m. 
acous (Ribbe) has a green basal spot and two red spots before the anal angle, whereas 
G. m. neyra (Rothschild) only has one red spot (1, 2, 7). 

Distribution Graphium mendana is endemic to the Solomon Archipelago. 
The four subspecies are distributed from Bougainville in the north-west to Malaita 
and Guadalcanal in the south-east. G. m. acous is only known from Bougainville 
Island (Papua New Guinea) whereas all of the other races occur in the Solomon 
Islands. G. m. neyra is found in the New Georgia group of islands where it has been 
recorded from New Georgia, Rendova and Vella Lavella, and the nominate 
subspecies G. m. mendana is known from Santa Isabel and the north-east of 
Guadalcanal. A fourth subspecies G. m. aureofasciatum Racheli was described in 
1979 from Dala, north-west Malaita (1,7). 

Habitat and Ecology Graphium mendana seems to be very rare throughout 
its range, and very poorly known. The foodplant is believed to be in the Piperaceae 
(3), but no further details of breeding or habitat are given in the literature. 
Nevertheless, it has been successfully bred in captivity (7). 

Threats G. mendana is to be found on most of the islands of the Solomon 
archipelago except San Cristobal. In the absence of habitat data it is difficult to 
catalogue threats in any detail, but forestry, agricultural activity and other forms of 
habitat destruction are certainly of most concern (4). G. mendana neyra may be 
vulnerable to logging operations in the New Georgia group (4). 

Conservation Measures More data on the habitat and ecology of Graphium 
mendana are needed before suitable conservation measures can be put forward. 

234 



Graphium mendana 

Breeding habits and details of habitat are essential prerequisites to a captive breeding 
programme that could benefit the butterfly and its breeders. Habitat data will also 
indicate where disruption of native vegetation is most likely to affect the species, and 
where protected areas would be most effective. Export of butterflies from the 
Solomon Islands requires a permit, but this is difficult to enforce, and collecting is in 
any case unlikely to constitute a major threat. 

References 

1. D'Abrera. B. (1971). Butterflies of the Australian Region. Lansdowne Press. Melbourne. 
415 pp 

2. Godman, F.D. and Salvin, O. (1888). New species of butterflies collected by Mr. CM. 
Woodford in the Solomon Islands. The Annals and Magazine of Natural History. (6)1: 
209-214. 

3. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

4. Macfadane, R. (1983). In litt., 15 March. 

5. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supp\ement 17: 1-51. 

6. Racheli,T. (1979). Newsubspeciesof Pap/V/o and Grap/j/ww from the Solomon Islands, with 
observations on Graphium codrus (Lepidoptera, Papilionidae). Zoologische Mededelingen 
54(15): 237-240. 

7. Racheli, T. (1980). A list of the Papilionidae (Lepidoptera) of the Solomon Islands, with 
notes on their geographical distribution. Australian Entomologists' Magazine. 7: 45-59. 



235 



Threatened Swallowtail Butterflies of the World 

Battus zetides Mumoe, 1971 VULNERABLE 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Battus zetides, the Zetides Swallowtail, is recorded from both 
Haiti and the Dominican Republic on the Caribbean island of Hispaniola. It occurs 
only very locally in stands of deciduous forest at high altitudes. With severe human 
population pressure and habitat destruction in Haiti, moderated only slightly in the 
Dominican Republic, the status of Battus zetides is cause for serious concern. The 
species may already be extinct in Haiti. 

Description The sexes are similar in general appearance, with a forewing 
length of 35-40 mm (males) or 39^4 mm (females) (1,3) (Plates 2.6 and 2.7). 
Battus zef/rf^5 was originally described as zeto (Westwood, 1847) (10), later found to 
be an invalid homonym (9). 

UFW with a submarginal row of yellow ochre spots over a dark, chocolate brown 
ground colour (1, 9, 10). UHW with a medium length tail and a scalloped edge, a 
fringe of yellow crescents and a subdiscal band of large orange or yellow ochre spots 
(1,9, 10), slightly darker than on the UFW (3). The tailless specimen figured in 
reference (9) is mutilated (see 1,6). 

LFW similar to the UFW, with orange or pale yellow submarginal spots and discal 
marks (1, 9, 10). LHW dark brown basally, with a boldly contrasting silvery-white 
discal band, a postdiscal row of red lunules and a row of white submarginal spots 
fringed with yellow (1). 

The recently-described caterpillars have yellow-orange longitudinal stripes on a 
tan or darker brown ground colour (13). Dorso-lateral tubercles are only found on 
the second and third thoracic segments and the first, eighth and ninth abdominal 
segments (13). The pupa is dark apple green, yellowish dorsally, with a tapering 
thoracic process (13). 

Distribution Battus zetides is found on the Caribbean island of Hispaniola 
in the Greater Antilles (9, 10). It has been recorded both from Haiti (9, 10) (although 
not since the 1950s) and the Dominican Republic (3), the two countries that share the 
island. The history of its capture and recent rediscovery in southern Dominican 
Republic is well described in recent papers by Gali and Schwartz (3) and Weintraub 
(13). The three presently known localities are in Pedernales Province, Indepencia 
Province and Peravia Province (13). 

Habitat and Ecology Battus zetides , for many years very poorly known , was 
recently found in some numbers on the southern slopes of the Massif de la Selle in 
Pedernales Province in the southern Dominican Republic (3). Two further localities 
with smaller populations have also been identified, one on the northern slopes of the 
Massif in Indepencia Province, the other in Peravia Province in the Cordillera 
Central (13). The habitat of the larger population on the Massif was an enclave of 
upland broad-leaved deciduous forest at an altitude of about 1000-1300 m, 
completely surrounded by forests of Pinus occidentalis (3, 13). On the northern 
slopes the species flies slightly lower, at 900-1000 m, and in the Cordillera Central the 
locality is at 1450 m (13). These deciduous woodlands are generally dense but the 
butterflies prefer to fly in sunny glades and rides, soaring slowly and deliberately at a 
general level of about 3.5 m, but ranging as high as 10 m and as low as 2 m (3). The 

236 



Battus zetides 

butterflies were not found in small patches of deciduous trees in a ravine on the Massif 
de la Selle, an observation that led to speculation that Battus zetides may require 
extensive stands of deciduous forest for its survival (3). However, both the colonies 
recently discovered were in medium-sized patches of remnant forest of less than 10 ha 
(13). Adults are on the wing between 9.00 a.m. and 4.00 p.m. except when the 
weather is overcast. Nectar plants include species from the Boraginaceae, Labiatae, 
Leguminosae and Verbenaceae. Adults have been recorded in March- April and 
July-October and are probably continuously brooded, although more common in the 
wet season (13). The foodplants are believed to be Aristolochia bilabiata and 
Aristolochia montana (13). The eggs are laid in clusters of up to 15 and are gregarious 
in the first two instars. 

Deciduous broad-leaved forests were once extensive in Hispaniola, growing at 
altitudes above 800 m in Haiti (2, p. 175) and at variable altitudes in the Dominican 
Republic, beginning at 500-900 m and reaching 1000-2200 m (2, p. 299). Pines may 
also be dominant at these levels, interdigitating in a complex pattern of stratification 
which seems to vary considerably according to rainfall, aspect, slope and other 
physical features. 

Threats Hispaniola as a whole has a very high population density and this is 
the basic reason for the threats to this butterfly and other forms of wildlife. The total 
human population of the island is now over 10 million, living in an area of 
76 150 sq. km, but the Haitian sector is more seriously overcrowded than the 
Dominican Republic, with over 4.8 million people (1978) living in 27 750 sq. km 
(2, 5, 8). This represents the highest population density in Latin America. Three- 
quarters of Haiti and a smaller, but nevertheless large, proportion of the Dominican 
Republic is mountainous, with a natural cover of forest. With a high proportion of 
rural people (85 per cent in Haiti (8)), demand for land and forest products is high. 
Many forests are degenerating through over-exploitation for charcoal, stakes and 
building timbers (2). Others are cleared and burnt for agriculture by subsistence 
farmers and for plantations of coffee and cocoa (2, 3). 

In Haiti, degradation of soils is particularly serious. Without proper conservation 
efforts on the mountain slopes, slash-and-burn agriculture exposes the top-soil to 
erosion by the heavy rains of 1000-2000 mm per year. Annual fires exacerbate the 
problem, preventing forest regrowth and resulting in a bare landscape of impover- 
ished grassland dotted with spiny legumes and cactus plants (2). Forest remains only 
in steep ravines. In Haiti total forest cover was reduced from 80 per cent to 9 per cent 
(including secondary fallow) between 1958 and 1978 (12, p. xi). An estimate in 1980 
shows a further reduction to only 1.7 per cent (2). In 1980 the estimated cover of 
broad-leaved forest in Haiti was 36 million ha, but the projection for 1985 was only 30 
million ha, a loss of nearly 17 per cent in five years (2). As stated above, Battus zetides 
has not been recorded in Haiti since the 1950s and may be extinct there. 

In the Dominican Republic the human population is less dense and the pressures 
on natural habitats are somewhat ameliorated. Nevertheless, there is no doubt that 
the rich broad-leaved deciduous forests and pine forests of middle altitudes are under 
increasing threat. On the slopes of the Massif de la Selle, where Battus zetides was 
recently found, most of the deciduous forest has been cleared for coffee plantations 
and subsistence crops (3). In 1980 the area of broad-leaved forest in the Dominican 
Republic was estimated at 444 million ha, spread between the eastern region, the 
Sierra de Neiba, the Cordillera Central and the Sierra de Baoruco (2). Projections for 
1985 were optimistic, with an expected reduction to 432 million ha, a loss of only 2.7 
per cent in five years (2). Nevertheless, there is good reason to suppose that Battus 

237 



Threatened Swallowtail Butterflies of the World 

zetides does not occur throughout these forests. It is very localized in relict patches of 
deciduous forests, none of which is protected and all of which are vulnerable to 
deforestation. The two smaller populations are particularly vulnerable to shifting 
agriculturalists, a proportion of which is comprised of illegal squatters crossing the 
border from Haiti (13). The larger population is situated on a bauxite mining 
concession and is currently relatively free of threat. However, the price of tin is 
depressed and if the concession is not worked it may be returned to the government. 
In this event the site would almost certainly be deforested and put to agricultural use. 
This site also contains populations of the Endangered Haitian Solenodon (Solenodon 
paradoxus) and the Indeterminate Hispaniolan Hutia {Plagiodontia aedium) 
(12, 13). 

Conservation Measures Haiti has such a large and widely distributed 
human population that it can only afford minimal reserved areas (5 , 12) . In 1980 there 
was one small and badly damaged reserve (La Citadelle) and a further five proposed 
(5). There is no evidence that Battus zetides survives in any of these areas, or indeed in 
Haiti as a whole. If it does survive, there can be little doubt that it is Endangered. If 
suitable relict patches of deciduous forest can be found, they should be afforded all 
possible protective measures. 

In the Dominican Republic there are five protected areas of which two, Armando 
Bermudez N.P. and Jose del Carmen Ramirez N. P. , He on the northern and southern 
slopes of the Cordillera Central (5). Battus zetides may occur in these parks, both of 
which include broadleaved hardwoods and Pinus occidentalis at higher altitudes (5). 
Confirmation of the presence of Battus zetides and other forest butterflies in these 
parks would be very valuable, not only for the butterfly but also for increasing the 
general appreciation of these important protected areas. Once found, studies of the 
foodplants and young stages of the butterfly would be required in order to assess 
possible management measures. At the same time, further data on the distribution of 
Battus zetides throughout the Dominican Republic are needed, together with an 
assessment of its status in Haiti. 

The most important known site for B. zetides is the mining concession site on the 
Massif de la Selle . All possibilities for declaring this locality a reserve for the butterfly 
should be explored. 



References 

1. D'Abrera, B. (1981). Butterflies of the Neotropical Region. Part 1. Papilionidae and 
Pieridae. Lansdowne Editions, Melbourne, in association with E.W. Ciassey, Faringdon. 
172 + xvi pp. 

2. FAO/UNEP (1981). Proyecto de Evaluacion de los Recursos Forestales Tropicales. Los 
Recursos Forestales de la America Tropical. FAO, Rome. 343 pp. 

3. Gali, F. and Schwartz, A. (1983). Battus zetides in the Republica Dominicana. Journal of 
the Lepidoplerists' Society 37: 170-171. 

4. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

5. lUCN ( 1982). lUCN Directory of Neotropical Protected Areas. Tycooiy, Dublin. 436 pp. 

6. Lewis, H.L. (1973). Butterflies of the World. Harrap, London. 312 pp. 

7. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supp\emenl 17: 1-51. 

8. Paxton, J. (1981). The Statesman's Year-Book. Macmillan Press, London. 1696 pp. 

9. Riley, N.D. (1975). A Field Guide to the Butterflies of the West Indies. Collins, London. 244 
pp., 24 col. pi. 

238 



Battus zetides 

10. Rothschild, W. and Jordan, K. (1906). A revision of the American Papihos. Novitates 
Zoologicae 13(3): 411-752. (Facsimile edition ed. P.H. Arnaud, 1967). 

11. Schwartz, A. (1984). In litt.. 16 May. 

12. Thornback,J.andJenkins,M. (1982). The lUCN Mammal Red Data Book Part LIVC^, 
Gland. 516 pp. 

13. Weintraub, J.D. (unpublished manuscript). Notes on the biology of Battus zetides Munroe 
(Lepidoptera: Papilionidae). 



239 



Threatened Swallowtail Butterflies of the World 

Parides ascanius (CrameT, 1775) VULNERABLE 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Parides ascanius, the Fluminense or Ascanius Swallowtail, is a 
strikingly beautiful butterfly which occupies only a small fraction of its potential 
'restinga' habitat (subcoastal swamps and thickets) in the state of Rio de Janeiro, 
south-eastern Brazil. It is a primitive species, lacking vigour, and most of its habitat is 
being drained and subdivided for residential, industrial, and recreational purposes. 
Many colonies active before 1970 are now extinct. Because of habitat pressure it is on 
the Brazilian list of animals threatened with extinction, the first insect so designated. 
Only about ten self-supporting colonies are now known, many in areas under 
pressure for development. Large colonies need to be located in regions suitable for 
establishment of conservation units. 

Description The sexes of this relatively large, tailed butterfly are similar in 
appearance although the female lacks the androconial fold found on the hindwing of 
the male (2, 7) (Plate 2.2). Parides ascanius is the nominate species of the ascanius 
species-group, which includes 1 1 species in published lists (4, 5), but is expected to be 
reduced to nine in a forthcoming checklist of neotropical Lepidoptera (1). 

UFW black with a broad white median band (2). 

UHW also black, with a deeply scalloped outer margin, a relatively long tail, and 
an extension of the median band of the forewing to the inner margin. This wing is 
washed with rose coloured scales, especially anally, and there is also a row of red, 
hourglass-shaped submarginal spots (2). 

Distribution Parides ascanius is presently known only from widely scat- 
tered points near the coast of the state of Rio de Janeiro, south-east Brazil, between 
the mouth of the Rio Parafba do Sul (to the north) and Itaguai (to the south-west) (3) . 
Formerly it may have occurred in favourable habitats on the coast of Sao Paulo state, 
but it is not known there today despite substantial searches (6). 

Habitat and Ecology Parides ascanius inhabits only subcoastal and lowland 
swamps and thickets where its larval foodplant Aristolochia macroura (Aristolo- 
chiaceae) is abundant, and suitable flowers (mostly Compositae and Verbenaceae) 
are available to adults throughout the year (6). Very patchy distribution and 
wide-ranging male promenading indicate dispersed, possibly unstable, colonies with 
extensive competition from the sympatric P. zacynthus and P. anchises nephalion, 
the two most advanced members of the genus , both strong and aggressive species (6) . 
P. ascanius occurs mostly on sandy soils in vegetation of low to medium height and 
flies only in the sun during the morning and late afternoon (6). It can be reared quite 
easily, and kept in captivity in proper conditions (6). The habitat is very rich in 
endemic plants and animals, almost all of which have far wider and less patchy 
distributions than P. ascanius, which seems to be a relict (6). 

Threats Habitat destruction throughout its range, but particularly near the 
city of Rio de Janeiro, has rendered P. ascanius increasingly scarce (6). Many 
colonies known before 1970 no longer exist because of swamp drainage for 
development into recreational areas, banana plantations, pasture or buildings (6). In 
addition, there is some danger from competition with other Parides species (6). As a 

240 



Parides ascanius 

primitive and relict species with unusual affinities, P. ascanius has value for scientific 
study. Its beauty is also widely recognised and specimens sell for a high price when 
available. 

Conservation Measures In an executive action (3481-DN) on 31 May, 1973, 
the Brazilian National Parks agency (through the IBDF, Brazilian Institute for Forest 
Development) placed P. ascanius on the official list of Brazilian animals threatened 
with extinction (as the synonym Battus orophobus D'Almeida). It is the only insect 
presently given this status, and is thereby protected from commerce (6). Eurytides 
lysithous harrisianus has also been proposed for listing (see separate review). 

The recently-established (1974) Federal Biological Reserve of Pogo das Antas, 
north-east of Rio de Janeiro, is a 5000 ha area which includes at least 1000 ha of ideal 
P. ascanius habitat. Suitable areas may be extended by river management. P. 
ascanius is known to live in the reserve in adequately large colonies (6). 

A small but permanent colony of P. ascanius is also present in a swampy forest of 
less than 2 ha in the Parque Reserva Marapendi, Restinga de Jacarepagua in Rio de 
Janeiro. However, the presumed source of this colony is a patch of the food plant with 
its own colony in an unprotected nearby swamp, which is due to be drained (6). The 
Parque Reserva Marapendi is too small and inadequately protected. Attempts to 
transplant the food plant into suitable parts of the reserve have failed . Key parts of the 
nearby swamp should be protected from draining and building in order to ensure the 
future of this small colony (6). Should this colony fail, however, the population left in 
the Reserve of Pogo das Antas cannot be assigned full and exclusive responsibility for 
preservation of P. ascanius. The main hope for the species lies in 1000 sq. km of 
almost impenetrable swamps in lowland Rio de Janeiro , south and west of Itaguai and 
Campos. Colonies may well be flourishing there, but this is not certain (6). When 
possible, surveys in this region should be carried out. 

Commerce in wild specimens should be discouraged internationally as well as 
nationally (6). Official captive breeding of the species in its habitat could be 
encouraged (6). Populations in reserves need to be extended and carefully moni- 
tored, to guarantee the well-being of this butterfly in coastal Rio de Janeiro (6). 

References 

1. Brown. K.S., Jr. (1984). In //»., 4 January. 

2. D'Abrera, B. (1981). Butterflies of the Neotropical Region. Parti. Papilionidae and Pieridae. 
Lansdowne Editions, Melbourne, xvi -t- 172 pp. 

3. D'Almeida, R.F. (1966). Catdlogo das Papilionidae Americanos. Sociedade Brasileira de 
Entomologia, Sao Paulo, Brazil. 366 pp. 

4. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

5. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
£«rowo/ogwr Supplement 17: 1-51. 

6. Otero, L.S. and K.S. Brown, Jr. (in press). Biology and ecology of Parides ascanius 
(Cramer, 1775) (Lepidoptera, Papilionidae), a primitive butterfly threatened with extinc- 
tion. Atala. 

1. Rothschild, W. and Jordan, K. (1906). A revision of the American Papilios. Novitates 
Zoologicae 13(3): 411-752. (Facsimile edition ed. P.H. Arnaud, 1967). 

This review has been adapted from an entry in The lUCN Invertebrate Red Data 
Book, whose authors and contributors are gratefully acknowledged. 



241 



Threatened Swallowtail Butterflies of the World 

Parities hahneli {Staudinger, 1882) RARE 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Parides hahneli, Hahnel's Amazonian Swallowtail, is an exquis- 
itely beautiful butterfly that has been known for nearly a century, but only in the past 
decade have any reasonably dense colonies been discovered. It occupies a very 
specialized habitat in the lower middle Amazon Basin of Brazil and only three 
localities have ever been found for it. Until the most recent locality was found, just 20 
collected specimens were known. A very primitive species, it may hold the key to the 
evolution of this swallowtail genus. 

Description Parides hahneli, a beautiful black and yellow butterfly with a 
wingspan of 80 -100 mm, is the only tailed member of its genus in the Amazon basin 
of Brazil. It resembles sympatric ithomiine butterflies {Methona, Thyridia), which it 
apparently mimics, possibly to gain protection. The sexes are similar (5, 6, 7, 10). 
The larva is dark brown with yellow rings and resembles that of P. pizarro steinbachi 
and P. vertiimnus (8, 9), other members of the same species-group. P. hahneli is the 
most primitive member of the aeneas species-group, which includes 25 species 
according to current lists (9, 10). However, the assemblage is probably artificial and 
will be divided up in the forthcoming checklist of neotropical Lepidoptera (3) , leaving 
only 12 species in the aeneas group. Six species in the group are possibly threatened, 
and are reviewed in the following pages. 

UFW/LFW black, long, narrow, and rounded, with three broad bands of 
yellow-grey patches (5). 

UHW/LHW also black, with a scalloped outer margin and a long spatulate tail. 
There is a large patch, yellower than those of the forewing, covering all but the 
borders of the hindwing. The LHW has a red spot near the tail (5). 

Distribution Until 1970 Parides hahneli was known only from the region of 
Maues, south of the middle Amazon Basin in Brazil, but it was then discovered in the 
Rio Arapiuns area more to the east. There are sight and unconfirmed capture records 
from the region of Manaus (and Manacapuru) to the north-west of Maues, but no 
other records have been located for this distinctive species in spite of extensive 
searching by lepidopterists and commercial collectors over many years (2, 5). The 
butterfly is very rare and has a patchy distribution. 

Habitat and Ecology Parides hahneli is apparently restricted to ancient 
sandy beaches now covered by scrubby or dense forest vegetation (2). Like many 
other troidine swallowtails using Aristolochia as a host plant, P. hahneli is probably 
extremely localized in occurrence, limited by the distribution and density of the plant. 
In addition, the adults need to feed on a continuous supply of nectar plants. The 
particular host species of Aristolochia for P. hahneli is not identified but it is also used 
at least by the sympatric P. chabrias ygdrasilla (1,9). Sympatric butterflies include 
Heliconius egeria, also partial to sandy areas and very local in conjunction with its 
foodplant, Passiflora glandulosa. Many plants are endemic to the Maues region. In 
general, the butterfly seems to occupy a poor but very specialized habitat which is not 
very diverse but has a high degree of endemism, demonstrated by this insect and 
other organisms (2). P. hahneli may suffer from food plant competition with 
sympatric troidine swallowtails (2). 

242 



Parides hahneli 

P. hahneli retains a primitive pattern, morphology and tails, and is related to 
species of southern Brazil and Central America. It seems to represent a bridge 
between these archaic Parides, now pushed to the margins of the Neotropics, and the 
modern species in the P. aeneas and P. anchises groups. It persists in a marginal 
biotope in the central Amazon, where it is extremely rare. This situation offers many 
opportunities for research on tropical ecological and evolutionary processes, which 
can only be solved in the context of the living organism (2). 

Threats Over-collecting for commercial purposes represents the only 
current threat. However, there is potential for deleterious habitat changes prior to its 
ecological requirements being fully understood (2). The species is beautiful (and 
rather bizarre), participates in mimicry rings not usual in other Parides, and currently 
sells for a very high price (2). It is possible to rear this species in captivity (11), but 
collecting sites are a closely guarded secret in view of its commercial value. Scientific 
work on the species has hardly begun. 

Conservation Measures The butterfly may occur in the Amazon National 
Park between Itaituba and Maues, but this is not likely. A search should be made in 
the small Ducke reserve near Manaus (2). A general study of the habitat and colonies 
of P. hahneli should be undertaken, with a view towards proposing a reserve near 
Maues or Arapiuns. Such a reservation would permit the survival of this swallowtail 
as well as the many other endemic organisms of the region, and would be a significant 
component of the northern Brazilian land conservation system (2). Repeatedly 
scarred by rivercourse changes and affected by Pleistocene drying, this area is unique 
in its biological properties. 

References 

1 . Brown Jr. , K.S. , Damman, A. J. and Feeny , P. ( 1981 ). Troidine swallowtails (Lepidoptera: 
Papilionidae) in southeastern Brazil: natural history and foodplant relationships. Journal 
of Research on the Lepidoptera 19: 199-226. 

2. Brown, K.S., Jr. (1982). In fill.. 19 January. 

3. Brown, K.S., Jr. (1983). In lilt. ,23 October. 

4. Brown, K.S., Jr. (1983). In liii., 19 September. 

5. D'Abrera, B. (1981). Butterflies of the Neotropical Region. Parti. Papilionidae and 
Pieridae. Lansdowne Editions, Melbourne, in association with E.W. Classey, Faringdon. 
172 + xvi pp. 

6. D'Almeida, R.F. (1966). Catdlogo dos Papilionidae Americanos. Sociedade Brasileira de 
Entomologia. Sao Paulo, Brazil. 366 pp. 

7. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

8. Moss, A.M. (1919). The papilios of Para. Novitates Zoologicae 26: 295-319. 

9. Moss, A.M. Manuscript material preserved with the Moss collection in the British Museum 
(Natural History). 

10. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supplement 17: 1-51. 

11. Rothschild, W. and Jordan, K. (1906). A revision of the American Papilios. Novitates 
Zoologicae 13(3): 411-752. (Facsimile edition ed. P.H. Arnaud, 1967). 

This review has been adapted from an entry in The lUCN Invertebrate Red Data 
Book, whose authors and contributors are gratefully acknowledged. 



243 



Threatened Swallowtail Butterflies of the World 

Parides pizarro Staudinger, 1884 INSUFFICIENTLY KNOWN 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Parides pizarro is a little icnown swallowtail from Peru and 
Brazil. There are two subspecies (three if P. steinbachi is included), one fairly 
common but the other known from very few specimens. Further research is needed to 
confirm its conservation status. 

Description Parides pizarro is a relatively large butterfly with sexes that are 
similar in general appearance (2,7). The forewing is entirely black, and the hindwing, 
which has a scalloped outer margin, is black with a cream-coloured discal patch 
consisting of three to four discal spots in the male, and three to six in the female (2,7). 

Distribution Two subspecies of P. pizarro, P. p. pizarro and P. p. 
kuhlmanni, are currently recognized in the literature, although some authorities 
believe that P. steinbachi (see next review) represents a third (1,5). Until new 
findings are published we adopt those of the most recent published work (4). The 
situation is confused by D'Abrera (2), who lists kuhlmanni as a subspecies of P. 
steinbachi, a view not accepted by other authorities (1,5). P.p. pizarro, is from the 
upper Amazon in north-eastern Peru (formerly part of Ecuador) and has been 
recorded in the Departments of Loreto, San Martin and UcayaU (3, 7). P.p. 
kuhlmanni is known from two specimens from the Department of Madre de Dios in 
eastern Peru and was originally described from a single male specimen found in the 
neighbouring state of Acre in western Brazil. A further two Brazilian specimens are 
held in the National Museum in Rio de Janeiro (1). P. pizarro is in the aeneas 
species-group (4, 6). 

Habitat and Ecology Parides pizarro occurs in lowland primary forest, 
probably below 500 m (5). The immature stages and foodplant are unknown, but the 
latter is likely to be Aristolochia. More information is needed on the life history, 
biology and ecology of P. pizarro. 

Threats Cannot be assessed without further data. 

Conservation Measures P. p. pizarro is said to be moderately common in 
the reaches of the upper Amazon of eastern Peru , but P. p . kuhlmanni has apparently 
been recorded only from two specimens from Peru and a small number from Brazil. 
The Peruvian specimens were both taken in the Tambopata Nature Reserve, a 
privately managed 5000 ha site on the Tambopata River, 25 km south-east of Puerto 
Maldonado (5). This extraordinarily rich reserve has so far yielded 893 species of 
butterflies (5). Although P. pizarro may be effectively protected by this Reserve, 
there is still insufficient information to be sure. Further research on its biology and 
habits are needed before conservation needs can be properly assessed. 



References 

1. Brown, K.S., Jr. (1983). In to., 23 October. 

2. D'Abrera, B. (1981). Butterflies of the Neotropical Region. Parti. Papilionidae and Pieridae . 
Lansdowne Editions, Melbourne, xvi + 172 pp. 

244 



Parides pizarro 

3. D' Almeida, R.F. (1966). Catdlogo dos Papilionidae Americanos . Sociedade Brasileira de 
Entomologia. Sao Paulo, Brazil. 

4. Hancock, D.L. (1983). The classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

5. Lamas, G. (1983). In litt.A April. 

6. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist StuppXcmsnX 17: 1-51. 

7. Rothschild, W. and Jordan. K. (1906). A revision of the American Papilios. Novitates 
Zoologicae 13(3): 411-752. (Facsimile edition ed. P.H. Arnaud, 1967). 



245 



Threatened Swallowtail Butterflies of the World 

Parides steinbachi Rothschild, 1905 INSUFFICIENTLY KNOWN 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Parides steinbachi is a very poorly known swallowtail from 
Bolivia. Its precise distribution is unclear and its taxonomic status questionable. 
Some authorities would place it as a subspecies of P. pizarro. More data are needed 
to assess the threats to this apparently very rare taxon. 

Description Parides steinbachi is a relatively large butterfly, with a 
forewing length of 35^0 mm (8). The female is larger than the male, but otherwise 
similar in appearance, with minor differences in wing markings (2,8). P. steinbachi is 
listed as a good species by Hancock (4) but is considered by a number of authorities 
to be a subspecies of P. pizarro (1,6) (see previous review) and is expected to be 
published as such in a forthcoming checklist of neotropical Lepidoptera. 

UFW/LFW ground colour black with a cream or white median patch (2, 8). 

UHW/LHW black with a scalloped outer margin, white marginal spots and a discal 
patch of red spots with an innermost white spot. LHW has a red spot in both sexes, 
close to the anal angle (2, 8). 

The caterpillars are likely to be fleshy with segmental tubercles, and the pupae 
with prominent abdominal ridges (7). Specimens of larvae and pupae are held in the 
Moss collection in the British Museum (Natural History) (1). 

Distribution This butterfly is confined to Bolivia (8). The localities given in 
(8) are rather confusing, the name Mapiri being not uncommon in Bolivia. However, 
the balance of evidence is that P. steinbachi is mainly or entirely in eastern Bolivia 
(3, 8). Zischka (10) reports the species from Santa Cruz and Buena Vista up to the 
Brazilian border. There is also a specimen from Santa Cruz province in the Tyler 
collection (9). 

Habitat and Ecology The original series was collected between February 
and June, apparently from altitudes between 200 and 1000 m in the eastern Andes. 
There is no information on the life history of the species, but in common with other 
troidine swallowtails, it probably feeds on Aristolochia (7). 

Threats Threats to this very rare and poorly known butterfly cannot be 
assessed at present. The economic situation in Bolivia is presently so serious that the 
entire national park system is believed to be under threat. Lack of funds, equipment, 
management presence and enforcement combine to negate the intended protection 
of Bolivia's wildlife. 

Conservation Measures No rational conservation measures can be pro- 
posed without more information on the biology, distribution and habitat of the 
species. Large areas of eastern Bolivia remain relatively undeveloped. The 
Huanchaca National Park is over 5000 sq. km in extent (5), but it remains to be seen 
whether Parides steinbachi occurs in the region. 

References 

1. Brown, K.S., Jr. (1983). In litt.,23 October. 

2. D'Abrera, B. (1981). Butterflies of the Neotropical Region. Part 1. Papilionidae and 

246 



Parides steinbachi 

Pieridae. Lansdowne Editions, Melbourne, xvi + 172 pp. 

3. D'Almeida, R.F. (1966). Catdlogo dos Papilionidae Americanos. Sociedade Brasileira de 
Entomologia. Sao Paulo, Brazil. 

4. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

5. lUCN (1982). lUCN Directory of Neotropical Protected Areas. Tycooly International, 
Dublin. 436 pp. 

6. Lamas, G. (1983). In litt.,4 April. 

7. Munroe. E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supplement 17: 1-51. 

8. Rothschild. W. and Jordan, K. (1906). A revision of the American Papilios. Novitates 
Zoologicae 13(3): 411-752. (Facsimile edition ed. P.H. Arnaud, 1967). 

9. Tyler, H.A. (1983). In litt., 30 September. 

10. Zischka, R. ( 1950). Catalogo de los insectos de Bolivia. Folia Universidad Cochabamba 4: 
51-56. 



247 



Threatened Swallowtail Butterflies of the World 

Parides coelus Boisduval, 1836 INSUFFICIENTLY KNOWN 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Parides coelus is an extremely rare and poorly known troidine 
swallowtail from the state of Cayenne in French Guiana. Further research is needed 
before conservation measures can be proposed. 

Description A large butterfly with a forewing length of about 49 mm in the 
male and 55 mm in the female (1). The sexes are similar but the male has prominent 
white androconial folds and smaller red spots on the hindwings (1, 4, 5, 6). P. coelus 
is a member of the aeneas species-group (5). 

UFW/LFW ground colour blackish-brown, lighter in the apical and subapical 
regions and with a diffuse white spot at the apex of the cell , slightly larger on the LFW 
(1,6). 

UHW/LHW with a deeply scalloped outer margin, no distinct tail but a discal band 
of four bright red spots (six on the LH W) contrasting with the blackish-brown ground 
colour. A large whitish or grey androconial fold is present on the anal margin of the 
male, replaced by an extra pair of red-orange spots in the female (1,6). 

Distribution Parides coelus is only known from French Guiana in South 
America. The type female was caught in the area of Cayenne (2). 

Habitat and Ecology Virtually nothing is known of this extremely rare 
species. The male and female were described as separate species, although they are 
fairly similar in appearance (1,2, 6). In common with most other troidine swallow- 
tails, the foodplant is likely to be Aristolochia, the larva fleshy with segmental 
tubercles, and the pupa with prominent edges or rows of tubercles (5). 

Threats French Guiana has immense tracts of forests and it seems unlikely 
that forest destruction could be responsible for the apparent rarity of this species. 
Nevertheless, a new highway has been built from the coast to the Brazilian border, 
bringing settlement and forest destruction in its wake. Although commercial logging 
is still Umited in extent, it is expanding rapidly. Further study is needed to assess what 
threats, if any, are affecting Parides coelus. Many troidine swallowtails feeding on 
Aristolochia have very patchy distributions. 

Conservation Measures So little is known of this rare butterfly that 
conservation measures cannot be proposed. It is first necessary to discover its 
distribution, habitat and biology. There is no information on its presence or absence 
in the two protected areas on the mainland of French Guiana, namely the Sinnamari 
and Basse Mana National Parks (3). The former is mainly mangrove and the latter 
coastal wetland (3). French Guiana is a French Overseas Department subject to 
French laws. If necessary, Parides coelus could be placed on the French list of 
protected species, but research and protection of its habitat would be more useful 
objectives at this stage. 

References 

1. D'Abrera.B. (1981). Butterflies of the Neotropical Region. Parti. Papilionidae and Pieridae . 
Lansdowne Editions, Melbourne, xvi + 172 pp. 

248 



Parides coelus 

2. D'Almeida, R.F. (1966). Catdlogo dos Papilionidae Americanos . Sociedade Brasileira de 
Entomologia. Sao Paulo, Brazil. 

3. lUCN (1982). lUCN Directory of Neotropical Protected Areas. Tycooly International, 
Dublin. 436 pp. 

4. Lewis, H.L. (1973). Butterflies of the World. Harrap, London. 312 pp. 

5. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
En/owo/ogwr Supplement 17: 1-51. 

6. Rothschild, W. and Jordan, K. (1906). A revision of the American Papilios. Novitates 
Zoologicae 13(3): 411-752. (Facsimile edition ed. P.H. Arnaud, 1967). 



249 



Threatened Swallowtail Butterflies of the World 

Parides klagesi Ehrmann, 1904 INSUFFICIENTLY KNOWN 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Parides klagesi is a small, very rare and poorly known troidine 
swallowtail from the northern and eastern foothills of the Bolivar highlands in 
Venezuela. Further research is needed before conservation measures can be 
proposed. 

Description Parides klagesi is in the aeneas species group (4, 6) and 
somewhat resembles P. coelus, but is a much smaller butterfly with a forewing length 
of only about 30 mm (2). The male and female are very similar in appearance, the 
former being only slightly darker and smaller (2). 

UFW/LFW black with a white median patch and the distal part of the forewing 
somewhat paler than the rest. 

UHW/LHW black with a scalloped outer edge, and a discal band of six pinkish 
spots, in the female or three in the male, which has a whitish anal hair-pouch. 

Distribution Parides klagesi is only known from northern and eastern 
Venezuela, where it has been recorded from the region of the Caura, Suapure and 
Caroni rivers, which flow north into the Orinoco from the highlands of Bolivar 
(1,3,7). 

Habitat and Ecology Little information on the habitat or ecology of this 
species has been recorded. As with most other troidine swallowtails, the foodplant is 
likely to be Aristolochia, the caterpillar fleshy and tuberculate, and the pupa with 
tubercles or prominent edges (6). Specimens have been taken in recent years in the 
region between the Pan American highway and the lower reaches of the River Caroni 

(1). 

Threats No information on this region of Venezuela has been obtained and 
with so little known about the butterfly it is not possible to assess threats at present. 
Undoubtedly the building of the Pan American Highway through eastern Venezuela 
will attract development and settlement, possibly reducing the range of the butterfly. 

Conservation Measures Parides klagesi, like its close relative in French 
Guyana, P. coelus, is too poorly known to permit rational conservation measures. 
Further research on its distribution, habitat and biology is needed. The only national 
park in the Bolivar highlands is Canaima, a 30 000 sq. km park 600 km to the east in 
La Gran Sabana (5). P. klagesi may occur in parts of the Park, but has not been 
recorded there so far. The Pan American Highway appears to bisect the Park; the 
effect of this on the wildlife remains to be assessed. 



References 

1. Brown, K.S., Jr. (1983). In lilt.. 23 October. 

2. D'Abrera, B. (1981). Butterflies of the Neotropical Region. Parti. Papilionidae and Pieridae . 
Lansdowne Editions, Melbourne, xvi + 172 pp. 

3. D'AImeida, R.F. (1966). Catdlogo dos Papilionidae Americanos . Sociedade Brasileira de 
Entomologia. Sao Paulo, Brazil. 

250 



Parides klagesi 

4. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1—48. 

5. lUCN (1982). lUCN Directory of Neotropical Protected Areas. Tycooly International, 
Dublin. 436 pp. 

6. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
£/7/owo/og/5/ Supplement 17: 1-51. 

7. Rothschild, W. and Jordan, K. (1906). A revision of the American Papilios. Novitates 
Zoologicae 13(3): 411-752. (Facsimile edition ed. P.H. Arnaud. 1967). 



251 



Threatened Swallowtail Butterflies of the World 

Parides burchellanusWestwood, 1872 VULNERABLE 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Parides burchellanus is a large, velvet-black butterfly which 
inhabits riverine forest in central Brazil. It is very rare and its habitat is subject to 
flooding and forest clearance. 

Description Parides burchellanus is a large, velvet-black butterfly with a 
forewing length of about 45 mm. The forewing is without markings except for a fringe 
of some small , white spots. The UHW has a row of small , red postdiscal spots which 
are smaller and paler on the lower surface. The hindwing has a scalloped outer 
margin, no tails, and a large hair-pouch on the anal margin of the male. Apart from 
this, the sexes are alike. The species is placed in the aeneas group by Munroe (5). It is 
believed to be very close to, and possibly even conspecific with, P. aeneas from 
Amazonas, with which a putative intermediate is known (1). 

Distribution Parides burchellanus is found in central Brazil from northern 
Goias state to western Sao Paulo (1, 3, 4, 6). The most recent specimens have come 
from the Rio Maranhao on the border of Goias state and the Federal District of 
Brazilia (2). A specimen described by Schaus as being from Bolivia requires 
confirmation. It was sent to him from a correspondent who had been resident in Rio 
de Janeiro (6). 

Habitat and Ecology Parides burchellanus is always associated with gallery 
(riparian) forest along rivers running through the cerrado landscape of central Brazil. 
Away from the rivers the land is dry for much of the year, but where ground-water 
occurs along ravines, streams and river-banks tall forest develops in a humid 
microclimate. P. burchellanus is rarely far from the river bank and may be seen flying 
a few centimetres above the water (1,3). Colonies contain between 10 and 50 
individuals in a favourable season; apparently six generations develop per year, as in 
other Brazilian Parides, with a low period in June-July (1). The foodplant remains 
unknown but, in common with other troidine swallowtails, is likely to be 
Aristolochia. 

Threats Very few individuals or colonies have been captured or seen since 
the species was described over a century ago. It is associated with the even rarer P. 
panthonus numa which has the same pattern and habits, and has not been seen since 
1920 (1). Colonies are subject to elimination by changes in water level. Natural floods 
are quite common in the cerrado landscape, but riverside clearing and agriculture 
increase their ferocity downstream. Changes in habitat and microclimate by cutting 
and opening up the gallery forest will also cause colonies to disappear. Colonies 
recently found along the Rio Maranhao may have since been destroyed by flooding 
(2). 

Conservation Measures More information is needed on the precise locali- 
ties of colonies and their breeding requirements. No conservation measures have 
been taken, but there are laws prohibiting the elimination of riparian vegetation. 
Once colonies have been located, these laws could be enforced. No captive breeding 
has been reported, but it should be feasible for river dwellers in the region (1). 

252 



Parides burchellamis 

However, the species is not particularly striking in appearance and may not be 
suitable for commercial development. 

References 

1. Brown, K.S., Jr. (1983). In litt.,6 April. 

2. Brown, K.S., Jr. (1983). In //«., 23 October. 

3. Brown, K.S., Jr. and Mielke, O.H.H. (1967). Lepidoptera of the central Brazil plateau. 1. 
Preliminary list of Rhopalocera (continued): Lycaenidae, Pieridae, Papilionidae, Hesperii- 
dae. Journal of the Lepidopterists' Society 21: 145-68. 

4. D'Almeida, R.F. (1966). Catdlogo dos Papilionidae Americanos. Sociedade Brasileira de 
Entomologia. Sao Paulo, Brazil. 

5. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supplement 17: 1-51. 

6. Rothschild, W. and Jordan, K. (1906). A revision of the American Papilios. Novitates 
Zoologicae 13(3): 411-752. (Facsimile edition ed. P.H. Arnaud, 1967). 



253 



Threatened Swallowtail Butterflies of the World 

Atrophaneura (Atrophaneura) luchti Roepke, 1935 RARE 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Atrophaneura luchti is a rare butterfly known only from a few 
specimens from the Ijen (Idjen) mountain range in eastern Java, Indonesia. Its 
biology and distribution are poorly known and more field information is needed as a 
basis for conservation assessment and planning. If the Indonesian Conservation 
Plan's recommendations are adopted, the species will be well protected in a large 
national park. 

Description Atrophaneura luchti is a large, handsome butterfly with a 
forewing length of 60 mm (male) to 65 mm (female). The sexes have similar patterns 
but the male is smaller, with more pointed forewings. Adults have a red stripe along 
the sides of the head, thorax and abdomen (1, 3). 

UFW/LFW elongated with a black ground colour fading to grey (3) or greenish 
grey (1) between the veins. 

UHW/LHW tailless and scalloped with a black-spotted, creamy-white (3) or pale 
yellow (1) submarginal/postdiscal band on the black ground colour. 

Atrophaneura luchti is sometimes treated as a subspecies oi A. priapus, of which 
the subspecies dilutus flies in a separate part of eastern Java. A . luchti is clearly a close 
relative oi A. priapus, and is believed to be an isolated relict of priapus stock (3). 
However, A. luchti is accepted as a good species by all recent authors (1, 2, 3) and a 
number of distinct characteristics are recognizable. The main difference from A. 
priapus is that both sexes have the sides of the abdomen covered by red hairs, rather 
than creamy yeUow hairs (3). 

Distribution This species is known only from the Ijen (Idjen) range of 
mountains in eastern Java, Indonesia. 

Habitat and Ecology Atrophaneura luchti was described less than fifty years 
ago and is still very poorly known. The only specimens come from Gunung (Mt) Ijen 
and apart from the holotype, the species remained uncollected until 1979/1980, when 
more specimens were taken (1, 3). The larval foodplants remain unknown. A. luchti 
is a member of the nox group, some other members of which are known to lay their 
eggs on Aristolochiaceae (3) in the manner of other troidine swallowtails. Adults 
have been collected in November, January and March, and may fly throughout the 
year, taking nectar from flowers in the forest canopy (3). 

The Ijen mountain range is volcanic in origin, with the highest points on Gunung 
Raung (3332 m) and Gunung Merapi (2800 m). This montane region has the lowest 
human population density in Java, most of the people being concentrated on the 
alluvial plains. Large areas of upland remain relatively undeveloped. The mountain 
ranges are mainly forested but with a high plateau, lakes and volcanic craters. The 
altitude at which A. luchti occurs is unknown, but is likely to be at medium or high 
elevations, most lowland habitat having been put to agricultural purposes. 

Threats Threats to this butterfly and its habitat are presently unknown. Its 
very restricted range on an island with one of the world's highest human population 
densities is cause for concern and there is no information on its distribution within 
protected areas. 

254 



Atrophaneura luchti 

Conservation Measures There are six protected areas in the Ijen mountains 
of eastern Java (4). Three of these are very small: Pacur Ijen Reserve (9 ha) is 
contiguous with the Maeleng Reserve (see below); Ceding Reserve (2 ha) is a forest 
in the Ijen crater, probably already destroyed; and Cerakmanis Sempolan Reserve 
(16 ha) is on the slopes of Gunung Raung. The other three protected areas are Kawah 
Ijen Merapi Ungup-ungup Reserve (2560 ha) which surrounds the Kawah Ijen crater 
lake and includes mountains such as Gunung Merapi; Maelang Taman Buru Reserve 
(70 000 ha), which includes extensive undisturbed lowland forest; and the Gunung 
Raung proposed Reserve (about 60 000 ha), which is a marvellous area of montane 
forest well buffered by pine and teak plantations. In the recent Indonesian 
Conservation Plan it has been proposed that the three larger reserves should be 
combined into one protected area for future development into a national park. The 
three smaller reserves would automatically be included (31). It is likely, although 
unconfirmed, that Atrophaneura luchti occurs in at least the Maelang Reserve. 
Adoption of the Indonesian Conservation Plan's recommendation would ensure the 
future of the species and should be implemented at the earliest opportunity. There 
are many reasons why the Ijen range requires protection, not least of which is to 
maintain essential watersheds and thus ensure a constant water supply in the highly 
seasonal climate of eastern Java. 

Atrophaneura luchti is clearly a rare, narrowly distributed, endemic species and 
further information on its biology and distribution is needed in order to ensure 
appropriate management measures. 

References 

1. D'Abrera, B. (1982). Butterflies of the Oriental Region. Pari 1 . Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi + 244 pp. 

2. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

3. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands. Vol. 1 
Papilionidae. (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 

4. UNDP/FAO National Parks Development Project (1982). National Conservation Plan for 
Indonesia. Vol 3. Java and Bali. FAO, Bogor. 



255 



Threatened Swallowtail Butterflies of the World 

Atrophaneura (Losaria) palu (Martin, 1912) INSUFFICIENTLY KNOWN 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Atrophaneura palu is a large and attractive black and white 
swallowtail known from very few specimens. It only lives in the area around Palu in 
Central Sulawesi , not far from the present site of the Lore Lindu Reserve . Field work 
is needed to assess the distribution, biology and conservation status of this apparently 
very rare species. 

Description Atrophaneura palu is a large butterfly with a fore wing length of 
up to 70 mm (2). As is usual in this genus the forewings of the female are a little 
broader and less angular than in the male, but otherwise the sexes ae similar in 
general appearance (2). Atrophaneura palu is a member of the coon species-group. 
Tsukada and Nishiyama consider palu to be a subspecies of coon (describing it as 
Losaria coon palu) (8) but Haugum et al. (4) follow Martin (5) in regarding it as 
specifically distinct. This judgement has been supported by Hancock, who recently 
examined the male genitalia (2). 

UFW/LFW elongated with the black ground colour forming a border around the 
wing, but becoming transparent light grey over the inner part. The veins remain 
heavily scaled in black and other black lines extend from the outer margin and apex 
between the veins and across the cell (4, 5,8). 

UHW/LHW elongated with a long tail (broad and rounded at the tip, narrow at the 
base), and a scalloped outer margin . The black ground colour has a postdiscal band of 
seven white wedge-shaped spots. Only the larger, inner four of these are visible on 
the upper surface (4, 5,8). 

Distribution Atrophaneura palu is an isolated species only known from 
Sulawesi (Indonesia). Other members of the coon group occur much further west in 
Sumatra, western Java, other parts of western Indonesia and Peninsular Malaysia, 
but not on neighbouring Borneo. The type specimen was captured at an altitude of 
2000 m at the village of Lewara, east of Palu on the west coast of Sulawesi. Further 
specimens, including the first male to be described, were taken at sea level west of 
Palu (2). 

Habitat and Ecology No published information is available on the habitat, 
breeding biology or foodplants oi Atrophaneura palu. The very wide altitudinal range 
of the butterfly is unusual; the type locality at 2000 m is presumably in a montane 
ecosystem quite different to the coastal habitat of the more recent discoveries. The 
foodplants of other members of the coon group are believed to be Aristolochiaceae 
(8). Palu, the provincial capital of Central Sulawesi, lies in a rain shadow with a 
rainfall of only about 700 mm per year, but the surrounding highlands have a normal 
rainy season from November to March, with a rainfall in excess of 3000 mm. The 
highlands are mainly forested, particularly on the slopes, but some valley bottoms are 
open grassland (1). According to unpubHshed information a breeding locahty has 
recently been found by Japanese collectors in the region west of Palu. Unconfirmed 
reports suggest that the butterfly is confined to an arid and rather barren habitat (3). 
This would be most unusual for a troidine clubtail and might suggest a foodplant 
outside the Aristolochiaceae. 

Sulawesi is one of the most distinctive and biologically interesting islands in 

256 



r 



Atrophaneura palu 

Indonesia. It has a fascinating swallowtail fauna recently reviewed by Haugum, 
Ebner and Racheli (4). Three species oi Atrophaneura are known from the island; 
none are common but /I. dixoni is fairly widespread and abundant in some localities 
(8). Atrophaneura kuehni lives on the north and east coast and is only slightly better 
known than paht. The northern subspecies, mesolamprus, is known only from the 
type specimens, and the eastern, nominate, form is known from very few collections 
(8). Other very rare species include PapiUo jordani (see separate review) and 
Graphuim dorcus, a rare but widespread species probably confined to high altitudes 
(8). 

Threats The precise habitat oi Atrophaneura pahi has yet to be established, 
making it difficult to assess the status of the species. Like Graphium dorcus , A . palu 
may be mainly distributed in the poorly known and thinly populated central 
mountainous areas of Sulawesi. Although the locality of the species is remote, the 
degree of environmental disruption, particularly deforestation, should not be 
underestimated. Some of the presently restricted ranges of Sulawesi endemics may at 
one time have been much more widespread. Large areas stretching between 
Makassar and the Northern and Central Districts are covered with the unpalatable 
and useless grass Imperata cylindrica (lalang), a sure sign of over-exploitation and 
impoverishment of the soil (7) . Natural habitats are increasingly being destroyed as a 
result of extensive, continuing and largely uncontrolled deforestation (7). Following 
the discovery of a new locality by Japanese coWeclors, Atrophaneura pahi has recently 
appeared in the European trade at grossly inflated prices. This is a possible 
confirmation of the rarity and vulnerability of the species and is of particular concern 
when no details of habitat and ecology have been published. 

Conservation Measures A UNDP/FAO development project has drawn up 
a conservation plan for Indonesia and made substantial recommendations for the 
conservation of the Sulawesi fauna (6). Lore Lindu Reserve, in Central Sulawesi, lies 
in the headwaters of the Palu River and is probably within 100 km of the type locality 
of Atrophaneura pahi. Further study within the reserve and in other areas around 
Palu is needed to verify the presence of the butterfly and to ascertain its breeding 
biology, foodplant and conservation status. 

References 

1. Anon, (undated). Indonesia, National Parks and Nature Reserves. Directorate Geneial of 
Tourism, Jakarta. 150 pp. 

2. Hancock, D.L. (1984). A note on Atrophaneura palu (Martin) 1912. Papilio International 
1(3): 71-72. 

3. Haugum, J. (1984). In litt., 14 April. 

4. Haugum, J., Ebner, J. and Racheli, T. (1980). The Papilionidae of Celebes (Sulawesi). 
Lepidoptera Group of 1968 Supplement 9: 21 pp, 1 map, 2 pi. 

5. Martin, Dr. (1912). Ein neuer Papilio aus Celebes. Deutsche Entomologische Zeitschrift 
"Iris" 26: 163-5. Plate given separately (1913) Deutsche Entomologische Zeitschrift "Iris" 27: 
Tafel 6. 

6. National Parks Development Project, UNDP/FAO (1982). National Conservation Plan for 
Indonesia Vol. VI: Sulawesi. FAO, Bogor. 

7. Straatman, R. (1968). On the biology of some species of Papilionidae from the island of 
Celebes (East-Indonesia). Entomologische Berichten 28: 229-233. 

8. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands Vol. 1 
Papilionidae. (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 

257 



Threatened Swallowtail Butterflies of the World 

Atrophaneura (Pachliopta) jophon Gray, 1852 VULNERABLE 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Atrophaneura jophon is a rare species confined to the rain forests 
of south-western Sri Lanka. Over the past 30 years the human population has 
doubled and these forests have been devastated by the combined impact of timber 
extraction and agriculture. The 11 000 ha Sinharaja Forest Reserve is probably one 
of the last strongholds of the butterfly, but selective logging and illicit encroachment 
have already reduced the virgin forest area to 5000 ha. Although the entire forest has 
now been declared a UNESCO Man and Biosphere Reserve, exploitation of forest 
products continues to threaten the butterfly's habitat. 

Description Atrophaneura jophon, the Ceylon Rose, is a large butterfly 
with a long tail to the hindwing, a forewing length of 60-64 mm, and similar sexes 
(Plate 3.1). 

Forewing black with creamy-white markings in the discal and postdiscal regions, 
less well defined on the UFW which has heavier suffusion of black scales (2, 3, 10). 

Hindwing deeply scalloped with a long spatulate tail. Black ground colour with 
creamy-white discal spots and pink submarginal spots, reduced to narrow lunules by 
heavy black suffusion on the UHW (2, 4, 10). 

Distribution Atrophaneura jophon is endemic to Sri Lanka. It was formerly 
considered to be conspecific with Atrophaneura pandiyana from southern India (10), 
but has been recognized by Munroe (8), and Hancock (6) as a separate species. 

Habitat and Ecology This butterfly flies at medium elevations 
(615-1230 m) in the wet, south-western zone of Sri Lanka (2, 10). Here the rainfall is 
over 2000 mm per year with no month completely dry and two main rainy seasons, 
from March to May and June to September (5). The natural vegetation is tropical 
evergreen rain forest, which is restricted to these south-western regions. The drier 
and more seasonal climate of the eastern and northern regions supports tropical 
savanna vegetation (5). Below 900 m the rain forest has a closed, dense canopy at 
22-27 m with emergents to 45 m. Typical trees include species of the genera 
Dipterocarpus, Shorea, Mesua, Doona, Hopea, Palaquium, Pygeum and others (5). 
From 900 to 1500 m montane semi-evergreen forests grow to 18-24 m, with 
emergents to 30 m (5). The boundaries beween these two forests and the montane 
wet evergreen forests above 1500 m are indistinct. 

The butterflies tend to fly low in the early morning and in the forest canopy later in 
the day (10). The eggs are yellow-brown, believed to be laid on the undersides of 
young leaves of Aristolochiaceae (10), as is the case in Atrophaneura pandiyana 
(2, 10). The caterpillars are purple-black with crimson tubercles and cream bands 
(10). Although usually rare and local, where it does occur it may be fairly numerous 
(10). 

Threats Since the last comprehensive inventory of Sri Lankan forest 
resources in 1959-60, widespread deforestation has occurred. Of 2.9 million ha of 
closed broadleaved forest in 1956, only about 1.6 million ha now remains, excluding 
forest fallows (5). Of this total, only about 0.43 million ha is not exploited, over half 
of this for reasons of physical inaccessibility, the rest for legal reasons (5). 

Deforestation in the country as a whole constitutes a very serious problem. The 

258 



Atrophaneura jophon 

main cause is the pressure for agricultural land and forest products caused by high 
population levels and population growth rates, factors unlikely to be alleviated in the 
near future. The human population has doubled since the 1950s and now stands at 
almost 15 million. 

The main threat to this narrowly endemic and local butterfly is the destruction of its 
habitat. Members of the subgenus all fly in dense forests and there is no evidence to 
suggest that they can survive serious disturbance. 

Conservation Measures Numerous protected areas have been set aside in 
Sri Lanka and during the first half of this century the country boasted one of the best 
conservation programmes in Asia (7). However, the great majority of the reserves 
were in the dry northern and eastern regions (1), where this butterfly is not found. 
According to D'Abrera (2), the forests between Ratnapura and Deniyaya alluded to 
by Woodhouse and Henry ( 13 ) and Woodhouse ( 12) as the habitat of jophon, are now 
almost completely cleared for timber. 

The last extensive area of primary lowland rain forest is the Sinharaja Forest 
Reserve, probably one of the last strongholds of Atrophaneura jophon . In the past the 
butterfly is known to have been well established here, but there is now evidence that 
the constant human disruption of the reserve is causing depletion of the butterfly 
populations (2). 

Although the Sinharaja forest is 11 000 ha in extent, only about 5000 ha remains 
undisturbed, the rest having been seriously disrupted by a selective logging operation 
in 1972-75, illicit agricultural encroachment in many places along the reserve 
boundaries, and constant pressure for minor forest products (3). In April 1978 about 
1822 ha of forest was declared a UNESCO Man and Biosphere Reserve and the rest 
of the area has since been included under this designation (9). A number of scientific 
studies have been carried out there. The reserve is the responsibility of the Forest 
Department of Sri Lanka, which has taken steps to ensure its protection, but policing 
the area is a continuing problem. 

A survey of the butterfly fauna, with emphasis on Atrophaneura jophon , would add 
to the already evident value of the reserve. Sri Lanka has 15 species of swallowtails 
including another endemic, Troides darsius, two other species of Atrophaneura, A. 
hector and A. aristolochiae and two 'PapiUo species restricted to Sri Lanka and 
southern India, P. polymnestor and P. crino. Formany of these species the Sinharaja 
Forest Reserve will become an increasingly important habitat resource. 

Sri Lanka's President has recently appointed a task force to examine the status of 
the island's natural resources in relation to their conservation and utilization for the 
future (11). Recommendations will be made for the development and implementa- 
tion of a National Conservation Strategy (11). Unless the decline in Sri Lanka's 
forests is arrested quickly, Atrophaneura jophon and many other species will soon 
become very seriously endangered. 

References 

1. Crusz, H. (1973). Nature conservation in Sri Lanka (Ceylon). Biological Conservation 5: 
199-208. 

2. D'Abrera, B. ( 1982). Buiterfties of the Oriental Region. Part 1. Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi + 244 pp. 

3. Davis, S. (in prep.). Sinharaja Forest. Draft review for Plant Sites Red Data Book. 

4. Evans, W.H. (1932). The Identification of Indian Butterflies. Bombay Natural History 
Society. 2nd ed. 454 pp. 

259 



Threatened Swallowtail Butterflies of the World 

5. FAO/UNEP (1981). Tropical Forest Resources Assessment Project. Forest Resources of- 
Tropical Asia. FAO, Rome. 475 pp. 

6. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

7. Hoffmann, T.W. (1983). Wildlife conservation in Sri Lanka. Paper presented at Bombay 
Natural History Centenary Seminar, Bombay (in press). 

8. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supp\ement 17: 1-51. 

9. Sri Bharathi, K.P. (1979). Man and Biosphere reserves in Sri Lanka. Sri Lanka Forester 
14(1-2): 37-38. 

10. Talbot, G. (1939). The fauna of British hidia, including Ceylon and Burma. Butterflies vol I. 
Taylor and Francis Ltd., London, reprint New Delhi 1975. 600 pp. 

1 1 . Wijayadasa, K.H.J. ( 1983). Formulating a Sri Lankan strategy. I UCN Bulletin Supplement 
4: 3. 

12. Woodhouse, L.G.O. (1950). The butterfly fauna of Ceylon. Second Edition. Colombo 
Apothecaries' Co. Ltd., Colombo. 231 + xxxii pp. 

13. Woodhouse, L.G.O. and Henry, G.M.R. (1942). The Butterfly Fauna of Ceylon. 
Colombo. 172 pp + 49 pis. 



260 



Atrophaneura schadenbergi 
Atrophaneura (Pachliopta) schadenbergi (Semper, 1886) VULNERABLE 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Atrophaneura schadenbergi is restricted to northern and central 
Luzon and the Babuyan Islands in the Philippines, where it occurs in grassland and 
wooded areas in foothills. It is threatened by loss of habitat caused by the demands of 
the rapidly expanding human population. 

Description Atrophaneura schadenbergi is a relatively large butterfly 
(forewing length 50-60 mm) with narrow elongated wings and long spatulate tails 
(1, 3) (Plate 3.2). The sexes are similar in appearance, the female sHghtly larger than 
the male (3). 

UFW/LFW white, heavily suffused with black scaling which covers the basal 
region, wing borders and veins, and occurs as stripes between all but the veins in the 
tornal region (1,3). The white tornal flash is conspicuous in flight (3). 

UHW/LHW scalloped and tailed, black with seven large submarginal spots, the 
lower three red and the remainder white. All these spots are white in the subspecies 
micholitzi (1, 3). 

Distribution Atrophaneura schadenbergi is endemic to the northern and 
central parts of Luzon and the Babuyan Islands in the Philippines. The nominate 
subspecies occurs in the Cordillera Central and Sierra Madre of northern Luzon 
(2, 3), reaching as far south as the Bataan Peninsula and the Wawa Dam, north-east 
of Manila (3). The second subspecies, /I. s. micholitzi is poorly known, but occurs at 
least in Camiguin , the Babuyan Islands, and possibly the northern tip of Luzon (1,3). 

Habitat and Ecology Atrophaneura schadenbergi is essentially an inhabi- 
tant of lowland foothills, although it is found at altitudes up to about 1200 m in the 
Sierra Madre of eastern Luzon , and on the slopes of the Cordillera Central in the west 
(3). It seems to have a variable habitat. At the Asin Hotspring near Baguio it flies in 
open wooded grassland, together with Atrophaneura kotzebuea (3). In the eastern 
and western foothills of the Sierra Madre (2) in eastern Luzon, and also in the 
limestone hills of Luzon's northern tip (3) it flies in heavily wooded secondary forests, 
while A. kotzebuea remains m the grassland (3). A. schadenbergi flies at heights of 
1-4 m, particulary during the morning and late afternoon (2). The main flight season 
in the Sierra Madre seems to be February to August, with a peak in April (2). Other 
records, mainly from the Cordillera Central, indicate adults on the wing rather later 
in the year, from May to November (3). Whether it is continually breeding or 
producing one or two broods per year remains uncertain (3) . The immature stages are 
unknown (3) and the foodplant, although unknown, is Hkely to be in the Aristolo- 
chiaceae. 

Threats Atrophaneura schadenbergi has a limited distribution within which 
it occurs only locally (3). According to C.G. Treadaway, an authority on Philippine 
butterflies, it is in danger of disappearing as a result of the persistent degeneration 
and elimination of its habitat for agricultural purposes (2). A graphic description of 
the loss of suitable woodland is given by Tsukada and Nishiyama (3), who describe 
the difficulties encountered by S. Yamaguchi in finding forests in northern Luzon, 
where "the bald mountains stretched around him as far as the eye could see" (3). 

261 



Threatened Swallowtail Butterflies of the World 

Fortunately the foodplant of the butterfly can evidently survive in secondary forest, 
otherwise the plight of this butterfly might be even more serious. 

Conservation Measures No measures have been taken to protect Atro- 
phaneura schadenbergi and there are no known functional protected areas in 
northern Luzon. As a denizen of the foothills of northern Luzon the butterflies are 
probably surviving by migrating between forest patches, colonizing secondary 
growth as it matures. As human population pressures inevitably increase still further, 
the butterfly will become even more local and disjunct, finally surviving in isolated 
and highly vulnerable relict forests. The only way to ensure the continuing survival of 
this attractive species is the designation and effective management of suitable 
protected areas. 

References 

1. D'Abrera, B. (1982). Butterflies of the Oriental Region. Part 1. Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi + 244 pp. 

2. Treadaway, C.G. (1984). In litt., 25 May. 

3. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands. Vol. I 
Papilionidae. (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 



262 



Atrophaneura atropos 
Atrophaneura (Pachliopta) atropos Staudinger, 1888 INDETERMINATE 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Atrophaneura atropos is a handsome black-winged swallowtail 
confined to the western Phihppine island of Palawan. Its status is uncertain, but it may 
be declining in the face of rapidly accelerating deforestation on the island. The 
national parks system of the Philippines is currently undergoing redevelopment. The 
Province of Palawan should protect large areas of its remaining wilderness as a resort 
for its unique flora and fauna, and also as resources for the growing tourist industry. 

Description Atrophaneura atropos has entirely black and grey wings and 
red markings on the abdomen. The elongated forewing, which has a length of 
45 - 55 mm, is grey with black margins, veins, parallel lines in the spaces between the 
veins, and lines in the cell. The hindwing is entirely black, and has a scalloped outer 
margin with a long spatulate tail. The sexes are alike, though the female is slightly 
larger. The lower surface is the same as the upper (3, 12). 

This species is a member of the poly dorus group of the genus Atrophaneura (6, 9), 
all of which have similar dark caterpillars with rows of dorsal and lateral tubercles, 
often red or red-tipped (6, 7). Some tubercles may be white and there is a white band 
on the abdomen (6, 7). 

Distribution Atrophaneura atropos is endemic to the Philippine island of 
Palawan where it is found in the southern, central and northern lowlands (12). 
Graphium megaera is also restricted to Palawan and is reviewed separately in this 
volume. 

Habitat and Ecology Palawan lies to the southwest of the main archipelago 
of the Philippines and just to the north of Sabah, East Malaysia. The island is 
dominated by a central mountain chain, with heavier and less seasonal rainfall on the 
eastern side. The lowland coastal plains in the east provide much of the island's arable 
land while in the west the mountains slope more directly to the coast. According to 
1972 figures almost 70 per cent of the land area of the province was forested (1), but 
more recent studies indicate that only 53 per cent of Palawan is still under forest ( 10) , 
a considerable reduction characteristic of all the Philippine islands (3). In the 
lowlands the forests are dominated by Dipterocarpaceae (notably Dipterocarpus, 
Shorea, Parashorea and Pentacme), which are highly prized timber trees (2, 3). In 
lower montane regions the dipterocarps gradually disappear to be replaced by oaks 
(Quercus) and a variety of other trees {Tristania, Hopea, Eugenia, Agathis, etc.) (3). 
On mountain tops upper montane mossy forests are found (3). 

Palawan is on the eastern edge of the Asian continental shelf. Its fauna and flora 
bear many resemblances to that of nearby Sabah, with which Palawan was contiguous 
during the lowered sea levels of the last Ice Age. This is demonstrated by the 
distribution of Atrophaneura neptunus, which occurs from Peninsular Malaysia 
across Sumatra and Borneo to Palawan, where dacasini, a distinctive subspecies, flies 
(12). 

Atrophaneura atropos flies in primary lowland rain forest throughout Palawan 
(10, 12). This species prefers humid forests and does not fly in the same forests as 
Atrophaneura neptunus, which prefers drier habitats (12). The foodplants of the two 
species are unknown, but likely to be in the Aristolochiaceae (5, 7, 9). Atrophaneura 

263 



Threatened Swallowtail Butterflies of the World 

atropos flies slowly through the forest, quite often close to the ground and in open 
areas, or along the forest edge. They can be seen even on cloudy days or in light rain, 
sometimes stopping to take nectar from flowers (9). Adults are often counted as a 
rarity, but are in fact fairly widespread, can be common and fly all year round (9). 
There is evidence that the species survives well in secondary growth (4). 

Threats By Asian standards Palawan is sparsely populated, with only 
370 000 people (in 1979) spread over nearly 1.5 million ha (1,2). Most of these 
people live in the eastern lowlands. In the less accessible central highlands, southern 
and western regions there are still large areas of forest and wilderness (1). 
Atrophaneura atropos probably still flies throughout these rain forest regions and is 
unlikely to be critically threatened at the moment. Nevertheless, the pace of 
deforestation is accelerating, particularly in the south (2) , with 44 per cent of the land 
area under logging concessions in 1976, the area currently leased must be well over 
half of the total. The main practice is selective felling but with 'kaingin' farmers, the 
shifting cultivators, using logging roads for access to fresh areas, there is often no 
opportunity for the forest to recover. Mining of sand and minerals such as mercury 
and chrome have a local effect on forests, particularly when the operations close 
down and the employees remain and set up new villages and farms (2, 8). Although 
A. atropos and its foodplant apparently survive in secondary growth, there is 
evidence that clearance for upland rice after decreasing periods of fallow is having an 
adverse affect on populations (4). 

Conservation Measures As has been stated in a number of other reviews of 
Philippine endemics, the national park system in the islands is undergoing a time of 
change. At one time as many as 63 parks were designated by the Philippine 
Government, 51 under the Bureau of Forest Development, but these are currently 
being restructured in favour of a reduced number of national parks to be set up by the 
Bureau and maintained to international standards (11). It is not clear whether any 
parks or wilderness areas will be sited in Palawan forests, although the St Paul's 
Underground River National Park will presumably remain. The regions around 
Cleopatra's Needle and Mt Mantalingajan have been suggested as wildlife sanctua- 
ries with access only for scientific purposes (11). This is an important area for A. 
atropos and many other forms of wildlife (8). There is great potential for new and 
important parks and wilderness areas in Palawan, not only to protect the unique flora 
and fauna but also as a resource for the growing tourist industry. It is also essential to 
the well-being of the Philippine economy and of the Palawan environment that 
exploitation of non-protected forests should be done in a controlled and sustainable 
manner. In this way much of the wildlife will be retained. 



References 

1. Bruce, M. (1980). The Palawan Expedition Stage 1. Traditional Explorations, Sydney. 
47 pp. 

2. Bruce, M. (1981). The Palawan Expedition Stage 2. Associated Research, Exploration and 
Aid, Sydney. 139 pp. 

3. D'Abrera.B. (1982). Butterflies of the Oriental Region. Parti. Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi + 244 pp. 

4. Dacasin, G.A. (1984). In litl., 25 April. 

5. FAO/UNEP (1981). Tropical Forest Resources Assessment Project. Forest Resources of 
Tropical Asia. FAO, Rome. 475 pp. 

264 



Atrophaneura atropos 

6. Hancock, D.L. (1980). The status of the genera Atrophaneura Reakirt and Pachliopta 
Reakirt (Lepidoptera: Papilionidae). Australian Entomologists' Magazine 7: 27-32. 

7. Igarashi, S. ( 1979). Papilionidae and Their Early Stages. Vol 1 : 219 pp. , Vol 2: 102 pp. of 
plates. Kodansho, Tokyo. (In Japanese). 

8. Jumalon, J.N. (1984). Inlitt., 10 July. 

9. Monroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
fw/owo/of (J/ Supplement 17: 1-51. 

10. National Environment Protection Council (1979). Philippine Environment 1979. NEPC 
Third Annual Report. NEPCMinistry of Human Settlements, Republic of the Philippines. 
158 pp. 

11. Pollisco, F.S. (1982). An analysis of the national park system in the Philippines. 
Likas-Yaman {Journal of the National Resources Management Forum) 3(12): 11-56. 

12. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands. Vol. 1 
Papilionidae. (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 



265 



Threatened Swallowtail Butterflies of the World 

Troides {Troides) andromache (Staudinger, 1892) INDETERMINATE 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Troides andromache is a birdwing which occurs at high eleva- 
tions in a restricted part of Sabah and Sarawak in northern Borneo. Little is known of 
its ecology or precise range, but there is no doubt that threats to its habitat are 
multiplying. It occurs on Mt Kinabalu, an important national park for a variety of 
wildlife, but economic pressures are changing the park adversely. More information 
on T. andromache is required to assess its conservation status more precisely, and the 
planning of land-use in the area needs to be more effective. 

Description Troides andromache is a large butterfly with a forewing length 
of about 65 mm (male) or 85 mm (female). The male is black, yellow, and grey in 
colour, while the female has additional brown and white scaling on the forewing 
(2, 3, 8, 9). Although two geographically separated subspecies, the nominate and T. 
a. marapokensis Fruhstorfer, have been distinguished by some authors, the latter is 
now regarded as a female form of the former (6). 

Male: UFW entirely black with a violet sheen; LFW with a band of large greyish 
distal spots dusted with yellow. 

Hindwing almost entirely yellow with black scaling narrowly over the veins, as a 
broad inner margin, and as large fringe spots producing an edentate submarginal 
band (2, 3, 8, 9). 

Female: Forewing greyish-white lightly dusted with brown scales; brown apical 
area, outer margin and veins (2, 3). 

Hindwing differs from male in the band of large, black discal/postdiscal spots. 
These join with the large fringe spots to leave very little of the yellow colour visible on 
the distal part of the hindwing (2, 3, 8, 9). 

Distribution Troides andromache occurs in East Malaysia where it is found 
in Sabah and Sarawak in northern Borneo. The nominate subspecies is well known as 
occurring on Mt Kinabalu, particularly on the Pinosuk Plateau (1,5, 6), but it has 
also been recorded from neighbouring mountains in the Crocker range. It has been 
suggested that it may also occur further south in Kalimantan, the Indonesian sector of 
Borneo, where there is plenty of suitable habitat but where little collecting has been 
done. T. a. marapo/:e«5;5 occurs further south in the area of Mt Marapok in Sarawak, 
a different mountain from the Mt Marapok south of Mt Kinabalu in Sabah, where the 
subspecies has not been found, despite incorrect statements to the contrary (2, 3, 8). 

Habitat and Ecology Troides andromache occurs at high altitudes, from 
about 1000 to 2000 m above sea level. It has been described as an alpine butterfly (9) 
but this is misleading since the mountains are densely forested at such altitudes. It 
flies throughout the year, particularly in the morning and evening. Adults nectar on 
Mussaenda flowers and females fly lower than males (9). 

Virtually nothing is known about the ecology of the early stages. The foodplant is 
an unidentified species ot Aristolochia vine, on which oviposition has been observed 
(9). 

Threats Destruction of the habitat of Troides andromache has occurred on 
a considerable scale (1, 4, 6). Parts of the Pinosuk Peninsula have been 'developed' 
for tourism, with construction of access roads, a golf course and a visitor centre, and 

266 



\ 



Troides andromache 

with other facihties planned. Other areas have become a tea estate and a cattle ranch , 
or have been altered by timber processing and hydroelectric projects. 

A large area of eastern Kinabalu, designated as a national park, has been leased 
(from 1973) for 30 years to a copper mining company. The Manu copper mine has 
changed the eastern face of Mt Kinabalu (4). For more details see the reviews of 
Graphium (Graphium) procles and Papilio (Princeps) acheron. 

T. andromache is not a well-known species in trade and is only occasionally offered 
by dealers, usually as old rather than recently-collected specimens (2). 

The province of East Kalimantan , where the species may possibly occur, has a low 
human population that is growing rapidly. Shifting agriculture and commercial 
logging are increasingly extensive in this region. 

Conservation Measures Little is known about the ecology of Troides 
andromache and research and surveys are needed as a basis for practical conservation 
measures. 

Although Mt Kinabalu was designated a national park in 1964 (7), economic 
developments in the short-term may reduce its value as an important wildlife area. 
Besides being the highest mountain in South East Asia, it has an exceptionally rich 
fauna and flora. T. andromache flies there with two other Rare swallowtails, 
Graphium procles and PapiUo acheron (see separate reviews). 

The undoubted reductions in the habitat of T. andromache do not give cause for 
immediate alarm (1), but if economic and tourist pressure increase, the butterfly may 
become more seriously threatened. It is clear that effective and sensitive planning and 
practical conservation are required to ensure the adequate survival of the fauna of 
which T. andromache is a part. 

Some mention may be made of conservation in East Kalimantan, even though 
there are as yet no records of T. andromache from this province. East Kalimantan is 
the least densely populated of the five Kalimantan provinces and is of especially high 
conservation importance because of its richness and diversity (10). The very large 
Kayan-Menterang reserve of 1.6 million ha, in particular, should be thoroughly 
surveyed for T. andromache . 

The species should be retained, until its status can be reviewed, under Appendix II 
of the 1973 Convention on International Trade in Endangered Species of Wild Fauna 
and Flora (CITES). Appendix II listing implies that commercial trade is allowed 
providing a permit from the country of export is obtained, this can provide a method 
of monitoring trade levels. 

References 

1. Barlow, H.S. (1983). In lilt.. 29 June. 

2. D'Abrera, B. (1975). Birdwing Butterflies of the World. Lansdowne Press, Melbourne. 
260 pp. 

3. D'Abrera, B. (1982). Butterflies of the Oriental Region. Part 1. Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi + 244 pp. 

4. Davis, S. (in prep.) Mount Kinabalu. Draft review for lUCN Plant Sites Red Data Book. 

5. Holloway, J.D. (1978). Butterflies and Moths. In Kinabalu Summit of Borneo. Sabah 
Society Monograph, 25-278. 

6. Holloway, J.D. (1983). In litt., 28 February. 

7. Luping, D.M., Wen, C. and Dingley, E.R. (eds) (1978). Kinabalu, Summit of Borneo. 
Sabah Society Monograph, Malaysia. 

8. Ohya, T. (1983). Birdwing Butterflies. Kodansha, Tokyo. 332 pp, 136 col. pis. 

9. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands. Vol. 1 

267 



Threatened Swallowtail Butterflies of the World 

Papilionidae. (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 
10. UNDP/FAO National Parks Development Project ( 1981). National Conservation Plan for 
Indonesia. Vol. 5: Kalimantan. FAO, Bogor. 



268 



Troides prattorum 
Troides (Troides) prattorum (Joicey and Talbot, 1922) INDETERMINATE 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Troides prattorum is a large black and yellow birdwing butterfly 
with particular well-developed structural coloration producing a blue-green sheen on 
the angled hindwing. The species occurs only at high elevation in the Indonesian 
island of Buru, in the Moluccas. Its ecology is unknown. It occurs with the endemic 
Delias apatela and probably with two other endemic Delias. Research and survey are 
the first priority for conservation but a reserve for the unique butterfly fauna of Buru 
should also be established. 

Description Troides prattorum, the Buru Opalescent Birdwing is a large, 
black butterfly with an unusual opalescent blue-green cast to the golden-yellow part 
of the hindwing. The female is larger with a forewing length of about 100 mm 
compared to about 85 mm in the male (1,2). 

Male: UFW black with narrow white borders to the veins on the upper part of the 
wing. 

UHW slightly scalloped, almost entirely golden-yellow with black scaling nar- 
rowly over the veins, as a broad inner margin, and as large fringe spots producing an 
edentate submarginal band. The golden-yellow patch shows the dramatic charac- 
teristic of this species, the form of opalescent scaling. When observed fron various 
angles the entire patch changes to many different tones of opalescent blue-green. 

LFW/LHW. The white scaling on the forewing is less extensive on the lower 
surface, which also has some black suffusion to the lower part of the hindwing 
golden-yellow patch (1, 2). 

Female: UFW with more extensive white scaling in the female , especially in the top 
of the cell. 

Hindwing with golden-yellow patch reduced by a broad discal band of black 
streaks to an irregular postdiscal line and a small patch around the cell apex. There is 
also more suffusion of dark scales distally but the opalescence is still present (1,2). 

Distribution Troides prattorum is known only from the Indonesian island of 
Buru in the Moluccas, which lies due east of Sulawesi. Its occurrence here is 
something of a zoogeographical mystery (1,2). 

Habitat and Ecology The habitat of Troides prattorum is the high plateau of 
the centre and west of the island. The type series of four specimens (one male, three 
female) was taken from 600 to 1600 m above sea level (3). 
Nothing is known about the foodplants, early stages or ecology. 

Threats Troides prattorum may be at risk because of its restricted area of 
occurrence, although as a high-altitude species it is perhaps less at risk than similar 
species at low elevations, which are frequently threatened by forestry and agricultu- 
ral intensification. It is known, for example, that logging operations on steep 
limestone hills to the south of Bara are having a serious environmental impact. 

Buru was apparently used from 1965 onwards as a prison island and access was 
particularly difficult (2). Conditions have eased recently and certain commercial 
collectors have been active on the island. Pairs of T. prattorum were very highly 

269 



Threatened Swallowtail Butterflies of the World 

priced in 1980 and were commercially much more valuable than almost any other 
species of Troides (7). The effects of commercial exploitation are not known. 

Conservation Measures Research on the ecology of T. prattorum is the first 
priority , together with an assessment of its precise distribution , current status and the 
effects of commercial collecting. 

An effective reserve for this species, and others, should be established. The 
uncommon Delias apatela Joicey and Talbot (Pieridae) is endemic to Buru and flies in 
the same plateau biotope as T. prattorum (1,5). D. rothschildi rothschildi Holland is 
more frequent (and occurs in Timor as a second subspecies) but is also an upland 
species. The only known specimen of D. dumasi Rothschild is also from the same or a 
similar biotope (1, 4). 

A potential reserve of 145 000 ha has been proposed for Gunung Kelaput Muda in 
north-west Buru, and given a high priority in the National Conservation Plan for 
Indonesia (6). It is not yet known whether Buru's endemic butterflies are to be found 
in this region. 

The species should be retained, until its status can be reviewed, under Appendix II 
of the 1973 Convention on International Trade in Endangered Species of Wild Fauna 
and Flora. Appendix II listing implies that commercial trade is allowed providing a 
permit from the country of export is obtained, this can provide a method of 
monitoring trade levels. 

References 

1. D'Abrera, B. (1971). Butterflies of the Australian Region. Lansdowne Press, Melbourne. 
415 pp. 

2. D'Abrera, B. (1975). Birdwing Butterflies of tlie World. Lansdowne Press, Melbourne. 
260 pp. 

3. Howarth, T.G. (1977). A list of the type-specimens of Ornitlwptera (Lepidoptera: 
Papilionidae) in the British Museum (Natural History). Bulletin of llie British Museum of 
Natural History (Entomology) 36: 153-169. 

4. Talbot, G. (1929). A Monograph of the Pierine Genus Delias. Vol. 3. Bale & Co. Ltd., 
London. Pp. 117-172. 

5. Talbot, G. (1937). A Monograph of the Pierine Genus Delias. Vol. 6. British Museum 
(Natural History), London. Pp. v -I- 261-656. 

6. UNDP/FAO National Parks Development Project (1981). National Conservation Plan for 
Indonesia. Vol. 7: Maluku and Irian Jaya. FAO, Bogor. 

7. Various trade catalogues (1980-1984). 



270 



Troides dohertyi 
Troides (Troides) dohertyi (Rippon, 1893) INDETERMINATE 

Subfamily PAPILIONINAE Tribe TROIDINI 

Summary Troides dohertyi is an unusually dark birdwing confined to the 
Talaud and Sangihe Islands north of Sulawesi, in Indonesia. The ecology of the 
species is very poorly known and research is needed. Human population pressure is 
very high on the islands and although the species may be protected in the Karakelang 
Hunting Reserve on the largest of the Talaud Islands, the proposed Gunung 
Sahendaruman Game Reserve on Great Sangihe Island should also be surveyed and 
gazetted. 

Description Troides dohertyi, the Talaud Black Birdwing, is a large black or 
black-brown butterfly with a forewing length of about 73 mm (male) or 82 mm 
(female). The male has golden-yellow markings, but these may be absent in the 
female (1,2, 5, 7) (Plates 3.3 and 3.4). 

Male: Almost entirely black, rarely with coloration on the upper surface. 
Whitish-grey scaling along, but not over, the veins of the upper part of the LFW. The 
LHW has a golden-yellow, central discal band. There is usually suffusion of black 
scales from the base, which may obliterate the two costal spots of the discal band. 

Female: Forewing black-brown ground colour with a variable amount of grey 
scaling. Grey scaling less diffuse on the LHW where it occurs around the veins. 

Hindwing with a slightly scalloped outer edge and small yellow marginal spots. The 
golden-yellow central discal patch on the UHW varies from being large and well 
defined to entirely blackened. A similar whitish-grey patch on the LHW has a 
variable amount of dark suffusion. 

Distribution Troides dohertyi is only known from northern Indonesia 
where it occurs on the Talaud Islands and Sangihe Island, between Sulawesi 
(Indonesia) and Mindanao (Philippines). 

T. dohertyi has been variously regarded as a good species (1,3,7), even a 
distinctive species (2), or as a subspecies of T. rhadamantus (Lucas), together with 
T.plateni Staudinger (4,5,6,7). T . rhadamantus is widely distributed in the 
Philippines and T. plateni is common, even in secondary forest, in Palawan. 

Habitat and Ecology Troides dohertyi is a lowland species, which has been 
observed flying at sea level and taking nectar from Mitssaenda flowers. It flies all the 
year round and throughout the daylight hours, though more particularly in early 
morning and late evening (7). 

Little has been recorded on the ecology of the early stages. They are probably 
similar to those of T. rhadamantus , which have been well illustrated in colour (5). 
Both T. plateni and T. rhadamantus larvae are said to be common and easily reared 
or ranched (7). The foodplant is likely to be Aristolochia tagala. 

Threats The precise degree of threat to Troides dohertyi has not been 
documented, hence the category Indeterminate is retained. That some degree of 
threat is present may be evidenced by two factors in particular. Firstly, human 
population pressure on the Sangihe and Talaud Islands is higher than in any other 
part of the province of Northern Sulawesi (8). The population growth rate in the 
province as a whole is 2.2 per cent per year (8). Pressure for land is likely to be most 

271 



Threatened Swallowtail Butterflies of the World 

heavy in the coastal lowlands, where the birdwing is known to fly. There is no 
information on the capacity of T. dohertyi to adapt to secondary vegetation 
formations. 

Secondly, the Sangihe and Talaud Islands are not particularly well served by 
protected areas. Two large hunting reserves, known jointly as the Karakelang 
Reserve, were recently established on the largest of the Talaud Islands (8). The main 
game animals are feral Balinese cattle (8) . If these areas are kept under fairly natural 
vegetation and exotic species are not introduced, the forests could serve as useful 
refuges for the endemic Talaud fauna, including T. dohertyi. However, such 
circumstances are far from being assured. There are no protected areas on the 
Sangihe Islands and the only large piece of remaining forest is in the south of Great 
Sangihe, on Gunung Sahendaruman (8). Troides dohertyi is likely to be at risk 
because of its extremely restricted area of occurrence . Whether it is a good species or 
not, the taxon is one which should be assessed for any threats, particularly land-use 
changes in its island habitats. 

Conservation Measures There is an immediate need for further data on the 
habitat requirements and conservation status of Troides dohertyi. As a preliminary 
measure, it should be sought in the Karakelang Hunting Reserve on the Talaud 
Islands, and in the proposed Gunung Sahendaruman Game Reserve on Great 
Sangihe. 

The Indonesian Conservation Plan proposes to develop strict controls on hunting 
parties in the Karakelang Hunting Reserve or, failing this, to upgrade the site to a 
Game Reserve (8). Such attention to monitoring of the site would be essential to its 
long-term stability. The proposed Sahendaruman Game Reserve, being the last 
remnant of forest on Sangihe, should clearly be gazetted as quickly as possible (8). 
With the growing human populations in the area, encroachment may otherwise be 
inevitable. Troides dohertyi is not listed as a protected species in Indonesia, but this 
may be because the taxon is assumed to be included in T. rhadamantits . The species 
should be retained, until its status can be reviewed, under Appendix II of the 1973 
Convention on International Trade in Endangered Species of Wild Fauna and Flora 
(CITES). Appendix II listing implies that commercial trade is allowed providing a 
permit from the country of export is obtained. This can provide a method of 
monitoring trade levels. 

References 

1. D'Abrera, B. (1975). Birdwing Butterflies of the World. Lansdowne Press, Melbourne. 
260 pp. 

2. D'Abrera, B. (1982). Butterflies of the Oriented Region. Part 1 . Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi -I- 244 pp. 

3. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

4. Haugum, J. and Low, A.M. (1983). A Monograph of the Birdwing Butterflies. Vol. 2 (2): 
105-240, col. pis. 5-12. Scandinavian Science Press, Klampenborg. 

5. Igarashi, S. (1979). Papilionidae and Their Early Stages. Vol. 1: 219 pp., Vol. 2: 102 pp. of 
plates. Kodansha, Tokyo. (In Japanese). 

6. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
En/owo/ogwr Supplement 17: 1-51. 

7. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands. Vol. 1 
Papilionidae. (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 

8. UNDP/FAO National Parks Development Project (1982). National Conservation Plan for 
Indonesia. Vol. 6. Sulawesi. FAO, Bogor. 

272 



Ornithoptera tithonus 
Ornithoptera tithonus De Haan, 1840 INSUFFICIENTLY KNOWN 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Ornithoptera tithonus is a large, attractive birdwing which is 
restricted to western Irian Jaya and a few neighbouring islands. Its ecology and 
conservation needs are very poorly known. It may be best classified Rare because of 
its restricted distribution and possible 'relict' nature. On the other hand, there is 
concern because it has been offered in substantial numbers in commercial trade, the 
effects of which have not been monitored. A primary need is for research on ecology, 
distribution and possible conservation measures, including the effectiveness of 
existing and proposed reserves. 

Description Ornithoptera tithonus is a relatively large birdwing, with a 
forewing length of 70-85 mm in the male and 105-110 mm in the female. The male is 
black, iridescent golden-green and golden-yellow, and the female is dark brown to 
black with yellow and white markings (Plates 3.5 and 3.6). The iridescent scaling of 
the male may vary from gold to green (2,5). O. t. waigeuensis differs slightly from the 
nominate subspecies described below (4), and a second subspecies occurring on 
Misool has yet to be formally described (1,7). O. t. misresiana has recently been 
synonymised with the nominate subspecies (7), but another subspecies, cytherea, has 
been described, which occurs in the Enarokei and Snow Mountains (Pegunungan 
Maoke) area (7). 

Male: UFW black with iridescent golden-green radial, cubital and anal bands. 
UHW elongated and ovoid with a distinct anal notch, a long creamy brush border, a 
large, golden-yellow patch bordered outwardly with green and three black 
postdiscal spots. The wing margin and the broad anal area are black. 

LFW iridescent greenish-yellow with black on the borders, apex, veins, postdiscal 
spots, as a large patch centred round the cell apex, and in a broad anal area (2, 5). 
LHW differs from UHW in narrower black margins and up to three extra postdiscal 
spots (2, 5). 

Female: forewing black, becoming dark brown with age. White markings include a 
transverse cell spot, subapical streaks, discal spots and a series of submarginal spots. 

Hindwing rounded with a slightly scalloped outer margin. Black with a broad, pale 
distal band containing large, black postdiscal spots (2, 5). 

Distribution Ornithoptera tithonus is known to occur only in the western 
part of Irian Jaya on the mainland of New Guinea and on the neighbouring islands of 
Waigeo (Waigeu), Salawati (Salwatty) and Misool (Mysol). The nominate subspe- 
cies occurs on mainland Irian Jaya, with centres of distribution in the Arfak 
Mountains, the Onin Peninsula and at the southern end of Geelvinck Bay; it has also 
been taken far to the east, low down in the Snow Mountains area (4). Subspecies 
waigeuensis occurs in Waigeo and Salawati and the other subspecies is on Misool. 
The distribution of O. tithonus is not well known but the butterfly appears to be 
absent where O. chimaera occurs and vice versa. It has been suggested that it is a 
relict species, the range of which has contracted in recent geological time (5). 

Habitat and Ecology Ornithoptera tithonus inhabits hill forest areas. It does 
not usually ascend as high as O. chimaera and has been found at sea level. However, 

273 



Threatened Swallowtail Butterflies of the World 

it normally ranges up to about 1250 m above sea level and occasionally up to 1900 m 
or above (5). 

Although virtually nothing has been recorded on the ecology, or even life history, 
of O. tithonus, the trade in apparently reared specimens implies that the biology is 
known locally in Irian Jaya. The field notes of C.B. Pratt indicate that both sexes 
nectar on particular trees, or that they assemble round a 'master tree' in the same way 
that some temperate butterflies do (5). 

Threats Only generalised threats to this species can be currently recog- 
nised. However, many pressures on areas where it is hkely to occur have been 
identified: for instance, logging, transmigration of people and prospecting for oil on 
Salawati Island (3), mining in Waigeo (9) and logging in the Arfak and Fakfak areas 

(9). 

Ornithoptera tithonus has a restricted area of occurrence and is possibly a 'relict' 
species, with a range which may continue to contract (5). These factors and the 
possibility of further natural loss of populations may mean that O. tithonus should be 
classed as Rare. 

However, there has also been a considerable trade in specimens, with one report 
that they have been collected in quantity (8). Prices have tended to be intermediate 
between those asked for O. rothschildi and those charged for much more widespread 
species such as O. goliath (8). The effects of commercial collecting on O. tithonus are 
not known. 

Conservation Measures Ornithoptera tithonus is one of the least well known 
birdwings and its conservation depends on adequate research being done to 
determine at least the main features of its ecology. 

Six nature reserves (Cagar Alam) in which O. tithonus has either been recorded or 
is likely to occur have either been established or proposed (9). Pulau Waigeo Barat 
Reserve (153 000 ha), Pulau Salawati Utara Reserve (62 000 ha) and Pulau Misool 
Reserve (105 000 ha) are all proposed nature reserves which should conserve both O. 
t. waigeuensis and the as yet undescribed Misool subspecies. A survey of Salawati to 
include groups other than birds, to make a more comprehensive biological inventory, 
and its wardening to combat the effects of hunting and human population pressure, 
have been recommended (3). On the mainland the most important areas proposed 
for protection are in Pegunungan Arfak (45 000 ha) and Pegunungan Fafak 
(51 000 ha). The nominate O. t. tithonus occurs in both areas, but is particulary 
numerous in the Arfak area (or perhaps has just been collected in greater numbers 
there). Overall priority for these four reserves is either one or two (9). It is possible 
that O. t. tithonus also occurs in the very large and important Lorentz Nature Reserve 
which has already been established (over 2 million ha, proposed reduction to 
1 675 000 ha) and in the small Meriam Hill area which is proposed as a recreation 
park (Taman Wisata). O. t. cytherea occurs in the Enarotali Nature Reserve, 
although it is proposed that this reserve be replaced by a new Weyland Mountains 
Nature Reserve to the west (7, 9) (see review of Ornithoptera paradisea). It thus 
appears that if the Conservation Plan for the Irian Jaya Province is adopted, this 
birdwing will be well catered for in nature reserves. A detailed study of the biology of 
the species is necessary to effectively realise this potential. 

O. tithonus is a protected species in Indonesia, and the effectiveness of this 
protection should be monitored. The species is listed under Appendix II of the 1973 
Convention on International Trade in Endangered Species of Wild Fauna and Flora 
(CITES). Appendix II listing implies that commercial trade is allowed providing a 

274 



Ornithoptera tithonus 

permit from the country of export is obtained. This designation can provide a method 
of monitoring trade levels and should be retained. 

References 

1. Anon. (1984). Review of Ohya (1983). Papilio International 1: 84. 

2. D'Abrera, B. (1975). Birdwing Butterflies of the World. Lansdowne Press, Melbourne. 
260 pp. 

3. Diamond, J.M. et al. (1983). Surveys of five proposed reserves in Irian Jaya, Indonesia: 
Kumawa Mts, Wandammen Mts, Yapen Island, Salawati Island and Batanta Island. Report 
to World Wildlife Fund and to the Directorate of Nature Conservation, Indonesia. 49 pp. , 8 
appendices. 

4. Haugum. J. (1983). In litt..2 June. 

5. Haugum, J. and Low, A.M. (1978, 1979). A Monograph of the Birdwing Butterflies. Vol. 1 
(3): 193-308. Scandinavian Science Press, Klampenborg. 

6. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist S\xpp\cmtnX 17: 1-51. 

7. Ohya, T. (1983). Birdwing Butterflies. Kodansha, Tokyo. 332 pp., 136 col. plates. 

8. Various trade catalogues (1978-84). 

9. UNDP/FAO National Parks Development Project (1981). National Conservation Plan for 
Indonesia. Vol. VII. Maluku and Irian Jaya. FAO, Bogor. 



275 



Threatened Swallowtail Butterflies of the World 

Ornithoptera rothscbildiKennck, 1911 INDETERMINATE 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Ornithoptera rothschildi is a little-known birdwing with a restric- 
ted distribution at high elevations in the mountains of the Arfak area, north-western 
Irian Jaya, Indonesia. Virtually nothing is known about its ecology, but considerable 
numbers of specimens have entered the commercial trade. The first needs are for 
research, survey and assessment of status and a thorough review of the effects of 
commercial exploitation. 

Description Ornithoptera rothschildi, Rothschild's Birdwing, is a relatively 
large birdwing with a forewing length of 63-80 mm in the male and 80-93 mm in the 
female. The male is black, iridescent pale green and golden-yellow, and the female is 
dark brown with creamy-white and yellow markings (1, 2, 4, 6) (Plates 4.1 and 4.2). 
O. okakeae Kobyashi and Koiwaya, is considered to be a natural hybrid between O. 
rothschildi and O. priamus poseidon (4). 

Male: UFW black with iridescent pale green markings. UHW with black margins 
and anal area; inner part golden-yellow with a large iridescent green patch and a 
series of black spots. 

LFW black with markings of iridescent yellow-green, including a broad median 
band, submarginal spots, subapical streaks and two streaks in the cell. LHW differs 
from the UHW in the narrow black margin (1, 2, 4). 

Female: Forewing blackish-brown with creamy-white markings including small 
subapical spots, small submarginal spots, fringe spots and three discal spots. The 
LFW may have more extensive maculation (2, 4). 

Hindwing elongated and rounded with a scalloped outer edge, an irregular black 
outer border, a very broad pale band which is cream-white discally becoming yellow 
distally and a complete series of black subdiscal spots (1, 2, 4, 6). 

Distribution Ornithoptera rothschildi is endemic to the Arfak Mountains 
area of the Beran Peninsula in north-western Irian Jaya (Indonesia). It has the 
smallest range of any of the mainland New Guinea birdwings (4). 

Habitat and Ecology Ornithoptera rothschildi is a montane species and is 
said to replace O. tithonus at high elevation (though the latter species is more widely 
distributed) (4). Most of the records of O. rothschildi are from about 1800-2450 m 
above sea level. The butterflies apparently prefer sheltered valleys and ravines which 
are sunny and protected from strong winds. The vegetation in this habitat is rich but 
shrubby, with some emergent larger trees (4). 

Almost nothing is known about the ecology of O. rothschildi. Flight and behaviour 
seem to be typical for species of Ornithoptera. The species has been supplied in large 
numbers by commercial collectors recently and at least some of the specimens appear 
to have been reared from collected larvae or pupae. Nevertheless, there are no 
recorded descriptions of the early stages or foodplants, nor of larval behaviour and 
natural mortality. 

Threats Ornithoptera rothschildi has a very restricted area of occurrence. 
Although no particular threats to its montane habitats have been identified, the area 
surrounding the Arfak Mountains is relatively densely populated and the cutting of 

276 



Ornithoptera rothschildi 

wood in the forests is a general threat to the region (9). However, O. rothschildi 
would probably be classed as Rare if it were not for doubts about the effects of 
commercial collecting. 

The very considerable trade in this species has caused comment and concern 
(3, 4, 7). High prices were originally quoted when specimens first became available, 
but by 1982 pairs were selling for £10 or less (8). There is no information on the effects 
of commercial exploitation on the butterfly and no means of knowing whether the 
resource is being managed or controlled in any way. The only indication that O. 
rothschildi is not threatened by commercial collecting is that it is not one of the species 
of birdwing which are protected in Indonesia, in contrast to O. chimaera, O. goliath 
and O. paradisea. 

Conservation Measures The primary need for the conservation of Orni- 
thoptera rothschildi is an adequate account of its habitat and ecology and an 
assessment of its status. Without at least an outline of its life history, and 
identification of its foodplants, natural enemies (if any) and actual threats to its 
abundance, no proposals for practical measures to conserve the species can be made . 

However, proposals for a nature reserve (Cagar Alam) of 45 000 ha, the 
Pegunungan Arfak Reserve, which would include part of the range of O. rothschildi, 
have been given a high priority (9). O. rothschildi is only one of many rare and 
endemic species in need of protection by establishment of a reserve here . Included in 
the proposals is a small staff of wardens to check the spread of human settlement in 
the reserve (9). 

The species should be retained, until its status can be reviewed, under Appendix II 
of the 1973 Convention on International Trade in Endangered Species of Wild Fauna 
and Flora (CITES). Appendix II listing implies that commercial trade is allowed 
providing a permit from the country of export is obtained; this can provide a method 
of monitoring trade levels. 

References 

1. D'Abrera, B. (1971). Butterflies of the Australian Region. Lansdowne Press, Melbourne. 
415 pp. 

2. D'Abrera, B. (1975). Birdwing Butterflies of the World. Lansdowne Press, Melbourne. 
260 pp. 

3. Haugum, J. (1983). //; //«.. 2 June. 

4. Haugum, J. and Low, A.M. (1979). A Monograph of the Birdwing Butterflies. Vol. 1 (3): 
193-308. Scandinavian Science Press, Klampenborg. 

5. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
£Ai/owo/og(i/ Supplement 17: 1-51. 

6. Ohya, T. (1983). Birdwing Butterflies. Kodansha, Toyko. 332 pp., 136 col. plates. 

7. Pasternak, J. ( 1981 ). On the rediscovery of Ornithoptera meridionalis tarunggarensis Joicey 
and Talbot on a new locality in Kamrau Bay, south west Irian Jaya, Indonesia. Transactions 
of the Himeji Natural History Association 1981: 2-14. 

8. Various trade catalogues (1977-84). 

9. UNDP/FAO National Parks Development Project (1981). National Conservation Plan for 
Indonesia. Vol. VII. Maluku and Irian Jaya. FAO, Bogor. 



277 



Threatened Swallowtail Butterflies of the World 

Ornithoptera chimaera (Rothschild, 1904) INDETERMINATE 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Ornithoptera chimaera is a specialised birdwing of montane 
biotopes and occurs throughout the mountainous areas of mainland Papua New 
Guinea; it is also found in a few mountain ranges of Irian Jaya. It is restricted to one 
species of foodplant and the density of larvae is low. Threats to the species have not 
been clearly defined, but its status should be effectively monitored. The butterfly is a 
good candidate for controlled utilisation to accompany its conservation. 

Description The male of Ornithoptera chimaera, the Chimaera Birdwing, is 
a large black, iridescent yellow-green and golden butterfly with a wingspan of 
120-155 mm. The female is larger, up to 190 mm, and is dark brown with pale 
markings (1, 2, 6, 11) (Plates 4.3 and 4.4). The description below is for the nominate 
subspecies. A second subspecies, O. c. charybdis, has males with more extensive, 
iridescent yellow-green scaling on the wings. A third subspecies, O. c. fiavidior, 
described in recent monographs (2,6), has now been synonymised with the nominate 
subspecies (9). 

Male: UFW black ground colour with iridescent yellow-green radial, anal and 
cubital bands. UHW rounded with a noticeable anal notch, a large golden area, a 
yellow-green submarginal band, two or three subdiscal black spots and a narrow 
black margin (1, 2, 6, 11). 

LFW lighter and more yellow, with black veins, wide black costal and outer 
margins and a row of black postdiscal spots (2,6). 

Female: forewing dark brown with white or grey markings including a small cell 
spot and small discal, subapical and submarginal spots. 

Hindwing broad, dark brown with a scalloped outer margin and a broad light 
yellow-grey distal band containing large, brown discal spots (1, 2, 6, 11). 

Distribution Ornithoptera chimaera is confined to mainland New Guinea. 
Subspecies chimaera is widely distributed along the central cordillera and has been 
recorded from the Finisterre Mountains and those of the Huon Peninsula, and from 
thirty-one 10 km squares in Papua New Guinea ( 10) . There is an outlying record from 
eastern Irian Jaya (6). 

O. chimaera charybdis is known from central Irian Jaya, from the Wandaman 
Mountains on the east coast of the Vogelkop (Berau Peninsula) through the Weyland 
Mountains (the main centre of occurrence) to an outlying locality in the Pegunungan 
Maoke (Snow Mountains) (6). 

Habitat and Ecology Ornithoptera chimaera is a montane butterfly, 
occurring in areas of tall but fairly open primary forest, often in moderately to very 
steep-sided valleys along water courses. Adults frequent forest margins to collect 
nectar. They may be found in regions from 1200 to 2800 m above sea level, but 
normally occur between 1600 and 1800 m (10). Males congregate round special trees 

(7). 

As far as is known, O. chimaera is monophagous on Aristolochia momandul 
throughout its range (10, 12). A. pithecurus (8) is thought to be synonymous with /I. 
momandul (9). The eggs, which are 4 mm in diameter, are laid on the underside of 
leaves (12), and the larvae are usually solitary (10). The young larvae attack the 

278 



Ornithoptera chimaera 

tender leaves and shoots of the foodplant but mature caterpillars eat older leaves and 
may occasionally chew the bark of the main stem. Feeding occurs mainly in the early 
morning and late afternoon (10). In the field, the egg stage lasts 14 days, the larval 
stage probably about 2 months, and 49-70 days are spent as a pupa ( 10) . Some larval 
cannibalism occurs and attack by the braconid wasp, Apanteles cf. vitripennis , may 
be heavy in some localities ( 10, 12). The early stages have been excellently figured in 
colour (8, 12). 

Adults appear to range widely, particularly females in search of oviposition sites. 
Although the data come from a small sample, females which have been dissected 
were found to be carrying either no eggs or from 6-10 (10). There are no records of 
vertebrate predation on adult butterflies (10). 

Threats Ornithoptera chimaera is restricted not just to the mainland of 
New Guinea but to areas of medium to high elevation, and to a single species of host 
plant. Its existing habitat in Papua New Guinea is limited but difficult to exploit 
commercially, e.g. by extraction of timber (10). Like many birdwings, O. chimaera is 
a K-selected species, producing few, well-protected offspring compared with many 
other butterflies. Although ecologically highly specialised, it can be quite common at 
times in some localities and cannot be considered as 'very rare' (10, cf. 1). 

In Irian Jaya, O. chimaera is much less generally distributed and its habitat in the 
Wandamen Mountains may be under some threat from forest fires, depending on 
how high into the montane habitat these fires extend (3). Illegal collecting of 
vertebrates occurs in the Wandamen Mountains, but it is not known if protected 
birdwings, of which O. chimaera is one, are also collected (3). 

O. chimaera is at risk in the Weyland Mountains area because if logging increases 
here, soil erosion would follow on the steep slopes of the mountains (13). 

Certainly O. chimaera is in demand as specimens for study and display. No legal 
trade is possible in Papua New Guinea and until recently the trade was only in O. c. 
charybdis. There is no information on whether trade, legal or illegal, has had any 
adverse effect on the abundance of the butterfly. 

Conservation Measures Although because of its high altitude habitat O. 
chimaera is not greatly threatened at the present time, its status needs continual 
monitoring in both Papua New Guinea and Irian Jaya. The effects of forest fires and 
timber extraction, where it occurs at such an elevation, need to be assessed. 

In Papua New Guinea, the establishment of five nature reserve areas should be 
sufficient to conserve O. chimaera, without the need to undertake specific 
management of habitat and populations (10). The suggested locations for these 
reserves are Telefomin, Bundi, Naniwe Mission, Tapini-Woitape and Central Huon 
(10). 

Two proposed nature reserves (Cagar Alam) in Irian Jaya almost certainly contain 
populations of O. chimaera. The Weyland Mountains Nature Reserve is proposed to 
replace the existing Enarotali Nature Reserve which lies to the east and is unsuitable 
because of its large human population (13) {see rewiew of Ornithoptera paradisea) . In 
elevation the proposed reserve varies from 900 to over 3800 m and so includes the 
entire altitudinal range of O. chimaera (10, 13). 

The proposed Pegunungan Wandamen/Wondiwoi Nature Reserve includes most 
of the mountainous part of the Wandamen Peninsula, ranging from sea level to more 
than 2200 m and comprises 79 500 ha (13). This area contains a notable fauna of 
montane birds, and it has been suggested that a small local industry of natural history 
tours could be established, particularly to see bowerbirds (Ptilonorhynchidae) and 

279 



Threatened Swallowtail Butterflies of the World 

birds of paradise (Paradisaeidae) (3). Ornithoptera chimaera could easily be included 
as a tourist attraction. 

Suggestions have been made that Ornithoptera chimaera should be brought back 
into the area of legitimate trade (10). This is not the retrograde step in conservation 
terms that it superficially appears to be: "Ironically it is now becoming an accepted 
fact that the very demand for Ornithoptera is one of the main assets which will ensure 
their future survival if they can be exploited in the correct way." (10). A possible 
alternative, or first step, might be careful husbandry in a Wildlife Management Area 
in Papua New Guinea (7). 

If O. chimaera were to be brought back into commerical trade , this should be under 
strict control and the effects on populations and abundance would need to be closely 
and effectively monitored. Information on the effects of trade on the numbers of any 
species of birdwing is urgently required. If controlled trade in O. chimaera were 
permitted, the protected status of the species would have to be rescinded, or at least 
suspended. If controlled utiHsation of the resource were successful, the species could 
be removed from Appendix II of CITES (10). 

References 

1. D'Abrera, B. (1971). Butterflies of the Australian Region. Lansdowne Press, Melbourne. 
415 pp. 

2. D'Abrera, B. (1975). Birdwing Butterflies of the World. Lansdowne Press, Melbourne. 
260 pp. 

3. Diamond, J.M. et al. (1983). Surveys of five proposed reserves in Irian Jaya, Indonesia: 
Kumawa Mts, Wandammen Mts, Yapen Island, Salawati Island, and Batanta Island. 
Report to World Wildlife Fund and to the Directorate of Nature Conservation, Indonesia. 
49 pp., 8 appendices. 

4. Fenner, T.L. (1983). In litt., 15 March. 

5. Haugum, J. (1983). /n litt., 2 June. 

6. Haugum, J. and Low, A.M. (1979). A Monograph of the Birdwing Butterflies. Vol. 1 (3): 
193-308. Scandinavian Science Press, Klampenborg. 

7. Hutton, A.F. (1984). In litt., I February. 

8. Igarashi, S. (1979). Papilionidae and Their Early Stages. Vo\. 1: 219pp., Vol. 2: 102 pp. of 
plates. Kodansha, Tokyo. (In Japanese). 

9. Ohya, T. (1983). Birdwing Butterflies. Kodansha, Tokyo. 332 pp., 136 col. plates. 

10. Parsons, M.J. (1983). A conservation study of the birdwing butterflies Ornithoptera and 
Troides (Lepidoptera: Papilionidae) in Papua New Guinea. Final report to the Department 
of Primary Industry, Papua New Guinea. 1 1 1 pp. 

11. Smart, P. (1975). The Illustrated Encyclopedia of the Butterfly World. Hamlyn, London. 
275 pp. 

12. Straatman, R. and Schmid, F. (1975). Notes on the biology oi Ornithoptera goliath and O. 
chimaera (Papilionidae). Journal of the Lepidopierists' Society 29: 85-88. 

13. UNDP/FAO National Parks Development Project ( 1981 ). National Conservation Plan for 
Indonesia. Vol. VII. Maluku and Irian Jaya. FAO, Bogor. 



280 



Ornithoptera paradisea 
Ornithoptera paradisea (Staudinger, 1893) INDETERMINATE 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Ornithoptera paradisea is a particularly attractive, sexually- 
dimorphic butterfly which is widespread, though very localised, throughout northern 
mainland New Guinea as six described subspecies, several ill-defined. It is apparently 
extinct in one well-known area, but no obvious specific threats are known. It should 
be surveyed to establish its status, reserves should be established, and the future for 
ranched specimens as a marketable resource should be considered. For the present, it 
should remain protected in Papua New Guinea and Irian Jaya, and remain on 
Appendix II of CITES. 

Description Ornithoptera paradisea is variously known as the Paradise 
Birdwing, Tailed Birdwing or Butterfly of Paradise. The male is a beautiful, black, 
iridescent yellow-green and golden, tailed butterfly with a wingspan of 100-130 mm 
(Plate 4.5). The dark brown female with yellow and white or creamish markings has 
an average wing span of 160 mm (2, 3, 6, 8, 10) (Plate 4.6). There is much variation 
both within and between subspecies. Six subspecies of O. paradisea have been 
described, including the nominate O. p. paradisea, but several are ill-defined, 
poorly-known, or both. The validity of the subspecies of this birdwing and the 
differences between them, must therefore be treated with caution, particularly as 
some populations have not yet been reliably referred to subspecies (6, 10). These 
include O. p. arfakensis, frequently encountered in the butterfly trade, and O. p. 
tarunggarensis formerly described as a subspecies of O. meridionalis (5). 

Male: UFW of the nominate subspecies broad and elongated with a pointed apex 
and a rounded outer margin, black with broad, iridescent light yellow-green radial 
and cubital bands and a relatively short anal band. UHW reduced in size and almost 
triangular with a distinct apex, anal notch, and a tail at vein two which is a further 
20-25 mm long (6) . Black scaling limited to narrow costal and outer margins, and to a 
relatively narrow anal area. There is a broad yellow-green submarginal border and a 
variable , large , golden-yellow patch . Long white or cream hairs form a brush-border 
to the anal margin. 

LFW predominantly covered with iridescent, light yellow-green scales, more 
golden than those on the upper surface. Slight black scaling around the veins, on 
narrow wing margins, and in a part of the wing apex (absent in eastern populations). 
LHW almost entirely Hght yellow-green and golden with a pale silver anal area 
(2,3,6,8,10). 

Female: forewing pale to very dark brown with pale cream markings, including 
relatively large subapical streaks, submarginal spots and sometimes up to three discal 
spots. Hindwing elongated with a pale discal band, white towards the cell and yellow 
distally, containing a row of relatively small, black subdiscal spots (2, 3, 6, 10). 

Distribution Ornithoptera paradisea is widely distributed on the mainland 
of New Guinea and has been reported more or less unreliably (sightings and dealers' 
records) from a number of adjacent islands (6). In Papua New Guinea, O. paradisea 
may be extinct in its well-known 19th century localities in Madang Province (9). 
There is a good sprinkling of localities in the north-west of the country in the Maprik 
and adjacent areas and recent records from the Lake Kutubu region (information 
about which post-dates the map in Haugum and Low (6, 9)). There are no recent 

281 



Threatened Swallowtail Butterflies of the World 

records from the eastern half of mainland Papua New Guinea, nor from any of the 
islands. In all, there are modern records from seventeen 10 km squares (9). 

In Irian Jaya, O. paradisea is also very local, but the butterfly is well distributed, 
particularly in the west (though not the extreme west) and in the north-east (f. 
borchi); it has not been taken in the south-east of the country. There are scattered 
records from the Penunungan Maoke (Snow Mountains) westwards to Kamrau Bay, 
with a fairly well known area of occurrence in the Arfak region of the north-west (ssp. 
arfakensis) (6). 

Habitat and Ecology Ornithoptera paradisea occurs in mature secondary 
and primary forest in hilly areas, where it frequents clearings, valleys, gulleys and 
gorges and also flies on slopes and ridges. It is normally found at altitudes between 200 
and 800 m but there are records of occasional butterflies being found much higher, 
even once at 2000 m (1, 4, 6, 9). 

The foodplant is a species of Aristolochia closely related to A . momandul, to which 
the manuscript name of ^4 . "pseudo-momandul" has been given for ease of reference 
(9). The eggs are normally deposited singly on the underside of the leaves of the 
foodplant, sometimes, but rarely, on other objects nearby (1). The eggs are 
particularly large (4 mm diameter) and so probably few eggs are carried by a female, 
perhaps 8-10 at any one time. Eggs hatch in 10-12 days, the larval stage takes 36-40 
days and the pupal stage 37 days (1). The immature stages therefore last about three 
months. The early stages and foodplant have all been well figured in colour (1,8). 

There is mortality in the egg stage from parasitic wasps. The larvae are taken by 
various vertebrate predators and attacked by braconid wasps. They are also 
destroyed by bad weather (1). Although mortality is said to be high, its significance 
has not been quantified. 

Adults appear to be fairly localised in their movements and do not move long 
distances. They use valleys and gorges as flyways, and collect nectar frequently on 
various flowers. They probably live about three months. 

Threats No clear threats to Ornithoptera paradisea have been identified. 
The possible extinction of the Madang populations gives rise to concern, but no 
plausible cause has been suggested. The birdwing may be Vulnerable but is best 
regarded as Indeterminate in status because of local extinction due to unknown 
causes, together with its patchy distribution. It should be classed as Rare if threats are 
not serious; although its area of distribution is not particularly narrow, it is very 
generally regarded as an uncommon butterfly. 

In addition, it is much in demand as specimens. In recent years, considerable 
numbers of O. p. arfakensis have come into trade, but there is no evidence that this 
has been a threat to its continued existence or numbers (12). 

Conservation Measures In Papua New Guinea, the conservation of this 
species falls into three parts. First , the distribution records and assessment of status of 
Ornithoptera paradisea appear to be incomplete. Survey of the butterfly is certainly 
necessary and should include a thorough field survey of the Madang area. Secondly, 
reserves need to be established under the authority of the Conservation Areas Act 
(1978), to prevent undue loss of habitat. Five National Reserve Areas have been 
suggested as the optimum number to conserve this and other Ornithoptera spp., at 
South Vanimo, Maprik, Frieda River, Lake Kutubu and East Erave (9). Thirdly, the 
utilisation of the resource , possibly as a tourist attraction , but particularly for farming 

282 



Ornithoptera paradisea 

of specimens, should be considered in the long term (7, 9). Even a limited, trial 
production of farmed specimens is clearly some years away. 

In Irian Jaya, O. paradisea appears to be present in several reserves or proposed 
reserves. Perhaps the most important is the proposed Pegunungam Arfak Nature 
Reserve (Cagar Alam), which has overall priority one for establishment (see review 
of O. tithomis) (11). O. p. arfakensis is the subspecies which would be protected by 
this reserve. Other populations of O. paradisea are likely to occur in the established 
Lorentz Nature Reserve (see under review of O. tithonus) and in the proposed 
Weyland Mountains Nature Reserve. This area of 228 000 ha has been recom- 
mended to replace the Enarotali Nature Reserve of 300 000 ha which lies to its east 
(1 1). The Weyland Mountains reserve has an overall priority of one for establishment 
and almost certainly contains at least some of the area's populations of O. paradisea 

(ID- 
AS in Papua New Guinea, an assessment of the birdwing's status and a survey of its 

ecological requirements are both necessary. Utilisation of the resource, both as a 

tourist attraction and in the form of carefully-controlled farming for the specimen 

trade, could also be considered. 

For the present, however, O. paradisea should retain its status as a fully protected 

species in Papua New Guinea, as a protected species in Irian Jaya (Indonesia), and 

the butterfly should be retained in Appendix II of the 1973 Convention on 

International Trade in Endangered Species of Wild Fauna and Flora (CITES). 

Appendix II listing imphes that commercial trade is allowed, providing a permit from 

the country of export is obtained; this can provide a method of monitoring trade 

levels. This situation should remain at least until effective conservation measures can 

be implemented. 

References 

1. Borch, H. and Schmid, F. (1975). The life cycle o{ Ornithoptera paradisea (Papilionidae). 
Journal of the Lepidopterists' Society 29: 1-9. 12 col figures. 

2. D'Abrera, B. (1971). Butterflies of the Australian Region. Lansdowne Press, Melbourne. 
415 pp. 

3. D'Abrera, B. (1975). Birdwing Butterflies of the World. Lansdowne Press, Melbourne. 
260 pp. 

4. Fenner, T.L. (1983). In litt., 15 March. 

5. Hancock, D.L. (1983). A note on the status of Ornithoptera paradisea tarunggarensis 
(Joicey & Talbot) (Lepidoptera: Papilionidae). Australia Entomologists' Magazina 8: 
93-95. 

6. Haugum, J. and Low, A.M. (1979). A Monograph of the Birdwing Butterflies. Vol. 1 (3): 
193-308. Scandinavian Science Press, Klampenborg. 

7. Hutton, A.F. (1984). In litt., 1 February. 

8. Igarsishi^S. (1979). Papilionidae and Their Early Stages. yo\. 1: 219pp., Vol. 2: 102 pp. of 
plates. Kodansha, Tokyo. (In Japanese). 

9. Parsons, M.J. (1983). A conservation study of the birdwing butterflies Ornithoptera and 
Troides (Lepidoptera: Papilionidae) in Papua New Guinea. Final report to the Department 
of Primary Industry, Papua New Guinea. Ill pp. 

10. Smart, P. (1975). The Illustrated Encyclopedia of the Butterfly World. Hamlyn, London. 
275 pp. 

11. UNDP/FAO National Parks Development Project (1981). National Conservation Plan for 
Indonesia. Vol. VII. Maluku and Irian Jaya. FAG, Bogor. 

12. Various trade catalogues (1978-84). 



283 



Threatened Swallowtail Butterflies of the World 

Ornithoptera meridionalis (Rothschild, 1897) VULNERABLE 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Ornithoptera meridionalis is a birdwing butterfly with a high 
degree of sexual dimorphism. It is restricted to New Guinea, where it is a rare and 
very localized species of lowland forest. It occurs mainly in south-eastern Papua New 
Guinea but is also known from Irian Jaya, Indonesia. It is threatened by increased 
extraction of timber from its localities, many of which are very accessible. Reserves 
should be established for O. meridionalis and these could be managed in part as 
tourist showpieces. Protected status for the species should be maintained. 

Description The male of Ornithoptera meridionalis has tailed hindwings, a 
wingspan of 80-115 mm (3) and is black, iridescent yellow-green and golden (Plate 
5.1). The male is, on average, much smaller than any other male Ornithoptera, 
including that of the closely-related O. paradisea. The female is considerably larger 
than the male, black with white and yellow markings (1, 2, 4, 6, 10) (Plate 5.2). 

Male: UFW similar to that of O. paradisea, black with broad, iridescent 
yellow-green radial and cubital bands and a short anal band. UHW curved inwards 
and much reduced in size, tailed, with a small, diamond-shaped tip, a reduced black 
anal area and an extensive yellow-gold patch. 

LFW iridescent yellow-green, more golden than on the upper surface. Black 
scaling covers the veins, the narrow costal and outer margins, a large cubital patch 
and the wing apex. LHW similar to the upper surface with a pale grey brush border 
and no black scaling (1, 2, 4, 6, 10). 

Female: Forewing black with white markings including a large cell spot, subapical 
streaks, submarginal spots and three large discal spots. 

Hindwing rounded, black with a broad distal band, extending over the discal 
region and part of the cell. It is yellow and suffused with black scales on the distal side 
of a row of large, black subdiscal spots, and white on the discal side (1, 2, 4, 6, 10). 

Distribution Ornithoptera meridionalis has been found only in the island of 
New Guinea. Until recently it was thought that the butterfly occurred only in Papua 
New Guinea. Females of a separate subspecies, tarunggarensis, occurring at 
Nomnangihe, 40 km south-west of Wanggar and highly disjunct with the Papua New 
Guinea populations, have been shown to be referable to O. paradisea, not O. 
meridionalis (3). However, O. meridionalis has been discovered recently to be truly 
present in Irian Jaya, in the region of Kamrau Bay about 200 km west of the Weyland 
Mountains, in the Weyland Mountains themselves and in the area around Lake 
Yamur (Jamur) to the east of the Weyland Mountains (3, 7, 9). 

However, the name tarunggarensis has been most confusingly applied to these 
populations (7, 9). If, as seems likely, they represent a subspecies distinct from O. m. 
meridionalis, it currently lacks a valid name. 

O. meridionalis is quite widely distributed in mainland Papua New Guinea, but 
very local. Its main area of occurrence is along the southern part of the south-eastern 
peninsula, but it has also been recorded from single localities in the Southern 
Highlands and East Sepik Provinces. It has not been reported from any of the islands 
and is known from only fourteen 10 km squares in the country (8). 

Habitat and Ecology Ornithoptera meridionalis occurs mainly in lowland 
rain forest, both primary and mature secondary, usually between 20 and 200 m above 

284 



Ornithoptera meridionalis 

sea level. However, it probably extends into hill forest ifitsfoodplant is present. The 
specimen from the Southern Highlands Province was captured at 800 m (8). In the 
Kamrau Bay population in Irian Jaya, the butterfly is also a mainly lowland species, 
though one population was found to be established at about 700 m (9). 

There is doubt as to the exact species of foodplants. Some accounts state that the 
larvae are monophagous on Aristolochia dielsiana (known in the literature as A. 
schlechteri), others that A. pithecurus (probably synonymous with A. momandul) is 
the main foodplant (5,6, 11, 12). Larvae in the Kamrau Bay area feed on A. 
dielsiana, a species oi Aristolochia indistinguishable from A. scfilechteri (9). 

The male butterflies fly poorly and are less often seen than the males of other 
Ornithoptera species; this is evidently correlated with the small size and unusual 
shape of the hindwing (9). Adult females seem to fly mainly in their home range, 
although they have also been seen flying along tracks in open secondary forest (8). 

Females deposit eggs singly on the undersides of the foodplant leaves. The eggs are 
large (3 mm diameter) and females carry very few, usually between five and seven 
(8, 11, 12). Females of the Kamrau Bay populations apparently do not oviposit on 
foodplants growing on steeply-sloping terrain. In this population, 50-60 per cent of 
eggs were attacked by parasitic flies (9), probably a species of Trichogramma 
(Hymenoptera: Chalcidoidea). The parasite was not observed to attack eggs of 
Ornithoptera priamiis on the same foodplant and may be specific to O. meridionalis 
(9). Feeding by mature larvae is characteristic: the central disc of the leaf is eaten 
from the apex, leaving a crescent-shaped area uneaten. Both larvae and pupae have 
been described and figured (6, 11. 12). 

Natural enemies of the larvae include birds, tree frogs, a species of lizard and 
invertebrates such as large reduviid bugs and spiders (9). 

Threats The Irian Jaya populations of O. meridionalis do not appear to be 
so much at risk as those of Papua New Guinea (8, 9). Indeed, it is probably the 
inaccessibility of the Irian Jaya localities, and paucity of human settlements, which 
has delayed their discovery. 

However, the Papua New Guinea populations are seriously threatened by habitat 
destruction and change. These low-lying localities are particularly subject to 
commercial extraction of timber. Some of the main areas of occurrence are close to 
Port Moresby and are accessible by road, an unusual circumstance in Papua New 
Guinea. Logging is increasing in the region of the Brown and Vanapa Rivers, which 
lies in the centre of the range of O. meridionalis in south-eastern Papua New Guinea 
(5,8). 

Specimens of O. meridionalis are much in demand by collectors. The birdwing 
occurs only in New Guinea and is totally protected in Papua New Guinea. It is not 
protected in Irian Jaya, probably because the butterfly was so little known in that 
country until recently. No threats are known to be posed by illegal collecting and 
trade, nor by legitimate dealing. However, the butterfly is one of the most valuable 
known; a pair was offered in Britain for £650 in 1980 (13). It can be expected that 
commercial trade in specimens from Irian Jaya will be developed now that the 
locaHties are known there. 

Conservation Measures Because of the main threat to O. meridionalis in 
Papua New Guinea, the most important measure to be taken for the butterfly's 
conservation is the establishment of National Reserve Areas under the Conservation 
Areas Act. The establishment of five reserves has been suggested, with priority given 
to the Brown and Vanapa Rivers areas (8). Because this area is easily accessible from 

285 



Threatened Swallowtail Butterflies of the World 

Port Moresby, the capital of Papua New Guinea, conservation could be combined 
with a tourist attraction by the establishment of a Wildlife Management Area. 
Enrichment of the habitat should be considered by planting the host of O. 
meridionalis , Aristolochia dielsiana, as has been done for the common birdwings by 
planting yl. tagala (8). 

The other four reserves in Papua New Guinea suggested for O. meridionalis are at 
Lake Kutubu, the Frieda River, Cape Rodney and Mamai Plantation. At Lake 
Kutubu, located in the Southern Highlands, three uncommon birdwing species 
(Ornithoptera goliath, O. paradisea and O. meridionalis) could be conserved (8). 

The Irian Jaya populations of O. meridionalis appear to lie mainly outside the 
boundaries of existing or proposed reserves. This exception may be the proposed 
Weyland Mountains Nature Reserve (Cagar Alam), which has been put forward to 
replace the unsuitable Enarotali Nature Reserve (see review of O. chimaera) (14). 
Although the conservation of O. meridionalis in a national reserve in Irian Jaya is 
desirable, the recent discovery of the species in new areas suggests that survey and 
exploration are the first priorities. 

In the long term, the future of O. meridionalis may be linked with its carefully 
controlled utilisation, both as a tourist showpiece, at least in Papua New Guinea, and 
as a sustainable resource of specimens (8). 

O. meridionalis is a totally protected species in Papua New Guinea, so that any 
controlled utilisation would require amending legislation. The butterfly is listed 
under Appendix II of the 1973 Convention on International Trade in Endangered 
Species of Wild Fauna and Flora (CITES). This status should be retained. Appendix 
II listing implies that commercial trade is allowed, providing a permit from the 
country of export is obtained. If enforced, this can provide a method of monitoring 
the amount of trade. 



References 

1. D'Abrera, B. (1971). Butterflies of the Australian Region. Lansdowne Press, Melbourne. 
415 pp. 

2. D'Abrera, B. (1975). Birdwing Butterflies of the World. Lansdowne Press, Melbourne. 
260 pp. 

3. Hancock, D.L. (1982). A note on the status of Ornithoptera meridionalis tarunggarensis 
(Joicey& Talbot) (Lepidoptera: PapWionidae). Australian Entomological Magazine93-95. 

4. Haugum, J. and Low, A.M. ( 1978-9). A Monograph of the Birdwing Butterflies. 
Vol. 1 (3): 193-308. Scandinavian Science Press, Klampenborg. 

5. Hutton, A.F. (1984). In litt., 1 February. 

6. Igarashi, S. (1979). Papilionidae and Their Early Stages. Vol. 1 , 219 pp. , Vol. 2, 102 pp. of 
plates. Kodansha, Tokyo. (In Japanese). 

7. Ohya, T. (1983). Birdwing Butterflies. Kodansha, Tokyo. 332 pp., 136 col. plates. 

8. Parsons, M.J. (1983). A conservation study of the birdwing butterflies, Orn/f/ioprera and 
Troides (Lepidoptera: Papilionidae) in Papua New Guinea. Final report to the 
Department of Primary Industry, Papua New Guinea. 1 1 1 pp. 

9. Pasternak, J. (1981). On the rediscove-ry of Ornithoptera meridionalis tarunggarensis 
Joicey & Talbot in a new locality in Kamrau Bay, south-west Irian Jaya, Indonesia. 
Transactions of the Himeji Natural History Association 1981 : 2-14. 

10. Smart, P. ( 1975). The Illustrated Encyclopedia of the Butterfly World. Hamlyn, London. 
275 pp. 

11. Straatman, R. (1967). Additional notes on the biology of Ornithoptera meridionalis 
(Rothschild). Transactions of the Papua New Guinea Science Society 8: 36-38. 

12. Szent-Ivanny, J.J.H. and Carver, R. A. (1967). Notes on the biology of some Lepidoptera 
of the Territory of Papua New Guinea with descriptions of the early stages of Ornithoptera 

286 



Ornithoptera meridionalis 

meridionalis Rothschild. Transactions of the Papua New Guinea Science Society 8: 3-35. 

13. Trade catalogue (1980). 

14. UNDP/FAO National Parks Development Project (1981). National Conservation Plan for 
Indonesia. Vol. VII. Maluku and Irian Jaya. FAO, Bogor. 



287 



Threatened Swallowtail Butterflies of the World 

Ornif/iopteraa/exandrae (Rothschild, 1907) ENDANGERED 

Subfamily PAPILIONINAE ^ Tribe TROIDINI 



Summary Ornithoptera alexandrae. Queen Alexandra's Birdwing, is the 
worid's largest butterfly. It is restricted to primary and advanced secondary lowland 
rain forest in or near the Popondetta Plain, a small area in the Northern Province of 
Papua New Guinea. Protected by law since 1966, the species is not often collected, 
but its habitat is now severely threatened by the expanding oil palm and logging 
industries. Conservation measures taken and proposed include the establishment of 
reserves and a wide range of ecological survey and research. 

Description Ornithoptera alexandrae is the world's largest butterfly, the 
dark brown females having a wingspan of up to 250 mm (Plate 5.4). The male is 
smaller (wingspan 170-190 mm) and is light blue, yellow, green and black 
(3, 4, 9, 12) (Plate 5.3). The average head and body length is about 75 mm; the 
abdomen of both sexes is bright yellow and the ventral wingbases are bright red. 

Male: UFW elongated, black with a long, broad, green radial band, a broad, 
blue-green anal band fused at both ends with a narrower cubital band to enclose a 
large, black sex brand. UHW elongated, black with a broad, blue-green submarginal 
band which is contined to the base along the costal and inner margins, and a broad, 
blue-green cell streak. 

LFW blue-green with more blue towards the anal region; black veins, narrow 
margin and subapical streaks. LHW yellow becoming bluish towards the anal region, 
with black veins and narrow margins. 

Female: forewing dark brown with relatively small, pale grey submarginal and 
discal spots which become smaller towards the apex. 

Hindwing elongated, with a band of seven pale grey, yellow-powdered, wedge- 
shaped patches separated by broad bands over the veins. All but the innermost 
contain brown discal spots, and all are more yellow on the UHW. 

Distribution The first specimen, a small, dull, atypical female, was col- 
lected in 1906 from the type locality high on the upper reaches of the Mambare River, 
well outside its present range (9, 13). To date, Ornithoptera alexandrae has on\y been 
recorded from nine 10 km grid squares on the Popondetta Plain in Northern 
Province, Papua New Guinea, and is known from only one other locaHty as a 
separate, high altitude population not far from the larger lowland population (18). It 
is reported that the 1951 eruption of Mt Lamington destroyed 250 sq. km of prime 
habitat, further fragmenting the already patchy distribution produced by agriculture 
and logging (10). 

Habitat and Ecology Ornithoptera alexandrae occurs with its larval food- 
plant, Aristolochia dielsiana (formerly known as Aristolochia schlechteri) , in secon- 
dary and primary lowland rain forest up to 400 m altitude on the volcanic ash soils of 
the Popondetta Plain, and in secondary hill forest on clay soils from 550 m to 800 m 
altitude in its other locality (8, 16, 17). It is strictly monophagous, although this is due 
to the oviposition specificity of the female as the larvae can mature equally well (and 
apparently even better) on the softer-leaved Aristolochia tagala, a vine which is 
common and far more widespread throughout Papua New Guinea (21,22). The 
much commoner Papua New Guinea birdwing, Ornithoptera priamus , uses the same 

288 



Ornithoptera alexandrae 

foodplant as O. alexandrae. Whether any competition occurs between the two 
species is uncertain. The duration of the early stages (from egg to aduh emergence) 
exceeds four months and adults can live up to a further three months in the wild. 
Adults are subject to little predation but eggs are attacked by ants and heteropterous 
bugs. The larvae are preyed upon by toads, lizards and birds such as cuckoos, drongos 
and crow pheasants. Parasitism of larvae by unidentified tachinid flies, and of pupae 
by parasitic wasps, has been reported (21). Opinions vary as to whether parasitism 
occurs commonly or rarely (19, 21) and this is a topic which requires much more 
research. However, it is believed that the aposematic (warning) coloration of the 
larvae and adults is an indication that they can probably store the toxins that their 
foodplants are known to contain, using them for their own protection against more 
general predators (15). Adults are strong fliers but appear to remain in home ranges, 
ignoring other available habitat. It has been established recently that male butterflies 
often swarm around a large timber tree, Intsia bijuga (Leguminosae, known locally as 
Kwila), when it is in flower (10). Observations indicate that flying females will not 
accept males unless they have visited the flowers (10). Experimental confirmation of 
the behaviour pattern is needed, but the distribution of the tree may account for the 
absence of the butterfly from certain apparently suitable areas (10), although /. bijuga 
is a common and widely distributed species. 

The eggs of O. alexandrae are extremely large (4 mm diameter) and it has been 
calculated that females, if their ovaries are continuously productive, have the 
potential to lay about 240 eggs during their lifetime (16). They possibly carry only 
15-20 (maximum 30) at any one time (11). 

Conventional mark-recapture methods cannot be used to estimate numbers of O. 
alexandrae as the species flies high and is too infrequently seen. Larval counts are also 
low (only one or two may be located during a day's survey) and the leaves of the 
foodplant vine are often 40 m high in the upper canopy, effectively precluding 
observation of larvae. 

Threats The greatest current danger is the expanding oil palm industry in 
the Popondetta region, although cocoa and rubber plantations have also been a 
problem in the past. These have already claimed large tracts of forest known to have 
been habitat for Ornithoptera alexandrae (1). Negotiations to exploit the reserves of 
wood in the Kumusi Timber Area are also in progress. Localized extinctions are 
occurring due to the clearing of forest to make food gardens. During the Second 
World War, Popondetta was an important air base, and at one time contained 26 
airstrips (7). 

O . alexandrae is greatly prized by collectors and some illegal trade has undoubtedly 
occurred from time to time (see below) . However, illegal collecting is not comparable 
with loss of habitat as a threat. 

Conservation Measures In 1966, the Fauna Protection Ordinance gave O. 
alexandrae and six other birdwings legal protection from collection (5). The law has 
been stringently enforced on several occasions, resulting in fines for nationals and 
deportation of expatriates. Surveys by the Division of Wildlife are establishing the 
presence or absence of O. alexandrae in defined areas. A large Wildlife Management 
Area (WMA) , comprising approximately 1 1 000 ha of grassland and forest, has been 
established north of Popondetta. Unfortunately, it is not at all clear how effective this 
WMA is in conserving wildlife in general, and O. alexandrae in particular. Reports 
suggest that the WMA has agriculture within its boundaries. Several thousand 
cuttings oiA. dielsiana are being prepared and an area of 4 ha of government owned 

289 



Threatened Swallowtail Butterflies of the World 

primary forest at the Lejo Agricultural Station is being planted as a future reserve and 
study area for O. alexandrae . The Wildlife Division has applied for a total of about 
40 ha of government land that has been rejected for use as oil palm plantations 
because of the deeply dissected topography. The aim is to create reserves for O. 
alexandrae on government land, which can be protected by law in perpetuity. A trial 
planting of ^4. dielsiana cuttings under tall, shady, mature (c. 14 years old) oil palms 
has been undertaken at the Popondetta Agricultural Training Institute to study the 
growth of the vines in this artificial habitat and to see whether O. alexandrae will 
eventually utilize them. Provincial wildlife officers regularly hold educational 
meetings with people in the Northern Province, to explain why the butterfly needs to 
be conserved. Representations for conservation of the species have been made to the 
Government of Papua New Guinea by several international bodies, including the 
lUCN/SSC Butterfly (formerly Lepidoptera) Specialist Group. 

There are well-defined plans for future conservation efforts. In particular, 
negotiations to establish new WMAs are in progress between the Wildlife Division 
and interested landowners. Proposals for three reserve areas within the Kumusi 
Timber area have been supported by the landowners and the timber company 
involved (Fletcher Forests, New Zealand). Implementation of the recent Conser- 
vation Areas Act (1978), which gives special protection to "sites and areas having 
particular biological, topographical, geographical, historic, scientific or social 
importance" , is being considered for certain sites. The Act also provides for the active 
management of such areas (2). It is hoped that portions of prime Ornithoptera 
alexandrae habitat will be considered for inclusion under this Act. 

The discovery of Aristolochia dielsiana on Siassi I. (= Umboi I.) in Morobe 
Province has prompted the suggestion that an establishment of O. alexandrae should 
be made there (19). Although O. priamus occurs on Siassi I. (and feeds on A. 
dielsiana), Troides oblongomaculatus , a potential competitor, does not. The 
suggestion of an establishment here is an important contribution to the positive 
conservation of O. alexandrae , but of course, a thorough survey of the island and the 
occurrence oiA. dielsiana on it is necessary before an estabHshment can be planned. 

O. alexandrae is the largest butterfly in the world and is aesthetically very 
attractive. The birdwings have long been held in high esteem by insect collectors and 
are in great demand worldwide. Species such as O. alexandrae , which are not only 
impressive but restricted in their range and hard to obtain, realise extremely high 
prices. Within the Division of Wildlife in Papua New Guinea, there is already a 
marketing agency which supplies insect dealers with the unprotected insects of the 
country (14). If the long term future of O. alexandrae is safeguarded, it could provide 
an extremely valuable income to the people of Papua New Guinea (6). Eventually the 
butterfly may become an added attraction to the growing tourist industry (20). 

The resource provided by this butterfly could be utilised even now. It has been 
suggested that some of the 5 1 ex-pupa papered specimens of O . alexandrae ( 19 males, 
32 females), confiscated as having been taken illegally, should be offered for sale, 
possibly by a 'sealed bid' method (19). These specimens are currently kept in papers 
at the Entomology Department, DPI, where they have no scientific purpose. Their 
sale could partially finance conservation programmes for this butterfly. 

It may be possible to breed O. alexandrae in captivity so that its biology and the 
reasons for its monophagy can be more closely studied. However, extremely large 
flight cages are required if the species is to behave normally in captivity, and the cost is 
prohibitive. Experiments to breed selectively for a culture of O. alexandrae which 
oviposits on Aristolochia tagala may prove rewarding (20). Despite its attractions, O. 
alexandrae is poorly known because it is so rare, and further research on its life 

290 



Ornithoptera alexandrae 

history, behaviour, natural enemies and population dynamics should be undertaken 
at the same time that conservation measures are put into effect. 

O. alexandrae is currently included, together with all other birdwings, in Appendix 
II of the Convention on International Trade in Endangered Species (CITES) . As the 
species is Endangered and totally protected in its only country of occurrence, it is 
more appropriate to include it on Appendix I if practical problems, such as its 
identification by custom officers, can be resolved (19). This change should not be 
formally proposed until consideration has been given to the partial financing of a 
practical conservation programme by the sale of confiscated specimens, mentioned 
above. 

References 

1. Anon. (1976). Appraisal of the Popondetta Smallholder Oil Palm Development. Report 
No. 1160, 25 September. Department of Primary Industry, Papua New Guinea. 

2. Conservation Areas Act (1978). No. 52 Independent State of Papua New Guinea. 12 
September. 

3. D'Abrera, B. (1971). Butterflies of the Australian Region. Lansdowne Press, Melbourne. 
415 pp 

4. D'Abrera, B. (1975). Birdwing Butterflies of the World. Lansdowne Press, Melbourne. 
260 pp. 

5. Fauna Protection Ordinance (1966). No. 19 Independent State of Papua New Guinea. 

6. Fenner, T.L. (1975). Proposal for experimental farming of protected birdwing butterflies 
with particular reference to Ornithoptera alexandrae . Unpublished manuscript. Depart- 
ment of Primary Industry. 5 pp. 

7. Fenner, T.L. (1983). In litt., 15 March. 

8. Haatjens, H.A. (ed.) (1964). General report on the lands of the Buna-Kokoda Area, 
Territory of Papua and New Guinea. C.S.I.R.O. Land Resources Series No. 10, 113 pp. 

9. Haugum, J. and Low, A.M. (1978). A Monograph of the Birdwing Butterflies. Vol. 1 (1): 
1-84. Scandinavian Science Press, Klampenborg. 

10. Hutton, A.F. (1982). In litt., 20 June. 

11. Hutton, A.F. (1984). In litt., 1 February. 

12. Igarashi, S. (1979). Papilionidae and Their Early Stages. Vol. 1: 219 pp.. Vol. 2: 102 pp of 
plates. Kodansha, Tokyo. (In Japanese). 

13. Meek, A.S. (1913). A Naturalist in Cannibal Land. T. Fisher Unwin, London. 238 pp. 

14. National Research Council (1983). Butterfly Farming in Papua New Guinea. Managing 
tropical animal resources series. National Academy Press, Washington D.C. 34 pp. 

15. Owen, D. (1971). Tropical Butterflies. Oxford University Press, Oxford. 214 pp. 

16. Parsons, M.J. (1980). A conservation study of Ornithoptera alexandrae Rothschild 
(Lepidoptera: Papilionidae). First report. Wildlife Division, Papua New Guinea. 89 pp. 

17. Parsons, M.J. (1980). A conservation study of Ornithoptera alexandrae Rothschild 
(Lepidoptera: Papilionidae). Second report. Wildlife Division, Papua New Guinea. 16 pp. 

18. Parsons, M.J. (1980). A conservation study of Ornithoptera alexandrae Rothschild 
(Lepidoptera: Papilionidae). Third report. Wildlife Division, Papua New Guinea. 15 pp. 

19. Parsons, M.J. (1983). A conservation study of the birdwing butterflies Ornithoptera and 
Troides (Lepidoptera: Papilionidae) in Papua New Guinea. Final report to Department of 
Primary Industry, Papua New Guinea. Ill pp. 

20. Pyle,R.M. and Hughes, S. A. (1978). Conservation and utilisation of the insect resources of 
Papua New Guinea. Report of a consultancy to the Wildlife Branch, Dept. of Nature 
Resources, Independent State of Papua New Guinea. 157 pp. 

21. Straatman, R. (1971). The life history of Ornithoptera alexandrae (Rothschild). Journal of 
the Lepidopterists' Society 25: 58-64. 

22. Straatman, R. (1979). Summary of survey on ecology of Ornithoptera alexandrae 
Rothschild. Consultancy report to the Department of Agriculture, Stock and Fisheries, 
July 1970. 5 pp. 

291 



Threatened Swallowtail Butterflies of the World 

Ornithoptera aesacus (Ney, 1903) INDETERMINATE 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Ornithoptera aesacus is a beautiful but little-known birdwing 
butterfly restricted to one small island. Obi, in the Moluccas (Maluku Province), 
Indonesia. Obi still has virgin forest on steep terrain around the highest peak in the 
centre of the island, but most of the lowland forest has been logged over, possibly a 
main cause of the apparent extreme rarity of the species. A national park has been 
proposed for Obi's central peak, an important conservation measure for this species. 
Surveys of the precise distribution of O. aesacus are needed in order to ascertain its 
ability to withstand habitat disturbance and its correct conservation status. 

Description The male of Ornithoptera aesacus is iridescent turquoise and 
black with some yellow and green on the underside (Plate 5.5). The larger female 
(wingspan 150 mm) is black with pale markings (1, 2, 5) (Plate 5.6). Despite early 
doubts about the status of O. aesacus as a good species, modern research has firmly 
established its specific status (4, 5, 6, 7). 

Male: UFW black with an iridescent light turquoise radial band and narrower anal 
and submarginal bands. The iridescent scaUng has a violet-blue reflection. UHW 
iridescent light turquoise-blue with a narrow black margin, a golden tinge distally and 
up to four small golden-yellow spots. 

LFW light blue becoming green towards the apex, with black wing margins, veins 
and markings. LHW iridescent turquoise-blue basally and in the cell, surrounded by 
green and yellow scaling. There is a narrow black margin and five large subdiscal 
spots and a golden-yellow anal patch, large subcostal spot and submarginal marks. 

Female: UFW black with prominent white markings including large subapical 
streaks, large discal and submarginal spots and a large cell spot. UHW with a 
scalloped outer margin and a large pale grey distal band with grey suffusion distally 
and small black discal spots. 

Despite earlier doubts about the status of O. aesacus as a good species, modern 
research has established its specific status without serious doubt (4, 5, 6, 7). 

Distribution Ornithoptera aesacus is known to occur only in the small island 
of Obi (or Ombira) in the Moluccas (Maluku Province) , Indonesia. Obi lies due south 
of Halmahera and due east of Sulawesi. The precise distribution of the butterfly 
within Obi is not known. 

Habitat and Ecology Virtually nothing is known about the early stages and 
life history of Ornithoptera aesacus, despite the fact that the original series of three 
specimens (one male, two females) was bred (by the collector John Waterstredt) 
from larvae collected in May 1902 (5). The foodplant is stated to be "Aristolochia 
species" (1), but it is not clear whether this is based on recent observations or on an 
assumption. Nothing has been published on the habitat and behaviour of O. aesacus; 
it remains one of the least known birdwings. 

Obi, the southernmost of the northern group of islands in the Moluccas is a 
mountainous island with broad coastal plains in the east and west, but steep coasts in 
the north and south. The native vegetation is seasonal monsoon forest and the island 
has remained forested with secondary formations, despite logging operations that 
have continued for many years (8). 

292 



Ornithoptera aesacus 

Threats Ornithoptera aesacus is listed as Indeterminate mainly because of 
the undoubted extensive land-use changes that have taken place in Obi. Although the 
island is said to be still forested with secondary or disturbed growth (8) there are 
insufficient data to be confident that O. aesacus, can adapt to these conditions. The 
extreme rarity of specimens mitigates against this. Apart from the primary threat of 
habitat destruction, there appears to be some more or less clandestine trade in this 
species (5), but this is unlikely to pose a significant threat. 

Conservation Measures No practical conservation measures for Ornithop- 
tera aesacus have yet been attempted. The first priority is an assessment of the 
distribution and precise conservation status of the species on Obi, with particular 
attention to its ability to survive forest disturbance. At present there are no protected 
areas on Obi, but there is a proposal to gazette a new nature reserve of 45 000 ha 
centred on Obi's main peak (8). The proposed reserve embraces an altitudinal range 
of 500-1616 m; the mainly steep terrain is unsuitable for logging and would require 
little protection (8). The reserve would protect the endemic dove PtiUnopus 
gramdifrons as well as O. aesacus and other fauna and flora unique to the island (8). 
The species should be retained, until its status can be reviewed, under Appendix II 
of the 1973 Convention on International Trade in Endangered Species of Wild Fauna 
and Flora (CITES). Appendix II listing implies that commercial trade is allowed 
providing a permit from the country of export is obtained. This can provide a method 
of monitoring trade levels. 

References 

1. D'Abrera, B. (1971). Butterflies of the Australian Region. Lansdowne Press, Melbourne. 
415 pp. 

2. D'Abrera, B. (1975). Birdwing Butterflies of the World. Lansdowne Press, Melbourne. 
260 pp. 

3. D'Abrera, B. (1983). In lilt., 12 March. 

4. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smiihersia 2: 1-48. 

5. Haugum, J. and Low, A.M. (1978-9). A Monograph of the Birdwing Butterflies. Vol. 1 
(1-3): 308 pp. Scandinavian Science Press, Klampenborg. 

6. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supp\ement. 17: 1-51. 

7. Smart, P. (1975). The Illustrated Encyclopedia of the Butterfly World. Hamlyn, London. 
275 pp. 

8. UNDP/FAO National Parks Development Project ( 1981). /I National Conservation Planfor 
Indonesia. Vol. 7: Maluku and Irian Jaya. FAO, Bogor. 



293 



Threatened Swallowtail Butterflies of the World 

Ornithoptera croesus Wallace, 1859 VULNERABLE 

Subfamily PAPILIONINAE Tribe TROIDINI 



Summary Ornithoptera croesus is a highly attractive birdwing butterfly 
with a unique golden-yellow coloration in the male. It is restricted, as two 
well-known subspecies and a little-known third, to a few islands in the Moluccas. It is 
at risk primarily because of deforestation but possibly also as a result of insecticidal 
spraying on a large scale. Conservation of O. croesus is inadequate and a thorough 
assessment of its status followed by protection of suitable habitat is needed. 

Description Ornithoptera croesus has a wingspan of 130-150 mm in the 
male and 160-190 mm in the female (4). The male is dark brown, orange, and golden 
with a green and black lower surface . The female of the nominate subspecies is brown 
with white markings (1, 2, 3, 4, 6). The subspecies O. c. /yd/u^, commonly seen in the 
butterfly trade, differs widely from the nominate subspecies described below and is 
illustrated in Plates 6. 1 and 6.2. The female is unique among the birdwings in being a 
mimic of unpalatable Danainae species. A third subspecies O. c. sananaensis is only 
known from a single female and is stated to be intermediate between the first two. 

Male: UFW ground colour very dark brown with a broad iridescent orange radial 
band and short anal streak (1, 2, 4, 8). UHW orange with a narrow black margin and 
a golden-yellow subcostal patch, discal and submarginal spots. 

LFW black with iridescent green submarginal and discai spots, radial band and a 
patch in the cell. LHW yellow-green with black veins, subdiscal spots and a narrow 
margin, a yellow anal area and golden areas as on the upper surface (1, 2, 3, 4, 6). 

Female : UFW dark brown ground colour with white markings including a cell spot, 
marginal fringe spots, submarginal and discal spots. UHW darker than forewing with 
yellow-brown distal patches and black subdiscal spots. LFW/LHW differs only in 
having paler markings (1, 2, 3, 4, 6). 

Distribution The nominate subspecies, Ornithoptera croesus croesus, has 
been found only on the island of Bacan (Bachan, Batjan) in the Moluccas (Maluku). 
It is said to be very localized but less rare in its chosen sites than has been previously 
thought (3) . O. c. lydius occurs in the neighbouring islands of Halmahera (also known 
as Jailolo Gilolo or Djailolo), Ternate, Tidore, and possibly Morotai (Morty) (3). 
Halmahera, by far the largest of these islands, is the most important locality, and is 
the source of most of the recent captures of this birdwing (9). 

The only known specimen of O. c. sananaensis was taken on Sanana, the most 
southerly of the Sula Islands (8) . Sanana lies about 240 km south-west of Bachan and 
this distribution requires confirmation. 

Habitat and Ecology Ornithoptera croesus is a lowland butterfly in Bachan, 
where it occurs in swamps and other wet places (4). Its habitat has been regarded as 
difficult of access since the time of Alfred Russel Wallace, the discoverer of the 
butterfly (11). The larva and pupa of O. c. croesus have been briefly described but the 
larvae oiO.c. lydius are stated to be unknown (3) , a surprising fact since many of the 
recently-collected specimens have been reared in captivity (9). Excellent figures of 
the larvae and pupae have been published, but of which subspecies is not stated (4). 
The foodplants are not comprehensively known but include Aristolochia gaudichau- 
dii (4, 8). 

294 



Ornithoptera croesus 

Threats In 1982 about 90 per cent of all forest in the northern and central 
Moluccas was under concession to large-scale commercial logging operations (7). In 
1980 the entire production of 1 .4 million cu. m of logs were exported and government 
control is reported to be slight (7). As a result, deforestation is being carried out in an 
irresponsible and unsustainable manner often on steep, easily eroded, slopes and 
with no reforestation — all in defiance of government regulations ( 7) . Indonesia is one 
of the world's biggest timber producers but such exploitation without regard to 
conservation can only be a very short-term development strategy. Ornithoptera 
croesus is only one of many Moluccan endemics that are threatened by the 
devastating rate and nature of deforestation in the region (8). 

Both O. c. croesus and O. c. lydius live in highly productive lowland forest, the 
most valuable timber concessions and the first areas to be deforested. There can be 
little doubt that O. croesus is declining in numbers and seriously at risk from further 
deforestation throughout its range. A further threat, the impact of which has not been 
assessed, is the reported use of large-scale insecticidal spraying in the swampy 
lowlands of Bachan (3), presumably as a mosquito control measure. 

O. croesus has always been a much sought-after species (4, 9). From about 1979, 
large numbers of O. c. /vt/n« appeared in trade. Prices were originally U.S. $90 a pair 
or more but by mid- 1982 had fallen to $24 a pair in the U.S. A. , perhaps indicating that 
the market had been satisfied (9). There have been no reports that O. c. lydius has 
been threatened by this increase in trade, but consideration might be given to the 
possibility of setting up a ranching programme on Halmahera. 

Conservation Measures The Halmahera group of islands, including Moro- 
tai, Bacan and Obi, has by far the greatest number of endemic species, the widest 
range of land form types and the most varied climate in the whole of the Moluccas 
(10). Sadly, there is not a single reserve established or even approved on any of these 
islands. Seven reserves have been proposed in the National Conservation Plan for 
Indonesia, including one on Obi, one on Morotai, one on Bacan and four on 
Halmahera (10). There is no information on the likelihood of Ornithoptera croesus 
habitat being found within these proposed reserves and a survey is an essential 
preliminary step towards more specific conservation measures. 

The species should be retained, until its status can be reviewed, under Appendix II 
of the 1973 Convention on International Trade in Endangered Species of Wild Fauna 
and Flora (CITES). Appendix II listing implies that commercial trade is allowed 
providing a permit from the country of export is obtained . This can provide a method 
of monitoring trade levels. 

References 

1. D'Abrera, B. (1971). Butterflies of the Australian Region. Lansdowne Press, Melbourne. 
415 pp. 

2. D'Abrera, B. (1975). Birdwing Butterflies of the World. Lansdowne Press, Melbourne. 
260 pp. 

3. Haugum, J. and Low, A.M. (1978-9). A Monograph of the Birdwing Butterflies. VoL 1 
(1-3): 308 pp. Scandinavian Science Press, Klampenborg. 

4. Igarashi, S. (1979). Papilionidae and Their Early Stages . Vol. 1: 219pp., Vol. 2: 102 pp. of 
plates. Kodansha, Tokyo. (In Japanese). 

5. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
£«towo/og(5r Supplement 17: 1-51. 

6. Smart, P. (1975). The Illustrated Encyclopedia of the Butterfly World. Hamlyn, London. 275 
pp. 

295 



Threatened Swallowtail Butterflies of the World 

7. Smiet, F. (1982). Threats to the Spice Islands. Oryx 16(4): 323-8. 

8. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands. Vol. 1 
Papilionidae . (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 

9. Various trade catalogues (1979-82). 

10. UNDP/FAO National Park Development Project ( 1981). National Conservation Plan for 
Indonesia. Vol. 7. Maluku and Irian Jaya. FAO, Bogor. 

11. Wallace, A.R. (1869). The Malay Archipelago. MacMillan, London. (Dover reprint 
edition, 1962, pp. 257-8). 



296 



Papilio homerus 
Papilio (Pterourus) homerus Fabricius, 1793 ENDANGERED 

Subfamily PAPILIONINAE Tribe PAPILIONINI 



Summary Papilio homerus, the Homerus Swallowtail, is a superb 
butterfly, rivalling in size some of the birdwings (Troidini) of South East Asia. This 
and the fact that it is uncommon and restricted to the island of Jamaica have made it 
particularly prized by collectors. However, the main threat seems to be destruction of 
habitat. The once continuous population is already divided into two isolated pockets 
and conservation measures are urgently needed. 

Description Papilio homerus is one of the largest species in the genus 
(comparable with Papilio antimachus) and the largest American swallowtail, with a 
forewing length of about 75 mm (Plate 6.3). It is easily recognizable by the large size 
and generally slow flight (8). The sexes are alike (1-6). 

UFW black or very dark brown with a broad yellow discal band extending across 
the wing, including a yellow bar across the cell, two subapical spots and two or three 
submarginal spots. 

UHW broad, elongated, with a relatively long spatulate tail. The wing is black or 
dark brown with a broad yellow discal band, powdery blue postdiscal spots and 
brick-red submarginal lunules. 

LFW/LHW similar to the upper surface with a dark brown ground colour. The 
hindwing has a much narrower yellow-brown discal band which is dusted with blue 
scales, and there are dark marks between the red and the blue spots (1-6). 

The eggs are spherical and light green in colour (8). The first three larval instars are 
essentially black and white; the last two instars are dark brown with extensive saddles 
of green and a pair of conspicuous lateral eyespots on the thorax (8). The pupa is 
shades of brown with eight small white, dorsal spots; the shape is blunt with only slight 
dorso-ventral flattening (8). 

Distribution Papilio homerus is confined to the Caribbean island of 
Jamaica, where it has been recorded from 7 of the 13 parishes (1). It is now restricted 
to two strongholds, the eastern population in St Thomas and Portland (where the 
Blue Mountains meet the John Crow range) and the western population in the 
'Cockpit Country' of Trelawny Parish (1, 4, 6). Each locality is only a few square 
kilometres of forest (8). A third population in central Jamaica probably became 
extinct around the turn of the century (8). There is no record on any other Caribbean 
island, despite suggestions to the contrary (2). The nearest relatives are apparently P. 
garamus and P. abderus in Mexico. 

Habitat and Ecology P. homerus is restricted to virgin forest on mountain 
slopes and in gullies at fairly low elevations (150-600 m, occasionally higher) (8). 
Flight is slow but powerful, and occurs throughout the day (0900-1800 hrs). The 
adults bask on high trees and bushes, visiting flowers daily, but only for short periods 
of time. Preferred nectar sources include species of Blechum, Bidens, Asclepias, 
Lantana, amalvaceousplant, and possibly a local /pomoea (Morning Glory) (7). The 
larval foodplants are two endemic Hernandia species H. catalpaefolia (eastern 
population) and H. troyiana (western population) (7). Oviposition has also been 
observed on an endemic species of the Camphorwood tree (Ocotea sp.) in both 
locations (7). Thespesia populnea (Seaside Mahoe) has been recorded as a foodplant 

297 



Threatened Swallowtail Butterflies of the World 

(4) but does not occur in the known distribution of the insect (7). It may have been 
confused with Hernandia which is locally known as Water Mahoe (7). SeasonaHty is 
not marked, but the adults fly during February to April, September and October (4). 
Little information is available on populations. P. homenis is said to be quite 
common in a few favoured localities, but the total population is certainly small. The 
population range is rapidly shrinking and the two populations are probably 
genetically isolated. Predation of larval instars by birds is high but the adult 
populations seem to be free of the marked fluctuations in numbers characteristic of 
some other Caribbean swallowtails. 

Threats Records for the past 20 years together with former and present 
collecting or confirmed visual records clearly establish that both populations are 
declining (8). The main reasons for this are the destruction and alteration of habitat, 
first for timber extraction and then for coffee gardens and pine plantations. The rate 
of habitat destruction in the east has slowed since 1952, but that population is still 
threatened and on the decline (7,8). The western population, although living in more 
mountainous country, is also suffering a reduction in range , and thorough surveys are 
needed. Collecting is difficult in such mountainous country, but commercial 
collecting by expatriates may be a minor threat since prices are high. The extent of 
collecting by Jamaicans is unknown. 

Conservation Measures There have been no conservation measures speci- 
fically for this species. A detailed survey of the distribution of the species and its 
ecological requirements is needed, in order that consideration may be given to 
conservation. Some form of habitat protection seems essential, probably in the form 
of a patrolled nature reserve or national park (8). The Natural Resources Conser- 
vation Department of Jamaica should take some responsiblity for the protection of 
this beautiful butterfly, and should take steps to ensure the permanent protection of 
its breeding grounds. P. homerus is of great aesthetic value and a rational farming 
programme would help conservation efforts, and be an appeaHng commercial 
venture. The species has been reared with some difficulty in captivity (7) and further 
research is needed. Commercial farming would lessen pressure on wild populations, 
and funds should be made available to provide suitable facilities (7,8). Collecting of 
wild specimens should be limited to those used for scientific purposes; permits should 
be issued only by the Natural History Division of the Institute of Jamaica (8). 

References 

1. Brown, F.M. and Heinemann, B. (1972). Jamaica and its Butterflies, E.W. Classey Ltd., 
Faringdon, U.K. 478 pp. 

2. D" Abrera, B . ( 1981 ) . Butterflies of the Neotropical Region Pari. 1, Papilionidae and Pieridae. 
Landsdowne Editions, Melbourne, xvi + 172 pp. 

3. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supplement 17, 51 pp. 

4. Riley, N.D. (1975). A Field Guide to the Butterflies of the West Indies. Collins, London. 

5. Rothschild, W. and Jordan, K. (1906). A revision of the American Papilios. Novitates 
Zoologicae 13: 411-752. 

6. Smart, P. (1975). The Illustrated Encyclopedia of the Butterfly World. Hamlyn, London. 
275 pp. 

7. Turner, T.W. (1982). In litt., 12 August. 

8. Turner, T.W. ( 1983). The status of the Papilionidae, Lepidoptera of Jamaica with evidence 
to support the need for conservation of Papilio (Pterourus) tiomerus Fabricius and Eury tides 
marcelUnus Douh\e:ddy . Unpublished report, 14 pp. 

298 



Papilio esperanza 
Papilio (Heradides) esperanza Beutelspacher, 1975 VULNERABLE 

Subfamily PAPILIONINAE Tribe PAPILIONINI 



Summary Little information is available on this extremely rare and 
recently described species from Mexico. The breeding locations are kept secret as a 
protection from commercial collectors, but a conservation plan is needed to ensure 
long-term protection and survival of the species. 

Description Papilio esperanza is an attractive, medium-sized black, yellow 
and orange swallowtail (Plate 6.4). The female, known only from one specimen, has 
a similar pattern to the male, but is slightly larger (7). The male forewing length is 
51-52 mm ( 1 ) whereas in the female it is 65 mm (7). The affinities of the species are in 
dispute. It was originally placed in the glaucus group (1,9), but was transferred to the 
thoas group in the recent revision by Hancock (3), who found the male genitalia to be 
very similar to those of P. androgens (4). However, a number of authorities familiar 
with the insect in the wild consider the nearest relative of P. esperanza to be P. 
palamedes, in the troiliis species group, thus placing it in the subgenus Pterourus 
(6,8). 

UFW black with a broad yellow median band and a row of small, yellow 
submarginal spots. 

UHW with a slightly scalloped outer margin with yellow fringe spots and a long 
tail. The broad UFW median band extends across the UHW, tapering sHghtly to the 
inner margin. There is an orange anal lunule, a further series of narrow, yellow 
submarginal lunules, and postdiscal groups of blue scales. 

LFW differs from the upper surface in having broad yellow lines in the cell, an 
orange submarginal region and orange scaling over the wing apex. 

LHW predominantly orange with yellow-bordered black veins, a concave black 
subbasal-abdominal line, and an irregular black postdiscal Hne with silvery-white 
scales over the middle of it ( 1 , 7, 2) . 

Distribution Papilio esperanza is only known from 1700 m above sea level 
at La Esperanza in the Sierra de Juarez, Oaxaca State, Mexico (1,7). 

Habitat and Ecology Papilio esperanza flies in montane cloud forest with 
an annual rainfall of over 4000 mm (6). The dominant tree species in the forest is 
Engelhardtia (Oreomunnea) mexicana (Juglandaceae), a relict species of the Ceno- 
zoic era (10). The adult butterflies are only active on sunny days from 11.00-15.00 
hours (6) and take nectar from the purple, aromatic flowers oi Eupatorium sordidum 
var. atrorubens, a herb that grows along streams and gullies (5). Two broods are 
produced each year, one in March and the other in August (6, 7). Adults from the 
two broods show slight differences in the depth of colour and width of the yellow 
wing-bands (7). The foodplant of the caterpillars is unknown. Associated species 
include Eurytides calliste, Papilio abderus, Anetia thirza, Paramacera chinanteca. 
Piscina zelys and Polygonia g-argenteum (6). In nine years only 50 specimens have 
been collected, and the total population is believed to be declining (6). 

Threats The distribution of the species is very limited. The precise 
breeding locations are kept secret in order to protect it from commercial collectors 
(6,11). Threats to the habitat are unknown, but the cloud forest is restricted in extent 
and vulnerable to human interference (6). 

299 



Threatened Swallowtail Butterflies of the World 

Conservation Measures The Sociedad Mexicana de Lepidopterologia has 
kept vigilant watch over the type locaHty of this exciting new discovery since 1975, 
preventing any commercial collecting (6). The Mexican Government plans to 
estabUsh a reserve in the Sierra de Juarez, Oaxaca, which will include the habitat of P. 
esperanza (6). National and international conservation bodies are encouraged to 
assist in this worthy proposal 

References 

1. Beutelspacher, C.R. (1975). Una especie nueva de Papilio L. (Papilionidae) Revista 
Sociedad Mexicana de Lepidopterologia 1(1): 3-6. 

2. D'Abrera, B. (1981). Butterflies of the Neotropical Region. Part I. Papilionidae and 
Pieridae. Lansdowne Editions, Melbourne, xvi + 172 pp. 

3. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

4. Hancock, D.L. (1984). In litt., 15 February. 

5. Maza E., J. de la (1984). In litt.,22 March. 

6. Maza, E., J. de la (1983). In litt.,4 October. 

7. Maza, E., J. de la, and Diaz Frances, A. (1979). Notas y descripciones sobre la familia 
Papilionidae en Mexico. Revista de la Sociedad Mexicana de Lepidopterologia 4(2): 51-56. 

8. Miller, L.D. (1984). In litt..6 January 

9. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supp\ement 17: 1-51. 

10. Rzedowski and Palacios (1977). El bosque de Engelhardtia (Oreomunnea) mexicana, en la 
region de la Chinantla (Oaxaca, Mexico), una reliquia del Cenozoico. Boletin de la 
Sociedad Botdnica de Mexico 36: 93-1 19. 

11. Tyler, H.A. (1983). In litt., 13 March. 



300 



Papilio aristodemus ponceanus 
Papilio (Heradides) aristodemus ponceanus Schaus, 191 1 ENDANGERED 

Subfamily PAPILIONINAE Tribe PAPILIONINI 



Summary Papilio aristodemus ponceanus , usually known as Schaus' Swall- 
owtail, was formerly locally common in Dade and Monroe Counties, Florida, 
U.S.A., but is now restricted to North Key Largo, Elliott Key and several smaller 
keys between them. The Key Largo population is severely threatened by develop- 
ment and insecticide spraying, despite being federally protected in the U.S.A. since 
1976. The Elliott Key population is protected by the Biscayne National Park, but 
suffers dangerously wide fluctuations in numbers. 

Description An attractive medium-sized, tailed swallowtail with a 
wingspan of 86-95 mm (Plate 6.5). The sexes are similar in general appearance, the 
female slightly larger than the male (10). 

UFW/UHW with brown ground colour, a bold, median yellow band, yellow 
submarginal spots, long spatulate tails and a red and blue anal spot (16). 

LFW/LHW duller with a wide orange-brown band on the LHW followed by a 
distinctive postdiscal row of blue lunules (16). 

The subspecies ponceanus is quite different from the nominate subspecies, being 
distinguished by the reduction in width of the dorsal oblique yellow band and more 
extensive yellow beneath. The cryptic larvae are brown mottled with tan, white and 
yellow, resembling bird droppings. 

Distribution This subspecies is only recorded from Florida, U.S.A. It was 
formerly locally common in areas of Dade County (including Miami) and Monroe 
County, including Elliott, Sands, Largo, Old Rhodes, Totten, Porgy, Adams, Upper 
and Lower Matecumbe and possibly Lignumvitae Keys (23). It has been destroyed in 
most of its range and is now restricted to localized colonies on North Key Largo in the 
south, Elliott Key to the north, and probably Old Rhodes, Totten, Adams, and Porgy 
Keys in between (21). Other subspecies are found on Cuba and the Cayman Islands 
(P. a. temenes), Hispaniola {P. a. aristodemus), the Bahamas (P. a. bjorndalae) and 
possibly Puerto Rico (7, 25)). 

The first colony of P. a. ponceanus was found in the Brickell Hammock of Miami 
(19), but was apparently destroyed by the city's growth (16). It was subsequently 
rediscovered in the Keys, but after a hurricane hit Lower Matecumbe in September 
1935 P. a. ponceanus was feared extinct (11). Surveys after World War II showed 
that this was not so (13, 14, 15) but the most recent report of distribution only lists 
breeding populations on North Key Largo and the larger keys of the Biscayne 
National Park, such as Elliott Key (10, 21). 

Habitat and Ecology Only tropical hardwood hammocks (patches of forest) 
containing the host plant Torchwood {Amyris elemifera) will support populations of 
Schaus' Swallowtail, although another rutaceous plant, the Wild Lime (Zanthoxylum 
fagara) has been observed as an oviposition site (19, 21). Torchwood and Wild Lime 
are pioneering shrubs or small trees in whose shade sprout the hardwood seedlings 
which eventually form the hammock. They are therefore abundant at the edge of 
hammocks, but scarcer within the understorey of the mature trees (18). Schaus' 
Swallowtail probably continually colonizes regrowth areas partially destroyed by 
storm or fire ( 18) . Females oviposit single green eggs on the underside of young leaves 

301 



Threatened Swallowtail Butterflies of the World 

at the tips of branches (18). Caterpillars hatch in four to seven days and moult four 
times at intervals of approximately 12 days. The final instar fastens itself vertically to 
a twig with silk and moults into a rusty brown or grey pupa (5). Emergence is 
normally slightly less than one year later, but some pupae in captivity have remained 
in diapause for two years (5, 18). Annual population fluctuations suggest that this 
behaviour occurs in the wild and may be an adaptation to avoid unfavourable 
conditions while making best use of good rainfall years (21). Reproduction is 
correlated with the beginning of the rainy season (April to June) which , perhaps with 
light intensify or day length, seems to trigger a synchronous emergence of adults. 
There is some evidence to suggest a partial second brood in some years (7), for 
example, specimens were discovered during September in 1969 (2). Adult Ufe span 
does not seem to exceed one month, but during that time the adults are quite capable 
of flying across open water to adj acent islands (5 ) . Adults take nectar from blossoms 
of Guava (Psidium guajava) and Wild Tamarind {Lysiloma latisiliqua) in the 
hammocks, or Cheese Scrub (Morinda roioc) on their edge (18). 

The Schaus' Swallowtail's normal population size seems to be small at all stages, 
although numbers may follow a cyclical pattern (17). Large numbers of adults (up to 
100 per day) were recorded in the keys of the Biscayne National Park in 1972 
(3, 4, 6, 9) but populations were small during surveys there from 1973 to 1981 (6, 17). 
However, a survey in May 1982 on Elliott Key recorded about 15 individuals, a cause 
for guarded optimism (8). Possible reasons for the population decline are given 
below. Recorded numbers on Key Largo have always been low (9), or perhaps 
locally common (20) and most authorities consider that population to be doomed to 
extinction. 

Threats The Schaus' Swallowtail is a rare example of an essentially tropical 
butterfly resident in a peripheral habitat in the only suitable area of the U.S.A. 
However, this is only one example of a number of less conspicuous tropical 
invertebrate species threatened in Florida by environmental degradation (12). From 
the 1940s until today the range of Schaus' Swallowtail has been progressively and 
irrevocably eroded by the destruction of hardwood hammocks by private develop- 
ment and the leisure industry. Schaus' Swallowtail is no longer present on Upper or 
Lower Matecumbe (21), and it is nearly lost from North Key Largo. In the latter 
locality prime habitat has been subjected to fires, development, and aerial spraying 
against mosquitoes (6). Despite its protection under federal law (23, 26, 27), the 
extinction of Schaus' Swallowtail on North Key Largo seems inevitable. Incidental 
injury to, or destruction of, deposited eggs, larvae or pupae of Schaus' Swallowtail 
was not an offence when the species was classified as Threatened(23), but the recent 
reclassification to Endangered status (26, 27) provides stricter protection (see 
below). Such incidental injuries are inevitable on building sites and the law has so far 
done little to protect the species' range. The extent of threat to Elliott Key, and to 
other smaller keys to the north of Key Largo, is presumably lessened by their 
inclusion in the Biscayne National Park. However, restriction to such a small range 
would inevitably increase the threat of extinction from natural disasters such as 
hurricanes, frost or disease (22). Natural disasters such as the 1935 hurricane, which 
struck only four months after the species was rediscovered on Lower Matecumbe 
Key, have threatened the swallowtail populations in the past (6). Successive 
droughts since the early 1970s, combined with hard winter conditions in 1977/78, 
have also been detrimental (21 ) . Throughout its chequered history Schaus' Swallow- 
tail has suffered at the hands of collectors (16). Such a rare species fetches high prices 
(16). 

302 



Papilio aristodemus ponceanus 

Conservation Measures On 17 April 1975 the U.S. Fish and Wildlife 
Service published a proposed ruling of Threatened status for Schaus' Swallowtail 
(22). It was recognised that the range of the butterfly was greatly depleted, but its 
status was limited to Threatened because the current protection of the population on 
Elliott and other keys in the Biscayne National Park was considered substantial (22). 
In the Final Rulemaking of 28 April 1976, the eggs and immature stages were 
protected from collectors, although not from incidental or inadvertent damage, and 
collection of adults was permitted outside protected areas (23). However, this 
legislation was reinforced by Florida state law, which banned collection of adults or 
immature stages of any federally listed species, except by special permit (1, 26). In 
1978 an authority considered the Biscayne National Park large enough to maintain 
Schaus' Swallowtail, unless natural catastrophe critically reduced the population (7). 
The staff of the National Park are aware of the butterfly, and co-operative in its 
conservation (7). Nevertheless, since the Final Rulemaking in 1976, the populations 
there have declined inexplicably. Recently, the Fish and Wildlife Service proposed a 
new Crocodile Lake National Wildlife Refuge on Key Largo (25). The report for the 
proposal notes the presence of Schaus' Swallowtail on North Key Largo, but it is not 
clear whether populations have been located within the proposed Refuge (25). 
Schaus" Swallowtail is classified as Endangered in the most recent assessment by 
authorities in Florida (12, 21) and a recovery plan by the Florida Game and Fresh 
Water Fish Commission recommended that Schaus' Swallowtail be federally listed as 
Endangered (25) . In a notice of proposed reclassification in 1983 the status of Schaus' 
Swallowtail was to be raised from Threatened to Endangered, giving it full protection 
(26). This proposal was accepted in 1984 (27) and it is to be hoped that the other 
recommendations of the recovery plan will be adopted as expeditiously as possible. 
The recovery plan for Schaus' Swallowtail gives details of the objectives which 
must be achieved to protect the butterfly from extinction (25). The priorities include 
protection of extant colonies and re-establishment of colonies within the species' 
historic range. The Biscayne National Park and Crocodile Lake National Wildlife 
Refuge will partially fulfil these aims, but should be expanded to include various 
important hammocks (25). Increased public awareness could result in voluntary 
conservation by informed developers and landowners in North Key Largo. Local 
authorities should inform planners of the butterfly's vulnerability. Scientific research 
should be encouraged, to facilitate large-scale artificial rearing and re-introductions 
(25). Suitable sites include Lignumvitae Key, a preserve of the Nature Conservancy 
(8), or the proposed Crocodile Lake Refuge on Key Largo (24). There is a great need 
for detailed scientific study of the population dynamics of Schaus' Swallowtail. 
Although habitat destruction and excessive collecting will harm the species, very 
little is known of the effects of parasites on young stages, or of fire and hurricane on 
maintenance of suitable habitat. Although only given subspecific rank, Schaus' 
Swallowtail is widely separated spatially from its conspecifics and natural reintroduc- 
tion from conspecific stock is an unlikely event. 

References 

1. Baggett,H.D. (1982). Schaus' Swallowtail. In Franz, R. (Ed.), The Rare and Endangered 
Biota of Florida. Vol. 6, Invertebrates. University Presses of Florida, Gainesville. Pp. 

73-74. 

2. Baggett,H.D.(1982).Statementfrom39-27.02oftheRulesoftheFloridaGanieandFresh 

Water Fish Commission. In litt. , 24 October. 

3. Brown, C.H, (1976) A colony of Papilio aristodemus ponceanus (Lepidoptera: Papilioni- 
dae) in the upper Florida Keys. Journal of the Georgia Entomological Society 11:117-118. 

303 



Threatened Swallowtail Butterflies of the World 

4. Brown, L.N. (1973). Populations of a new swallowtail butterfly found in the Florida Keys. 
Florida Naturalist April 1973: 25. 

5. Brown, L.N. (1973). Populations of Papilio andraemon bonhotei Sharpe and Papilio 
aristodemus ponceanus Schaus (Papilionidae) in Biscayne National Monument, Florida. 
Journal of the Lepidopterists' Society 27: 136-140. 

6. Brown, L.N. (1974). Haven for rare butterflies. National Parks and Conservation Magazine 
July 1974: 10-13. 

7. Covell, C.V., Jr. (1976). The Schaus Swallowtail: a threatened subspecies? Insect World 
Digest 3: 21-26. 

8. Covell, C.V., Jr. (1978). Project Ponceanus and the status of the Schaus Swallowtail 
(Papilio aristodemus ponceanus) in the Florida Keys. Atala 5 (1977): 4-6. 

9. Covell, C.V., Jr. (1982). In liti., 18 May. 

10. Covell, C.V., Jr. and Rawson,G.W. (1973). Project Ponceanus: a report on first efforts to 
survey and preserve the Schaus Swallowtail (Papilionidae) in southern Florida. Journal of 
the Lepidopterists' Society 27: 206-210. 

11. D'Abrera, B. (1981). Butterflies of the Neotropical Region. Part I. Papilionidae and 
Pieridae. Lansdowne Editions, Melbourne, xvi + 172 pp. 

12. Franz, R. (Ed.), (1982). The Rare and Endangered Biota of Florida, Vol. 6, Invertebrates. 
University Presses of Florida, Gainesville. 131 pp. 

13. Grimshawe,F.M. (1940). Place of sorrow: the world's rarest butterfly and Matecumbe Key. 
Nature Magazine 33: 565-567, 611. 

14. Henderson, W.F. (1945). Papilio aristodemus ponceana Schaus (Lepidoptera: Papilioni- 
dae) Entomological News 56: 29-32. 

15. Henderson, W.F. (1945). Additional notes on Papilio aristodemus ponceana Schaus 
(Lepidoptera: Papilionidae). Entomological News 56: 187-188. 

16. Henderson, W.F. (1946). Papilio aristodemus ponceana Schaus (Lepidoptera: Papilioni- 
dae). Entomological News 51: 100-101. 

17. Klots, A.B. (1951). Field Guide to the Butterflies. Houghton Mifflin, Boston. 349 pp. 

18. Loftus, W.F. and Kushlan, J. A. (1982). The status of Schaus' Swallowtail and the Bahama 
Swallowtail butterflies in Biscayne National Park. Report M-649, National Park Service, 
Everglades N.P., Homestead, Florida. 

19. Rutkowski, F. (1971). Observations on Papilio aristodemus ponceanus (Papilionidae). 
Journal of the Lepidopterists' Society 25: 126-136. 

20. Schaus, W. (1911). A new Papilio from Florida, and one from Mexico (Lepid.) 
Entomological News 22: 438-439. 

21. Spencer Smith, D. (1983). In litt., 25 February. 

22. U.S.D.L Fish and Wildlife Service (1975). Proposed threatened status for two species of 
butterflies. Federal Register 40{1S): 17757. 

23 . U.S.D.L Fish and Wildlife Service ( 1976) . Determination that two species of butterflies are 
Threatened species and two species of mammals are Endangered species. Federal Register 
41(83): 17736-17740. 

24. U.S.D.L Fish and Wildlife Service (1980). Ascertainment report: Crocodile Lake National 
Wildlife Refuge, Monroe County, Florida. 25 pp. 

25. U.S.D.L Fish and Wildlife Service. (1982). Schaus' Swallowtail butterfly recovery plan. 
U.S.D.L Fish and Wildlife Service, Atlanta, Georgia. 57pp. 

26. U.S.D.L Fish and Wildlife Service (1983). Proposed delisting of Bahama Swallowtail 
butterfly and reclassification of Schaus Swallowtail butterfly from Threatened to Endan- 
gered. Federal Register 48(168): 39096-7. 

27. U.S.D.L Fish and Wildlife Service (1983). Endangered and Threatened wildlife and plants; 
final rule to deregulate the Bahama Swallowtail Butterfly and to reclassify the Schaus 
Swallowtail Butterfly from Threatened to Endangered. Federal Register 49(171): 34501-4. 



This review has been adapted from an entry in The lUCN Invertebrate Red Data 
Book, whose authors and contributors are gratefully acknowledged. 

304 



Papilio caiguanabus 
Papilio (Heraclides) caiguanabus Poey , 1851 INDETERMINATE 

Subfamily PAPILIONINAE Tribe PAPILIONINI 



Summary Papilio caiguanabus , sometimes known as Poey's Black Swall- 
owtail, is endemic to Cuba, the largest island in the Caribbean. It is rare throughout 
its range but there is insufficient information to assess the extent of threats to the 
survival of the species. 

Description Papilio caiguanabus is a tailed butterfly whose sexes are similar 
but with slight colour differences (1, 6, 7). The forewing length is 48-50 mm (6). 

UFW with a submarginal row of equal-sized yellow spots, (pale yellow in the 
female) over the black ground colour. 

UHW markings include a postdiscal row of larger spots, deep yellow in the male, 
white in the female, and a red eye-spot. 

LFW/LHW similar to the upper surface but with two red spots below the hindwing 
cell and a row of small blue discal lunules (3, 5, 6)). 

P. caiguanabus is a member of the thoas species group and the mature caterpillar is 
black dorsally, dark brown along the side, with bold white markings (6). 

Distribution Papilio caiguanabus is endemic to the Caribbean island of 
Cuba, where it is found up to an altitude of about 300 m (2). It is fairly widespread, 
occurring in Matanzas and Oriente provinces (6), but is rare everywhere. Records 
and sightings have been more frequent in the eastern provinces (6). 

Habitat and Ecology The food plant has been reported as Securinega 
acidothannus (6), synonymized in 1919 with Securinega acidoton (L.) Fawcett & 
Rendle. This Caribbean species is known, at least in Jamaica, as Green Ebony. In the 
family Euphorbiaceae, S. acidoton may be a shrub of 2-3 m height or a tree of up to 
6 m. 

Feeding on Euphorbiaceae is very unusual for a species in Munroe's section IV (5) 
(Hancock's Heraclides species-group (3)), which generally use Rutaceae or Piper- 
aceae, rarely Umbelliferae (3). Further research is needed to confirm this observa- 
tion and to elucidate details of the habitat and breeding pattern of the butterfly. 

Threats There is insufficient information to assess the extent of threat to 
Papilio caiguanabus , but its rarity throughout Cuba is certainly a cause for concern. 

Conservation Measures There is little information available on the Cuban 
national parks legislation (4), and no information on the presence of Papilio 
caiguanabus within park boundaries. Cuba is the largest island in the Caribbean, with 
adequate area and resources for conservation programmes. Surveys of the distribu- 
tion of this species are needed, followed by biological studies and an assessment of the 
degree of threat. 

References 

1. D'Abrera,B. (1981). Butterflies of the Neotropical Region. Parti. Papilionidae and Pieridae . 
Lansdowne Editions, Melbourne, xvi -I- 172 pp. 

2. D'Almeida, R.F. (1966). Catalogo dos Papilionidae Americanos. Sociedade Brasileira de 
Entomologia. Sao Paulo, Brazil. 

305 



Threatened Swallowtail Butterflies of the World 

3. Hancock, D.L, (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

4. lUCN/CNPPA (1982). lUCN Directory of Neotropical Protected Areas. Tycooly, Dublin. 
436 pp. 

5. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
£n/owo/og/.sr Supplement 17: 1-51. 

6. Riley, N.D. ( 1975). A Field Guide to the Butterflies oftiie West Indies. Collins, London. 244 
pp., 24 col. pi. 

7. Rothschild, W. and Jordan, K. (1906). A revision of the American Papilios. Novitates 
Zoologicae 13(3): 411-752. (Facsimile edition ed. P.H. Arnaud, 1967). 



306 



Papilio aristor 
Papilio (Heraclides) aristor Godart, 1819 INDETERMINATE 

Subfamily PAPILIONINAE Tribe PAPILIONINI 



Summary Papilio aristor, the Scarce Haitian Swallowtail, was until 
recently known only from a few specimens collected near Port-au-Prince, Haiti, and 
Monte Christi, north-western Dominican Republic. Surveys are urgently needed in 
order to define its present distribution more precisely, particularly in the xeric areas 
which it is now known to inhabit. 

Description This black swallowtail is similar to P. caiguanabus from Cuba 
(see previous review) but with additional discal spots on the UFW and rather smaller 
postdiscal spots of colour on the UHW. The female is larger (forewing length 50 mm) 
than the male (forewing length 40 mm), and the yellow spots on both wings are 
somewhat paler, but the sexes are otherwise similar (1, 7, 8). 

UFW with a diagonal row of five small discal yellow spots from the costal margin, 
converging with a submarginal row of larger yellow spots. 

UHW tailed with a postdiscal row of large yellow spots, yellow marginal crescents 
and a red anal spot. 

LFW/LHW differs from the upper surface in having a diffuse yellow spot in the 
forewing cell, and paler postdiscal spots on the hindwing. The hindwing also has two 
red spots below the cell and a discal row of blue lunules. The immature stages are 
unknown (7) but, like Papilio machaonides , P. caiguanabus , and P. esperanza, P. 
aristor is a member of the thoas species group (3,6). This implies a mottled brown 
mature larva with pale yellowish or white patches, an ornate metathoracic band and a 
raised band on the first abdominal segment (3). 

Distribution This species is endemic to the Caribbean island of Hispaniola 
and is known from very few specimens. Most have their origin in the area around 
Port-au-Prince in Haiti (1, 2, 7, 8) but there is also a record from Monte Christi in 
north-western Dominican Republic (5). Recent studies in the Dominican Republic 
have discovered a number of new localities, but these have not yet been revealed (9). 

Habitat and Ecology Papilio aristor is known to inhabit xeric areas of 
lowland country in the Dominican Republic, and probably in Haiti also (9). Studies in 
recent years by A. Schwartz and his colleagues indicate that the species is not as rare 
as has been implied by its common name (7). However, only one female and few 
males have been recorded in the literature so far (7 , 8) . The foodplants are unknown , 
but as a member of the thoas species-group P. aristor is likely to feed on Rutaceae, 
Piperaceae or Umbelliferae (3). Nothing is known of the young stages or breeding 
cycle, but adults are known to be on the wing in July. Details of the habitat of P. 
aristor, although known to lepidopterists who visit the region, have not yet been 
published. 

Threats There are no data specifically addressing the threats to Papilio 
aristor. Nevertheless the threats to all forms of wildlife on Hispaniola, and 
particularly in Haiti, are well documented ( 10, and see review of Battus zetides) . Land 
development for agriculture, timber extraction and cash crop plantations by a large 
and rapidly expanding population has taken a heavy toll on wildlife. Degradation and 
erosion of soils through poor agricultural practices is exacerbating the problem, 

307 



Threatened Swallowtail Butterflies of the World 

causing a constant search for new land. P. aristor is said to inhabit xeric, lowland 
areas (9), possibly unsuitable for agricultural expansion. Nevertheless, natural 
habitats on Hispaniola are threatened to such a degree that Indeterminate category is 
retained for this species until more substantial data on its distribution and ecology are 
available. Xeric biomes are by no means immune from environmental damage; 
over-grazing or lowering of the water table may have a catastrophic effect. 

Conservation Measures An assessment of the present distribution of 
Papilio aristor is required, together with a careful study of its ecological require- 
ments. The only national park in Haiti is La Citadelle and this is poorly protected (4). 
The del Este and Isla Cabritos National Parks in the Dominican Republic contain 
subtropical dry forest characterized by mesquite, giant milkweed and several cactus 
species (4), and may possibly include suitable habitat for the butterfly. 

References 

1. D'Abrera, B. (1981). Butterflies of the Neotropical Region. Part I. Papilionidae and 
Pieridae. Lansdowne Editions, Melbourne, xvi + 172 pp. 

2. D'Almeida, R.F. (1966). Catdlogo dos Papilionidae Americanos. Sociedade Brasileira de 
Entomologia. Sao Paulo, Brazil. 

3. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

4. lUCN/CNPPA (1982). lUCN Directory of Neotropical Protected Areas. Tycooly, Dublin. 
436 pp. 

5. Miller, L.D. (1984). In to., 6 January. 

6. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
£«rowo/ogisr Supplement 17: 1-51. 

7. Riley, N.D. (1975). A Field Guide to the Butterflies of the West Indies. Collins, London. 244 
pp., 24 col. pi. 

8. Rothschild, W. and Jordan, K. (1906). A revision of the American Papilios. Novitates 
Zoologicae 13(3): 4n-752. (Facsimile edition ed. P.H. Arnaud, 1967). 

9. Schwartz, A. (1984). In litt., 16 May. 

10. Thornback, J. and Jenkins, M. (1982). The lUCN Mammal Red Data Book. Parti. lUCN, 
Gland. 516 pp. 



308 



Papilio garleppi 
Papilio (Heradides) garleppi Staudinger, 1892 INSUFFICIENTLY KNOWN 
Subfamily PAPILIONINAE Tribe PAPILIONINI 



Summary Papilio garleppi is a very rare species known from only a few sites 
in Peru, Brazil, Bolivia and Guyana. Nothing is known of its biology or the reasons 
for its rarity and disjunct distribution. 

Description Papilio garleppi is in the torquatus species-group (4) . The male 
has a forewing length of 45-50 mm. The female is not described in the literature. 

Male: UFW black-brown with a broad yellow postdiscal band which may or may 
not be interrupted by a black band from outer margin to the costal margin. 

UHW with yellow band broadening across the discal zone; red anal spot preceded 
by two or three yellowish-buff spots. 

LFW/LHW lighter in colour, with a row of rufous red discal spots , one small bluish 
dot and a row of yellow submarginal spots on the hindwing (2,5). 

Distribution Three subspecies have been described (2). P. g. garleppi is 
known from Mateo, Rio Juntas and Rio Chapare in Bolivia (3). P. g. interruptus 
occurs in eastern Peru as far south as the Rio and Cordillera de Carabaya, and in the 
Upper Amazon region of Brazil around Sao Paulo d'Olivenqa and Benjamin 
Constant, near the Peruvian and Colombian borders (3). The third subspecies, P. g. 
insidiosus, is recorded from the Maroni River in French Guiana, Surinam and 
Guyana (2,3). No explanation for this disjunct distribution has been published. 
However, it has been suggested by K.S. Brown Jr. that P. garleppi could possibly be a 
hybrid between P. torquatus and P. astyalus, two sympatric species (1). This would 
explain the disjunct distribution of P. garleppi and is worthy of further research. 

Habitat and Ecology Very few specimens of this species have ever been 
collected and no data on its biology or habitat have been published. In common with 
other members of the torquatus species-group, the caterpillars probably feed on 
Rutaceae (4) . The type specimen came from the Rio Chapare , which lies between 200 
and 500 m altitude in the eastern foothills of the Bolivian Andes. Similarly, 
specimens from western Brazil and eastern Peru seem to be from low-lying areas with 
an altitude of less than 100 m. One southern Peruvian locality however, Carabaya, is 
near Lake Titicaca at an altitudinal range of 3000-5000 m . It is unusual for a species to 
embrace such a diversity of habitats, but the range might be possible for an ancient 
species in this group (7). A mislabeUing of the specimen might also be suspected. 

Threats No specimens of Papilio garleppi are held in museums in Peru (6) 
and very few are held elsewhere, suggesting that the species is very rare. However, 
Zischka (8) claims that it is not rare in Chapare district, taken to be the region just to 
the north of Cochabamba and Santa Cruz. This observation certainly requires 
confirmation. Without details of its habitat or the precise limits of its distribution it is 
impossible to assess threats. 

Conservation Measures Details of the biology, habitat and distribution of 
Papilio garleppi are needed before any conservation measures can be proposed. 
There is no information on the occurrence of the species within the national parks and 
protected areas of Peru, Brazil, Bolivia or Guyana. 

309 



Threatened Swallowtail Butterflies of the World 

References 

1. Brown, K.S., Jr. (1984). In litt.,4 January. 

2. D'Abrera, B. (1981). Butterflies of the Neotropical Region. Fart I. Papilionidae and Pieridae. 
Lansdowne Editions, Melbourne, xvi + 172 pp. 

3. D'Almeida, R.F. (1966). Catdlogo dos Papilionidae Americanos . Sociedade Brasileira de 
Entomologia. Sao Paulo, Brazil. 

4. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
£«to/r2o/ogi5? Supplement 17: 1-51. 

5. Rothschild, W. and Jordan, K. (1906). A revision of the American Papilios. Novitates 
Zoologicae 13: 411-752. (Facsimile edition ed. P.H. Arnaud, 1967). 

6. Tyler, H. (1983). In litt., 13 March. 

7. Tyler, H. (1983). In litt., 30 September. 

8. Zischka, R. (1950). Catalogo de los insectos de Bolivia. Folia Univ. Cochabamba 4: 51-56. 



310 



Papilio himeros 
Papilio (Heraclides) himeros Hopffer, 1866 VULNERABLE 

Subfamily PAPILIONINAE Tribe PAPILIONINI 



Summary Papilio himeros occurs in the south-eastern states of Brazil and 
possibly Argentina, and is known to be disappearing from its former haunts. 
Although healthy populations are known, the localities are restricted. There is 
insufficient information on its distribution and biology to propose conservation 
measures. 

Description Papilio himeros is a member of the torquatus species-group 
(4, 7). It is a relatively large butterfly with a forewing length of 44-50 mm, the female 
being larger than the male (Plates 6.6 and 6.7). 

The wings of the male have a blackish-brown ground colour with a broad yellow 
median band from the forewing subapical region which continues to cover the entire 
discal and basal part of the hindwing. The hindwing also has a red and blue eye-spot 
and a row of yellow submarginal spots. The outer margin is scalloped and the long 
spatulate tail has a yellow spot at the apex (2, 8). 

The female differs in having a narrower median band which does not cover the base 
of the hindwing and submarginal lunules which are yellow near the apex but 
otherwise red (2, 4, 7, 8). 

Distribution Brazil and, according to D'Almeida, also from Corrientes in 
Argentina (3). There are two subspecies: P. h. baia from Bahia State, and P. h. 
himeros from Minas Gerais, Rio de Janeiro (3) and Espfrito Santo (1). This is the 
subspecies which is also reported from Corrientes in Argentina, although the locality 
is a long way south of the Brazilian sites (5). There is no published explanation for this 
apparently disjunct distribution. 

Habitat and Ecology Generally found in open areas at low altitudes, 
particularly in February, August and September (3). The female is very rare (3). The 
species flies quickly and with an intense, tremulous flight (3). Its habits are similar to 
the other members of the thoas, torquatus and anchisiades groups in Munroe's (7) 
section IV (called Heraclides by Hancock (4)). The larvae presumably feed on 
Rutaceae and, as in other members of the torquatus group, are probably tuberculate 
(7) and solitary (9). Adults seek nectar at flowers, but apparently have some highly 
specialized requirements which prohibit them from persisting in most areas (1). 

Threats The species has disappeared from the Cavalao area near Rio de 
Janeiro city, where many were captured earlier this century ( 1 ) . P. himeros persists in 
some numbers around Linhares in Espfrito Santo (1), but coastal development is 
probably gradually reducing the available habitat. 

Conservation Measures Although some localities are known, the full range 
of this species has not been properly determined. There is no information on its 
occurrence in Brazilian national parks, of which there are several in the south-eastern 
region (6). Studies of its biology are needed in order to assess its precise feeding 
requirements. Only then can proper conservation measures be put forward. No 
information on captive breeding is available. 

311 



Threatened Swallowtail Butterflies of the World 

References 

1. Brown, K.S., Jr. (1983). In litt.,6 April. 

2. D'Abrera,B. (1981). Butterflies of the Neotropical Region. Parti. Papilionidae and Pieridae . 
Lansdowne Editions, Melbourne, xvi + 172 pp. 

3. D'Almeida, R.F. (1966). Catdlogo dos Papilionidae Americanos. Sociedade Brasileira de 
Entomologia. Sao Paulo, Brazil. 

4. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

5. Hayward, K.J. (1967). Genera et Species Animalium Argentinorum. 4. Insecta. Lepidoptera 
(Rhopalocera). Familiae Papilionidarum et Satyridarum. Guillermo Kraft Lbla, S.G.; 
Bonariae (General editors Rossi, J.A.H. and Lillo, M.). 447 pp. 

6. lUCN (1982). lUCN Directory of Neotropical Protected Areas. Tycooly International, 
Dublin. 436 pp. 

7. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
£«rowo/og/i/ Supplement 17: 1-51. 

8. Rothschild, W. and Jordan, K. (1906). A revision of the American Papilios. Novitates 
Zoologicae 13: 411-752. (Facsimile edition ed. P.H. Arnaud, 1967). 

9. Tyler, H.A. (1975). The Swallowtail Butterflies of North America. Naturegraph Publishers, 
viii + 192 pp. 



312 



Papilio maroni 
Papilio (Heraclides) maroni Moreau, 1923 INSUFFICIENTLY KNOWN 

Subfamily PAPILIONINAE Tribe PAPILIONINI 



Summary Papilio maroni is only known from the region around St Laurent 
on the River Maroni in western French Guiana. More information is required on the 
taxonomic status and biology of this apparently very rare species. 

Description Papilio maroni is a relatively large butterfly with a forewing 
length of 52-55 mm. The sexes are similar though the female tends to be slightly 
larger (1, 4, 5). There is some doubt concerning the taxonomic status of this species 
(2). One authority considers it to be a form of Papilio anchisiades, a very variable 
species (9); others believe it may be a subspecies of P. chiansiades (7) or P. isidorus 
(8). 

UFW black with a white central-median spot which is smaller in the male and is 
positioned closer to the inner margin (1,4, 5). 

UHW black with a deeply scalloped outer margin and no distinct tail. There are 
three long, red discal-postdiscal spots forming a short band from the inner margin. 
These may be divided, by black suffusion, into separate discal and postdiscal spots, 
and in the female this band may be extended towards the costal margin by additional 
submarginal spots, and another discal spot (1,4, 5). 

Distribution Papilio maroni occurs only in western French Guiana. It has 
been recorded from the area around St Laurent, on the River Maroni, which 
constitutes French Guiana's western border (3). The species is known from very few 
specimens. 

Habitat and Ecology Papilio maroni is placed in the P. anchisiades group by 
Munroe (6) and Moreau (4). The larvae are probably tuberculate and gregarious 
when mature, feeding on Rutaceae (5). 

Threats These cannot be assessed without further biological and distribu- 
tional data. French Guiana still contains vast tracts of undisturbed forest and it would 
appear unlikely that forest destruction has contributed to the rarity of Papilio maroni. 

Conservation Measures Verification of the taxonomic status of this species 
is needed. As yet there are insufficient data on its habitat, biology and distribution to 
pursue conservation measures. The Reserve Naturelle de Basse Mana is on the 
north-west coast. It runs from above the lower Mana River as far as the river's mouth 
at Maroni (3) . The reserve is mainly coastal wetland , but it includes a variety of other 
habitats and could contain Papilio maroni. French Guiana is a French Overseas 
Department subject to French laws. Papilio maroni could be placed on the French list 
of protected species if it proved to be seriously threatened, but research and 
protection of habitat are more useful objectives at this early stage. 

References 

1. D'Abrera, B. (1981). Butterflies of the Neotropical Region. Parti. Papilionidae and Pieridae . 
Lansdowne Editions, Melbourne, xvi + 172 pp. 

2. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

313 



Threatened Swallowtail Butterflies of the World 

3. lUCN (1982). lUCN Directory of Neotropical Protected Areas. Tycooly International, 
Dublin. 436 pp. 

4. Moreau, E. (1923). Un Papilio nouveau de la Guyane Fran§aise. Bulletin de la Societe 
Entomologique de France 1923: 144, 215. 

5. Moreau, E. (1924). Description de Pap/V/owaroni male de Guyane Frangaise. Bulletin de la 
Societe Entomologique de France. 1924: 93-94. 

6. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supplement 17: 1-51. 

7. Racheli, T. (1984). In litl., 18 June. 

8. Smart, P. (1975). The Illustrated Encyclopedia of the Butterfly World. Hamlyn, London. 
275 pp. 

9. Tyler, H.A. (1983). In litt., 30 September. 



314 



Papilio maraho 
Papilio (Chilasa) maraho Shiraki & Sonan, 1934 VULNERABLE 

Subfamily PAPILIONINAE Tribe PAPILIONINI 



Summary Papilio maraho is a beautiful tailed butterfly endemic to Taiwan. 
Its habitat in the mountains is under severe pressure from growing human 
populations needing land for agriculture and forestry. Specimens are much in 
demand by collectors and sell for high prices. A combination of the destruction of 
critical habitats and excessive, intensive collecting gives cause for grave fears about 
the survival of the species. 

Description Papilio maraho is an attractive black, red and cream butterfly 
with slender forewings and deeply indented edges to the hindwings giving the 
impression of large spatulate tails. The sexes are similar with a forewing length of 
about 60 mm (1). The species is believed to be in a mimetic association with troidine 
swallowtails of the genus Atrophaneura, possibly A. polyeiictes ox A. aristolochiae , 
both of which are similar in general appearance. 

UFW brown-black, darker along the veins. 

UHW brown-black ground colour with a creamy-yellow cell and a row of bold, red 
submarginal lunules. 

P. maraho was originally described as a subspecies of the Chinese species 
P. elwesi, but in a recent reassessment it was afforded full species status (2). 

Distribution Papilio maraho is endemic to the island of Taiwan (Republic 
of China), lying about 145 km from mainland China and straddling the Tropic of 
Cancer. P. maraho is a replacement species for P. elwesi, a close relative that is found 
on the mainland and is very similar in appearance. 

Habitat and Ecology The island of Taiwan is 385 km long and 145 km wide , 
almost 36 000 sq. km in area (5). The eastern half is very mountainous, with over 60 
peaks exceeding 3000 m. The highest point is Yu Shan (Mt Morrison), 3950 m. The 
climate and vegetation of the island vary widely, from tropical monsoon forests to 
rocky snow-capped mountains (5). Although small in size, Taiwan has a diverse 
lepidopteran fauna, reflecting the diversity of habitat type. Shirozu (6) listed 361 
species of butterflies in 161 genera. About lOpercent of these species are endemic to 
Taiwan (3). 

Very little is known about the habitat and ecology of P. maraho except that it 
favours montane habitats. 

Threats Taiwan is an island with a large and growing population. Most of 
the 17 million or more people have traditionally concentrated in the lowlands, but 
increasing numbers have been moving to the mountains. As a result, the virgin 
hardwood and coniferous forests are now being cleared for agriculture, forestry and 
other developments, following the already almost total destruction of forests in the 
lowlands (5). The precise altitude preferred by P. maraho is unknown, but likely 
habitat in the hardwood forests below 2000-2300 m is now under severe pressure of 
replacement by conifer plantations (R.D. Schultz, in ref. 5). Destruction of critical 
habitat is probably the single most important factor in causing the decline of 
P. maraho and other butterflies in Taiwan today (5). 
Taiwan has the world's largest and most intensive industry in butterflies (7) . Details 

315 



Threatened Swallowtail Butterflies of the World 

of the extent of the trade have been presented in section 5 of this book. P. maraho is a 
rarity that is never exploited in the low quality, high volume curio trade. Specimens 
sell to specialist collectors at high prices, generally £20-£50 per individual. As has 
been emphasized throughout this book, small-scale butterfly collecting alone is 
unlikely to have a serious impact on populations. However, when destruction of 
natural habitat is so extensive that only small areas of critical habitat remain, 
intensive and commercial collecting within these critical habitats can be disastrous. 
Because of soaring prices and decreasing habitat , P. maraho is very much in demand. 
Some dealers have expressed fears that it may already by seriously endangered and 
close to extinction (4). 

Conservation Measures The conservation movement in Taiwan has 
gathered considerable pace in recent years (4), despite the lack of international 
support. The future of Papilio maraho and the many other endemic species of Taiwan 
depends upon the designation of protected areas. Four national parks are already 
established, but there is uncertainty about the inclusion of P. maraho range within 
their boundaries. A survey of the habitat of P. maraho must be a high priority. 
Biological studies would be of immense value in planning management strategies and 
developing farming or ranching. 

For the time being, capture and trade of Papilio maraho in Taiwan should be 
discouraged by whatever means seem most appropriate. In international trading 
circles, dealers should consider a voluntary ban on trading in this species. The British 
Association of Entomological Suppliers has already done this for Troides aeacus ka- 
guya, another Taiwanese endemic. 

Taiwan has a "critical fauna" for international conservation of the Papilionidae 
(see section 4). Of the 32 swallowtail species known from the island, five are endemic 
(15 per cent). These are Atrophaneurafebanus, A . horishanus, Papilio thaiwanus, P. 
maraho and P. hoppo. Reports on the status of all these species are required. 

References 

1. D'Abrera, B. (1982). Butterflies of the Oriental Region. Part 1. Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi + 244 pp. 

2. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

3. Marshall, A.G. (1982). The butterfly industry of Taiwan. Antenna 6(2): 203-4. 

4. Morris, M.G. (1985). In litt., 11 January. 

5. Severinghaus, S.R. (1977). The butterfly industry and butterfly conservation in Taiwan. 
Atala 5(2): 20-23. 

6. Shirozu, T. (1960). Butterflies of Formosa. T. Shoten, Tokyo. 65 pp. 

7. Unno, K. (1974). Taiwan's butterfly industry. Wildlife 16: 356-359. 



316 



Papilio osmana 
Papilio (Chilasa) osmana Jumalon, 1967 VULNERABLE 

Subfamily PAPILIONINAE Tribe PAPILIONINI 



Summary Papilio osmana is an extremely rare swallowtail from southern 
Leyte and north-eastern Mindanao in the southern Philippines. Very little is known 
about this butterfly, but its habitat is being rapidly exploited and degraded by human 
activities. Further surveys and biological studies are needed in order that conser- 
vation methods and protected areas may be recommended. 

Description Papilio osmana is a relatively large tailless butterfly belonging 
to the veiovis species-group. The sexes are similar with a fore wing length of 
47-52 mm (7). The species is believed to be in a mimetic association with Salatura 
melanippus (Nymphalidae: Danainae), and some individual variation is to be 
expected (7). 

Male: UFW/UHW bluish grey ground colour scattered with blue scales over the 
basal half of the forewing. The veins are covered by brownish-black scales and there 
are also black wing margins, forewing apex and a broad, black distal border with 
elongated bluish-white submarginal-postdiscal spots. The hindwing has a small, 
orange anal spot (3, 7). 

LFW/LHW differs in being lighter than above due to a reduction in the dark 
scaling. The anal spot is absent (3, 7). 

Female: More robust and darker appearance with slightly more diffuse markings 
and the anal spot yellower (3, 7). 

Distribution P. osmana was discovered relatively recently in southern 
Leyte and north-eastern Mindanao in the south-eastern Philippines (3,7). New 
localities are to be expected from future work, particularly in Samar, Mindanao and 
possibly other southern Philippine islands, but Papilio osmana is still one of the rarest 
swallowtails in the Philippines, second only to P. carolinensis (7). The holotype was 
captured at Sitio Katigahan (450 m) on the outskirts of the village of Catman near St 
Bernard in southern Leyte (3). In Mindanao the locality is not precisely given, but 
appears to be inland from Lianga, on the north-east coast (7). Further studies may 
extend the known distribution into the forests of central and southern Mindanao. 

The veiovis species-group is believed to have had its origins on the Asiatic mainland 
and the isolation of the present distribution of P. osmana (and P. veiovis in Sulawesi 
and P. carolinensis in Mindanao) is somewhat enigmatic. Some Philippine forms 
have only survived the retreat of the ice caps by adopting montane habits (e.g. P. 
chikae, P. benguetanus), but P. osmana, P. carolinensis and P. veiovis are lowland 
species. The distribution of P. osmana and P. carolinensis may prove to be larger than 
is presently known, but the reason for their absence from Luzon remains a mystery 
(4). 

Habitat and Ecology Collecting records for this very rare species are limited 
to April, May, June and July and there is presumed to be only one brood per year 
(3,7). The larval foodplant and early stages are unknown (3). Mature larvae of other 
species in the subgenus are often dark with pale bands or patches and orange 
segmental spots and spiny tubercles, and are commonly feeding on Lauraceae (2). 
The type specimens were taken on logging roads in lowland dipterocarp rain forest at 
altitudes between 150 m and 500 m (3). 

317 



Threatened Swallowtail Butterflies of the World 

Threats Papilio osmana and P. carolinensis (see next review) are both 
extremely rare and elusive rain forest swallowtails. The extent and rate of logging, 
agricultural extension, shifting cultivation and other forms of forest exploitation in 
the Philippines are such that some degree of threat to both species must be assumed 
(6). Their ranges are very small and with modern technology the habitat could be 
virtually destroyed in a matter of decades. 

According to a recent FAO/UNEP report the Philippines and Sabah are the 
tropical countries with the highest intensities of logging and deforestation. The 
Phihppine government has introduced a policy of local processing of logs, which is 
likely to steady the level of production and exploitation by logging companies, but 
that level is still extremely high. Estimates for deforestation rates in lowland 
dipterocarp forest by all causes were 100 000 hectares per year for 1976-80 and are 
90 000 hectares per year for 1981-85 (1). This latter figure is a little over 1 per cent of 
the total area of dipterocarp forest (8.9 million hectares) remaining in the Philippines 
in 1980. A further 3.4 milHon hectares are in forest fallow (1). 

Although it is difficult to assess the threat from deforestation of the habitat of P. 
osmana and P. carolinensis , their main area of distribution, Mindanao, produces 
three quarters of the total registered commercial log production for the Philippines 
(1). By implication, the threat to Mindanao forest endemics from logging must be 
greater than almost anywhere else on earth. 

A further threat only recently being fully realized is the serious drought of 1982-3 
that has affected eastern Mindanao, eastern Borneo and perhaps other islands. In 
the wake of the drought, huge fires have destroyed vast areas of forest. In Mindanao 
whole mountains have been burnt through and both Mt Apo and Mt Katanglad have 
suffered badly (5). The full extent of defohation is not yet known and will only be 
discovered from aerial or satellite survey. 

Over the Philippines as a whole logging is not the prime reason for loss of forest. 
The main cause is the increasing population of shifting cultivators 'kaingineros' and 
the spread of agriculture (1). Pressure is such that forest regeneration is unable to 
occur within the cycles imposed by the cultivators, and degeneration of both 
vegetation and soils follows (1). The number of 'kaingineros' is being swelled by 
landless tenants, labourers and unemployed rural people who need to find land to 
grow food for their families. As long as there are thousands of landless people in the 
basically rural economy of these islands, there will be no end to the destruction of the 
forest. Southern Leyte and northern Mindanao are also areas of permanent 
pineapple and coconut plantations, a source of some of the Philippines' most 
important exports. During 1980-85 the total forest area is projected to decrease by 
about 1 .5 million hectares, about 0.45 milHon being virgin or near-virgin dipterocarp 
forest (1). 

Conservation Measures Further surveys are needed to assess the distribu- 
tion of P. osmana, particularly within Mindanao and Samar. At present there are no 
projected or existing national parks in Samar or in eastern Mindanao within the 
butterfly's known range. Once the butterfly's habitat has been surveyed and its 
biology studied, areas of forest should be set aside for its protection. Many other 
endemic species of invertebrates, vertebrates and plants would also benefit from 
such actions. 

References 

1. FAO/UNEP (1981). Forest Resources Assessment Project. Forest Resources of Tropical 
Asia. FAO, Rome. 475 pp. 

318 



Papilio osmana 

2. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

3. Jumalon, J.N. (1967). Two new papilionids. Philippine Scientist 1(4): 114-118. 

4. Jumalon, J.N. ( 1969). Notes on the new range of some Asiatic papilionids in the Philippines. 
The Philippine Entomologist 1(3): 251-257. 

5. Lewis, R.E. (1984). In litt., 17 April. 

6. Treadaway, C.G. (1984). In litt., 25 May. 

7. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands. Vol. I 
Papilionidae. (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 



319 



Threatened Swallowtail Butterflies of the World 

Papilio (Chilasa) carolinensis Jumalon, 1967 VULNERABLE 

Subfamily PAPILIONINAE Tribe PAPILIONINI 



Summary Papilio carolinensis is an extremely rare and little known 
swallowtail from north-eastern Mindanao in the southern Philippines. A denizen of 
the rich dipterocarp forests of the island, it is believed to be threatened by rapid 
deforestation in the region. Surveys of its distribution, biological studies and 
presence in protected areas are needed. 

Description Papilio carolinensis is similar in general appearance to P. 
osmana (previous review). The sexes are similar with a forewing length of about 
50 mm (2, 5). 

Male: differs from that of P. osmana in having more extensive black scaling and 
orange-brown markings. The submarginal and subapical spots are white on the 
forewing , and orange-yellow on the hindwing. The dark scaling of the hindwing is 
chocolate brown and the extent of it varies, but it broadly covers the veins. Between 
the veins is diffuse orange-yellow scaling to the inside of the distal border (1, 2, 5). 

Female: similar but may have a lighter ground colour (5). 

When first discovered, P. carolinensis was presumed to be related to P. agestor 
because of its chocolate brown hindwings (2). However, closer examination of its 
morphology and genitalia revealed its close association with the other members of the 
veiovis group, P. osmana and P. veiovis (1, 5). P. carolinensis is a danaid-mimic, as 
distinguished, according to Jumalon, by the rows of white spots on the abdomen (2). 
Of the Danaidae with which it has been found to associate, it most closely resembles 
Danaus luzonensis (2). 

Distribution Papilio carolinensis was only discovered relatively recently on 
Mindanao in the south-eastern Philippines. It is known only from Agusan province in 
the north-east and Bukidnon province towards the centre of Mindanao (2). Tsukada 
and Nishiyama refer to it as the rarest Philippine swallowtail, being represented in 
collections only by five males and one female (5). In north-eastern Mindanao the 
distribution overlaps with P. osmana (5). Both P. carolinensis and P. osmana are 
presumed to be relict species of the last glacial era, but their present distribution in 
lowland tropical habitat is puzzling (3, and see previous review). 

Habitat and Ecology All specimens were collected between February and 
May and the species is presumed to be univoltine (2). Foodplants and early stages are 
unknown (2) . Mature larvae of other species in this subgenus are commonly dark with 
pale bands or patches and orange segmental spots and spiny tubercles, generally 
feeding on Lauraceae (1). Tsukada and Nishiyama expect that more specimens will 
be found in the rain forests of east-central Mindanao (2). 

Other species with which P. carolinensis has been found to associate include 
Atrophaneura mariae, Danaus luzonensis and D. vitrina (2). Graphiiim idaeoides, a 
species which is also considered to be threatened (see separate review), has been 
found at lower elevations in the same region (2). 

Threats Three localities for this species are known so far. The type locality 
is at 615 m near a small village called Barrio Kitcharao on the east shore of Lake 
Mainit , north-west of the Diwata Range in Agusan Province (2) . Behind the village is 

320 



Papilio carolinensis 

a mountain with bare slopes except for relict forest patches in rocky areas. The 
presumed rain forest habitat of the butterfly, which once covered the mountain, is 
now almost entirely destroyed. The second locality is in the mountains to the 
north-east of Malaybalay, the capital of Bukidnan Province, where large remnants of 
forest still remain. Even so, it seems that suitable habitat is very restricted, most of the 
Bukidnan Province being dry highland grasslands (2). The third and most southerly 
locality is Masara Mine in Bukidnon Province. The region is reported to be 
undergoing development and has poor prospects for this butterfly (2). 

The comments on destruction of lowland dipterocarp forest made in the review of 
P. osmana are equally applicable to P. carolinensis. Deforestation in the Philippines 
is proceeding faster than in any other state with the possible exception of Sabah . The 
main causes are shifting agriculture and logging. The effects of the extensive forest 
fires that followed the drought of 1982-3 have not yet been assessed. Aerial and 
satellite surveys are needed (4). 

Conservation Measures Further surveys are needed to assess the distribu- 
tion of P. carolinensis and other butterflies on Mindanao. At present there are no 
projected or existing national parks in eastern Mindanao within the butterfly's range. 
Biological studies of the butterfly are needed and suitable protected areas should be 
set up for this and other species native to Mindanao's forests. 

References 

1. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

2. Jumalon, J.N. (1967). Two new papilionids. Philippine Scientist 1(4): 114-118. 

3. Jumalon, J.N. ( 1969). Notes on the new range of some Asiatic papilionids in the Philippines. 
The Philippine Entomologist 1(3): 251-257. 

4. Lewis, R.E. (1984). In litt., 17 April. 

5. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands. Vol. I 
Papilionidae. (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 



321 



Threatened Swallowtail Butterflies of the World 

Papilio (Chilasa) toboroi Ribbe, 1907 RARE 

Subfamily PAPILIONINAE Tribe PAPILIONNI 



Summary Papilio toboroi occurs on Bougainville (Papua New Guinea), 
Holibara, Santa Isabel and Malaita (Solomon Islands). Although locally common on 
Bougainville and apparently adaptable to human interference , it is little collected and 
very poorly known. More data on its ecology, the extent of its distribution, and 
threats to its survival are required. 

Description This species is a large and beautiful short-tailed butterfly in the 
laglaizei group (7). The forewing length is about 60 mm in the female, 56 mm in the 
male, and the wingspan from apex to apex is about 110 mm. The sexes are similar, 
though the male tends to be smaller (1, 11, 12). Two subspecies have been described 
(10). 

UFW ground colour black-blue with a blue silky sheen and two narrow, curved 
green-blue bands (1, 11). 

UHW with a short tail, a slightly scalloped outer margin and a ground colour the 
same as the UFW. The curved blue-green median band is continuous with the inner 
band of the UFW, and there is a yellow spot at the anal angle (1, 11). 

LFW/LHW ground colour duller than the upper surface, but otherwise broadly 
similar on the forewing. Basal third of the LHW grey with blue-black veins. A 
postdiscal band is composed of seven large spots merged together; the inner five 
yellow, the rest pale grey (1, 11). 

The final instar larva is brown with black on the head, legs, prothorax, anal 
segment and prolegs (5, 13). White spots are present on thorax and abdomen and the 
tubercles are 8-9 mm long, stiff and black with white spots at the base (5, 13). The 
pupa is yellow with black on the underside and laterally on some abdominal segments 
(5,13). 

Distribution Papilio toboroi is confined to the eastern side of the Solomon 
Archipelago in the south-west Pacific Ocean 91,2,3,4,10,11). The nominate 
subspecies appears to be widespread on Bougainville (Papua New Guinea) 
(1, 2, 4, 8, 10, 11), and a second subspecies, P. t. straatmani Racheli, occurs further 
to the south-east in the Solomon Islands (9, 10). It has been recorded from Holibara 
in south Santa Isabel, and also from Malaita (9, 10). It may possibly have been 
overlooked on Choiseul Island. 

Habitat and Ecology Papilio toborio lays its eggs in large masses, as do the 
other members of the group (13). The larvae are strongly gregarious, the largest 
recorded group consisting of 700 individuals (13). Feeding is at night (13) on the 
leaves of its main food plant Litsea irianensis (Lauraceae), or alternatively Flindersia 
sp. (Rutaceae) (4). Straatman recorded caterpillars on unidentified host plants on 
high hills at about 1300 m above sea level (13). The location was in secondary growth 
far from water but near village gardens where the adults were feeding on flowers. The 
larvae are heavily predated by toads, frogs, geckos, spiders and ants, only a fraction 
reaching a spot suitable for pupation . Of these , about 20 per cent have been observed 
to produce parasitic species of Tachinidae (Diptera) and Braconidae (Hymenoptera) 
(13). 

322 



Papilio toboroi 

Threats The most significant threat to Papilio toboroi is likely to be forest 
destruction , but it is difficult to be precise about the extent of that threat . Hundreds of 
square kilometres of potential habitat have never been searched. In addition, 
Straatman's notes (12) indicate a degree of adaptability to human interference of its 
habitat. 

On Bougainville P. toboroi is said to be fairly widespread, and common in certain 
places (4). However, the fifth instar caterpillars are highly sought after as a gourmet 
food item. They are easily collected since 300 or more larvae collect in forks of the 
food plant (4) . f. laglaizei is used for the same purpose (4) . This practice is unlikely to 
constitute a serious threat to the species unless it is carried out on a commercial scale, 
for which there is no evidence at the moment. Natural losses of larvae are anyway 
known to be high, mainly to reptiles, amphibians and invertebrate predators (12). 

Conservation Measures The Solomon Archipelago has an interesting 
endemic insect fauna, of which Papilio toboroi is just one example. Given more 
information on status and distribution, there is potential for farming of selected 
rarities for the butterfly market and for release into the wild. Occasional specimens 
are already collected for sale (6, 8). A survey of the Solomon Islands insects was 
proposed some years ago, but so far no funding has been available (3) . Papilio toboroi 
may be more widespread than is apparent from its known distribution and presence in 
collections. The Solomon Islands Government requires a permit for export of 
butterflies, but this is difficult to enforce. 

References 

1. D'Abrera, B. (1971). Butterflies of the Australian Region. Lansdowne Press, Melbourne. 
415 pp. 

2. Fenner, T.L. (1983). In litt., 15 March. 

3. Holloway, J.D. (1983). In litt., 28 February. 

4. Hutton, A.F. (1983). In //«., 28 March. 

5. Igarashi, S. (1979). Papilionidae and Their Early Stages. Vol 1: 219 pp. Vol 2: 102 pp. of 
plates. Kodansha, Tokyo. (In Japanese). 

6. Insect Farming and Trading Agency (1979, 1980). Special butterfly price list. Div. of 
Wildlife, Bulolo, Morobe Province, Papua New Guinea. 

7. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supplement 17: 1-51. 

8. Parsons, M.J. (1983). In litt., 2 March. 

9. Racheli, T. (1979). New subspecies of Papilio and Graphium from the Solomon Islands, 
with observations on Graphium codrus (Lepidoptera, Papilionidae). Zoologische Med- 
edelingen 54(15): 237-240. 

10. Racheli, T. (1980). A list of the Papilionidae (Lepidoptera) of the Solomon Islands, with 
notes on their geographical distribution. Australian Entomologists' Magazine. 7: 45-59. 

11. Ribbe,C. (1907). ZweineuenPapilioformen von derSalomo-Insel Bougainville. Deutsche 
Entomologische Zeilschrift "Iris" 20: 59-63. 

12. Smart, P. (1975). The Illustrated Encyclopedia of the Butterfly World. Hamlyn, London. 275 
pp. 

13. Straatman, R. (1975). Notes on the biologies of Papilio laglaizei and P. toboroi 
(Papilionidae). Journal of the Lepidopterists' Society 29: 180-187. 



323 



Threatened Swallowtail Butterflies of the World 

Papilio (Chilasa) moerneri Aurivillius, 1919 VULNERABLE 

Subfamily PAPILIONINAE Tribe PAPILIONNI 



Summary Papilio moerneri is an extremely rare species known only from 
New Ireland, although it may occur in unexplored regions of other islands in the 
Bismarck Archipelago (Papua New Guinea). The butterfly has not been seen since 
1924 and surveys are needed to assess its present range. If it could be found and its 
conservation assured, it has great potential for farming or ranching. 

Description This elusive butterfly in the laglaizei group has only been 
recorded from the male which has a wingspan of about 108 mm (3, 7) (Plate 7.1). 

UFW ground colour black with a dark blue sheen to all but the costal and outer 
margins. A broad, relatively straight, grey-green diagonal band runs from the inner 
margin to the middle of the cell apex (1, 3). 

UHW upper surface is all black with a dark blue sheen and a very short tail (1,3). 

LFW black with a yellow submarginal band that curves inwards near the apex and 
crosses the subapical region to the costal margin (1,3). 

LHW grey-yellow with a broad, black discal band, black veins, and a wavy line 
inside the outer margin. There is also an orange-yellow streak in the abdominal 
region (1,3). 

Distribution Papilio moerneri is only known from New Ireland in the 
Bismarck Archipelago (Papua New Guinea), north-east of mainland Papua New 
Guinea. The species may also occur on the nearby islands of New Britain, and New 
Hanover, large areas of which are still unstudied (5) and very poorly known for 
butterflies. In 1939 Bang-Haas described a race of P. moerneri from New Britain and 
called it mayrhoferi, but it has not been seen since (2). 

Habitat and Ecology Very little is known about the biology and habitat of 
Papilio moerneri. In addition to the 1919 and 1924 specimens mentioned by D'Abrera 
(3), Straatman notes that there is one specimen in the Australian National Collection, 
Canberra, and that a few more were taken in 1968 (10). It has not been reported by 
lepidopterists who have visited the region in more recent years (5,6,8,9). The 
natural vegetation of the Bismarck Archipelago is rain forest, much of it in hilly or 
mountainous country, and the butterfly is presumed to inhabit these forests. 
Members of the laglaizei group are mimetic of Uraniidae (7). In other species the 
larvae are black with orange tubercles or have orange segmental bands and yellow 
spots (7). The foodplant is unknown, possibly lauraceous (7), and in other members 
of the group the eggs are laid in large masses and the larvae are gregarious (10) (see 
review of Papilio toboroi) . 

Threats The distribution of P. moerneri is too poorly known to be sure of 
the major threats to its survival. Its extreme rarity even early in this century implies 
that timber exploitation and other forms of habitat destruction may not be a prime 
threat. Nevertheless, logging is now extensive in New Ireland and must represent an 
encroachment on what is almost certainly a denizen of the primary forest. The 
Bismarck Archipelago, with recently latent but still active volcanic phenomena, has 
some highly fertile soils and the islands are responsible for most of the agricultural 

324 



Papilio moerneri 

production of the country (4) . Many parts of New Britain and New Ireland have been 
severely altered (9). 

Conservation Measures A survey of potential Papilio moerneri habitat is 
needed in order to assess its conservation status thoroughly. The Whiteman Range 
and the high interior of the Gazelle Peninsula on New Britain, and the south-eastern 
lobe of New Ireland, have wilderness areas which may reward a careful search (9). 
This species is certainly one of the rarest and least known of all Papua New Guinea 
Papilionidae and would be worthy of international conservation funding. Once 
re-located, proposals for its management and protection would be required. Once its 
conservation is assured there would be great potential for farming, and studies of 
foodplants and other requirements would be quickly repaid. World demand for 
farmed specimens of this attractive but Httle-known species would certainly be high. 
The Insect Farming and Trading Agency at Bulolo already has the experience 
required to market insects without threatening their wild populations. 

References 

1. Aurivillius, C. (1919). Eine neue Papilio-Art. Entomologisk Tidskrift 1919: 177-178. 

2. Bang-Haas, O. (1939). Neubeschreibungen und Berichtigungen der Exotischen Macrole- 
pidopteren fauna II. Entomologische Zeiischrifi 52(39): 301-302. 

3. D'Abrera, B. (1971). Butterflies of the Australian Region. Lansdowne Press, Melbourne. 
415 pp. 

4. FAO/UNEP (1981). Tropical Forest Resources Assessment Project. Forest Resources of 
Tropical Asia. FAO, Rome, 475 pp. 

5. Fenner, T.L. (1983). In litt., 15 March. 

6. Hutton, A.F. (1983). In litt.,2H March. 

7. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supplemenl 17: 1-51. 

8. Parsons, M.J. (1983). In litl.,2 March. 

9. Pyle, R.M. and Hughes, S. A. ( 1978). Conservation and utilisation of the insect resources of 
Papua New Guinea. Report of a consultancy to the Wildlife Branch, Dept. of Nature 
Resources, Independent State of Papua New Guinea. 157 pp. unpublished. 

10. Straatman, R. (1975). Notes on the biologies of Papilio laglaizei and P. toboroi 
(Papilionidae). Journal of the Lepidopterists' Society 29: 180-187. 



325 



Threatened Swallowtail Butterflies of the World 

Papilio (Papilio) hospiton Guenee, 1839 ENDANGERED 

Subfamily PAPILIONINAE Tribe PAPILIONNI 



Summary Papilio hospiton is usually known as the Corsican Swallowtail, 
despite the fact that it occurs on the island of Sardinia (Italy) as well as Corsica 
(France). It is found at altitudes above 600 m in the mountains and has declined 
dramatically through the impact of habitat destruction, commercial collecting and 
destruction of its foodplants, which are poisonous to sheep. Protection of habitat, 
surveys of distribution and detailed biological studies are urgently required. There is 
reason to believe that this is one of Europe's most seriously endangered butterflies. 

Description Papilio hospiton (known as the Corsican swallowtail, le 
Porte-Queue de Corse or the Korsicher Schwalbenschwanz) is a short-tailed black 
and yellow swallowtail with blue and red markings. The male and female are similar 
in appearance, with a wingspan of 72-76 mm (6) (Plate 7.2). 

UFW light yellow with black markings dusted with yellow scales. Black areas 
include the basal region, veins, large spots inside and above the cell, and a distal band 
that has rows of yellow submarginal and fringe spots. 

UHW short-tailed and pale yellow with a black distal band, basal region, and inner 
margin. Markings include yellow outer marginal fringe crescents and submarginal 
lunules, blue postdiscal spots, and a small red eye spot. 

LFW differs from the upper surface in that the postdiscal band is composed of 
black-bordered grey lunules, and the narrow submarginal band has a zig-zag edge. 

LHW slightly paler than the upper surface and perhaps with a little more red on the 
hindwings, but otherwise very similar (6). 

Distribution This butterfly is restricted to the islands of Corsica (France) 
and Sardinia (Italy) (1, 6), where it is extremely localized. 

Habitat and Ecology The habitat of Papilio hospiton is open, mountainous 
country between 600 and 1500 m above sea level (4,6). Although there is a 
considerable area of mountainous country on the two islands, the breeding sites for 
Papilio hospiton are extremely localized on Corsica (3), and presumably also on 
Sardinia. Precise localities are often kept secret. The larval foodplants are 
Umbelliferae, but there is some dispute as to which species are utilised. Fennel 
(Foeniculum vulgare) is commonly cited (6), although two authors give Giant fennel 
{Foeniculum {Ferula) communis) (8,9). Others claim that fennel (and carrot) leaves 
are refused by caterpillars in captivity (3). Fausser rejects claims that both 
Foeniculum communis and Ruta Corsica are used, but cites Peucedanum paniculatum 
as the true foodplant (3). Information gathered by Heath at first agreed in indicating 
Peucedanum as a foodplant (4) , but has since been refuted (5). Clearly there is a need 
for further research on the subject. Young stages in Corsica may be attacked by the 
parasitic ichneumonid wasp Trogus violaceus (3). Adults generally emerge between 
May and June and are on the wing until early August. P. hospiton is generally 
believed to be single-brooded. However, Fausser (3) has found numerous fresh 
adults and even caterpillars in Corsica in early August, and firmly believes that two 
broods are possible. In support of this contention he describes morphological 
differences in the appearance of the early and late generations (3). An alternative 
suggestion is that some over- wintered pupae could emerge very late in the season, but 

326 



Papilio hospiton 

all the evidence indicates that emergence of the over-wintered brood occurs in April , 
May or June. Hybridisation with the lowland Papilio machaon is a fairly regular 
occurrence, at least in Corsica (3). 

Threats The threats to Papilio hospiton appear to be similar in Corsica and 
Sardinia. Habitat destruction is probably primarily responsible for the decline of the 
species (4). Much of the habitat, and particularly the status of the foodplants, is 
adversely affected by agricultural practices. The umbelliferous foodplants are 
believed to be poisonous to sheep and are destroyed by fires started by local people 
(4). More localized, but also more permanent, damage to habitat is done by 
developments such as the growth of skiing resorts on Corsica (4). In addition, it is 
widely believed that excessive collecting for private and commercial purposes is 
causing severe declines in butterfly numbers. Both adult and immature stages are 
taken by local and foreign collectors (4, 5) who are aware of the rarity of the species. 
This even happens on Corsica, where collecting is forbidden by French law. 

Conservation Measures The conservation status of Papilio hospiton is 
widely understood and international concern is reflected in a number of laws and 
reports (4). The Corsican population of Papilio hospiton is protected under a French 
decree published on 22 August 1979. This is the same decree that protects Papilio 
phorbanta on Reunion and, as stated in the review of that species, the legislation 
draws attention to the species' plight, but fails to have any impact on the protection of 
its habitat and foodplants. Under the 1980 Endangered Foreign Species Act of the 
Netherlands, Dutch citizens may not import or purchase P. hospiton. In a report on 
the status of European butterflies commissioned by the Council of Europe and 
published in 1981, P. hospiton was classified as endangered and urgent action called 
for (4). The Sardinian population is not protected under Italian law and there is an 
urgent need for regional, if not national, recognition of the problem. Steps taken by 
the Piemonte Regional Council in protecting Carabiis olympiae (Coleoptera: 
Carabidae) and encouraging the preparation of a recovery plan have set an excellent 
precedent for such an effort (2). No measures have been taken specifically to protect 
the habitat of the butterfly and the designation of suitable nature reserves on both 
islands is urgently needed (4). Detailed surveys of the populations on both islands are 
required as a matter of urgent priority. These should be followed by an immediate 
ecological study to evaluate foodplants and habitat requirements, with a view to 
gazetting and management of protected areas and, if necessary, additional protective 
legislation. 

References 

1. Bretherton, R.F. and De Worms, C.G. (1963). Butterflies in Corsica 1962. Entomologists' 
Record and Journal of Variation 75: 93-104. 

2. Collins, N.M. (1984). Piemonte, Italy, protects a ground beetle. Oryx 18: 69. 

3. Fausser, J. (1980). Observations concernant Papilio hospiton Gene en Haute-Corse. 
Bulletin Liaison l Association Entomologique d'Evreux 5: 18-19. 

4. Heath, J. (1981). Threatened Rhopalocera (Butterflies) in Europe. Nature and Environment 
Series (Council of Europe) No. 23, 157 pp. 

5. Heath, J. (1984). Summary of comments received from a number of authorities during 
preparation of reference (3). In litt. , 27 February. 

6. Higgins, L.G. and Riley, N.D. (1980). A Field Guide to the Butterflies of Britain and Europe. 
4th ed., revised. Collins, London. 384 pp., 63 pi., 384 maps. 

7. Panchen, A.L. and Panchen, M.D. (1973). Notes on the butterflies of Corsica, 1972. The 

327 



Threatened Swallowtail Butterflies of the World 

Entomologist's Record 85: 149-153, 198-202. 

8. Watson, A. (1981). Butterflies. Kingfisher Books, London. 

9. Whalley, P. (1981). The Mitchell Beazley Guide to Butterflies. Mitchell Beazley, London. 
168 pp. 



328 



Papilio benguetanus 
Papilio (Princeps) benguetanus Joicey and Talbot, 1923 VULNERABLE 

Subfamily PAPILIONINAE Tribe PAPILIONNI 



Summary Papilio benguetanus is a butterfly which may be locally common , 
but has a very restricted distribution within the Cordillera Central of northern Luzon , 
Philippines. There are no known protected areas in this region. Studies are needed to 
assess the foodplant, limits of distribution, and the extent of threats to the survival of 
this butterfly. 

Description Papilio benguetanus (mistakenly called benguelana by 
D'Abrera (2)) is a lovely butterfly with short tails and a forewing length of 44-48 mm 
(Plate 7.3). The sexes are similar in appearance, but the female is slightly larger (8). 

UFW/UHW yellow with wide brown veins in the basal and discal areas, brown in 
the postdiscal and submarginal areas. A row of submarginal lunules is present on each 
wing, yellow on the forewings, orange on the hindwings, with a bluish tinge above the 
lunule on the anal angle. The female has brighter orange lunules on the hindwing and 
broader, paler brown-black streaks on its wings (8). 

LFW/LHW similar to uppersides but paler, with less brown in the postdiscal and 
submarginal areas. 

Distribution Papilio benguetanus is known only from northern Luzon in the 
Philippines (8). Its distribution is essentially the same as Papilio chikae, between 
Bontoc and Baguio in the Cordillera Central. To the east the River Cagayan valley is 
a great rice-growing region which separates the Cordillera Central from the Sierra 
Madre. The butterfly is not known from the Sierra Madre, although there could be 
access at a fairly high level at the southern end. Mt St Thomas and Mt Paoai are the 
most famous localities for this species. 

Habitat and Ecology Papilio benguetanus is a relict species of continental 
origin, as is Papilio chikae (see separate review) (8). At times it has been considered 
an island race of the continental species P. xuthus, but it is quite distinct in 
appearance. Vnhke xuthus, it has no known seasonal variation (8). On Mt St Thomas 
and Mt Paoai the most famous localities for the species, the butterflies occur in some 
numbers from March to June, when many individuals can be found flying along 
pathways and hill-topping between 7 a.m. and 11 a.m. (7, 8). Males sometimes fight 
each other quite fiercely in their territorial battles (7). In fine weather it associates 
with Parantica phyle (Danaincie). Appias phoebe (Pieridae), Celastrina spp. (Lycae- 
nidae) and Hestina spp. (Nymphalidae) (8). Both Parantica phyle and Hestina bear a 
superficial resemblance to P. benguetanus on the wing (8). Females are more 
secretive than males and the hostplant and immature stages are unknown (4,8). 
Apart from the nominate species P. benguetanus is the only other member of the 
xuthus group (3). P. xuthus has a green eye-spotted caterpillar with segmental orange 
spots, feeding on Rutaceae (5). 

Threats This attractive and rare butterfly clearly has a very limited 
distribution in northern Luzon and although it is rather more common than Papilio 
chikae, it may nevertheless suffer from a similar level of disturbance. It is listed under 
category Vulnerable in recognition of the human population pressures that are 
resulting in extensive habitat alteration for agriculture and forestry in Luzon , and also 

329 



Threatened Swallowtail Butterflies of the World 

because the species is coveted by collectors (1). There are some indications that P. 
benguetanus may thrive in partially cleared areas and even in areas around vegetable 
growing operations, possibly even deriving benefit from man's activities (7). In this 
event its conservation status might require review. As stated in the review of P. 
chikae, the precise extent of habitat disruption in the Cordillera Central remains 
unknown. Baguio is an important summer resort and the surrounding region may be 
subject to recreational pressure as well as agricultural and forestry expansion. Much 
may depend upon the habitat of the foodplant, and its ability to withstand 
disturbance. 

Conservation Measures Papilio benguetanus rarely appears on commercial 
Usts but it may be the subject of substantial private collecting operations. It is to be 
hoped that entomologists visiting Luzon will exercise restraint. Rather than collect 
numerous specimens, visitors to the region could be encouraged to search for the 
foodplant and young stages, and to visit remote regions of the Cordillera Central in 
order to assess the distribution of the species more fully. It has not been possible to 
assess whether there are any functional national parks within the range of this 
butterfly, but this seems improbable . The Philippine national parks system is curently 
under review (6). More data are needed on the extent of habitat disruption at high 
altitude in the Cordillera Central. This information should be used to propose a site 
for a new national park, not only as a conservation area for wildlife, but also as a 
pubHc amenity and for protection of the watersheds. 

References 

1. Ae, S.A (1983). In lilt., March 18. 

2. D'Abrera, B. (1982). Butterflies of the Oriental Region. Part 1. Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi + 244 pp. 

3. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

4. Iwase, T. (1965). How Papilio chikae was found and named. Tyo To Ga (Transactions of the 
Lepidopterists' Society of Japan) 16: 44-47. 

5. Monroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
EHtowo/ogwr Supplement 17: 1-51. 

6. PoUisco, F.S. (1982). An analysis of the national park system in the Philippines. 
Likas-Yaman. Journal of the Natural Resources Management Forum 3(12): 56 pp. 

7. Treadaway, C.G. (1984). In /»/., 25 May. 

8. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands. Vol. 1 
Papilionidae. (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 



330 



Papilio acheron 
Papilio (Princeps) acheron Grose-Smith, 1887 RARE 

Subfamily PAPILIONINAE Tribe PAPILIONINI 



Summary Papilio acheron is a rare montane species from the northern 
highlands of Borneo. Its known distribution is from Mt Kinabalu in Sabah to Mt Dulit 
in Sarawak, but research may reveal more localities further inland. Whilst some 
national parks appear to guarantee the livelihood of the species, vigilance is needed 
to ensure the integrity of the park boundaries. 

Description Papilio acheron is a large, mainly black butterfly with a 
wingspan of 58-66 mm. Both sexes appear to mimic the sympatric Alrophaneura nox 
(3,7,8). 

Male: upperside black with no markings but lighter bluish scales scattered over the 
discal part of the hindwing between the veins (3, 7, 8). Hindwing with a slightly 
scalloped outer margin but no tail. Underside also black, with a basal red spot on the 
LHW and a short golden or buff distal band enclosing three black postdiscal spots. 
Diffuse, blue postdiscal lunules extend from this band to the costal margin (3, 8). 

Female : very similar to the male but with a paler ground colour; usually resembUng 
the female oi A. nox in having pale brown scaling bordering the veins of the upper 
part of the UFW. Occasionally a uniformly black male-like example is found (3, 8). 

Distribution Papilio acheron is found in the mountains of Sabah, Sarawak 
and Brunei in north-western Borneo. It occurs at altitudes over 1000 m on Gunung 
(Mt) Kinabalu, south along the Crocker Range to Gunung Pagonprick and Bukit 
Retak in eastern Brunei and into northern Sarawak where it is known from Gunung 
Mulu and Gunung Dulit (2, 3, 5, 7, 8). It may also occur on other mountains in 
Sarawak, notably the Tamabo and Penambo Ranges, the Linau Balui Plateau and the 
Hose Mountains, and further east in the border area of the Eastern Province of 
Kalimantan (Indonesia), but such localities have not yet been reported. 

Habitat and Ecology Papilio acheron is a rare and localized species from 
lower montane forest. Males fly low and rapidly on patrol routes along streams, often 
stopping to drink (7, 8). Females are much more scarce and the immature stages and 
foodplants are unknown (8). P. acheron is a member oi X\\q memnon group, in which 
the foodplant is normally rutaceous and the caterpillars have raised bands on the 
metathorax and first abdominal segment, the former ending in a prominent eye-spot 
(4,6). 

Threats Montane habitats in Borneo tend to be rather better protected 
than lowland ones. Both Kinabalu and Mulu are designated as national parks. At 
present Papilio acheron is not a matter for great conservation concern, but it must be 
emphasized that even montane habitats are not free of threat. Kinabalu National 
Park is under pressure from small farmers, copper mining concessions and tea 
plantations. The cutting of virtually all the forest on the Pinosuk plateau for cattle 
farming, sawmills and hydro-electric projects was a particularly significant loss ( 1 ) . In 
the Eastern Province of Kalimantan where it borders with Sarawak lies the giant 
recently declared Sungai Kayan — Sungai Mentarang Reserve (1.6 million ha). The 
reserve is largely hill forest with some lowland and montane forest (elevation range 
200-2558 m) and may well include Papilio acheron. However, the area has always 

331 



Threatened Swallowtail Butterflies of the World 

supported quite a high density of people; their influence can be seen in the extensive 
grasslands and the effects of hunting (9) . (See also the review of Graphium procles) . 

Conservation Measures Further research in the highlands of northern 
Borneo may extend the known distribution of P. acheron. In addition to distribu- 
tional study, vigilance is needed to ensure that national park boundaries are not 
violated by farmers, and that parcels of land are not excised for mining, large-scale 
plantations of cash crops, or extensive tourist facihties, particularly on Kinabalu. In 
the Kayan-Mentarang Reserve in Kalimantan surveys are needed not only to check 
for the butterfly, but also with a view to revisions of the boundaries to exclude 
disturbed areas and include more lowland areas (9). 

References 

1. Barlow, H.S. (1983). In //«., 29 June. 

2. Cassidy , A.C. ( 1982). An annotated checklist of Brunei butterflies including a new species of 
Catapaecilma (Lycaenidae). Brunei Museum Journal 5: 202-272. 

3. D'Abrera, B. (1982). Butterflies of the Oriental Region. Part 1. Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi + 244 pp. 

4. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

5. Holloway, J.D. (1978). Butterflies and Moths. \n Kinabalu Summit of Borneo SabahSoaeVy 
Monograph, 25-278. 

6. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Enfowo/ogwr Supplement 17: 1-51. 

7. Robinson, J.C. (1975-6). Swallowtail butterflies of Sabah. Sabah Society Journal 6: 5-22. 

8. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands. Vol. I 
Papilionidae. (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 

9. UNDP/FAO National Parks Development Project (1981). National Conservation Plan for 
Indonesia Vol. 5: Kalimantan. FAO, Bogor. 



332 



Papilio jordani 
Papilio (Princeps) jordani Fruhstorfer, 1902 RARE 

Subfamily PAPILIONINAE Tribe PAPILIONNI 



Summary Papilio jordani is a large and striking swallowtail from the 
Minahassa Peninsula in northern Sulawesi, Indonesia. It is extremely rare and its 
life-cycle and habits are very poorly known. Hopefully it breeds within the bounda- 
ries of the new Dumoga-Bone National Park. Surveys of the region and studies of 
the butterfly's breeding and feeding habits are needed. 

Description Papilio jordani is a beautiful, large, tailless butterfly which was 
placed in the fuscus group by Munroe (5) but has since been transferred to the polytes 
group (2, 4). The male differs markedly from the female, which mimics Idea blan- 
chardii{4, 8). The forewing length is 75-80 mm (male) or 80-85 mm (female) (1,8). 

Male: almost entirely black-brown with a scalloped outer edge to the hindwing. 
There is a series of white, outer marginal fringe spots on both wings, those on the 
forewing becoming larger towards the tornus, and a band of large, elongated, white 
discal spots on the hindwing (1, 8). 

Female: white with black or dark brown scaling over the veins, forming lines or 
streaks in the cell, and between the veins of the forewing (1,8). The hindwing has 
some dark submarginal wedge-shaped spots and some smaller discal ones, with dark 
scales scattered between them. The forewing also has dark scales scattered over the 
distal region (1,8). 

Distribution Papilio jordani is now found only on the Minahassa Pen- 
insula, which is the northern limb of Sulawesi (Celebes), Indonesia. The two known 
localities are Tolitoli in the north-west and Doluduo near Manado in the extreme 
north-east of the island (1). The type locality, however, is apparently in south- 
eastern Sulawesi (3). The reason why the type locality is so far removed from the area 
where P. jordani is now found remains unexplained. Either the species' range has 
retracted quite drastically in the past 85 years, or the type specimen was a vagrant or 
mistaken locality. 

Habitat and Ecology This extremely rare and local butterfly is very poorly 
known. Males have been observed to fly slowly and straight along sunny streams 
keeping to a height of about 3 m (8). Females mimic Idea in their flight as well as their 
appearance, fluttering gently between the banks of streams, but when alarmed can 
fly off swiftly in the manner of other Papilio species (8). P. jordani associates with P. 
ascalaphus but whilst y'oriifl/i; usually remains near streams in primary forest, ascala- 
phus inhabits areas near villages (8). Tsukada and Nishiyama note that mimics of 
Idea are often rare (8). The mimics Graphium idaeoides (see separate review), 
Papilio veiovis and Elymnias kunstleri (Satyridae) are certainly rare, but Graphium 
delesserti is often common (8). The larval foodplant of P. jordani is unknown and the 
young stages undescribed. However, other members of t\\t polytes group have larvae 
with metathoracic and abdominal bands present, and vestigial eye-spots (2). The 
pupa is rough and curved and the foodplants are in the Rutaceae (2) . Adults occur all 
year round, with a peak of emergence in November and February (8). 

Threats Sulawesi is well known for its richness and high levels of endemi- 
city in Lepidoptera, but many butterfly habitats are being destroyed by extensive 
deforestation (7). Large areas stretching between Makassar and the Northern and 

333 



Threatened Swallowtail Butterflies of the World 

Central Districts are already covered with the unpalatable and useless grass Imperata 
cylindrica (lalang), a sure sign of over-exploitation. Sulawesi has 10.4 million people 
in a land area of 190 000 sq. km, but they are not evenly distributed. Settlement is 
concentrated on the southern arm and the Minahassa Peninsula (6), where this 
butterfly lives. Nevertheless, Papilio acheron is too poorly known to be sure of the 
extent to which this concentration threatens its survival. The peninsula is mountai- 
nous with a natural vegetation of lowland and montane forest. Logging, shifting 
cultivation and some irrigated rice-growing in the valley bottoms are the main uses to 
which the land is put. Irrigation projects in the valleys of the Dumoga and Bone rivers 
in the centre of the Minahassa Peninsula have received substantial support from the 
World Bank. 

Conservation Measures At the end of the 1970s, a WWF/IUCN project in 
northern Sulawesi identified the catchment areas of the Dumoga and Bone rivers as 
the most pertinent location for a large conservation area that would afford protection 
to the rich and varied flora and fauna of the province (6). This culminated in an 
agreement between the Government of Indonesia and the World Bank, whereby the 
latter would give considerable financial support towards the development of the 
proposed Dumoga-Bone National Park, in co-operation with WWF. The project 
aims to increase threefold the production of rice on 13 000 ha of prime agricultural 
land, whilst maintaining the vital watersheds which feed the land by preserving 
280 000 ha of hilly country as the new national park. The project began in 1980. WWF 
personnel were employed to write a management plan and the park has now been 
properly gazetted. Encroachment, which was at first severe, has now stabilized. 

In 1985 the Royal Entomological Society of London is celebrating its 150th 
anniversary by organizing 'Project Wallace" in this very important park. With over 
100 entomologists taking part, P. jordani will certainly be tracked down if it occurs 
there. Once found, it will hopefully be possible fully to elucidate the life-cycle and 
biology of the species. 

References 

\. D'Abrera, B. (1982). Butterflies of the Oriental Region. Part 1. Papilionidae and Pieridae. 
Hill House, Victoria, Australia, xxxi + 244 pp. 

2. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

3. Haugum, J. (1984). In Hit., 14 April. 

4. Haugum, J., Ebner, J. and Racheli, T. (1980). The Papilionidae of Celebes (Sulawesi). 
Lepidoptera Group of 1968 Supplement 9: 21 pp, 1 map, 2 pi. 

5. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
E/jtowo/ogK/ Supplement 17: 1-51. 

6. National Parks Development Project, UNDP/FAO (1982). National Conservation Plan for 
Indonesia. Vol. VI: Sulawesi. FAO, Bogor. 

7. Straatman, R. (1968). On the biology of some species of Papilionidae from the island of 
Celebes (East-Indonesia). Entomologische Berichten 28: 229-233. 

8. Tsukada, E. and Nishiyama, Y. (1982). Butterflies of the South East Asian Islands. Vol. 1 
Papilionidae. (transl. K. Morishita). Plapac Co. Ltd., Tokyo. 457 pp. 



334 



Papilio weymeri 
Papilio (Princeps) weymeriNiepeh, 1914 RARE 

Subfamily PAPILIONINAE Tribe PAPILIONNI 



Summary Papilio weymeri is endemic to Los Negros Island and Manus 
Island, Papua New Guinea. There is little information on its biology and distribution. 
Further studies and surveys are needed. The island has extensive rain forests and 
secondary growth. Plans for sustainable development should include conservation 
measures for this endemic butterfly and other forms of wildlife. 

Description Papilio weymeri is a large butterfly with a forewing length of 
about 70 mm (male) or 75 mm (female) (Plate 7.4). The Papuan group of swallow- 
tails of which this species is a member was described by Munroe as the aegeus group 
(7), but Hancock has made a few changes and renamed it the gambrisius group (4). 
Papilio weymeri is used here in preference to the synonym cartereti Oberthiir, 
although seniority may be disputed. Both names were published in the first six 
months of 1914(2). 

Male: UFW/UHW black with a cream postdiscal band of large spots across the 
forewing, and a discal band of cream streaks over the hindwing. Both wings have 
white fringe spots. The hindwing is slightly scalloped and has no tail (1, 10). 

Female: UFW blackish-brown relieved by a postdiscal band of white streaks. 
These become very diffuse and are suffused with dark scales in the submarginal 
region where they terminate as narrow yellow spots. There are also some cream 
fringe spots (1, 10). 

UHW blackish-brown with yellow submarginal lunules, relatively large, iridescent 
blue postdiscal spots, and an orange-red eyespot. Blue scaling occurs on the inside of 
the eye-spot, yellow spots inside some of the blue postdiscal spots, and there are 
cream fringe spots (1, 10). 

Distribution Papilio weymeri is endemic to Manus and Los Negros Islands 
of the Admiralty group in the northern part of Papua New Guinea. Manus Island is 
about 80 km long and 32 km wide, with an approximate area of 2100 sq. km (6). Los 
Negros is much smaller. 

Habitat and Ecology The young stages of P. weymeri have been collected 
and photographed but not described in print (5). The adults apparently have the 
unique habit of laying their eggs on the upperside of the hostplant's leaves (5). The 
early stages have been seen on Micromeliim mimitum (Rutaceae) (8), and possibly on 
Flindersiasp. (Rutaceae) (3). Many of Manus Island's forests were disturbed during 
the Second World War, but have since returned to secondary forest and bush (9). It 
seems likely that the rutaceous shrubs used by P. weymeri can regenerate in 
secondary growth. It is not clear whether P. weymeri is a denizen of rain forest on the 
island, or of forest edges and areas of regeneration. Females have been recorded 
flying slowly in small , stunted brush ; males fly erratically and never seem to settle (2) . 
Females are rarer than males; June to August are good flight months (2). 

Threats Since Papilio weymeri has adapted to areas disturbed during 
wartime (9), it would appear to be a relatively resilient species. The human 
population of Manus is relatively large, but is concentrated along the shorelines (9). 
The interior is hundreds of square kilometres of wild and sparsely populated country 

335 



Threatened Swallowtail Butterflies of the World 

with large expanses of natural and regenerating vegetation (3, 6, 9). Nevertheless, 
other endemics on Manus Island are threatened and there is a need for careful 
monitoring of Papilio weymeri. The Manus Green Tree Snail Papustyla pulcherrima 
(Gastropoda: Camaenidae) is endemic to the Manus forests and threatened by 
logging of the trees upon which it lives, and by large scale collecting (11). 

Conservation Measures Basic details of habitat and life cycle are needed for 
this butterfly. It is known to feed on Rutaceae, but whether disturbance of natural 
forest is likely to threaten or encourage the species remains unknown. Studies on 
Manus are a clear priority, followed by monitoring of butterfly populations and 
human development on the island. 

It has been proposed that a Wildlife Management Area could be designated in 
central Manus, where the rain forest is relatively undisturbed (9). Wildlife Manage- 
ment Areas are an expression of local as well as national concern for wildlife, being 
set up on privately owned or traditionally held land. Such areas could effectively 
serve to protect Papilio weymeri, the Manus Tree Snail and other wildlife (9). There 
is already a certain amount of ranching of P. weymeri on Manus (9). This could be 
further encouraged as a measure for both conservation and development. Most of 
Manus Island is unsuitable for arable or tree crops, or pasture improvement (6). The 
comprehensive 1980 report by the Papua New Guinea Office of Environment and 
Conservation on the resources of Manus Province (6) is an ideal starting point for the 
development of the island's resources in an ecologically sustainable manner. 

References 

1. D'Abrera, B. (1971). Butterflies of the Australian Region. Lansdowne Press, Melbourne. 
415 pp. 

2. Ebner, J. A. (1971 ). Some notes on the Papilionidae of Manus Island, New Guinea. Journal 
of the Lepidopterisls' Society 25: 73-80. 

3. Fenner, T.L. (1983). In lilt., 15 March. 

4. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

5. Hutton, A.F. (1983). In litt.,28 March. 

6. Kisokau, K. M. ( 1980) . Manus Province, a physical resource inventory. Report of the Office 
of Environment and Conservation, Papua New Guinea. 73 pp. 

7. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supplement 17: 1-51. 

8. Parsons, M.J. (1983). In lit!., 2 March. 

9. Pyle, R.M. and Hughes, S. A. (1978). Conservation and utilisation of the insect resourcesof 
Papua New Guinea. Report of a consultancy to the Wildlife Branch, Dept. of Nature 
Resources, Independent State of Papua New Guinea. 157 pp. unpublished. 

10. Smart, P. (1975). The Illustrated Encyclopedia of the Butterfly World. Hamlyn, London. 275 
pp. 

11. V^eUs, S.M., Pyle, R.M. andCo\Vms,N.M. (1983). The lUCN Invertebrate Red Data Book. 
lUCN, Gland. L + 632 pp. 



336 



Papilio sjoestedti 
Papilio (Princeps) sjoestedti Aurivillius, 1908 RARE 

Subfamily PAPILIONINAE Tribe PAPILIONNI 



Summary Papilio sjoestedti, sometimes known as the Kilimanjaro Swall- 
owtail, flies in the montane forests of Mt Meru. Mt Kilimanjaro and Ngorongoro in 
north-eastern Tanzania. It has a very restricted range but is well protected in national 
parks. 

Description Papilio sjoestedti is a relatively large tailless swallowtail of the 
cynorta group (5, 7) with a forewing length of 45-54 mm (Plate 7.5). The female is 
slightly larger than the male and has a different pattern; like most other members of 
the cynorta group, the female mimics members of the genus Amauris (Danaidae) 
(1,2,7,8). 

Male: ground colour blackish-brown with white markings consisting of a straight 
median band of postdiscal spots and a series of fringe spots on the forewing (1, 3, 8). 
The hindwing has a slightly broader, unbroken, white discal band and a series of 
white fringe spots (1, 3, 8). 

Female: The forewing has fewer, more widely spaced white postdiscal spots and 
several discal spots. The hindwing has a series of large fringe spots and a very broad 
yellow-ochre discal median band (1,3. 8). 

The lower surfaces are darker and mimetic of Acraeinae (5). The female has a 
truncated orange-brown basal area on the LHW (1). 

Distribution Papilio sjoestedti occurs as two subspecies in north-eastern 
Tanzania. The nominate form P. s. sjoestedti is found on Mt Meru, the highlands 
around the rim of Ngorongoro Crater, and further south in the hills near Mbulu (6), 
while P. s. atavus is only known from Mt Kilimanjaro (1, 3, 6, 8). 

Habitat and Ecology Papilio sjoestedti flies in montane forest at altitudes 
between 1900 and 2800 m (1, 6). The immature stages and foodplants are unknown, 
but in other members of the cynorta species-group the caterpillars are green when 
mature, with metathoracic and abdominal bands and no eye-spots, feeding on 
Rutaceae. Possible foodplant genera include Calodendron, Vepris and Teclea (4). 
The mountains occupied by P. sjoestedti are all volcanic, and have a much more 
recent origin than the Usambaras, Uzungwas and other smaller massifs of eastern 
Tanzania. - 

Threats Papilio sjoestedti is listed here because it has a very limited range 
within an East African vegetation type that is being depleted daily. In general, the 
main threats are forest clearance for agriculture and plantation forestry. However, in 
this case it is heartening to note that the range of the species lies almost entirely within 
the boundaries of protected areas. Highly priced specimens of this species have 
recently appeared in dealers' lists. 

Conservation Measures Mt Meru, Mt Kilimanjaro and the Ngorongoro 
Crater are all national parks, open to visitors but protected from incursion by 
agriculturalists and foresters. Although there is no immediate threat to the forested 
areas in these parks, there is increasing evidence of inadequate management 
resources within the Tanzanian National Parks authority. This is particularly 

337 



Threatened Swallowtail Butterflies of the World 

manifest in the disturbing levels of poaching reported from Ngorongoro Crater, a 
park not only designated as a Biosphere Reserve under the Man and the Biosphere 
programme of UNESCO , but also proposed as a World Heritage Site of international 
importance. 

References 

1 . Carcasson , R. H . ( 1960) . The swallowtail butterflies of East Africa ( Lepidoptera . 
Fapilionidae). Journal of the East African Natural History Society Special Supplement 6: 33 
pp. + llpl. (Reprinted by E.W. Classey.Faringdon, 1975). 

2. Carcasson , R. H . ( 1980) . Collins Handguide to the Butterflies of Africa. Collins, 
London. Hardback edition, 188 pp. 

3. D'Abrera, B. (1980). Butterflies of the Afrotropical Region. Lansdowne Editions. 
Melbourne, xx + 593 pp. 

4. Hancock, D.L. (1979). Systematic notes on three species of African Papilionidae 
(Lepidoptera). /4rnoW(a 8(33): 1-6. 

5 . Hancock , D . L. ( 1983) . The classification of the Papilionidae (Lepidoptera) : a 
phylogenetic approach. Smithersial: 1-48. 

6. Kielland, J. (1983). In litt. , 6 March. 

7. Munroe,E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist Supplement 17: 1-51. 

8. Williams, J. G. (1969). A Field Guide to the Butterflies of Africa. Collins, London 
238 pp. 



338 



Papilio manlius 
Papilio (Princeps) manlius Fabricius, 1798 INDETERMINATE 

Subfamily PAPILIONINAE Tribe PAPILIONNI 



Summary Papilio manlius is said to be quite common in Mauritius, despite 
the widespread loss of native vegetation there. The caterpillars are able to survive on 
cultivated Citrus, but native habitat is virtually restricted to the Black River Gorges. 

Description The male of this short-tailed butterfly in the nireus species- 
group (8) is black with bright greenish-blue markings. The female, at first described 
as a separate species (9), is paler in colour. The forewing length is 42-52 mm, and the 
wingspan is 70-89 mm. 

Male: UFW ground colour black with a median band of iridescent greenish-blue 
spots and yellow fringe spots. 

UHW slightly scalloped with a short tail and similar greenish-blue spots. 

LFW/LHW blackish-brown with lighter wing margins and on the hindwing a 
yellow tinge, a white band conspicuous in flight (15), white submarginal spots and a 
white lunule (4,9, 11). 

Female: ground colour brown with pale bluish-green spots in a similar pattern 
(4, 9, 11), lacking the white band found on the underside of the male (15). 

Distribution Papilio manlius is endemic to Mauritius, in the Mascarene 
Islands, Indian Ocean. It is a close relative of P. phorbanta of Reunion (see separate 
review), P. epiphorbas of Madagascar and the other members of the nireus 
species-group, or 'blue-banded papilios" of the mainland (3, 5). 

Habitat and Ecology The vegetation types of Mauritius range from coastal 
marshes and palm savannas to lowland, upland and dwarf forests (14). The lowland 
forests were said to be extremely productive and were one of the main reasons for the 
settlement of the island (10). They once covered much of the island below about 
220 m, where the rainfall is 1000-2500 mm per year (10). The upland forests once 
covered most of the fertile uplands of the Plaine Wilhelms and the south of the island, 
where the rainfall is heavy (about 3800 mm per year), and plant growth luxuriant, 
although the soils are poor (10). 

Little is known of the habits and biology of Papilio manlius in its natural state, 
mainly because so little of its habitat remains (see Threats). As in other members of 
the nireus species group, the natural foodplant would probably be a wild member of 
the family Rutaceae, but this has not been ascertained. P. manlius caterpillars are 
bright green at all stages, with a red osmeterium (5). The adults are reported to be 
common throughout the island and in no danger of extinction because they can breed 
on Citrus (15). However, the populations on natural vegetation are believed to be 
small and are a matter of conservation concern. 

Threats The majority of the natural vegetation of Mauritius has been cut 
down and the land developed for agricultural purposes, particularly for growing 
sugar cane. Only 2 per cent of the 1865 sq. km of Mauritius remains under native 
vegetation (6). The lowland forests are nearly extinct, only a few impoverished 
remnants remain on the western slopes of the mountains between Mt du Rempart 
and Chamarel (10). Upland forest is restricted to about 1600 ha in the south-west of 
the island, around the Black River Gorges (10). These gorges are believed to be one 

339 



Threatened Swallowtail Butterflies of the World 

of the few remaining areas of natural forest suitable for the butterfly. Although the 
gorges are strictly protected (1,5) they may be doomed because of the encroachment 
of introduced competitors such as Chinese guava {Psidium cattleianum) and Privet 
{Ligustrum walkeri), which strangle the understorey and prevent tree regeneration 
(5). The major tree species are hardy plants adapted to cyclones. They are long-lived 
and robust, but reproduce at a low rate and at irregular intervals. As a result they 
must be considered highly vulnerable to disturbance of this kind (1 , 10). The situation 
is a parallel of the problems of Papilio aristophontes with encroaching bananas in the 
forests of Grande Comore, but far more serious (see separate review). 

Because of its ability to breed on cultivated Citrus, P. manlius itself is seemingly 
reprieved from extinction, even if its natural habitat disappears completely. 
Nevertheless, it is reviewed here and given Indeterminate status in order to draw 
attention to its circumstances. It is certainly undesirable that a species should be 
threatened with the possibility of being entirely dependent upon man and his crops. 
There are many pitfalls to such an existence. For example, insecticidal measures are 
no doubt periodically taken against the more serious pest P. demodocus which, 
however, has an assured existence elsewhere. Developments in the efficacy of 
insecticides, or of biological control measures, could threaten P. manlius. In 
addition, P. demodocus is known to be a competitive and aggressive species, capable 
of ousting other species. It has been suggested that P. manlius could suffer in this way 
(13). However, records indicate that P. demodocus was absent from Mauritius in 
1833 (2) and 1866 (12), but present in 1908 (7). During the past three quarters of a 
century some sort of balance between the species has presumably been struck. 

Conservation Measures Because of its ability to feed on Citrus, it seems 
unlikely that P. manlius is threatened with extinction. Nevertheless, the likelihood 
that the remaining naturally-breeding populations are restricted to native forests that 
are declining in the face of encroachment of exotic species is a matter of concern not 
only for P. manlius but for many other forest endemics as well. Mauritius has a wealth 
of endemic plants, invertebrates and vertebrates, all of which are under threat 
through habitat destruction. Three butterfly subspecies, Hypolimnas dubius drucei, 
Antanartia borbonica mauritiana and Salanis angustina vinsoni are believed to be 
already extinct on Mauritius, although the first of these still survives on the Comoro 
Islands (5). Cyclyrius mandersi, an endemic lycaenid butterfly of coastal regions, may 
be threatened with total extinction (5). 

It is clear that careful management and recovery plans are needed for the 
remaining native vegetation of Mauritius. Every encouragement should be given to 
research programmes that seek further to facilitate the protection of these interna- 
tionally important localities. 



References 

1. Barnes. M.J.C. (1982). In litt., 2 August. 

2. Boisduval, J.B.A.D. de (1833). Faune Entomologique de Madagascare, Bourbon et 
Maurice. Paris. 

3. Carcasson, R.H. (1980). Collins Handguide to the Butterflies of Africa. Collins, London. 
Hardback edition, 188 pp. 

4. D'Abrera, B. (1980). Butterflies of the Afrotropical Region. Lansdowne Editions, 
Melbourne, xx + 593 pp. 

5. Davis, P.M.H. and Barnes, M.J.C. (in press). The butterflies of Mauritius. Journal of 
Research on the Lepidoptera. 

340 



Papilio manlius 

6. Jones, C.G. (1980). Parrot on the way to extinction. Oryx 15: 350-354. 

7. Manders, N. (1908). The butterflies of Mauritius and Bourbon. Transactions of the 
Entomological Society of London 1907: 429-454. 

8. Munroe. E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist S\XY)p\tmtnX 17: 1-51. 

9. Paulian, R. and Viette, P. (1968). Faune de Madagascar. XXVII Insectes Lepidopteres 
Papilionidae. O.R.S.T.O.M. and C.N.R.S., Paris. 97 pp., 19 pi. (2 col.), 34 figs. 

10. Procter, J. and Salm, R. (1974). Conservation in Mauritius. Report to lUCN/WWF. 

11. Smart, P. (1975). The Illustrated Encyclopedia of the Butterfly World. Hamlyn, London. 
275 pp. 

12. Trimen,R.( 1866). Notes on the butterflies of Mauritius. Transactions of the Entomological 
Society of London 1866: 329-344. 

13. Vane-Wright, R.I. (1983). Pers. comm., 9 June. 

14. Vaughan, R.E. and Wiehe, P.O. (1937). Studies on the vegetation of Mauritius, I. A 
preliminary survey of the plant communities. Journal of Ecology 25: 289-343. 

15. Vinson, J.M. (1938). Catalogue of the Lepidoptera of the Mascarene Islands. Mauritius 
Institute Bulletin December 1938 : 1-69. 



341 



Threatened Swallowtail Butterflies of the World 

Papilio (Princeps) phorbanta Linnaeus, 1771 VULNERABLE 

Subfamily PAPILIONINAE Tribe PAPILIONNI 



Summary Papilio phorbanta, the Papillon La Pature , is protected by law on 
the island of Reunion where it is now endemic, having been extinct in the Seychelles 
since 1890. Without adequate protection of its habitat and food plants the species 
could still be very seriously threatened, despite the legislation. Surveys of its habitat 
and designation of protected areas are urgently required. 

Description This beautiful short-tailed butterfly in the nireus species-group 
(10) has a forewing length of 40-55 mm. The male is black and blue and the female is 
brown with white markings (6, 11) (Plates 8.1 and 8.2). This difference between the 
sexes is unique in the nireus species group (4), and may be caused by mimicry of 
Euploea goudotii (Danainae) (7). 

Male: UFW/UHW black with a broad median band and blue spots. 

LFW/LHW ground colour blackish brown becoming yellowish at the wing 
margins, with whitish submarginal spots (11). 

Female: UFW/UHW red-brown with white submarginal spots (2, 11). 

LFW/LHW brown with a pale grey marginal band, black triangular spots and a 
grey mark on the hindwing (2, 11). 

Distribution This species is now endemic to the island of Reunion , a French 
Overseas Department 900 km east of Madagascar in the Indian Ocean (11). The 
much smaller subspecies nana Oberthiir is known only from a single pair from the 
Seychelles and is believed to be extinct since 1890 (6). 

Habitat and Ecology Reunion is a volcanic island with a remarkable 
topography. The highest point is 3069 m above sea level and 61 per cent of the land 
surface is at altitudes over 1000 m. Because of the altitude and ruggedness of the 
terrain, large areas of upland vegetation remain unaffected by man. In the lowlands 
however, virtually no forest remains. Papilio phorbanta is a forest-dwelling butterfly 
which, like other members of the nireus species group, feeds on Rutaceae. The 4 cm 
long green or yellow-green caterpillars (11) are nowadays usually found on Citrus 
trees, to which the species has apparently adapted . The 3 cm long pupae are known to 
be collected and raised in captivity for commercial purposes. P. phorbanta has long 
used Citrus as a food source , having been recorded in garden fruit trees by Manders in 
1908 (7) and by Boisduval as long ago as 1833 ( 1 , quoted in 5). However, the extent of 
its present colonization of Citrus groves has not been reported. 

Threats It has been suggested that the pair of Papilio phorbanta nana from 
the Seychelles was either artificially introduced or wind-blown from Reunion (2, 6), 
but in view of the unusually small size of the pair, such a provenance seems unlikely. 
Although the origin of the subspecies was probably Reunion, the small size suggests a 
lengthy period of isolated evolution and natural selection. In the absence of further 
evidence there is no reason to suppose that nana was anything but a rare subspecies of 
P. phorbanta, now extinct for unknown reasons. Such an event may act as a warning 
to prevent the same fate for the nominate subspecies on Reunion. 

Well into the 1950s Papilio phorbanta was still common on Reunion (13). 
However, with the rapidly expanding human population, more and more land has 

342 



Papilio phorbanta 

been turned over to agriculture, mainly for sugar (9). Although this has caused severe 
reductions in the butterfly's range, there are still significant areas of upland forest on 
Reunion, theoretically protected due to the rugged relief (13). Evergreen rain forest 
still covers a large belt around Piton des Neiges (3069 m) and Piton de la Fournaise 
(2631 m) (9). Reunion is the largest and still the least ecologically disturbed of the 
Mascarene islands, but deforestation continues, often in favour of reafforestation 
with Japanese red cedar {Cryptomeria japonica). A number of vertebrates have 
become extinct in the 300 years since human settlement (9) and there is no reason to 
suppose that many invertebrates have not disappeared also. The ability of the larvae 
of P. phorbanta to feed on Citrus trees might be expected to ensure the butterfly's 
survival even in the face of destruction of its natural forest habitats, but three factors 
mitigate against this. Firstly, feeding on Citrus is no insurance against destruction 
because of the stringent insecticidal measures taken against species that are regarded 
as pests of that crop. Secondly, there is a suspicion that the introduction of a parasitic 
tachinid fly 10-15 years ago to control the caterpillars of P. demodocus may have also 
taken a toll of P. phorbanta (13). Thirdly, P. demodocus itself may competitively 
exclude P. phorbanta from the Citrus groves, as it is suspected of doing for P. manlius 
on Mauritius. Papilio demodocus , like P. demoleus, is an aggressive butterfly that 
actively chases other species (2) . Even were these fears to prove unfounded, it is quite 
unacceptable for wild species to decline so far that they are only able to survive as a 
minor pest of man's agricultural labours. 

Conservation Measures In recognition of the increasing threat to the 
species the French Ministers of Environment and Agriculture declared on 22 August 
1979 that P. phorbanta would henceforth be a protected species (8). The decree 
outlawed the destruction or removal of eggs, caterpillars, pupae and adults and 
forbade trade in the species. Such legislation has served to draw attention to the plight 
of P. phorbanta, but failed to have any impact in the more important matter of 
protecting the butterfly's natural habitat or food plants. In addition, it has not 
encouraged much-needed biological studies, nor facilitated ranching or farming of 
specimens for release into the wild. During the five years since the decree was passed, 
there is no evidence of any conservation effort being made on behalf of this species. 
There are no recent reports of the butterfly's status, but a new survey is currently 
under way (3). 

References 

1. Boisduval, J.B.A.D. de (1833). Faune Entomologique de Madagascare, Bourbon et 
Maurice. Paris. 

2. D'Abrera, B. (1980). Butterflies of the Afrotropical Region. Lansdowne Editions, 
Melbourne, xx + 593 pp. 

3. Dove, H. (1983). In litt., 18 April. 

4. Hancock, D.E. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic 
approach. Smithersia 2: 1-48. 

5. Hancock, D.E. (1983). In litt., 21 June. 

6. Legrand.H. (1959). Note sur la sous-espece nana Ch. OberXhur de Papilio phorban Linne 
des lies Seychelles [Lep. , Papilionidae] . Bulletin de la Societe Entomologique de France 64: 
121-123. 

7. Manders, N. (1908). The butterflies of Mauritius and Bourbon. Transactions of the 
Entomological Society of London 1907: 429-454. 

8. Ministere de Tenvironnement and Ministere de I'agriculture (1979). Liste des insectes 
proteges on France. Journal Officiel 22 August: 93-94. 

343 



Threatened Swallowtail Butterflies of the World 

9. Moutou, F. (1984). Wildlife on Reunion. Oryx 18: 160-162. 

10. Munroe, E. (1961). The classification of the PapiHonidae (Lepidoptera). Canadian 
Entomologist Supplement 17: 1-51. 

11. Paulian, R. and Viette, P. (1968). Faune de Madagascar. XXVll Insectes Lepidopteres 
PapiHonidae. O.R.S.T.O.M. and C.N.R.S., Paris. 97 pp. 

12. Smart, P. (1975). The Illustrated Encyclopedia of the Butterfly World. Hamlyn, London. 
275 pp. 

13. Viette, P. (1983). In litt.,6 July. 



344 



"^^^^2^ 6ot/U-t9-^''^'> n-^-^L Papilio aristophontes 

PapUio (Princeps) aristophontes Oberthur, 1897 INDETERMINATE 

Subfamily PAPILIONINAE Tribe PAPILIONNI 



Summary Papilio aristophontes is endemic to the forests of Anjouan, 
Moheli and Grande Comore, Comoro Islands. So far the impact of subsistence 
agriculture has been mainly in the lowlands, but rapid population growth requires 
that forest reserves be designated in order to protect the wildlife, particularly on 
Grande Comore. The butterflies still survive in upland forests, but research is needed 
to study the biology and distribution of the species and to locate suitable areas of its 
habitat for protection. 

Description Papilio aristophontes was originally described as a full species 
but was subsequently considered to be a subspecies of P. (P.) nireus, albeit very 
distinct (1, 5). D'Abrera, following Carcasson, recently reinstated it to full species 
rank (3), a recognition which concurs with the views of lepidopterists familiar with the 
butterfly in the wild {2,6). P. aristophontes is in the nireiis group (4), and the male and 
female differ somewhat in appearance. The male is black with iridescent blue 
markings and a brown lower surface, whilst the female is brown and olive-green. The 
forewing length is 48-52 mm, and the wingspan is 80-90 mm. The young stages are 
unknown. 

Male: UFW black with an iridescent blue median band and five blue spots. 

UHW also black with an iridescent blue band, a row of blue submarginal spots, a 
scalloped margin to the hindwing and virtually no tail. 

LFW/LHW. The LFW is entirely blackish-brown; LHW red-brown with black 
veins and a narrow cream submarginal band with a silver reflection. 

Female: UFW/UHW brown with a dull olive green median band and a submarginal 
band of yellowish-olive lunules, continuous over both wings. 

LFW/LHW red-brown with a silvery submarginal band continuous over both 
wings; a fawn brown median band to the hindwing with pinkish-grey scales on the 
outside, and a white abdominal spot. 

Distribution Papilio aristophontes is endemic to the forests of the Comoro 
Islands, between Mozambique and Madagascar. The main stronghold is Grande 
Comore, but there are also populations on Moheli and Anjouan (2). 

Habitat and Ecology The general ecology of the four Comoro Islands has 
been briefly described in the review of Graphium levassori. Papilio aristophontes is 
another inhabitant of the forests of La Grille (1087 m) and Karthala (2560 m), the 
two volcanic regions of Grande Comore. In 1980 and 1983 it was seen flying at 
Oussoudjou and other places in the massif of La Grille (7, 8) and at Nioumbadjou 
(550 m) (8), Hantsongoma (1000 m) and other localities up to 1700 m (6) on 
Karthala. The species may be locally abundant , flying along the edges of clearings and 
paths (7). The larval feeding habits have not been documented but, like the other 
members of the nirens species group, the caterpillars probably feed on wild Rutaceae 
(4). They have also been reported on wild-growing trees of the domesticated lemon 
(2). 

Threats Although slash and burn agriculture is comparatively uncommon 
in the Comoros, the forests on Karthala and more particularly on La Grille are 
retreating from a more insiduous threat, the prevention of forest regeneration by the 

345 



Threatened Swallowtail Butterflies of the World 

planting of a dense understorey of banana trees. Bananas are one of the staple crops 
of the rapidly growing population of the Comoros. The main agricultural areas are 
coastal and lowland, but population pressure is forcing agricultural development to 
altitudes of up to 1200 m (7), where bananas, guavas, vanilla and ylang ylang 
{Cananga oderata) can still be grown. On the smaller islands of Anjouan and Moheli 
there is much less high ground than on Grande Comore and therefore agriculture is 
even more extensive. The distinctive population of P. aristophontes on Moheli is 
under serious threat from agricultural conversion and intensification (2). 

Conservation Measures Despite increasing modification of its habitat, P. 
aristophontes still survives in good numbers on La Grille and Karthala (Grande 
Comore). However, the species is more seriously threatened by loss of habitat on 
MoheH and Anjouan (2). Throughout the Comoros the human population is growing 
rapidly, and it is inevitable that the threats to this and other endemic forest species 
will gradually increase. At the moment there are no protected areas on the Comoro 
Islands and it is clearly necessary to designate representative reserves at the earliest 
opportunity. Already the forests of La Grille are seriously damaged, and suitable 
areas of undisturbed middle altitude forest is hard to find on Karthala (8). 
Nevertheless, some low altitude forest could be set aside for natural regeneration. As 
proposed in the review of Graphium levassori. all land over 1200 m altitude on 
Karthala might be given protected status. This would inconvenience few people and 
would ensure the survival of many forest species (8). 

Conservation measures specifically for Papilio aristophontes should be incorpo- 
rated into efforts to conserve representative biotopes. Studies of its distribution and 
reproductive biology are needed. 

References 

1. Carcasson, R.H. (1960). The swallowtail butterfliesof East Africa (Lepidoptera, Papilioni- 
dae) . Journal of the East African Natural History Society Special Supplement 6: 33 pp. + 11 
pi. (Reprinted by E.W. Classey, Faringdon, 1975). 

2. Collins, S. (1983). In litt.. 12 July. 

3. D'Abrera, B. (1980). Butterflies of llie Afrotropical Region. Lansdowne Editions, Mel- 
bourne. XX + 593 pp. 

4. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian 
Entomologist S\ipp\ement 17: 1-51. 

5. Paulian, R. and Viette, P. (1968). Faune de Madagascar. XXVII Insectes Lepidopteres 
Papilionidae. O.R.S.T.O.M. and C.N.R.S., Paris. 97 pp., 19 pi. (2 col.), 34 figs. 

6. Turlin, B. (1983). In litt., 1 July. 

7. Turlin, B. (1983). In litt., 15 September. 

8. Viette, P. (1980). Mission lepidopterologique a la Grande Comore (Ocean Indien 
occidental). Bulletin de la Societe Entomologique de France 85: 226-235. 



346 



Papilio desmondi teita 
Papilio (Princeps) desmondi teita van Someren, 1960 ENDANGERED 

Subfamily PAPILIONINAE Tribe PAPILIONINI 



Summary The Taita (Teita) Hills in southern Kenya contain a number of 
endemic plants, birds, reptiles, amphibians and insects, including Papilio desmondi 
teita , and are the type localities for many more . During this century deforestation has 
been extensive due to increasing population pressure, and only relict patches of 
natural forest remain. P. d. teita and many other forest species are confined to these 
relict forests, which should be protected from further interference. 

Description When it was first found, this species in the nireus species group 
(13) was thought to be a race of Papilio brontes desmondi (17), now known simply as 
Papilio desmondi. Later, when more material from the Usambara Mountains and the 
Chyulu Hills was available, the Taita specimens were noted to be distinctive in wing 
pattern, although close to P. desmondiin the form of the genitalia (18). After much 
confusion, it was considered sufficiently different from its neighbours to be described 
as a full species, Papilio teita. Subsequently the species was questioned as being 
similar to P. nireus (15), but that superficial examination took no account of the 
totally different genitalia. In recent studies, D'Abrera (5) and Carcasson (1) reduced 
teita to a subspecies of Papilio desmondi, a course which is followed here. 

As in other blue-banded papilios, the overall wing ground colour is black, with a 
broad median band of blue across the UFW and UHW and rows of blue submarginal 
spots, larger on the UHW (Plate 7.6). The lower side bears a characteristic 
continuous cream submarginal band on the LHW, fading towards the leading edge of 
the LFW. Also characteristic are the squat forewings and broad-base triangular UFW 
median area. Length of forewing 42 mm, hindwing40 mm (18). The larvae have not 
been described but are probably green in the later stages. 

Distribution Papilio desmondi teita is confined to the Taita (Teita) Hills in 
Taveta District, coast Province, Kenya. It has been found in forest on Mt Mbololo, 
Ngangao, Ronge, Chawia and Bura Bluff (4, 5, 14, 16). It was also once recorded 
12 km away on Mt Sagala ( 18) , but this observation has not been repeated. The forest 
on Ngulia Hill in Tsavo West was visited by N.M. Collins and M.P. Clifton in July 
1983 and appeared to be largely unsuitable for the butterfly. 

Habitat and Ecology The Taita Hills consist of an almost circular inselberg 
of 27 km diameter with three main peaks, Vuria (2228 m), Ngangao (2149 m, also 
recorded as Ngao Ngao or Ngoa Ngoa (5)) and Mbololo (2209 m) (also known as 
Mraru). The hills lie about 1 10 km east-south-east of Mt Kilimanjaro and 25 km west 
of Voi, overlooking to the north, east and west the plains of Tsavo West National 
Park 1000 m below. As Moreau (12) has pointed out, the name Taita Hills is a 
ridiculous diminutive for such an imposing massif. 

The importance of these hills lies in the probability that montane rainforest has 
grown on them for tens of thousands of years. During the last Ice Age, when eastern 
Africa was probably (but not certainly (12)) much drier than it is today (10), all forests 
were severely reduced in size. However, relict patches remained and acted as vital 
refugia for forest species of fauna and flora. Such refugia were probably found on a 
string of mountains through the Usambaras and Ulugurus to Mt Mlanje in Malawi 

347 



Threatened Swallowtail Butterflies of the World 

(21). Each montane forest block has therefore tended to evolve a unique flora and 
fauna, isolated as they are from other patches of forest. 

The Taita Hills include both upper montane evergreen rain forest and lower 
montane woodland (6), ahhough their extent is now severely limited (see Threats). 
Only the former is suitable habitat for the butterfly, consisting of closed canopy forest 
with camphorwood {Ocotea usambarensis) as a common tree (6). A list of trees is 
given by Dale (6), who does not, however, mention the two endemics of the Taita 
Hills, Memecylon teitense (Melastomataceae)