I may add that a fossil encrusting coralline is the same with one now living in the bay.
Of these shells it is almost certain that twelve species (and the coralline) are absolutely identical with existing species; and that four more are perhaps so; the doubt partly arising from the imperfect condition of the specimens. Of the seven remaining ones, four are minute, and one extremely imperfect. If I had not made a collection (far from perfect) of the shells now inhabiting Bahia Blanca, Mr. Sowerby would not have known as living kinds, five out of the twelve fossils: therefore, it is probable, if more attention had been paid to collecting the small living species, some of the seven unknown ones would also have been found in that state. The twelve first shells, as well as the four doubtful ones, are not only existing species, but nearly all of them inhabit this same bay, on the shores of which they are likewise found fossil. Moreover, at the time, I particularly noticed that the proportional numbers appeared closely similar between the different kinds,—in those now cast up on the beach, and in those embedded with the fossil bones. Under these circumstances, I think, we are justified (although some of the shells are at present unknown to conchologists) in considering the shingle strata at Punta Alta, as belonging to an extremely modern epoch.
From the principle already adduced, namely, the regular and gradual elevation of this part of the continent, I should have judged from the small altitude of the beds at Punta Alta, that the formation had not been very ancient. The conclusion here arrived at, concerning the age of these fossil mammalia, is nearly the same, with that, inferred respecting those entombed in the Pampas; and it will hereafter be shown, that some of the species are common to the two districts. We may suppose, that whilst the ancient rivers of the Plata occasionally carried down the carcasses of animals existing in that country, and deposited them in the mud of the estuary; other animals inhabited the plains round the Sierra de la Ventana, and that lesser streams, acting together with the currents of a large bay, drifted their remains towards a point, where sand and shingle were accumulating into a shoal. The whole area has since been elevated: the estuary mud of the former rivers has been converted into wide and level plains; and the shoals of the ancient Bahia Blanca now form low headlands on the present coast.
The third locality, which I have to specify, is Port St. Julian, in latitude 49° 15′ on the coast of Southern Patagonia. The tertiary plains of that country are modelled into a succession of broad and level terraces, which abut one above the other; and where they approach the coast, are generally cut off by a line of precipitous cliff. The whole surface is thickly covered by a bed of gravel, composed of various kinds of porphyries, and probably originating from rocks situated within the Cordillera. The lower part of the formation consists of several varieties of sandstone, and contains many fossil shells, the greater number of which are not found in a living state.
The south side of Port St. Julian is formed by a spit of flat land, of nearly a hundred feet in height; and on its surface existing species of littoral shells are abundantly scattered. The gravel is there covered (a circumstance which I did not observe in scarcely any other locality) by a thin but irregular bed of a sandy or loamy soil, which likewise fills up hollows or channels worn through it. In the largest of these channels the remains of the single fossil quadruped, which was here discovered, were embedded. The skeleton probably was at first perfect; but the sea having washed away part of the cliff, has removed many of the bones,—the remaining ones, however, still occupying their proper relative position to each other. I am inclined to attribute the origin of this earthy matter, to the mud which might have accumulated in channels, and on the surface of the gravel, if this part of the plain had formerly existed as a harbour, such as Port St. Julian is at the present day. The Guanaco, the only large animal now inhabiting the wild plains of Patagonia, often wanders over the extensive flats, which are left dry at the head of the harbour during ebb tide: we may imagine that the fossil animal, whilst in a like manner crossing the ancient bay, fell into one of the muddy creeks, and was there buried.
I have stated that existing species of shells are scattered over the surface of this plain; namely, Mytilus Magellanicus; a second and undescribed species, now living on the beach; M. edulis; Patella deaurata; and on another part of the coast, but having similar geological relations, Fusus Magellanicus; Voluta ancilla; and a Balanus:—these shells are among the commonest now living on this coast. Although they must have been lying exposed to the atmospheric changes for a very long period, they still partially retain their different colours. From these facts we know, with certainty, that the superficial deposit, containing the remains of the quadruped, has been elevated above the sea, within the recent period. From the structure of the step-like plains, which front the coast, it is certain that each step must have been modelled, subsequently to the elevation of the one standing above it; and, as the same recent shells occur on two higher plains, we may, with safety, conclude, that the earthy matter, forming the surface of this lower one, together with its embedded skeleton, was deposited long after the existence of the present species, still inhabitants of the sea. According, therefore, to the chronology, taken from the duration of species among the molluscs, the fossil quadruped of Port St. Julian must have been coeval, or nearly so, with those from Bahia Blanca.
Having now briefly described the principal circumstances in the geology of the three districts, to which I at first alluded, I will conclude, by observing, that the fossil mammalia of La Plata, Bahia Blanca, and Port St. Julian, must all have lived during a very modern period in the geological history of the world. It is not the proper place in this work to enter on any speculations, concerning the cause of the extinction of so many gigantic animals. I will only here add, that there is the strongest evidence against admitting the theory of a period of overwhelming violence, by which the inhabitants of the land could have been swept away, and destroyed. On the contrary every thing indicates a former state of tranquillity, during which various deposits were accumulating near the then existing coasts, in the same manner, as we may suppose others are at this day in progress. The only physical change, which we know has taken place, since the existence of these ancient mammalia, has been a small and gradual rising of the continent; but it is difficult to believe, that this alone could have so greatly modified the climate, as to have been the cause of the utter extermination of so many animals. Mr. Owen will mention the exact locality where the remains of each quadruped were discovered; and, at the conclusion, it will be easy to specify by name those, which, from being embedded in the same deposit, are known formerly to have co-existed on the continent of South America.
FOSSIL MAMMALIA.
BY MR. OWEN.
It may be expected that the description of the osseous remains of extinct Mammalia, which rank amongst the most interesting results of Mr. Darwin’s researches in South America, should be preceded by some account of the fossil mammiferous animals which have been previously discovered in that Continent. The results of such a retrospect are, however, necessarily comprised in a very brief statement; for the South American relics of extinct Mammalia, hitherto described, are limited, so far as I know, to three species of Mastodon, and the gigantic Megatherium.
One of the above species of Mastodon (Mast. Cordillerarum) was established by Cuvier on remains discovered by Humboldt, in Quito, near the volcanic mountain, called Imbaburra, at an elevation of 1200 toises above the level of the sea; and likewise at the Cordilleras of Chiquitos, near Santa Cruz de la Sierra, a locality which is near the centre of South America. A second species (Mastodon Humboldtii, Cuv.) is indicated by molar teeth, stated to have been discovered by the same philosophic traveller, in Chile, near the city of Concepcion. The third species of Mastodon appears to have once ranged in vast troops over the wide empire of Peru: numerous teeth were brought thence to Paris by Dombey, and similar teeth, together with a humerus and tibia from Santa Fé de Bogota were placed by Humboldt at the disposal of Cuvier, who considered them to belong to the Mastodon angustidens, a species of which the fossil remains are by no means uncommon in several localities of Europe. Cuvier is also disposed to refer to the same species the teeth of the Mastodon from Brazil and Lima, mentioned by Dr. W. Hunter in his observations on the animal incognitum from the Ohio. The Megatherium has been scientifically described and illustrated in the works of Bru, Cuvier, and D’Alton, whose accounts are founded on a nearly complete skeleton of this stupendous quadruped which has existed in the Royal Museum at Madrid for more than half a century. The few deficiencies in its osteography have recently been supplied by the descriptions and figures given by Dr. Buckland and Mr. Clift, taken from remains of the Megatherium, brought by Sir Woodbine Parish from Buenos Ayres, and which were discovered in the bed of the Rio Salado, a tributary of the Rio Plata. Sir Woodbine Parish’s collection from the same locality, includes also remains of other species of extinct Edentata, which have not yet been described. M. D’Orbigny, in his travels in South America (vol. i. p. 96.), states that, in the banks of the Parana, he found the fossil remains of a large quadruped, of the size of an Ox,—another quadruped of the size of a Cat, apparently of the carnivorous order;—and a third, a Rodent as large as a Rat.
This meagre condition of the historical part of the subject of South American fossils by no means arises from their actual scarcity. The writings of some of the old Spanish authors, for instance, Torrubia, Garcillasso, and others, contain frequent allusions to the bones of giants, who in times of old dwelt in Peru. Legentil, also, in 1728, speaks as an eye-witness of these Peruvian remains; and his guides pointed out to him the traces of the thunder-bolts, by which the Anaks of the New World had been exterminated. Bones and teeth of the Mastodon are, according to Humboldt, so abundant in a locality near Santa Fé de Bogota in Columbia, that to this day it bears the name of the “Field of Giants.”
But independently of these indications, the abundance and variety of the osseous remains of extinct Mammalia in South America are amply attested by the materials for the following descriptions, collected by one individual, whose sphere of observation was limited to a comparatively small part of South America; and the future traveller may fairly hope for similar success, if he bring to the search the same zeal and tact which distinguish the gentleman to whom Oryctological Science is indebted for such novel and valuable accessions.
It is remarkable that all the fossils, collected by Mr. Darwin, belong to herbivorous species of mammalia, generally of large size. The greater part are referrible to the order which Cuvier has called Edentata, and belong to that subdivision of the order (Dasypodidæ) which is characterized by having perfect and sometimes complex molar teeth, and an external osseous and tesselated coat of mail. The Megatherium is the giant of this tribe; which, at the present day, is exclusively represented by South American species, the largest (Dasypus Gigas, Cuv.) not exceeding the size of a Hog. The hiatus between this living species and the Megatherium, is filled up by a series of Armadillo-like animals, indicated more or less satisfactorily by Mr. Darwin’s fossils, some of which species were as large as an Ox, others about the size of the American Tapir. The rest of the collection belongs, with the exception of some small Rodents, to the extensive and heterogeneous order Pachydermata; it includes the remains of a Mastodon, of a Horse, and of two large and singular aberrant forms, one of which connects the Pachydermatous with the Ruminant Order; the other, with which the descriptions in the following pages commence, manifests a close affinity to the Rodent Order.
A DESCRIPTION OF THE CRANIUM OF TOXODON PLATENSIS; A gigantic extinct mammiferous animal, referrible to the Order Pachydermata, but with affinities to the Rodentia, Edentata, and Herbivorous Cetacea.
The cranium, which is the subject of the present description, was found in the Sarandis, a small stream entering the Rio Negro, and about 120 miles to the N. W. of Monte Video: it had been originally embedded in a whitish argillaceous earth, and was discovered lying in the bed of the rivulet, after a sudden flood had washed down part of the bank.
The zoological characters deducible from this cranium, forbid its association, generically, with any known Mammiferous animal, and it must therefore be referred to an extinct genus, which I propose to call Toxodon, from the curved or arched form of the teeth, as will afterwards be described. The specific name, in the absence of other means of knowing the peculiarities of the animal than those afforded by the skull, may be most conveniently taken from the district (La Plata), in which its remains were first discovered.
The dimensions of the cranium of the Toxodon Platensis amply attest that the animal to which it belonged was of a magnitude attained by few terrestrial quadrupeds, and only to be compared, in this respect, with the larger Pachyderms, or the extinct Megatherium. The length of the skull (of which a base view of the natural size is given in Plate I.) is two feet four inches: the extreme breadth one foot four inches. The other requisite admeasurements are given in the table at the conclusion of this description.
The general form of the skull, as seen from above, is pyriform; but viewed sideways, and without the lower jaw, it is semi-ovate; it is depressed, elongate, of considerable breadth, including the span of the zygomatic arches, but becoming rather suddenly contracted anterior to them, the facial part thence growing narrower to near the muzzle, which again slightly expands.
Among the first peculiarities which strike the observer, is the aspect of the plane of the occipital foramen, and of the occipital or posterior region of the cranium, the latter of which inclines from below upwards and forwards at an angle of 50° with the basal line of the skull. This slope of the back part of the skull is one of the characteristics of the Dinotherium; it is common to all the Cetacea, and is met with in a slighter degree in many Rodentia, and in the great Ant-eater and some others of the Edentate order. The corresponding aspect of the foramen magnum presents nearly the opposite extreme to man in the occipital scale, proposed by Daubenton to determine the diversities of the form of the cranium, as a gage of the intelligence of different animals; and the indication of the limited capacity of the Toxodon, thus afforded, is strengthened by the very small proportion, which the bony walls of the cerebral cavity bear to the zygomatic and maxillary parts of the skull, and to the size of the vertebral column, as indicated by the condyloid processes, and foramen magnum.
The zygomatic arches are of remarkable size and strength; they commence immediately anterior to the sides of the occipital plane, increase in vertical extent as they pass outwards, forwards and downwards, and are suddenly contracted as they bend inwards to abut against the sides of the sockets of the two posterior molar teeth.
The cranial cavity is remarkably narrow at the space included by the zygomatic arches; being, as it were, excavated on each side to augment the space for the lodgment of the temporal muscles, so that its diameter at this part is less than that of the anterior extremity of the upper jaw. The upper surface of the cranium expands to form the post-orbital processes, and again contracts anterior to these.
The muscular ridges, or other characters, at the top of the skull, cannot be precisely determined, as a great proportion of the outer table of the bone is broken away, exposing a coarse and thick diplöe. There seems, however, to have been a strong ridge separating the occipital from the coronal or upper surface of the cranium. The form of the remaining parts, which are modified in relation to the attachment of the muscles of the jaws, indicates that these were powerfully developed both for the offices of mastication and prehension. The general form of the skull, while it presents certain points of resemblance to that of the aquatic Pachydermata, and even of the Carnivora, has much that is peculiar to itself; but, in the facial part, it approaches the nearest to that of the Rodentia; and the dentition of the Toxodon, as exhibited in the upper jaw, corresponds with that which characterizes the Rodent Order.
The teeth of the Toxodon consist of molars and incisors, separated by a long diastema, or toothless space. In the upper jaw the molars are fourteen in number, there being seven on each side; the incisors four, one very large, and one small, in each intermaxillary bone.
The general form and nature of the teeth are indicated by the sockets; and the structure of the grinders is exhibited in a broken molar, the last in the series on the left side of the jaw of the present cranium (See a figure of the grinding surface restored of this tooth, fig. 2, Pl. I.), and by another perfect molar, the last but one on the right side of the upper jaw, which, though not belonging to the same individual as the skull here described, undoubtedly appertains to the same species. This latter tooth (Fig. 3, Pl. I.; figs. 2 and 3, Pl. IV.) was found by itself, embedded in the banks of the Rio Tercero, or Carcarana, near the Parana, at the distance of a hundred and eighty miles from the locality where the head was discovered. Fragments of a molar tooth of a Toxodon, apparently the seventh of the left side, upper jaw, were also found at Bajada de St^a Fé, in the province of Entre Rios, distant forty miles from the mouth of the Rio Tercero.
All the molar teeth are long and curved, and without fangs, as in most of the herbivorous species of the Rodent Order: in those, however, with curved grinders, as the Aperea or Guinea-pig, and Cavia Patachonica, the concavity of the upper grinders is directed outward, the fangs of the teeth of the opposite sides diverging as they ascend in the sockets; but, in the Toxodon, the convexity of the grinders is outward, and the fangs converge and almost meet at the middle line of the palate, forming a series of arches, capable of overcoming immense resistance from pressure. (See the upper view of the skull, Plate III., in which the fractures expose to view a part of the series of these arched sockets.)
Of the incisors, the two small ones (the sockets of which are indicated at s s, Pl. III.) are situated in the middle of the front of the upper jaw, close to the suture between the intermaxillaries, and the two large ones in immediate contiguity with the small incisors, which they greatly exceed in size. The sockets of the two large incisors (t t, Pl. III.) extend backwards, in an arched form, preserving a uniform diameter, as far as the commencement of the alveoli of the molar teeth: the curve which they describe is the segment of a circle; the position, form, and extent of the sockets of these incisors are the same as in those of the corresponding teeth of the Rodentia.
The matrix, or secreting pulp of the large incisors, was lodged, as in the Rodentia, in close proximity with the sockets of the anterior molars; and we are enabled to infer, from the form of the incisive sockets, notwithstanding the absence of the teeth themselves, that the pulp was persistent, and that the growth of these incisors, like those of the Rodentia, continued throughout life.
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