Part 7
This Table shows the great affinity of the Malayan with the Indian Papilionidæ, only three out of the nineteen groups ranging beyond, into Africa, Europe, or America. The limitation of groups to the Indo-Malayan or Austro-Malayan divisions of the archipelago, which is so well marked in the higher animals (see ‘Journal of Linnean Society,’ vol. iv. 172, and ‘Journal of the Royal Geographical Society,’ 1863, p. 230), is much less conspicuous in insects, but is shown in some degree by the Papilionidæ. The following groups are either almost or entirely restricted to one portion of the Archipelago:—
_Indo-Malayan Region._
_Nox_-group. _Coon_-group. _Macareus_-group (nearly). _Paradoxa_-group. _Dissimilis_-group (nearly). _Brookeanus_-group. LEPTOCIRCUS (genus).
_Austro-Malayan Region._
_Priamus_-group. _Ulysses_-group. _Erechtheus_-group.
The remaining groups, which range over the whole archipelago, are, in many cases, insects of very powerful flight, or they frequent open places and the sea-beach, and are thus more likely to get blown from island to island. The fact that three such characteristic groups as those of _Priamus_, _Ulysses_, and _Erechtheus_ are strictly limited to the Australian region of the archipelago, while five other groups are with equal strictness confined to the Indian region, is a strong corroboration of that division which has been founded almost entirely on the distribution of Mammalia and Birds.
If the various Malayan islands have undergone recent changes of level, and if any of them have been more closely united within the period of existing species than they are now, we may expect to find indications of such changes in community of species between islands now widely separated; while those islands which have long remained isolated would have had time to acquire peculiar forms by a slow and natural process of modification.
An examination of the relations of the species of the adjacent islands will thus enable us to correct opinions formed from a mere consideration of their relative positions. For example, looking at a map of the archipelago, it is almost impossible to avoid the idea that Java and Sumatra have been recently united; their present proximity is so great, and they have such an obvious resemblance in their volcanic structure. Yet there can be little doubt that this opinion is erroneous, and that Sumatra has had a more recent and more intimate connexion with Borneo than it has had with Java. This is strikingly shown by the mammals of these islands—very few of the species of Java and Sumatra being identical, while a considerable number are common to Sumatra and Borneo. The birds show a somewhat similar relationship; and we shall find that the group of insects we are now treating of tells exactly the same tale. Thus:—
Sumatra 21 sp.│20 sp. common to both islands; Borneo 29 sp.│ „ ──────────────┼────────────────────────────── Sumatra 21 sp.│11 sp. common to both islands; Java 27 sp.│ „ ──────────────┼────────────────────────────── Borneo 29 sp.│20 sp. common to both islands; Java 27 sp.│ „
showing that both Sumatra and Java have a much closer relationship to Borneo than they have each other—a most singular and interesting result when we consider the wide separation of Borneo from them both, and its very different structure. The evidence furnished by a single group of insects would have had but little weight on a point of such magnitude if standing alone; but coming as it does to confirm deductions drawn from whole classes of the higher animals, it must be admitted to have considerable value.
We may determine in a similar manner the relations of the different Papuan Islands to New Guinea. Of thirteen species of Papilionidæ obtained in the Aru Islands, five were also found in New Guinea, and eight not. Of nine species obtained at Waigiou, five were New Guinea, and four not. The five species found at Mysol were all New Guinea species. Mysol, therefore, has closer relations to New Guinea than the other islands; and this is corroborated by the distribution of the birds, of which I will only now give one instance. The Paradise Bird found in Mysol is the common New Guinea species, while the Aru Islands and Waigiou have each a species peculiar to themselves.
The large island of Borneo, which contains more species of Papilionidæ than any other in the archipelago, has nevertheless only two peculiar to itself; and it is quite possible, and even probable, that one of these may be found in Sumatra or Java. The last-named island has also two species peculiar to it; Sumatra has not one, and the peninsula of Malacca only one. The identity of species is even greater than in birds or in most other groups of insects, and points very strongly to a recent connexion of the whole with each other and the continent. But when we pass to the next island (Celebes), separated from them by a strait not wider than that which divides them from each other, we have a striking contrast; for with a total number of species less than either Borneo or Java, no less than eighteen are absolutely restricted to it. Further east, the large islands of Ceram and New Guinea have only three species peculiar to each, and Timor has five. We shall have to look, not to single islands, but to whole groups, in order to obtain an amount of individuality comparable with that of Celebes. For example, the extensive group comprising the large islands of Java, Borneo, and Sumatra, with the peninsula of Malacca, possessing altogether 45 species, has about 21, or less than half, peculiar to it; the numerous group of the Philippines possess 21 species, of which 16 are peculiar; the seven chief islands of the Moluccas have 27, of which 12 are peculiar; and the whole of the Papuan Islands, with an equal number of species, have 17 peculiar. Comparable with the most isolated of these groups is Celebes, with its 24 species, of which the large proportion of 18 are peculiar. We see, therefore, that the opinion I have already expressed, in the papers before quoted, of the high degree of isolation and the remarkable distinctive features of this interesting island, is fully borne out by the examination of this conspicuous family of insects. A single straggling island, with a few small satellites, it is zoologically of equal importance with extensive groups of islands many times as large as itself; and standing in the very centre of the archipelago, surrounded on every side with islets connecting it with the larger groups, and which seem to afford the greatest facilities for the migration and intercommunication of their respective productions, it yet stands out conspicuous with a character of its own in every department of nature, and presents peculiarities which are, I believe, without a parallel in any similar locality on the globe.
Briefly to summarize these peculiarities, Celebes possesses three genera of mammals (out of the very small number which inhabit it) which are of singular and isolated forms, viz., _Cynopithecus_, a tailless Ape allied to the Baboons; _Anoa_, a straight-horned Antelope of obscure affinities, but quite unlike anything else in the whole archipelago or in India; and _Babirusa_, an altogether abnormal wild Pig. With a rather limited bird population, Celebes has an immense preponderance of species confined to it, and has also five remarkable genera (_Meropogon_, _Streptocitta_, _Enodes_, _Scissirostrum_, and _Megacephalon_) entirely restricted to its narrow limits, as well as two others (_Prioniturus_ and _Basilornis_) which only range to a single island beyond it.
Mr. Smith’s elaborate tables of the distribution of Malayan Hymenoptera (see ‘Proc. Linn. Soc.’ Zool. vol. vii.) show that, out of the large number of 301 species collected in Celebes, 190 (or nearly two-thirds) were absolutely restricted to it, although Borneo, on one side, and the various islands of the Moluccas on the other, were equally well explored by me; and no less than twelve of the genera are not found in any other island of the archipelago. I have just shown in the present paper that, in the Papilionidæ, it has far more species of its own than any other island, and a greater proportion of peculiar species than many of the large groups of islands in the archipelago—and that it gives to a large number of the species and varieties which inhabit it, 1st, an increase of size, and, 2nd, a peculiar modification in the form of the wings, which stamp upon the most dissimilar insects a mark distinctive of their common birth-place.
What, I would ask, are we to do with phenomena such as these? Are we to rest content with that very simple, but at the same time very unsatisfying explanation, that all these insects and other animals were created exactly _as_ they are, and originally placed exactly _where_ they are, by the inscrutable will of their Creator, and that we have nothing to do but to register the facts and wonder? Was this single island selected for a fantastic display of creative power, merely to excite a child-like and unreasoning admiration? Is all this appearance of gradual modification by the action of natural causes—a modification the successive steps of which we can almost trace—all delusive? Is this harmony between the most diverse groups, all presenting analogous phenomena, and indicating a dependence upon physical changes of which we have independent evidence, all false testimony? If I could think so, the study of nature would have lost for me its greatest charm. I should feel as would the geologist, if you could convince him that his interpretation of the earth’s past history was all a delusion—that strata were never formed in the primeval ocean, and that the fossils he so carefully collects and studies are no true record of a former living world, but were all created just as they now are, and in the rocks where he now finds them.
I must here express my own belief that none of these phenomena, however apparently isolated pr insignificant, can ever stand alone—that not the wing of a butterfly can change in form, or vary in colour, except in harmony with, and as a part of, the grand march of nature. I believe, therefore, that all the curious phenomena I have just recapitulated are immediately dependent on the last series of changes, organic and inorganic, in these regions; and as the phenomena presented by the island of Celebes differ from those of all the surrounding islands, it can, I conceive, only be because the past history of Celebes has been to some extent unique and different from theirs. We must have much more evidence to determine exactly in what that difference has consisted. At present, I only see my way clear to one deduction, viz., that Celebes represents one of the oldest parts of the archipelago, that it has been formerly more completely isolated both from India and from Australia than it is now, and that, amid all the mutations it has undergone, a relic or substratum of the fauna and flora of some more ancient land has been here preserved to us.
It is only since my return home, and since I have been able to compare the productions of Celebes side by side with those of the surrounding islands, that I have been fully impressed with their peculiarity, and the great interest that attaches to them. The plants and the reptiles are still almost unknown; and it is to be hoped that some enterprising naturalist may soon devote himself to their study. The geology of the country would also be well worth exploring, and its recent fossils would be of especial interest as elucidating the changes which have led to its present anomalous condition. This island stands, as it were, upon the boundary-line between two worlds. On one side is that ancient Australian fauna which preserves to the present day the facies of an early geological epoch; on the other is the rich and varied fauna of Asia, which seems to contain, in every class and order, the most perfect and highly organized animals. Celebes has relations to both, yet strictly belongs to neither; it possesses characteristics which are altogether its own; and I am convinced that no single island upon the globe would so well repay a careful and detailed research into its past and present history.
In the following catalogue of the Malayan species of Papilionidæ I have included those from Woodlark Island, collected by M. Montrouzier, as that island comes fairly within the limits of the archipelago; while I exclude New Caledonia as belonging more to the Australian and Pacific fauna. I have given full particulars of the variation of the several species, and have described all new species, forms, varieties, and undescribed sexes. The distribution of each species is noted chiefly from my own observations[11]. As the full synonymy and references to almost every work on Lepidoptera are given in the British Museum List of Papilionidæ, I have not thought it necessary to do more than to refer to a good figure and description in well-known works; and I have quoted Boisduval’s ‘Species Général des Lépidoptères’ throughout. In all cases, however, where I have myself corrected the synonymy, or determined sexes which had been before improperly located, I have given much fuller references.
Footnote 11:
Species collected by myself have (Wall.) after the localities where I have found them.
I have found it necessary to describe and name twenty new species, and to separate six or seven more which have been hitherto considered as varieties or sexes of other species. I have also described and separated twenty-five local forms or races, and twenty polymorphous forms or sexes, as well as several simple varieties. On the other hand, I have reduced fourteen species, which figure in some of our latest lists, to the rank of sexes or local or polymorphic forms of other species. For convenience of reference, I add a list of these, with a reference to the page where will be found the reasons for not adopting them.
Ornithoptera Pronomus, _G. R. Gray_, = O. Poseidon, _Db._ (var.), p. 36. Ornithoptera Archideus, _G. R. Gray_, = O. Poseidon, _Db._ (var.), p. 36. Ornithoptera Euphorion, _G. R. Gray_, = O. Poseidon, _Db._ (♀ var.), p. 36. Ornithoptera Amphimedon, _Cr._, = O. Helena, _L._ ♀, p. 38. Papilio Hegemon, _G. R. Gray_, = P. Polyphontes, _Bd._, p. 43. Papilio Melanides, _De Haan_, = P. Theseus, _Fab._ (♀ form), p. 53. Papilio Romulus, _Cr._, = P. Pammon, _L._ (♀ form), p. 52. Papilio Rumanzovia, _Eschsch._, = P. Emalthion, _Hübn._ (♀ form), p. 48. Papilio Polytes, _L._, = P. Pammon, _L._, ♀, p. 51. Papilio Orophanes, _Bd._, = P. Ambrax, _Bd._, ♀, p. 54. Papilio Elyros, _G. R. Gray_, = P. Alphenor, _Cr._ (♀ form), p. 53. Papilio Amanga, _Bd._, = P. Ormenus, _Guér._ (♀ form), p. 55. Papilio Onesimus, _Hewits._, = P. Ormenus, _Guér._ (♀ form), p. 55. Papilio Drusius, _Cr._, = P. Gambrisius, _Cr._, ♀, p. 58.
As the arrangement of the species of _Papilio_ which I have adopted in this paper is somewhat new, and I hope will be found to be more natural than those which have been previously used, I here add lists of the Indian and Australian species arranged in the same manner. Those already included in my Malayan list will be indicated thus, (Mal.), and printed in _italics_.
_List of the_ PAPILIONIDÆ _of the Indian Region_.
1. Teinopalpus imperialis, _Hope_.
2. Ornithoptera Darsius, _G. R. G._ (Ceylon).
3. —— Rhadamanthus, _Bd._
4. —— _Pompeus_, Cr. (Mal.).
5. —— _Amphrisius_, Cr. (Mal.).
_Papilio_ (Sect. A).
_Nox group._
6. Papilio _Varuna_, White (Mal.).
7. —— Aidoneus, _Db._
8. —— Philoxenus, _G. R. G._
9. —— Polyceutes, _Db._
10. —— Dasarada, _Moore_.
11. —— Ravana, _Moore_.
12. —— Minereus, _G. R. G._
13. —— Icarius, _Westw._
14. —— Bootes, _Westw._
15. —— Janaka, _Moore_.
_Coon group._
16. Papilio Doubledayi, _Wall._
_Polydorus group._
17. Papilio Jophon, _G. R. G._ (Ceylon).
18. —— _Diphilus_, Esp. (Mal.).
19. —— Alcinous, _Klug._
20. —— Mencius, _Feld._
21. —— Hector, _L._
_Papilio_ (Sect. B).
_Protenor group._
22. Papilio Protenor, _Cr._
23. Papilio Elphenor, _Db._
24. —— Rhetenor, _Westw._
25. —— Sakontala, _Hewits._
_Peranthus group._
26. Papilio Crino, _Fab._ (Ceylon).
27. —— Bianor, _Cr._
28. —— Polyctor, _Bd._
29. —— Ganesa, _Db._
30. —— Arcturus, _Westw._
31. —— Paris, _L._
32. —— Palinurus, _Fab._?
33. —— Krishna, _Moore_.
_Memnon group._
34. Papilio _Androgeus_, Cr. (Mal.).
35. —— Polymnestor, _Cr._ (Ceylon).
36. —— Demetrius, _Cr._
_Helenus group._
37. Papilio _Helenus_, L. (Mal.).
38. —— Chaon, _Westw._
39. —— Castor, _Westw._
40. —— _Nephelus_, Bd. (Mal.).
_Pammon group._
41. Papilio _Pammon_, L. (Mal.).
_Demolion group._
42. Papilio _Demolion_, Cr. (Mal.).
_Papilio_ (Sect. C).
_Erithonius group._
43. Papilio _Erithonius_, Cr. (Mal.).
_Paradoxa group._
44. Papilio Telearchus, _Hewits_.
45. —— Slateri, _Hewits_.
_Dissimilis group._
46. Papilio dissimilis, _L._
47. —— Panope, _L._
48. —— Lacedæmon, _Fab._
49. —— Pollux, _Westw._
_Papilio_ (Sect. D).
_Macareus group._
50. Papilio _Macareus_, God. (Mal.).
51. —— _Leucothoë_, Westw. (Mal.).
52. —— Megarus, _Westw._
53. —— Agestor, _G. R. G._
54. —— Epytides, _Hewits_.
55. —— Xenocles, _Db._
_Antiphates group._
56. Papilio _Antiphates_, Cr. (Mal.).
57. —— Agetes, _Westw._
58. —— Anticrates, _Db._
59. —— Orestes, _Fab._
60. —— Alebion, _G. R. G._
61. —— Glycerion, _G. R. G._
_Eurypylus group._
62. Papilio Gyas, _Westw._
63. —— Evan, _Db._
64. —— Cloanthus, _Westw._
65. —— _Sarpedon_, L. (Mal.).
66. —— Chiron, _Wall._
67. —— _Jason_, Esp. (Mal.).
68. —— _Agamemnon_, L. (Mal.).
69. —— _Rama_, Feld. (Mal.).
4. Chinese species.
61. Indian species.
4. Ceylon species.
_List of the_ PAPILIONIDÆ _of the Australian Region_.
_Ornithoptera (Priamus group)._
1. Ornithoptera _Poseidon_, Db. (Mal.).
2. —— Richmondia, _G. R. G._
_Papilio_ (Sect. A).
_Polydorus group._
3. Papilio _Leodamas_, Wall. (Mal.).
4. —— _Liris_, Godt. (Mal.).
5. —— Godartianus, _Bd._ (Pacific Islands).
_Papilio_ (Sect. B).
_Helenus group._
6. Papilio Capaneus, _Westw._
7. —— Ilioneus, _Don._
_Ulysses group._
8. Papilio Ulyssinus, _Westw._
9. —— Montrouzieri, _Bd._ (New Caledonia).
_Pammon group._
10. Papilio Canopus, _Westw._
_Erectheus group._
11. Papilio Erectheus, _Don._
12. —— Amyntor, _Bd._ (New Caledonia).
_Papilio_ (Sect. C).
_Erithonius group._
13. Papilio _Erithonius_, Cr. (Mal.).
_Anactor group._
14. Papilio Anactor, _McL._
_Papilio_ (Sect. D).
_Antiphates group._
15. Papilio Leosthenes, _Db._
16. —— _Permatus_, G. R. G. (Mal.).
_Eurypylus group._
17. Papilio _Sarpedon_, L. (Mal.).
18. —— Gelon, _Bd._ (New Caledonia).
19. —— Lycaon, _Westw._
20. —— Macleayanus, _Leach._
21. —— Scottianus, _Feld._ (Ash Islands).
22. Eurycus Cressida, _Fab._
6. Pacific Islands.
16. Australia.
_Catalogue of Malayan_ PAPILIONIDÆ.
ORNITHOPTERA (Boisd.).
[Illustration:
Fig. 1.
Anal valves of _O. Amphrisius_. ]
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.