A History of the Birds of New Zealand.
The Ancient Avifauna
The Ancient Avifauna.
The first Moa-bone of which we have any record was a mere fragment of a femur six inches in length, with both extremities broken off, which was brought to England in 1839, and offered for sale at the Royal College of Surgeons by an individual who stated that he had obtained it in New Zealand from a native who told him that it was the bone of a great Eagle. Professor Owen, on its being first submitted to him, assured the owner that “it was a marrow-bone like those brought to table wrapped in a napkin”; but on subsequent and more critical examination he arrived at the conviction that it had in reality come from a bird, that it was the shaft of a thigh-bone, and that it must have formed part of the skeleton of a bird as large as, if not larger than, the full-sized male Ostrich, with this very striking difference, that whereas the femur of the Ostrich, like that of the Rhea and the Eagle, is “pneumatic,” or contains air, the huge bone, of which a fragment was now submitted to him, had been filled with marrow like that of a beast. The price asked for this unique specimen was only ten guineas, and although Professor Owen strongly recommended its acquisition, the Museum Committee declined to purchase the “unpromising fragment.” Much against the advice of his scientific contemporaries Owen insisted on publishing his conclusions, announcing boldly—“So far as my skill in interpreting an osseous fragment may be credited, I am willing to risk the reputation for it on the statement that there has existed, if there does not now exist, in New Zealand a Struthious bird, nearly, if not quite, equal in size to the Ostrich.”
After the publication of Professor Owen’s paper the bone was purchased by Mr. Bright, M.P. for Bristol, and many years subsequently came into the possession of the British Museum, where this historic relic is now carefully preserved.
More than three years elapsed before any confirmatory evidence was received from New Zealand; and then came a letter from the Rev. W. Cotton to Dr. Buckland, followed by another from the Rev. W. Williams, giving an account of the discovery of large numbers of these fossil remains and accompanied by a box of specimens, which triumphantly established the accuracy of Owen’s prevision. The specimens transmitted by Mr. Williams were, as a matter of course, confided by Dr. Buckland to the learned Professor for determination; and these materials, scanty as they were, enabled him to define the generic characters of Dinornis, as afforded by the bones of the hind extremity. An examination of a second and richer collection sent home by Mr. Williams, together with three additional specimens lent by Dr. (afterwards Sir John) Richardson of Haslar Hospital, enabled him to discriminate six distinct species of the genus, ascending respectively from the size of the Great Bustard to that of the Dodo, of the Emu, of the Ostrich, and finally attaining a stature far surpassing that of the last-named biped.
The first of these was a Cursorial bird which, on account of the agreement of its tibia in its general characters with the same bone in the larger species, he referred at that time to the genus Dinornis, but which subsequent investigations proved to belong to another genus, characterized by page xix the presence of a strong hind toe, for which the name of Palapteryx was proposed. As this bird had something of the appearance of the Great Bustard, he called it Dinornis otidiformis.
It may be here mentioned that in the Ostrich, Rhea, and Cassowary there is no vestige of a hind toe or hallux.
The next was a three-toed Struthious bird differing from the other species of Dinornis in its relatively shorter and broader metatarsus, in which characters it appeared closely to resemble the extinct Dodo (Didus ineptus) of the Isles of France and Rodriguez; and as it could not have been greatly superior in size to that bird, he named it Dinornis didiformis. Judging by its skeleton, this bird stood a little under four feet in height, or of intermediate size between the Cassowary and the Dodo. In the metatarsal of this bird, as with the larger species of Dinornis to be presently mentioned, there was not the slightest trace of the articulation of a fourth posterior toe, the generic distinction from Didus and Apteryx being thus distinctly indicated.
The next species described, which appears to have attained the average height of the Ostrich (about seven feet), with a more robust and stronger build, he named Dinornis struthioides, and pointed out characters which placed the fact of its being a good and true species beyond all cavil or doubt.
Another species, which attained the height of nine feet, he provisionally named Dinornis ingens; but, as will appear further on, this bird was also subsequently referred to the genus Palapteryx. Then came the discovery of a still larger form, standing ten feet in height if not more, which he distinguished as Dinornis giganteus. A fair idea of the size of this gigantic bird, in comparison with the stature of an ordinary-sized man, may be obtained from the accompanying woodcut, which is a reduction from the lithograph forming the frontispiece to my first edition*. The representation of the skeleton is from a photograph of the magnificent specimen in the Canterbury Museum, and the figure of the Maori, clothed in a dogskin mat and “wrapt in contemplation,” is taken from the portrait of the old Ngapuhi chief, Tamati Waka Nene, as given in Angas’s ‘New Zealanders illustrated.’
Yet another form, with a stature of about five feet, had to be discriminated, and this Owen named Dinornis dromioides, on account of its similarity in size to the Emu.
Not content with this large addition to the hitherto known Struthious birds of the world from one small area of land, the learned Professor made a happy forecast of further discoveries yet in store, for he then wrote:—
“Already the heretofore recorded number of the Struthionidae is doubled by the six species of Dinornis determined or indicated in the foregoing pages; and both the Maori tradition of the destruction of the Moa by their ancestors and the history of the extirpation of the Dodo by the Dutch navigators in the Isles of Maurice and Rodriguez, teach the inevitable lot of bulky birds unable to fly or swim, when exposed, by the dispersion of the human race, to the attacks of man. We may therefore reasonably articipate that other evidences await the researches of the naturalist, which will demonstrate a further extent of the Struthious order of Birds anterior to the commencement of the present active cause of their extinction.”
Among the most important contributors to the history of Dinornis at this early period were the Rev. William Colenso, F.R.S., who not only collected specimens of the bones, but published a very interesting memoir on the subject in the ‘Tasmanian Journal’ (vol. vii., 1843), and the Rev. Richard Taylor, who, in 1844, wrote as follows:—
“Early in 1843 I removed from the Bay of Islands to Wanganui, and my first journey was along the coast of Waimate. As we were resting on the shore near the Waingongoro stream, I noticed the fragment of a bone which reminded me of the one I found at Waiapu. I took it up and asked my natives what it was. They replied ‘a Moa’s bone; what else? Look around and you will see plenty of them.’ I jumped up, and to my amazement I found the sandy plain covered with a number of little mounds entirely composed of Moa-bones; it appeared to me to be a regular necropolis of the race. I was struck with wonder at the sight, but lost no time in selecting some of the most perfect of the bones. I had a box in which my supplies for the journey were carried; this I emptied, and filled with the bones instead, to the amazement of my followers, who exclaimed ‘What is he doing? What can he possibly want with these old Moa-bones? One suggested’ hei rongoa pea’ (to make medicine perhaps); to this the others consented, saying ‘koia pea’ (most likely).”
Other stray collections continued to arrive from time to time, till at length Mr. Percy Earl, in 1846, unearthed from the turbary deposits of Waikouaiti and sent to England a more extensive series of bones than any other collector had succeeded in bringing together. These collections all found their way, more or less directly, into Professor Owen’s hands, and he was thus enabled to rectify or confirm many of his former deductions. He was also enabled to add several new species. One of these was Dinornis casuarinus, nearly agreeing in size with D. dromioides, and combining the stature of the Cassowary with more robust proportions and especially more gallinaceous characters in the feet. A mutilated femur of this bird he had previously regarded as belonging to a young individual of the last-named species, and when he afterwards corrected the error he pointed out that it was a mistake on the safe side, “the caution which refrains from multiplying specific names on incomplete evidence being less likely to impede the true progress of zoological science than the opposite extreme.” The most abundant of the remains collected by Mr. Earl belonged to this species (D. casuarinus); but there were also in the collection bones of another very remarkable species, which was named Dinornis crassus, in allusion to the strength of its osseous frame. It was intermediate in size between Dinornis ingens and D. struthioides, and, with a stature equal to that of the Ostrich, the page xxi femur and the tarso-metatarsus of this bird present double the thickness in proportion to their length. Of this species Prof. Owen writes:—“It must have been the strongest and most robust of Birds, and may be said to have represented the pachydermal type and proportions in the feathered class.” A third new species, following next, in order of size, to Dinornis didiformis, and strictly confined in its range to the North Island, was named Dinornis curtus.
These more complete materials contained indubitable proof that Dinornis dromioides possessed, in common with D. ingens, the character of a distinct hind toe. Among the true forms of Dinornis this member was reduced (as in the Apteryx) to a high-placed hallux of diminutive size and functionless character, the attachment of this rudimentary toe being merely ligamentous. In most of the skeletons of Dinornis hitherto found there was no trace whatever of a hallux; but Professor Owen has, with every show of probability, ascribed this absence to the extremely small size of these bones and the ease with which they could be overlooked or lost rather than to their non-development, although at an earlier date he was inclined to make it a character of generic importance.
In 1850, Sir George Grey, who had been actively collecting Moabones in the district lying under Tongariro mountain, forwarded his collections to the British Museum; and two years later, Professor von Hochstetter, the naturalist attached to the Expedition of the Imperial Austrian frigate ‘Novara,’ who had undertaken a topographical examination of the North Island, obtained a rich collection from the same locality.
Most of these remains were found to belong to Palapteryx ingens, of which the annexed imaginary sketch is given in Prof. Hochstetter’s ‘Neu-Seeland’ (1863, p. 438).
Up to this period of our narrative the remains discovered appear to have belonged exclusively to birds of the Struthious Order; and, as Professor Owen had on more than one occasion explained, the existing Apteryx, notwithstanding the inferiority of size, modified structure of the palate, and page xxii different proportions of the beak, was the nearest living representative of these extinct and comparatively ancient forms*.
But new discoveries of a most interesting kind were yet in store for the great comparative anatomist, by which he was afterwards able to demonstrate further links of connection between the extinct types and still existing forms.
In 1852–55 it fell to the lot of Mr. Walter Mantell (at that time a Government Land-Purchase Commissioner) to explore the Moa-bone deposits at Waingongoro, in the North, and at Waikouaiti, in the South Island, and the extensive collections which he then made and transmitted to England not only “excited the delight of the natural philosopher, and the astonishment of the multitude,” but, having been deposited in the British Museum, these new materials, in hitherto unknown abundance, enabled the Professor not only to verify some of his former conclusions, but to establish the characters of several new genera†. No doubt the most important result was the discovery of Dinornis elephantopus, “a species which, for massive strength of the limbs, and the general proportion of breadth or bulk to height of body, must have been the most extraordinary of all the previously restored wingless birds of New Zealand, and unmatched, probably, by any known recent or extinct member of the class of Birds”‡.
The excellent woodcut on the next page, showing this skeleton as articulated in the British Museum, is copied by permission from Dr. Thomson’s ‘Story of New Zealand,’ p. 32.
* As to the affinities of the Apteryx, deducible from its anatomy, Prof. Owen says:—“Commencing with the skeleton, all the leading modifications of that basis of its structure connect it closely with the Struthious group. In the diminutive and keelless sternum it agrees with all the known Struthious species, and with these alone. The two posterior emarginations which we observe in the sternum of the Ostrich are present in a still greater degree in the Apteryx; but the feeble development of the anterior extremities, to the muscles of which the sternum is mainly subservient, as a basis of attachment, is the condition of a peculiarly incomplete state of the ossification of that bone of the Apteryx; and the two sub-circular perforations which intervene between the origins of the pectoral muscle on the one side, and those of a large inferior dermocervical muscle on the other, form one of several unique structures in the anatomy of this bird.”
† “The kainya (at the stream now known as Awamoa) which we found in 1852 afforded further unmistakable proof of the coexistence of man with the Moa—the bones and egg-shells of Dinornis and its kindred, mixed with remains of every available variety of bird, beast, and fish used as food by the aborigines, being all in and around the umus (or native ovens) in which they had been cooked. Although my collection from this place reached England in 1853, it remained unopened until after my arrival there in 1856, when I caused it to be conveyed to the crypts of the British Museum, and there unpacked it in the presence of the great authority on our gigantic birds, Professor Owen. With the exception of two small collections which were selected for me by Professor Owen, and which I gave, one to the Museum of Yale College, U.S., and the other to that of the Jardin des Plantes, the whole of this collection is now in the British Museum. The fragments of egg-shells from these umus varied in size from less than a quarter of an inch of greatest diameter to three or four inches. These, after careful washing, I had sorted; and having, with some patience, found the fragments which had originally been broken from each other, and fitted them together, I succeeded in restoring at least a dozen eggs to an extent sufficient to show their size and outline. Six or seven of the best of these I gave to the British Museum after their purchase of the collection; one is in the Museum of the College of Surgeons; the rest, including one very beautiful egg with a polished ivory-like surface, are still in my ownership somewhere in England. Some idea of the labour entailed by this attempt to rehabilitate eggs may be gathered from the fact that several of those restored consisted of between 200 and 300 fragments. I may add that in the markings, size, and so forth (making allowance for the ulteration of the former towards the ends of the eggs) I made out about twenty-four varieties, of which I have specimens.”—Mantell.
‡ “By the side of the metatarsus of Dinornis elephantopus, that of D. crassus shrinks to moderate if not slender dimensions. But the peculiarities of the elephant–footed Dinornis stand out still more conspicuously when the bones of its lower limbs are contrasted with those of D. giganteus.”—Owen.
The abundant remains of Aptornis enabled the Professor to discriminate two species, namely Aptornis otidiformis (originally, from the examination of a single bone, referred to Dinornis) and the still larger Aptornis defossor, of such size and strength that, to quote his own words, “the civil engineer might study perhaps with advantage the disposition of the several buttresses, beams, and arched plates which support the iliac roof of the pelvis, and strengthen the acetabular walls, receiving the pressure of the thigh-bones in this huge and powerful Woodhen.”
I may here mention that an Aptornis skull, dug up by Mr. W. W. Smith at Albury, near Oamaru, affords slightly larger measurements than any hitherto recorded of A. Otidiformis, and that Sir Richard Owen, to whom I presented the specimen, wrote to me saying:—“The facial part exceeds in size that of figd. 2 and 3 (pl. xliii. of the 4to vols.), but so little as not to support a distinct species, unless the rest of the skeleton affords corroborative characters†. The specimen you have kindly presented and which, for your sake, I shall value while the brief remainder of life lasts, is evidently, from its specific gravity, from a bird that has long passed away. I should, however, rejoice if confirmatory characters justified me in introducing to our Zoological Society an Aptornis bulleri.”
Then followed the discovery, in succession, of Dinornis geranoides, D. gravis, D. rheides, and D. robustus. Of the last-named species there is now an almost perfect skeleton in the museum at York, in a remarkable state of preservation, with portions of the integuments and quill-part of the feathers still attaching to the sacrum, and the legs still preserving some of the ligaments and interarticular cartilages. With this valuable series, which will be found fully described by Mr. Allis in the ‘Proc. Linn. Society’ (vol. viii. p. 50), were found the rudimentary wing-bones which had so long been an object of diligent search. The skeleton is probably that of a male bird, because lying immediately beneath it, buried in the heap of sand with which the remains were covered, were the bones of four young ones, presumably the whole of a clutch. This deduction as to the sex seems a fair one from analogy, inasmuch as the male Apteryx, and indeed that sex in the majority of Struthious birds, alone performs the duty of incubation.
* Dr. Meyer, from a careful comparison of the bones, concludes that the South-Island bird is a distinct species, for which (in his ‘Abbildungen von Vögel-Skeletten’) he has proposed the name of Notornis hochstetteri.
† In relation to measurements Prof. Owen says:—“These may perhaps be deemed by some ornithologists to be slight or trivial differences; yet, taken in connection with the greater breadth and thickness of the bone, in proportion to its length, they unquestionably support the conclusions of specific distinction deducible from those proportions.”
Subsequently another species, coming from the extreme North, was determined by Prof. Owen, and named Dinornis gracilis, on account of the remarkable length and slenderness of its legs.
But there seemed to be practically no limit to the ornithic wonders revealed by the Post-pliocene deposits of New Zealand. Professor Owen had already well nigh exhausted the vocabulary of terms expressive of largeness by naming his successive discoveries ingens, giganteus, crassus, robustus, and elephantopus, when he had to employ the superlative in Dinornis maximus to distinguish a species far exceeding in stature even the stately Dinornis giganteus. In this colossal bird, as the Professor has well remarked, some of the cervical vertebræ almost equal in size the neck-bones of a horse. The skeleton in the British Museum, even in an easy standing posture, measures eleven feet in height, and there is evidence that some of these feathered giants attained to a still greater stature.
A fair idea may be gained of its proportionate size from the accompanying woodcut, which appeared some years ago in ‘The Illustrated London News,’ representing the entire left leg of a Moa (now in the Madras Government Museum) obtained by Major Michael, of the Madras Staff Corps, from the Glenmark swamp, about 40 miles from Christchurch, where it was found in situ, at a depth of four feet, by a party of workmen who were cutting a drain. The measurements are:— Femur 1 ft. 6 in.; tibia 3 ft. 3 in.; tarsus 1 ft. 8 in.; outer toe 9 ¾ in.
The corresponding right leg was exhumed a considerable time afterwards, when Mr. Fuller was conducting a search on behalf of the Canterbury Museum, and this specimen, with the phalanges complete, is now in my private collection.
In November 1878, Mr. H. L. Squires of Queenstown, South Island, obtained and forwarded to the British Museum the head of a Moa with a continuous part of the neck, with the trachea enclosed and covered by the dried integument, and exhibiting even the sclerotic bone-ring of the dried eyeballs; also the bones of both legs with the feet covered by the dried skin, with some feathers adhering to it, and with the claws intact.
It was this specimen that enabled Sir Richard Owen to characterize his Dinornis didinus, and we may imagine the delight with which the veteran scientist embraced this opportunity of examining, for the first time, a specimen in which the characters could be studied as in a living or recent bird, and the value of his deductions from the study of single bones thereby tested, as well as the satisfaction with which he found his general conclusions so amply verified. This bird was scarcely larger than Dinornis didiformis, but presented characters of sufficient importance to separate it specifically from that form. The result of a close comparison of this dried head with that of existing Struthious birds was that “the Moa is found to repeat most closely, in the form and proportions of the beak, and. in the shape, relative positions, and dimensions of the narial, orbital, and auditory apertures, the Emus page xxv of the Australian continent.” Two points of considerable interest were established by this specimen, namely, the existence, at any rate in this species of Moa, of a strong hind toe with almost grasping power, and secondly the remarkable fact that the tarsus was feathered right down to the toes.
The other newly-discovered species (D. parvus) was founded on a nearly complete skeleton procured during the construction of a new road, about forty miles to the north-west of Nelson. The opportunity thus afforded of examining the entire osteology of a single bird was of extreme importance in the final determination of the generic characters. In size Dinornis parvus was scarcely superior to the Bustard (Otis tarda); and, although the smallest known member of this race of Struthious birds, it had proportionately the largest skull of all the Dinornithidœ. On this curious fact Owen thoughtfully remarks that if the peculiarly nutritious roots of the common fern contributed, together with buds or foliage of trees, to the food of the various species of Moa, the concomitant gain of power in the locomotive and fossorial limbs does not appear to have called for a proportionate growth or development of brain or of bill.
But the turbary deposits of New Zealand had not yet yielded up the whole of their wonderful story of the past. In the year 1868, it was discovered that the Glenmark swamp was a veritable necropolis of extinct birds. It is said that portions of no less than eighteen skeletons were dug up from the spot whence Major Michael obtained his leg of Dinornis maximus and within an area of about ten square feet. Under the able direction of the late Sir Julius von Haast, and with indefatigable zeal, these fossil remains were exhumed literally by thousands, sent to the Canterbury Museum in waggon-loads, sorted and classified there, and then distributed among the museums of the world, producing in return, by a judicious system of interchange, some £20,000 worth of specimens of various kinds, and helping materially to place the Canterbury Museum in the proud position which it now occupies in the Colony.
Sir Julius von Haast worked out the collections which he had formed in a very painstaking manner, and published the results in an Address to the Philosophical Institute of Canterbury. His minute observations and measurements over a wide field of specimens had the effect of confirming in a very remarkable manner the conclusions arrived at previously by Sir Richard Owen as to generic and specific distinctions, although these were not unfrequently based on a single bone or fragmentary part of a skeleton.
But perhaps the most important discovery was that of the existence, contemporaneously with the Moa, of a gigantic bird of prey, far exceeding the Golden Eagle in size, to which Haast gave the name of Harpagornis moorei. The evidence of this furnished by the fossil remains was fully discussed by the discoverer in a paper published in the Transactions of the New-Zealand Institute*; and it has become a favourite theme of speculation whether the true function in life of this great Raptor was not to prey upon the smaller species of Dinornis, or the chicks or young broods of the more gigantic forms.
A second and smaller species was afterwards described, under the name of Harpagornis assimilis; but it is not unlikely that this was the male of H. moorei, the disparity in size, which is the only difference, being thus easily explained.
* Vol. iv. pp. 192–196.
In a paper published in 1874, Sir Julius Haast proposed a new classification of the extinct Struthiones, which, so far, does not appear to have met with general acceptance. He divided them into two Families, which he named respectively the Dinornithidœ and the Palapterygidœ, each with two genera, the former comprising Dinornis and Meionornis, and the latter Palapteryx and Euryapteryx. He made the total absence of hind toe or hallux the distinguishing character of the first-named Family, thus following the broad line by which Owen had already differentiated his genera Dinornis and Palapteryx. He ventured, moreover, to characterize his Family Palapterygidœ as one in which the anterior limbs are entirely absent; but his conclusions on this head are far from being decisive. It would appear more likely, from the analogy of the case, that in those species in which the wings are supposed to have been wholly absent they existed only in a very rudimentary form, and that the small bones have perished, leaving no trace behind for the modern student of palæontology. It seems to be placed beyond doubt that in all the so-called Wingless Birds, by long-continued disuse of the anterior limbs through many successive generations, these organs had become enfeebled and ultimately atrophied and dwarfed to the condition of mere rudiments, as is now conspicuously apparent in the existing species of Apteryx. Professor Owen has suggested that in the case of Dinornis “the degree of atrophy, which seems to have been carried to a total loss of the limb-appendages of the scapulo-coracoid arch, implies the operation of the influence of disuse through a period of pre-Mäori æons greatly exceeding the time during which the Lamarckian law has operated on the Cassowary, the Rhea, and the Ostrich.”
Following this came the discovery by Sir James Hector of the remains of an extinct Goose, of very large if not gigantic proportions, and undoubtedly flightless. This proved to be the bird a few detached bones of which Professor Owen had previously referred to a genus “hitherto unknown to science,” and supposed to be of the Struthious Order, for which he proposed the name of Cnemiornis calcitrans. The first tolerably complete skeleton of this Anserine form, which was certainly contemporaneous with the colossal Moas, was obtained by the Hon. Captain Fraser in the Earnsclough caves, and was afterwards presented by him to the British Museum. Another coeval species determined by Professor Huxley, was the giant Penguin (Palœeudyptes antarcticus), of which the bones were discovered by Mantell in the Oamaru limestone in 1849. To the same species are doubtless referable the fossil remains more recently found by Mr. James Duigan at Hokitika. These were discovered imbedded in a reef exposed only at low water and forming part of the Seal Rock, a bold headland which protects the anchorage of Woodpecker Bay. The bones were thoroughly mineralized, resembling the condition in which fossil reptilian bones are usually found†.
* Trans. Zool. Soc. vol. xii. p. 171.
† Trans. N.-Z. Instit, vol. iv. pp. 341–346.
As already mentioned Sir Julius Haast added Dinornis oweni to the list of species; and he likewise discovered an extinct form of Apteryx, far exceeding in size those existing at the present day, to which he gave the name of Megalapteryx hectori.
Doubtless other forms, perhaps as interesting and remarkable as any yet brought to light, remain entombed to reward the zeal and enterprise of the future explorer.
Bearing on the question of the geographical relations of the New-Zealand Avifauna, one very important fact presents itself to us. In the same way that, as at the present day, certain well-marked species in the North Island are represented in the South Island by closely allied but yet specifically distinct forms, so it was also with the extinct Avifauna. The strong-limbed Moas with bulky frames were Dinornis gravis, D. crassus, D. elephantopus, D. robustus, and D. maximus, and these appear to have been strictly confined to the South Island. Six species having proportionately thin leg-bones and a slighter frame, namely, Dinornis didiformis, D. dromioides, D. gracilis, D. struthioides, D. ingens, and D. giganteus, appear to have been restricted in their range to the North Island. Dinornis rheides, which appears to have inhabited both Islands, is intermediate in the strength and thickness of its limbs; and two species remarkable for their small size—Dinornis geranoides and D. curtus—have hitherto been found only in isolated localities.
As remarked by Sir Richard Owen, in one of his latest Memoirs, Dinornis maximus is specially remarkable for its great size and strength even in a race of giants. One specimen exhibits such extreme measurements that Owen has suggested the existence of a yet taller species, for which he selects the provisional name of Dinornis altus.
Having had opportunities of examining very large series of bones, exhibiting an almost continuous gradation of size from the largest to the smallest, my own belief is that some at least of the birds differentiated above are mere varieties or conditional states of one and the same species; but their discrimination is not the less interesting and important from a scientific point of view. Even Professor Hutton, whose paper “On the Dimensions of Dinornis Bones” (Trans. N.-Z. Inst. vol. vii. pp. 274–278) goes far to establish this view, especially as to the impossibility of defining any strict line of demarcation between Dinornis elephantopus and D. crassus, is constrained to add:—“Still, notwithstanding all that I have said, I am convinced that it will be necessary to retain the names both of crassus and elephantopus to mark both ends of the series as characterized by the proportions of the metatarsus, the length of which in D. crassus is more than four times the breadth of the middle of the shaft, while the length is less than four times the breadth in D. elephantopus and D. gravis.”
* “We possess, amongst others, the leg-bone of a specimen of Dinornis maximus which is in size only second to the largest bones we have, but in which this immature character in the metatarsus is not yet quite effaced.”—Von Haast.
Not the least interesting fact connected with these giant Wingless Birds is that they have passed away within the historic period. The remains of all the species mentioned above have been discovered intermingled with human bones; they have been found, calcined and chopped, amid the rejectamenta of old Maori feasts in the ancient kitchen-middens of both Islands—facts which, quite apart from Maori tradition, prove incontestably that they were coeval with the early native inhabitants, and that their final extirpation was accelerated, if indeed it was not occasioned, by human agency.
The only question remains—At what period of history did they cease to exist? The late Sir Julius von Haast, who had devoted years of study to the subject, came to the conclusion that the extinction of the various species of Dinornis dates back perhaps a thousand years, and that the association with man, as proved by the numerous kitchen-middens and cave-habitations which he himself explored, had relation to a prehistoric or autochthonous race which, in the remote past, inhabited New Zealand*. He wrote an elaborate paper, on “Moas and Moa-hunters,” in support of this contention; but I do not think this view of the subject has obtained much support. To my mind the evidences of the comparatively recent existence of, at any rate, several species of Dinornis are overwhelming. The circumstance already mentioned of the discovery of a skeleton with a portion of the skin and feathers attached, in such a climate as that of New Zealand, is entirely opposed to the theory of remote antiquity.
Then, again, the comparatively recent date of the bones of even the larger species is attested by their chemical condition and the large amount of animal matter they contain. As compared with a recent tibia of the Ostrich, containing 26·51 of animal matter, the fossil femur of Dinornis didiformis has been found to contain 25·99. According to another comparative analysis, a recent femur of the Ostrich contained 34·86 of animal matter, and a fossil femur of Dinornis struthioides 37·86. As Professor Owen has already remarked, this superabundance of animal matter in the bone of the extinct bird is due chiefly to the fact of its being a marrow bone, whilst that of the Ostrich contains air.
* Even this champion for the great antiquity of the Moa would appear to have latterly somewhat changed his views on this subject. I have a letter in my possession from him stating that having read the report of my speech in the Native Land Court, as Counsel for the Ngatiapa in the Rangatira case, with the Appendix containing an account of Apa-hapai-taketake and the pet Moa of the Ngatituwharetoa tribe—a story accepted by both the contending parties as true—he felt almost constrained to abandon the ground he had so persistently taken up.
The following is an extract from the evidence given on that occasion by the leading witness, Hue Te Huri:—“I have heard the name of Apa-hapai-taketake an ancestor of the Ngatiapa tribe. He was the original source of the quarrel. Apa-hapai-take-take stole a Moa, which was a pet bird of the Ngatituwharetoa. While doing so he fell over a cliff and broke his thigh and was thenceforward nicknamed ‘Hapakoki’ (‘Hop and go one’). He got off with the Moa in spite of this. Then Ngatituwharetoa heard of it, and they went down upon his place and carried off his wife Hinemoatu in payment (utu) for the Moa which he had stolen. Then Hapakoki in great wrath went and seized the kumaras of Kawerau; and Ngatituwharetoa, in equal wrath, made an attack on the Ngatiapa. Then the Ngatiapa left and came to Maunganui, on the Upper Rangitikei; for all this happened at Putauaki, near the Awa-o-te-Atua, in the Bay of Plenty. The Ngatituwharetoa pursued them and attacked them at Maunganui. Ngatiapa moved on south and settled on the north-east side of the Taupo Lake; but they were followed up and again attacked, and they again moved on to Tawhare-Papauma and Moturoa, south of Taupo, and close to Rotoaira, on the edge of the lake of that name.”
The Hon.Walter Mantell, whose opinions on this point are entitled to every consideration, assigns a higher antiquity to the Moa-bones that are found under the stalagmite which forms the flooring to certain limestone caves, similar in character to those bone-caves in which traces of the early animals that inhabited Great Britain have been preserved to us; and he has declared his conviction that the more ancient species of Moa were extirpated by a race which inhabited New Zealand before the arrival of the aboriginal Maoris†.
Many of these bones have been found under a considerable depth of fluviatile deposits which may be of Quaternary or even of Pliocene age.
There can be doubt, however, from the evidences already mentioned, that some of the species, even of the largest stature, existed contemporaneously with the ancestors of the present race; and Mr. Mantell himself, during his early explorations in the South Island, discovered, drawn upon the walls of a cave in the Waitaki valley, a rude likeness of the Moa by some aboriginal artist of a bygone generation, painted with red ochre on the face of the rock, probably soon after the arrival of the first Maori immigrants.
Mr. Dallas, who, in 1865, described (Proc. Zool. Soc.) the feathers of Dinornis robustus, was the first to establish the fact that the feathers in some of the species of Moa possessed a large accessory plume or double shaft, as in the Emus and Cassowaries of Australia and the Indian Archipelago. In these feathers “the barbs consist of slender flattened fibres, bearing long silky and very delicate barbules, without any trace of barbicels.”
* Mr. Frederick Chapman (to whom I am indebted for those ‘Moa-stones’) writes :—“ When we came upon the ground disturbed by the wind (the soil being shifting sand) we soon found a number of distinct groups of gizzard-stones. It was impossible to mistake them. In several cases they lay with a few fragments of the heavier bones. In all cases they were in distinct groups; even where they had become scattered, each group only covered a few square yards of ground, and in that space lay thickly strewn…… The peculiar feature of the stones was that they were almost all opaque white quartz pebbles. In one place I found a small group of small pebbles of different colour, more like the few brown water-worn pebbles which may be picked up hereabouts. These lay with a set of bones much smaller than the very large bones I found with most of the clusters of pebbles. I did not gather these brown pebbles, as I thought it uncertain whether they were gizzard-stones or not, though it is possible that the species to which the smaller bones belonged was not so careful in selecting white stones. A glance at the pebbles lying about in the surrounding country showed that the quartz-pebbles were not collected here…… Mr. Murdoch and I collected three sets of pebbles, and these I can pronounce complete or nearly so. It is beyond question, too, that each set belongs to a distinct bird. No. 1 weighs 3 lb. 9 oz.; No. 2 weighs 4 lb.; while No. 3 weighs no less than 5 lb. 7 oz. This giant set contains individual stones weighing over 2 oz.; indeed, I have picked out 8 stones weighing almost exactly 1 lb.” (Trans. N.-Z. Inst. vol. xvii. pp. 173, 174.)
† Trans. N.-Z. Inst. vol. i. pp. 18,19.
At a somewhat later date other Moa-feathers, in an equally fresh condition, were found in a locality between Alexandra and Roxburgh; and these, according to Hutton, are distinguished by the presence of barbules to the tips, from which it may be inferred that they belonged to a less typical Struthious form.
In 1871 Dr. (now Sir James) Hector described a remarkable specimen from the same district, being the neck of a Moa, apparently of the largest size, upon the posterior aspect of which the skin, partly covered with denuded feathers, was still attached by the shrivelled muscles and ligaments*. This unique specimen was found by a gold-miner in a cave, or under an overhanging mass of micaschist, and is now in the Colonial Museum at Wellington.
* Sir James Hector, writing on this subject, says :—“The above interesting discoveries render it probable that the inland district of Otago, at a time when its grassy plains and rolling hills were covered with a dense scrubby vegetation or a light forest growth, was where the giant wingless birds of New Zealand lingered to latest times. It is impossible to convey an idea of the profusion of bones which, only a few years ago, were found in this district, scattered on the surface of the ground, or buried in the alluvial soil in the neighbourhood of streams and rivers. At the present time this area of country is particularly arid as compared with the prevalent character of New Zealand. It is perfectly treeless—nothing but the smallest sized shrubs being found within a distance of sixty or seventy miles. The surface-features comprise, round-backed ranges of hills of schistose rock with swamps on the top, deeply cut by ravines that open out on basin-shaped plains formed of alluvial deposits that have been everywhere moulded into beautifully regular terraces, to an altitude of 1700 feet above the sea-level. That the mountain-slopes were at one time covered with forest, the stamps and prostrate trunks of large trecs, and the mounds and pits on the surface of the ground which mark old forest land, abundantly testify, although it is probable that the intervening plains have never supported more than a dense thicket of shrubs, or were partly occupied by swamps. The greatest number of Moa-bones were found where rivers debouch on the plains, and that at a comparatively late period these plains were the hunting-grounds of the aborigines, can be proved almost incontestably…… Still clearer evidence that in very recent times the natives travelled through the interior, probably following the Moa as a means of subsistence, like natives in the countries where large game abounds, was obtained in 1865–6 by Messrs. J. and W. Murison. At the Maniatoto plains bones of several species of Dinornis, Aptornis, Apteryx, large Rails, Stringops, and other birds are exceedingly abundant in the alluvium of a particular stream, so much so that they are turned up by the plough with facility…… A desire to account for the great profusion of Moa-bones on a lower terrace shelf nearer the margin of the stream, led the Messrs. Murison to explore the ground carefully, and by excavating in likely spots they found a series of circular pits partly lined with stones, and containing, intermixed with charcoal, abundance of Moa-bones and egg-shells, together with bones of the dog, the egg-shells being in such quantities that they consider that hundreds of eggs must have been cooked in each hole. Along with these were stone implements of various kinds, and of several other varieties of rock, besides the chert which lies on the surface. The form and contents of these cooking-ovens correspond exactly with those described by Mantell in 1847, as occurring on the sea-coast; and among the stone implements which Mantell found in them, he remembers some to have been of the same chert which occurs in situ at this locality, fifty miles in the interior. The greater number of these chert specimens found on the coast are with the rest of the collection in the British Museum…… The above facts and arguments in support of the view that the Moa survived to very recent times are similar to those advanced, at a very early period after the settlement of the colony, by Walter Mantell, who had the advantage of direct information on the subject from a generation of natives that has passed away. As the first explorer of the artificial Moa-beds, his opinion is entitled to great weight. Similar conclusions were also drawn by Buller, who is personally familiar with the facts described in the North Island, in an article that appeared in the ‘Zoologist’ for 1864.”
* A representation of this feather appeared in the Trans. N.-Z. Institute (vol. xviii. pl. ii.); but the figure is a misleading one, as it represents the barbs thickly furnished with barbules, whereas in the feather itself they do not exist at all.
But a still more recent instance is afforded by the very interesting specimen of the Moa’s foot in the University Museum at Cambridge, obtained in the Hector. Ranges, Otago, in 1884. It was brought to England by Mr. W. J. Branford, who stated that he had himself found it in a cave where, as he believed, there was the entire skeleton of this bird and some more beside.
Professor Newton having kindly lent me this unique specimen for the purpose of having it photographed, I submitted it to Sir Richard Owen, who unhesitatingly pronounced it the metatarsal of Dinornis elephantopus*
The bone is in a perfectly fresh condition, with about four square inches of dark brown integument, having a tuberous surface and with underlying dried tendons of a maximum thickness of 1·5 inch, adhering to the proximal extremity and representing the true heel; the astragalus (or a bone that performs the same function in the Apteryx, though not hitherto recorded in Dinornis) is in position, two of the phalanges are still articulated to the metatarsal by means of dried ligament, and a portion of the tough covering of the sole, nearly half an inch in thickness and of a yellowish-brown colour, is still attached to the lower surface.
* In the foregoing pages I have made free use of Sir Richard Owen’s name in connection with the successive discoveries of the Dinornithidæ; I have stated his views as they were developed from time to time, and I have given publicity to one of his letters to myself. Under these circumstances, I thought it right to submit the proof-sheets to him before going to press. The learned Professor returned them to me without a single alteration, but accompanied by a letter which I am glad (with his permission) to place on permanent record, the more so as he assures me that the active work of his life is ended and his last contribution made to the Royal Society, of which body he has been so distinguished an ornament for upwards of half a century.
“Sheen Lodge, Richmond Park, 22nd March, 1888.
“My dear Buller,—
“Seldom have I enjoyed a morning more, in the quiet period of my existence, than during the perusal of the sheets of your ‘Introduction,’ so kindly submitted to me.
“Conclusions and inferences which bad escaped my memory have been brought back, and I seem to be repeating or living again an active period of my zoological life.
“The Moa-bone from Hector Ranges, Otago, is a metatarsal of Dinornis elephantopus……
“I do not recall anything that I could add which would heighten the pleasure your friendly visit has given me.
In 1866 two more eggs were discovered in the alluvial sandy loam of the Upper Clutha plains, Otago. One of these was two feet from the surface, the other only about a foot apart from it and three inches deeper. Of the first and more perfect one pieces were fitted together, making nearly one complete side of the egg, which was estimated to measure 8·9 inches in length by 6·1 in breadth. It contained the bones of an embryo chick, which are now preserved in the Colonial Museum. The shell had been eroded by the solvents of the soil, but on the granular surface so produced the characteristic linear air-pores were distinctly visible. The shell yielded 0·9 per cent. of organic matter, showing that it had not been long enough in the soil to part with all its soluble constituents†.
No one who takes the trouble to examine the skeleton of Dinornis parvus which now stands in the Palæontological gallery of the British Museum, exhibiting bleached but not fossilized bones, some of them still retaining their inherent “grease,” will be able to resist the conclusion that the bird to which they belonged was living at a comparatively recent date.
The well-marked footprints of the Moa in the sandstones of Poverty Bay—models of which are now to be seen in most of our public museums—are interesting historically, but their presence in such a formation is quite consistent with the alleged antiquity of the bird. The case is different, however, with the round cakes of excrement collected by Mr. Taylor White, with other Moa remains, in the Wakatipu cave, the condition of which was such that undigested fragments of fern-stalk and other vegetable matter could still be detected§.
Further evidence of the comparatively recent existence of Dinornis is afforded by the fact that mixed with its remains are the bones of many species of birds still inhabiting New Zealand. Among these I may mention the following genera:—Nestor, Stringops, Platycercus, Himantopus, Hæmatopus, Limosa, Larus, Diomedea, Rallus, Porphyrio, Anas, Phalacrocorax, and Eudyptula.
* This egg was submitted to me for examination soon after its exhumation, and I made the following notes:—It is perfectly oval and measures 9·3 inches in length by 7 inches in breadth; of a pale cream-colour, stained on one side with yellowish brown as if it had been smeared with the yoke, an appearance which may have been due to contact with decomposed animal matter. The egg has a solid appearance, the surface looking more like half-polished Moa-bone than egg-shell, and its thickness is about that of a new shilling. The entire surface is covered with short linear air-pores, or minute puncta, as if made with the point of a pen-knife, and disposed longitudinally; being filled with darker matter than the shell they present the character of pencilled markings, varying in extent from mere points to lines one tenth of an inch in length; there are some irregular dark markings on one portion of the egg having the appearance of obscure marbling, but these do not seem to be inherent in the shell.
† Hector in Proc. Zool. Soc. 1867, p. 996.
§ Trans N.-Z. Instit. vol. viii. p. 99.
From what we know of the range and habits of the Struthiones in other parts of the world, it cannot be supposed that the extinct race of Moas, comprising twenty, if not more, species or varieties, some of them attaining to colossal dimensions, were always confined within the geographic limits of modern New Zealand. The Ostrich inhabits the arid deserts of Africa, the Rhea (of which there are two, if not three, species, each occupying a separate district) is spread over a great portion of America, extending from Patagonia to Peru, two species of Emu and a Cassowary occupy the Australian continent, the range of each being well defined, and eight other species of Cassowary are limited to New Britain, New Guinea, and the islands of the Indian Archipelago, each inhabiting a separate area. It may be safely assumed that the Moas of the remote past roamed over a wide continent now submerged, and that when, by the gradual subsidence of their domain beneath the waters of the Great Pacific, they were driven as it were into a corner and overcrowded, the struggle for existence became a severe one and the extinction of the race commenced; that the more unwieldy giants, thus “cabined and confined,” were the first to succumb; and that the smaller species, perhaps in course of time differentiated from their ancestors by the altered physical conditions of their environment, continued to live on till their final extirpation by man within recent historic times.
Professor Owen compares New Zealand to one end of a mighty wave of the unstable and ever-shifting crust of the earth, of which the opposite end, after having been long submerged, has again risen with its accumulated deposits in North America, showing us in the Connecticut sandstones of the Permian period the footprints of the gigantic birds which trod its surface before it sank; and he surmises that the intermediate body of the land-wave, along which the Dinornis may have travelled to New Zealand, has progressively subsided, and now lies beneath the Pacific Ocean. But Professor Hutton, in his treatise ‘On the Geographical Relations of the New Zealand Fauna,’ considers it necessary to account for the phenomenal number of Struthious species inhabiting New Zealand, as compared with the other much larger areas of the earth’s surface. He supposes the existence of an ancient continent, with one or two species of Dinornis; then, by some convulsion of nature, this continent sinks beneath the ocean, leaving its mountain-ranges exposed, in the form of islands, and the only refuge for the surviving Moas; after a sufficiently long period to allow of specific changes, there is an elevation of the land and the differentiated birds are mingled together; then follows the final subsidence, when New Zealand as the central mountain-chain becomes a “harbour of refuge” for them all. In support of this bold hypothesis he refers to the remarkable fact of five or six distinct species of Cassowary inhabiting isolated localities extending from New Britain and New Guinea to the Molucca Islands. His general conclusion is thus expressed:—“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 page xxxv 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.”
Mr. A. R. Wallace, commenting on this, in his ‘Island Life,’ says:—“If New Zealand has really gone through such a series of changes as here suggested, some proofs of it might perhaps be obtained in the outlying islands which were once, presumably, joined with it. And this gives great importance to the statement of the aborigines of the Chatham Islands, that the Apteryx formerly lived there, but was exterminated about 1835. It is to be hoped that some search will be made here and also in Norfolk Island, in both of which it is not improbable remains either of Apteryx or Dinornis might be discovered. So far we find nothing to object to in the speculations of Captain Hutton, with which, on the contrary, we almost wholly agree; but we cannot follow him when he goes on to suggest an Antarctic continent uniting New Zealand and Australia with South America, and probably also with South Africa, in order to explain the existing distribution of Struthious birds.… The suggestion that all the Struthious birds of the world sprang from a common ancestor at no very remote period, and that their existing distribution is due to direct land communication between the countries they now inhabit, is one utterly opposed to all sound principles of reasoning in questions of geographical distribution…… We have direct proof that the Struthious birds had a wider range in past times than now. Remains of extinct Rheas have been found in Central Brazil, and those of Ostriches in North India; while remains, believed to be of Struthious birds, are found in the Eocene deposits of England. As the intervening sea appears to be not more than about 1500 fathoms deep it is quite possible that such an amount of subsidence may have occurred. It is possible, too, that there may have been an extension northward to the Kermadec Islands, and even further to the Tonga and Fiji Islands, though this is hardly probable, or we should find more community between their productions and those of New Zealand. A southern extension towards the Antarctic continent at a somewhat later period seems more probable, as affording an easy passage for the numerous species of South-American and Antarctic plants, and also for the identical and closely allied freshwater fishes of these countries.”
Professor von Haast, in his ‘Anniversary Address to the Philosophical Institute of Canterbury’ (1874), objected to the above theory on the ground that the geological record of these islands, so far as we are acquainted with it, does not warrant our assuming such repeated changes in the level of the land. He thus states the case:—“An unfortunate country, such as New Zealand, of which a good number of the species of its fauna and flora show great resemblance to other species from distant countries, has to be dipped down and brought up again a great many times in order to establish connexions in various directions, so that a bird or fish, a shell, insect, or centipede might cross from the one to the other, moreover, without allowing any other species from the same country to pass.” But Professor Hutton, with a much broader grasp of the subject, returns to the discussion in an able article ‘On the Origin of the Fauna and Flora of New Zealand’ (‘Journal of Science,’ 1884, vol. ii. parts 1 and 6), in which, after qualifying his former theory by abandoning the idea of an extensive Antarctic continent, and substituting a South-Pacific continent connecting New Zealand with South America, he defends his views with considerable force of argument*.
* Contending all through that in Miocene times New Zealand was represented by a cluster of twenty or more islands, on which the various species of Moa were probably developed, Professor Hutton thus sums up his conclusions:—“Our general results then are that in early Mesozoic times New Zealand, Eastern Australia, and India formed one biological region, land probably extending continuously from New Zealand to New South Wales and Tasmania. At the close of the Jurassic period the New-Zealand Alps were upheaved and the geosynclinal trough between New Zealand and Australia was formed. During the Lower Cretaceous period a large Pacific continent extended from New Guinea to Chili, sending south from the neighbourhood of Fiji a peninsula that included New Zealand. Nearly all the southern part of America was submerged. Western Australia and Eastern Australia formed two large islands lying at some distance from the continent. This continent supported dicotyledonous and other plants, insects, land-shells, frogs, a few lizards, and perhaps snakes and a few birds, but no mammals. In the Upper Cretaceous period New Zealand became separated and reduced to two small islands; the South Pacific continent divided in the middle between Samoa and the Society Islands and—the eastern portion being elevated while the centre sank—it ultimately became what we know now as Chili, La Plata, and Patagonia. In the Eocene period elevation commenced in our district; Eastern Australia was joined to New Guinea, which stretched through New Britain to the Solomon Islands. New Zealand was also upheaved and extended towards New Caledonia, but the two lands were divided by an arm of the sea. The mainland of New Guinea had by this time been invaded from the north by a large number of plants, birds, lizards, snakes, &c., which migrated south into Eastern Australia and a few passed over the New-Caledonia channel and reached New Zealand. But still no mammals. In the Oligocene period New Zealand again gradually sank, carrying with it the sparse flora and fauna it had received, and in Miocene times was reduced to a cluster of islands; Eastern Australia all this time receiving constant additions to its fauna and flora through New Guinea. In the Pliocene period elevation once more took place; New Zealand extended towards the Kermadeo Islands, and the continent of Australia was formed; after which subsidence again occurred in New Zealand.”