Geologists divide the history of the earth, since life first appeared upon it, into four general eras, the Proterozoic, Paleozoic, Mesozoic, and Cenozoic, that is, into eras of first life, ancient life, middle life, and recent life. These divisions were made long ago by geologists when it was believed that extraordinary changes, great cataclysmic revolutions, marked their limits.
With a fuller knowledge of the life of the past we know that evolution has been continuous and uninterrupted; possibly accelerated or retarded at times, but without break. Were the earth’s history to be written anew, with our present knowledge, and with an unbiased mind, it is very doubtful whether many of the time divisions would have the same limits that they have now—whether the Paleozoic would terminate with the Carboniferous, or the Permian, or the Trias, or whether indeed we should think it necessary to make any primary divisions whatsoever. In other words, our greater knowledge of living and extinct organisms, and of the rocks which contain fossils, has made the problems of classification much more complex than they seemed to be formerly. It is much easier to classify organisms or rocks, or anything else, when we know only a few isolated kinds—much easier to draw divisional lines. Geological history is like a volume in which pages, leaves, and even whole chapters either are missing or are printed in languages which we understand only imperfectly. Where the lost or unknown parts belong, the largest divisions may be made, and possibly such may have been epochs of unusual activity, of diastrophic changes which greatly accelerated organic evolution. No one can say just where the dividing line should be drawn between the rocks of Paleozoic and Mesozoic age, or between the Mesozoic and Cenozoic, for there is none; the most that we can hope for is to make the divisions everywhere in the world conform to those first made for local reasons.
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Fig. 23a.—Range of the Reptilia.Heavy lines indicate occurrence in North America.
The periods of the Paleozoic era are the Cambrian, Ordovician, Silurian, Devonian, Carboniferous, and Permian, in the order as given; those of the Mesozoic era are the Triassic, Jurassic, and Cretaceous; those of the Cenozoic era, the Eocene, Oligocene, Miocene, Pliocene, Pleistocene, and Recent. As a relic of an old classification we still often divide the Cenozoic into two quite arbitrary divisions, the Tertiary and the Quaternary, the latter including the Pleistocene and Recent only. The same may be said regarding the limits of each of these periods as of the eras; the sole problem is to make each period contemporaneous throughout the world, an exceedingly difficult problem, because no faunas or floras have ever been the same over the whole earth. Indeed, with the exception of some of the lowliest and most generalized forms, or man himself, no species are the same throughout the earth today. Inasmuch as we must depend upon the fossils in the rocks for the determination of the ages, where none is quite the same in strata of remote localities the identification becomes very difficult or even impossible. Nor are the periods, as accepted, of equal or even approximately equal duration; the Cretaceous period, for instance, was longer than all the remainder of the Mesozoic, longer perhaps than all the time which has elapsed since its close.
The earliest animals with a backbone, or rather the earliest that we call vertebrates—for some vertebrates have no vertebrae—began their existence, so far as we know, in late Ordovician times, as attested by fish bones in Ordovician rocks of Colorado and Utah. The first evidences of the existence of air-breathing vertebrates in geological history are footprints preserved in the uppermost Devonian rocks of Pennsylvania. We call them amphibian because they resemble footprints associated with amphibian skeletons in later formations, and because the foot itself is still the most important difference we know between fishes and the higher animals.
Fig. 24.—Permocarboniferous landscape (adapted from Neumayr) with restoration of Eryops, a stegocephalian amphibian ancestrally allied to the reptiles; and Limnoscalis, a cotylosaur (in water).
In the rocks of the next great time division, the Mississippian, as we call it in America, corresponding more or less closely with the Lower or Subcarboniferous of other parts of the world, numerous footprints of amphibians have been discovered, but no fossil remains except a few from near its close in Scotland. From the Upper Carboniferous, or Pennsylvanian, however, not only numerous footprints but the actual skeletons, or impressions of skeletons, have long been known in Europe and America. Until recently all these footprints and skeletons were supposed to be exclusively amphibian. We are now almost sure that some of them belonged to reptiles of lowly type, the earliest coming from near the middle of the Pennsylvanian of Linton, Ohio. The amphibians of this period were, for the most part, salamander-like creatures of from a few inches to two or three feet in length. They all belong to the group collectively known as the Stegocephalia, except that very near the close of the period there appeared small, slender, small-legged aquatic forms which seem to be the ancient representatives of the real salamanders of modern times. Some of the Stegocephalians had become greatly specialized as legless, snake-like, or eel-like creatures.
Fig. 25.—Restoration of Seymouria, the most primitive of known cotylosaur reptiles. From the Permian of Texas, about two feet long.
By the beginning of Permian times tremendous changes had taken place in the land life. The small amphibians of the Carboniferous types dwindled away, soon to disappear, and their places were taken by others of peculiar types, for the most part larger; and by many and diverse kinds of reptiles—water reptiles, marsh reptiles, land reptiles, and even climbing tree reptiles. From the uppermost Carboniferous and Lower Permian rocks of the United States more than fifty genera and twice that many species of amphibians and reptiles have been made known in recent years, and doubtless as many more will be discovered in the future. From other parts of the world the history of reptiles of the Lower Permian is yet scanty, two or three forms from South America, as many more from Africa, and a half-dozen or so from Europe are all; and of these very few are known at all well.
Fig. 26.—Captorhinus, a cotylosaur reptile from Texas.
We classify all the known forms of reptiles from the Lower Permian under three or four orders, the Cotylosauria, Theromorpha or Pelycosauria, Proganosauria, and possibly the Protorosauria, but the classification is yet provisional, representing merely the present stage of our knowledge. The Proganosauria and Protorosauria, including distinctively aquatic reptiles, will be more fully described in the following pages. To give even a brief description of the more terrestrial reptiles of this, the earliest known reptilian fauna, would be beyond our purpose; the accompanying life restorations by the author of some of the more typical and better known forms, based upon nearly perfect skeletons, will suffice.
From the reptiles and amphibians of the Lower Permian of Texas and New Mexico to the ichthyosaurs of the Middle Triassic of California there is a complete gap in the records of the land life of North America. We do not know what became of all the remarkable animals of the Permian. There are few traces of their descendants elsewhere known, unless it be in South Africa. From the Middle and Upper Permian of South Africa and Russia, a marvelous reptilian fauna has been made known in recent years. More than a hundred species of six or seven groups, and at least two orders have been described. Of these the Cotylosauria are the continuation of the American order, but include more specialized forms, the Pareiasauria and the Procolophonia, all of them, like the more primitive American forms, characterized by the imperforate temporal region. The Therapsida, likewise, seem to be the continuation of the American Theromorpha, so closely allied to them that it is difficult to draw a distinguishing line between them. On the other hand, these African reptiles merge through the Theriodontia into the mammals in the Triassic. They are all terrestrial, crawling reptiles, except a few which are described on a later page under the Anomodontia.
Fig. 27.—Restoration of Labidosaurus, a cotylosaur reptile from Texas, about three feet long.
The records of the lower part of the Triassic period are scanty everywhere in the world, save perhaps in Africa. Before the close of the period, however, probably every important group of cold-blooded air-breathing animals had made its appearance in geological history, if we except the snakes; even the mammals had appeared, and possibly the birds. The Cotylosauria, Theromorpha, and Therapsida disappeared, the latter giving birth to the mammals; the nothosaurs and plesiosaurs, the ichthyosaurs, dinosaurs, crocodiles, phytosaurs, rhynchocephalians, lizards, and turtles have all left records of their existence in Upper Triassic rocks; and the pterodactyls had also, in all probability, begun their career, though none is surely known till the Jurassic.
During Jurassic times all these orders of reptiles waxed prosperous and powerful, and branched out in many ways and in countless numbers; many new kinds of each appeared—the marine crocodiles, the quadrupedal dinosaurs, etc.—but no order or suborder, so far as we know, disappeared before its close. And this prosperity continued on into the Lower Cretaceous and for many even into the Upper Cretaceous. The largest dinosaurs disappeared in the Lower Cretaceous, so far as our knowledge goes, but the old-fashioned crocodiles continued on into the Upper, to give place to the new-fashioned kinds. The ichthyosaurs lingered on for a while on the western continent, but the mosasaurs appeared, and the plesiosaurs reached their highest evolution and continued to the end. The flying reptiles attained the zenith of their evolution, but disappeared before the close. The marine turtles attained the maximum of specialization and size. The upright-walking dinosaurs continued on unabated to the close of the period; and a new kind of dinosaurs appeared near the end.
Fig. 28.—Restoration of Dimetrodon,
a pelycosaur reptile from the Permian of Texas;
about eight feet long.
With the opening of the next great era—the Cenozoic or Tertiary—the reptiles dwindled away to their present insignificant position, while the birds and mammals appeared in great numbers and varied forms. The Age of Reptiles was closed and the Age of Mammals had begun.
The history of the reptiles during the Cenozoic is an uneventful one; they ceased their dominion upon land, in the water, and in the air. Their remains are scanty, for the most part, in the rocks of the Tertiary, and such as are known differ only in details from those now living. The land tortoises only, like the mammals of Oligocene and Miocene times, seized the opportunities of open prairies and prospered. A few of the late Mesozoic forms continued a short while into the Eocene. No new groups, perhaps few new families, came into existence during the greater part of this time; it was the age only of land tortoises and the poisonous snakes among reptiles.
The oldest known fossil reptile of North America, or indeed of the world, is represented by a single specimen, lacking the skull, from black shales of Middle Pennsylvanian age overlying a coal seam at Linton, Ohio. The specimen was originally described as an amphibian, but was later recognized by Professor Cope as a true reptile. It was more fully described by the writer under the name Eosauravus Copei, who agreed with Cope as to its reptilian nature. Until the skull is discovered, however, the precise relationships of the animal must remain doubtful.
The next later rocks that have yielded reptilian remains are those of Illinois and Texas formerly supposed to be of Permian age. Later evidence, furnished by invertebrates, however, seems to prove that the lowermost of the strata are of uppermost Carboniferous age. The Illinois deposits, so far as known, are of very limited extent, consisting practically of a single bone-bed in black shale in the immediate valley of the Kaskaskia River near Danville. The known fossils from this bone-bed—all isolated bones—are preserved in the museum of the University of Chicago, and include the types of several genera later recognized in the Texas deposits.
The deposits of Texas, extending northward through Oklahoma to the south line of Kansas, are of considerable extent, for the most part lying along the Wichita River and its tributaries, north of Seymour, Texas. They are composed chiefly of red clays and sandstones of fresh-water or delta origin, perhaps eight hundred feet in total thickness. Beds of like character and yielding similar fossils are also known from northern New Mexico on the tributaries of the Chama River. Their chief characters, as well as restorations of some of the more noteworthy forms, have already been given.
No vertebrate fossils are known in America from the Upper Permian and Lower Triassic. Marine limestones of Middle and Upper Triassic age of Nevada and northern California have yielded numerous remains of primitive ichthyosaurs, the only known remains of the thalattosaurs, and a few others of doubtful affinities, all of which have been described by Dr. Merriam. The Upper Triassic exposures, of considerable extent, occur between the Pitt River and Squaw Creek in Shasta County, California. Reptilian remains from the Middle Triassic are so far known only from the limestones of West Humboldt and New Pass regions of western and central Nevada.
Fig. 29.—Restoration of Varanops, a
theromorph reptile from the Permian of Texas;
about four feet
long.
Land reptiles of Middle and Upper Triassic age are known from many widely separated localities in the United States, but chiefly from the extensive “red beds” of the Rocky Mountain region. The fossils from these beds occur for the most part at least in the horizon called the Shinarump. Its age is usually considered to be Upper Triassic, but the character of the fossils seems to indicate possibly the Middle Triassic. Aside from the stereospondylian amphibians, the last of the Stegocephalia, the vertebrates from this horizon and these regions are chiefly Phytosauria. A few anomodonts, or what seem to be anomodonts—the only record of their occurrence outside of Africa—are known from Wyoming and Utah. And a single specimen from the Wind River red beds, described by the writer as Dolichobrachium, may represent reptiles allied to the dinosaurs. Phytosaur fossils of this horizon have been discovered in Utah, the Wind River Mountains, and near Laramie City in Wyoming; in southwestern Colorado; in western Texas; and in various places in New Mexico and Arizona. Doubtless when these fossiliferous beds are more thoroughly explored many new and interesting reptiles will be discovered.
Phytosaur remains, probably of about the same age as the Rocky Mountain ones, have long been known from the Triassic of North Carolina. From somewhat more recent Triassic deposits in Connecticut and Massachusetts, several skeletons of small carnivorous dinosaurs, and various parasuchian remains have been described by Marsh, Lull, and Talbot. And these beds have long been famous in Massachusetts for their footprints, for the most part originally referred to birds, but now pretty well known to have been made by dinosaurs and amphibians.
No vertebrate fossils of Lower or even Middle Jurassic age are known from North America. From the Baptanodon beds of Wyoming, limestones of about two hundred feet in thickness, four genera of plesiosaurs, the very peculiar ichthyosaur from which the beds take their name, and a few bones of an ancient crocodile are known.
Immediately overlying the Baptanodon beds, the Morrison beds, of from two hundred to four hundred feet in thickness, probably of Uppermost Jurassic and Lowermost Cretaceous age, have yielded an exceedingly rich vertebrate fauna, consisting chiefly of dinosaurs. Discovered first in the vicinity of Morrison, Colorado, in 1877, hundreds of tons of bones have been collected from these beds for various museums. The dinosaurs include many genera of all three suborders, varying in size from that of a cat to some of the largest known land animals. Of other reptiles a very few jaws of a true rhynchocephalian, a fragment of a wing bone of a pterodactyl, numerous turtles, and crocodiles, only, are known. The beds are predominantly black-clay shales, intercalated with sandstones, and all are of fresh-water origin.
From beds definitely known as Lower Cretaceous (Trinity) in Oklahoma, a few bones of a sauropod dinosaur are known, and from nearly corresponding rocks in southern Kansas, plesiosaurs, crocodiles, turtles, and carnivorous dinosaurs are known from sparse remains. Doubtless the Potomac beds of Virginia, which have yielded bones of various dinosaurs, are also of Lower Cretaceous age.
Fig. 30.—Restoration of Casea, a
theromorph reptile from the Permian of Texas,
about four feet long.
With the exception of a single vertebra of doubtful affinities and the cast of a turtle-shell no vertebrate fossils have ever been discovered in the extensive sandstones of Dakota age, the lowermost of the Upper Cretaceous. From the next horizon above the Dakota, the Benton Cretaceous, chiefly marine limestones, at least three genera of plesiosaurs are known from Kansas, Texas, and Arkansas, with two or three more from the limestone shales of Wyoming. A few specimens of armored dinosaurs, two genera of ancient crocodiles, nearly the last of their kind, some marine turtles, and a few vertebrae of ichthyosaurs, the last of the order known anywhere in the world, are also known from the Benton Cretaceous of Wyoming.
Continuous with the Benton limestones above in Kansas are the famous beds of Niobrara chalk; perhaps no fossil deposits in the world are more famous. Exposures covering hundreds of square miles in western Kansas, almost pure chalk, have furnished fossil-hunters during the past forty years literally thousands of specimens of mosasaurs, hundreds of pterodactyls, and scores of plesiosaurs and marine turtles, in addition to the famous birds with teeth and countless fishes of diverse kinds. Two or three specimens of spoon-billed dinosaurs have been found in these deposits, but no other reptiles of any kinds. Beds of like age in Colorado and New Mexico have furnished a few specimens of mosasaurs.
From the marine beds of Fort Pierre age, next above the Niobrara in the west, have come some excellent specimens of two genera of mosasaurs, three or four forms of plesiosaurs, a few pterodactyls, the largest of all marine turtles, and still fewer specimens of dinosaurs, in Kansas, South Dakota, Wyoming, and Montana. From deposits of approximately like age in Mississippi, Alabama, and New Jersey, many incomplete specimens were found years ago of mosasaurs, plesiosaurs, and turtles, the last of the amphicoelian crocodiles, the first of the procoelian crocodiles, and the famous specimen of Hadrosaurus which served for the Hawkins restoration, the first attempt of its kind.
From the uppermost Cretaceous beds of America, the Lance, Judith River, or Belly River beds as they are variously called, have come the remains of a marvelous reptilian fauna. These beds may be grouped together though not all contemporaneous, and there is dispute about their age, some excellent paleontologists insisting that the uppermost are really of Eocene age. From Colorado east of Denver, from eastern Wyoming, from Montana, and especially from the vicinity of Edmonton in Canada, as also occasionally in western Texas and New Mexico, have come many marvelous specimens of dinosaurs, huge bipedal carnivorous dinosaurs, great spoon-billed aquatic dinosaurs, armored stegosaurian dinosaurs, and many kinds of the great horned dinosaurs, the Ceratopsia, so far known only from these beds. Here at the very close of the Age of Reptiles, at the close of the Age of Dinosaurs, are found the ultimate specializations of all the chief groups of dinosaurs except the long-necked quadrupedal dinosaurs which gave up the ghost in Lower Cretaceous times. Many were provided with horns and spines, some indeed seemed to have bristled with spines throughout, a sure sign that they were approaching the end of their career. The modern type of crocodiles had usurped the ancient forms of the early Cretaceous, and reached the largest size of their race perhaps, though but few specimens are known. Here also in these beds we find the first representatives of lizards and snakes in America, though snakes have been described from earlier strata, perhaps, in Brazil. Those archaic, old-fashioned rhynchocephalians described on a later page as the Choristodera appeared also for the first time in these beds, and persisted for a little while in the Eocene, in Europe and America. And with all these there has very recently been described the last of the plesiosaurs, whose race went out with the dinosaurs at the very close of the Mesozoic. It is needless to say that the turtles also occur, for, as a general rule, wherever vertebrate fossils are found, in rocks of the land or the sea, marine or fresh-water, there will be some bones of turtles among them.
With the beginning of the Cenozoic the record of the reptiles becomes relatively scanty in America. In the warm waters of the old Eocene lakes and rivers of Wyoming lived countless crocodiles, true crocodiles of modern aspect and of large size. But, as the climate of North America grew progressively colder, the crocodiles retreated to the south, till, in the Oligocene, the scanty remains of the last crocodiles are found in the American Tertiary. On the other hand, as the open lands appeared toward the close of the Eocene, and in the Oligocene and Miocene, the land tortoises throve and grew greatly in size. In the Bad Lands of South Dakota one may see their remains in almost incredible numbers. And in equally great numbers are these land tortoises, in shape much like the common box tortoise of today, but vastly larger, found in the rocks of the late Miocene or early Pliocene age in western Kansas. And these are the last records of the big tortoises in North America; their descendants are perhaps yet living in the Galapagos Islands.
The history of the lizards and snakes, the only other reptiles found in the Cenozoic rocks of America, is very brief. A few specimens from the Lower Eocene of Wyoming; a few skinks and amphisbaenas from the Oligocene Bad Lands of South Dakota, and some bones of a python-like snake in the early Eocene of Wyoming are about all that we know of the Squamata in the Tertiary. Doubtless snakes and lizards were just as abundant then as now, though but few were preserved, for they are and always have been distinctly terrestrial animals, that only by accident fell into places where they could be fossilized.
The author has collected reptile bones from nearly all of the horizons here mentioned and believes that the list is complete.