3/OLOGV RA 6 THE STORY OF REPTILE LIFE THE FRINGED GECKO (See p. 80). THE STORY OF REPTILE LIFE BY W. P. PY CRAFT F.Z.S., A\L.S., &c. AUTHOR OF " THE STORY OF BIRD LIFE," "THE STORY OF FISH LIFE" 1905 LONDON: GEORGE NEWNES, LTD. SOUTHAMPTON STREET, STRAND " »;• » /• ' BlOLOGt ••---• — :;••.%." ur; TAe rights of translation and of reproduction are reserved. PREFACE. AMONG the higher animals at least, probably none are so generally and so universally disliked and mistrusted as the Eeptiles. Tradition has done much to inculcate this antipathy ; and the natural dread inspired by such as the snakes, has provided justification for its perpetuation. If, however, it be true that " a fellow feeling makes us wondrous kind,7' then, surely, many of those who now regard the Eeptiles as nature's outlaws, should be induced to reconsider their harsh verdict ; since, as it is the purpose of this little book to show, like ourselves, these creatures are also called upon to battle with nature for a hold on life. In the preparation of these pages I have received much kindly help from Dr A. Smith Woodward, F.E.S., of the British Museum of Natural History ; and from Mr G-. A. Boulenger, F.E.S., also of the Museum. For the most valuable assistance which they have giyen me I am grateful. Those who may wish to pursue their enquiries into the story of Eeptile Life yet further, will find a mine of information in Dr A. Smith Wood- ward's "Palaeontology of Vertebrates," and Dr Gadow's volume on " Eeptiles " in the Cambridge Natural History. W. P. PYCRAFT. LONDON, 1905. '-.52884 CONTENTS. CHAP. PAGE INTRODUCTION 9 I. "BEAKED" LIZARDS 11 II. TORTOISES AND TURTLES ... o 19 III. CROCODILES o 46 IV. GECKOS, LIZARDS, AND CHAMELEONS . . 72 Y. SNAKES 106 VI. DOMESTIC LIFE 130 VII. REPTILIAN LIVERIES . 146 VIII. FLYING-DRAGONS 165 IX. EARTH DRAGONS 175 X. DRAGONS OF THE DEEP . 193 LIST OF ILLUSTRATIONS. PAGE The Fringed Gecko (p. 80) . . Frontispiece FIG. 1 — The Tuatera Lizard (Sphenodon punctatum) 1 3 ,, 2 — The upper portion of the Shell — Carapace — of a Tortoise 21 „ 3— The Skeleton of a Turtle ... 24 ,, 4 — The Big-headed Tortoise ... 30 ,, 5 — Armadillo rolled up .... 32 ,, 6 — Outline restoration of an extinct Crocodile 52 ,, 7 — Legs of different Species of Lizards . 93 ,, 8— The Flying-lizard (Draco volans) . . 101 , , 9 — Outline drawing of Chameleons ( Chameleon) 1 03 ,, 10 — Head of a Poisonous Snake dissected . 125 ,, 11 — The Frilled-lizard (Chlamydosaurus kingi) 158 ,, 12 — Outline restoration of an extinct Flying- dragon (Pteranodori) . . . .173 ,, 13 — Outline restoration of the Skull of an extinct Mammal-like Reptile Cyno- gnathus . . . . . .177 ,, 14 — Restoration of Land -dragon . . . 185 ,, 15 — Outline restoration of an extinct gigantic Land -dragon . . . . .188 ,, 16 — Outline restoration of an extinct long- necked Sea-dragon .... 195 ,, 17 — Outline restoration of an extinct Fish- lizard 199 „ 18— A. Left-arm of Crocodile, etc. . . 203 THE STORY OF EEPTILE LIFE. INTEODUCTIOK THE story of Eeptile Life is largely the story of a people, if we may so use the word, whose glory has departed. Just as in the study of the human race we are able to follow the histories of nations, their rise, glorious zenith, and decadence; so with the Keptile folk we may trace a similar course of evolution. The survivors of to-day are but a remnant ; a feeble tribe, spurned and despised. Time was when they were the domi- nant forms of life upon the earth, so that we speak of the " Age of Eeptiles " : for millions of years — from the Permian to the end of the Jurassic era — they held sway, but were at last outnumbered and overpowered by the present reigning type of animal life — the mammalia. But their defeat is without ignominy, since, as we shall show later, their conquerors are at the same time their descendants. Similarly, the Eeptiles are the descendants of the race which they in turn displaced. Lowly in origin, and restricted in influence, these creatures nevertheless rapidly came to the fore ; they spread themselves over the face of the earth, and took possession of the waters 10 THE STORY OF REPTILE LIFE. thereof, and of the heavens above, many exceed- ing in stature all creatures that have ever lived either before or since. Waxing numerous they branched out into many tribes, each carving a way for itself. What the members of these several tribes were like, and how they have overcome their enemies ; what weapons they have employed ; what subterfuges they have re- sorted to ; how they have conducted themselves as parents ; and what their pedigrees disclose, is to be the theme of the following chapters. But before w^e proceed further, it is essential that a clear idea should be formed of what con- stitutes a reptile. Save among experienced naturalists, only the very vaguest notions appear to exist on this head. Generally, a group of animals totally distinct is included in the popular idea of the class reptiles. These alien forms are the frogs and toads, newts and salamanders. Now between the reptile and the creatures just referred to there is a very wide difference. Both agree in being "cold-blooded," in having a back- bone, and four limbs adapted for walking : but they differ fundamentally in that the young of the frog-tribe — certain peculiar exceptions apart — when they leave the egg do so in the form of "larvae." That is to say they differ from their parents in that they are obliged to live in water and to breathe by means of gills like the fishes. Later, these gills disappear and breathing by lungs is substituted. But in this adult stage they still differ from the reptile in that the skin is naked, and richly supplied with glands for the purpose of keeping the skin moist — hence the "BEAKED LIZARDS." 11 " sliminess " which makes these creatures so objectionable to many. Among the reptiles the young leave the egg in the form of the parent — that is to say, they never pass through a gill- breathing stage. Further, the body is invested in a scaly covering, and the skull moves upon the backbone by means of a single bony knob, instead of two such knobs, or " condyles " as they are called, as in the frog and its allies. Other characters there are whereby these two groups — the Reptiles and the " Amphibia" — may be distinguished, but we do not intend to discuss them here. They are of too technical a character for this little book. CHAPTER I. "BEAKED LIZARDS." AFTER the elimination of the alien tribes re- ferred to in our Introduction, we have left four large groups, the Crocodiles, the Tortoises and Turtles, the Lizards, and the Snakes, and lastly, a fifth, represented only by a single species, the Tuatera lizard of New Zealand. This creature, with certain fossil forms, constitutes the order Rhynchocephalia, or beaked " lizards." Though but remnants of a fallen race, as we have just remarked, these four groups are so distinct one from another that it will be neces- sary to tell the story of their rise and evolution in as many separate chapters. 12 THE STORY OF REPTILE LIFE. The distinctions which enable us so easily to discern the reptile from the amphibian, are based upon observations made upon living ani- mals. A study, however, of the skeletons of the two groups would have led to practically the same results. But supposing there had been no survivors either of the Amphibia or the Eeptiles, then the dividing line would have been very difficult to draw ; for though the skeleton of any of the modern Amphibia differs conspicuously from that of any reptile now living, the same is by no means true of the more primitive types of these two groups. Indeed, even at the present day the greatest experts are not agreed as to the class to which certain very ancient fossils belong. One places them with the Amphibia, regarding them as members of the group known as " Laby- rinthodonts," another with the Keptiles. These creatures, it is not surprising to learn, are ances- tral types from which have probably sprung the living reptiles of to-day. Other and allied forms of the problematical types probably gave rise to some of the fossil species — the "Dragons" of the later chapters of this book. Be this as it may, belonging to the same geological era — the Permian — remains have been found of an un- doubted reptile, known as Palseohatteria, which is represented to-day by an actual living de- scendant, the " Tuatera " of New Zealand. This "living fossil," as it has been called, is thus one of the most remarkable of existing reptiles, and forms in itself a quite distinct group, the fifth, to which reference has just been made. With an ancestry traceable for millions of years, this .2 3 •a « « •§ g O ^H 14 THE STORY OF REPTILE LIFE. wonderful creature, according to human stand- ards, is one of the very bluest blood. Yet, measured by the cold and impartial standards of scientific criticism, it ranks among the lowest instead of the highest of the Eeptile people. The proud position of precedence is given to the much less ancient house of the Crocodiles, these having risen highest in the scale of evolution. Whether, as some hold, the Tuatera, or Hatteria as it is also called, represents the stock from whence our reptiles of to-day have been derived, or whether it and its ancestor the Palseohatteria are offshoots derived, in common with the remaining living species, from the problematical reptiles, is a point which we do not propose to discuss here. Those who would examine the evidence on this matter will find much information ready to hand in Dr Gadow's book, to which reference has already been made. Whether the Tuatera is rightly regarded or not as the representative of the ancestral stock from which the existing reptiles are derived, it is probably the most primitive of all living species of this class. On this account, then, it is fitting that it should be described, at least briefly, in this opening chapter ; for many of its peculiarities appear again, with modifications, in other groups. Although the Tuatera is generally referred to as a lizard, it is really nothing of the kind. It is no more permissible to speak of it as a lizard, than it would be to call it a tortoise or a croco- dile. It stands by itself in a group apart from "BEAKED LIZARDS." 15 all its contemporaries. The evidence on which this decision is based is of too technical a char- acter for discussion here : we need only say that it rests upon the structure of the skeleton, and of certain internal characters related to the organs of generation. Of the skeletal structures which appear again in other groups, sometimes strangely modified, we may mention one or two of the more im- portant. The ribs, for example, in the Tuatera are remarkable for the presence of curious hook- like processes which project backwards from the middle of the upper portion of each rib, to overlap the rib next behind it. These " uncinate " processes occur elsewhere only among the Croco- diles and the birds. Behind the breastbone are found numerous rod-like bones embedded in the muscles of the belly. These are commonly known as the " abdominal ribs " ; they occur again in the Crocodiles and the ancient " Labyrinthodonts " and " fish-lizards," and it is probable that from the fusion of similar bones the singular shield on the belly of the Tortoise has been derived. The teeth are quite remarkable. Numerous and irregular in size, they occur not only along the edges of the jaws, but also on the bones of the palate. These teeth are not renewed when worn out as in most other reptiles, but fusing with the edges of the jaws gradually wear away till, in very old individuals, the jaws become toothless. It is on this account that the name beaked lizards has been bestowed. Large males attain a length of about two feet and a half. Both sexes are alike save only in that the males have 16 THE STORY OF REPTILE LIFE. a larger head and a stronger nuchal crest. Of a dark olive green, with small spots of white on the sides, the general appearance of the animal is sober enough. It is enlivened, however, by a row of pointed and slightly erectile spines along the ridge of the back and extending on to the tail. The under surface is clothed in moderately large scales, whilst those on the sides are so small as to give a granular appearance. The eye is large and has a vertical pupil. The tail is thick and compressed, and like that of many lizards, and at least one species of mammal — a mouse, which occurs in the island of Cyprus — is easily shed. This is an ingenious device which enables the owner to escape when seized by this organ. Among the lizards at least, when threatened, the tail is temptingly paraded, when, as soon as it is grasped, the creature makes a dash for liberty, the tail becoming detached with the slightest shock. The lost appendage is speedily renewed, and with it a new chance of escape from unpleasantly close encounters is gained. A similar regeneration does not, however, appear to take place with the mouse in question. But perhaps more than on any other account the Tuatera is celebrated for the presence of a third eye seated in the middle of the roof of the skull. It should be mentioned, however, that this organ is now quite f unctionless and has shrunk to quite insignificant proportions. It is connected by a long stalk with what is known as the pineal gland, an outgrowth of the " primary fore-brain." Coming to the surface through a special aperture in the skull, between the parietal "BEAKED LIZARDS." 17 bones — hence the name parietal eye — it is covered by a horny scale. At one time this eye was undoubtedly functional. But whether it served some peculiar need in the days of long ago, a need which has now ceased to exist, or whether it merely supplemented the normal lateral eyes which were then less perfect than now, no man can tell. Perchance it is a survival of pre-reptilian days, that is to say of some larval condition, before the lateral eyes were developed. The discovery of this eye is quite a modern achievement; and immediately the fact of its existence became known a vigorous search was instituted by anatomists the world over, with the result that a similar, but yet more vestigial eye was found in many other living reptiles, and in a large number of extinct forms. In many of the latter, indeed, this eye seems to have been of considerable size. This is especially the case in the old fish lizards, Ichthyosaurus. Besides the reptiles, however, the amphibia and fishes also show traces of this organ. The Tuatera is, alas, verging on extinction. Bush-fires, wild-pigs, dogs and cats, and reptile- eating Maories and the advancement of civilisa- tion have swept the Tuatera from the mainland of New Zealand, so that to-day it is to be found only in a few uninhabited islands. Here it dwells in the seclusion of a burrow which it digs for itself, and into which a hasty retreat is beaten on the slightest sign of danger. The Tuatera is a sociable animal, sharing its burrow with various kinds of Petrels, though, be it noted, B 18 THE STORY OF REPTILE LIFE. members of its own species are summarily evicted should they presume to attempt a lodgement. The birds, by a mutual arrangement, occupy the left, and the Tuatera the right side of the chamber. It is interesting to note that whilst the dark burrow serves all the purposes of a nursery for the bird, which incubates its eggs, it is unsuitable for this purpose for the pro- prietors of the burrow, who, owing to their more phlegmatic temperament are compelled to seek the aid of the sun to bring their young into being. The hatching time covers a rela- tively enormous period — thirteen months, and is rendered further remarkable by the fact that the development of the embryo is, for a time, during the winter months, suspended so that the embryo may be said to hybernate within the egg. Somewhat lazy in their movements as a rule, crawling at a slow pace and dragging the body and tail along the ground, when animated by the excitement of the chase they lift the whole trunk off the ground, and move with some speed. This is kept up, however, only for a few yards, when they grow weary and stop. During the greater part of the day they sleep ; and are fond of lying in the water, being able to remain submerged for hours without breathing. They feed only upon other animals, and these they will take only when alive and moving about. During the night, and especially during the pairing season, they are said to croak or grunt. CHAPTEE II. TORTOISES AND TURTLES. FAMILIARITY certainly blinds our eyes to muck that is wonderful in Nature. Perhaps in no- instance is this more true than in the case of the -creatures which form the subject of the present chapter. To the world generally the Tortoise is regarded as a somewhat uninteresting creature, the type of sluggishness and the source of the familiar ornamental commodity known as "tortoise-shell," whilst its cousin the Turtle has acquired undying fame on account of the excellent soup it affords when boiled ! Yet, if we look but a little below the surface, really as well as metaphorically, we shall discover that these "ugly ducklings" occupy a really unique position in the animal kingdom. This isolated position these creatures owe to the remarkable coat of mail which they have developed as a protection against their enemies. This armour, in the typical tortoise, takes the form of an inflexible shell investing the trunk, but leaving the head, tail, and limbs free. These, when danger threatens, can be drawn into the front and hinder apertures of the shell so that, being also armoured, a practically invulnerable mass is presented to the enemy. Critically examined, this shell is found to be composed of an outer layer of horny plates or shields, superimposed upon a closely fitting series. 19 20 THE STORY OF REPTILE LIFE. of bony plates, which, like the horny covering shields, are also symmetrically disposed. Generally, in armour-clad animals where the protecting covering is made up of horny shields covering bony plates, the horny outer layer exactly corresponds to the bony plate beneath it. This is not the case with the tortoises and turtles, or, to use a more comprehensive term, embracing all the members of this order, the Chelonians. In the specimen from which this figure was drawn, the horny shields in question were removed from one side of the carapace, as this shell is called, so that their relationship one to another and to the bony elements beneath them, can be readily made out. If the exposed portion of the underlying bony shell be examined two sets of patterns can be made out on its surface. One is formed by shallow grooves, which represent the impress of the edges of the horny shields, the other by a series of curiously ziz-zag lines. These represent the sutures or rough jagged edges of distinct bones, which have a very remarkable history, inasmuch as one set — those which may be seen running down the middle of the back — are formed by table-like expansions of what are known as the " spines " of the vertebrae, or back- bone, and therefore belong to the skeleton ; whilst the remainder, those which take the form of bands, running at right angles to the backbone, and those which form the margin of the shell, are derived from bones originally embedded in the skin. They are this, and something more. Since the day when they formed nodules of TORTOISES AND TURTLES. 21 bono embedded in skin, many and strange changes have taken place. The order of these changes none can tell, only the result thereof is ours, and this, without exaggeration, may be called startling. When we come to take FIG 2. —The upper portion of the shell— carapace— of a tortoise, to show the form and position of the horny plates and underlying bones. The horny plates are indicated on the right hand side, the impres- sion of their edges made upon the underlying bones is shown by dots on the left side. The bones are shown by jagged lines. an inventory of the structures pertaining to the outside of these creatures — structures which, without previous experience, we might expect to find, since they are found in other reptiles — one of the first things to be missed would be "22 THE STORY OF REPTILE LIFE. the skin, and the next the thick layer of muscles underlying this. Both are wanting in the trunk of the Tortoises ! When we come to look deeper into this anomalous state of things we find that the bands of bone which we have just described as running at right angles to the long axis of the skeleton, that is to say the bands of bone running from side to side, have a strangely complex history. Originally, as we have said, nodules of bone embedded in skin, they have, with the wasting of the underlying muscles, come to lie at first directly upon the ribs, and later, fusing therewith, have ultimately replaced even these almost completely. So much so, that all that re- mains to-day, of what appear to be complete ribs, is the head thereof, or the portion which joins the backbone and the tip, or that portion which supports the little bones which form the margin of the shell. This we know, because in the very young tortoise comp]ete ribs are present, but as development proceeds, all save the two ends become absorbed and replaced by the bones, origin- ally, as we have said, belonging to the skin. Exactly how this is done requires an intimate knowledge of the changes which may take place in the tissues of animals, and these we cannot attempt to deal with here. Those who would verify these statements must consult more technical works than this little book, and bring with them at the same time the results of a scientific training. The history of the breastplate of the tortoise is no less strange. Like the shell this is made up, externally, of symmetrically arranged plates TORTOISES AND TURTLES. 23 of horn, covering equally symmetrically arranged bony elements. There is no skin, save that represented by the horny shields, and no inter- mediate layer of flesh. The bony elements repre- sent in part the bones of the shoulder girdle of other animals, and in part the peculiar " abdominal ribs " which we have already mentioned as a notable feature in the Tuatera " lizard." There is apparently nothing equivalent to the breast- bone of other reptiles to be found in the Tortoises. The development of the shell has been accom- panied by many other and profound changes in the form of the skeleton. Thus, the backbone, between the base of the neck and the base of the tail, has become virtually suppressed, its originally separate elements being greatly reduced, and immovably fused one with another. The reduc- tion of the spine to its present almost vestigial condition has been a long process. The decline was inevitable directly the bony shell had acquired rigidity enough to prevent the movement of the backbone. As soon as this stage was reached a new order of things became established, and the material of the superseded spine became available for absorption and reincorporation into the newly developing structures. This is the fate of all useless organs in Nature ; they are broken up, like old buildings, to contribute to the growth of new ones. Yet another remarkable feature is the position of the limb-girdles. In all other animals the shoulder-blades lie on the outside of the foremost ribs ; but in the Chelonians these bones are placed on the inside of the ribs, i.e. within the shell. 24: THE STORY OF REPTILE LIFE. Similarly the haunch-bones which support the hind-limbs, in other animals lie behind the last rib, and near the surface of the body, in the Chelonians they lie, like the shoulder-blades, within the shell. The anomalous position of FIG. 3. — The skeleton of a turtle, viewed from the inside, after the removal of the breastplate. Note the position of the shoulder and hip girdles, lying beneath the ribs instead of above them. these bones, however, is only a feature of adult life. In very young tortoises, where the develop- ment of the shell has only just begun, the shoulder- blades lie a little in front of the first rib, whilst the hip-bones are to be found just behind the last rib. But the broadening of the ribs caused by the excessive development of the external bony shields gradually creeps over the shoulder- blades in front, and the hip-bones behind, so that TORTOISES AND TURTLES. 25 at last they came to lie, as we see, entirely within the shell. It is interesting to note that some few tor- toises seem to have found that the inflexible bony shell, such as we have just described, needed yet further modification to make it a thoroughly trustworthy fortress against attack. This need has been met by developing a hinge either across the carapace, or across the plastron or breast shield. By this means one or other or both of the apertures of the shell can be closed completely as by a portcullis. The hinged carapace, be it noted, is found only in these species belonging to the genus Cinyxis, found in tropical Africa, whilst the hinged plastron has been independently acquired by several different tortoises in widely different parts of the world. Thus the little Spider-tortoise of Madagascar has the plastron so hinged that the front portion can be drawn up, so as to completely close the mouth of the shell. The Iberian tortoise of Spain and Morocco closes the hinder aperture of the shell in a similar manner. In this species, however, the hinge does not appear till comparatively late in life, and is best developed in females. Some six species of the genus Testudo, found in India, Madagascar, and S. China, can also .close this end of the shell by raising the hinder half of the plastron. But the N. American species of the genus Cistudo have made a decided improvement on the mechanism adopted by the foregoing, inasmuch as these can raise both ends of the plastron, so that when the head, legs and tail are drawn in, and the "oak is sported," the 26 THE STORY OF REPTILE LIFE. hungry enemy must either pass on to other game, or sit down and endeavour to take his prize by siege ! As these creatures are able to live for long periods without food, siege tactics -are not likely to succeed. Before we leave the subject of the shell we must briefly comment on the remarkable cara- pace and plastron of the rare Leathery Turtle {Dermochelys) of the West Atlantic and Indian Oceans — the largest of all living Chelonians. x Like that of the Tortoises we have already examined, the shell is of dermal origin. That is to say, it is made up of bony growths developed in the outer skin ; but here the resemblance ceases. For whereas in the Tortoises and Turtles gener- ally these bony plates are symmetrically disposed, and, in the carapace sink down on to and become inseparably fused with the skeleton, in the Leathery Turtle they take the form of innumer- able small plates* interlocked one with another to form a mosaic. The carapace of this turtle is furthermore peculiar in that it remains per- manently distinct from the skeleton, so that, when removed the ribs and vertebrae are re- vealed, muscle-covered, as distinct as in other reptiles. The horny shields which cover the shell generally in other Chelonians are here con- spicuous by their absence; instead, the shell is •covered with a smooth, leathery skin — hence the name, " Leathery Turtle." The fundamental differences between the shell of the Leathery Turtle and that of other Chelo- nians has an important bearing upon the ques- tion of the origin of the two groups. The TORTOISES AND TURTLES. 27 mosaic-like plates of the carapace of the former resemble the plates of the bony armour of the crocodiles, and are probably derived from a primitive armature of this kind. If this be so, then the bony plates of other Chelonians must be regarded as a later development, the origin of which is yet to be discovered. This is the view most generally favoured at the present time, and accordingly, we must look upon the Leathery Turtle as the sole survivor of a primitive and independent group. Parallels are always interesting, and it is seldom that they cannot be found in the animal world, however remarkable the instance we may have to match. Thus, though the shell of the Tortoise has no counterpart among the living reptiles, we find a very close resemblance thereto in the shell of cer- tain gigantic and extinct mammals — the S. Ameri- can Armadillos, known as Glyptodonts. These creatures were encased, like the Chelonia, in a bony shell, which in some forms is as much as five feet long, and an inch in thickness. The struc- ture of this shell resembled that of the Leathery Turtle, in that it was made up of a series of small bones closely interlocked to form a mosaic, but different therefrom in that each of these bony plates was covered by a horny shield. As in the Chelonia, the separate bones of the verte- bral column were welded together to form a tube. The limb-girdles, however, did not occupy the anomalous position which they held in the Chelonia, inasmuch as they bore the same rela- tion to the skeleton as in other animals. 28 THE STORY OF REPTILE LIFE. In the structure of the skull several points are to be observed whereby the Chelonia differ from other reptiles. With these, in detail, we have no concern here ; for our purpose it is enough to notice, firstly, that teeth are con- spicuous by their absence. Their work is per- formed by horny sheaths which encase the jaws as in birds. That the ancestral Chelonia had teeth is very probable, and doubtless some day this fact will be established by the discovery of a fossil skull with teeth implanted in the jaws. In one other point the Chelonia and the birds agree — though this of course by no means im- plies relationship — and this is in the form of the lower jaw, which, instead of being made up of two separate halves, is fused into a single bone. The breathing of the Chelonia has acquired certain peculiarities, inasmuch as, on account of the rigid walls of the shell, expansion of the chest cavity by the movement of the ribs and abdomen has become impossible. The lungs, which are complicated, spongy structures, are filled and emptied partly by the movement of the neck and limbs, which by their movement act as pistons, and partly by the action of the tongue bones, which are of great size. By these, when the neck is stretched out, the throat is alternately inflated and emptied by air drawn in through the nostrils. The deflation of the throat causes the air to be forced down the windpipe, the valves of the nostrils preventing its escape by any other way. In every great group of animals we find that the struggle for existence has caused a gradual TORTOISES AND TURTLES. 29 dispersal of the members thereof, compelling them to seek a livelihood in an environment quite different to that of the centre from which they started. Thus we get an exchange con- stantly taking place between the inhabitants of the sea and those of the land. We find the crab deserting his natural element to climb palm-trees for cocoa nuts, and mammals which have adopted the life of fishes. Such an exchange, however, can only take place under certain conditions — the emigrants must adapt themselves to the require- ments of their environment ; and this brings about a more or less complete transformation of the body. Among the Chelonia we have many instances of this. Originally terrestrial, some have adopted a fresh water habitat, others have taken to the sea. The modification which these aquatic forms have undergone are sufficiently well marked to- render them easily distinguishable from their stay-at-home relatives. Hence we get Land Tor- toises, Water Tortoises, and Turtles. What the ancestral Tortoise may have been like we do not know, but its descendants do not- appear to have found any great necessity to change their form after once the general archi- tecture of the body had been determined on. This much is to be gathered from the fact that the fossil remains of these creatures, which occur in remote geological formations — the earliest known Chelonian occurring in the Upper Keuper of Wiirtemberg — -differ but little from the same parts of the skeleton of its nearest modern repre- sentative. It is only in minor characters, divid- 30 THE STORY OF REPTILE LIFE. ing the larger or smaller groups of species one from another, that modifications occur among the living Chelonia. This conservatism is really very remarkable when we reflect that they were already an ancient group long before we have any record of the advent of the birds upon the earth. It seems difficult to realise that the conditions of life through such enormous periods of time can have affected them so little. Fig. 4.— The big-headed tortoise. This creature is remarkable for the large size of the head and the extraordinary flatness of the shell. In the history of the past, many animals have sought protection for the body by encasing it in armour-plate. It would seem, however, that such protection either ends in bringing about the extermination of the species by which it has been adopted, or in being discarded by later generations. The Chelonia afford an exception to this rule. Judging by the conditions of life which obtain to-day, it is not easy to see why this should be so, inasmuch as the Tortoises and Turtles do not appear to be beset by any more formidable enemies than their unprotected rep- tilian neighbours. The key to the mystery may TORTOISES AND TURTLES. 31 perhaps rest in their slowness of motion. Their unarmoured neighbours escape by flight, a way- which is impossible to these sluggish creatures. Land-tortoises may be readily distinguished by their feet. The toes are very short, with no trace of webs between them, and the hind-feet at least are club-footed. The peculiar form of these feet is the outcome of adaptation to the support of the remarkably heavy and inflexible body. To the weight and immobility of the trunk we may attribute the fact that the creatures are confined to the earth. Tree-climbing to them is an impossibility. They may delve below the surface, but they cannot rise above it. But this by the way. The outer surface of the legs is covered with little horny scales, covering, in many cases, bony nodules, so that when drawn into the shell the mouth and hinder apertures thereof are effectually protected against attack. In some cases, as we have already remarked, these apertures are closed by raising the ends of the breast-plate. Comparisons are said to be odious. Applied to human affairs this is often true. In natural history it is otherwise. Let us then contrast the method of barricading practised by the Tortoise with that of the remarkable South American mammal, the Armadillo. This animal, like the Tortoise, is encased in bony armour covered with horny plates. The great back-shield differs from that of the Tortoise among other things in its great flexibility, so that, having no breast-plate, the animal can curl itself up into a ball, leaving but a single aperture, which is closed by the tail 32 THE STORY OF REPTILE LIFE. and head-shield. How wonderfully this is done can be seen in the accompanying illustration. Of the Land Tortoises, one of the most familiar, at least in Great Britain, is the " Greek Tortoise " Fig. 5. — Armadillo rolled up. (Testudo iberia), this being the species commonly offered for sale on barrows in the streets, from whence it finds its way into our gardens, and also, not seldom, our houses, being imported into the latter under the mistaken idea that it will kill the "black beetles." TORTOISES AND TURTLES. 33 The home of the Tortoise is 1ST. Africa and S. W. Asia. It also occurs in Southern Andalucia, breeding in the sandy pine forests of the Marismas, near the mouth of the G-uadalquiver. There are three claimants, it should be men- tioned, to the title of " Greek " Tortoise. The second, Testudo Grceca, is a close ally of T. iberia, and occurs in the northern half of the Balkan Peninsula, Asia Minor, Syria, Italy, and the Islands of the Mediterranean. The third is the genuine Greek Tortoise, Testudo marginata, since it is restricted to Greece proper, and is the only land tortoise which is found there. These three species are very much alike in habits, and are exclusively vegetable feeders. They are very fond of basking in the sun, and extremely averse to getting wet, retreating to some sheltered place on the fall of the first few drops of rain. On the approach of winter they bury themselves in a hole in the ground, or in a heap of decaying leaves and there hybernate till spring. The " Gopher " Tortoise of the South-eastern States of North America is a particularly interest- ing species on account of its curious habit of living in a burrow. The burrow, at its mouth, is only sufficiently wide to admit the body of the animal, and runs slightly downwards to a length of about four feet. The whole passage may be as much as two yards long, and gradually widens from the mouth inwards, terminating in a spacious chamber lined with branches of fir trees. Each burrow is inhabited by a pair of tortoises. When C 34 THE STORY OF REPTILE LIFE. the dew is on the grass, or after rain, they emerge to feed upon the grass, succulent vege- tables, or fruit, varying this with gum which exudes from trees, especially the gum of the pine. This species is easily captured by the simple expedient of digging a hole in front of the entrance to the burrow, so that the animal when leaving its house immediately falls into the pit. Beauty of form the Tortoise has not, but the coloration of the horny plates of the back, in some species, is certainly striking. This is especially true of a small group known as " elegant " tortoises. The majority belong to S. Africa ; but perhaps the most beautiful of all is the "starred" tortoise (Testudo elegans) of India and Ceylon. All the members of this group are conspicuous for their extremely convex carapace. The horny covering shields are either black with bright yellow lines radiating from the centre of each, or yellow with black radiating lines. The effect of this black-and-gold scheme of coloration is, in many species, heightened by the elevation of the scales into prominent bosses. It might be imagined that such a plan of coloration would render the wearer extremely conspicuous. As a matter of fact, the reverse is the case ; for it has been shown that these tor- toises are very difficult to distinguish from the rocky ground of the grass jungles in which they live. According to an old Sanskrit legend, the world is supported on the back of a gigantic elephant, TORTOISES AND TURTLES. 35 whose feet are planted on the back of a still more gigantic tortoise. What the tortoise took its stand upon the legend does not so much as hint at ! This old legend, like so many legends, has a central particle of solid truth, inasmach as large species of tortoise occur both in Europe and North America, as far back as the Eocene, one species, indeed, from the Siwalik Hills of India having a carapace nearly six feet long. These ancient giants have long since ceased to exist on the continents either of the Old or New World. They were, says Dr Gunther, "unable to survive the changes of climate in the northern latitudes, or to coexist with the large carnivora, and especially with man, in the more congenial south. But there were two spots on the earth's surface where they continued to flourish to within a century or two of our time — viz. Mada- gascar and the neighbouring islands of the Western Indian Ocean, and the Galapagos Archipelago in the easternmost part of the Pacific." They do not appear to have lived in Mada- gascar within historic times, having probably been cleared off from the inhabited parts of the island at the time when the first Europeans landed. But their skeletons occur in consider- able numbers throughout the island. In the islands north of Madagascar, however, these creatures dwelt in absolute security for ages, none of these islands, save the Comoro group, being inhabited either by man or large mammalia. As a consequence, with absolute freedom from enemies, they had nothing to do 36 THE STORY OF REPTILE LIFE. but to eat, to multiply, to grow in stature, and possess the land. Thus, not only the larger islands of the Aldabra group — the Seychelles, Keunion, Mauritius, and Rodriguez — but also the smaller islands became peopled in incredible numbers. The discovery of these island fastnesses by Europeans speedily brought about the downfall of these harmless creatures. Their vast numbers melted like the snow. Proving more wholesome and more toothsome food than turtle, every passing ship stopped to bear away as many as she could carry. Later, a still further drain upon their ranks was imposed by the settlement of naval and military forces, until speedily the supply became exhausted. Recourse was then had to importation from neighbouring islands, and we gather from the reports of the French India Company that in 1759 four small vessels were accordingly employed in bringing tortoises from Rodriguez to Mauritius. One vessel carried a cargo of no less than 6000, and altogether more than 30,000 were imported into Mauritius in less than eighteen months ! As a result the dawn of the nineteenth century witnessed the practical extinction of these wretched creatures on all the islands save the south island of the Aldabran atoll. Here a few stragglers still possibly remain, thanks to the rugged char- acter of the land. Of the tortoises of the Galapagos Islands much the same story must be told. At the beginning of the sixteenth century immense numbers existed in these islands. Now only three, Abing- TORTOISES AND TURTLES. 37 don, Albemarle, and Duncan Island harbour a few survivors. Such is the lamentable history of these helpless victims, as collected with infinite pains by Dr Gunther some five years since. Isolated by the submergence of the lower land, these tortoises were prevented from inter- breeding and the swamping effect of intercrossing, so that in time each group of islands, and in the case of the Galapagos, almost every island came to possess its own peculiar species. Of the many species of these Galapagos tor- toises which have been described, two only can be mentioned here. The first, and perhaps the most interesting of all, is Daudin's Tortoise, Testudo daudini, from the south island of Aldabra. A specimen recently in the collection of the Hon. Walter Eothschild was the largest living tortoise known. The length of its shell was 55 inches, or 67^- over the curve, and the weight 560 Ibs. The species known as Testudo abingdoni is peculiar on account of the thinness of its shell, which is extremely delicate. A curious feature about the carapace of these Galapagos tortoises, or at least of the majority of the species, is the great size of the opening of the front of the shell, which presents a cave-like appearance, very different from the narrow crescentic aperture of the typical tortoise. Water is hard to find in the islands where these tortoise live, and travellers have often found relief for their parching thirst in the fluid contained in the pericardium or membrane sur- rounding the heart. The naturalist Baur relates 38 THE STORY OF REPTILE LIFE. an instance where lie and his five companions, when thus suffering, found relief by killing one of the species known as Testudo vicinia, a native of Albemarle Island. They found no less than five cups of clear fluid in this receptacle. Simi- larly Darwin relates in his "Voyage of the Beagle " how that the contents of the bladder are also, under pressure, greedily drunk. The taste of the fluid is said to be bitter, whilst that of the pericardium is tasteless. The difference between the fresh- water or pond Tortoises and Terrapins and their cousins of the dry land are generally by no means such as would strike one at first sight. So closely do they resemble one another, indeed, that some surprise might naturally be expressed that such different environments should have effected so little change. The only difference between the land and aquatic forms appears to be in the form of the feet, the aquatic species having webbed feet, which may even become paddle-shaped. A more careful examination of one of these pond-tortoises would, however, reveal modifications which are obviously special adaptations to their peculiar mode of life. Besides the change in the form of the feet to facilitate movement through the water, special breathing organs have been developed to permit of prolonged submersion. Thus in certain " soft- shelled " tortoises of the sub-order Trionychoidea, the mucous membrane of the throat is beset with thread-like processes richly supplied with blood. These act like the gills of fishes, fresh water being constantly taken in through the mouth and passed over the delicate blood-filled pro- TORTOISES AND TURTLES. 39 cesses which exchange the carbon dioxide of the blood for oxygen. But there is this differ- ence between the breathing of the tortoise and that of the fish. In the former the vitiated water is expelled through the mouth, in the fish through one or more slits at the sides of the throat. The water breathing of these tor- toises is supplementary to that of the lungs, and is effected in a perfectly rhythmical manner some sixteen times per minute, by the move- ment of the hyoid or tongue bones. Other water tortoises, such as the European Pond- Tortoise (Emys orbicular is), and the Sculptured Terrapin of North America (Clemmys insculpata), have developed accessory breathing organs in the shape of thin-walled bladders which open into the hinder end of the gut. The inside of these bladders is richly ^supplied with blood- vessels which are bathed by a constant exchange of fresh water, the bladder being incessantly filled and emptied through the vent. If one of these tortoises is suddenly taken out of the water the contents of these vessels will be instantly squirted out. Although this habit is well known, the source of the water is gener- ally misunderstood, inasmuch as it is usually supposed to be the urine from the bladder. Yet another extremely interesting modification which the pond-tortoises have undergone is that displayed by a large number of species forming the sub-order Pleurodira. These all have ex- tremely long necks, and on this account ap- parently are unable to retract them so as to the head within the shell. Instead, they 40 THE STORY OF REPTILE LIFE. curve the neck round to the right or left bring- ing it and the head under the eaves of the "carapace," hence the name side-necked tor- toises. Of course in this way the whole of one side of the head and face is exposed. In most land-tortoises this would be a grave danger, but apparently in the aquatic regions the need for such effective protection is less urgent. When, probably owing to overcrowding, some of the more adventurous tortoises essayed to make a living by the margins of pools and shelving banks of streams, and eventually in the water itself, they apparently found it ex- pedient to exchange a vegetable for an animal diet, inasmuch as the water-tortoises of to-day are almost entirely carnivorous. To the car- nivorous habits we may trace further peculiarities in the development of new traits of character called forth by the very different nature of the food. Preying on other living creatures, often more active and highly organised than them- selves, their only chance of success is, not seldom, by cunning. An admirable instance of this is furnished by the Snapping Turtle or " Snapper " (Chelydra serpentina), one of the largest of the pond- tortoises. Fishes are decoyed within striking distance by the artful display of two temptingly worm-like filaments protruding from the tip of the tongue, the rest of the animal being con- cealed in the mud. The consequences of touching these are far more serious than treading on the tail of the Irishman's coat! Larger prey this diabolical monster captures by stealthily ap- TORTOISES AND TURTLES. 41 preaching under the disguise of an old rotten log. This disguise is afforded by the fresh water algse which grows luxuriantly on its shell, and on the mud which settles there. The Snapper measures more than three feet from the snout to the tip of the tail, and has a geographical range extending from the Canadian Lakes, east of the Rockies, through the United States to Central America. It is held in whole- some fear, even by man himself, on account of the severe bites which it is capable of inflicting, and is besides cordially hated for the destruction it causes amongst food-fishes and water-fowl. For this destruction, however, the Snapper pays a heavy toll, inasmuch as young Snappers are caught in considerable numbers by his arch- enemy man, for table purposes. Their capture is effected by a hook baited with pieces of fish ; but the tackle used must be of the strongest, for a hooked Snapper is not taken without a struggle. Only the young are esteemed, the adults being uneatable owing to the strong odour of musk which pervades the flesh. The "Alligator Turtle" (Macroclemmys tem- mincki), a very near relative of the Snapper, and very like it in size and appearance, is even more ferocious, perhaps we should say courageous. When danger threatens, instead of retreating within its shell it assumes a defiant attitude. Eaising itself on its legs, with open mouth, it throws itself upon its assailant with great spirit, shooting out the head as far as the long neck will allow, and at the same time throwing the body forwards, often with such impetus as to 42 THE STORY OF REPTILE LIFE. bring it to the ground should the object aimed j at be missed. The strength of the jaw is sur- j prising. One of these creatures has been known to bite a piece clean out of an inch plank. The cunning of the disguise of the Snapper is equalled only by one other pond-tortoise — the " Mata-mata " (Chelys fimbriata). The back of the shell in this animal, as in the Snapper, bears a close resemblance to an old sunken log, but the guise has been acquired by somewhat different means, large conical bosses, divided from one another by deep valleys, giving the appearance of rough bark, and thus taking the place of the algae on the shell of the Snapper. Like the Snapper, the Mata-mata has assumed a disguise in order to increase the effectiveness of a lure by which unwary fishes and other prey may be brought within reach of the mouth. The lure in the present case takes the form, not of brightly-coloured filaments from the tongue, but of ragged-looking flaps of skin projecting from the head and neck. The ear-flaps and the flaps of skin on the throat are kept in constant motion, and thereby attract the attention of passing fishes and other curious creatures which, drawing closer and closer, are at length brought near enough to the mouth to be suddenly engulfed by the inrush of water down the throat of the artfully concealed monster. The Mata-mata is a native of the rivers of Guiana and Northern Brazil, and is perhaps the most bizarre-looking of all its tribe. It is a really big tortoise, attaining a length of more than three feet when fully adult. TORTOISES AND TUETLES. 43 From the pond-tortoises we pass, by a very natural transition, to the Marine Turtles. In them we see the final results of adaptation to jin aquatic life. Whilst the general form of juhe body has undergone little or no change, phe limbs have become completely transformed iinto swimming paddles. In the fore-limbs the J3xtent of the changes is extremely marked, ^either in the fore nor the hind limbs are iligits any longer to be distinguished; in ac- bordance with their new functions in both limbs ohey are enclosed within a common skin, so fthat the once walking limbs have now become < • paddles," superficially bearing the strongest resemblance to the paddles of the old fish [lizards — the Plesiosaurs and Ichthyosaurs, to )e described later — the Penguins among the Ipirds; or the Whale tribe among the Mam- |nals. In all these cases, just as in the turtles, bhe paddles have been evolved by the modifica- tion of limbs originally used in quite other ways. [Another point of interest with regard to the paddles of the turtles is, that whilst those of Ihe fore-limbs were of great length, the hinder |)air were extremely short. The explanation of [his, of course, is obvious — the long paddles were lised as propellers, the short ones as rudders. IChe same applies also to the ancient Ichthyo- laurs, and to the modern whales. In the latter, Indeed, the hind-paddles have disappeared al- |ogether, the work of steering being undertaken >y the tail. In some other cases, to be discussed 'S we proceed, we shall find this arrangement xactly reversed — the hind-limbs developing at 44 THE STORY OF REPTILE LIFE. the expense of the fore, so that these became id time reduced to the merest vestiges. Ill-fitted as they are for a life ashore, yet the females, at any rate, are obliged to sojourn here awhile, at least once a year, when they come tq deposit their eggs. Other aquatic reptiles, such as the ancient Plesiosauria and Ichthyosauria.; seem to have avoided this necessity by retaining! the eggs within the body until they hatched — j that is to say, they were viviparus. Gigantic as some of the tortoises have become, I they are surpassed by their sea-dwelling cousin* the turtles, the species known as the Leathery! Turtle, sometimes weighing as much as a ton.j This animal, however, is like other giant forms, verging on extinction. We shall see indeed as] we proceed how often Nirvana has been achieved! by the reptile-people through the gateway oil over-growth. The Green-turtle or " edible-Turtle " (CMonA my das), and the Hawks-bill Turtle (Chelone imbril cata), in so far as the general shape of the body! and paddles is concerned, are extremely like the! Leathery Turtle just described, yet, as has been! pointed out earlier in this chapter, they are only! in a very remote degree related — their similarity! is due to what is called " convergence of de- velopment," and not to community of descent. It is this same convergence, this adaptation to environment, which has produced the community of likeness, which has been pointed out between such widely different forms as the turtle, the fish- lizards, and the whales. The Green-turtle is the species which plays TORTOISES AND TURTLES. 45