# The Zoology of the Voyage of H.M.S. Beagle/Part 1

Described by
RICHARD OWEN, ESQ. F.R.S. F.G.S. F.L.S.
PROFESSOR OF ANATOMY AND PHYSIOLOGY TO THE ROYAL COLLEGE OF SURGEONS IN LONDON; CORRESPONDING MEMBER OF THE ROYAL ACADEMY OF SCIENCES OF BERLIN; OF THE ROYAL ACADEMY OF MEDICINE, AND PHILOMATHIC SOCIETY OF PARIS; OF THE ACADEMY OF SCIENCES OF PHILADELPHIA, MOSCOW, ERLANGEN.
WITH
A GEOLOGICAL INTRODUCTION,
BY CHARLES DARWIN, ESQ. M.A. F.G.S. &c. &c.
CORRESPONDING MEMBER OF THE ZOOLOGICAL SOCIETY.
GEOLOGICAL INTRODUCTION.
BY MR. DARWIN.
MR. OWEN having undertaken the description of the fossil remains of the Mammalia, which were collected during the voyage of the Beagle, and which are now deposited in the Museum of the College of Surgeons in London, it remains for me briefly to state the circumstances under which they were discovered. As it would require a lengthened discussion to enter fully on the geological history of the deposits in which these remains have been preserved, and as this will be the subject of a separate work, I shall here only give sufficient details, for the reader to form some general idea of the epoch, at which these animals lived,—of their relative antiquity one to the other,—and of the circumstances under which their skeletons were embedded. All the remains were found between latitudes 31° and 50° on the eastern side of South America. The localities may conveniently be classed under three divisions, namely—the Provinces bordering the Plata; Bahia Blanca situated near the confines of Northern Patagonia; and Southern Patagonia.
The first division includes an enormous area, abounding with the remains of large animals. To the eastward and southward of the great streams, which unite to form the estuary of the Plata, those almost boundless plains extend, which are known by the name of the Pampas. Their physical constitution does not vary over a wide extent; — the traveller may pass for many hundred miles on a level surface, without meeting with a single pebble, or discovering any change in the nature of the soil. The formation consists of a reddish argillaceous earth, generally containing irregular concretions of a pale brown, indurated marl. This stone, where most compact, is traversed by small linear cavities, and in several respects resembles the less pure fresh-water limestones of Europe. The concretions not unfrequently become so numerous, that they unite and form a continuous stratum, or even the entire mass.
At Bajada de Sta. Fé, in the Province of Entre Rios, beds of sand, limestone, and clay of different qualities, containing sharks' teeth and sea-shells, underlie the Pampas deposit. The shells, although numerous, are few in kind. Mr. George B. Sowerby informs me that they appear to belong to one of the less ancient tertiary epochs; they consist of Venus nov. spec. near to V. cancellata; Arca nov. spec. near to A. antiquata ; a very large oyster, probably an extinct species; an imperfect specimen of a second species of oyster near to O. edulis ; and a Pecten near to P. opercularis. These beds pass upwards into an indurated marl, and this again into the red argillaceous earth of the Pampas, containing the remains of those extinct quadrupeds, which every where characterize that deposit. To the southward of the Plata level plains of an uniform composition, interrupted only at wide intervals by hills of crystalline rock, extend to a distance of about three hundred miles; and to the northward for at least an equal space, and probably much further. As might have been expected from the perfectly level surface, wherever a continuous section is presented on the banks of the great rivers, very slight changes of colour show, that the deposit has been accumulated in strata as horizontal as the land, or as the water-line at the base of the cliffs.
In the province of Banda Oriental (to the N. and N. E. of the Plata), and in part of that of Entre Rios, the land, though very low and level, has a foundation of granitic and other primary rocks. These older formations are partially covered, in most parts, by a reddish earthy mass containing a few small calcareous concretions; while in other parts, they are concealed by more regular strata, of indurated marl passing into limestone, of conglomerates, and ferruginous sandstone. The entire formation probably belongs to the same epoch with that of the Pampas deposit. In the earthy mass, even where it is of little thickness, and where it might readily be mistaken for detritus produced from the underlying granites, remains of large quadrupeds have several times been discovered.
On the shores of the Plata and in the neighbouring districts, proofs of a change of level having taken place between the land and the water within a recent period, may be observed. Both near Monte Video and Colonia del Sacramiento, beds of shells are lying on the beach at the height of several feet above the present tidal action. Near Maldonado I saw estuary shells of recent species embedded in clay, and raised above the level of a neighbouring fresh-water lake.
On the banks of the Parana, a shell identical with, or most closely resembling an estuary species (Potamomya labiata, now living in that part of the Plata, where the water is brackish) is accumulated in great masses, which are found some miles inland, and are elevated several yards above the level of the river. Sir Woodbine Parish, also, has in his possession, shells procured from an extensive formation near Ensenada de Barragan (south of Buenos Ayres), which is quarried for lime. Mr. George Sowerby has examined these fossils, and says the following are identical with living kinds; Voluta colocynthis, Dillwyn : V. angulata, Swainson: Buccinum globulosum, Kiener : a variety of Oliva patula: a Cytheræa closely resembling or identical with C. flexuosa, and a fragment of a second species, probably C. purpurascens; Potamomya labiata ; and fragments of oysters. There is, however, a species of Mactra in very great numbers, with which Mr. Sowerby is wholly unacquainted. I may observe that I found recent shells of the first five species inhabiting the coast, a short distance to the southward. Some shelly limestone from the same place, which Sir Woodbine Parish had the kindness to show me, resembles that which I saw at Bajada, and in Banda Oriental. These beds, therefore, probably form parts of the Pampas deposit, and are not merely indicative of the period of its elevation. Nevertheless, on the opposite shores of the Plata, near the mouth of the Uruguay, I found lines of sand dunes, where the Mactra and Cytheræa flexuosa were lying in such quantities on the bare surface, that the inhabitants, by merely sifting the sand, collect them for burning into lime.
After these facts we may feel certain, that at a period not very remote, a great bay occupied the area both of the Pampas and of the lower parts of Banda Oriental. Into this bay the rivers which are now united in the one great stream of the Plata, must formerly have carried down (as happens at the present day) the carcasses of the animals, inhabiting the surrounding countries; and their skeletons would thus become entombed in the estuary mud which was then tranquilly accumulating. Nothing less than a long succession of such accidents can account for the vast number of remains now found buried. As their exposure has invariably been due to the intersection of the plain by the banks of some stream, it is not making an extravagant assertion, to say, that any line whatever drawn across the Pampas would probably cross the skeleton of some extinct animal.
At Bajada, a passage, as I have stated, may be traced upwards from the beds containing marine shells, to the estuary mud with the bones of land animals. In another locality a bed of the same mineralogical nature with the Pampas deposit, underlies clay containing large oysters and other shells, apparently the same with those at Bajada. We may, therefore, conclude that at the period when the Arca, Venus, and Oyster were living, the physical condition of the surrounding country was nearly the same, as at the time when the remains of the mammalia were embedded; and therefore that these shells and the extinct quadrupeds probably either co-existed, or that the interval between their respective existences was, in a geological point of view, extremely short. In this part of South America there is reason to believe that the movements of the land have been so regular, that the period of its elevation may be taken as an element in considering the age of any deposit. The circumstance, therefore, that the beds immediately bordering the Plata, contain very nearly the same species of molluscs, with those now existing in the neighbouring sea, harmonizes perfectly with the more ancient (though really modern) tertiary character of the fossils underlying the Pampas deposit at Bajada, situated at a greater height, and at a considerable distance in the interior. I feel little doubt that the final extinction of the several large quadrupeds of La Plata did not take place, until the time when the sea was peopled with all, or nearly all, its present inhabitants.
Bahia Blanca, situated in latitude 39°, and about 250 miles south of the Plata, constitutes the second district, in which I found the remains of quadrupeds. This large bay is nearly surrounded by very low land, on which successive lines of sand dunes mark in many parts the retreat of the water. At some distance inland a formation of highly indurated marl, passing into limestone, forms an escarpment. Beyond this, rocks of the same character extend over a wide and desolate plain, which rises towards the flanks of the distant mountain of the Sierra de la Ventana, composed of quartz. On the low shores of this bay, only two places occur, where any section of the strata can be seen; and at both of these I found fossil remains.
At Monte Hermoso, a line of cliff of about 120 feet in height, consists in the upper part of a stratum of soft sandstone with quartz pebbles; and in the lower of a red argillaceous earth, containing concretions of pale indurated marl. This lower bed has the same mineralogical character with the Pampas deposit; and possibly may be connected with it. The embedded bones were blackened, and had undergone more chemical change than in any other locality, which I examined. With the exception of a few large scattered bones, the remains seemed to belong chiefly to very small quadrupeds.
In another part of the bay, called Punta Alta, about eighteen miles from Monte Hermoso, a very small extent of cliff, about twenty feet high, is exposed. The lower bed seen at ebb tide, extends over a considerable area; it consists of a mass of quartz shingle, irregularly stratified, and divided by curved layers of indurated clay. The pebbles are cemented together by calcareous matter, which results, perhaps, from the partial decomposition of numerous embedded shells. In this gravel the remains of several gigantic animals were extraordinarily numerous. The cliff, in the part above high-water mark, is chiefly composed of a reddish indurated argillaceous earth; which either passes into, or is replaced by, the same kind of gravel, as that on which the whole rests. The earthy substance is coarser than that at Monte Hermoso, and does not contain calcareous concretions. I found in it a very few fragments of shells, and part of the remains of one quadruped.
From the bones in one of the skeletons, and likewise from those in part of another, being embedded in their proper relative positions, the carcasses of the animals, when they perished, were probably drifted to this spot in an entire state. The gravel, from its stratification and general appearance, exactly resembles that which is every day accumulating in banks, where either tides or currents meet; and the embedded shells are of littoral species. But from the skeleton, in one instance, being in a position nearly undisturbed, and from the abundance of serpulæ and encrusting corallines adhering to some of the bones, the water, at the time of their burial, must have been deeper than at present. This conclusion might also have been inferred from the fact, that in the neighbouring cliff the same bed, with its shells, has been uplifted some yards above high-water mark. On the coast to the southward abundant proofs occur, of a recent elevation of the continent. In the gravel, nearly all the pebbles are of quartz, and have originally proceeded from the lofty range of the Ventana, distant between forty and fifty miles. Besides the pebbles of quartz, there are a few irregular masses of the same indurated marl, of which the escarpment of the neighbouring great plain is composed. Hence the gravel beds must have been deposited, when the plain existed as dry land; and on it probably those great animals once lived, of which we now find only the remains. The indurated marl forming the plain, is the same kind of rock with that occurring over a wide extent of the Pampas; and there is no reason to doubt, they are parts of one great formation. Nevertheless, the gravel bed of Bahia Blanca, although subsequent to the calcareous formation, may be of the same age with those parts of the Pampas, which stand at a low level near the Plata. For on this whole line of coast, I believe, as the land has continued rising, fresh littoral deposits have been formed; and each of these would often owe part of its materials to the degradation of the one last elevated.
With respect to the relative age of the Monte Hermoso and Punta Alta beds, it is not possible to speak decidedly. A certain degree of similarity in the nature of the strata containing quartz pebbles, and those of the reddish indurated earth; and the short distance between the two localities, would indicate that no long interval had intervened. The beds at Monte Hermoso, certainly were deposited more tranquilly, and probably in a deeper sea; so that even skeletons of animals, no larger than rats, have been perfectly preserved there. In some parts of the surrounding country, obscure traces of a succession of step-formed terraces may be observed; and each of these indicates a period of repose during the elevation of the land, at which time the strata previously existing were worn away, and fresh matter deposited. The Monte Hermoso beds were, perhaps, formed during one such interval, anterior to the accumulation of the shingle bank at Punta Alta.
Mr. G. Sowerby, who has been good enough to examine the shells which were found with the remains of the quadrupeds, has given me the following list.
1. Voluta angulata.
2. -------- colocynthis.
3. Oliva Brasiliensis.
4. ------- Nearly related to O. patula, but specimen imperfect.
5. ------- Nearly related to O. oryza; less nearly to small species now living at Bahia Blanca.
6. ------- Nov. spec.
7. Buccinum cochlidium.
8. ------------ globulosum.
9. ------------ One or two minute species, perhaps young specimens, — unknown.
10. Trochus Nov. spec. (?) same as one now living in the bay.
11. ---------- Nov. spec. (?) nearly related to last ; differs in not being granular on the surface.
12. Assiminia (?) Minute species, identical with one living in the bay.
13. Bulinus nucleus.
14. Fissurella Probably same as a kind (nov. spec. ?) living in the bay.
15. Crepidula muricata.
16. ------------ Nov. spec.
17. Cytheræa Closely related to, or identical with C. purpurascens.
18. Modiola Same as recent kind (nov. spec.) living in the bay.
19. Nucula Near to N. margaritacea.
20. Corbula Minute species, unknown.
21. Cardita Ditto ditto
22. Pecten Nov. spec. (?) very imperfect specimen.
23. Ostrea Oysters of the same size now live in the bay.
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 :—all 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 coexisted on the continent of South America.