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Chapter II: Part 2

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The happy perception, by the Danish Professor J. Reinhardt, in 1843[20], of the resemblance of the beak of the Dodo to that of the tropical Doves, generically separated by Cuvier under the name _Vinago_, on account of their proportionately larger, more strongly arched, and compressed beak than in other Pigeons, and the still closer resemblance, in miniature, of the beak of the Samoan Dove to that of the great Mauritian bird, which led Titian Peale to give to the former the generic name _Didunculus_, directed the ornithologist and ornithotomist to the family in which the most instructive comparisons might be made; and the results of these, so far as relates to the head and foot and the bones of those parts, published by the authors of the above-cited work (p. 4), left little doubt of the “striking affinity which exists between this extinct bird and the Pigeons”[21].

Whatever doubt, indeed, may have lingered in the minds of naturalists as to this affinity will probably be finally set at rest by the results of the comparison of the large proportion of the skeleton of the _Didus ineptus_ which has at length been transmitted from the island of Mauritius to London, under the following circumstances.

In 1863, I was favoured by Miss A. Burdett Coutts with an introduction to the Bishop of Mauritius, then in this country, and I endeavoured to interest his lordship in aiding or promoting the acquisition, by the British Museum, of the zoological rarities of Madagascar, and especially of any remains of the Dodo which might be discovered in the island of Mauritius, to which his lordship was about to return.

How speedily and successfully the Bishop has fulfilled my latter desire will be shown by the following letter, with which I was favoured in November, 1865.

“St. James, Port Louis,
“October 7, 1865.

“+My dear Sir+,—when I had the pleasure of conversing with you for
a short time in London two years ago, I promised to acquaint you
with any facts or discoveries which might come to my knowledge,
likely to interest you in connexion with Madagascar. I have not
anything as yet to communicate definitely respecting that island in
the way of natural history, but I have strong reasons to believe
that a discovery has been made here recently which will gratify you
very much. Mr. George Clark, who has for many years devoted himself
to the work of teaching in this island with great success, is an
ardent student of natural history, and has explored many parts of
the island in search of information on the subject. From careful
observation he was led to conclude that no remains of the Dodo were
likely to be found in any of our watercourses, because of their
steep descent and the immense rush of water which sweeps down them
at times. But he had also frequently expressed his opinion that in
certain marshes, with high banks of sand between them and the sea,
such remains would probably be found. In one of these places he has
found several of the bones of the Dodo (as he believes), and is now
forwarding them home for your inspection[22].

“At his request, I write these lines to ask for your kind care of
his interests in securing any reward which may accrue to him. It
would be a great pleasure to me to find that his discovery was
really important, and likely to be useful to himself; for he
has pursued these and similar investigations with an amount of
intelligence, skill, and diligence, in his vacation-times (by no
means extensive), which deserves much credit and encouragement.

“The book which you kindly sent me on the Aye-Aye has been read
by many, and especially by medical men, with much interest. I
entrusted the other copy to Mr. John Douglas for the Society here.

“I remain, my dear Sir,
“Your very faithful Servant,
(Signed) “+Vincent N. Mauritius+.”
“_Professor Owen._”

This letter was accompanied with the following “Statement” by Mr. George Clark, Master of the Government School at Mahébourg, Island of Mauritius:—

“On the estate called ‘Plaisance,’ about three miles from
Mahébourg, in the island of Mauritius, there is a ravine of no
great depth or steepness, which, apparently, once conveyed to the
sea the drainings of a considerable extent of circumjacent land,
but which has been stopped to seaward, most likely for ages, by
an accumulation of sand extending all along the shore. The outlet
from this ravine having been thus impeded, a sort of bog has
been formed, called ‘La Mare aux Songes,’ in which is a deposit
of alluvium, varying in depth, on account of the inequalities of
the bottom, which is formed of large masses of basalt, from three
to ten or twelve feet. The proprietor of the estate a few weeks
ago conceived the idea of employing this alluvium as manure; and
shortly after, the men began digging in it; when they had got to
a depth of three or four feet they found numerous bones of large
tortoises, among which were a carapace and a plastron pretty nearly
entire, as also several crania.

“When I heard of this, it immediately struck me that the spot was
one of the most likely possible to contain bones of the Dodo, and I
gave directions to the men working there to look out for any bones
they might find. Nothing, however, was turned up but a fragment of
what I supposed to be the humerus of a large bird. This encouraged
me to look further; and my search was rewarded by the discovery of
several tibiæ, more or less perfect, two tarsi, one nearly perfect
pelvis, and fragments of three others.

“These were found imbedded in a black vegetable mould, the
lighter-coloured specimens being near the springs. My reasons
for believing these to be remains of the Dodo are:—the certainty
that that bird once existed in Mauritius; the similarity of these
bones to what the representations of the Dodo which I have seen
would lead one to expect, particularly the breadth of the pelvis,
the stoutness of the tibiæ and tarsi, and the shortness of the
latter; the favourable nature of the spot in which they were found
for the haunts of such birds when living—a sheltered hollow with
two springs in it; the non-existence, actual or traditional, in
Mauritius of any bird to which bones such as these could have
belonged; the indubitable antiquity of these bones, proved by the
deposit of alluvium which covered them.

“During nearly thirty years that I have inhabited this colony, I
have made frequent inquiries of old people as to the finding of
the bones of large birds, and have offered liberal rewards for
such; and I have consulted with the late Dr. Ayres as to the spots
most likely to contain them. We agreed that the floods which sweep
the hill-sides and the ravines in the rainy season would be most
likely to carry any remains into the sea; and this would doubtless
have been the case here, but for the stoppage occasioned by the
sand-down.

(Signed) “+George Clark.+ 1865.”

The above “Statement” was authenticated by the following testimony:—

“Having visited the place with Mr. Clark, I can vouch for the truth
of the facts herein mentioned.
(Signed) “+William Thomas Banks+,
“Civil Chaplain, Mauritius.”

“The Rev. W. T. Banks, Civil Chaplain at Mahébourg, in this
diocese, and Mr. George Clark, Master of the Government School at
Mahébourg, are well known to me, and deserving implicit credit for
their statements as to matters of fact.

(Signed) “+Vincent N. Mauritius.+ Oct. 6, 1865.”

§ 2. _Description of the Skeleton._ (Plate III.)

The bones of the Dodo (_Didus ineptus_, Linn.) discovered by Mr. Clark, under the above circumstances, which have reached me up to the present date (December 20th, 1865) are the following:—

Name. Number of bones or parts.

Cranium and lower jaw, in parts 14
Vertebræ and pelvis 30
Ribs 22
Sternum 2
Scapular arch, in parts 7
Humerus, ulna, radius 6
Femora 5
Tibiæ 6
Fibulæ 4
Metatarsals 4
———
Total number of parts of skeleton of the Dodo 100
===

The known characters of the skull and metatarsus of the _Didus ineptus_ served to identify those bones as belonging to that species: the agreement in relative size, colour, condition, and locality left no room for hesitation in referring the other bones in the above list to the same species[23]. They belong, however, to four or five individuals varying somewhat in size. With the bones of the Dodo were the end of the lower jaw of a broad-billed Parrot, two bones (radius) of a small Mammal, and part of the skull of a large Tortoise[24].

To the description of the Dodo’s bones I now proceed.

_Vertebræ._ (Plates III., IV., V., VIII., XI.)

The dorsal vertebræ are chiefly represented, in this series of bones, by three which are anchylosed together by their bodies and neural arches (Pl. V. figs. 1–5): the posterior articular surface of the body of the last of these vertebræ (ib., fig. 4, _c_) is subquadrate, longer in the vertical than the transverse direction, concave vertically, convex transversely, almost fitting, but being rather too small for, the anterior articular surface of the body of the first of the sacral series (Pl. VII. fig. 1, _c_). The difference is such as to indicate that only one dorsal vertebra may have intervened; and I conclude that the last of the three coalesced vertebræ is the penultimate dorsal. The anterior articular surface of the foremost of the three (Pl. IV. fig. 1, _c_) is 11 lines in transverse, and 4 to 5 lines in vertical diameter: it is concave transversely for the middle three-fifths, and convex transversely at the two outer fifths of its extent: it is more or less convex vertically throughout its extent. The bodies of these vertebræ are compressed and wedged-shaped, slightly expanded at their coalesced ends, produced below into subquadrate hypapophyses in the first and second (Pl. V. fig. 1, _hy_); while this process is restricted to the fore part (ib. _hy_ 3), or may be represented only by a slight anterior production of the lower edge of the wedge, in the third (ib. fig. 5, _hy_ 3).

The hypapophysis of the first of the three expands at its termination (Pl. IV. fig. 1, _hy_), with the hinder angle bent back to coalesce with the front one of the next hypapophysis, which is somewhat longer, and bent forward with a similar terminal expansion: a full elliptical space is intercepted by this terminal confluence of these hypapophyses (Pl. V. figs. 1 & 5, _hy_). Each vertebra shows an elliptical articular cavity (ib. figs. 1 & 5, _p_, _p_ 3) for the head of the rib, near to the anterior articular surface; the long axis of this costal surface is directed from above obliquely downward and forward. The surface of the rib’s tubercle cuts obliquely the lower part of the free end of the diapophysis (Pl. IV. fig. 1, _d_).

The neural arch circumscribes a canal the anterior outlet of which (ib. fig. 1, _n_) is oval with the small end downward, 5 lines in vertical, and 3½ in transverse diameter: the sides of the neural canal slightly project inward above the lower third: the posterior outlet (Pl. V. fig. 4, _n_) is more regularly elliptical in form, and rather narrower in proportion to its vertical diameter. The neurapophysis sends off from the outer and fore part of its base a stout process, which expands and divides into zygapophyses (Pl. IV. fig. 1, _z_) and diapophyses (ib. _d_); the articular surface of the former is of a full oval shape, flat, looking obliquely upward and inward; the diapophyses extend outward and a little backward: the articular surface for the tubercle of the rib is transversely elliptical and nearly flat. The hinder part of the neurapophysis expands into the postzygapophyses: these have coalesced with the præzygapophyses in the succeeding vertebra (Pl. V. fig. 2, _z_), as has happened also between this and the third vertebra. In the last of the three vertebræ the postzygapophyses are entire (ib. _z_ 3), and show very slightly concave, oval articular surfaces, looking obliquely downward and outward (ib. fig. 4, _z_). The conjugational foramina, continuously surrounded by bone, are a full ellipse, and large, the anterior one (ib. figs. 1 & 5, _f_) being 5½ lines in vertical diameter; the second (ib. _f′_) is somewhat less: these foramina are also rather larger in one of the specimens than in the other. The length of the three coalesced dorsals is the same in both, viz. 2 inches 3 lines. The neural spines have run together into a continuous ridge in fig. 1, _ns_; in fig. 5 the summit is broken off in both, leaving only the anterior angle of the foremost entire; in both this inclines forward; the hinder border of the third vertebra (fig. 1, _ns_) has the same vertical parallel as the back part of the centrum. The anterior margin of the base of the spine shows a rough surface for the attachment of ligament (Pl. IV. fig. 1, _ns_). A small foramen behind the base of each of the coalesced zygapophyses (Pl. V. fig. 2, _z_ _z_) leads to a canal descending to the neural one, and indicates superiorly the limits of the otherwise continuously ossified neural arches.

In the series of detached vertebræ, one (Pl. V. figs. 6 & 7) indicates by its neural spine and hypapophysis a position at the base of the neck. The centrum is barely an inch in length; its anterior surface (ib. fig. 7, _c_) is narrow vertically, broad transversely; both fore and hind surfaces indicate freedom and extent of flexure. The hypapophysis has a broad, bituberculate base (ib. _hy_), but is limited in fore and aft extent to the middle third of the under surface of the centrum: its length is shown in fig. 6, _hy_. The parapophysis (fig. 7, _p_) is slender, and expands at both attachments, with an indication of a terminal surface. The diapophysis (_d_) has a larger costal surface: it sends forward a convex ridge midway between the di- and zygapophysis (_z_). The neural canal (fig. 7, _n_) has wider and more fully elliptical outlets than the hinder dorsal vertebræ, in relation to the greater extent of motion at the fore part of the series. I conclude that a free pleurapophysis (_pl_) existed, indicating the present to be the first of the dorsal series, as shown in Pl. III. The neural spine is short, broad, obtusely pointed, with a vertically oblong syndesmotic surface (fig. 7) before and behind. Each postzygapophysis (fig. 6, _z′_) supports an anapophysial tubercle (_a_).

A cervical vertebra from a position just in advance of the above has lost the neural spine, but retains the hypapophysis. This process (ib. figs. 8 & 9, _hy_) is compressed and directed obliquely downward and forward for an extent of 6 lines; the extremity is rounded: the length of the centrum of this vertebra is 1 inch 3 lines; the anterior articular surface is longest transversely, and concave in that direction, convex vertically; the proportions and curvatures are transposed in the posterior surface (fig. 9, _c_). The parapophysis (ib. _p_) is continued from the anterior border of the centrum to the middle; it is a depressed plate, confluent with the rib (ib. _d_). The diapophysis forms a short, obtuse projection above its anchylosis with the rib (ib. _pl_): this projects backward 7 lines in length, terminating obtusely, and circumscribing a vertebrarterial foramen (ib. _v_) of a full elliptic shape, 5½ lines in long diameter. The surfaces of the præzygapophyses (_z_) are larger, and look more upward and less inward, than in the preceding and the dorsal vertebræ: they are very slightly concave. Those of the postzygapophyses (fig. 8, _z′_), with a downward and slightly outward aspect, are in a similar degree convex. The neural canal, as usual in the cervical series, expands at its outlets, most so posteriorly (fig. 9, _n_); the middle of the upper surface of the neural arch is impressed by an elliptical, rough, ligamentous surface, which slightly rising in the middle is the sole indication of a neural spine. The upper surface of each postzygapophysis developes a tuberous anapophysis (figs. 8 & 9, _a_).

The three cervicals that succeed the axis show progressively sinking neural spines, which subside in the six following vertebræ (Pl. III.). The third cervical has also the hypapophysis (Pl. XI. fig. 3, _hy_).

In all the other cervicals of the present series the hypapophysis is wanting, but each parapophysis developes a plate (Pl. V. figs. 10 & 11, Pl. VIII. fig. 1, _p_) to form the sides of the hæmal canal through which the carotids ran; and the position of such vertebræ in the cervical series is indicated, respectively, by the degree of convergence of these processes, in none of which, where entire, have they met so as to circumscribe the canal: in some of these vertebræ, however, they are mutilated. They differ chiefly in the position and shape of the anapophyses (fig. 10, _a_), which advance from above the postzygapophyses (_z′_), converging towards the middle of the upper surface of the neural arch, being arrested, save in one instance, at the sides of the ligamentous surface occupying the common position of the base of the neural spine.

In the axis vertebra (Pl. V. figs. 12 & 13) the posterior articular surface, concave vertically, and 3 lines in that extent at its middle part, is very convex transversely, being continued upon the sides of the posterior part of the centrum; a thick obtuse hypapophysis (fig. 13, _hy_) descends below this surface: the anterior or odontoid surface presents the usual form in birds; the odontoid process (ib. _x_) has a pit at its apex. The prezygapophyses (fig. 12, _z_), of very small size, project from the outer and fore border of the neural arch, with their articular surface looking outward and slightly upward; a ridge is continued from their back part to the base of the postzygapophyses: the surface (fig. 13, _z′_) in these, 4½ lines in long diameter, is three times the size of the anterior one; it is concave transversely, and looks downward and a little outward. The anapophyses (ib. fig. 12, _a_) are large tubercles rising above the articular surfaces. The base of the neural spine, 9 lines in length (ib. _ns_), is coextensive with the neural arch; the spine rises posteriorly to a height of 6 lines, with a thickness of 2 lines, having a convex upper margin (Pl. III.).

The relative size and position of the cervical vertebræ, as coadjusted in the position and degree of flexure of the neck represented in Sir Hans Sloane’s life-size painting of the Dodo, in the British Museum, are given in Plate III. with the varying proportions of the pleurapophyses and other processes.

_Ribs._ (Plates III. & IV.)

The specimens of ribs include both vertebral and sternal portions; that which appears to be the second or third on the right side (Pl. IV. figs. 7, 7 _a_) is 4 inches 4 lines in length (following the outer curve), and expands to a breadth of 7 lines at its lower part; the interval between the articular surfaces of the head and tubercle is 6 lines. The appendage (ib. _a_) has coalesced with the middle of the hind margin of the shaft. The neck is compressed, with a thin upper margin; the lower one is continued with a curve upon a strong internal buttress-like ridge (ib. _b_), which runs to near the fore part of the flattened body of the rib, where it meets the ridge continued from the tubercle, about 2 inches down the rib: there is a shallow channel between these ridges, contracting to their confluence. The inner surface of the rib is impressed by a deeper and broader channel behind the buttress: the posterior border expands in the form of a triangular plate, with a base of about an inch in extent, due to the complete confluence there of the epipleural process. The anterior border is thicker, and is almost straight. Towards the sternal end the pleurapophysis contracts and thickens, terminating in a rough syndesmotic elliptical surface, 3 lines by 2 (fig. 7, _f_), for the attachment of the hæmapophysis or sternal rib.

A vertebral rib (ib. fig. 2) which is entire, measures 9 inches in length (following the outer curve). The head and tubercle are at the same distance as in the preceding, but the tubercle is broader. The characters of the body of the rib are very similar; but it is narrower, not attaining a breadth of 5½ lines at its lower end; the narrowing and thickening to the articular surface for the sternal rib is more gradual.

A last vertebral rib is adapted, by the longitudinal extent and partial division of the tubercle, to the vertebra which forms the first of the coalesced series of sacrals; and the body of the rib, instead of preserving the regular outward curve of the antecedent ones, is more suddenly bent soon after it emerges beyond the margin of the ilium; the lamelliform part thence continued is straighter, and, moreover, shows upon its outer surface a flattened facet, indicative of pressure or friction by the movements to and fro of the thigh over a rib in such position. Beyond this surface the rib curves in a way not shown in the other specimens; the distal end has the flat syndesmotic articular surface to which had been attached a hæmapophysis not reaching the sternum. In this last (eighth) free rib there is no epipleural process, nor any definitely marked ligamental surface on the posterior margin indicative of the attachment of such process.

The body of a posterior vertebral rib (Pl. IV. fig. 10) shows a fracture which has been healed, with some irregular ossific deposit on the inner surface. All the ribs have a pneumatic foramen (ib. figs. 2, 7, 8, _p_) at the fore part of the neck, near the base of the tubercle.

The eight left vertebral ribs (Pl. III.) and the five right ones do not, either of them, constitute a consecutive series, but have come from different individuals, of different sizes, as exemplified in the third rib figured in Plates III. and IV.

The sternal ribs (P. IV. figs. 3 & 12) are characterized by the two facets, nearly or quite meeting at an open angle, into which their sternal end expands (ib. fig. 3, _c_). One of these ribs, which is entire, shows the single, elliptic syndesmotic surface at the opposite end (ib. _b_); it is 3½ inches in length, with a greatest breadth of 5 lines, and is straight. Another and longer specimen (ib. 12) shows a moderate degree of curvature. A third specimen is 6 inches in length: the proximal end has a breadth of nearly half an inch (the penultimate rib in Pl. III.).

Five successive sternal ribs are indicated by gradational size and curvature, and a sixth, which does not reach the sternum. Before describing this bone I shall proceed with the account of the sacral vertebræ, and the expanded hæmal arches of such as complete the pelvis.

_Pelvis._ (Plates III. & VII.)

The pelvis of the Dodo is chiefly remarkable for the flatness and great breadth of the posterior half, corresponding with the characteristic proportions of that part of the body in Pl. I. fig. 2, and in the old woodcuts of the Dutch “Dodaersen”[25]. It includes sixteen coalesced sacral vertebræ, with which the iliac bones are continuously confluent.

The first sacral shows the transversely extended and concave articular surface of the centrum (Pl. VII. fig. 1, _c_); the subcircular pit (ib. _p_) for the head of the rib is behind the middle of the side of the centrum, at its upper part; the inferior surface is ridged lengthwise; and a transverse low but sharp ridge defines the posterior boundary, the depressions in front of which indicate the hindmost origins of the subvertebral muscle (longus colli?). The anterior outlet of the neural canal (ib. _n_) is subcircular in one specimen, vertically elliptic in others, and 3 lines or less in transverse diameter. From the sides of the neurapophyses stretch out the strong buttresses of bone which blend with the under part of the ilia, giving off from the fore part of their base the præzygapophyses (ib. _z_), and from the back part of their apex the surface (ib. _d_), or part of it, for the tubercle of the last moveable rib, the ilium in the latter variety affording the rest of that surface. The fore part of the strong neural spine (ib. _ns_) is roughened by a syndesmotic surface; it rises to a height of 14 lines, curving forward, and is confluent at its summit with the approximated anterior margins of the ilia. A continuous track of bone, forming a smoothly obtuse longitudinal ridge, represents the summits of the succeeding sacral spines (ib. fig. 2, _ns_) to the hindmost vertebra of the series, without any trace of their primitive division; but this track rises, posteriorly, above the shallow channel on each side, in which are the foramina (ib. _o_), indicating most of the constituent vertebræ.

The second sacral vertebra abuts against the ilium by a pleurapophysis (ib. fig. 1, _pl_ 2), as well as a diapophysis (ib. _d_ 2); but the former is a slender, straight filament, or narrow plate of bone, confluent at both ends.

In the next two vertebræ the pleurapophysis (ib. _pl_ 3 & 4) assumes more breadth and robustness, but is short and straight, abutting against the inner surface of the ilium an inch in advance of the acetabulum. The first of these rib-buttresses inclines forward, and is completely confluent with the ilium; the thicker one (ib. _pl_ 4) has retained part of its primitive ligamentous attachment to the ilium: the proportions of both are subject to some variety.

These are succeeded by three or four vertebræ in which the pleurapophysis is not developed, the attachment to the ilia being by diapophyses only (ib. _d_ _d_), which are short slender lamellæ, directed upward and backward; below and between them are the double orifices for the separate motory and sensory roots of the sacro-spinal nerves. In the next vertebra the pleurapophysis (ib. _pl_ 8) reappears, longer but more slender than in the fourth sacral, extending obliquely backward, and expanding at its extremity to abut against a prominence on the underside of the ilium, opposite the hind part of the acetabulum, with which prominence the rib has completely coalesced by an expanded end. The under part of all these vertebræ is traversed by a sharp median longitudinal ridge, which is more feebly and interruptedly continued to near the end of the sacral series.

Eight vertebræ, abutting by diapophyses only (Pl. VII. _d d_) against the ilia, succeed the one last described; their coalesced bodies are less than half the breadth of those of the preceding vertebræ: they gradually diminish in depth to the last, without loss of breadth. The diapophyses proceed obliquely outward and backward, are lamelliform, about 9 lines in length, and intercept oblong cavities of the same extent and direction, into which open the orifices (ib. fig. 2, _o_) noticed on the upper surface of that part of the pelvis. The articular surface of the body of the last sacral is transversely elliptic, 4 lines by 2 lines, and very slightly convex. The outlet of the neural canal, above it, is circular, and about a line in diameter, the whole vertical extent of the last sacral being 5 lines, while that of the first sacral is 2 inches 2 lines.

The ilium is divided, as usual, into two parts by the ridge on its upper or outer surface (ib. fig. 2, _r_), extending obliquely backward to behind the acetabulum—the anterior division being narrower and concave, the posterior broader and convex but in a minor degree. The anterior (slightly thickened) border of the ilium is curved with the convexity forward, extending 8 or 9 lines in advance of the fore part of the neural spine of the first sacral vertebra. The ilia almost meet above that of the second and third sacrals, with which they coalesce, and then diverge to the oblique boundary ridge, which is thence continued, in some with an angular bend, more directly outward. At this angle the bone is so confluent with the sacrum that the orifices leading to the ileoneural canals[26] are almost or quite obliterated. These canals are, here (ib. _i ï_), the longitudinally extended cavities intercepted between the fore parts of the ilia and the continuous coalesced sacral spines and diapophyses, widening to their anterior outlets. The extent of that part of the ilium in advance of the acetabulum is 3 inches 8 lines; the breadth at its middle part is 2 inches. As the ilium approaches the acetabulum it increases in thickness, and is grooved at the outer margin by a vessel which leaves impressions of its ramifications upon the upper concave surface of the bone (ib. fig. 2, +62+). The acetabulum (ib. _a a_) is circular, 11 lines in the diameter of its outlet, 9 or 10 lines in that of its inner circumference, being widely open, as usual in birds, towards the cavity of the pelvis; the trochanterian surface (ib. _t t_) above the acetabulum is elliptic, with the long axis lengthwise, 9 lines by 6 in its diameter, with its upper border sharp and produced; the anterior border (ib. _b_) of the acetabulum is slightly produced; the position of this articular cavity is about midway between the fore and hind ends of the pelvis. The oblique external ridge of the ilium terminates in the outer margin of the broader part of the bone (ib. _r′_), 7 lines above the sharp and prominent margin of the trochanterian surface (ib. _t_). The ilia have diverged from each other for the extent of an inch and a half behind the beginning of the boundary line (ib. _r_), which interval is occupied exteriorly by lateral ossification from the neural spines to the diapophyses of that part of the sacrum: the mesial borders of the ilia (ib. fig. 2, 62′) slightly converge to the fifteenth sacral vertebra, where they are separated by an interspace of 1 inch, and then again diverge to the last sacral; they coalesce with the diapophyses (ib. fig. 2, _d_ _d_). The inner or under surface of the ilium is thickened into a kind of buttress (ib. fig. 1, _e_), terminating behind the ischiadic foramen. The breadth of the iliac bones and intervening sacrals, 1 inch behind the acetabulum, is 5 inches; at the back part of the pelvis it is 4 inches. The outer border of the posterior part of the ilium (ib. fig. 2, _g_) projects as an obtuse ridge above the ischiadic foramen and the succeeding expanded and confluent part of the ischium (ib. 63), which is vertically concave externally: the ilium, ischium, and pubis (ib. fig. 1, 64) have completely coalesced around the acetabulum. The pubis, which in this part is 7 lines thick, contracts as it becomes free to a diameter of 4 lines; it is smooth and convex below, and has been broken off near the acetabulum on both sides; the fracture shows its pneumatic structure. The ischium, as it recedes from the acetabulum, contracts to a trihedral column, with a vertical diameter of 4 lines; it is concave outwardly, convex inwardly, and suddenly expands below, about an inch from the acetabulum, to form part of the posterior boundary of the obturator foramen (ib. fig. 1, _f_), which is 9 lines in length, and is situated one half in advance of, and the other half beneath, the ischiadic foramen (ib. _m_). This latter is oval, with the large end forwards, 1 inch 3 lines by 10 lines in its principal diameters. Behind this foramen the ischium is confluent with the ilium for an extent of 2 inches, or perhaps rather more, as the posterior margin of the pelvis is not entire in any of my specimens. The inner surface of the ischium forms a low, obtuse longitudinal ridge towards the pelvic cavity, losing thickness as it recedes from the acetabulum. The chief pneumatic foramina in the pelvis are on the inner surface, above the acetabulum, behind the trochanterian articulation, and behind the iliac confluence of the last sacral pleurapophyses,—also at the hinder part of the ilium, on each side of the transverse buttress (ib. _e_) near its posterior junction with the ischium. The prærenal fossa (between _pl_ 4 & _pl_ 8, fig. 1) is deep and subdivided by the diapophysial plates: the postrenal fossa is wide and shallow.

_Sternum._ (Plates III., IV., VI., XI.)

Of this instructive and determinative bone there are two specimens, the one most entire (Pls. III., IV. fig. 4, & VI.) measuring in a straight line, from the costal process to the hind border, 7 inches. The extreme breadth between the lateral processes (Pl. IV. _h_) is 4½ inches; from this diameter the bone contracts anteriorly to a breadth of 3½ inches at the costal processes (ib. _d_), and posteriorly it contracts more rapidly to an obtuse, horizontally flattened apex (Pl. VI. fig. 3). The anterior border of the sternum (Pl. IV. fig. 4) is widely and rather deeply emarginate at the middle (_e_), less deeply so on each side: the breadth of the mid notch (_b_ _e_ _b_) is 1 inch 9 lines, that of each side notch (_b_ _d_) is 1 inch 2 lines. The sternum is deeply hollowed above (Pl. XI. fig. 4), correspondingly convex beneath (ib.); the keel (_s_) is low and thick, commencing by a pair of broad obtuse ridges (Pls. IV. fig. 4, & VI. fig. 1, _r_ _r_) from the mesial ends of the outer walls of the coracoid grooves (ib. _b′_), which gradually rise from the surface of the bone as they extend backward, converging to form the beginning of the keel about 2 inches from the anterior emargination (_e_): the keel gains a depth of ¾ of an inch at the middle of the sternum, then gradually sinks to the level of the bone, as it extends backward, at 1½ inch from the hind end (Pl. VI. fig. 3), a little increasing in thickness as it subsides: its free border describes a pretty regular convex curve (Pl. III.); it is thick, flat, partially canaliculate: the sides of the base of the keel expand, to be continued gradually into the body of the sternum (Pl. XI. fig. 4). Behind the costal surface (Pl. VI. _c_), on each side, extends a lamelliform process (Pls. III. & VI. _h_), ½ an inch in breadth, upward and a little outward, slightly expanding to its free termination, which, however, is not entire in either specimen: the longitudinal extent of this characteristic process, where it is best preserved, is 1 inch; it is conjecturally restored in Plate III.; it answers to the ectolateral process (_h_) of the gallinaceous sternum (Pls. III. & XII. fig. 3): there is no trace of an entolateral process (ib. _i_). The thin margin of the Dodo’s breast-bone, behind the ectolateral process (Pls. III. & VI. _h_), is entire and uninterrupted to the obtuse apex, and the body of the sternum is imperforate: the notch (_f_) behind the process (_h_) represents the ectolateral notch of the gallinaceous sternum (Pl. XII. figs. 1 & 3, _f_). The costal border (Pl. VI. fig. 2, _c_) is 1 inch 9 lines in extent, and 6 lines across its broadest part; it shows articular surfaces for five sternal ribs, of which the four posterior (2–5) are bilobed, the anterior one (_c_ 1) simple, and limited to the outer half of the border; the second sternum shows some variety in this respect: the deep interspaces, in both, are perforated by pneumatic foramina. The costal process (_d_)[27] in advance of these surfaces expands, as it rises upward and a little outward and forward, to the extent of nearly an inch; the hinder and outer side is impressed by a concavity, continued from the costal border; the inner side is smooth and convex: it is not quite entire on either side. The coracoid grooves (Pl. IV. fig. 4, _b_ _b′_) are small in proportion to the sternum, and are divided from each other by an interspace of about an inch; the outer wall of the groove (_b′_), 9 lines in extent, is moderately produced and convex; it appears to be a continuation of one of the initial ridges (_r_) of the keel: the inner wall of the groove (_b_) is deeper, and is formed by the obtuse angle of the anterior border of the sternum, between the medial and lateral emarginations. External to each coracoid groove is a large elliptical pneumatic foramen (_p_) or depression. There is no episternal process. On the convex outer surface of the body of the sternum the “pectoral” ridge (Pl. VI. fig, 1, _k_)[28] is feebly indicated, extending from the outer end of the coracoid groove backward and inward to near the posterior third of the keel. The concave surface of the sternum (ib. fig. 2) shows a number of small pneumatic foramina, chiefly along the middle line to near the posterior third. Behind the costal border the substance of the sternum gradually increases in thickness from the sharp lateral margins to the middle, above the base of the keel, and shows there a fine pneumocancellous texture (Pl. XI. fig. 4).

_Scapular Arch._ (Plates III. & VIII.)

This consists of the scapula (Pl. VIII. figs. 6, 7, 8 & 9, 51), coracoid (ib. figs. 4 & 5, 52), and clavicle (ib. 58), the latter ending in a point and here tied by ligament to its fellow, to form a furculum. I have received the elements of this arch in three conditions:—one in which the bones, though of full size, are separate; a second, in which the scapula and coracoid are confluent, but the clavicle distinct; a third, in which the three bones are confluent at the ends converging to the humeral articulation. The scapula (ib. figs. 6, 7, 8 & 9, 51), 3 inches 7 or 8 lines in length, has the usual sabre-shaped body, slightly expanding and decurved at its free extremity, the breadth of which is 7 lines: it terminates obtusely: varieties of shape are shown in figures 6 & 8. The outer surface of the bone, at the two posterior thirds of its extent, is slightly concave and marked by muscular attachments; the inner surface of that part is smooth and slightly convex: the bone increases in breadth, with some diminution of thickness, towards the articular end, and is remarkable for sending off from the lower border, at 7 or 8 lines from that end, a short process (ib. 51); between this process and the articulation the breadth of the bone is little more than 3 lines; the breadth of the articular end is 9 lines. Nearly one-half of it is occupied by the almost flat, subcircular humeral surface (fig. 8, _a_), with a diameter of 4½ lines, and directed upward, outward, and a little forward. From this is continued an oblong, much narrower coracoidal surface, beyond which the acromial process (fig. 6, _c_) extends forward, curving toward the coracoid, and terminating obtusely.

The coracoid (ib. figs. 4, 5, 8 & 9, 52), averaging a length of 3 inches 7 lines, expands to a breadth of 1 inch 3 lines at its sternal end (52), of which the articular surface (_e_) occupies an inch; the non-articular part forms the outer angle (_m_), and extends in advance of the pneumatic foramen (Pl. IV. fig. 4, _p_) at that part of the breast-bone: the outer border which extends from this free angle to the body of the bone, into which it subsides, at one-third of the extent of the bone, is sharp; the inner border is obtuse to near the inner angle (Pl. VIII. figs. 4 & 5, _n_). The outer surface of the expanded sternal end is smooth and convex; the inner surface is flatter and more irregular, perforated by pneumatic foramina; the diameter of the subcylindrical part of the shaft is 4 lines: the extremes of difference in the distal expansion of the coracoid are shown in figs. 4 & 8, 52, Pl. VIII. A muscular ridge and rough surface (ib. fig. 9, _r_) mark the back part below the middle of the shaft. The bone then expands to its upper articular end, which is obliquely truncate from within outward: it shows, first, the oblong surface for the scapula, which is extended upon the inner prominence of that end; next, the larger and full oval surface for the humerus (_h_), from which the thick, obtuse, inner continuation of the scapular end projects inward, forward, with a slightly upward curve, and shows the narrow oblong surface for the articulation and ultimate confluence of the clavicle (58). The coracoid unites with the scapula at an angle of 100°.

The clavicle (ib. figs. 4 & 5, 58), at its scapular end, is slightly expanded, compressed, with an obtuse recurved termination articulating with the above-named surface of the coracoid, and in one instance coalescing therewith, and by extended ossification with the “acromion scapulæ” (ib. figs. 8 & 9). As the clavicle descends it curves slightly and contracts to a point. The angle at which the pair meet is shown in figs. 4 & 5.

_Bones of the Wing._ (Pls. III. & VIII. figs. 12–17.)

Of the humerus the series contains two specimens, both measuring 4 inches 3 lines in length, one right, and the other left (Pl. VIII. figs. 12–14), but differing slightly in their proportions and in colour—one being of the olive-brown tint with which most of the bones are stained, the other black. The articular head (ib. _a_) is an elongate oval convexity, with the larger end toward the radial side, prominent toward the back and rather flattened toward the front of the bone, which there swells out beyond the base of the articular surface. The radial tubercle is small, and descends from the radial end of the head for about 5 lines; the pectoral process (ib. _b_) is triangular, obtuse, short, and bent, or directed toward the front side of the bone: the ulnar tuberosity (ib. _c_) is more produced in that direction; it is oblong, obtuse, with its base impressed by a large pit both above (fig. 12, _h_) and below—the lower one (ib. _g_) being the deepest, and perforated by a pneumatic foramen; the convex, broad, ulnar border of this tuberosity has two slightly produced processes, an upper or posterior (ib. fig. 12, _c_) and a lower and internal (ib. _g_), which is the smallest. The breadth of the proximal end of the humerus, across the tuberosities, is 1 inch 5 lines, beyond them the bone contracts to a smooth subcylindrical shaft, showing at the back part of the proximal third a longitudinal ridge (fig. 12, _r_), half an inch in length; it gradually expands at the distal third to a breadth of 10 lines, where the articulations offer the usual avian characteristics of the elbow-joint. The head of the humerus is occupied by a fine cancellous structure: into the large vacuity below this, crossed in the section figured (Pl. XI. fig. 5) by a transverse slender bar of bone, the small pneumatic foramina at the bottom of the wide and deep fossa for the axillary air-cell open. The part of the hollow proximal end giving off the pectoral and other processes for the attachment of muscles is strengthened by similar abutments. The pneumatic cavity of the main part of the shaft of the humerus is simple, with a compact wall thicker than at the ends of the humerus, but not exceeding that which is characteristic of the long air-bones in birds. The portion of the distal end chiefly serving for muscular attachments and the antibrachial articulation are also cancellous.

The _radius_ (Pls. III. & XII. fig. 15) is a straight and slender bone, 3 inches 1 line in length, and 2 lines in chief diameter of the shaft. The proximal articular surface is subcircular, 3 lines in diameter, moderately concave; the distal end expands to the same extent, but is compressed, as usual.

The _ulna_ (Pls. III. & VIII. figs. 16 & 17) is 3 inches 1 line in length, of the usual ornithic character, with a well-defined, narrow, elliptic, rough muscular depression, 8 lines in length (fig. 16, _c_), extending upon the shaft from below the anterior or palmar angle of the proximal articular surface. This bone has no pneumatic foramen; the orifice for the medullary artery is above the middle of the same palmar surface, the canal inclining distad. The shaft of the bone is nearly straight; the back or anconal surface, which is slightly convex, shows feeble impressions of the attaching ligaments of the alar plumes, which are represented in all the figures of the entire or living bird. A second ulna is 3 inches 3 lines in length.

There was no carpal or pinion bone in the collection of remains submitted to me: this part of the wing is conjecturally restored in dotted outline in Plate XV.

_Bones of the Leg._ (Pls. III., IX., X. & XI.)

Of the five _femora_ in the above defined series of remains of the Dodo, two measure 6 inches 3 lines in length; one (Pl. IX.) is 6 inches 4½ lines; the shortest is a little under 6 inches, with proportionate differences in the diameter of the shaft. All of them show a small pneumatic foramen (Pl. IX. figs. 1 & 2, _p_) on the inner side of the anterior ridge of the great trochanter (ib. _c_), and on the same transverse line with the head of the bone. This part shows an oblong depression (ib. figs. 2 & 3, _a_) for the “ligamentum teres” at the upper and back part. The articular surface on the same aspect of the neck (ib. fig. 3, _b_), adapted to the trochanterian prominence of the pelvis (Pl. VII. _t_), is well-defined. The trochanter (Pl. IX. fig. 1, _c_) rises, ridge-like, above the level of the head, and is continued from behind the middle of the articular surface on the neck, forward, with a convex outline upon the fore and outer part of the shaft, where it gradually subsides; a narrow intermuscular ridge (ib. fig. 1, _r_), inclining to the middle of the fore part of the shaft, is continued from the trochanterian one. The small trochanter (ib. fig. 3, _d_) is a small subcircular tuberosity, in some specimens a ridge, 3 to 4 lines in length, on the inner side of the shaft, about an inch below the head. The muscular impressions on the fore part of the bone are well defined. A minute medullary canal (ib. fig. 3, _m_) perforates the middle of the back part of the shaft; the popliteal fossa (ib. fig. 3, _o_) shows a few small pneumatic orifices; a triangular rough flat surface divides the fossa from the outer condyle. Above the fibular depression (ib. fig. 3, _g_) there is a well-defined, slightly raised, rough surface (ib. _k_) for the head of the ectogastrocnemius muscle. The ridge (ib. _n_) extending to the back part of the inner condyle is not sharp; the rotular groove (ib. fig. 1, _p_) is deep and moderately wide, with the inner boundary, formed by the narrow anterior part of the inner condyle (ib. fig. 5, _e′_), most produced. The breadth of this end of the longer femora is 1 inch 9 lines; the character of the distal articular surface is shown in Pl. IX. fig. 5.

The head, neck, and great trochanter (Pl. XI. fig. 6) are occupied by a pneumatic cancellous structure, with a thin compact wall on the upper part and sides: this begins to gain thickness at the under part of the neck and at the lower and back part of the trochanter, the compact wall acquiring a thickness of a line at the beginning of the shaft, where the cancellous structure is confined to the outer side of the pneumatic cavity; this structure gives way to a few delicate filaments of bone crossing the cavity of the major part of the shaft, and is not resumed until the bone expands to form the distal condyles (ib. fig. 7).

The five _tibiæ_ of _Didus_ in the same collection range in length from 8 inches 8 lines to 9 inches. The procnemial ridge (Pl. X. figs. 1, 2, 4, _p_) is a triangular plate, with the base longest and the apex rounded off: it inclines outwardly, and does not extend much more than half an inch from the level of the proximal end of the bone: the length of its base rather exceeds an inch: on its inner side a triangular muscular surface is well defined by an irregular inferior line or ridge (ib. fig. 2, _n_). The ectocnemial process (ib. figs. 1, 3, 4, _e_) is thicker, shorter, and terminates roughly and obtusely. There is a low, narrow ridge (ib. fig. 2, _g_), about half an inch in length, on the inner side of the proximal end of the shaft, beginning about 9 lines below the articular surface at that end. The fibular ridge (ib. figs. 1 & 3, _h_), beginning 1 inch 8 lines from the proximal end, extends about 2 inches down the outer side of the shaft. The epicnemial ridge (ib. figs. 1 & 4, _k_) is obtuse, and but little produced above the upper articular surfaces or condyles (_t_ _d_) of the tibia: the breadth of that end of the bone, in the longest specimen, is 2 inches 3 lines. The tendinal canal at the fore part of the distal end is bridged by bone (ib. fig. 1, _l_), and is situated on the inner half of that aspect of the shaft; the lower opening is subcircular and close to the anterior end of the inner lower condyle (ib. _a_), which is more produced forward than the outer one (ib. _b_). Their hind ends project very little beyond the level of that aspect of the shaft of the tibia. An intermuscular ridge (ib. fig. 1, _r_) strengthens into a tuberosity (_r′_) at the inner side of the tendinal groove.

The cancellous structure in the tibia is limited to an extent of about half an inch below the proximal articular surfaces (Pl. XI. fig. 8), and to about an inch and a half from the distal end of the line (ib. fig. 9): the shaft is occupied by a large air-cavity, with a compact wall of half a line in thickness at the upper third, gradually increasing to about a line at the lower fourth, until the cancellous structure is reestablished; the transverse direction of a plate of this structure indicates the extent of the original distal epiphysis of the tibia (fig. 8).

The _fibula_ (Pl. X. figs. 6–8) presents the usual ornithic characters of the bone: it varies from 4 inches 4 lines to 4 inches 6 lines in length, with a greatest proximal breadth of 8 lines. No adequate gain would result from a detailed description or comparison of this bone; and the rest of the bones of the foot have received every requisite attention in this way in the excellent work on the Dodo and its kindred, already quoted. A longitudinal section of the _metatarsus_, taken in the direction from side to side (Pl. XI. fig. 10), shows the loose cancellous texture of the common epiphysis of the three long metatarsals, and the remnant of their contiguous coalesced walls reduced to a thin lamella of bone. As the moiety of the bone figured is the posterior one (of the left metatarsus), the usual oblique position of the middle metatarsal (_iii_), with its proximal end nearer the back part and its distal end nearer the fore part of the coalesced series, produces a corresponding direction of the section, with narrowing and termination of the exposed part of the medullary canal about one-third from the distal end of that metatarsal. The medullary canal of the outer metatarsal (_iv_) is wider and descends lower before the breaking up of the inner surface into decussating lamellæ or filaments, than that of the inner metatarsal (_ii_): the peripheral compact wall of the inner is twice the thickness of that of the outer metatarsal. I may remark that the more posterior position of the middle metatarsal at its proximal end, from which and the corresponding part of the common epiphysis the calcaneal process is developed, is related to the greater share taken by the middle toe in the act of walking and scratching. I will only remark that of the four metatarsals of as many Dodos in the present series, one exceeds by a line the length of that figured in plate xi. _op. cit._, and one falls short thereof to the same trifling amount.

_Skull._ (Plates III. & XI. fig. 1.)

Of the skull of the Dodo, the series of bones transmitted to me include the cranial part with the detached upper mandibular bone (more or less mutilated) of two mature birds, and the lower mandible of three individuals. In the latter the dentary elements (Pl. XI. fig. 1, 32), confluent at the “gonys,” are distinct from the hinder halves of the rami formed by the confluent, or perhaps connate, articular, surangular and angular elements (ib. 31): if the “splenial” were ever distinct, it has coalesced with the dentary, where its upper boundary is indicated by a linear groove or series of small foramina.

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Memoir on the Dodo (Didus ineptus, Linn.)Chapter II: Part 2

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