OOLOGICAL SOCIETY OF LONDON. Vor. Hi —Parr 3. oe wD MOONES Pp a ale ee on crwengeto LONBON: PRINTED FOR THE SOCIETY, 7 “A ND-SOLD “AT THE. SQ AEX'S ROUSE, IANOVFR SQUARE; SOLD ALSO BY LONGMAN, BROWN, GREEN, AND TONGMANS, {\NOSTER-LOW, nud J.B. Taylor, Printers Red Lion Court, Flect &treer. vier, J ''SH! WILLIAM DILLER MATTHEW | GIFT OF | WILLIAM DILLER MATTHEW '' '' '' wey aid a EARTH , SCIENCE Say staan us ary » ie } ul 2 a Sea ao y es > pea > ‘ 37.9) i X. On Dinornis', an extinct Genus of tridactyle Struthious Birds, with descriptions of portions of the Skeleton of five Species which formerly existed in New Zealand. By Professor Own, M.D., F.R.S., Z.8., &c. &c. (Part t.) Communicated November 28th, 1843. Introduction. THE brief history of the discovery of the Dinornis, a genus of gigantic terrestrial birds, which appears to have become extinct within the historical period in the North Island of New Zealand, like the Dodo in the island of Mauritius, will be found in the Proceed- ings of the Zoological Society for November 1839, and in the Society’s Transactions, vol. ili. p. 32, pl. 3. These papers contain the inferences deduced from the structure of the shaft of a femur, which led to the first announcement of the former existence in New Zealand of a large Struthious bird ‘of a heavier and more sluggish species than the Ostrich.” As the full development and confirmation of this idea is included in the following pages, I am induced, in vindication of the fruitful principle of physiological correlations, the value of which as an instrument in the interpretation of organic remains there has been a tendency to depreciate in an otherwise estimable osteological work’, to pre- mise the abstract of my former communication published four years ago :— “The fragment is the shaft of a femur, with both extremities broken off. The length of the fragment is six inches, and its smallest circumference is five inches and a half. The exterior surface of the bone is not perfectly smooth, but is sculptured with very » shallow reticulate indentations ; it also presents several intermuscular ridges. One of these extends down the middle of the anterior surface of the shaft to about one-third - from the lower end, where it bifurcates ; two other ridges or line aspere traverse lon- gitudinally the posterior concave side of the shaft; one of them is broad and rugged, the other is a mere linear rising. ‘‘The texture of the bone, which affords the chief evidence of its ornithic character, presents an extremely dense exterior crust, varying from one to two lines in thickness ; then there occurs a lamello-cellular structure of from two to three lines in thickness. The lamelle rise vertically to the internal surface of the dense wall, are directed ob- liquely to the axis of the bone, decussate and intercept spaces which are generally of a rhomboidal form, and from two to three lines in diameter. This coarse cancellated structure is continued through the whole longitudinal extent of the fragment, and im- 1! Aewds, surprising, opves, bird. 2 The ‘ Osteographie’ of Prof. De Blainville. VOL. III.—PART III, a1 ® 767407 ''285 : PROFESSOR OWEN ON THE GENUS DINORNIS. res es apediataly bounds the medullary cavity of the bone, which is about one inch in diameter *" "St the middle, and slightly expands towards the extremities. There is no bone of similar size which presents a cancellous structure so closely resembling that of the present bone as does the femur of the Ostrich ; but this structure is interrupted in the Ostrich at the middle of the shaft where the parietes of the medullary, or rather air-cavity, are smooth and unbroken. From this difference I conclude the Struthious bird indicated by the present fragment to have been a heavier and more sluggish species than the Ostrich ; its femur, and probably its whole leg, was shorter and thicker. It is only in the Ostrich’s femur that I have observed superficial reticulate impressions similar to those on the fragment in question. ‘The Ostrich’s femur is subcompressed, while the present frag- ment is cylindrical, approaching in this respect nearer to the femur of the Emeu; but its diameter is one-third greater than that of the largest Emeu’s' femur, with which I have compared it. “ Ib. fig. 2: 3 Ib. fig. 4. ''PROFESSOR OWEN ON THE GENUS DINORNIS. 251 of Dinornis didiformis, characterized by superiority of size, or to a distinct species of Dinornis. Comparing the femora f 6, f 13 (Pl. XXIII. fig. 1.) and f16 (Ib. fig. 2.) with each other, it was obvious that one of them differed in its proportions from the rest, f 13 being relatively thicker, as is shown in the plate and in the table of admeasurements. This femur corresponded much more closely with the femur f 12 (Pl. XXI. fig. 3.) in its general form, its ridges and tuberosities ; but these were less strongly developed, and the manner and extent of abrasion of both proximal and distal articular surfaces would well accord with the supposition of their having been in that cartilaginous or less completely ossified state which characterizes the femur of a bird not quite fully arrived at maturity. The state of development of the muscular ridges and tuberosities forbids the reference of this femur to a very young bird, but supports the conclusion that the bone had belonged to an individual as far advanced in growth as is indicated by the difference in size be- tween it and the femur f 12. The different condition and proportions of the two remaining femora, of 93 inches in length, f 6 and f 16, establish their specific distinctions from the femora f 13, f 12 and f 2. Of this I think no doubt can be entertained by any anatomical naturalist who may inspect the plate (Pl. XXIII.) containing the figures of f 13 and f 16, selected for the comparison, or who may give due consideration to the following statement of their differential characters. These bones are of equal length but of unequal thickness: the shape of the shaft of the bone is also different ; the relative antero-posterior diameter of f 13 is much greater than that of f 16, especially at the proximal end and trochanterial enlargement of the shaft, and just above the inner condyle: the anterior surface of the proximal part of the shaft presents a shallow equable concavity in f 16 which is not present inf 13. In f 16 a pretty sharp ridge leads from the middle of the posterior surface of the shaft ob- liquely to the upper and posterior angle of the inner condyle, and the posterior surface of the expanded shaft above the condyles is regularly excavated by a moderate concavity which is continued uninterruptedly into the inter-condyloid depression. In f 13 an oblong rough tuberosity, with its long axis parallel with that of the bone, exists in the place where we find the oblique ridge in the other bone, the tuberosity being separated from the upper and posterior angle of the inner condyle by a smooth channel or de- pression, which leads to an oval depression much deeper and more circumscribed than is the corresponding concavity in f 16. The complete development of the muscular ridges and tuberosities, with the better preserved state of the articular extremities, show the femur f 16 to be a more mature bone than f 13; the differences in proportion and configuration prove it to belong to a distinct species from Dinornis struthoides. We next come to the question whether the femora f 6 and f 16 belong to the species Dinorms didiforms, founded on the femora f 7, f 8, f 17, the tibia ¢ 3, t 4, ¢ 5, t 8, t 9, t 10, and the metatarsi m 4, m 5 and m 6, and whether the femora f 6 and f 16 VOL. I11.—PART III. 2 1 ''252 PROFESSOR OWEN ON THE GENUS DINORNIS. represent a sexual superiority of size, or are specifically distinct from the shorter femora. If the discrepancy of the thickness of the shaft as compared with the length of the bone be sufficiently obvious in femora of equal length, like f 13 and f 16 (Pl. XXIII), it becomes still more striking when the more robust proportions are exhibited in a femur of shorter size, which is one of the first differences that strike the eye in comparing f 6 and f 16 (Pl. XXII.) with f 7, f 8 and f 17 (Pl. XXIV.). The table of admeasurements shows that the femur f 17, which is one inch and five lines shorter than the femur f 16, has very nearly an equal circumference of the middle of the shaft, and a quite equal breadth of the distal end, the antero-posterior diameter of the condyles being also the same in both. If the comparison of these two femora be pursued into further details, it is seen that the anterior margin of the great trochanter is more produced but narrower in f 16 than in f 17, that the anterior surface of the shaft is more convex, and that the anterior curve of the outer condyle is shorter in the longer femur: the antero-posterior diameter of the great trochanter and of the shaft, especially of that part leading to the outer condyle, is less in the longer femur. With regard to the configuration of the popliteal space, the same differences exist between f 16 and f 17 as have been already pointed out between f 16 and f 13, viz. a circumscribed tuberosity (d, fig. 2. Pl. XXIV.) in place of a continuous ridge (d, fig. 2. Pl. XXII.), a deeper and smaller instead of a shal- lower and larger concavity, &c. In regard to the relation of these differences to sex, it is to be observed, that the male Ostrich slightly exceeds the female in size, and the difference between the two sexes of the Apteryx is relatively greater, yet the femora and other bones of the leg do not differ at all in proportions or configuration, but only in size, corresponding in degree with the rest of the body. I am not, indeed, aware of a single fact in the osteology of ex- isting birds which would justify the conclusion that the differences presented by the femur f 16, as compared with f 17, were merely sexual. It has already been shown that the differences between f 16, as compared with the femora f 13 and f 12, cannot depend upon nonage, and, @ fortiori, the femur f 16 cannot be regarded as belonging to a young individual of the gigantic species: there remains then only the conclusion that it must represent a fifth distinct species, of which there are neither tibiz nor metatarsi in the present collection. I venture to surmise that the tibia, and especially the tarso- metatarsus of this species, will be found relatively longer and more slender than in the Dinormthes struthoides and didiformis: so much may be anticipated from the more slender proportions of the femur, which, moreover, resembles the femur of the Emeu in some of the characters by which it differs from the above species of Dinornis, viz. in the sharper anterior border of the great trochanter, the more equable and deeper concavity between this border and the head of the femur, and in the uninterrupted ridge leading from the middle of the back part of the bone to the inner condyle. The generic cha- racters of Dinornis are, however, manifested in the absence of the air-hole and air-cavity ''PROFESSOR OWEN ON THE GENUS DINORNIS. 253 of the femur, the greater robustness of the bone in this the least robust of the genus, the much higher trochanter, the much wider distal extremity, and especially the wider and shallower cavity for the patella. From the equality of size of this femur with that of the Emeu, the species which it indicates may be termed Dinornis dromeovdes. Pelvis. (Plates XIX. XX. & XX a.) The first portion of the pelvis here described consists of twelve anterior anchylosed vertebra of the sacrum, with a portion of the right ilium and acetabulum (Pl. XIX. fig.1.). Of the size of this fine fragment an idea will be given by the subjoined table of its dimensions, compared with those in a full-sized Ostrich. Dinornis,p 1. — Struthio. In. Lin. In. Lin. Height of the first sacral vertebra . . . ee 4 6 Breadth of the articular surface of the body af aie ‘ae i. § Breadth of the seventh sacral vertebra. . . . . . 3 8 Le Length of the. first seven sacral vertebra...) s .. «4. 0 8 5.38 The last admeasurement shows that the anterior part of the sacrum, including the first series of vertebree provided with double transverse processes on each side!, is shorter in proportion to its height and breadth compared with the Ostrich ; and these proportions are shown to characterize the entire pelvis by the smaller specimen, subse- quently to be described. The under surfaces of the first seven vertebra are flattened, and form a smooth and slightly concave platform in the remaining four. The inferior transverse processes pass out horizontally to the lower border of the ilium, which de- scends to the level of the under surface of the bodies of the sacral vertebree. In the Ostrich they ascend obliquely upwards to join the upper transverse processes, before abutting against the lower border of the ilium, which does not descend so low as the bodies of the vertebre. In the Ostrich the first two inferior transverse processes of the sacrum retain their primitive condition of detached ribs, and three transverse processes succeed them before the commencement of the os pubis. In the great Dinornis the second sacral rib is an- chylosed as a transverse process, and four other processes succeed this before the one which abuts against the beginning of the pubis: this is much thicker and stronger than the preceding ones, and it is succeeded by four confluent sacral vertebra, which have no lower transverse processes. In the Ostrich the transverse processes of the sixth sacral vertebra abut against the part of the innominatum from which the pubis is continued, and the transverse processes of the four succeeding vertebre abut against the origin of the ischium, parallel with the lower part of the acetabulum ; then a single vertebra with- out a lower transverse process or sacral rib intervenes before these are again developed, to abut against the posterior part of the acetabulum. ! See description of the sacrum in -Birds, in the ‘ Cyclopedia of Anatomy,’ art. Avzs, p. 271, 242 ''7 254 PROFESSOR OWEN ON THE GENUS DINORNIS. The four ribless sacral vertebra, which in the Dinornis are interposed between those which send their anchylosed ribs to abut upon the os innominatum anterior to the ace- tabulum, and those which strengthen in like manner the posterior part of the acetabu- lum, are very short ; their bodies have coalesced into a single mass of bone, smooth and flattened below, rounded at the sides, and only recognizable as distinct bones by the orifices for the nerves at the sides of the anchylosed mass: these orifices are double, as in the sacrum of other birds’, the two roots of the nerves escaping separately, the motor root issuing by the lower, the sensitive root by the upper orifice. The upper transverse process of the first sacral vertebra is a broad and thick piece of bone, extending from the body and anterior articular process of the vertebra, and having a deep and smooth excavation at its anterior part: in the Ostrich the correspond- ing part is much smaller and is reticulated by the bars of bone dividing the orifices by which the air is admitted into the interior of the vertebra. I shall not swell the extent of the present paper by pursuing -farther the description of the structure of the pelvis of the Dinornis, as exhibited in the present striking frag- ment, but proceed to notice the other specimens of pelvic bones which have been enu- merated as forming parts of the present collection. The large portion of the right os innominatum, including the entire acetabulum (Pl. XX. fig. 1.), must have belonged to a bird of rather smaller size than the one to which the above-described portion of the sacrum belonged. The part of the ilium before and above the acetabulum rises with a steep slope and a slight general concavity to meet its fellow above the spinous crest of the anterior part of the sacrum: behind the acetabulum the outer surface of the ilium is divided into two facets, the upper one nearly horizontal, the lower one vertical, save where it arches out to the flat arti- cular surface behind the acetabulum. The ridge dividing these two facets commences anteriorly above the middle of the acetabulum, and describes a regular curve in its course backwards, the convexity being downwards: in the Ostrich the correspond- ing ridge forms two curves, meeting at an angle above the prominent articular sur- face behind the acetabulum, and the convexity of both curves is upwards; from the angle an obsolete ridge extends down to the prominent articulation, and divides the ante- rior from the posterior vertically concave surfaces of the ilium: in the great Dinornis the corresponding surfaces are uninterruptedly continuous above the acetabular promi- nence. The posterior wall of the acetabulum (f) is incomplete, as in other birds; the smooth articular surface is continued upon an oblong prominence above and behind the cavity. The pubis (d), a slender bone, as usual in Birds, springs from a protuberance at the lower part of the acetabulum. The ischium (e) is continued more directly from the lower and back part of the cavity : a very slight ridge indicates the posterior boundary of the notch for the tendon of the obturator internus, and the upper border of the notch * Cyclop. of Anat., art. Aves, p. 271. The Ostrich is the only exception to this rule with which I am acquainted. ''PROFESSOR OWEN ON THE GENUS DINORNIS. 255 is nearly straight. In the Ostrich this part is concave, and a well-developed process extends down, but does not join the pubis at the back part of the obturator notch. The Apteryx resembles the great Dinornis in this part of the pelvis. The ischium be- comes compressed and gradually expands vertically as it extends backwards, its lower margin forming almost a straight line. In the Ostrich the ischium maintains its trie- dral form for a much longer extent and suddenly expands, the lower margin curving down to join the pubis (Pl. XIX. fig. 4. e): there is no indication of such a junction in the present specimen, nor does the superincumbent ilium curve down, as in the Bus- tard, to join the ischium: both the ischiadic and the obturator notches seem to have been unclosed by bone in the Dinornis as in the Apteryx. The third specimen of the pelvis of a Dinornis, p 4 (Pl. XIX. fig. 2. & Pl. XX. figs. 2 & 3.), is more entire, but much smaller than the foregoing. It seems to include all the sacral vertebree, which are eighteen in number : seven anterior ones with the lower trans- verse processes, four without those processes, and seven in which they reappear, ex- tending obliquely outwards and backwards to the line of junction of the ilia with the broad posterior part of the sacrum. The most important feature in the present pelvis is the demonstration of what was obscurely indicated in the foregoing specimen, viz. that the ilia do not, as in existing Struthious birds, including the Apteryx, approximate one another along the whole length of the sharp and narrow ridge formed by the spines of the sacrum, but that they diverge above the acetabula, to give place to a broad hori- zontal expanse of bone developed from the posterior sacral spines (Pl. XX. fig. 3. 6), as in the Bustard and most other birds. This surface forms a smooth shallow concavity, perforated as usual by two lateral series of small foramina. From the pelvis of the Bustard that of the Dinornis differs in the greater relative depth and verticality of the anterior plates of the ilia, which meet above to form a ridge, as in the existing Struthio- nid@ : the posterior expanded part of the pelvis is relatively shorter than in the Bustard, and the difference is extreme which this part of the pelvis of the Dinornis presents, as compared with that of the Apteryx', the Ostrich, the Emeu, and @ fortiori the Rhea, in which the ischiadic bones meet, and are united for a considerable extent below the posterior part of the sacrum, which there becomes almost obliterated. The acetabula are relatively nearer to each other than in the Bustard, but farther apart than in the Ostrich, Emeu, and relatively than in the Apteryx. There is likewise another difference in the relative position of the acetabula as compared with the Ostrich : in this bird those cavities are so situated that their posterior wide orifice exposes to view the neural arches and spinous processes of the intervening sacral vertebrae. In the Dinornis only the lower part of the bodies of the corresponding vertebra are seen by looking directly into the acetabulum (Pl. XX. fig. 2. f), and below these we have the open cavity of the pelvis: the Apteryx and Emeu resemble the Dinornis in this respect : nothing but the cavity of the pelvis is seen on looking directly through the acetabula 1 See Zool. Trans. vol. ii. p. 291. ''256 PROFESSOR OWEN ON THE GENUS DINORNIS. in the Bustard. ‘The body of the third sacral vertebra is carinate below in the Bustard, and none of the vertebra abut by their transverse processes against the anterior part of the acetabulum. The smaller pelvis of the Dinornis, p 4, when compared with the portions of the larger pelves, p 1, p 2, presents so many differences besides those of size as to leave no doubt about the specific distinction of the birds to which they belonged. The first sacral vertebra in the smaller pelvis (Pl. XX. fig. 2.) has a narrower and deeper body, and there is not the deep excavation on the anterior part of the upper trans- verse process : I do not lay much stress on the fact that the lower transverse processes of the first two sacral vertebre retain the condition of ribs articulated to depressions at the upper part of the intervertebral spaces (2b. a) ; but every other part of the present pelvis manifests the characters of maturity. These ribs, as well as their anchylosed ana- logues, the transverse processes which succeed them, come off higher up than in the large pelvis. ‘The lower border of the ilium is thin, and does not form a broad convex surface, increasing the width of the pelvis anterior to the acetabulum, as in the large Dinornis : the four inter-acetabular vertebrz without inferior transverse processes (ib. 6) are carinate along their under surface, not flattened as in the great Dinornis. The upper facet of the posterior part of the ilium is more horizontal, and forms a right angle with the vertical facet (Pl. XX. fig. 2. c): this is also divided from the anterior concave wall of the ilium, as in the Ostrich, by an angle formed by an obsolete ridge: the articular prominence behind the acetabulum is relatively longer in the axis of the pelvis, but less deep in the smaller species. The root of the ischium where it forms the upper part of the obturator notch is concave, and an angular process descends towards the pubis, forming a well-marked posterior boundary to the notch. In this character the smaller pelvis more resembles the pelvis of the Emeu than does that of the larger one; but the ischial process does not quite reach, as in the Emeu, the pubic bone. The ischium re- sembles, in its gradual expansion and straight direction, that of the larger species, and the more perfect condition of the smaller pelvis proves that the extremity of this bone (ib. e) projects freely backwards, as in the Apteryx and Emeu. The portion of a somewhat smaller pelvis, p 5 (Pl. XIX. fig. 3. Pl. XX. fig. 4.), than the preceding is less complete, but manifests characters which prove it to belong to a distinct species of Dinornis, and apparently to an older bird, since the second sacral rib on the left side is anchylosed to the vertebral interspace. This anchylosis sufficiently demonstrates that the smaller pelvis is not of a younger bird than the larger one ; and, besides the difference of size, there are the following differences of configuration :—In the smaller pelvis the second and third sacral ribs arise nearer the lower surface of the bodies of the vertebrze, a character by which the smallest pelvis approximates the largest one, from which it differs, in having the bodies of these vertebrae relatively less broad and flat. The extent occupied by the four posterior orifices forming the interspaces of the lower transverse processes of the third to the seventh sacral vertebrae inclusive, is three lines greater in the smallest pelvis, p 5, than in the one next in the order of size, ''PROFESSOR OWEN ON THE GENUS DINORNIS. 257 p4. The bodies of the four vertebra without lower transverse processes (b) are flatter below in the smallest pelvis than in p4. The vacuity at the sides of these vertebre, into which the posterior aperture of the acetabulum opens, is relatively much smaller in the p 5 than in p 4; but the two transverse processes of the twelfth and thirteenth vertebre which abut against the posterior part of the acetabulum are absolutely much thicker in p 5 than in p4. For example, the first of these transverse processes in the larger pelvis p 4 is one inch seven lines long and one and a half line broad: in the smaller pelvis it is ten lines long and four lines broad. Such differences are not mani- fested in the pelvis of individuals of the same species in other birds: they are associated in the present instances with minor differences in the shape of the acetabulum, espe- cially of its posterior and inferior border, and in the relative breadth of the bodies of the posterior sacral vertebre ; the latter, however, might be a sexual difference. Seven- teen of the sacral vertebra are preserved in the specimen p 5, and the expanded spi- nous plate of the posterior ones is more perfectly preserved than in the preceding specimen, p 4. The following are some of the dimensions of p 4 and p 5, compared with those of the pelvis of an Emeu :— Dinornis, p 4. Dinornis, p5. Dromaius. In. Lin. In. Lin. In. Lin. Lepeth of togelvis.. ee a o O° Fou Greatest breadth at post-acetabular protuberance . . 6 9 5 10 4 0 depth at the origin of pubic bones ees a 0 570 Breadth of the pelvis at the posterior part of the ilia . 3 6 uD Lg After a summary of the characters which the different pelves of the Dinornis present in common, it is obvious that the genus recedes furthest from the Struthious type and makes the nearest approach to the tridactyle Gralle in the structure of this part of the skeleton. There remains to be determined, to which of the species of Dinornis already esta- blished by the bones of the leg, the three specifically-distinct pelves are to be referred. We cannot take the relative lengths of the pelves and femora of existing Struthionide as a guide, on account of the disproportionate length of the pelvis in them as compared with the Dinornis. The pelvis of the Bustard is also relatively longer than in the Di- nornis. The size of the acetabulum is not so good a guide as in Mammalia, because it is always relatively larger than the head of the femur in Birds: thus, although the ace- tabulum in the portion of the pelvis p 2 is larger than the head of the entire femur f 2, the size of the post-acetabular articular prominence corresponds sufficiently closely with the articular surface upon the neck and trochanterian enlargement of that femur to render it probable that they belonged to the same species, if not individual. The fragment of the huge pelvis p 1 might well have belonged to a Dinornis of the largest stature. ‘The two remaining specimens of pelvis are too small to be adapted to the femur f 12, but of the species indicated by that and the tarso-metatarsal bone m 3, there appears to be a small portion of a pelvis in the present collection. ''258 PROFESSOR OWEN ON THE GENUS DINORNIS. This is a part of the right os innominatum, p 3 (Pl. XX. fig. 5.), including the posterior and inferior angle of the acetabulum, the origins of the pubis and ischium, which form the obturator notch, and a fractured continuation of the latter bone. The fragment has belonged to a pelvis intermediate in size between p 2 and p 4, but is nearer the former. From this it differs in the concavity of the upper boundary of the ischiadic notch, and the descending process forming its posterior boundary which almost touches the pubis. The posterior margin of the wall of the acetabulum is straight, and ascends at a right angle with the horizontal ischium. In the larger pelvis, p 2, as in the smaller one, p 4, this margin curves back at less than a right angle. The ischium is thinner and less convex internally. The pelvis p 4 agrees in the proportions of its acetabula and the form of the posterior articular protuberance with the femora of Dinornis dromaoides ; and the smaller pelvis p 5 offers the same correspondence with the femora of the Dinornis didiformis. If a species of Dinornis intermediate in size between the D. struthoides and D. didi- formis had not been indicated by the femora f 6 and f 16, I must have heen led to the same conclusion by the two pelves, p 4 and p 5, as to the existence of such a species. Since the foregoing description of the pelvic bones was put into type I have been favoured by William Cotton, Esq., F.R.S., with the view of some specimens of the bones of the Dinornis, very recently transmitted by his son the Rev. Wm. Cotton, M.A., from New Zealand, one of which is a fractured pelvis', corresponding in length and in so many other characters with p 5, as to lead to the conclusion that it belongs te the same species, and that the differences between them are attributable to sex. These differences are the following. The bodies of the first two sacral vertebre in Mr. Cotton’s specimen, which I shall call p 6, are flatter on their under surface and broader ; the form of the anterior articular surface of the first sacral vertebra more nearly resembles that in the largest fragment, p1. The spines of the seven anterior vertebre and the co-ascending plates of the iliac bones are less elevated in p6. Thus, from the end of the transverse process of the sixth sacral vertebra to the summit of the ilium in p 5, is three inches ten lines, whilst in p 6 it is only two inches nine lines. The length of the part of the pelvis formed by the first seven sacral vertebra is precisely the same in both. But whilst the height of p 5 is greater its breadth is rather less, espe- cially immediately behind the acetabulum. The expanded horizontal thin plate of bone at the back part of the pelvis, between the diverging iliac bones, is well preserved in Mr. Cotton’s specimen. The breadth of this at the back part of the acetabula is five inches five lines ; the breadth at the hinder end of the pelvis is three inches nine lines. Of the two pelves which correspond in length and in most of the characters by which the one first described differs from p 4, we may regard the higher and narrower speci- men, p 5, as belonging to the male, and the lower and broader one, p 6, to the female. 1 PL. XXa. fig. 1. ''PROFESSOR OWEN ON THE GENUS DINORNIS. 259 Vertebra. (Plates XVIII. & XVIII a.) Of the five vertebree in the present collection, only one is of a size which surpasses in a marked degree that of the corresponding vertebra in a full-grown Ostrich ; but all present much stronger proportions, especially of the spinous process, which is un- usually robust. The largest vertebra’, v1, is a cervical one, probably from below the middle of the neck, anterior to those which are distinguished by a median inferior spine. The following are its dimensions as compared with the twelfth cervical of a full-sized Ostrich :— Dinornis. — Struthio. Ino Ein. « Tn? Ea Length, at the middle of the terminal articular surfaces . 2 9 22 Breadth; at the middie of the bodys... 2). . 425.4 eG 8 Height. of the, middlesef the: bady, ie.) fi. 6)' serel 8 Height from anterior base of spine to the lower part ‘a ihe anteriggasticular sipface «Geta ds ee SO be 20 Length: of the neural ately <1, ihe i eta ON i bh 46 Breadthel do. se vey Ai eh ee ee ee Oi i Every process and prominence of this specimen of the vertebree of the Dinornis is broken off, with the exception of the right posterior oblique process. The texture every- where presents large reticulate cancelli, which communicate with the outer surface by an orifice on each side the neural arch, behind the upper transverse process. The body of the vertebra is square-shaped, with a broad and flat, or slightly concave under surface: the anterior part of this surface is divided from the anterior articular surface by a transverse channel, that surface being raised to a higher level. This structure does not exist in the corresponding vertebre of the Ostrich: it is slightly indicated in those of the Apteryx. The spinal canal presents the usual infundibular expansion at both extremities : it is not larger at its middle contracted part than in the Ostrich. The remains of the base of the spinous process show this to have been almost square-shaped, and much thicker relatively as well as absolutely than in the Ostrich. Two other vertebrae belong to the base of the neck, and correspond with those few cervical vertebre at that part which, in most birds, have a median inferior process for the more advantageous origin of the great longus colli anticus muscle*. ‘These two ver- tebree must have come from the same or from closely contiguous parts of the neck ; but they present differences of configuration and proportion which are incompatible with the identity of the species of Dinornis to which they respectively belonged. Both manifest the generic massive proportions, the squareness of the body, the great 1 Pl. XVIII. figs. 1, 2, 3. 2 None of the cervical vertebre present this character in the Ostrich or Emeu, but we find it in the last cervical of the Rhea, and in the last three cervicals of the Apteryx and Bustard. VOL. III.—PART III. 2m ''260 PROFESSOR OWEN ON THE GENUS DINORNIS. breadth of its under surface, and the thick four-sided spinous process ; but one (v 2)’ is broader in proportion to its length than the other (v3). The more slender vertebra* has a thicker spinous process, which, at the same time, is more compressed from be- hind forwards: the cavity behind the spine is deeper and more angular, as is also the notch between the posterior oblique processes. The anterior articular processes are raised higher above the body in the more robust vertebra, v 2. The anterior articular surface of the vertebra v 3 has a much less vertical extent than in the thicker vertebra ; and the inferior spine (fh) is narrower, but of greater antero-posterior extent, and is situated nearer the posterior part of the body. Both these vertebre have the orifices at the sides of the neural arch which communicate with the interior loose cancellous structure. These are not present in the Apteryx, the corresponding vertebra of which in other respects more nearly resemble the present in general form and proportions’* than do those of the other existing Struthiomde. The vertebra v 4,* from New Zealand, transmitted to me by Dr. Richardson, the author of the ‘ Fauna Boreali-americana,’ belongs to the same species as the vertebra v 2. It is either the first or second of the dorsal series: the inferior transverse processes manifest part of the concavity for the articulation of the head of the rib, and there is a spinous process (h) from the under surface of the body of the vertebra, which, as in the anterior dorsal of the Apteryx, is less broad and flattened than in the anterior cervicals. Of the difference of the character of this vertebra, as compared with the correspond- ing one in the Ostrich, the figures’ give a better idea than can be conveyed by verbal description. The upper transverse processes are continued, as in the first and second dorsals of the Apteryx, from the anterior part of the whole side of the neural arch, not, as in the Ostrich, from near the summit; these processes also, as well as the spinous process, are considerably thicker and stronger than in the Ostrich. In regard to the spinous process, the Dinornis, in the squareness of that part, differs as much from the Apteryx, in which the dorsal spines are compressed laterally and extended antero- posteriorly, as from the Ostrich. The last vertebra, v 5, of the Dinornis in the present collection that remains to be noticed, is from the middle of the dorsal region : it belongs to a smaller species than the preceding ; most probably to the Din. didiformis. The body® is laterally compressed, and terminates below in a median carina, which has a concave outline: it has the characteristic shortness as compared with the breadth of the vertebree in this genus ; the anterior articular surface’ is more concave from side to side, and the posterior surface more convex in the same direction than in the corre- sponding vertebre of the Ostrich or Apteryx: both these surfaces have an unusual ver- tical diameter in proportion to their breadth. | Pl. XVIII. figs. 4, 5, 6 ® Pl. XVIIL. figs. 7, 8, 9. $ Pl. XVIII. fig. 10. + Pl. XVII a. figs. 1, 2, 3. > Pl. XVIlla. figs. 3 & 4. ° Pl. XVIllLa. figs. 6 & 9. 7 Pl. XVIII a. fig. 8. ''PROFESSOR OWEN ON THE GENUS DINORNIS. 261 The following are comparative admeasurements of this vertebra, and a corresponding one of a full-grown Ostrich :— Dinornis. Struthio. In. Lin. In. Lin. ESR. a us Re a ea 20 Depth of anterior anicutat SUING 6 a ee 0 7 Breadth, including costal articulations . . . : 1 6 20 From the lower margin of posterior articular pine to the upper one of posterior oblique process . . . . 2 1 i 8 The spinous process of this vertebra is strong and square-shaped’, and shows, like the preceding dorsal, that there was no blending together of the spines, nor any union by continuous splint-like ossifications, as in many birds, and especially in those that fly. The dorsal region in the skeleton of the Dinornis, by the intervals separating the spinous processes, must have resembled that in the large existing Struthionide, and have differed from the same part in the Apteryx, in which the dorsal spines are con- tiguous though not confluent ; but the Dinornis surpassed all known birds in the thick- ness and squareness of its upright spinous processes. Of the length of these processes none of the five vertebre afford an exact idea, all being more or less fractured. The spinal canal is proportionally more contracted than in the Ostrich, or even in the Apteryx, where it is rather smaller than usual. This character in the Dinornis indi- cates, of course, a more slender spinal chord, in which respect it betrays a closer ap- proach to the Reptilia. We may associate, with such a condition of the spinal marrow, less delicate perception, and less energetic muscular action; and the vertebra thus confirm the induction from the texture of the femur, that the Dinornis was a more sluggish or less active bird than the Ostrich. CONCLUSION. Physiological indications of the nature and proportions of the Anterior or Pectoral Members. Had the Dinornis wings? To this question I was led to give a negative reply after the examination of the first fragment of that bird’s bone which came into my hands’. It has appeared strange and almost incredible to some, that the cancellous texture of the shaft of a thigh-bone should give, to speak mathematically, the presence or absence of wings. But if the negative had been premature and unfounded, a guess rather than a demonstration, its fallacy might have been exposed by the very next bone of a Dinor- nis transmitted from New Zealand. A bird of flight has as many wings as legs ; it has two humeri as well as two femora, two radii as well as two tibiz, two ulnz as well as 1 Pl. XVII a. fig. 7. 2 Absence of the organs of flight is the essential character of a Struthious bird, more especially of one ‘‘ heavier than the Ostrich.” 2m 2 ''262 PROFESSOR OWEN ON THE GENUS DINORNIS. two fibulz ; the humerus and radius are usually, and the ulna is always, longer and larger than their analogues in the hind extremities ; then also there are the two distinct carpal bones, a metacarpus and characteristically modified phalanges. ‘The chances were thus greater that the next bone of an extremity discovered in the alluvium of New Zealand would have been one of the anterior members, had these been developed to serve as wings in the Dinornis. But what is the fact? Eighteen femora, eleven tibiz and six tarso-metatarsi, with two toe-phalanges, have been consecutively discovered, and not a trace of any part of the osseous framework of a wing: not a fragment of scapula, of hu- merus, or of the bones of the forearm or hand. The doctrine of chances thus adds its proof, were such required, to the inferences of physiological correlation, that the Dinornis had no wings. We may next inquire to what extent, short of the faculty of flight, the anterior or pectoral members were deve- loped in this extinct genus, with the same confidence in the laws of correlation as a guide to the determination of this question. The anterior members present very different degrees of arrested development in the different existing species of the Struthionide, and always retain, under even their most rudimental condition, the characteristic modifications of form and structure by which they are adapted to serve the office of flight in ordinary birds. In these, as is well known, the body is made specifically lighter, and in a direct ratio with the powers of flight, by a proportionate extension of the air-cells through the muscular and osseous systems. A much greater proportion of the skeleton is permeated by air in the Swallow than in the Quail. The Rhea and the Ostrich have the largest and most wing-like anterior members of all the Struthionide ; they use them to aid in their swift progression : throwing their body forwards beyond the centre of support afforded by the hind legs, they partly sustain it by the flapping of the curtailed wings, whilst the legs, to the extent to which they are thus relieved from the act of sustaining, are free to exert additional force in propelling the body: and it may be said of the Ostrich at full speed that half the body flies and half runs. Now we find that in these semivolant Struthionide the warm and expanded air of the respiratory cavities is freely admitted into the bones of the skull, the vertebra, the ribs, the sternum, the coracoids, the pelvis and the femora. In the Emeu and Cassowary, whose pectoral members are much reduced in size, use- less for anything like flight, and serving, so far as is known, only for some feeble actions of defence, the air is less freely admitted to the bones of the trunk, but still penetrates the femur. In the Apteryx the rudimental wings are so minute, that the fact of their retention of the typical structure requires careful dissection for its demonstration : and in this species we find the lungs confined to the thoracic-abdominal cavity, and not extended into any part of the skeleton. The Dinornis presents an intermediate condition between the Apteryx and the Emeu in regard to the extension of the air-cells, which penetrated the ''PROFESSOR OWEN ON THE GENUS DINORNIS. 263 vertebral column, as is shown by the pneumatic foramina in the vertebra, but were not continued into the femora. We may infer, therefore, from the known relations of the development of the air-cells to that of the anterior members in existing Struthionide, that these were more rudimentary in the Dinornis than in the Emeu, but not quite so minute in proportion to the body as in the Apteryx. The size of the bones on this inference, even in the Dinornis giganteus, must have been small enough to prevent any surprise at their not having yet been recovered ; especially when it is remembered that no part of the sternum nor any of the ribs, which doubtless surpassed the scapulz and humeri in size, appear hitherto to have been found. Stature of the different species of Dinornis. (Pl. XXX.) The height of the hind leg of the Dinornis giganteus in the ordinary standing posture, from the sole of the foot to the upper ridge of the trochanter, being given by the bones of the pelvic extremity in the present collection, the total altitude of the bird may be approximatively determined by the analogies of the existing Struthionide. In these the neck varies slightly in its relative length, being longest in the Ostrich and Emeu, in which it includes 18 or 19 vertebra, and shortest in the Cassowary and Apteryx, which have respectively 16 and 15 cervical vertebre ; but in all the species it is of sufficient length to enable them readily to pick up substances from the ground by a slight rotation or bending down of the trunk and pelvis upon the hip-joints. In estimating the height of the Dinornis giganteus by the standard of the Ostrich, I have taken the latter at eight feet four inches, which is the altitude given by the skeleton of one with a tibia two feet in length’. The distal end of the metatarsus being raised in the living bird one inch and a half from the ground, the tarso-metatarsal bone, tibia and femur, placed at the angles which they form with one another in the standing pos- ture, rise to the height of four feet four inches ; and from the level of the highest point of the femur to the top of the head with the neck erect is four feet. The longest tibia of the Dinornis giganteus, with its extremities entire, measures two feet eleven inches : this bone articulated with a femur of sixteen inches and a tarso-metatarsal bone of eighteen inches in length, at angles corresponding to those in the Ostrich, and with an allowance of three inches for the natural angle of the toes and the callous integuments beneath the distal joint of the metatarsal bone, makes the height of the hind leg to the highest point of the femur five feet six inches: from the level of this point to the top of the head, supported upon an erect neck of the same proportions as in the Ostrich, is five feet, making the total height of the Dinornis giganteus ten feet six inches. If the tarso-metatarsal bone of the Dinornis had borne the same proportion to the tibia as in the Ostrich, its height would have been nearly twelve feet, but the acquisition of 1 The tibize of mature specimens of the Ostrich in the Museum of the Royal College of Surgeons measure respectively 1 foot 8 inches, 1 foot 9} inches, and 1 foot 11 inches in length. The accurate and learned authors of the ‘Gardens and Menagerie of the Zoological Society’ state that the Ostrich “is generally from six to eight feet in height.’”,—Vol. ii. p. 51. ''264 PROFESSOR OWEN ON THE GENUS DINORNIS. the tarso-metatarsal belonging to the largest tibia fortunately prevented this error of exaggeration. But since the Cassowary and Apteryx, as compared with the Ostrich and Emeu, combine shorter tarso-metatarsals with their shorter necks, the Dinornis is much more likely to have resembled these birds than the Ostrich in the proportionate length of its neck, and we know that it resembled the Apteryx much more than the Ostrich in the robust proportions of the cervical vertebra. In the Apteryx, however, the peculiar length of the bill compensates for the relative shortness of the neck ; and until we have proof to the contrary, we must suppose the Dinornis to have had a bill of the ordinary proportions which it presents in the large existing Struthionide. I, therefore, conceive the Cassowary to offer the best term of comparison by which to calculate the height of the Dinornis. In the skeleton of a full-grown Cassowary' the tarso-metatarsal bone measures eleven inches in length: allowing an inch for the callous integuments beneath its distal articulation, the tibia and femur, articulated at the angles natural in the standing posture, rise to the height of two feet nine inches. From the level of the top of the trochanter to the top of the cranial crest is two feet three inches, and to the base of the crest two feet. We have no evidence that the Dinornis had that peculiar de- fence upon the head, and therefore, from the ground to the summit of the trochanter of | the Dinornis giganteus being five feet six inches*, from this level to the top of the head, according to the proportion of the uncrested Cassowary, would be four feet, making the total altitude nine feet six inches. Thus, if we take the average of the altitudes of the Dinornis giganteus, as given by the analogies of the existing Struthionide, we are compelled to restrict our ideas of its height in the ordinary upright posture to ten feet. The Dinornis struthoides*, with a femur of eleven inches, a tibia of twenty-two inches, and a tarso-metatarsus of twelve inches in length, must have stood, according to the analogies of the Cassowary, six feet nine inches in height ; according to those of the Ostrich, seven feet four inches: we may therefore regard its height to have not exceeded seven feet, or to have been about equal to that of a moderate-sized Ostrich, but of a more robust and stronger build. The fragment of the femur first described by me in 1839 belongs to this species. The Dinornis didiformis, with a tibia as long as that of the Cassowary, viz. sixteen inches, but with a femur of eight inches and a tarso-metatarsus of only seven inches in length’, would, by the analogy of the Cassowary, be a little under four feet in height, or of intermediate size between the Cassowary and the Dodo. The femur of nine inches in length, with similar proportions of the tibia and meta- tarsus, which latter would probably be relatively longer, gives the height of five feet to the species which, from its similarity in size to the Emeu, I have called Dinornis 1 PLL XXX. fig. 2. 2 Ib. fig. 5. 3 Ib. fig. 3. * Ib. fig. 1. The tarso-metatarsal bone of the Dinornis didiformis in Mr. Cotton’s collection measures seven inches ten lines in length (Pl. XX a. fig. 2.); it is in other respects identical in character with the analogous bones described in the text, and indicates a sexual superiority of size. ''PROFESSOR OWEN ON THE GENUS DINORNIS. 265 dromeoides. The tibia of the Dinornis ingens’ indicates that species to have attained the height of nine feet. Comparison of the bones of the feet of the Dinornis with the American Ornithichnites. In 1836 Prof. Hitchcock*® published his remarkable discovery of impressions in the New Red Sandstone of the valley of the river Connecticut, Massachusetts, which he con- ceived to be the foot-prints of birds, the largest belonging to a species with three toes, surpassing the Ostrich in size. The epoch of these impressions is as ancient as that of the Cheirotheria or Labyrinthodont footsteps in Europe, and more ancient than those of the oolites and lias, from which the remains of our most extraordinary extinct reptiles have been obtained: but no fossil bones of birds have been found associated with the Labyrinthodont and Thecodont reptiles, nor with those of the lias or oolites, the Ptero- dactyles of which were once mistaken for birds. The Wealden is the oldest formation in which true ornitholithes have hitherto been discovered. The ancient foot-prints of the Connecticut sandstones were for the most part supposed to be those of Gralle ; but the high geological antiquity of those sandstones, and the inferences which might be deduced from the low character of the air-breathing animal creation, as indicated by fossil bones, of the condition of the atmosphere during the deposition of the oolites, lias and new red sandstones, led me to express a doubt in my report on British Fossil Reptiles whether foot-prints alone were adequate to support the inference that the animals that impressed them actually possessed the highly-developed respiratory organization of a bird of flight’. One could hardly in fact venture to reconstruct in imagination the stupendous bird which, on Dr. Hitchcock’s hypothesis, must have left the impressions called Ornithichnites giganteus ; for, before 1843, the only described relic of the extinct New Zealand bird did not warrant the supposition of a species larger than the Ostrich’. The species of Dinornis, in fact, to which that relic belonged, we now know not to have exceeded seven feet in height, which is the average stature of the Ostrich. But the bones of the Dinornis giganteus subsequently acquired demonstrate the existence, at a comparatively recent period, of a bird whose tridactyle foot-prints, as will be pre- sently shown, surpassed the Ornithichnites giganteus of Prof. Hitchcock. The length of this foot-print from its hind part to the extremity of the impression of the claw of the middle toe is sixteen inches; the breadth of the hind part is four inches six lines. The toes were broad and thick, and we may plainly discern that the bird supported itself, like the Ostrich, upon the under surface of the toes, from their extremities to the cushion beneath the distal end of the proximal phalanges ; and that in making the impression, the foot did not quite sink as far as the end of the metatarsal bone. 1 Pl. XXX. fig. 4. 2 American Journal of Science and Arts, vol. xxix. No. 2. 3 Report on British Fossil Reptiles, Part II., Trans. British Association, 1841, p. 208. + Zoological Proceedings, November 1839, p. 170. ''266 PROFESSOR OWEN ON THE GENUS DINORNIS. The length of a corresponding impression of the foot of the Ostrich is eight inches ; the breadth of the posterior part of the impression three inches ; the breadth of the distal end of the tarso-metatarsal bone two inches and a half. According to these pro- portions, the breadth of the distal end of the tarso-metatarsal bone of the tridactyle bird that impressed the Ornithichnites giganteus must have been three inches nine lines ; but the breadth of the distal end of the tarso-metatarsus of the Dinornis giganteus is five inches. According, therefore, to the proportions of the Ornithichnites giganteus, the breadth of the hind part of the foot-print of the Dinornis giganteus must have been six inches, and its length twenty-one inches and a half. The genus Dinornis was characterized by a relatively broader foot than the Ostrich, as we know by the tarso-metatarsal bones; and this bone in the Dinornis struthoides, the third species in point of size, indicates that its bulky body was supported by feet calculated to leave impressions nearly as large as those of the Ornithichnites giganteus. That the toes were as long in proportion to the breadth of the metatarsal bone as in the Ornithichnites, is shown by the two phalanges transmitted by Mr. Williams, the description of which I have reserved for this place. The largest of these phalanges is 34 inches long and 17 inch broad across the proximal joint. This does not present the median vertical ridge which the corresponding groove in the articular surface of the metatarsal indicates the proximal phalanx to possess, and I regard it, therefore, to be a second phalanx, which, as in the middle toe of the Ostrich, would then differ from the first phalanx in the equable concavity of the proximal arti- cular surface. In the second or outer toe of the Ostrich the median eminence is wanting on the proximal end of the first phalanx, but the want of symmetry in that bone shows that it cannot be the analogue of the phalanx of the Dinornis in question, which is almost quite symmetrical. From this character it may be referred to the middle toe: compared with the second phalanx of that toe in a full-grown Ostrich it is relatively longer, less depressed or flattened, the depth of the bone being equal to its breadth ex- cept at the distal articulation, which nevertheless is much less expanded and depressed than in the Ostrich. In this bird the length of the second phalanx of the middle toe is 24 inches, the breadth of the distal end is 14 inch, and its depth at the middle of the bone 8 lines. In the phalanx of the Dinornis the breadth of the distal end is le inch, its depth at the middle 10 lines. The size of the phalanx of the Dinornis, re- garded as the second of the middle toe, agrees well with that of the tarso-metatarsal of the Dinornis struthoides. The length of the second phalanx in the Ornithichnites gigan- teus is indicated by the articular eminences in the cast of that impression, and it is a little shorter than the phalanx of the Dinornis above described. The smaller of the two phalanges has an unsymmetrical figure, and its proximal arti- cular concavity is continuous with an oblique notch which divides the lower border into two tuberosities. This structure is slightly indicated at the corresponding part of the proximal phalanx of the outer toe in the Ostrich, and in the Bustard is as strongly ''PROFESSOR OWEN ON THE GENUS DINORNIS. 267 marked in the proximal phalanx of both the outer and inner of the three toes as in the phalanx of the Dinornis. This phalanx measures one inch ten lines in length, one inch two lines across the proximal end, and ten lines across the distal end: the articular surface here is impressed by a vertical groove, as in the proximal phalanges of the outer and inner toes in the Bustard, and it agrees in its general figure with that of the outer toe of the left foot, but is much thicker in proportion to its length. The proximal articulation matches in size with, but is not adapted by its configuration to, the outer trochlea of the trifid metatarsal of the Dinornis didiformis. The foot-print of this spe- cies was probably about the size of the Ormthichnites tuberosus of Prof. Hitchcock. From the foregoing comparison of the bones of the feet in the different species of Dinornis with the impressions left by the ancient extinct birds of the American conti- nent, it must not, however, be concluded that these were species of Dinornis. Agree- ment in the size of the foot and number of the toes does not constitute specific or even generic identity in Ornithology, as the living Emeu, Rhea and Cassowary testify ; and though we may admit that the discovery of tridactyle terrestrial birds of a size more gigantic even than that indicated by the Ornithichnites giganteus and Ornithichnites ingens tends greatly to remove the scepticism with which such evidences of the extinct animals of the Triassic period had been previously received, yet the recognized suc- cession of varying vertebrated forms in the interval between that period and the present forbids the supposition that the same species or genus of birds could have maintained its existence throughout the several great changes which the earth’s surface has under- gone during that vast lapse of time. We see, in fact, how diversified are the few existing forms of Struthionide: almost every species now represents a distinct genus. We know that this order has suffered greater diminution within the time of man than any other in the class of Birds, perhaps than any other in the whole animal kingdom. What, then, may not have been the extent and variety of the wingless terrestrial birds in times anterior to man’s dominion over the earth! , - Already the heretofore recorded number of the Struthionide 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 ex- tirpation 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 di- spersion of the human race, to the attacks of man. We may, therefore, reasonably anticipate that other evidences await the researches of the naturalist, which will demon- strate a further extent of the Struthious order of Birds anterior to the commencement of the present active cause of their extinction. And since the texture of the bones of the former gigantic tridactyle Struthionide of New Zealand proves that they resembled the Apteryx, in the comparatively low 1 The Maoris or Aborigines of New Zealand call the Dinornis ‘ Moa’ or ‘ Movie.’ VOL. III.—PART III. 2N ''268 PROFESSOR OWEN ON THE GENUS DINORNIS. and reptile-like condition of the respiratory apparatus, we are thereby further jus- tified in admitting the evidence of the co-existence of similar apterous and low orga- nized birds with the cold-blooded and slower-breathing Ovipara, which swarmed in such plenitude of development and diversity of forms during what has been termed the ‘Age of Reptiles.’ The remarkable geographical distribution of the birds of the Struthious order, which have no power of transporting themselves to distant isles or continents, either through the air or the ocean!, irresistibly leads us to speculate on the cause of that distribution, and its connexion with. the former extent and importance of the wingless terrestrial birds. Hereupon it may first be remarked that those species, now in existence, which have the least restricted powers of locomotion, enjoy the most extensive range for their exercise. The Ostrich is spread over nearly the whole of Africa, from the Cape to the deserts of Arabia; beyond which the species is unknown. The Rhea ranges over a great part of the southern extremity of the Western hemisphere. ‘To the Emeu has been assigned the vast mainland of Australia. The heavier Cassowary is limited to a few of the islands of the Indian Archipelago. The Dodo appears to have been confined to the Mauritius and the small adjoining Isle of Rodriguez. ‘The Apteryx still lingers in New Zealand, where alone any specimens of that most anomalous species of the Struthious order have been discovered. New Zealand was, also, at one period, the seat of a seventh genus of Struthionide ; and it is worthy of remark that the Fauna of no other island, nor of any of the great continents, has yet furnished an analogous example of two distinct genera of that group of birds. Moreover the most gigantic as well as the most diminutive species of the wingless group—always to Ornithologists most remarkable for the great size of its spe- cies—formerly occupied their place amid the fern-thickets and turbaries of New Zealand. And, again, the number of the species of Struthionide@ in this island equalled that in all the rest of the world, as registered in the catalogues of Ornithology. Now, since all the larger existing Struthious birds derive their subsistence from the vegetable kingdom, we may hope to receive from the botanist an elucidation of the circumstances which favoured the existence of so many large birds of this order in the remote and restricted locality where alone their remains have hitherto been found. It seems, at least, most natural to suppose that some peculiarity in the vegetation of New — Zealand adapted that island to be the seat of apterous tridactyle birds, so unusually numerous in species and some of them of so stupendous a size. The predominance of plants of the Fern-tribe, and the nutritious qualities of the roots of the species most common in New Zealand, are the characteristics of its Flora which ! The Rhea and Emeu have been seen to take water for the purpose of crossing rivers and narrow channels of the sea; but almost the entire body sinks below the surface, and their progress is slow, as might be anticipated _ from the absence of the swimming-webs in their feet. See Darwin, ‘ Voyage of the Beagle,’ vol. iii. p. 105. ''PROFESSOR OWEN ON THE GENUS DINORNIS. 269 appear to have been the conditions of the former peculiarities of the Fauna of this island. Some at least of the characters of the skeleton of the Dinornis may well have related to rhizophagous habits. The unusual strength of the neck indicates the appli- cation of the beak to a more laborious task than the mere plucking of seeds, fruits, or herbage. The present small Apteryx of New Zealand has a relatively stronger neck than any of the existing Struthionide, in relation to the needful power of perforating the earth for the worms and insects which constitute its food. Such small objects can- not be supposed to have afforded sustenance to the gigantic Dinornithes: but the still more robust proportions of their cervical vertebre, and especially of their spinous pro- cesses,—so striking when contrasted with the corresponding vertebre of the Ostrich or . Emeu,—may well have been the foundation of those forces by which the beak was asso- ciated with the feet in the labour of dislodging the farinaceous roots of the ferns that grow in characteristic abundance over the soil of New Zealand’. The great strength of the leg, and especially of the metatarsal segment, which is short- ened, as in the burrowing Apteryx, almost to the gallinaceous proportions, must have had reference, especially in the less gigantic species, to something more than sustaining and transporting the superincumbent weight of the body, and this additional function is indicated by both the analogy of the Apteryx and the Rasorial birds to be the scratch- ing up the soil. Thus far, at least, the positive facts justify the attempt to restore, and, as it were, to present a living portrait of the long-lost Dinornis ; and, without giving the rein to a too exuberant fancy, we may take a retrospective glance at the scene of a fair island, offering, by the will of a bountiful Providence, a well-spread table to a race of animated beings peculiarly adapted to enjoy it ; and we may recall the time when the several spe- cies of Dinornis ranged the lords of its soil—the highest living forms upon that part of the earth. No terrestrial Mammal was there to contest this sovranty with the feathered bipeds before the arrival of man’. Without laying undue stress on the native tradition of the gigantic Eagle or ‘ Movie,’ cited by Mr. Rule’, or on that of the great creature of the cavern, called ‘ Moa,’ which first attracted the attention of Mr. Williams to the remains of the Dinornis; and ad- mitting with the cautious scepticism due to second-hand testimony, the tale of the still- existing nocturnal gigantic bird which scared the whaling seamen on the hill at Cloudy ' « New Zealand is favoured by one great natural advantage, namely, that the inhabitants can never perish from famine. The whole country abounds with fern; and the roots of this plant, if not very palatable, yet contain much nutriment.’”’ Voyage of the Adventure and Beagle, vol. iii. ‘ Darwin,’ p. 504. ¢ Mr. Darwin says, “‘ It is a most remarkable fact that so large an island, extending over more than 700 miles in latitude, and in many parts 90 miles broad, with varied stations, a fine climate, and land of all heights from 14,000 feet downwards, with the exception of a small rat, should not possess one indigenous mammal.”—Loc. eepeolk. | 3 Polytechnic Journal, July 1843. ou 2 ''270 PROFESSOR OWEN ON THE GENUS DINORNIS. Bay,—the evidence of the chemical condition of the bones themselves!, and their allu- vial bed, favour the hypothesis of their comparatively recent date. It is not altogether improbable that the species of Dinornis were in existence when the Polynesian colony first set foot on the island; and, if so, such bulky and probably stupid birds, at first without the instinct and always without adequate means of escape and defence, would soon fall a prey to the progenitors of the present Maoris. In the absence of any other large wild animals, the whole art and practice of the chase must have been concentrated on these unhappy cursorial birds*. The gigantic Dinornis, we may readily suppose, would be the first to be exterminated: the strength of its kick would less avail, than its great bulk would prejudice its safety by making its con- cealment difficult ; at all events, the most recent-looking bones are those of the smaller species. The closely allied, but comparatively diminutive Apteryx still survives by vir- tue of its nocturnal habits and subterraneous hiding-place, but in fearfully diminished and rapidly diminishing numbers. When the source of animal food from terrestrial species was reduced by the total extirpation of the genus Dinornis to this low point, then may have arisen those cannibal practices which, until lately, formed the oppro- brium of a race of men in all other respects much superior to the Papuan Aborigines of the neighbouring continent of Australia, and very little inferior to the Polynesian natives of the most favoured islands of the Pacific. 1 T have been favoured with the following analyses by my friend Thomas Taylor, Esq., author of the Cata- logue of the Calculi and other Animal Concretions in the Museum of the Royal College of Surgeons :— ** Recent Tibia of Ostrich. Fossil Femur of Dinornis didiformis. ATMMal Matter ec Oe. 26°51 Animal matter 20.80. Se ee, 25°99 Pabephore crime 6 6.00 ls wee. 65°69 Phosphate of lime with phosphate of magnesia 66°19 Phosplate Of magnesia... ev. eee 0°95 Carbondievor ime s.100 0... ss cee ees 4°51 Carbonate of lime: tis .c06s oe ok a ey, 6:22 Peroxide. of iO | ails ice, ee a 2°81 Sulphate and carbonate of soda, with a Wa PAVUPAINE Sh ye oe a 0:22 Ofmuriatesss.. err 66 Sulphate, carbonate, and muriate of soda.... 0°32 Sulphate of lime, a trace. Fluorine, a trace. 99°49 Sulphate of lime, a trace. Fluorine, a very distinct trace. 100°04”’ The subjoined comparative analysis, kindly undertaken by Dr. G. Day, also shows the large proportion of animal matter in the bones of the Dinornis :— ** Recent femur of Ostrich. Animal matter i:s45;-..... 84°86 Inormanic matter |, .2.....-. 65°65 100-00 Femur of Dinornis struthoides. eens Gusiepel site oe eerie Gees 37°86 Ope iss elleeo eects cleo ccc e 62°94 100:00” The superabundance of animal matter in the bone of the extinct bird depends upon its being a marrow-bone, whilst that of the Ostrich contains air. * As the Maoris prize the skin and feathers of the Apteryx for the manufacture of ornamental robes, it might be worth inquiry whether any of the natives preserve remains of their ancestors’ dresses composed of feathers of unknown and larger species of birds. Such relics of a Dinornis might in this way be recovered. ''PROFESSOR OWEN ON THE GENUS DINORNIS. 271 ADDENDUM. J. R. Gowen, Esq., a Director of the New Zealand Company, has obligingly forwarded to me the subjoined indication of a further discovery of the bones of the Dinornis, from a new locality in New Zealand :— “ Extract of a letter from Colonel William Wakefield to J. R. Gowen, Esq., dated Wellington, 19th September, 1843. ‘ 2S i . PROFESSOR OWEN ON THE GENUS DINORNIS. 273 . Back or upper view of the same pelvis: b, the confluent spines of the poste- rior sacral vertebree. . Side view of pelvis of Dinornis didiformis. . Fragment of the pelvis of Dinornis struthoides. f. The acetabulum. PLATE XX a. Pelvis of male Dinornis didiformis, natural size. . Tarso-metatarsal bone of ditto, ditto. PLATE XXI. Femora, natural size. Back view of femur of Dinornis ingens. . Lower or distal end of femur of Dinornis ingens. Back view of femur of Dinornis struthoides. Distal end of femur of Dinornis dromeéoides. PLATE XXII. Femur of Dinornis dromeéoides, natural size. . Front view. Back view: d, ridge above the internal condyle ; e, fossa above the external condyle. PLATE XXIII. Side view of femur of a young Dinornis struthoides. . Side view of femur of Dinornis dromeoides. PLATE XXIV. Femur of Dinornis didiformis, natural size. . Front view. NS = . Back view: d, tuberosity above the internal condyle ; e, fossa above the ex- ternal condyle. . Distal end. PLATE XXV. Front views of tibiz, natural size. . Tibia of Dinornis ingens. ''274 Fig. Fig. or yb oe 6. Aer bh WwW DH nw — PROFESSOR OWEN ON THE GENUS DINORNIS. Distal trochlea of ditto. . Tibia of Dinornis didiformis. . Distal trochlea of ditto. . Tibia of Dinornis otidiformis. . Distal trochlea of ditto. PLATE XXVI. Back views of tibiz, natural size. Tibia of Dinornis ingens. Proximal end of ditto. . Tibia of Dinornis didiformis. Proximal end of ditto. Tibia of Dinornis otidiformis. Proximal end of ditto. The following letters indicate the corresponding parts in each figure :— Fig. Cb = Oo Ov Fic. 1. 3. a. Concavity anterior to the articular condyles. b. Process for the attachment of the rotular ligament. c. Anterior crista at the proximal end. d. Ridge for the attachment of the fibula. e. Internal condyle or prominence of the distal trochlea. PLATE XXVII. Tarso-metatarsal bones, natural size’. . Front view of the tarso-metatarsal of Dinornis giganteus. . Front view of the tarso-metatarsal of Dinornis struthoides. . Front view of the tarso-metatarsal of a female (?) Dinornis didiformis. . Back view of ditto. . Upper or proximal end of ditto: a, the outer concavity ; b, the inner one. . Lower or distal end of ditto. PLATE XXVIII. Tarso-metatarsal bones, half the natural size. Front view of the metatarsus of a young Ostrich (Struthio Camelus). . Proximal ends of the three metatarsal bones, which are still separate at this part. Front view of the metatarsus of a young Dinornis giganteus. ' The mirror was not used in drawing these bones ; they appear, therefore, to belong to the right leg. ''Fig. 1. PROFESSOR OWEN ON THE GENUS DINORNIS. 275 . Proximal ends of the three metatarsal bones not united together at this part. . Side view of the tarso-metatarsal bone of the Dinornis struthoides. . Side view of the tarso-metatarsal bone of the Dinornis didiformis. . A transverse section of ditto, at the part corresponding with the three separate metatarsals of the young Dinornis giganteus, fig. 4. PLATE XXIX. Internal structure of certain bones. Section of the femur of Dinornis didiformis, natural size: h, an accidental de- pression at the back of the cervix, not leading into the interior medullary cavity ; 2, the depression in the popliteal space, without any opening into the interior of the bone. . Section of the femur of an Ostrich, half the natural size: h, the orifice and oblique canal conducting into the interior pneumatic cavity of the bone ; 2, the outlet of the same cavity in the popliteal space. Section of the femur of the Apteryz, in which no air is admitted into the me- dullary cavity ; natural size. . Section of the tibia of the Apteryx, natural size. Section of the tarso-metatarsal bone of Dinornis didiformis, natural size: h. the obliterated line of union of the tarsal epiphysis. PLATE XXX. Restoration of the Dinornis giganteus, and scale of altitude of that and other species, Piet. oo wo according to the standard of the Cassowary. The three principal bones—femur, tibia, and tarso-metatarsus—of the hind extremity of Dinornis didiformis. Skeleton of Casuarius galeatus (Pander & D’ Alton). The femur, tibia, and tarso-metatarsus of Dinornis struthoides. The femur, tibia, and tarso-metatarsus of the Dinornis ingens. . The pelvis, femur, tibia, tarso-metatarsus, and restored outline of the Dinor- nis giganteus. VOL. III, —PART III. 20 '' '' Soins Soot Soc Ce. 3, PMY 5 eeee eeee Day bHughe, lath to the ucen J Eraleben Le 4 / eece '' '' J Fraleben, Zine. 15 0:9 & De ee > Ri ore » . ; Day bllag he bth to the Qucen, t ¢ , '' '' Spano Leol Ae Vt EP 14, fr. 253. 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S XQ Xs = Ne ; SS } yQ cy < < — y Nat. Sige. '' '' 3/29 pid, Vol Py ee : wy , ith! to the Juten Di zy & Hagh leben, Zine ETH JL 4 CENYK. Gy 4, Coe Me ey, ) ihe teh We bs @ CIA ae phon z '' '' ewe wee me ee eee eee _ é. ip 0. Scale af Eng: feet. J Erdebar, Zinc. Qo be '' '' ''RETURN TO DESK FROM WHICH BORROWED EARTH SCIENCES Lipase. This book is d eral Library versity of California Berkeley ''wi A CO34bLb7237 ''X. On Dinornis, an extinct Genus of tridactyle Struthious Birds, with descripiian. of of . I ” o ~, . « ° « ee portions af the Skeleton of five Snectes which’ formerly existed in New Zealand. By Professor Owen, M.D., FLR.S., F.Z.S8,, Se. &c. page 242 EB wes AT ''