Threatened Swallowtail Butterflies of the World Tn@ IUCN [eked DalaBoo coun Saal ee Tee se Pics OF) PAPILIO INT. 3 (4), 1986; 207 - 226 ISSN 0109-7008 date of publication, July 1987 207 PAPILIO HIPPONOUS IN THE IUCN SWALLOWTAIL RED DATA BOOK, A CORRECTION J. Haugum 6 N. M. Collins Owing to an unfortunate misunderstanding the ranges of P. (Princeps) hipponous Felder & Felder 1862 and P. (Princeps) pitmani Elwes & de Nicéville 1886 given in the IUCN’ Red Data Book "Threatened Swallowtail Butterflies of the World" (Collins 6& Morris, 1985) were mixed up. The correct entries are as follows: (per (uae ook > oo, 10 4 Digitized by the Internet Archive in 2010 with funding from UNEP-WCMC, Cambridge http://www.archive.org/details/threatenedswallo85coll Threatened Swallowtail Butterflies of the World The IUCN Red Data Book INTERNATIONAL UNION FOR CONSERVATION OF NATURE AND NATURAL RESOURCES Threatened Swallowtail Butterflies of the World The IUCN Red Data Book. N. MARK COLLINS IUCN Conservation Monitoring Centre, Cambridge, U.K. MICHAEL G. MORRIS Chairman, IUCN/SSC Butterfly Specialist Group Institute of Terrestrial Ecology, Wareham, U.K. With the help and advice of the network of IUCN Commissions and other experts throughout the world. Photographic plates by N. MARK COLLINS PUBLISHED BY IUCN, GLAND, SWITZERLAND AND CAMBRIDGE, U.K. 1985. IVECN IUCN (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, [UCN 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. IUCN 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 IUCN, 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. CITIBANK®> © 1985 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 IUCN Red Data Book. 1UCN, Gland and Cambridge. vii + 401pp. + 8 pls. 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, Smee U.K. Contents How to use this book Acknowledgements The swallowtail butterflies: an introduction to the family and its conservation Swallowtails of the world: their nomenclature,distribution and conservation status Swallowtails of the world: an annotated species list Geographical index to species Analysis of critical faunas Trade in swallowtail butterflies Reviews of threatened species BARONIINAE 1. Baronia brevicornis PARNASSIINAE Parnassiini 2. Parnassius autocrator 3. Parnassius apollo Zerynthiini 4. Bhutanitis mansfieldi 5. Bhutanitis thaidina 6. Bhutanitis ludlowi 7. Luehdorfia chinensis 8. Luehdorfia japonica PAPILIONINAE Leptocercini 9. Teinopalpus imperialis 10. Teinopalpus aureus 11. Eurytides (Protesilaus) marcellinus 12. Eurytides (Protesilaus) lysithous harrisianus 13. Eurytides (Eurytides) iphitas 14. Graphium (Pathysa) epaminondas 15. Graphium (Pathysa) idaeoides 16. Graphium (Pathysa) megaera 17. Graphium (Arisbe) aurivilliusi 18. Graphium (Arisbe) weberi 19. Graphium (Arisbe) levassori Page Number —_— Contents V1 20. Dike DON 23. 24. Troidini WS. 26. Dil. 28. 29. 30. Bie 32. 33. 34. 35. 36. Bie 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. Graphium (Graphium) procles Graphium (Graphium) meeki Graphium (Graphium) stresemanni Graphium (Graphium) sandawanum Graphium (Graphium) mendana Battus zetides Parides ascanius Parides hahneli Parides pizarro Parides steinbachi Parides coelus Parides klagesi Parides burchellanus Atrophaneura (Atrophaneura) luchti Atrophaneura (Losaria) palu Atrophaneura (Pachliopta) jophon Atrophaneura (Pachliopta) schadenbergi Atrophaneura (Pachliopta) atropos Troides (Troides) andromache Troides (Troides) prattorum Troides (Troides) dohertyi Ornithoptera tithonus Ornithoptera rothschildi Ornithoptera chimaera Ornithoptera paradisea Ornithoptera meridionalis Ornithoptera alexandrae Ornithoptera aesacus Ornithoptera croesus Papilionini 49. 50. Syl 525 D3 54. 55). 56. Sie 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. Papilio (Pterourus) homerus Papilio (Heraclides) esperanza Papilio (Heraclides) aristodemus ponceanus Papilio (Heraclides) caiguanabus Papilio (Heraclides) aristor Papilio (Heraclides) garleppi Papilio (Heraclides) himeros Papilio (Heraclides) maroni Papilio (Chilasa) maraho Papilio (Chilasa) osmana Papilio (Chilasa) carolinensis Papilio (Chilasa) toboroi Papilio (Chilasa) moerneri Papilio (Papilio) hospiton Papilio (Princeps) benguetanus Papilio (Princeps) acheron Papilio (Princeps) jordani Papilio (Princeps) weymeri Papilio (Princeps) sjoestedti Papilio (Princeps) manlius 224 226 228 231 234 236 240 242 244 246 248 250 252 254 256 258 261 263 266 269 271 273 276 278 281 284 288 292 294 297 299 301 305 307 309 311 313 315 317 320 822 324 326 329 331 333 335 337 339 69. 70. TAL WZ: 1B 74. 12% 76. Tike 78. Papilio (Princeps) phorbanta Papilio (Princeps) aristophontes Papilio (Princeps) desmondi teita Papilio (Princeps) antimachus Papilio (Princeps) morondavana Papilio (Princeps) grosesmithi Papilio (Princeps) leucotaenia Papilio (Princeps) mangoura Papilio (Princeps) chikae Papilio (Princeps) neumoegeni Appendices A. List of Papilionidae in threatened categories B. Papilionidae that require further monitoring and research Index Plates Contents 342 345 347 351 354 356 358 362 364 367 369 371 375 Vii Bh - Maetden i: I wah ive sis in ry ‘ 4 -saiprechera iene ope hog, con on At prunesice rn or a ae | es Pe depen never , oe a) i 7 ? | se wire ety “eae yy : ae -— . , ; Ps of a. ae oe Pe oe balsas: ieade 7 ris : a 7 ee A 7 =e ."s A ast elw 7 a ee (LC SO ie gh a i) ee “te en ut ee eo a =a Loe 7 ne re yas Tus en . * a m t p ; ¥ r =" Danhhen 2-apalbenn wiper den 9 eee : 7 “ Pooeiia Up oral ve) La wil ee Te = 7 ices Wie ’ “jut freA >) Amp ratte | D ia) . hs an i) Oa i ie wre ite irate irvates pie Deo gga” Saale lens | HO a oaptet staan 7 e i desi, pn iy Baie. f 77, re A, *y Ra Mee eh ae ee - By Aan ais. anon plate A ea it 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 list 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 [UCN 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 IUCN 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 2 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 unlikely 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 (1) 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 ier. 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. 4 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. 5 in A XB 10 =| Sy 5 aes, LESS, V UFW: _ upper forewing LFW: lower forewing UHW: — upper hindwing LHW: _ lower hindwing Mi submarginal 8. outer margin 2. postdiscal oF anal or inner margin 1&2. distal 10. tail 3h discal iB cell 4. basal 12: apical 5: apex 15) subapical 6. anal angle or tornus 14. subcostal if costa or costal margin 15%, median YES) The following terms apply particularly to Ornithoptera males 16. radial band ie 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 IUCN/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 IUCN Conservation Monitoring Centre and the Institute of Terrestrial Ecology, Furzebrook, who gave enthusiastic encouragement and helpful advice on many matters. Michael Morton of Brunel 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 P.R. Ackery S.A. Ae H.S. Barlow M.J.C. Barnes A. Bedford—Russell M.J. Braga F.M. Brown K.S. Brown Jr. J.P. de Carvalho P.B. Clark Sir C.A. Clarke M.P. Clifton M.J. Coe S. Collins T.C.E. Congdon G.A. Dacasin E.W. Diehl P. Dollinger F.M. Dori M.J. Dourojeanni H. Dove K. Eltringham R.D. Estes P. Feeny T.L. Fenner R.W. Garrison M. Ghilarov A.K. Ghosh S.K. Ghosh H.C.J. Godfray R. Guilbot N. Gunatilleke D.L. Hancock J. Haugum T. Hidaka J.D. Holloway W. Hsu A.F. Hutton M. Ishii J.N. Jumalon J. Kielland J. Kingdon Kim Hon Kyu G. Lamas T.B. Larsen R.R.B. Leakey R.E. Lewis D. Lowe R. Macfarlane D.K. Mandall A.G. Marshall J.de la Maza E. O.H.H. Mielke J.S. Miller L.D. Miller T. Milliken J. Nabhitabahata T.R. New D. N’Sosso A.U. Oboite S.N. Okiwelu P.A. Opler G.W. Otis D.F. Owen M.J. Parsons R. Paulian N.D. Penny M.enC.H. Pérez A. Pinratana R.M. Pyle C.D. Quickelberge T. Racheli J. Riley A.H.B. Rydon T. Saigusa R.L. Smiles D. Spencer-Smith T.T.Struhsaker B. Turlin T.W. Turner A. Tye the late H.Tyler R.I. Vane—Wright L. Vazquez G. P. Viette M.V. Walker B.M. Wankhar D. Wankhar P.G. Waterman J.D. Weintraub J. G. Williams M.L. Winston A. Youdeowei 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 Z, 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 (Meandrusa), 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 7 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 aslit 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 8 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 Ornithoptera 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 Eurytides 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 2.1) (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 60—70°N and 40—S0°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 9 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 Islands Papilio phorbanta f. Papilio dardanus f. Papilio dardanus f. Papilio dardanus f. Papilio dardanus f. Papilio cynorta f. Papilio cynorta f. Papilio echerioides f. 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 Papilio rex Danaus formosa South America Eurytides thymbraeus aconophos Parides alopius Eurytides thymbraeus Parides photinus Eurytides branchus Parides arcas mylotes Eurytides belesis Parides polyzelus Eurytides euryleon Parides iphidamas phalias Eurytides euryleon clusoculis Parides lycimenes Eurytides pausanias 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 Papilionidae 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)). Latitude® 70 60 50 40 30 20 10 80 80 ” 0 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)). ae a ena 0 10 20 30 40 50 60 Number of species ilil Threatened Swallowtail Butterflies of the World Fig. 2.3. Latitudinal gradients in the species richness of the Indo—Australian and Eastern Asian Papilionidae (adapted from Scriber (53)). DS ee Latitude® | 70 > 60 50 40 30 123 SEE Se ee 0 10 20 30 40 50 60 70 80 90 100 110 120 130 Number of species Fig. 2.4. Latitudinal gradients in the species richness of the Papilionidae of the world. Latitude® 70, 60 50 40 30 20 10 50 0 10 20 30 40 50 60 70 80 90 100110 120 130 140 150 160 170 180 190 200 210 220 230240 250 260 Number of species Wy 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 Nymphalidae. 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 ofa 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 Rauwenhoffia (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 regularly 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 facilitated 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 Sericinus/Bhutanitis (Parnassiinae) in western and eastern Asia respectively, Eurytides (Protesilaus), Papilio (Pterourus) 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 delicate 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 acommon 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 Magnoliales (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 Magnoliales 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 ina 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 linked 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 IUCN 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 manilius 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 Réunion, 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 Eurytides 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/hain Burma to 90 cu. 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 swallowtai!s are more widespread in their distribution. However, deforestation throughout its range, but particularly in India and Nepal, is placing the Kaiser-I-Hind, Teinopalpus imperialis, 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. Eurytides 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 fertilizers. With the very limited 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 mote 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 Eurytides 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 Réunion isa 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 Réunion 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 published 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 IUCN 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 valid 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 1(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 IUCN/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-listing as Endangered under the Endangered Species Act. Schaus’ Swallowtail has been the subject 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 IUCN was divided into a Butterfly Specialist 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 in litt., 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 4: 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 Seiler, W. (1979). Biomass burning as a source of atmospheric gases CO, H,, Nx0, NO, CH3Cl 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. Part 1. Papilionidae and Pieridae. Hill House, Victoria, Australia. xxxi + 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 21. 22. 23. 24. 25: 26. Alle 28. 29. 30. SH 32. 33: 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. Swallowtail introduction Ehrlich, P.R. and Raven, P.H. (1964). Butterflies and plants: A study in coevolution. Evolution 18: 586-608. Eisner, T. and Meinwald, Y.C. (1965). Defensive secretion of a caterpillar (Papilio). Science 150: 1733-1735. Eisner, T., Pliske, T.E., Ikeda, M., Owen, D.F., Vazquez, L., Pérez, H., Franclemont, J.G. and Meinwald, J. (1970). Defense mechanisms of arthropods. XX VII. Osmeterial secretions of papilionid caterpillars (Baronia, Papilio, Eurytides). Annals of the Entomolo- gical Society of America 63: 914-915. FAO/UNEP (1981). Tropical Forest Resources Assessment Project. (3 volumes). FAO, Rome. 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. 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. Fraenkel, G.S. (1959). The raison d’étre of secondary plant substances. Science 129: 1466-1470. Global Environment Monitoring System, UNEP (1982). The global assessment of tropical forest resources. GEMSPAC Information Series No. 3, 14 pp. Groombridge, B. and Wright, L. (1982). The IUCN Amphibia—Reptilia Red Data Book. Part 1. 1UCN, Gland. xliii + 426 pp. Hancock, D.L. (1983). Classification of the Papilionidae (Lepidoptera): a phylogenetic approach. Smithersia 2: 1-48. Heath, J. (1981). Threatened Rhopalocera (Butterflies) in Europe. Nature and Environ- ment Series No. 23. Council of Europe, Strasbourg. IUCN Commission on Ecology (1984). Population and Natural Resources. Commission on Ecology Occasional Paper Number 3, 12 pp. IUCN/UNEP/WWE. (1980). World Conservation Strategy. 1UCN, Gland. Igarashi, S. (1979). Papilionidae and Their Early Stages. Vol. 1219 pp., Vol. 2 102 pp. of plates. Kodansha, Tokyo. (In Japanese). Larsen, T. (1984). Butterflies of Saudi Arabia and its Neighbours. Stacey International, London. 160 pp. 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. Lucas, G. and Synge, H. (1978). The IUCN Plant Red Data Book. Y\UCN, Gland. 540 pp. Miller, L.D. and Brown, F.M. (1981 (83)). Butterfly taxonomy: a reply. Journal of Research on the Lepidoptera 20: 193-198. 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. Moriarty, F. (1975). Pollutants and Animals. Chapter 6: Where have all the butterflies gone? Pp. 81-97. Allen and Unwin, London. ; Morris, M.G. (1981). Conservation of butterflies in the United Kingdom. Beiheft Veroffentlichung Naturschutz Landschaftspflege Baden—Wiirttemberg 21: 35-47. Munroe, E. (1961). The classification of the Papilionidae (Lepidoptera). Canadian Entomologist Supplement 17: 1-51. Munroe, E. and Ehrlich, P.R. (1960). Harmonization of concepts of higher classification of Papilionidae. Journal of the Lepidopterists’ Society 14: 169-175. Myers, N. (1979). Conversion Rates in Tropical Moist Forests. Report to National Academy of Sciences. National Research Council, Washington D.C. 205 pp. National Research Council (1983). Butterfly Farming in Papua New Guinea. Managing Tropical Animal Resources Series. National Academy Press, Washington D.C. 30 pp. Nikusch, I. (1981). Die Zucht von Parnassius apollo Linnaeus mit jahrlich zwei Generationen als Moglichkeit zur Erhaltung bedrohter Populationen. Beiheft Veréffentli- chung Naturschutz Landschaftspflege Baden—Wiirttemberg 21: 175-176. Owen, D.F. (1971). Tropical Butterflies. Clarendon Press, Oxford. 214 pp. 31 Threatened Swallowtail Butterflies of the World 49. 50. 51. 52. 53: 54. 55. 56. Sil: 58. 59: 60. 61. 62. 63. 64. 32 Troides (Lepidoptera: Papilionidae) in Papua New Guinea. Final report to the Depart- ment of Primary Industry, Papua New Guinea. 111 pp. Pérez 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 México 42, Ser. Zool. (1): 63-72. Poore, D. (1976). Ecological Guidelines for Development in Tropical Rain Forests. 'UCN, Gland. 39 pp. Pyle, R.M. (1984). The Audubon Society Handbook for Butterfly Watchers. Charles Scribner’s Sons, New York. 274 pp. Pyle, R.M., Bentzien, M. and Opler, P. (1981). Insect conservation. Annual Review of Entomology 26: 233-258. Scriber, J.M. (1973). Latitudinal gradients in larval feeding specialization of the world Papilionidae (Lepidoptera). Psyche 80: 355-373. Seiler, 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. 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 Slansky, F. (1972). Latitudinal gradients in species diversity of the New World swallowtail butterflies. Journal of Research on the Lepidoptera 11(4): 201-218. Smart, P. (1975). The Illustrated Encyclopedia of the Butterfly World. Hamlyn, London. 275 pp. Thornback, J. and Jenkins, M. (1982). The IUCN Mammal Red Data Book. Part 1.1UCN, Gland. xl + 516 pp. UNDP/FAO National Parks Development Project (1981/1982). National Conservation Plan for Indonesia. Vols 1-8. FAO, Bogor. Vane-Wright, R.I. (1976). A unified classification of mimetic resemblances. Biological Journal of the Linnean Society 8: 25-56. Wells, S.M., Pyle, R.M. and Collins, N.M. (1983). The IUCN Invertebrate Red Data Book. IUCN, Cambridge and Gland. L + 632 pp. Whalley, P. (1980). Butterfly watching. Severn House, London. White, F. (1981). The history of the Afromontane archipelago and the scientific need for its conservation. African Journal of Ecology 19: 33-54. 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. Swallowtails of the world e 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 common 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 list 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 Order: Swallowtails of the world LEPIDOPTERA (Butterflies and moths) Family: PAPILIONIDAE (Swallowtail butterflies) Subfamily: BARONIINAE Genus: Baronia Salvin Baronia brevicornis Salvin, 1893 Rare—refer to section 6, p. 182. Restricted to a very small area of Mexico and particularly important 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 Hibner Archon apollinus (Herbst, 1798) 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 Ménétriés Hypermnestra helios (Nickerl, 1846) A narrow range in Afghanistan, Pakistan (Baluchistan), Iran and U.S.S.R. (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. Genus: Parnassius Latreille Species-group: szechenyii Parnassius szechenyii Frivaldszky, 1886 Western China (north-eastern Xizang Zizhiqu (Tibet), Qinghai, Gansu, 35) Threatened Swallowtail Butterflies of the World 36 10 Sichuan (Szechwan) and Yunnan). No threats known but apparently rather rare (264). Possibly eleven distinguishable subspecies (265). Other refs: LSI AGo 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. 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 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. /atonius) (80). Other refs: 1, 4, 31, 71, 74, 169, 259, 264. 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). Parnassius patricius Niepelt, 1911 U.S.S.R. (Kirghizia); a rather narrow range but not known to be threat- ened. Five subspecies (265). Other refs: 1, 31. 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. 11 12 13 14 iS 16 17 Swallowtails of the world Species-group: imperator Oberthir Parnassius imperator Oberthir, 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. augustus 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: 1a ol 200. Species-group: charltonius Gray 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. Parnassius inopinatus Kotzsch, 1940 Found only in mountain ranges in north-western Afghanistan where it is very localised (264). Little is known about it, butit 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. Parnassius loxias Pingeler, 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. 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 inthe 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 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 Parnassius acco Gray, 1853 37 Threatened Swallowtail Butterflies of the World 38 18 19 20 21 22 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. Parnassius przewalskii Alpheraky, 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). Parnassius hannyngtoni Avinoff, 1916 North-eastern India (Sikkim), China (Xizang Zizhiqu (Tibet)) and Bhutan (2). 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 amended 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 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 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 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. 23 24 25 26 2 28 Swallowtails of the world Parnassius clodius Ménétriés, 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 listed 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. Parnassius eversmanni Ménétriés, 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. 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. Parnassius ariadne Lederer, 1853 (= clarius Eversmann, 1843) U.S.S.R (Altay mountains and Tadzhikistan) and western Mongolia. Not known to be threatened. Localized in distribution (264) and rare in collections. Two subspecies (265). Other refs: 1, 31, 169. 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. 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 its range in the U.S.S.R. and listed in the U.S.S.R. Red Data Book (8, 260). 39 Threatened Swallowtail Butterflies of the World 40 29 30 31 32 33 34 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. Parnassius stubbendorfi Ménétriés, 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. 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 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. 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. Parnassius bremeri Bremer, 1864 Eastern U.S.S.R., northern China (Heilongjiang, Hebei and Shanxi), North Korea, South Korea where it 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 overcollection 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. Parnassius jacquemontii Boisduval, 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. 35 36 37 38 39 Swallowtails of the world Parnassius epaphus Oberthir, 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. Parnassius actius (Eversmann, 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 Shy 745 1691239. Parnassius phoebus (Fabricius, 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. Parnassius tianschanicus Oberthir, 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. 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 Mongolia) (?)), 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 of 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 listed 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 declining 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: PARNASSITINAE Tribe: Zerynthiini 42 41 Genus: Sericinus Westwood Sericinus montela Gray, 1843 (= telamon Donovan, 1798) Eastern U.S.S.R. (Primorskiy Kray), China (Heilongjiang, Jilin, 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 43 44 45 46 47 Swallowtails of the world species of Zerynthia found in Asia Minor. This course has been followed by Hancock (109). Genus: Allancastria Bryk 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. Allancastria deyrollei Oberthtr, 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. 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 of A. cerisy by most authors (31, 141). Larsen considers it to be distinct (160) as does Turlin (264). Conservation status needs clarification. 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 Zerynthia polyxena (Denis & Schiffermiiller, 1775) 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. 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 44 48 49 50 51 52 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 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. 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. 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. Bhutanitis ludlowi Gabriel, 1942 Insufficiently Known—refer to section 6, p. 196 Bhutan: Trashiyangsi Valley only. The type-series is unique (1). Other ref: OL. Genus: Luehdorfia Criiger 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. 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 53 54 Swallowtails of the world stages. The above status seems appropriate until its specific identity and possible threats can be assessed. Two subspecies (141). 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 of 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. 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: 55 56 57 Leptocircini Genus: Iphiclides Hibner 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 protected in seven of them, and Indeterminate throughout Europe (123). Six subspecies including J. 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 J. podalirius (125). Common names: Scarce Swallowtail (125, 253, 282), Sail Swallowtail (253), Pear-tree Swallowtail (158). Other refs: 141, 202. Iphiclides podalirinus (Oberthiir, 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 46 58 59 60 61 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. 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 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 Jachinus 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. Meandrusa payeni (Boisduval, 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. 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 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. 62 63 64 65 66 67 68 69 70 Swallowtails of the world Eurytides (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. 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. 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. 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: Di Sls 55234- 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: Sle a02 alia pease Eurytides (Protesilaus) anaxilaus (C. & R. Felder, 1864) (=arcesilaus Lucas, 1852) Northern Venezuela and Colombia. Not known to be threatened (234). Other ref: 51. 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: $1559 3234: 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 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 48 71 72 73 74 18 76 77 78 79 80 81 Species-group: protesilaus Linnaeus Eurytides (Protesilaus) agesilaus (Guérin 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. 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. 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. 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. 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. 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. Eurytides (Protesilaus) aguiari (D’ Almeida, 1937) Brazil, from Belém (Para) to Benjamin Constant (Amazonas). No known threats but more data required. Refs: 51, 55. Eurytides (Protesilaus) embrikstrandi (D’ Almeida, 1936) Brazil. A little known and fairly recently described species. No threats known. Refs: 51, 55. Eurytides (Protesilaus) travassosi (D’ Almeida, 1938) Brazil. Another little known and relatively recently described species. Not known to be threatened Refs: 51, 55. 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. 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) 82 83 84 85 86 87 88 89 90 91 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). 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- liaceae (115). 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. 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 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. Eurytides (Protesilaus) microdamas (Burmeister, 1878) Paraguay and adjacent areas of Argentina and Brazil. Little known but no threats recognized. Refs: 51, 55, 122, 234. 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. 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. 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. Eurytides (Protesilaus) ilus Fabricius, 1793 Colombia, Panama and northern Venezuela. Rare and little known but not recognized as threatened. Refs: 51,55, 234. 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 50 92 93 94 95 96 97 98 99 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). Eurytides (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. 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. 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. 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. Eurytides (Protesilaus) xynias (Hewitson, 1875) Ecuador, Bolivia and Peru. Not uncommon and not threatened. Two subspecies. Refs: 51, 55, 234. Eurytides (Protesilaus) phaon (Boisduval,1836) Mexico, Guatemala, Honduras, Belize, Nicaragua, Costa Rica (likely, 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. 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. 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. Eurytides (Protesilaus) protodamas (Godart, 1819) Southern Brazil, Paraguay and Argentina. Not known to be threatened. 100 101 102 103 104 105 106 107 108 109 Swallowtails of the world Two forms, not regarded as subspecies. Ranched in Brazil (24). Other refs: S555; 122; 234: Species-group: thyastes Drury Eurytides (Eurytides) 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. Eurytides (Eurytides) thyastes (Drury, 1782) Eastern Ecuador, Peru, Bolivia and Brazil. Not uncommon and not threat- ened. Ranched in Brazil (24). Three subspecies. Other refs: 51,55, 234. 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. Eurytides (Eurytides) leucaspis (Godart, 1819) Ecuador, Colombia, Peru and Bolivia. Common and not threatened. Two subspecies. (222). Other refs: 51, 55, 234. Eurytides (Eurytides) lacandones (Bates, 1864) Mexico, Central America, Colombia, Ecuador, Peru and Bolivia. Possibly conspecific with E. (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. 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 Eurytides (Eurytides) serville (Godart, 1824) Colombia, Venezuela, Brazil, Ecuador, Peru and Bolivia. Possibly conspe- cific with E. (E.) columbus (222). Common and not threatened. Two subspecies. Other refs: 51, 55, 234. Eurytides (Eurytides) columbus (Kollar, 1850) Colombia, north-western Ecuador and possibly Venezuela. Possibly con- specific with FE. (E.) serville (222). No known threats. Other refs: 51955234: 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. 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 52 110 111 112 113 114 115 116 Eurytides (Eurytides) callias (Rothschild and Jordan, 1906) Eastern Ecuador and Peru. Not known to be threatened and not uncom- mon. Refs: 51, 55, 234. Eurytides (Eurytides) 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. Eurytides (Eurytides) 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 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 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. 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 Species-group: alebion Gray Graphium (Pazala) eurous (Leech, 1892-94) 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 117 118 119 120 121 122 123 124 125 Swallowtails of the world Kashmir, and the Himalayas). Common name: Six-bar Swordtail (80, 251). Other refs: 4, 52, 141, 259. Graphium (Pazala) mandarinus (Oberthir, 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. 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. Graphium (Pazala) tamerlanus (Oberthiir, 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 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. 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. 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. 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. Graphium (Pathysa) androcles (Boisduval, 1836) Indonesia (Sulawesi and Sula (262)). Apparently not uncommon and not known to be threatened. Two subspecies. Other ref: 121. Graphium (Pathysa) epaminondas Oberthiir, 1879 Insufficiently Known—refer to section 6, p. 213. 53 Threatened Swallowtail Butterflies of the World 54 126 127 128 129 130 131 132 133 Confined to the Andaman Is; distribution within the group poorly known. Although 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). 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). 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 (2). 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. 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. 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 Graphium (Pathysa) phidias (Oberthiir, 1896) Vietnam, Laos (?). Recorded from only a small area and possibly Vulnerable but very little information available. Refs: 52, 262. Graphium (Pathysa) encelades (Boisduval, 1836) Indonesia (Sulawesi). Not uncommon and not recognized as being threat- ened. Refs: 52, 262. Graphium (Pathysa) idaeoides (Hewitson, 1853) Rare—refer to section 6, p. 215. 134 135 136 137, 138 139 140 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 Jdea leuconoe. Much prized by Japanese and other collectors (274). Other refs: 52, 147, 148, 262. Graphium (Pathysa) delesserti (Guérin, 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. 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. 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. Graphium (Pathysa) ramaceus (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. 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, Bali 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. 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. Graphium (Pathysa) stratocles (C. & R. Felder, 1862) Philippines (Luzon, Mindoro, Marinduque, Bohol, Mindanao, Calamians and Palawan) (52, 262). Not uncommon and not known to be threatened. Three subspecies. 55 Threatened Swallowtail Butterflies of the World 56 141 142 143 145 146 147 148 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. 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 Jdeopsis juventa and Tirumala hamata. One subspecies with three forms (104). Other ref: 48. Subgenus: Arvisbe Hubner Species-group: angolanus Goeze 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. Graphium (Arisbe) angolanus (Goeze, 1779) (= pylades F., 1793) Southern and tropical Africa: distributed in woodland over most of the continent. Common, often abundant and not threatened. Three subspe- cies. Common names: Angola White Lady Swallowtail (282, 285), White Lady Swallowtail (34, 44, 214). Other refs: 20, 33, 50, oes LAF 165. See note after References. 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. 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 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 Graphium (Arisbe) philonoe (Ward, 1873) Forest and woodland in East Africa: Kenya, Tanzania, southern Sudan, northern Uganda, south-west Ethiopia, Mozambique and Malawi. Als. Connex Lee F Lepudep Soe a ba prO (1 y & a 149 150 151 152 153 154 155 156 157 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 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. 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. 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. 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. 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. 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. 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. 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. 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 58 158 159 160 161 162 163 164 165 threatened. Two or three subspecies. Common name: Milky Graphium (34). Other refs: 33, 50, 285. 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. 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. 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. 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: 2033; 503151, 285. Graphium (Arisbe) simoni (Aurivillius, 1899) Cameroon, Gabon, Congo and Zaire. Possibly a form of G. (A.) ucalegon (250), although 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: J/eonidas Fabricius 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. 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. Graphium (Arisbe) pelopidas (Oberthir, 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 Tomé and 166 167 168 169 170 171 172 Swallowtails of the world Principé 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). 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 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. 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 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. 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. 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. 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 60 173 174 175 176 177 178 (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 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 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. 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. 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 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 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: 2053356397, 165; 214, 215) 179 180 181 182 183 184 185 186 Swallowtails of the world Graphium (Graphium) evombar (Boisduval, 1836) Madagascar, distributed over the sitiale Se Common and not threat- ened. Refs: 33, 34, 50,206. tho Comer Pha >I far hess Species-group: eurypylus Linnaeus 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. 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. Graphium (Graphium) chiron (Wallace, 1865) Nepal, northern India (Assam), Burma, southern China, Thailand, Laos, Vietnam, Kampuchea and Peninsular Malaysia (76). Formerly regarded as asubspecies of G. (G.) bathycles but raised to specific level by Saigusa et al. (237). There is doubt concerning the validity of the name chiron. Hancock (115) considers it to be ahomonym 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. Graphium (Graphium) leechi (Rothschild, 1895) China. Little information available and not known to be threatened. Ref: 188. 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. Graphium (Graphium) meyeri (Hopffer, 1874) Indonesia (Sulawesi and Sula) (262). Not a well-known species, but not considered to be threatened. Two subspecies (52). 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 62 187 188 189 190 191 192 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. 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 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. 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. 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 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. 193 194 195 196 197 198 199 200 Swallowtails of the world 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. 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. 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. 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, Kaiand 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. 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.) codrus (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. 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.) clymenus 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). 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. Graphium (Graphium) monticolum (Fruhstorfer, 1897) Indonesia (southern Sulawesi). Regarded by some as a subspecies of G. (G.) sarpedon (121, 262), but by others asa 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 64 208 209 210 215 216 Swallowtails of the world Species-group: philenor Linnaeus Battus philenor (Linnaeus, 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. Battus zetides Munroe, 1971 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. Battus devilliers (Godart, 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 Battus polydamas (Linnaeus, 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. Battus streckerianus (Honrath, 1884) Peru; uncommon, but no threats known. Refs: 51, 55, 234. Battus archidamas (Boisduval, 1836) 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. 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. Battus philetas (Hewitson, 1869) Northern Peru and Ecuador. Not uncommon and not recognized as threatened. Two subspecies. Refs: 51, 55, 234. 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 Threatened Swallowtail Butterflies of the World 66 217 218 ZA9 220 221 Zoe 223 224 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. 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. 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. 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. 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 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 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. 226 227 228 229 230 231 pV 233 Swallowtails of the world “Genus: Parides Hiibner Species-group: ascanius Cramer Parides gundlachianus (C. & R. Felder, 1864) Restricted to forests in Cuba. Not exceptionally rare but requires moni- toring. Parides columbus (Herrich-Schaeffer, 1862) is a homonym. Common name: Gundlach’s Swallowtail (230). Other refs: 51,55, 234. Parides alopius (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. Parides photinus (Doubleday, 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 photinus. Other refs: 51, 55, 62, 234. 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. Parides phalaecus (Hewitson, 1869) Eastern Ecuador and possibly Peru. Very scarce and perhaps meriting Rare status, but more information required. Refs: 51,55, 234. Parides agavus (Drury, 1782) Brazil, Paraguay and Argentina. Not uncommon and not thought to be threatened. Ranched in Brazil (24). Other refs: 51, 55, 234. Parides proneus (Hubner, 1825) South-eastern Brazil. Not uncommon and not known to be threatened. Refs: 26, 51, 55, 234. Parides ascanius (Cramer, 1775) (= orophobus D’Almeida, 1942) Vulnerable—tefer 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. Parides bunichus (Hiibner, 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 68 234 235 236 237 238 239 240 241 242 243 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 Parides perrhebus (Boisduval, 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. 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. 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. 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. 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. Parides pizarro (Staudinger, 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. 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. 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. 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. 244 245 246 247 248 249 250 251 252 253 Swallowtails of the world Parides aeneas (Linnaeus, 1758) Colombia, Guyana, Surinam, French Guiana, Brazil, Ecuador, Peru and Bolivia. Not uncommon and not known to be threatened. Parides schuppi (Rober, 1927) appears to be a form of Parides aeneas (109). Six subspecies. Other refs: 51, 55, 234. 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. 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. 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. 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. 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. 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. Parides iphidamas (Fabricius, 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. 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. 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 70 254 255 256 257 258 259 260 261 262 263 264 Parides lycimenes (Boisduval, 1870) Mexico, Central America, Colombia, Venezuela and Ecuador. Fairly common and not threatened. Three subspecies. Refs: 55,57, 61, 62, 234, 269. 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. Parides drucei (Butler, 1874) Ecuador, Peru, Brazil and Bolivia. Not uncommon and not threatened. May be a subspecies of P. anchises. Refs: 55, 234. Parides erlaces (Gray, 1852) Ecuador, Peru, Brazil, Bolivia and northern Argentina. Not uncommon nor threatened. Five subspecies. Refs: 51,55, 122, 234. 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. 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. 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: /ysander Cramer Parides panthonus (Cramer, 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. Parides aglaope (Gray, 1852) Brazil, southern Peru and eastern Bolivia. Not common, but far from rare and not threatened. Refs: 26, 51, 55, 234. Parides castilhoi (D’ Almeida, 1967) Brazil (Castilho, Rio Parana, Sao Paulo) (51). More information is required on this little known species. Parides lysander (Cramer, 1775) Colombia, Brazil, Guyana, Surinam, French Guiana, Ecuador and Peru. Often common (55) and not threatened. Two subspecies. Other ref: 51. 265 266 267 268 269 270 271 272 Swallowtails of the world 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. 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. Parides neophilus (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. Parides eurimedes (Stoll, 1780) (= arcas 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), arcas 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. Parides timias (Gray, 1852) Ecuador. Fairly common, despite the restricted range. Two subspecies, possibly conspecific with P. eurimedes. Refs: 51, 55, 234. Genus: Atrophaneura Reakirt Subgenus: Pharmacophagus Haase Species-group: antenor 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: /atreillei Donovan Atrophaneura (Atrophaneura) daemonius (Alpheraky, 1895) China. Very little information available and none on current status. Further research required. Ref: 109. Atrophaneura (Atrophaneura) plutonius (Oberthir, 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 monitcring. The subspecies pembertoni (Pemberton’s Chinese Windmill (80)) and tyéleri (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 72 273 274 275 276 Pid 278 279 280 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 listed 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. 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, A. (A.) /. kabrua is protected by law in India (182). Common name: Rose Windmill (80, 251). Three subspecies. Other refs: 4, 141, 259. Atrophaneura (Atrophaneura) polla (de Nicéville, 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 Nicéville’s Windmill (80, 94). Other ref: 109. Atrophaneura (Atrophaneura) crassipes (Oberthiir, 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. 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. 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. 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. 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.) nevilli and may be a distinct species (52). No information on threats; more research needed. 281 282 283 284 285 286 287 288 289 290 Swallowtails of the world Atrophaneura (Atrophaneura) impediens (Rothschild, 1895) China including Sichuan (Szechwan) (207) and Taiwan. No data on conservation status; more research needed. Two subspecies (52). Atrophaneura (Atrophaneura) febanus (Fruhstorfer, 1908) Restricted to Taiwan. Apparently common, but status should be monitored in view of the limited distribution. D’Abrera lists febanus as a subspecies of impediens but this has not been demonstrated adequately (52). Not listed by Hancock (109). Other refs: 141, 246. 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 of A. (A.) dasarada. 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. 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 /ama (Oberthtr) 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 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. Atrophaneura (Atrophaneura) kuehni (Honrath, 1886) Indonesia (eastern and northern Sulawesi). Little known about status, possibly very rare (121). Itis 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. 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 A. (A.) priapus (118). Other refs: 52, 262. Atrophaneura (Atrophaneura) hageni (Rogenhofer, 1889) Indonesia: restricted to high plateau of Sumatra. Uncommon, but not known to be threatened. Refs: 52, 141. 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 74 291 292 293 294 295 296 297 298 information and close monitoring are needed. Two (52) or three (262) subspecies. Common name: White-head Batwing (3). 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. (A.) priapus). 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. 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. 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. 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. 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). Atrophaneura (Losaria) neptunus (Guérin—Méneville, 1840) Southern Burma, southern Thailand, Peninsular and Eastern Malaysia, Philippines (Palawan), Brunei and Western Indonesia (Sumatra, Nias, 299 300 301 302 303 304 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: ASO 2 ao) Sle 68S 2594262: Atrophaneura (Losaria) palu (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 (114, 121). Believed to be restricted to the west coast where it is rare and localized (172). 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 palu. Common name: Common Clubtail (45, 80, 282). Other refs: 4, 87, 107, 141, 168, 259. 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 facies within the genus. Both genera have in the past been called Polydorus. 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. Atrophaneura (Pachliopta) jophon (Gray, 1852) Vulnerable—refer to section 6, p. 258. Sri Lanka. Declining due to loss of habitat (52,53). A. (P.) pandiyana from southern India is sometimes regarded as a subspecies of A. (P.) jophon. Common name: Ceylon Rose (80). Other refs: 259, 287. 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 76 305 306 307 308 309 310 311 312 313 314 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. 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. 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. Atrophaneura (Pachliopta) schadenbergi (Semper, 1886) Vulnerable—refer to section 6, p. 261. Philippines: Luzon and the Babuyan Is. A locally distributed species with seasonally limited occurrence in lowland and medium elevation wooded areas. Threatened by habitat destruction. Two subspecies. Refs: 52, 177, 262. 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. 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. 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 (1930) strandi from 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 is sabinae (Seyer) (107). Other refs: SAME: 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. 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). Atrophaneura (Pachliopta) aristolochiae (Fabricius, 1775) Afghanistan (118), India, Nepal, Sri Lanka, Burma, Thailand, southern and eastern China (including Hainan (Guangdong prov.)), Hong Kong, 315 316 317 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. Atrophaneura (Pachliopta) polydorus (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. 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. 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. 77 Threatened Swallowtail Butterflies of the World 78 318 319 320 321 322 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 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 Hubner Species-group: amphrysus Cramer Troides (Troides) cuneifer (Oberthtir, 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. 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. 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. 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. 323 324 325 326 327 328 6290 Swallowtails of the world 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 Hsu 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) aeacus (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 (118), In 1974 the Entomological Suppliers Association of Great Britain agreed not to trade in the endemic Taiwan subspecies T. a. kKaguya 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 asubspecies of T. (T.) rhadamantus (118, 262). Other refs: 141, 147, 281. ides) dohertyi Rippon, 1893 Vulnerable—refer to section 6, p. 271. Indonesia (Talaud 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 80 330 331 332 333 334 335 Protected in Indonesia under the name T. (T.) rhadamantus, of which it has been considered to be a subspecies (118, 246). Common name: Talaud Black Birdwing. Other refs: 49, 52, 119, 141. 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. 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. Itis 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. 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. 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-like, 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. 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. 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. 336 337 338 339 340 341 Swallowtails of the world 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). 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. 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, 119) 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 IT 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 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. 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 Ornithoptera goliath Oberthir, 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), but 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. Ornithoptera tithonus de Haan, 1840 Insufficiently Known—refer to section 6, p. 273. Indonesia (Irian Jaya, Waigeo, Salawati and Mysol (Miso6l) 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. 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 82 342 343 344 345 346 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. Ornithoptera akakeae Kobayashi and Koiwaya, 1978 from 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. 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. Ornithoptera paradisea Staudinger, 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. 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 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, 119, 141. 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 (Abbé Allotte’s Birdwing) from the Solomon Archipelago (Bougainville and the south of 347 348 349 350 Swallowtails of the world Malaita) is generally regarded as a hybrid of O. victoriae x O. priamus 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. 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. 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. lydius is unusual in that the female mimics unpalatable species of Danainae. Other refs: 49, 141. Ornithoptera priamus (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 (Frihstorfer) (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 (Guérin—Méneville), 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, Gor l76, 217, 255) Ornithoptera euphorion (Gray, 1853) Cooktown to Mackay, northern Queensland, Australia. Listed as a subspecies of 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 351 should be monitored because of its restricted distribution in an area of extensive habitat destruction: Common name: Cairn’s Birdwing (49). 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: 84 352 353 354 355 356 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: gl/aucus Linnaeus 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. Papilio (Pterourus) alexiares Hopffer, 1886 Restricted to eastern Mexico, but not known to be threatened. Two subspecies. Refs: 51, 55, 62, 175, 234, 269. 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. 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 validity (137). Common name: Western Tiger Swallowtail (216). Other refs: 55, 62, 141, 234. Papilio (Pterourus) eurymedon Lucas, 1852 South-western Canada (British Columbia), western U.S.A. (Washington, Idaho, Montana, Oregon, Wyoming (?), California, Nevada, Utah, 357 358 359 360 361 362 363 364 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 Papilio (Pterourus) pilumnus Boisduval, 1836 Mexico and Guatemala; occasional records from adjacent U.S.A. (Texas and southern Arizona), Honduras and E| Salvador. Not threatened. Refs: 51, 55, 62, 137, 169, 216, 234, 269. 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. 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. 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. Papilio (Pterourus) neyi Niepelt, 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 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 asynonym. Refs: 51, 55, 169, 234. Papilio (Pterourus) bachus C. & R. Felder, 1865 Colombia, Ecuador, Peruand Bolivia. Not uncommon and not known to be threatened. Two subspecies. Refs: 51, 55, 234. Species-group: scamander Boisduval 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 86 365 366 367 368 369 370 SH 372 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. 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. 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 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. 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. 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. Papilio (Pterourus) victorinus Doubleday, 1844 Mexico and Central America (excluding Panama). Rare in Costa Rica. Flies at canopy level (61). Not believed to be threatened. Three subspecies. P. (P.) diazi Racheli and Sbordoni, 1975 was recently discovered in Mexico (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. 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 syedra 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. 373 374 ai) 376 377 378 7 319 380 381 Swallowtails of the world 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. Papilio (Pterourus) homerus Fabricius, 1793 Endangered—refer to section 6, p. 297. Jamaica. This much-prized species was formerly classified as Vulnerable 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. Papilio (Pterourus) warscewiczi Hopffer, 1886 Ecuador, Peru and Bolivia. Not uncommon and not threatened. Three subspecies. Refs: 51, 55, 234. 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. 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 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. Papilio (Heraclides) andraemon Hibner, 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. 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. 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 88 382 383 384 385 386 387 388 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 Hubner is a synonym. Common name: the Lycophron (9). Other refs: 26, 51, 55, 62, 137, 234, 269. Papilio (Heraclides) androgeus 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 subspecies. Common names: the Androgeus (9), Androgeus Swallowtail (230), Queen Page (9, 282). Other refs: 26, 51, 55, 62, 234, 269. 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. 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, ponceanus 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). Papilio (Heraclides) caiguanabus Poey, 1851 Indeterminate—refer to section 6, p. 305. Restricted to Cuba, where it is rare (230): more information onits status and prospects is required. Common name: Poey’s Black Swallowtail (230). Other refs: 51, 55, 234. 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. 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. 389 390 391 392 393 394 395 396 397 Swallowtails of the world Papilio (Heraclides) cinyras Ménétriés, 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. 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 seis: 1, 55),254. 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. 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: torquatus Cramer 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. 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. tolus 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. 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. 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. 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 declining (23). Other refs: 26, 51, 122, 139, 234. 89 Threatened Swallowtail Butterflies of the World 90 398 399 400 401 402 403 404 405 406 407 Species-group: anchisiades Esper 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. 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: S595, 254 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. 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. Papilio (Heraclides) erostratus Westwood, 1847 Eastern Mexico (not western as given by D’Almeida (55)), Guatemala and Belize. Not common, but not known to be threatened. Other refs: 51, 62, 234, 269. 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. 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. Papilio (Heraclides) chiansiades Westwood, 1872 Ecuador and Peru (Upper Amazon). Generally common and not threat- ened. Refs: 51, 55, 169, 234. Papilio (Heraclides) dospassosi Riitimeyer, 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 Eurytides trapeza (221). It could be a candidate for conservation concern if shown to be a good species. Papilio (Heraclides) anchisiades Esper, 1788 Southern U.S.A. (Texas), Mexico, Central America, South America (excluding Chile and Uruguay) and Trinidad. Widespread and generally 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. 408 409 410 411 412 413 414 415 Swallowtails of the world 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. 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. Papilio (Heraclides) isidorus Doubleday, 1846 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. 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 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: e/wesi Leech 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. 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 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 92 416 417 418 419 420 421 422 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. 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 Papilio (Chilasa) veiovis Hewitson, 1853 Indonesia (Sulawesi). Not uncommon, but not known to be threatened. Refs: 52, 262. 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. 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 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. 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. 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 423 424 425 426 427 428 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: /aglaizei Depuiset Papilio (Chilasa) laglaizei Depuiset, 1877 Indonesia (Irian Jaya, Waigeo and Aru) and Papua New Guinea. Local but not threatened. Refs: 14, 48, 141, 256. 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. 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 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. Papilio (Papilio) hospiton Guenée, 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. Papilio (Papilio) machaon Linnaeus, 1758 Entire Palearctic region through the U.S.S.R. to China (164, 207) and Japan (including Nepal, Bhutan and Taiwan) as far south as some oases in Saudi Arabia and Oman and the high mountains of Yemen (161); Pakistan, northern India, northern Burma, Canada and Alaska. Often common and not threatened as a species. Thirty seven subspecies (141). P. (P.) m. 93 Threatened Swallowtail Butterflies of the World 94 429 430 431 432 433 434 saharae flies in Yemen and parts of the Sahara (161). Larsen (161) listsit 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. 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). 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 asa 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. 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. 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. 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. 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. 435 > 436 437 438 439 440 441 Swallowtails of the world 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. 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. Papilio (Papilio) coloro Wright, 1905 (=rudkini 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. 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: xuthus Linnaeus Papilio (Princeps) xuthus 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. Papilio (Princeps) benguetanus 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 Papilio (Princeps) euchenor Guérin—Méneville, 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 96 443 444 445 446 447 448 449 450 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. 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 Kalimantan) (52, 191, 262). Fairly common and not threatened. Three subspecies. Common names: Banded Swallowtail (3, 45), Burmese Banded Swallowtail (80). Other ref: 141. 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 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. 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. 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 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.) jJanaka—see below). Common name: Tailed Redbreast (80). Other refs: 4, 52, 141, 259. 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. 451 | 452 453 454 455 456 457 458 459 460 Swallowtails of the world Species-group: memnon Linnaeus 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. 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. 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. Papilio (Princeps) oenomaus Godart, 1879 9 2") s wah } Indonesia (Lesser Sunda Is: Timor, Wetar, Moa, Leti, ‘Kisar and Romang) (262). Common (48) and presumably not threatened. Two subspecies. 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. 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. Papilio (Princeps) deiphobus Linnaeus, 1758 Indonesia (Moluccas). Not common but not thought to be threatened. At least two subspecies. Ref: 109. 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.) rhetenor Westwood, 1841. Common name: Redbreast (80, 251). Other refs: 4, 52, 207, 259. Papilio (Princeps) thaiwanus Rothschild, 1898 Taiwan only. Common, despite its restricted range and not therefore listed, though its status should clearly be monitored. Refs: 52, 141, 246. 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 98 461 462 463 464 465 466 467 468 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). Papilio (Princeps) memnon Linnaeus, 1758 North-eastern India (Sikkim, Assam and Nagaland), 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. 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 Papilio (Princeps) noblei de Nicéville, 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. 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. 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. 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. 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. 469 470 471 472 473 474 475 Swallowtails of the world 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. 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. 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 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. 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.) walkeri Janson 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. 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. 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 476 477 478 479 480 481 Australia. Collected for trade in Papua New Guinea (36). Not rare and not threatened. Five subspecies. Other refs: 14, 40, 48, 141, 176. Papilio (Princeps) phestus Guérin—Méneville, 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 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. Papilio (Princeps) nephelus Boisduval, 1836 Nepal, north-eastern India (including Assam and Sikkim), Bhutan (7), 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 ahybrid, 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. 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. 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. 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 > CR = ENS | Wo Wc | ae Papilio (Princeps] hipponous C. 6 R. Felder, 1862 * hilippines: Palawan, Banggi. Bazilan, Luzon, Mindoro, Panay is Negros, Cebu, Bohol, Indonesia: Sangihe and Talsut Sulawesi as vagrants. Leyte, Samar, Mindanao (Hancock 1985), Is., and Possibly northern Two subspecies, 484 485 486 487 488 Swallowtails of the world species is urgently needed. Up to six subspecies. Other refs: 52, 110, 118, 121, 262. 963. Papilio (Princeps) pitmani Elwes 6 de Nicéville, 1866. South Burma: Tenasserim, Tavoy. Thailand, N. Vietnam (110). ? Laos. China: Hainan. Possibly three subspecies. TLLUMNCUUDS. 21 Lampe asa uae 2 rans PPIeS Vireo aueuIUINs LU au Wve, 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. Papilio (Princeps) fuscus 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. andamanicus (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.) fuscus x P. (P.) tydeus (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. 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. 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. 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. 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 Threatened Swallowtail Butterflies of the World 489 490 491 492 493 494 495 496 102 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 placed as a full species near the 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 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. Papilio (Princeps) schmeltzii Herrich—Schaffer, 1869 Fiji. Despite its restricted range, the species is apparently common (48) and not threatened. Other ref: 111. 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 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. 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. Papilio (Princeps) weymeri 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 trade as ex pupae (36). P. (P.) cartereti Oberthiir, 1914 is asynonym, 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. 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. | 497 498 499 500 501 502 503 504 Swallowtails of the world Papilio (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. 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 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. Papilio (Princeps) plagiatus 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. Papilio (Princeps) zoroastres Druce,1878 Cameroon, Equatorial Guinea (Bioko, formerly known as Fernando P6o 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. 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. Papilio (Princeps) jacksoni Sharpe, 1891 Kenya, Uganda, Zaire, eastern Congo (?), Burundi (?), Rwanda, western Tanzania, Zambia and Malawi. Commonin 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. 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 (pv bv Threatened Swallowtail Butterflies of the World 505 Papilio (Princeps) sjoestedti Aurivillius,1908 Rare—refer to section 6, p. 337. Known from montane forest in only three localities, all in northern Tanzania: Mt Meru, Ngorongoro Crater and, as a different subspecies, Mt Kilimanjaro, where it is abundant. Mt Meru and Mt Kilimanjaro are both well protected national parks indicating that this species, although very narrowly distributed, is not threatened at the moment. Common name: Kilimanjaro Swallowtail (285). Other refs: 33, 34, 43, 50, 151. Species-group: rex Oberthtr 506 Papilio (Princeps) rex Oberthiir, 1886 Highland forest in Nigeria, Cameroon, Sudan, Ethiopia, eastern Congo, Zaire, Uganda, western Kenya, Rwanda, Burundi and Tanzania. Not 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 ee A a5 7 - \ ot _ 507 Papilio (Princeps) epiphorbas Boisduval, 1833 pase) shtOS —9 Af Madagascar. Well distributed over the whole island (and Comoro Is, e OOS. 0, according to D’Abrera (50)) and not threatened. Other refs: 34, 204, 206. = (ev’~ 508 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 Réunion. 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 Réunion 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 I.), 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 rep S12 513 514 515 518 Swallowtails of the world subspecies is reported to be rare in Tanzania (151). Common name: Horniman’s Swallowtail (285). Other refs: 33, 34, 50. 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. 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. 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). Itis not known to be threatened though more information is desirable. Common name: Abyssinian Blue-banded Swallowtail (285). Other refs: 34, 50, 116. Papilio (Princeps) nireus 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. Papilio (Princeps) aristophontes Oberthiir, 1897 Indeterminate—refer to section 6, p. 345. 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. Papilio (Princeps) oribazus Boisduval, 1836 Madagascar. Well distributed, except in the west, and apparently quite common and not threatened. Refs: 50, 204, 206. Papilio (Princeps) thuraui Karsch, 1900 East Africa: highland forest in southern Tanzania, Malawi and northern Zambia (115). Considered to be scarce and very local (43, 50, 285), perhaps deserving conservation status and certainly requiring monitoring. It has been reported to be rare in Tanzania (43, 151) and possibly threatened by deforestation if this proceeds unchecked, although it is quite firmly 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 Bok on A VinshovV. 208 ve | SOL Threatened Swallowtail Butterflies of the World holy) +1 ferwe t < ~ 106 519 520 521 ~ hick ly 522 523 524 previously placed as subspecies of P. (P.) bromius. Common name: Blue-spotted Black Swallowtail (285). Other refs: 34, 97. 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 Gifford, 1961 was areplacement name for P. (P.) brontes Godman, 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. occidua 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, PD Papilio (Princeps) interjecta 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 (109, 116). Conservation status not known; further research required. Papilio (Princeps) bromius Doubleday, 1845 Forests in Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin, Nigeria, Cameroon, Central African Republic, Sudan, Ethiopia (7), Equatorial Guinea (?), Sao Tomé, Gabon, Congo, Zaire, Uganda, Kenya, 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. 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 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 scarce) and species not known to be threatened. Often placed in the genus Iterus. Common names: Giant Blue Swallowtail (285), Great Blue Papilio (34). Other refs: 20, 50, 105. Species-group: zenobia Fabricius Papilio (Princeps) gallienus Distant, 1879 Lowland forest in Nigeria, Cameroon, Gabon, Equatorial Guinea (?), Congo and Zaire. Uncommon (285), but not markedly so and not 525 526 527 528 529 530 Sei Swallowtails of the world threatened. Common name: Large White-banded Papilio (34). Other ref: 20. 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. 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 Pé6o I.), 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. 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. Papilio (Princeps) andronicus Ward, 1871 Cameroon and possibly adjacent areas. Uncommon but not known to be threatened. Refs: 50, 133. 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 Papilio (34). Other refs: 20, 33, 50, 285. 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 listed 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: antimachus Drury 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, formerly known as Fernando Péo I.), Gabon, Congo, Zaire, Uganda, Rwanda and Angola (58). Males not uncommonly collected, females very scarce. Often placed in the genus Druryia. 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 532 533 534 108 535 53i/ 539 Species-group: menestheus Drury 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. 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. Papilio (Princeps) lormieri Distant, 1874 / Forests up to 2000 m in Nigeria, Cameroon, Central African Republic, south-western Sudan, Equatorial Guinea (?), Gabon (?), Congo, Zaire, Uganda, western Kenya and Angola. Not uncommon and not threatened. Replaced by P. (P.) menestheus west of the Niger and by P. (P.) ophidicephalus from the Kenya coast to South Africa. Sexes similar, three subspecies. Common names: Western Emperor Swallowtail (285), Emperor Swallowtail (34). Other refs: 33, 50. Species-group: demoleus Linnaeus 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. 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. Papilio (Princeps) erithonioides Grose—Smith, 1891 Madagascar only, mainly in the west of the island. Not uncommon and apparently not threatened. Refs: 34, 50. Papilio (Princeps) demodocus Esper, 1798 Tropical and southern Africa, Saudi Arabia, Yemen, South Yemen, Oman; introduced to Madagascar, Mauritius and Réunion. Abundant in open habitats and not threatened. A minor pest on Citrus. Aggressive towards other butterflies flying in its vicinity (50). Sexes alike, 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. Papilio (Princeps) demoleus Linnaeus, 1758 Oman, United Arab Emirates, Saudi Arabia, Kuwait, Bahrain, Qatar, western and possibly also eastern Afghanistan, Sri Lanka, India, Nepal, 540 541 542 543 544 545 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, Alor and Sumba), Papua New Guinea, Australia (including Lord Howe I. (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, 118, 141, 148, 149, 165, 176, 246, 262. Species-group: Jeucotaenia Rothschild 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. 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.) mangoura with female P. (P.) delalandei, which has similar yellow-barred wings (263). Other refs: 34, 50, 195. Species-group: phorcas Cramer Papilio (Princeps) constantinus 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. 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. 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 109 Threatened Swallowtail Butterflies of the World 110 546 547 548 549 550 551 552 359 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 Papilio (Princeps) euphranor Trimen, 1868 Restricted to highland forest in South Africa (Cape Province, Natal and Transvaal). Uncommon (34); but no threats known. Has been proposed for inclusion in a Red Data Book for South Africa (192), although no such work has been started. Common names: Forest Kite, Bush Kite (285), Natal Swallowtail (34). Other refs: 33, 50, 63, 214. 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. Papilio (Princeps) hesperus Westwood, 1843 Lowland forest in Ivory Coast, Ghana, Togo, Benin, Nigeria, Cameroon, Central African Republic, southern Sudan, Equatorial Guinea (Bioko, formerly known as Fernando Poo I.), Congo, Zaire, Uganda, Rwanda (?), Burundi (?), western Tanzania and north-western Zambia. Replaced by P. (P.) pelodorus in Malawi and Tanzania and by P. (P.) horribilis from the 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. Papilio (Princeps) horribilis Butler, 1874 West Africa; Sierra Leone, Liberia, Ivory Coast and Ghana. Givenasa 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. Papilio (Princeps) nobilis Rogenhoffer, 1891 Highland forest in southern Sudan, Kenya, Uganda, Burundi, Rwanda, Tanzania and Kivu Province, Zaire (34, 285). Not uncommon and not recognized as being threatened. Three subspecies. Common name: Noble Swallowtail (34, 285). Other refs: 33, 50. Species-group: palinurus Fabricius 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. 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. Papilio (Princeps) buddha Westwood, 1872 Southern India. Locally common and not rare (118). Protected by law 554 555 556 57) 558 559 560 561 Swallowtails of the world (182), but not known to be threatened. Common name: Malabar Banded Peacock (80). Other refs: 4, 52, 259. 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. 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 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. Papilio (Princeps) maackii Ménétriés, 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.) bianor. 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. Papilio (Princeps) bianor 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. Papilio (Princeps) syfanius Oberthir, 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. Papilio (Princeps) polyctor Boisduval, 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. Papilio (Princeps) dialis (Leech,, 1893) Southern China (including Hainan (Guangdong prov.)), north-eastern 111 Threatened Swallowtail Butterflies of the World 112 562 563 564 565 566 567 568 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. 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. 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. 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. Papilio (Princeps) arcturus 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. 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. 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 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. 569 570 571 572 a 4 , Swallowtails of the world 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. 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. 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 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, PALI PAUS Papilio (Princeps) montrouzieri Boisduval, 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): NYDN FWN Ke . Parnassius pythia Rothschild, 1932 . Parnassius rothschildianus Bryk, 1932 . Parnassius stenosemus Honrath, 1890 . Eurytides chibcha (Fassl, 1912) . Eurytides hipparchus (Staudinger, 1884) . Graphium sisenna (Mabille, 1890) . 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. 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). Jn litt., 18 March. 113 Threatened Swallowtail Butterflies of the World ie*) 27. 28. 29. 30. 31. 32. 114 . Anonymous (1980-1984). Various dealers’ lists. . Antram, C.B. (1924). Butterflies of India. Thacker, Spink & Co., Calcutta and Simla. 226 pp. . Arora, G.S. and Nandi, D.N. (1980). On the butterfly fauna of Andaman and Nicobar Islands (India). 1. Papilionidae. 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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. schaffgotschi, 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. olbrechtsi 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. Itis 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. Parnassius 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. Parnassius 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(?), 36-3 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, 34-36, 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 #, 1O;17(y, 19, 21, 35, 50754, 57, 59°60, 135, 198,272, 274( 7), 284.2550), 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 Jiangx1) 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 Tran Iraq Japan (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) 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, 41, 54, 273, 428, 446-448, 557, 558 (14 species) South Korea 29, 30, 33, 39, 41, 54, 203(?), 273, 428, 446-448, 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 105 17, 20; 21, 39, 57,116, 10 F 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 C A ae Le BAe ic 60, 114, 115, 120, ENON 126, 128, 129, 432) 134, 136-138, 141, 143, 180%" 5 Lite 181, 1852 190, 194-20), ms, 205, 224, 287-291, 296-300,G05-G072314-316, + (434% GIB-321 (324, 326, 320-334 633-344, 347-349, 415441? 422, 423, 442-444, we 45), 452. 455-457, 462, 466) 467, 469-475 477, 478,482, 484-486, 489, 493, (496-498, 539,552) 554, 563, 564, 5683572 (121 species) 195. Graphium f Graphium) stresemanni RARE (Moluccas) D> (AD BAD WB GD OD CGY BH HL? =A \ Mes >, eng Ps,” : Gu! SAr 15 | 99 lus $I (GA>) «KI “i CX Threatened Swallowtail Butterflies of the World 288. Atrophaneura (Atrophaneura) luchti RARE (Java) 299. Atrophaneura (Losaria) palu (NSUFFICIENTLY KNOWN (Sulawesi) 324. Troides (Troides) prattorum INDETERMINATE (Buru) 329. Troides (Troides) dohertyi VOLNERABLE (Talaud Is and Sangihe) 340. Ornithoptera tithonus INSUFFICIENTLY KNOWN (Irian Jaya) 341. Ornithoptera rothschildi INDETERMINATE (Irian Jaya) 342. Ornithoptera chimaera (NDETERMINATE (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 ! (Sabah and Sarawak) aR 60, 114, 115, 120, 126-130,134, 136-138, 180, 181, 184) 186-189, 197, 203, 296, 298, 314, 316, 320-322) 330, 416, 422, 444,\453, 461(?), 4p, 467, 470, 473, 478, 484, 554, 564 (42-43 species) a 184. Graphium (Graphium) procles INDETERMINATE (Sabah) 322. Troides (Troides) andromache INDETERMINATE 453. Papilio (Princeps) acheron RARE Peninsular Malaysia (including Singapore) 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, 484, 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, 423-425, 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 (NDETERMINATE 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) 128 133. 139. 202. 308. 312: 418. 419. 441. 556. Swallowtails of the world Graphium (Pathysa) idaeoides RARE Graphium (Pathysa) megaera INDETERMINATE (Palawan) Graphium (Graphium) sandawanum VULNERABLE (Mindanao) Atrophaneura (Pachliopta) schadenbergi VULNERABLE (Luzon, Babuyan) Atrophaneura (Pachliopta) atropos INDETERMINATE (Palawan) Papilio (Chilasa) oomana VULNERABLE (Leyte and Mindanao) Papilio (Chilasa) carolinensis VULNERABLE (Mindanao) Papilio (Princeps) benguetanus VULNERABLE (Luzon) 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. 204. Graphium (Graphium) meeki RARE 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 Faso (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) SBhile Angola Papilio (Princeps) antimachus RARE (Benin (?), Ghana, Guinea, Ivory Coast, Liberia, Sierra Leone and Togo (?)) 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) 5311 Burundi Papilio (Princeps) antimachus RARE 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 Chad Congo Djibouti Equatorial Guinea Awe 5 Ethiopia Gabon Kenya Malawi 144, 147(?), 149, 151, 152, 155, 157, 160, 161(?), 167(?), 168(?), 172, 178, 499(?), 500, 513, 515, 521, 523, 525-527, 529, 531, 534, 538, 544, 545, 548 (24-29 species) 531. Papilio (Princeps) antimachus RARE 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) (including Bioko (formerly Fernando P6o)) aS 144, 149(?), 152(?), 157(?), 160(?), 164, 172, 178, 501, 510, 515, 521(?), * 523(?), 524(?), 526, 531, 534(?), 538, 544, 545, 548 (13-21 species) 531. Papilio (Princeps) antimachus RARE 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 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 144, 145, 146, 148, 164, 171-173, 177, 178, 502-504, 512, 515, 518, 519, 532, 543-545, 547 (22 species) Mozambique Rwanda Somalia 130 144, 146, 148, 164, 169, 171-173, 177, 178, 502, 532, 538, 543-545 (16 species) 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) leucotaenia VULNERABLE 144, 164, 171, 172, 177, 178, 514, 515, 538, 543-545 (12 species) Sudan Swallowtails of the world 144, 148, 149, 164, 172, 178, 500, 501, 506, 512, 515, 521, 525, 526, 534, 538, 544, 545, 548, 550 (20 species) Tanzania Uganda Zaire Zambia Canada Mexico U.S.A. (including Zanzibar and Pemba Island) 144, 145, 147-149, 161, 164, 165, 171-173, 176-178, 501-506, 511,512,515, 518, 519, 521, 532, 538, 543-545, 547, 548, 550 (34 species) 505. Papilio (Princeps) sjoestedti RARE 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) antimachus RARE 540. Papilio (Princeps) leucotaenia VULNERABLE 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 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 23, 24, 37, 61, 208, 352, 334-356, 359, 391, 428, 430, 432-436 (18 species) 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 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. Eurytides (Protesilaus) marcellinus 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. Eurytides (Protesilaus) lysithous harrisianus ENDANGERED 112. Eurytides (Eurytides) 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, 211, 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, 411, 432 (59 species) Ecuador 71, 75, 76, 80, 83, 84, 92-98, 100, 101, 103, 104, 106, 107, 110, 111, 211, 215, 218, 219, 221, 229, 238, 244, 247, 248, 251-257, 259, 264, 267-269, 360-363, 366, 369, 375-377, 382, 383, 388, 389, 392, 394, 401, 405, 407, 410, 411, 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, 261, 264, 265, 267, 268, 360(?), 369, 382, 383, 388, 393, 394, 398, 407 (30-31 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 (Heraclides) 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 (Heraclides) 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(?), 111, 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 “|? 166, 507(2), 516, 545 (3-4 species) 166. Graphium (Arisbe) levassori VULNERABLE 516. Papilio (Princeps) aristophontes INDETERMINATE Madagascar 143, 163, 179, 270, 507, 517, 535-538, 541, 542, 545 (13 species) 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 Réunion 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 | Minho pal fhe Wet i ma. “ten Heri ahi inst 5 aris apes | nn) eee Ch vinta eabeeraia Suing. - * Wee te) a) : iM See Mena ae ABS, 262, BOR, MH?) "Peli eel sen dro _ ef | Sy thai sak soslgiiR, si. iF « apfeiaiy 1) SB: ; a ache | : Aiea wed ea na ies mm, spr, aa BR PF agit aia DAS, oe 1254, ann? 1289, Ih, ey, 28, its tet a } la ie ¢ ee aan Nosh "4a, AN? 35. 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