A History of the Birds of New Zealand.
Apteryx Australis. — (South-Island Kiwi.)
Apteryx australis, Shaw and Nodder, Nat. Misc. xxiv. pls. 1057, 1058 (1813).
Dromiceius novœ zealandiœ, Less. Man. d’Orn. ii. p. 210 (1828).
Apteryx mantelli, Bartlett, P. Z. S. 1850, p. 275.
Apteryx fusca, Potts, Trans. N.-Z. Inst. vol. v. p. 196 (1873).
Native names.—Kiwi and Tokoeka.
Ad. similis A. bulleri, sed major, pallidior, et magis grisescens; tergo tantùm vix castaneo tincto: scapis plumarum haud conspicuis, itaque ptilosi molliore distinguendus.
Adult. Differs from Apteryx bulleri in its larger size and in the lighter colour of its plumage, the feathers being of a sandy or greyish brown, with darker margins, those of the upper parts only slightly tinged near the tips with rufous. The plumage of the nape and back of the neck is less hairy; and the feathers of the back and hind parts are destitute of the lengthened and stiffened points which characterize the other species.
Male. Total length, following the curvature of the back, 22 inches; bill, along the ridge 3·75, along the edge of lower mandible 4·1; rudimentary wing, to end of hook, 1; tarsus 2·25; middle toe and claw 3·5; hallux ·75.
Female. Total length (measured as above) 27 inches; bill, along the ridge 5·5, along the edge of lower mandible 5·8; rudimentary wing, to end of hook, 1·5; tarsus 3; middle toe and claw 3·75; hallux ·8.
Obs. As a rule the South-Island birds are larger than those from the North Island; but occasionally examples of Apteryx bulleri are met with fully equal in size to the largest specimens of Apteryx australis; and this is therefore of little or no value as a specific character. It may be observed that in this species the long facial hairs or feelers are, generally speaking, far less abundant than in the North-Island Apteryx.
Young. Has the head and hind neck dark grey, and the rest of the plumage greyish brown, lighter on the under-parts, each feather with a narrow streak of fulvous along the shaft; on the feathers of the upper parts this streak is darker towards the tip, and the terminal filaments are black, whereas on the underparts of the body both the tips and filaments are light brown or fulvous; the bill, which measures two inches in length, is light horn-colour; the legs and feet are light brown, the metatarsi being covered anteriorly with thin scales, scarcely definable to the eye. In this young condition the quill-tubes are very minute, and the plumage of the body is extremely soft to the touch.
In the Rowley collection at Chichester House, Brighton, there is a specimen of the chick, apparently younger than that described above, and differing from it in the lighter tone of its plumage, especially on the upper parts.
The nestling has a wing-claw of the same character as in the adult, although scarcely more than a decimal of an inch in length.
Partial albino. In the Canterbury Museum there is a partial albino, in which the crown and sides of the head, the throat and the whole of the fore neck, and the front of the thighs are yellowish white.page 323
The first example of the Apteryx of which there is any record was obtained in New Zealand about the year 1813, by Captain Barclay, of the ship ‘Providence,’ and afterwards deposited in the collection of the late Lord Derby. This bird was first described, under the above name, by Dr. Shaw (Nat. Misc. l. c.), and afterwards, at greater length, by Mr. Yarrell, in the ‘Transactions of the Zoological Society’ (vol. i. p. 71, pl. 10). On the 10th December, 1850, a series of specimens was exhibited before the Zoological Society of London, when Mr. Bartlett pointed out characters which, as he contended, established the existence of two species, hitherto confounded under the specific name of Apteryx australis. Mr. Bartlett stated, at this meeting, that an Apteryx belonging to the late Dr. Mantell having been placed in his hands by that gentleman, he had remarked its dissimilarity to ordinary examples, and that, after a careful comparison with a number of other specimens, he had come to the conclusion that it was a new species. On comparing Dr. Mantell’s bird, however, with the original specimen in the Earl of Derby’s collection, he had found that they were identical. He accordingly referred his supposed new species to Apteryx australis, and distinguished the more common bird as Apteryx mantelli, for which he proposed the following characters:—“its smaller size, its darker and more rufous colour, its longer tarsus, which is scutellated in front, its shorter toes and claws, which are horn-coloured, its smaller wings, which have much stronger and thicker quills, and also its having long straggling hairs on the face” (Proc. Zool. Soc. 1850, p. 276).
In a paper read before the Wellington Philosophical Society on the 12th November, 1870*, I pointed out that the characters by which Mr. Bartlett had distinguished the species would not stand. I showed that the sexes differed from each other both in size and in the tone of their plumage, that the arrangement of the tarsal scutella differed according to age and other circumstances, that the peculiarity in the cubital quills was not a specific character, the “soft slender quills” indicating only immaturity, and that the length of the “straggling hairs on the face” varied in almost every individual. I stated further that an inspection of the drawings illustrative of the supposed specific distinctions (published by the Zoological Society) had only tended to confirm me in the opinion expressed above.
After that paper was written I had an opportunity of examining several fine series of South-Island Apteryges, and of comparing them with examples from the North Island; and I was then convinced that there are in reality two species of brown Apteryx, readily distinguishable from each other by a very remarkable difference in the structure of their plumage. In the South-Island Kiwi the feathers of the upper parts are soft and yielding when stroked against the grain, whereas in the North-Island bird, owing to a peculiarity in the structure of the shaft, they have stiffened points, and are harsh and prickly to the touch. This character (apart from a slight difference in the colour of the plumage) is constant in all the specimens I have examined; and I have no hesitation in giving it a specific value. In this course I am supported by the unanimous opinion of several of the best ornithologists in England, to whom I have submitted specimens for examination.
I take this opportunity of saying that the credit of this discovery belonged to the late Sir Julius von Haast, who, on receiving from me a North-Island bird for comparison with the specimens in the Canterbury Museum, detected this structural difference in the plumage, and informed me of it long before I had an opportunity of verifying the fact for myself.
Dr. Otto Finsch, however, has arrived at an opposite conclusion, although he seems to have practically conceded the point by admitting the North-Island Kiwi to the rank of a “variety” (i. e. Apteryx australis, var. mantelli)†.page 324
According to the now generally accepted view of what constitutes a “species,” the amount of difference is quite immaterial, provided it be constant and readily distinguishable. If (as is certainly the case) all the known examples from the North Island are referable to “var. mantelli (Finsch),” then, for all practical purposes, the bird must be regarded as distinct, and is, I submit, as much entitled to recognition as any other species on our list.page 325
Professor Sir Richard Owen aptly remarks:—“The Apteryx presents such a singular and seemingly anomalous compound of characters belonging to different orders of Birds as may well make the naturalist pause before he ventures to pronounce against the possibility of a like combination of peculiarities in the historical Dodo. It seems, as it were, to have borrowed its head from the Longirostral Grallœ, its legs from the Gallinœ, and its wings from the struthious order. It is clothed with a plumage having the characteristic looseness of that of the terrestrial birds deprived of the power of flight; its feathers resemble those of the Emu in the general uniformity of their size, structure, and colour, but they are more simple than in any of the tridactyle Struthionidœ, as they want the accessory plumelet… . When the trunk is stripped of its plumage, the body of the Apteryx presents the form of an elongated cone gradually tapering forwards, from the broad base formed by the haunches to the extremity of the attenuated beak. The wings appear as two small crooked appendages projecting about an inch and a half from the sides of the thorax, and terminated by a curved, obtuse, horny claw 3 lines long: the antibrachium is retained in a state of permanent flexion by the surrounding integument of the wing; and it cannot be brought by forcible extension beyond an angle of 45° with the humerus. Nine quasi-quill-plumes, not exceeding in length the ordinary body-feathers, but with somewhat thicker shafts, are arranged in a linear series along the ulnar margin of the antibrachium; the terminal ones are the largest, and in one specimen they presented a structure differing from that of the ordinary plumes, consisting of a shaft from which radiated a series of flattened horny filaments of nearly equal length.” (Prof. Owen’s ‘Memoir on Apteryx australis,’ Trans. Z. S. ii. p. 257.)
Professor Hutton, in his valuable essay on the “Geographical Relations of the New Zealand Fauna” (Trans. N.-Z. Inst. vol. vi. p. 232), says:—“The Apterygidœ have a more generalized 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.” And, again, in his review of my ‘Birds of New Zealand’ (first edition) in the ‘New-Zealand Magazine,’ p. 99, Professor Hutton says:—“We must take exception to the Kiwi being considered as the living representative of the Moa, or, as Dr. Buller puts it in his preface, ‘the only living representative of an extinct race.’ No doubt the Kiwi and the Moa have several features in common; but it is certain that both the Emu and the Cassowary are far more nearly related to the Moa than is the Kiwi.” Professor Mivart has since read a paper before the Zoological Society of London on the axial skeleton of the Struthionidæ, which effectually settles the question at issue. He pointed out that, judging by the characters of the axial skeleton, the Emu presents the least differential type, from which Rhea diverges most on the one hand, and Apteryx on the other; that the resemblance between Dromœus and Casuarinus is exceedingly close, while the axial skeleton of Dinornis is intermediate between that of Casuarinus and Apteryx; its affinities, however, with the existing New-Zealand form very decidedly predominating.
Still later, Professor Newton (in his article “Ornithology” in the ‘Encyclopædia Britannica’) thus referred to the subject:—“Some systematists think there can be little question of the Struthiones being the most specialized and therefore probably the highest type of these Orders, and the present writer is rather inclined to agree with them. Nevertheless the formation of the bill in the Apteryges is quite unique in the whole Class, and indicates therefore an extraordinary amount of specialization. Their functionless wings, however, point to their being a degraded form, though in this matter they are not much worse than the Megistanes, and are far above the Immanes—some of which at least appear to have been absolutely wingless, and were thus the only members of the Class possessing but a single pair of limbs.”
It will be seen, therefore, that I was fully justified in referring to the existing species of Apteryx as “the diminutive representatives of colossal ornithic types that have disappeared.”
An able paper communicated by Professor Huxley to the Zoological Society on June 2, 1882, contains some interesting information on the respiratory organs of Apteryx, from which I extract the page 326 following:—“The question whether Apteryx presents any real approximation to mammals in the structure of its breathing-apparatus is of considerable interest from its bearing upon the general problem of the affinities of birds to other groups of vertebrated animals. Having recently examined a specimen of Apteryx (which, although it had been many years in spirit, was still in a very fair state of preservation) with reference to this point, I have come to the conclusion that its respiratory organs differ in no essential respect from those of other birds, though they exhibit those peculiarities which are peculiar to and characteristic of the class Aves in a less developed condition than that which obtains in all those Carinatæ and Ratitæ which have been carefully studied… . . The respiratory organs of Apteryx are thoroughly ornithic, differing from those of other birds chiefly in the greater width and smaller aggregate surface of the respiratory passages, in the rudimentary condition of the pneumatic sacs, and in the much greater strength of the pulmonary and septal aponeurotic expansions. Neither in Apteryx, nor in any other bird, has either of these the slightest real resemblance to a mammalian diaphragm. For, as has been seen, the heart lies altogether behind both, and the muscular digitations of the pulmonary aponeurosis are supplied by the intercostal nerves, the phrenic being absent. The vertical and oblique septa really answer to the fibrous tissue of the posterior and middle mediastinum in mammals. In this, as in all other cases, the meaning of ornithic peculiarities of structure is to be sought, not in mammals, but in reptiles. It is only among reptiles that we meet with pneumatic bones similar to those of birds (Crocodilia, Pterosauria, Dinosauria), pulmonary airsacs (Chamæleonidæ), and membranous expansions which are comparable to the septa in birds.” (Proc. Zool. Soc. 1882, pp. 560–568.)
Comparatively few specimens of this species are now brought in by collectors in the South Island, whereas the supply of Apteryx oweni is undiminished; and the conclusion is irresistible that Apteryx australis, perhaps the most interesting bird in the Southern Hemisphere, is fast becoming extinct.
Mr. Reischek informs me that on the 25th September he captured a male bird of this species sitting on a single fresh egg on a loose nest composed of grass and dry leaves under the shelter of a stone at an elevation of 2000 feet above the sea. The egg, unfortunately, got broken through the kicking of the bird when resisting capture. The sex was determined by dissection, and the bird was of unusual size, equalling the measurements which I have given for the adult female.
A specimen of the egg in Mr. Philip Crowley’s collection at Croydon is exactly similar to that of Apteryx bulleri, but rather larger than ordinary examples of the latter, measuring 4·75 inches in length by 3·05 in breadth.
Chick of Apteryx bulleri. (See page 315.)
* Trans. N.-Z. Inst. vol. iii. pp. 37–56.
† Since the publication of my first edition, I have examined numerous examples of both forms, and I have seen no reason to change or modify the views expressed above as to the specific value of the North-Island Kiwi, as compared with Apteryx australis of the South Island. It is desirable, however, to have the arguments on both sides stated fully, and I have therefore taken the trouble to translate, from the German, Dr. Finsch’s last published remarks on this subject in the ‘Journal für Orni-thologie,’ from which it will be seen that this naturalist is still opposed to the recognition of the North-Island bird as a distinct species:—
“As hitherto I have had no opportunity of examining any reliable specimens from the North Island, it naturally was not possible for me to make sure about the value of certain characters. I am indebted now to the kindness of Dr. Buller for two specimens from the North Island, so that I am able to make a direct comparison of specimens from both islands. Besides the two specimens from the North Island, I have four old birds (two male and two female) and a young one from the South Island before me; also an old one and a half-grown bird, without any definite locality—consequently a total of nine specimens in different stages and conditions of age and sex. To refer, in the first place, to the tinge of colour. I had, before this, opportunities of observing that in specimens from the South Island the colour is by no means constant, but on the contrary varies from greyish brown to rusty-rod brown. The latter tone of colour, as is well known, is produced by the terminal third part of the feathers being of that shade. Each individual feather is coloured either dark brownish grey or brown, changing gradually towards the tip into rusty brown; the single filaments or barbs of the feathers, which stand far apart from each other, terminate, however, in black hair-like tips, which impart to the whole plumage the peculiar bristle-like character. In this fundamental point of colouring the specimens from both islands absolutely agree, and the feathers which I have before me, and which have been carefully pulled out, do not betray differences of any kind. Only, as I have already said, the intensity of the rust-brown on the third part of the tip of each feather is sometimes stronger, sometimes feebler, and on this depends the general colouring of the specimen. One specimen from the North Island shows the same darker and of a more vivid rust-brown than examples from the South Island. It does not, however, appear quite so dark as a specimen in the Bremen collection, without a positively defined locality, of which I have already made mention. The other specimen from the North Island, however, so perfectly agrees, in regard to the rust-brown tone of colour, with specimens from the South Island that, in point of fact, not the slightest difference is observable. Consequently the tinge of colouring as a specific character must be considered as absolutely worthless. The case is different, however, in regard to the relative hardness or softness of the plumage, which is perceptible to the touch. I am in a position to confirm the statement that in general the specimens from the North Island possess more strongly developed feather-shafts, which project beyond the barbs in the shape of naked tips, and consequently appear more like bristles and have a harsher feel. This peculiarity is very perceptible on stroking the feathers the wrong way, or on carefully feeling them; but cannot be distinguished on stroking with the palm of the hand along or in the direction of the feathers. If stroked in this way even the most delicately sensitive hand would be unable to detect any difference at all between certain specimens from the North and South Islands respectively. It is worth mentioning here that on patting the plumage of Apteryx oweni (in the manner described) the same difference as compared with Apteryx australis becomes at once apparent. What has been said in regard to the relative hardness or softness due to the more or less pronounced development of the projecting naked shaft-tips, which differ again in Apteryx oweni, has reference moreover to the plumage of the upper side of the rump. With that which covers the hind head and neck the case is different; and here perhaps might be found a single criterion, or distinguishing mark, which is appreciable not merely to the touch but also to the eye, and which might be considered as a sufficient specific character for the North-Island Apteryx. The feathers of the back of the head and the back of the neck have stronger and more projecting shafts, with the barbs composing the webs further apart and consequently less numerous. These hair-like barbs not only feel harder to the touch, but the longer and protruding hair-like filaments are quite apparent to the eye. This peculiarity I find borne out in all the specimens before me. If therefore one intends to acknowledge the Apteryx of the North Island as a distinct species, a distinguishing character could only be found in this visible difference of plumage on the hind head and back of neck. On the front and sides of the neck the peculiarity I have described is scarcely perceptible. Still, I do not venture as yet to set up this character as a constant one, as possibly there may be exceptions. Besides, this character alone does not appear to me of sufficient importance to differentiate a species. In my judgment therefore, for the present, this Apteryx of the North Island is only a slightly deviating form of the known Apteryx australis. I doubt whether it will be possible to define with certainty specimens the origin of which is not warranted, without direct comparison in all cases.”
Professor Huxley, in his ‘Characters for Classification,’ notices the absence of continued shafts as characteristic of the genus Apteryx. The abnormal character of A. bulleri in this respect is very curious.
Professor Parker has called my attention to another distinguishing feature, which appears to be constant: in Apteryx bulleri the claw on the wing is strongly curved and black; in A. australis it is less curved and whitish; in A. oweni it is much smaller and lighter coloured.