Art. XXVI.—On the Geographical Relations of the New Zealand Fauna.
[Read before the Wellington Philosophical Society, 4th and 11th September, 1872.]
I know of no part of the world that presents such a promising field to the student of Nature as New Zealand. Although small in size it contains a fauna and flora so peculiar that several naturalists consider it as a separate biological province apart from the rest of the world. Isolated from any large continental area longer probably than any other portion of the earth, it contains the remnant of the population of a continent that existed before the Mammalia had overspread the world, and to that has at various times been added, principally from Australia, a colonist population which culminated not many hundreds of years ago in the advent of man. New Zealand, therefore, presents us with what I may call the elements of a continental fauna, or a
continental fauna in its simplest state, and consequently in that state which is most advantageous for studying the mutual relations of the animals composing it.
Both Mr. Darwin and Mr. Wallace call New Zealand an “oceanic island” from a zoological point of view, owing to the absence of terrestrial Mammals, and the meagreness of its fauna and flora; that is to say they consider it as an island that has never formed part of a continental area since its last emergence from the sea. But I think that the Struthious birds have certainly as much weight in determining this point as terrestrial Mammals, for they have no superior means of dispersion, and New Zealand also possesses a frog, which is one of the great characteristics of a continental fauna. From a geological point of view I do not see how any land, except volcanic and coral islands, could have originated except as part of a large continental upheaval. I think, therefore, that the New Zealand fauna may be correctly called the remnants of a continental fauna, and that a close study of it will throw great light on many of the most important, but at the same time most obscure, problems in zoology. It will, however, be long before this can be accomplished. The describing and naming of the different animals, which is the foundation upon which all other researches must rest, is as yet far from being completed; the determination of what species are the original inhabitants, or the descendants of the original inhabitants, of the former continent, has hardly been attempted, but all this must be settled before any sound deductions can be drawn as to the reasons of extinction, variation, or permanency of type, of the animals.
It is to this latter point that I wish to draw attention, not that I am in possession of information sufficient to prove any one, perhaps, of the points that I shall raise, but because I think that sufficient is known to establish with great probability the main features in the zoological history of these islands, and this sketch, which I now presume to offer you, will I hope induce others to examine the subject more in detail, and will give a systematic direction to their observations. I propose to take first the zoological evidence—to point out the principal facts that have to be accounted for, and the deductions that they lead to. I will then rapidly glance at the geological and palæontological evidence, and finally I will draw up from the whole the hypothesis that appears best able to account for all the phenomena.
Of our two bats one (Scotophilus tuberculatus), although not found elsewhere, is closely allied to those of Australia, while the other (Mystacina velutina) forms the only species of a genus peculiar to New Zealand, but related to bats living in South America.
Two species of seal frequent our shores; the sea leopard (Stenorhynchus leptonyx) which is also found on ice floes in the Antarctic seas, and occasionally extends to Australia, and the fur seal (Arctocephalus cinereus), which is supposed to occur also on the southern coasts of Australia, and is closely related to, if not identical with, a species found at the Falkland Islands, Cape Horn, South Shetland, and South Georgia. In the Otago Museum there is also a skull that appears to belong to the sea elephant (Morunga proboscidea). Mr. Purdie informed me that it was picked up a long way inland.
Of the Cetacea some twelve or thirteen species are known, belonging to the six different families into which the marine members of this order have been divided, and it is remarkable that two thirds of them are endemic, that is not found anywhere else. Our two or three species of whale-bone whale have, up to the present, been found nowhere else. The sperm whale of our northern coasts is probably the same species as that found in Australia and the South Pacific (Catodon australis). It is certainly distinct from the northern sperm whale (C. macrocephalus) as the lower jaw is much narrower.*
Our ziphoid whales, of which we have three or four species, are all endemic, and two of them (Berardius arnuxii and Mesoplodon hectori) belong to genera not found elsewhere. None, however, of our Delphiniidœ are confined to New Zealand. Delphinus novœ-zealandiœ inhabits the antarctic seas, and perhaps Tasmania; Lagenorhynchus clanculus is found throughout the Pacific Ocean, but not in Australia, and Orca capensis, a lower jaw of which is in the Auckland Museum, ranges from the Cape of Good Hope through the Southern Ocean to Chili, and is also found in the North Pacific and Tasmania. The black-fish (Globiocephalus macrorhynchus) is found in the South Pacific and Japan, but not in Australia. Our Cetacea therefore, contrary to what might have been expected, show a nearer relation to the Pacific and Antarctic Oceans than they do to Australia, and it is remarkable that no species of porpoise has as yet been described as found in New Zealand, although two inhabit Tasmania.
The absence of terrestrial Mammalia is one of the chief points of interest in New Zealand zoology, as it proves that there has been no land communication between this country and Australia since the latter was inhabited by Marsupials, for I consider that the so-called Maori rat and native dog were both introduced by human agency.†
[Footnote] * Capt. Cook remarks in his first voyage that rats were “so scarce that many of us never saw them.” (Hawkesworth's “Coll. of Voy.,” III., p. 34.) He makes no mention of them ever being used for food, and I am not aware of any remains of rats having been as yet found in Maori cooking places.
[Footnote] † A lower jaw of the New Zealand sperm whale in the Auckland Museum is 17 ft. 7 in. in length, and only 2 ft. 2 in. in width at the condyles; there are 23 teeth on each side, 4 of which are rudimentary only; the length of the largest tooth is 7.4 in.
Sir George Grey informs me that he sent to the British Museum some grey “Maori rats” which had been caught in the interior of the South Island in 1847 by Mr. Torlesse, and that Dr. Gray had said that they were identical with a rat found in Polynesia, by which he must have meant the black rat (Mus rattus) for none of the islands in the Pacific possess an indigenous rat. Dr. Buller also collected a considerable amount of evidence to show that the “kiore-maori” was identical with a rat—now in the Colonial Museum—which he described (Trans. N.Z. Inst., III., p. 1) under the name of Mus novœ-zealandiœ, but which is certainly Mus rattus. Mr. Colenso says (“Proc. R. Soc. of Van Diemen's Land,” 1851, p. 301), in a letter to R. Gunn, Esql., dated 4th Sept., 1850, that after considerable trouble he had procured two specimens of the native rat, which he describes as “smaller than our English black rat (M. rattus) and not unlike it.” Against this we have the statement of Dr. Dieffenbach, who says (“New Zealand,” II., p. 185) that it was the English and not the Norway rat that killed off the “kiore-maori.” This, I think, must be a mistake, as all the Maoris attribute the destruction of the edible rat to the brown rat, and it could only have been from Maoris that Dr. Dieffenbach got his information. Mr. Murray also states (“Distr. of Mammals,” p. 277) that the Norway rat (M. decumanus) was not introduced into New Zealand in 1843, but he gives no evidence of the truth of this statement, and it is unquestionably erroneous.* The whole of the reliable evidence that we have, therefore, goes to prove that the Maori rat was no other than M. rattus.
The so-called “native dog” has been determined by Dr. Gray to be Canis familiaris (“Pro. Zool. Soc.,” 1868, p. 508), and not the Australian species, or variety, called Canis dingo, which is the strongest possible evidence of its being merely an escaped domestic breed; indeed, I am not aware that any naturalist believes in an indigenous native dog except Dr. Haast, who has argued (Trans. N.Z. Inst., IV., p. 88) that a wild dog existed in New Zealand before the domesticated one, because in certain old Maori cooking places he has found remains of the dog but no gnawed bones, while in others, which he considers as of later date, he finds gnawed bones.† But I am not aware that
[Footnote] * Since reading this paper Mr. Nichol has informed me that the brown rat was common in Nelson when he first arrived in the early part of 1842, and that he never saw any other kind there except a single specimen of a very large and slightly striped variety.
[Footnote] † The skulls of dogs found in old Maori cooking-places prove undoubtedly that Canis familiaris existed in New Zealand long before Europeans came here. Captain Cook says (21st October, 1769) that the dogs were “small and ugly,” and Mr. Anderson (“Cook's 3rd Voyage,” I., 153) calls it a “sort of fox-dog.” Capt. Cook also says in his first voyage that the dog was used for no other purpose than to eat. The fact that the inhabitants of the Friendly Islands have the same name (kuri) for the dog as the New Zealanders is strong evidence that the latter brought it with them, for if not they would have lost the name as they have done that of the fowl.
he has any proof of the existence of a dog in New Zealand before the arrival of man, and the difference of date of these cooking-places for which Dr. Haast contends, is denied by many observers, and his argument derived from the presence or absence of ground stone implements has, I think, been successfully controverted. I can therefore attach no weight to the absence of gnawed bones. On the other hand, there is the fact that no indigenous dog or rat has ever been found on an island that was not inhabited by other Mammalia, and when we remember that Marsupials came into existence long before rats and dogs, it is difficult to see how the latter could possibly get to any country without the former coming also. It is evident that neither Banks, nor Solander, nor the Forsters, considered the dog and rat that they found in New Zealand as a new species, or they would certainly have mentioned it; neither did Lesson in 1827, nor Quoy and Gaimard in 1831. Dr. Dieffenbach, in 1842, was the first to state that a frugivorous rat, distinct from M. rattus, existed in New Zealand; he, probably, not being aware that M. rattus is entirely frugivorous. I am therefore of opinion that both the rat and the dog were brought by human agency, and it is worth remarking that the Maori traditions relate that they brought both with them. (Travers, Trans. N.Z. Inst., IV., p. 58.) The specimen of Mus gouldi in the Auckland Museum (see Trans. N.Z. Inst., III., p. 3) was caught, I believe, at the Thames in January, 1853, and as a mission station had been established there some years previously this specimen was no doubt brought over from Australia in their vessel.
The animal seen at Dusky Bay by some of Capt. Cook's sailors (2nd Voyage, I., 98) was probably a dog, as none on board had at that time seen a dog in New Zealand.
The evidence of a kind of otter inhabiting the South Island rests upon some foot-prints seen by Dr. Haast, and mentioned by him in his first presidential address to the Canterbury Philosophical Society (“Nat. Hist. Rev.,” 1864. p. 30). In the same address he also mentions having seen tracks in great numbers of a small jumping mammal in the riverbed of the Hopkins, but as no further evidence of the existence of these creatures has been adduced, although eight or nine years have since elapsed, it is impossible for me to take any further notice of them in this paper.
The first point that claims our attention here is the great development of the Struthious birds. This division can be subdivided into two families, one (Apterygidœ) containing only the kiwis, and the other (Struthionidœ) including all other living forms as well as the extinct moas. The kiwis in the structure of the egg-shell have an affinity with the Carinate division of birds. Their
short legs, and the presence of a hind toe elevated above the level of the others, shows an approach to the Gallinaceous order, while their long bill, with its slightly swollen tip, resembles in some measure that of the Scolopacidœ, which have also the same habit as the kiwi of feeling about on the ground with their bill. Gallinago pusilla moreover lives in holes, and only comes out at night (Travers, see Art. xxii).
Thus the Apterygidœ have a more generalised structure than the other Struthious birds; they therefore belong to an older type, and cannot with any degree of correctness be said to represent the extinct race of moas. The relations between the second family, or the Struthiones proper, are very complicated, but Dinornis, which alone concerns us here, appears to be intermediate between the rheas of South America, and the emus and cassowaries of Australia and the adjacent islands. It approaches the rhea in the structure of its egg-shell and in having only three pairs of sternal ribs, while the emu, the cassowary, and also the kiwi, have four, and the ostrich five pairs. In the structure of its feathers, and in the shape of its pelvis and skull the moa approaches the emu. The Struthious birds exhibit a type of structure intermediate in many respects between the Carinate birds and the extinct Dinosaurians, and this leads naturalists to suppose that they are but the remnant of a race that once spread over the whole earth. About twelve species are known outside New Zealand; while here, besides our four species of Apteryx, Professor Owen has determined fourteen species of Dinornis, three of Aptornis, and one of Cnemiornis, thus making a total of twenty-two species of Struthious birds, belonging to four different genera, living in New Zealand only a few hundred years ago, that is to say, nearly twice as many as are found in all other parts of the world put together.
Probably, however, some of Professor Owen's species of Dinornis are but the young of others, and it seems to me very doubtful whether Aptornis and Cnemiornis should be regarded as Struthious birds at all. It is evident that these two genera are closely related, and if the wing bones placed upon Cnemiornis calcitrans really belong to the legs of the same bird we must suppose that the sternum had a keel sufficiently developed to support muscles of a size proportionate to the wings; for although we can understand how the kakapo (Stringops), belonging to an order of deeply keeled birds, may have lost, by disuse of the pectoral muscles, the keel on its sternum, we cannot possibly explain how a Struthious bird could have had large wing bones developed unless it had also sufficiently powerful muscles to use them. I also observe that Aptornis defossor now wears a skull similar to that of the late Dinornis casuarinus, which skull Mr. W. K. Parker says undoubtedly belonged originally to a Notornis. But omitting these two genera, and making a due allowance for doubtful species of Dinornis, the great number of
species living on so small an island is very remarkable when contrasted with other parts of the world. The continent of Africa, including Arabia, contains but one, or according to some naturalists two, species of ostrich. South America, from Patagonia to Peru, has but three species of rhea, each inhabiting a separate district. Australia possesses two species of emu, one on the eastern and the other on the western side, and one species of cassowary on the northern, while five other species of cassowary inhabit other detached islands, from New Britain and New Guinea to the Molucca Islands. I believe that outside of New Zealand no two species of Struthious bird are found living in the same district, while here we have now four species of kiwi and not long ago had at least half-a-dozen species of moa as well. How can this be accounted for? The solution is readily found by examining the distribution of the cassowaries. Here we have six species inhabiting six isolated localities. If now this region of the earth were to be elevated these six species might mingle, and if it were subsequently to sink again, all six species would undoubtedly be driven to the higher lands, and we should have in this supposed island a representation of New Zealand inhabited by six species of Struthious bird.
In order, therefore, to account for the numerous species of Dinornis we must suppose an ancient continent, inhabited by one or two species, to sink, and the birds to take refuge on the different mountain ranges left as islands above the water. We must suppose that they remained thus isolated from one another for a sufficiently long period to allow of specific changes being brought about; that then, by an elevation of the land they once more mingled together, and that, on subsidence again taking place, New Zealand as the central mountain chain formed a harbour of refuge for them all.
Whether this isolation of species points to some cause as yet unrecognized, by which in the struggle for life no two species of Struthious bird can live in close proximity I will not venture to give an opinion, but it is a fair subject for inquiry, and one on which the careful study of the relative ages of moa bones might throw considerable light, and enable us perhaps to understand the great mortality that must have taken place amongst the moas when confined to these small islands long before man set his foot here.
The distribution, therefore, of the Struthious birds in the Southern Hemisphere points to a large Antarctic Continent stretching from Australia through New Zealand to South America, and perhaps on to South Africa. This continent must have sunk, and Australia, New Zealand, South America and South Africa must have remained isolated from one another long enough to allow of the great differences observable between the birds of each country being brought about. Subsequently New Zealand must have formed part of a smaller continent, not connected either with Australia or South America,
over which the moa roamed. This must have been followed by a long insular period, ending in another continent still disconnected from Australia and South America, which continent again sank and New Zealand assumed somewhat of its present form.
Passing now to the Carinate division of birds the first thing that strikes us is the fragmentary nature of this part of our avi-fauna (if we exclude the Grallæ and Web-footed birds), thus strongly contrasting with the Struthious division.
Of the first six orders we possess, excluding the Chatham and Auckland Islands, forty-five species, thirty of which are endemic. These have been referred to thirty-one genera, ten of which are found nowhere else, and these thirty-one genera belong to twenty families, one of which (Stringopidœ) is peculiar to New Zealand. Two families only, the honey-eaters (Meliphagidœ) and the starlings (Sturnidœ) contain more than two genera. The first shows affinity to Australia, but it must be remarked that out of the four species of this family, belonging to four different genera, one genus only (Zosterops) is found in Australia, and the little bird (the “white-eye”) that belongs to this genus is known to be quite a recent arrival in this country. The Sturnidœ on the other hand show an affinity with Polynesia, for one species only (Calornis metallicus) of this family is found in the north of Australia and New Guinea. It should, however, be noticed that three other species are found in the latter island. In this family also our three species belong to three different genera, two of which (Creadion and Heteralocha) are found nowhere else, while the other (Aplonis) is very characteristic of Polynesia, and Aplonis caledonicus, which is said to have been found in New Zealand, occurs also in Norfolk Island and New Caledonia.
It is remarkable that our two owls should both be peculiar to New Zealand, and that one of them (Sceloglaux albifacies) should belong to a genus not found elsewhere, for the owls are usually widely spread birds, more so indeed than the hawks. It is also worthy of notice that Strix delicatula, which extends its range over most of the Pacific Islands and Australia, should be absent from New Zealand.
Our parrots present several points of interest. The kakapo (Stringops habroptilus) is found nowhere else, the genus Nestor extends only to Norfolk Island, while our paroquets, although belonging to a genus (Platycercus) equally plentiful both in Australia and Polynesia, show a greater affinity to the latter, one species (P. novœ-zealandiœ) ranging not only to Norfolk Island but also to New Caledonia. It is remarkable that we have no representatives of the cockatoos and grass-paroquets so common in Australia and Tasmania, for our own climate is quite suitable for them. The absence of Polynesian forms is not so remarkable as they belong chiefly to more tropical
genera, and the members of the genus Coriphilus are said to live only on bananas.
That we should have two cuckoos which migrate regularly to other countries, each more than a thousand miles distant, is a fact that deserves special attention, for I know of no parallel case in any other part of the world, the distance across the Mediterranean being less than half that travelled over by our summer visitants. The phenomenon of a bird at a certain season of the year flying out to sea to an island more than a thousand miles distant is remarkable enough, but is rendered still more so in the case of the little shining cuckoo (Chrysococcyx lucidus), which is supposed to come from Australia, by there being no apparent necessity for it. For this bird migrates east and west, and not from a warmer to a colder climate, and two other closely allied species which inhabit Australia never leave the country at all. Even in the case of the long-tailed cuckoo (Eudynamis taitiensis) which comes to us from the equable climate of the South Sea Islands, we cannot suppose that its migrations are caused either by alteration of temperature or by want of food, and the question forces itself upon us—How could this habit have arisen? The only reasonable hypothesis is, I think, that at one time the different lands to and from which these birds fly were connected, or nearly so; that the distance between them gradually increased, and that the habit, so common amongst birds, of resorting each year to the same place to breed, was not lost but gradually merged into a regular migration. From this point of view the arrival of the shining cuckoo indicates a connection with Australia or perhaps New Guinea, while that of the long-tailed cuckoo indicates one with Polynesia, and it must be noticed that while the latter bird is identical with specimens from Polynesia, the former shows such differences in the colouring of the tail feathers from the birds inhabiting Australia that it is considered by many naturalists to be a distinct species. Another remarkable fact, that has been quite lately brought to light, is that the shining cuckoo of the Chatham Islands is not the same variety as that visiting New Zealand, but is almost, if not quite, identical with an Australian species (C. plagosus). This curious fact proves how strong must be the force of habit, for these birds in their migration to and from the Chatham Islands must pass over, or at least in sight of New Zealand, but instead of stopping, after a journey of 1,400 miles, they continue on for 450 miles more, until they reach the little island that they have selected as their home.
A more difficult fact to account for is the presence of different species of grass-bird (Sphenœacus) in both Australia and New Zealand, for this bird has such feeble powers of flight that it could not cross a river, and must almost of necessity have travelled by land. It must, however, be noticed that this
genus extends through the Indian Archipelago into India, and I have not been able yet to compare our grass-birds with those of Australia and the Archipelago so that I am not able to say what amount of difference there is between them. The genus Keropia has most affinity with South American birds, while Graucalus melanops, which is closely related to our G. concinnus, is said to extend from Australia into New Guinea.
In the order Grallœ, or Waders, we come to birds more widely spread than any others, some indeed being almost cosmopolitan, but even amongst these the isolated character of our fauna is still marked, for out of twenty-eight species, belonging to seventeen genera, eight species and two genera are found nowhere else. The most noticeable feature in this order is the existence of a curious genus of rails (Ocydromus) quite unable to fly. Of this genus we possess four species, one in the North and three in the South Island, while a fifth species is found in Lord Howe Island, and a sixth in New Caledonia. Notornis, although somewhat like the pukeko (Porphyrio melanotus) in the bill, has the feeble wings, thick legs, and short toes of Tribonyx mortierii of Tasmania and Australia. Of our other rails two (R. pectoralis and O. tabuensis) are spread over Australia and Polynesia, while another (O. affinis) although not found elsewhere is closely related to a species from Australia (O. palustris). In the godwit (Limosa uropygialis) we have another migratory bird that probably comes from Polynesia, but as it is also found in Australia we cannot feel any certainty about it. New Zealand also displays the peculiarity of being the only country in the world inhabited by two species of stilt-plover (Himantopus) one of which (H. novœ-zealandiœ) is found nowhere else. This is probably owing to the length of time that New Zealand has been isolated, and to its having had during the whole of the period a stilt-plover on it, which gradually changed until it attained that remarkable jet black plumage which is so different from any other species, while the later colonist from Australia (H. leucocephalus) displays the colour usual to the genus. This view is rendered the more probable by the fact that the young of the black stilt-plover have the same pied plumage that is exhibited by the adults of those species from one of which I suppose it to have been derived.
In the crook-bill (Anarhynchus frontalis) we have another curious anomaly which as yet has received no explanation; and it must also be noticed that Cape Horn, the Cape of Good Hope, Australia, and New Zealand, each possess a black oyster-catcher (Hœmatopus), which are considered specifically distinct.
Among the herons the only very remarkable fact is the occurrence of the little bittern (Ardea pusilla), a bird found only in Australia and Natal. Our snipe (G. pusilla) very much resembles in plumage G. stricklandi from Tierra del Fuego, but it has a shorter bill.
Among the web-footed birds, the first thing that claims our attention is the oceanic family of the petrels (Procellaridœ), for although by no means peculiar to New Zealand, * the great number of species in the southern oceans, in comparison with the small number in the northern, is very noticeable. The northern and tropical species have all closely allied forms in the southern hemisphere, while many of the southern petrels, such as Ossifraga, Halodroma, Majaqueus, Pterodroma, Daption, and Prion have no representatives in the northern seas. This leads to the inference that the northern species have been derived from stray southern birds, and that the southern hemisphere has been the centre from which most oceanic birds have spread, while land birds, on the contrary, have spread chiefly from northern areas, and this leads to the further inference that the southern hemisphere has been for many ages more oceanic in character than the northern. The next most remarkable point is the great development of the cormorants, New Zealand possessing nine species, four of which are found nowhere else. No other country in the world possesses so many, and the phenomenon can only, I think, be accounted for in the same way as the numerous species of moa, that is, by the former existence of several small islands which have since been elevated to form the present New Zealand. The wide dispersion, however, of two of our cormorants is rather against this view, one (G. carunculatus) being found at the Crozet Islands and at Cape Horn, and the other (G. carbo) in Australia, China, and Europe. I must, however, remark that the identity of the first has not yet been perfectly established, and that the second, although very closely resembling specimens from Europe, shows at the same time some difference. It may also be useful to remark here that our gannet (Dysporus serrator), although a far better flying bird than the cormorants, is not found at the Chatham Islands, and Dr. Finsch informs me that it is undoubtedly different from the species (D. capensis) that occurs at the Cape of Good Hope. The occurrence of G. brevirostris and G. melanoleucus in New Zealand presents a parallel case to the two species of stilt-plover, with, however, this difference—that, judging from the colours of the young bird, it is probable that G. melanoleucus has been derived from G. brevirostris, owing to its having been isolated in Australia, and that its descendants have migrated back again to New Zealand.
Of the gulls we possess a species (L. pomare) which is found nowhere else, a peculiarity of which few countries can boast, but which can perhaps be accounted for by the fact that this gull only frequents fresh-water lakes, and seldom comes down to the sea. Our other gulls are widely spread, but it is a most remarkable fact, which at present appears to me to be quite inexplicable, that neither gulls nor cormorants occur in any of the Polynesian Islands.
Of ducks we possess nine species, four, or perhaps five of which are
[Footnote] * Procellaria parkinsoni is peculiar to New Zealand.
endemic; one, the blue duck (H. malacorhynchus), belonging to a curious genus found only in New Zealand, but related to a genus (Malacorhynchus) in Australia. The others are all found in Australia, one (P. gibberifrons) ranging through New Caledonia and the Indian Archipelago, and another, the common grey duck (A. superciliosa), spreading over Polynesia, as far north as the Sandwich Islands. The most remarkable circumstance connected with our ducks, is the presence of a species of Fuligula, a genus found neither in Australia nor Africa, but belonging properly to the northern parts of America, Europe, and Asia, although one species is found in South America. The occurrence, however, of a northern species (F. cristata) in the Pelew Islands points out to us perhaps the route along which the ancestors of our species travelled.
The Chatham Islands possess thirty-two species of birds, omitting the gulls, penguins, and petrels, of which six are found nowhere else. All the others are found in New Zealand, except the shining cuckoo (C. plagosus), which, as already stated, migrates to and from Australia. No genera, however, are peculiar to these islands, except perhaps a rail (Rallus? modestus) which is evidently incapable of flight, and which will probably have to be placed in a genus by itself. This curious form must not, however, be regarded as a change produced by long isolation, but rather as an old form preserved from destruction by isolation. The most noticeable circumstance in the Chatham Island fauna is the absence of Raptores, with the exception of an occasional visit from the harrier (Circus gouldi), which does not however appear to inhabit the islands, or at any rate is exceedingly rare there.
The Auckland Islands possess twelve birds, three or four of which are endemic, the remainder all belonging to New Zealand. The most remarkable facts are the occurrence of a species of merganser (Mergus australis), a genus found only in high northern latitudes, and of a duck (Nesonetta aucklandica) with very short wings, belonging to a genus found nowhere else.
On Norfolk Island we know of twenty-six birds. Of these two (Aplonis caledonicus and Platycercus novœ-zealandiœ) are found in New Zealand and New Caledonia; five others are common to New Zealand and Australia; a species of Nestor (N. productus) used to inhabit Philip Island close by, and the remainder show an affinity to Australia.
Lord Howe Island possesses only six land birds, two of which (Charadrius bicinctus and Ocydromus sylvestris) show a connection with New Zealand, while the rest show an affinity to Australia.
A review of the facts disclosed by a study of the distribution of the Carinate birds shows that although the affinity is greater with Australia than with any other place, there is yet a decided leaning towards Polynesia, and when we remember that a large portion of Australia lies in the same latitude
as New Zealand, while the whole of Polynesia is far away to the north, I think the difference is not so great as might have been expected. * The distribution of the genus Ocydromus proves that land communication must once have existed between New Zealand, Lord Howe Island, and New Caledonia, but the absence of cockatoos, grass-paroquets, pigeons, night-jars, and finches, indicates that this connection did not extend to Australia. With the exception of Sphenæacus, which has very feeble powers of flight, all our Australian birds could have crossed over a strait of considerable width. The phenomena of the paroquets, starlings, and long-tailed cuckoo of Polynesia, being associated in New Zealand with the honey-eaters, grass-bird, and gold-cuckoo of Australia, indicate that New Zealand was connected with a tract of land intermediate to both, but perhaps not connected with either; at-the same time the absence of the more tropical Polynesian birds is no evidence, that this tract of land did not extend into Polynesia, and in Zosterops lateralis, and Dendrocygna eytoni, both of which have appeared since Europeans came into the colony, we have positive evidence that our islands can even now be colonized from Australia by many kinds of birds, although 1,400 miles distant. It would also appear that this transfer of birds to New Zealand took place sufficiently long ago to allow of changes of generic value having taken place, while the Chatham and Auckland Islands have been isolated from New Zealand for a time sufficient only for changes of specific value.
The Reptiles of New Zealand are not numerous. We possess about eight species of lizards, four of which belong to widely spread genera of the family Scincidæ, but the species are all endemic. Three others belong to the Geckoidæ, and form a genus (Naultinus) which is found nowhere else. Of these one (N. pacificus) is said to be found in some of the Pacific Islands, but the other two are peculiar to New Zealand. Our eighth species, the curious tuatara (Sphenodon punctatum), which is now found only on a few rocky islets in the Bay of Plenty, and near Tory Channel in Cook Strait, is placed by Dr. Günther in a separate order from all other lizards on account of the affinity that it shows to the crocodiles. This remarkable form has no copulatory organs, and has uncinate processes on its ribs like birds. It has also nearly twice as many abdominal as true ribs, which protect the abdomen when being dragged along the ground, for, like the crocodile, the hind legs are too weak to support the hinder parts of the body. Dr. Günther also suggests that they may use them for locomotion, as snakes do. It is also remarkable
[Footnote] * The distribution of the Megapodidœ shows that Polynesia, Australia, the Indian. Archipelago as far as the Strait of Lombok, North-west Borneo, and the Philippine Islands, were united before the spread of the Mammals.
that this animal, which lives in holes and only comes out during warm weather, should have the dorsal crest that is so characteristic of tree lizards.
I omit all reference to Norbea? isolata, supposed to come from White Island, in the Bay of Plenty, because its true locality is not sufficiently well established; if, however, another specimen should be obtained, it would be most important evidence in the present discussion.
But one species of lizard is found on the Chatham Islands, which is very variable, but which I consider to belong to the species Mocoa zealandica; it is, however, larger, and shows some slight differences in the shape of its cephalic shields.
A ringed sea-snake, probably Platurus scutatus, of Australia and Polynesia, is sometimes washed alive on to our coasts as far south as the mouth of the river Waikato, but it is not yet ascertained whether it is an inhabitant of our seas. A peculiar variety of Pelamis bicolor, which as yet has not been found in any other locality, has also been taken on our shores.
The amphibious animals are worse represented even than the reptiles; one species of frog (Liopelma hochstetteri) being the only member of the class. This frog has now been found in three distinct localities, all, however, in the province of Auckland; these are the Cape Colville ranges from Coromandel to Puriri, the Huia on the north side of the Manukau harbour, and in the mountains behind Opotiki in the Bay of Plenty. It belongs to a genus not found elsewhere, but its nearest ally is Telmatobius peruvianus from Peru, and it should be remembered that the frogs of Australia are also allied to South American forms. It is evident that the absence of other Batrachians cannot be accounted for by the unsuitability of climate or want of food, for the common green frog of Australia (Litoria aurea), which has been introduced, has spread with great rapidity around both Auckland and Christchurch.
The evidence of the reptiles is, therefore, that New Zealand has had land communication with some of the Pacific Islands at a later date than with Australia, for in the first case there is no specific difference between forms found in both places, while in the latter the species are now quite distinct. Our frog proves a connection with South America at a period so remote that changes have since taken place of generic value.
Up to the present time about 134 species of marine fish are known to inhabit the shores of New Zealand. Of these 51, or 37 per cent, are found nowhere else. Thirty-eight extend to the Australian and Tasmanian seas, but no furthe, six range to the Pacific Islands, five inhabit South America, four South Africa, and one Kerguelen Land and the Auckland Islands. There
Hutton.-—On the Geographical Relations of the N.Z. Fauna. are also four others that are common to both Australia and South America, five common to Australia and South Africa, two common to Australia and the Pacific Islands, and one common to Australia and the Auckland Islands. Thus the total number of our sea fishes found in Australia is fifty, in South America and the Cape of Good Hope nine each, three (Prosopodasys cottoides, Trygon kuhlii, and Ostracion fornasini) are not found nearer than the Indian Archipelago (the identification, however, of the latter is doubtful), and one (Halargyreus johnsoni) has been obtained at Madeira only. The remaining thirteen are widely ranging species. These 134 species have been distributed among 114 different genera, eleven of which are not found elsewhere. The connection with Australia is here, as might be expected, so well marked that I need not dwell upon it, but will proceed to examine the affinities of New Zealand to other countries. Our former connection with South America is indicated by Mendosoma lineata, Notothenia cornucola, Merluccias gayi, and Genypterus blacodes; with South Africa by Trigla kumu, Gonorhynchus greyi, and Bdellostoma cirrhatum, while the occurrence of Gonorhynchus greyi and Congromuræna habentata at St. Paul's shows that that little volcanic island was also probably connected. The occurrence in New Zealand of species belonging to the southern genera Pseudorhombus, Bovichthys, Agriopus, Chilodactylus and Scorpis points to the extension of a former antarctic continent, of which these islands formed a part, while Acanthurus triostegus, Dascyllus aruanus, Chanos salmoneus, Peltorhamphus novæ-zealandiæ, a species of stingaree allied to Trygon thalassia, and species of the genera Labricthys and Trachelochismus, show an affinity for the islands of the Pacific.
I have already remarked that three of our fishes are not found nearer than the Indian Archipelago, and it is probable that our species of Torpedo and Doryichthys came from that direction also. But a still more curious affinity to Japan is shown by the presence of the genera Lotella and Ditrema, and another little fish (Calloptilum punctatum) which is found at the mouth of the river Thames, and which has its nearest allies in the genus Bregmaceros from China and the Philippine Islands. Gonorhynchus greyi and Clupea sagax are also both found in Japan, but they occur in Australia as well. Our species of Ditrema differs from D. læve of Japan in having teeth on its palate, and a band of teeth in each jaw instead of a single row. Platystethus cultratum, from Norfolk Island, is also closely allied. This connection with China and Japan is, I consider, the chief point of interest in the distribution of our marine fish.
In the genus Trypterygium, which is found only in the Mediterranean, we have an anomaly which is parallel to the cases of Fuligula and Mergus among the birds, and as we proceed we shall find many other similar cases cropping up.
The fresh-water fish naturally supply more important evidence as to the former distribution of land than those inhabiting the sea. Of these, New Zealand possesses fifteen species, belonging to seven genera, of which six species, or 40 per cent., and one genus, are found nowhere else. That the percentage of the endemic fresh-water fish should be nearly the same as that of the marine fish is a remarkable and unexpected result, for the number of species of marine fish inhabiting New Zealand and found also in other countries depends partly on permanency of specific characters since New Zealand was isolated, and partly on the power possessed by fishes of migrating to us from other countries, while among the fresh-water fish the proportion depends entirely on permanency of specific characters; consequently, this permanency of specific characters must be greater in fresh-water than in salt-water fish, and this is the more remarkable as-our fresh-water fish are far more variable, especially Galaxias attenuatus and Eleotris gobioides, than the marine, and Galaxias attenuatus being found both in South America and Tasmania must have had a longer specific existence than any of the others. It is therefore evident that a great amount of variability is not inconsistent with great specific longevity under certain conditions. The conditions in this case are, I believe, the absence of any large rapacious fish preying on the smaller variable ones, and thus tending to fix those varieties which are best adapted to elude the observation of the enemy. These conditions will soon no longer exist in our rivers on account of the introduction of the trout, and I should like to draw attention to the fact that descriptions and figures of all the varieties of fish occurring now in one or more of our rivers would be a most valuable contribution to science as material for future naturalists.
Of our fresh-water fish found beyond New Zealand, Retropinna richardsoni is found in the Chatham Islands; Galaxias fasciatus in both the Chatham and Auckland Islands; Galaxias attenatus in the Chatham Islands, Tasmania, Patagonia, and South America; Galaxias olidus in Australia; Anguilla aucklandii in the Auckland Islands; Anguilla australis in the Auckland Islands, Tasmania and Timor; Anguilla latirostris in the Chatham Islands, Europe, Egypt, China, and the West Indies; Geotria australis in Australia; and Geotria chilensis in Western Australia and-Chile. Thus four of our freshwater fish are found in the Chatham Islands, and three in the Auckland Islands, which are all the fresh-water fish known to inhabit those places; three are found in Australia, two in Tasmania, two in South America, one in the Island of Timor, and one is spread from China to Europe and the West Indies. The Australian grayling also (Prototroctes maræna), although a distinct species, much resembles our own (P. oxyrhynchus); and another closely related genus (Haplochiton) is found in South America.
The genus Eleotris is widely spread in tropical countries. Its head quarters
Hutton.—On the Geographical Relations of the N.Z. Fauna. are in the Indian Archipelago, and it ranges west to Madagascar, east to Mexico and the West Indies, north to Japan, and south to New Zealand, but is not found in Africa. The nearest ally of our species (E. gobioides) is E. obscura from Japan and China.
The evidence, therefore, to be derived from the fresh-water fish goes to prove that a close connection has existed between Australia, New Zealand, and South America. The fact of two species of the same genus of grayling being found in Australia and New Zealand respectively, while South America is inhabited by a closely allied but distinct genus, indicates either that our connection with Australia was later than with South America, or that in the old continent New Zealand and Australia were inhabited by one, and South America by another species of the same family. The fresh-water fish also prove that our connection with the Chatham and Auckland Islands was much later than with Australia. The distribution of Anguilla latirostris, which is not found nearer than China,* adds its testimony to that of Lotella and Ditrema of a former connection with that part of the world not by way of Australia, and we shall find that this remarkable connection with China and the Indian Archipelago, thus dimly shadowed out by the fishes, gets stronger and stronger as we review the invertebrate animals.
Of the New Zealand Mollusca about 460 species are now known, of which about one-half are found nowhere else. They show, as might be expected, a marked affinity with Australia, but are still very distinct. We miss Olivella, Vanikoro, Eutropia, Perna, Trigonia, and others; while Mitra, Columbella, Marginella, Natica, Scala, Conus, Cypræa, and Cardium, are very feebly represented with us. On the other hand Australia does not possess Buccinum, and Fuscus, Imperator, Purpura, Turritella and Pecten are much less developed than in New Zealand. As, however, the affinity is decided I shall here limit myself to pointing out our connection with other countries.
Of Cephalopoda we possess eleven species, only two of which are peculiar to New Zealand. Onychoteuthis bartlingii, Ommastrephes sloani, Nautilus pompilius, and Argonauta nodosa, are all found in the Indian Ocean, and the two last in the Pacific also, but none of them in Australia.
Of marine Gasteropods and Conchifera, omitting the marine air breathers, we have 330 species, about 160 of which are endemic. Of these Cyclina kroyeri, Mytilus magellanicus, and Anomia alectus are only found in South America, as also is the genus Solenella. Chione mesodesma is found at Valparaiso and the Philippine Islands, Barbatia pusilla in Peru and Australia,
[Footnote] * Dr. Günther has lately described A. obscura, a closely allied species, from the Fiji Islands.
Myodora ovata in the Philippines and Australia, Mytilus smaragdinus and Anomia cytæum in China; while we also have a small Cypræa which appears to me to differ from C. punetata, from the Philippines, only in the absence of red spots. Bankivia varians is found in South Africa and Tasmania. Our common pipi (Chione stutchburyi) is found in Kerguelen Land, while Ranella vexillum, which is also found in Tasmania, is closely allied to R. argus from the Cape of Good Hope, and to R. proditor from St. Paul Island. The genera Phorus, Rotella, and Calyptræa are found in the Philippine Islands and China, but not in Australia. The genus Lyonsia, of which we possess one species, extends from Europe and India to the Philippine Islands and Borneo, and is also found in Peru and the West Indies. A few of our shells are almost cosmopolitan, as Lucina divaricata, Saxicava arctica, Crypta unguiformis, and Lima squamosa; while Nucula margaritacea inhabits Europe. Dosinia subrosea is said to have been found in the Persian Gulf, and the genus Solemya is found only in Australia and the Mediterranean. While, therefore, our marine shells show a decided affinity to Australia, they also show a slight connection with South Africa, Kerguelen Land, St. Paul's, and South America, and point more decidedly to a connection with the Philippine Islands and China.
Of land and fresh-water shells, including the marine air-breathers, we possess 114 species, of which 97 are not found elsewhere. These show many striking and important facts in distribution. Three only, Helix subrugata, H. sydneyensis, and H. rapida are found in Australia, and of these the second is so like H. cellaria, of Europe, that it has only lately been distinguished from it by Dr. Cox, and is also closely allied to H. glaberrima from the Solomon Islands. Helix rapida is also found at Erromanga, one of the New Hebrides. Helix coniformis* inhabits the Louisade Islands, H. radiaria the Solomon Islands, and H. vitrea the Admiralty Islands. Cassidula mustelina is found at Singapore and Pulo-penang, and Amphibola avellana in New Caledonia. But the distribution of some of the genera is more important even than that of the species. Nanina spreads from India to China, the Philippines, Indian Archipelago, and Polynesia, and is also found in Madagascar and the Mauritius, but not in Australia. Amphibola extends over Australia and Polynesia to Burmah. Lymnæa extends from Europe to India, China and Java, and is also found in North America but not in Australia. Assiminea is found in England, India, Celebes, Molucca Islands, and the Navigator and Friendly Islands, but not in Australia. The family Ancylinæ, or fresh-water limpets, of which we possess two species, is found
[Footnote] * I am indebted to His Honour T. B. Gillies for the information that H. coniformis, H. radiaria, H. subrugata, and H. vitrea inhabit New Zealand. Mr. Gillies collected the specimens in the northern portion of the province of Auckland, and they were determined by Prof. Macalister, of Trinity College, Dublin.
Hutton.—On the Geographical Relations of the N.Z. Fauna. only in North and South America, Europe, and Madeira; and our common slug (Milax antipodarum) belongs to a genus found only in Europe and the Island of Teneriffe. Testacella, of which we also possess a species, is only found in Europe and Teneriffe.
Our former connection with Australia, however, is shown in the family of bitentaculate slugs (Janellidæ), a family which is found only in Australia and New Zealand, and also in the marine air-breathing limpets (Siphonaria), three of our species being found in Australia and Tasmania.
The land and fresh-water univalves therefore show a stronger affinity to Polynesia and the Philippine Islands, by way of New Caledonia, the New Hebrides, Solomon Islands and the Indian Archipelago, than they do to Australia, although the distribution of the genus Janella shows that land communication once existed with Australia also. To South Africa and South America they exhibit no special affinity. Like the birds and fishes they also show a slight anomalous affinity to Europe without any intermediate steps.
From the Chatham Islands eighty-two species of Molluscs are known, of which nine appear to be peculiar to those islands; the rest are all found in New Zealand, including Janella bitentaculata and Siphonaria scutulata.
I know of two shells only from the Auckland Islands (Patella illuminata and Vitrina zebra), both of which are endemic.
Of Brachiopods we possess eight or nine species, of which two only (Kraussia lamarkiana and Magas cumingi) are found in Australia, the latter being also reported to occur in China. The genus Rhynchonella is only known living in the arctic portions of North America and Japan, but this anomaly is not surprising when we remember that this genus existed during the Lower Silurian Period, but it is interesting as affording us the clue by which other similar anomalies may be explained.
The New Zealand Tunicata are as yet but little known. The genera Ascidia, Boltenia, and Botryllus, are only found in Europe and North America. Doliolum denticlatum is found at the Molucca Islands.
Of the Polyzoa I am acquainted with eighty-nine species, of which thirty-one have been found nowhere else as yet, but it is probable that their range is very imperfectly known. Twenty-three of our species are found in European seas, while the intervening tropical seas appear to be almost destitute of this form of life. The chief point of interest in our Polyzoa is the great development of the massive species of Cellepora, and of the coral-like family Idmoneidæ, which recall to mind the crag formation of England; indeed one of our species, Hornera striata, is found fossil in the crag; it is, however, also found fossil at Oraki, near Auckland, in beds of still older date. Considering how little
attention has been paid to our Polyzoa, the number of known species indicates a rich fauna, and, indeed, the entire class seems to be more abundant in the southern than in the opposite hemisphere, and, like the petrels, contains many forms quite unrepresented in the north.
No New Zealand naturalist who has collected insects on however small a scale in Europe, can, I think, fail to be struck with the paucity in New Zealand, not only of species, but in some orders of individuals also. It is remarkable that in this country, whose indigenous warm-blooded animals are limited to birds and bats, on entering the bush instead of finding the masses of decaying wood and leaves swarming with life, we find hardly a living creature,* while at the same time we are attacked by myriads of blood-thirsty mosquitos (Culex acer). It would certainly seem that abundance of food does not produce abundance of individuals in some orders (e.g. Coleoptera), neither does an absolute dearth of food in the imago state prevent the increase of individuals in others (e.g. Diptera). The swarms of sand-flies (Simulium cæcutiens), also, that greet us on the coast, from the North Cape to the Bluff, where could they possibly have found food before the advent of man? Where indeed do they find it now in sufficient quantities?
Of beetles about 200 species inhabiting the land are described, the whole of which, I believe, are found nowhere else. These species are distributed into about 110 genera, of which about thirty-five are peculiar to New Zealand. A remarkable contrast to this is shown in the water-beetles, of which four only are known, two (Cybister hookeri and Colymbetes refimanus) being, I believe, endemic, and the other two (Colymbetes notatus and Gyrinus natator) being found in Britain. The genera best represented are Elater with twelve, Feronia with eight, Mecodema with nine, Xylotoles with seven, Cincidela with six, Anchomenus and Maoria with five each, and Coptoma with four species. Few beetles can be called abundant, the little green species (Pyronota festiva) so destructive to our fruit trees, and a small brown species (Colaspis brunnea), common on the manuka (Leptospermum) in December and January, are, perhaps, the only two that deserve the name, although many can be called common. The beetles as a whole are, according to Mr. Pascoe, most closely allied to those of Australia.
The Hymenoptera are very poorly represented, about eighteen species only being as yet known. All are, I believe, endemic. Most of the genera are widely spread, but Orectognathus, and Dasycolletes, are peculiar to New Zealand. The poorness of our fauna in this order cannot be owing to
[Footnote] * My experience in this respect in New Zealand is very different to that of Mr. Wallace in Singapore and Borneo, but similar to his in Celebes and Cerain.
Hutton.—On the Geographical Relations of the N.Z. Fauna. unsuitableness of climate, for the honey-bee (Apis mellifica) which was introduced about thirty years ago, has spread over both Islands*
The Diptera are more numerous than the Hymenoptera, sixty species being known. This is just opposite to what obtains in most countries, including Australia and South America. Of these Tipula senex is found in Australia; Musca taitensis in Polynesia; and Musca læmica in both Australia and Polynesia. Although most nearly allied to Australia, our dipterous fauna must have been derived from other localities as well, for the genus Diphysa occurs only in Mexico and Brazil; Actina in Europe; Cænosia and Sapromyza in Europe and North America; and Opomyza in Europe and the Mauritius. No genus is endemic. Of the earwigs we possess one endemic species (For- ficula littorea), found only near the sea shore.
Of the Lepidoptera I know hardly anything, and prefer waiting until Mr. Fereday has published his promised descriptions of the species, before examining their bearing on the present subject. But one fact stands out prominently, viz, that out of more than three hundred species, only eight belong to the butterfly section (Fereday, Trans. N.Z. Inst. IV., p. 217), and of these several are world-wide stragglers.
Of Neuroptera about fifteen species are known. Of these, Perla opposita is found in Tasmania; and our representative of the white ants (Calotermes insularis) in Australia. This order appears to have more affinity with Tasmania than with Australia, and it is remarkable that the wide spread genus Perla, which is found throughout North and South America, and from Europe through India to China and Japan, is also found in New Zealand and Tasmania, but not in Australia. Leptocerus has also the same range, with the exception of not being known in China and Japan. Hermes extends from India to China and Java; it is also found in tropical Africa and South America, but not in Australia nor Tasmania. Palingenia is found in Europe, India, North Africa, and North and South America; while Philanisus is peculiar to New Zealand. The Heteroptera are remarkable for their fragmentary character, and wide distribution. The thirteen known species belong to thirteen different genera, and nine families. Arma schellembergii is found in Australia and the Philippine Islands; Cermatulus nasalis in. Australia and Tasmania; Platycoris immarginatus and Rhaphigaster amoyti in Australia; Lygans pacificus in Australia, Tasmania, and India; and Nysius zealandicus in Tasmania; thus leaving not more than seven endemic species, three of which have not yet been properly examined, and may therefore be found to be identical with species inhabiting other countries. One of the endemic species (Rhopalimorpha obscura), however, belongs to a genus found nowhere else.
[Footnote] * Mr. W. T. L. Travers informs me that the honey-bee was introduced into Nelson in 1842, and that wild bees were common in 1850.
In strong contrast to this stand the Homoptera, which include nineteen species, all endemic, and belonging to three genera only; Cicada having twelve, and Cixius seven species.
The number of species of Orthoptera I do not know, but in comparison with other orders it is well represented by both winged and wingless members, and the genera, as a rule, contain several species.
Whilst, therefore, the insect fauna as a whole shows its greatest affinity towards Australia it also exhibits a connection with other countries, more especially China and Europe. But the most remarkable fact is the great difference shown in this respect by the different orders. Whilst the Diptera, Neuroptera, Homoptera and Orthoptera present the appearance, in part at least, of an old fauna, the Heteroptera are nearly all stragglers, and this strongly suggests the inference that at the time of the spreading of the former orders the Heteroptera were not in existence. The same thing is seen in the difference between the moths and the butterflies, suggesting also that the latter were developed at a later period than the former, and there can be no doubt but that when our insects are better known a careful comparison of them with similar faunas of other countries will afford a most instructive lesson.
With the exception of the Indian (Blatta orientalis) and American (B. americana) cockroaches, neither of which are common, the flea (Pulex irritans), the bed-bug (Cimex lectularius), several Aphides, the slug-worm (Tenthredo cerasi), and the house-fly (Musca domestica), I am not aware of any insect that has been introduced unintentionally by man during the progress of colonization, for the ring-legged mosquito, which is supposed in Auckland to have been introduced by the troops from India, belongs to a species (Culex argyropus) not found elsewhere, and was sent home by Dr. Sinclair before the troops arrived. The only exceptions may perhaps be the black field-cricket, which, although inhabiting fields with us, and but rarely entering houses, appears to be identical with the house-cricket of Europe (Acheta domestica) and to have spread quite lately; and also a small dark-brown beetle belonging to the genus Elater, which is abundant in Auckland, but, to the best of my knowledge, is not found more than twenty miles out of that town.
Of Centipedes nine or ten species are now known, all of which are endemic. The genus Lithobius extends from North America, Europe, and North Africa to Singapore, but is not found in Australia. Henicops is found only in Chile and Tasmania, Cryptops only in North America and England, while Cermatia and Cormocephalus have wider ranges, and are both found in Australia.
Of Spiders we have about 100 species, but my knowledge of them is very limited. Mr. Pickard-Cambridge, in a letter to me remarks, “all the spiders you now send (from the Auckland province), except one or two, are strikingly European in appearance, nothing tropical-looking among them.” Perhaps the most remarkable fact is the occurrence in the Chatham Islands of a species of water-spider (Argyroneta) of which only one other species, inhabiting Europe, is known. Spiders are very numerous in New Zealand, owing no doubt to the abundance of Diptera, on which order they chiefly prey.
Of Crustaceans 106 species have been described as coming from New Zealand, but my knowledge of this class also is at present very limited. Professor Dana has remarked that New Zealand has a greater resemblance to Great Britain in its Crustacea than to any other part of the world; but our common salt-water crayfish (Palinurus lalandii) is found at the Cape of Good Hope and the Island of St. Paul.
Our marine Annelids have up to the present been almost entirely neglected. Of terrestrial forms we have two species of earthworm (Lumbricus) and a member of the peculiar genus Peripatus, found only in South America, the Cape of Good Hope, and the West Indies.
The most remarkable fact in this class is the occurrence of two or three species of land Planarians, the so-called “land-leeches,” one or two of which belong to the genus Bipalium, found only in India, China, and Japan.
Of Echinoderms we have seventeen species of star-fish, eight sea-urchins, and eight holothurians. Of these twelve star-fish, six sea-urchins and all the holothurians appear to be endemic. Of the others Ophionereis fasciatus is found at the Chatham Islands, Pentagonaster pulchellus at the Chatham Islands and in China, Othilia luzonica in the Philippine Islands and Vera Cruz, while we also possess species apparently identical with Astropecten armatus of South America, and Henricia oculata of Europe. It is worthy of special remark that although Australia possesses several species of Pentagonaster, the Chinese species is not found there, so that it must have migrated to us direct, and not have come via Australia. We also possess a species of Pteraster, a genus found only in South Africa and Northern Seas. Of the sea-urchins, Cidaris tubaria, and Echinobrissus recens are both found in Australia, but the latter appears to be very rare in New Zealand, as I have only seen one specimen, which is in the Colonial Museum.
Cælenterata and Protozoa
Of these very little is known. Our seven species of corals are all peculiar, as also appear to be many species of Sertularians and sponges, but I know of no facts among these lower animals that will help out the present investigation except in the case of Cryptolaria, a genus belonging to the family Sertularidæ, and consisting of two species, one of which is found in New Zealand and the other in Madeira.
If now we review the evidence adduced, and select the more important points we find in the distribution of the Struthious birds, the frogs, freshwater fishes, several shells (such as Cyclina kroyeri, Mytilus magellanicus, Anomia alecto, Barbaia pusilla, Chione stutchburyi, and Ranella vexillum), in the genus Hemicops among the Centipedes, and Perripatus among the Annelids, evidence of a former great extension of land in the Southern Hemisphere, for these cases cannot all be accounted for by drifting icebergs. With the exception of the shells and two fresh-water fishes no species however is common to New Zealand and South America on the one hand, nor to New Zealand and South Africa on the other, for I omit from consideration the species of marine fish, as they might perhaps have crossed at a later date. In the frogs the genera, and in the birds the families, are different. This perhaps indicates a very long interval since the separation of these countries took place, but differentiation of form, even in closely allied species, is evidently a very fallacious guide in judging of lapse of time, and a surer one is afforded us in the absence of Mammalia from New Zealand, for it is evident that if the Marsupials that now inhabit Australia, or the placental Mammals that inhabit South America, had been in existence at the time of the distribution of the Struthious birds some members would have found their way to New Zealand, and would have remained upon it with the Moas. This antarctic continental period must therefore have preceded the spread of the Mammalia into the Southern Hemisphere. Besides this continental period we have evidence in Eudynamis taitiensis, Naultinus pacificus, Amphibola avellana, Musca taitensis, and in the genera Ocydromus and Nestor, of a Polynesian continent quite unconnected with Australia, but including Lord Howe Island, Norfolk Island, and New Caledonia, while by Helix coniformis, H. rapida, H. radiaria and H. vitrea, we can prove a close connection with the New Hebrides, Solomon Islands, Louisade Archipelago, and the Admiralty Islands. By Nanina among land shells, and Assiminea among fresh-water shells, we prove a connection also with the Navigator and Friendly Islands, and these genera take us north through the Molucca Islands, Celebes, Borneo and the Philippines, to China, where we again come across many New Zealand species and genera. The
most important are Ditrema, Torpedo, and Anguilla latirostris among fishes; Mytilus smaragdinus, Phorus, Rotella, Calyptræa, Cassidula mustilina, Lymnæa, and Rhynchonella among shells; Perla and Hermes among insects; Lithobius among centipedes; Bipalium among the Scolecida, and Pentagonaster pulchellus and Othilia luzonica among the star-fish; none of these, it must be remembered, being found in Australia. The absence of Mammalia, however, in New Zealand shows that this line of communication was never continuous land, but the absence from Australia of the forms that I have mentioned shows that the connection along the whole line was closer at every point than it was with that continent, and this leads to the further conclusion that this line of communication existed at a later date than the connection of New Zealand with Australia.
The close relationship of the Chatham and Auckland Islands in all their natural productions to those of New Zealand, and the far greater difference between New Zealand and the islands more to the north, as well as the large number of species of moa lately inhabiting these islands, shows that another and smaller continent, or perhaps a large island, existed at a still later period, but has since subsided, and this must bring us nearly to the recent period, or the difference between New Zealand and the Chatham Islands would be greater.
The geographical distribution, therefore, of the New Zealand fauna points to the following conclusions:—
1. A continental period, during which South America, New Zealand, Australia, and South Africa were all connected, although it is not necessary that all should have been connected at the same time, but New Zealand must have been isolated from all before the spread of the Mammals, and from that time to the present it has never been completely submerged. This continent was inhabited by Struthious birds, and by Hymenolaimus, Notornis, Hinulia, Mocoa, Galaxias, Prototroctes, Liopelma, Janella, Amphibola, Hemicops, and Peripatus, and further to the north by Megapodius; and probably also by many forms peculiar to New Zealand, such as Stringops, Keropia, Xenicus, Heteralocha, Anarhynchus, Naultinus, etc. Of course in mentioning these names I do not mean that all the forms were the same then as now, but that the ancestors of these genera lived on the old antarctic continent.
2. Subsidence followed, and the evidence then points to a second continent stretching from New Zealand to Lord Howe Island and New Caledonia, and extending for an unknown distance into Polynesia, but certainly not so far as the Sandwich Islands. The fact of Mammals being found in the New Hebrides, Solomon Islands, and New Ireland, shows that between New Caledonia and the New Hebrides a narrow strait must have existed, cutting off land communication, but these were connected with China either direct or
by a chain of islands. This second continent received from the north those forms already enumerated together probably with Sphenæacus, the rails, and the starlings; at the same time it received from Australia the honey-eaters, Certhiparus, Gerygone, Petroica, Rhipidura and others, and from that time to the present has been occasionally receiving additional birds. It will also be noticed that very few of the birds of the middle palæotropical region came down this line of communication, no pheasants, woodpeckers, grackles nor finches, while Australia in its wood-swallows (Artamus), pittas, quails, and numerous finches, shows now some affinity to this region. This can be best explained by supposing that the New Zealand line of communication was broken up before these birds came into existence, and that further changes have since taken place in the lines of easiest communication; indeed, the fact of such forms as the elephant, tiger, and bear being found in Sumatra and Borneo; Marsupials in Celebes, the Moluccas, Solomon Islands, and New Hebrides; and the presence of an emu in New Guinea, and a cassowary in Australia, prove that changes in the distribution of land have since taken place, but it is foreign to the object of this paper to speculate on these here. This second continent was also inhabited by most of the orders of insects, although perhaps not in great abundance, but Heteroptera and the butterfly section of the Lepidoptera were absent.
3. Subsidence again followed, and New Zealand was reduced for a long time to a number of islands, upon many of which the moa lived. This was followed by—
4. Elevation; these islands were connected and a large island existed disconnected from Polynesia. This was once more followed by—
5. Subsidence, and the geography of this part of the World assumed somewhat of its present form.
Such are, I think, the deductions that may be fairly drawn from a study of our fauna. It remains now to examine the geological and palæontological evidence and see whether it agrees with that derived from zoology, and then try to fix with as much accuracy as possible the dates of the principal movements of the earth's surface which have gradually led to the present state of the New Zealand fauna.
Hardly anything is yet known of the palæozoic rocks of New Zealand. The earliest fossil shells described are almost identical with those living in Europe during the triassic period, but the only known plant is Dammara australis (Hochstetter's “New Zealand,” p. 57), a genus still living in New Zealand, but also found in Australia, New Caledonia, New Hebrides, Fiji, and the Indian Archipelago.
An interval then occurs, and the next formation probably belongs to the jurassic period. In this we find Belemnites aucklandicus, which can hardly be distinguished from B. canaliculatus, and Astarte wollumbillaensis. The ferns, too, found so plentifully near Port Waikato, in the Clent hills, at the Mataura, and at Waikawa harbour, are considered by Professor McCoy to be identical with Australian ferns from the same formation. At the close of this period movements on an extensive scale commenced in New Zealand, the land was upheaved, and an extensive mountain chain formed. Á long blank now occurs in our geological record (see Geo. Reports, 1872, p. 105), the next formation belonging to quite the uppermost part of the secondary epoch, later I believe than the white chalk of England. In it we find remains of dicotyledonous plants and large Saurians belonging to the genera Crocodilus? and Plesiosaurus. Here also we find three fossil shells (Dentalium majus, Lucina americana and Cucullœa alta), similar to those found in South America, one of which, Lucina americana, is found in the lower cretaceous rocks of Tierra del Fuego, and the other two in the miocene formations of Patagonia and Chile; thus showing that during this blank in our geological record an intimate connection had existed between New Zealand and South America. The disposition, however, of these beds shows that the New Zealand Alps were not submerged. A long interval now follows, during which New Zealand was again upheaved, and the next rocks that we find are of upper eocene date (Geo. Rep., 1872, p. 182). From that time until the close of the miocene period New Zealand was greatly depressed, and divided into several islands, but at the close of the miocene period it was once more upheaved. During this period we find several South American miocene shells not met with in the older formation, as well as several Australian ones. During the newer pliocene period it again subsided, and the Wanganui beds were deposited. From that time I can see no evidence of the land having ever stood at a higher level than it does at present, but as the later changes in the physical geography of New Zealand have a most important bearing on the present condition of its fauna, beyond the scope of my present inquiry, I propose treating the subject in a separate paper. * The geological evidence is, therefore, that since the jurassic period there have been three principal upheavals in New Zealand, in the lower cretaceous, lower eocene, and older pliocene periods respectively, and that these, were divided by two insular periods, viz., during the upper secondary (Danien), and from the commencement of the upper eocene to the close of the miocene, thus agreeing completely with the zoological evidence.
The dates assigned by the geological evidence also agree well with those derived from zoology. We have seen that it is necessary to suppose that the first great antarctic continental period was anterior to the date of the spread
[Footnote] * Vide post, “On the Date of the last Great Glacier Period in New Zealand.”
of the Mammals southwards. Now a few Marsupials are known in the triassic period, but it is quite possible either that they spread very slowly, or that barriers existed that prevented any southward migration. In the eocene period, however, some placental Mammals were in existence, although Marsupials, not of Australian types however, still formed in Europe the principal mammalian life; and if the supposed barriers to a southward migration were still in existence, we know, from what happened in the Northern Hemisphere, that the whole, or nearly the whole, of the Marsupials would have been exterminated. The Marsupials, therefore, must have migrated south not later than the eocene period, and as we know that our connection with Australia and South America must have been before that migration, it follows that the first, or lower cretaceous period of upheaval, must have been the time of the antarctic continent. This is rendered still more probable by the fact that our jurassic fossils show a connection with Australia only, while our upper secondary fossils show for the first time a relation to South America. The fact, too, of the cretaceo-oolitic rocks of Tierra del Fuego having been largely disturbed, metamorphosed, and broken through by dykes of green-stone, shows that extensive elevatory movements have taken place there, also, since they were deposited. It is therefore to the lower cretaceous period that we must probably look for the time of the dispersion of the Struthious birds. With regard to the date of the second, or Polynesian continental period, the only zoological evidence we have is that it probably preceded the wide dispersion of the Hemiptera, and the butterfly section of the Lepidoptera. This, therefore, could not have been later than the eocene, for a fossil butterfly (Vanessa pluto) has been found in the lower miocene deposits of Radaboj in Croatia, and fossil Heteroptera in the miocene beds of Œningen in Switzerland. The elevation during the lower eocene period was therefore probably the one which formed the continent that I have described as including New Caledonia and some of the Pacific Islands. At this period probably Northern Australia was submerged, and the southern portions of Australia and Tasmania formed one large island, while New Guinea, including the Solomon Islands and New Hebrides on the south, and the Molucca Islands on the north, formed another large island, divided from the New Zealand island, or continent, by the straits between New Caledonia and the New Hebrides.
This was the time of the migration from China southwards, and it is worthy of notice that at the same time a large ocean existed from southern Europe to China, in which the nummulitie limestone was being deposited. Would it be too bold to speculate that it was along the shores of this ocean that those fish, crustaceans, and shells migrated, which are now found in the North Atlantic or Mediterranean on the one hand, and in China or Japan on the other, but not on the southern shores of Asia; and that the anomalous
distribution of European forms of fish, shells, etc., in New Zealand may be traced to the same route? This same period of sea communication between Europe and Japan will also probably have been the time of the land connection that once existed between India, Madagascar, and Africa (the Lemuria of Dr. Sclater), as proved by the recent fresh-water fish, and birds, as well as by the miocene Mammalia, * and to this period we may also refer the origin of the curious affinity between some of the birds of Celebes and Africa. The long insular period during the upper eocene and miocene times will, therefore, be the period of specific change in the moas, while the older pliocene upheaval will be the time of the mingling of the various species in New Zealand, and the peopling of the Chatham and Auckland Islands. The newer pliocene was the time when the two islands of New Zealand were divided, and also the period when the Chatham and Auckland Islands were separated from them, but the latter occurrence probably preceded the former by a long interval.
Such appears to me to be the hypothesis most capable of accounting for the present fauna of New Zealand.
The objection, however, may be fairly raised that, if it is true, evidence of its truth ought to be also found in the flora of the country, which is not the case. I fully acknowledge the force of this argument, but think that some slight evidence can be found in the phænogamic flora. The distribution of Eucalyptus for instance, is somewhat parallel to that of the Marsupials, and can be only explained in the same way. Stilbocarpa polaris has its nearest allies in China and the Himalaya Mountains; while the distribution of Metrosideros, Ligusticum, Angelica, and perhaps Veronica, implies a connection between New Zealand and Asia not by way of Australia. This connection is obscured by the great preponderance of Australian and South American forms, but still furnishes an indistinct copy of the bolder outline sketched out by the fauna. This is owing to the wider distribution of genera among plants than among animals, and to me it appears to prove that the flora of a country, as a whole, is of a more ancient date than its fauna. Among the cryptogamic plants no trace of this outline can be discerned, as also is the case with the lower classes of the animal kingdom, owing to the genera having been, so to say, universally spread before the last migration from Asia took place.
That the facies of a fauna and flora should date back from so long a period as I suppose, is certainly at variance with ordinarily received opinion, but from a study of the fauna and geology of New Zealand I do not see how we can escape from the conclusions that I have arrived at. I am well aware,
[Footnote] * Professor Huxley thinks (“Quar. Jour. Geo. Soc.” 1870. Ann. Address, p. 56.) that the land communication between India and South Africa was caused by the upheaval of the nummulitic sea, but it seems to me more probable that the land communication was by the shores of that Sea.
however, that much more has to be done in the geology and natural history, not only of our own islands, but also of the surrounding countries, before they can be considered as satisfactorily proved; but I think that it will be easier afterwards to prove this hypothesis, or to disprove it and point out a more correct one, than it would be to detect it if the discussion had been postponed to a future period, when the more salient points will probably be obscured by the mass of facts which will then have accumulated. Such at least is my hope, but whether I am mistaken or not I leave others to judge.