Variation and Distribution in the Malayan Papilionidae

Catalogue of Malayan Papilionidæ

Ornithoptera (Boisd.)

The characters in the larva and pupa which have been supposed to distinguish this genus from Papilio are erroneous, or at least do not exist in all the species. My own observations on O. Poseidon show that the larva has no “external sheath” to the thoracic tentacles, and that the suspending thread passes round the pupa, and is not “fastened on each side to a silky tubercle.” There remain therefore only the characters of the perfect insect, the most important of which are the anal valves in the male. These are very large, ovate or rounded, coriaceous, and not hairy, and are furnished with projecting points or spines (sometimes very conspicuous) which serve to attach the male more firmly to the female in copulâ. In several species I have observed, these points or hooks were buried in the protruded anal gland of the female, and thus effectually prevented the great weight of the insects causing them to separate upon suddenly taking flight. The great strength and size of these insects, the thick texture of their wings, their long curved and stout antennæ, their peculiar form, colour, and distribution, are the only other characters that separate them from Papilio. Though these may not perhaps be technically sufficient, I think it advisable and convenient to retain a genus so well known and long established.

Ornithoptera is pre-eminently a Malayan genus, seventeen species inhabiting the archipelago, one (Rhadamanthus, Bd.) India and China, one (Darsius, G. R. Gray) peculiar to Ceylon, one (Richmondia, G. R. Gray) North Australia. O. Victoriæ, G. R. Gray, from some island east of New Guinea, should probably be included in the Malayan list; and Æacus, Felder, from an unknown locality. The following are the well-established Malayan species.

A. Priamus Group

1. Ornithoptera Priamus, Linnæus.

♂. Papilio Priamus, L.; Cram. Pap. Ex. t. 23. f. A, B; Godart, Enc. Méth. ix. p. 25. O. Priamus, Bd. Sp. Gén. Lép. p. 173. ♀. P. Panthous, L.; Cram. Pap. Ex. t. 123. f. A, t. 124. f. A.

This may be at once distinguished from all the allied species with which it has been often confounded—in the male, by the more rounded and deeply scalloped hind wings, with larger black spots and a broader border, the upper wings with no green on the median nervure or its branches, and the sooty patch extending only to the second median nervule; in the female, by the very constant and peculiar light olive-brown colour, the absence of any spots in the discoidal cell of the upper wings, and the broad shallow scallops of the hinder margin.

Hab. Amboyna and Ceram, probably also Bouru (Wall.).

2. Ornithoptera Poseidon, Doubleday.

O. Poseidon, Db. Ann. of Nat. Hist. xvi. p. 173; Westwood, Cat. of Orient. Ent. pl. 11, 14.

The numerous specimens of Ornithoptera which I obtained in various parts of New Guinea and the adjacent islands show so much instability of form, colouring, and even of neuration, no two individuals being exactly alike, that I am obliged to include them all in one variable species, to which I believe must also be referred O. Pronomus, G. R. Gray, from Cape York, O. Euphorion, G. R. Gray, from North Australia, O. Archideus, G. R. Gray (ex Boisd.), erroneously said to be from Celebes, and O. Boisduvalii, Montrouzier, from Woodlark Island.

Var. a, Aru Islands (Wall.). O. Arruana, Feld. Lep. Frag. p. 24.

Individuals from this locality differ in the arrangement of the nervures; in some the third subcostal nervure of the upper wings branches from the same point with the upper disco-cellular, in others considerably beyond it; the points from which the subcostal nervures branch also vary. The amount of green colour on the median nervure and its branches varies. In some specimens there is a spot at the anal angle of lower wings beneath, agreeing with O. Pronomus, G. R. Gray; but this is generally wanting.

Var. b, Dorey, Salwatty, south-west coast of New Guinea (Wall.).

These agree very closely with O. Poseidon, as figured by Westwood; they differ individually in the same manner as the last, and also in the length of the lower disco-cellular nervure on the under wings. They have generally no golden spots beneath the wings. They vary also in the outline of the under wings, the outer and anal angles being more acute in some specimens than in others. Some have the under wings of a uniform green entirely without spots, while others have a range of black spots more or less fully developed.

Var. c, Waigiou (Wall.). Archideus, G. R. Gray, ♀.

This agrees with the last; but the male is of a more delicate green than any of the others, and has less of that colour on the median veins. On the under side there are no golden spots. The whole surface has a golden tinge, and the central portion of the lower wings is tinged with amber-brown.

The females of all the above vary extremely, much more even than the males, and from the same locality two specimens are rarely alike. The discoidal cell is in some specimens more than half occupied by a whitish patch, while in others there are only a few small spots. One of my specimens from Salwatty almost exactly agrees with that figured by Westwood (Cat. of Or. Ent. pl. 14) as from Cape York. One of the Waigiou specimens is the same as Archideus, G. R. G., figured by Boisduval (Voy. de l’Astrolabe, t. 4. f. 1, 2); and another, from New Guinea, differs very little from Euphorion, G. R. G. (Brit. Mus. Cat. Lep. pt. 1. pl. 2. f. 3), from North Australia.

From these facts I am led to conclude that we have here a variable form spread over an extensive area, and kept variable by the continual intercrossing of individuals, which would otherwise segregate into distinct and sharply defined races. The same area is inhabited by many species of birds common to all parts of it; and just as the birds of Ceram and Amboyna are almost all distinct species from those of New Guinea, so do we find those islands inhabited by the Ornithoptera Priamus, a well-marked and constant species, readily distinguishable in either sex from the inconstant forms of New Guinea proper. The same parallel holds in North Australia. Many New Guinea species of birds extend, with very slight variation, to the country about Cape York; but when we reach the Moreton Bay district all these have disappeared, and we find only true Australian species. So the variable forms of O. Poseidon reach North Australia and Cape York, while in the Moreton Bay district we find the comparatively well-marked species O. Richmondia. Similar causes, whether geographical or climatal, have thus produced an analogous distribution in these widely separated groups of animals.

3. Ornithoptera Crœsus, Felder.

O. Crœsus, Feld. Wien. Ent. Monats., Dec. 1859. O. Crœsus, G. R. Gray, Proc. Zool. Soc. 1859, p. 424.

Hab. Batchian (Moluccas) (Wall.).

Local form, a.—Male: has the orange colour of the upper surface of a much deeper fiery-red hue; on the under surface, the black spots of the lower wings are nearer the margin, and the yellow spots below them are entirely absent; there is also a green line between the subcostal nervure and the margin; on the under surface of the fore wings the green patch in the discoidal cell extends to its base, and is reflexed in a narrow line along its upper edge.

Female: differs still more from that sex in O. Crœsus; the white markings on all the wings are so large as almost to fill up the spaces between the veins, the lower part of the discoidal cell in both upper and under wings being also occupied with a whitish patch; the range of spots occupying the posterior margin are of a dusky yellow colour.

Hab. Ternate (♂), Gilolo (♀) (Wall.).

This well-marked local form is no doubt peculiar to Gilolo and the small adjacent islands, as the original species is to Batchian.

I was three months in the island of Batchian before I obtained a specimen of this fine insect, which I had seen once or twice only flying high in the air. I at length came upon it flying about a beautiful cinchonaceous shrub with white bracts and yellow flowers (Mussænda, sp.); and having cleared a path round about, I visited the place every morning on my way to the forest, and once or twice a week had the satisfaction of capturing a fine male specimen of O. Crœsus. The females were more plentiful and more easily caught. I afterwards sent out one of my men with a net every day to look after this insect only. He would stay out all day long, wandering up a broad rocky torrent, where the males flew up and down occasionally or settled on the rocks which just appeared above the water. He generally brought me one, and sometimes even two or three specimens; and thus, with those that I myself captured at the flowers, I secured a fine series of this richly coloured species.

4. Ornithoptera Tithonus, De Haan.

O. Tithonus, De Haan, Verh. Nat. Gesch. Ned. t. 1. f. 1.

Hab. S.W. Coast of New Guinea (Leyden Museum).

This remarkable species must be very rare, as I never saw it in any part of the New Guinea district that I visited; nor was it seen during the exploration, a few years ago, by a Dutch steamer which visited the part of the coast where the only specimen known was said to have been obtained.

5. Ornithoptera Urvilliana, Guérin.

Papilio Urvilliana, Guér. Voy. de la Coquille, Lép. t. 13. f. 1, 2, ♂.

O. Urvilliana, Boisd. Sp. Gén. Lép. p. 175.

Hab. New Ireland (Paris Museum).

B. Pompeus Group

6. Ornithoptera Remus, Cramer.

Papilio Remus, Cr. Pap. Ex. t. 135. f. A, t. 136. f. A (♀), t. 386. f. A, B (♂); Fab. Syst. Ent. iii. 1. p. 11.

O. Remus, Bd. Sp. Gén. Lép. p. 176. Papilio Panthous ♂, Clerck, Icon. t. 18 (♀).

Hab. Amboyna, Ceram, Gilolo, Morty Island, Sulla Island, Celebes (Wall.).

The specimens above quoted agree well with Cramer’s figures. The female from the Sulla Islands differs only in having more yellow towards the anal angle of the lower wings. The specimens figured by Cramer in pls. 10, 11, under the name of “Hypolitus” seem to be a remarkable variety, in which the female has much of the character of the male. Messrs. Doubleday and G. R. Gray have adopted Panthous as the specific name of this insect; but this name was first used by Linnæus for the female of Priamus only, in the 10th ed. of the ‘Systema Naturæ’ (1758). Clerck (in 1759) adopted the name, but supposed he had found the male in the female of Remus. Linnæus, in Mus. Lud. Ulric. (1764), and in the 12th ed. of the ‘Systema Naturæ’ (1766), adopts this error, so far as referring to Clerck’s two figures; but in both these works his description refers only to the female of P. Priamus, indicating that the supposed other sex (P. Remus) was not known to him personally. The name of Panthous must therefore altogether drop, it having been applied to this species only through a double error—first, that of Linnæus, in supposing his Panthous to be distinct from Priamus, and then that of Clerck, in thinking that a female Remus was the male of the Linnean Panthous.

7. Ornithoptera Helena, Linnæus.

♂. P. Helena, Cram. Pap. Ex. t. 140. f. A, B. O. Helena, Boisd. Sp. Gén. Lép. p. 177.

♀. P. Amphimedon, Cram. Pap. Ex. t. 194. f. A. O. Amphimedon, Boisd. Sp. Gén. p. 176.

Hab. Amboyna and Ceram (Wall.).

The females from these localities are always sooty, with the spots and markings on the hinder wings of a dull buff-colour even in the freshest specimens.

a. Local form Bouruensis.—Male: exactly resembles the Amboyna specimens, except that the yellow patch is more variable in form and extent.

Female: nearly black, and with the markings on the lower wings almost as pure and deep yellow as in the males: size a little smaller than in the type.

Hab. Bouru (Wall.).

b. Local form Papuensis.—Female: sooty black, the two first branches of the subcostal nervure margined with whitish near their origin; markings of the lower wings of the same tint of orange-yellow as is O. Helena ♂, but not so glossy.

Male not known.

Hab. New Guinea, Salwatty (Wall.).

c. Local form Celebensis.—Male: wings a little more pointed than in O. Helena; yellow patch of lower wings extending nearer to the posterior margin, and bounded towards the abdominal margin by the first branch of the median nervure. Beneath, having the nervures between the discoidal cell and the outer border ashy-margined.

Female not known.

Hab. Macassar (Celebes) (Wall.).

Remarks.—Of these three local modifications of O. Helena, the first is very distinct in the female, but not separable in the male sex. Of the second and third, only one sex is known; and they may very probably prove to be well-marked species when more materials are obtained.

8. Ornithoptera Leda, n. s.

Male: upper wings elongate, triangular, glossy black, quite uniform and immaculate; the outer margin delicately white-marked at the termination of the nervures. Lower wings yellow, as in the allied species, with a black border about the same width as in O. Pompeus on the outer and abdominal margins, narrower on the inner margin; the posterior scalloping of the yellow patch not so deep as in O. Pompeus, and having a spot at the anal angle connected more or less with the margin.

The under surface differs from that of O. Pompeus by the ashy margins of the veins of the upper wings being entirely absent, and in having much less white on the outer edge. There are no submarginal spots except the anal one, much red at the base of the wings, and no black spots on the abdomen.

Female: this sex varies very much, some having the upper wings immaculate, while others have the veins about the end of the discoidal cell broadly margined with whitish. The marginal series of spots on the lower wings vary as they do in O. Pompeus and O. Amphrisius. The best distinction from O. Pompeus (♀) seems to be the more elongated wings, the less crenellated margin, and the more produced outer angle of the lower wings. The yellow patch is also of a deeper colour both on the upper and under surfaces.

Hab. Celebes (Macassar and Menado) (Wall.)

9. Ornithoptera Pompeus, Cramer.

P. Pompeus, Cr. Pap. Ex. t. 25. f. A (♂). P. Minos, Cr. Pap. Ex. t. 195. f. A (♀). P. Heliacon, Fab. Ent. Syst. 3. i. p. 19, 60.

O. Heliacon, Boisd. Sp. Gén. Lép. p. 178.

Hab. Sumatra, Borneo, Java, Lombock (Wall.), India (var.).

Remark.—The form that occurs in India, in its more elongate wings and darker colouring, approaches very closely to O. Rhadamanthus.

10. Ornithoptera Nephereus, G. R. Gray.

P. Astenous, Eschscholtz, Voy. Kotzebue, t. 4. f. A, B, C. (nec Fab.).

O. Nephereus, G. R. G., List of Lep. B. M. Papilionidæ, p. 6.

Hab. Philippine Islands.

Remark.—This is quite distinct from O. Rhadamanthus, Bd., with which it has generally been identified.

11. Ornithoptera Magellanus, Felder.

O. Magellanus, Feld. Lep. Nov. Phil. p. 11.

Hab. North of Luzon (Philippines).

Remark.—This fine species has a beautiful opalescent glow on the lower wings when viewed obliquely.

12. Ornithoptera Criton, Felder.

O. Criton, Feld. Lep. Fragm. p. 49.

Hab. Batchian, Ternate, Gilolo, Morty Island (Wall.).

13. Ornithoptera Plato, n. s.

Male: resembles O. Criton in the form and extent of the yellow patch, but the upper wings differ in having the outer half of a lighter tint; on the under surface this outer half of the wing is of a light ash-colour. Abdomen almost wholly black beneath. No red patches at the base of the wings, or any red collar.

Female unknown.

Hab. Timor (Wall.).

This is a very distinct species, though at first sight resembling several others. I obtained a single male specimen only.

14. Ornithoptera Haliphron, Boisduval.

O. Haliphron, Bd. Sp. Gén. Lép. p. 181 (♂); Felder, Lep. Fragm. p. 37, Taf. ii. f. 2, 3 (♂, ♀).

Hab. Macassar (Celebes) (Wall.).

15. Ornithoptera Amphrisius, Cramer.

P. Amphrisius, Cr. Pap. Ex. t. 219. f. A; Godardt, Enc. Méth. ix. p. 27, pt.

O. Amphrisius, Bd. Sp. Gén. Lép. p. 178.

Hab. Malacca, Java, Borneo (Wall.).

This may be readily distinguished from the allied species by the upper wings in the male being yellow-marked, and by the absence of red spots at the base of the wings beneath in both sexes.

C. Brookeana Group

16. Ornithoptera Brookeana, Wallace.

O. Brookeana, Wall. Proc. Ent. Soc. 1855, p. 104; Hewitson, Ex. Butt. Papilionidæ, i. f. 1. Papilio Trogon, V. Voll. Tijdschrift voor Ent. 1860, p. 69, pl. 6.

Hab. Borneo (Sarawak) (Wall.), Sumatra (Leyden Museum).

Remarks.—I have been in much doubt about the position of this remarkable species, and was for some time inclined to place it among the Papilios. It agrees, however, far better with Ornithoptera in the form and stoutness of the wings, the long stout and curved antennæ, the red collar and patches at the base of the wings beneath, the abdominal fold, and the flight and general appearance. It is powerful on the wing, and occasionally settles on the ground in damp sunny places. It inhabits the interior of North-west Borneo and the mountains of West Sumatra. The female is unknown. It is peculiar in the great length of the discoidal cell of the wings and its altogether unique style of coloration, and must be considered as the type of a distinct group of the genus Ornithoptera.