R.LYDEKKER FE ann R 1 aicanaishnnititi bl PSG egg ETL ee ihiopopewenne ‘pvienecs gonmnrenaealewee Fenanaecs ALBERT R. MANN LIBRARY NEW York STATE COLLEGES OF AGRICULTURE AND HoME ECONOMICS AT CORNELL UNIVERSITY aPATE DUE feta hice.) 8 (Gee Cornell University Library The original of this book is in the Cornell University Library. There are no known copyright restrictions in the United States on the use of the text. http :/Awww.archive.org/details/cu31924003022518 “KNOWLEDGE” SERIES. LIFE AND ROCK: a COLLECTION OF ZOOLOGICAL AND GEOLOGICAL ESSAYS. \ By R. LYDEKKER, B.A. Cantab, F.GS., F.ZS, AUTHOR OF ‘‘ PHASES OF ANIMAL LIFE” AND ‘‘ HORNS AND HOOFS”’3 JOINT- AUTHOR OF ‘‘THE STUDY OF MAMMALS” AND ‘( ’ MANUAL OF PALAONTOLOGY”; EDITOR AND CHIEF AUTHOR OF ‘‘THE ROYAL NATURAL HISTORY,” ETC., ETC. NUMMULITE. London: TITE UNIVERSAL PRESS, 326, HIGIT HOLBORN, W.C. 1894. A PREFACE. In presenting to the public the following series of essays, which have already appeared in serial form, the Author has but few words to say by way of preface. It will be seen from the chapters themselves that the greater portion of the work relates to Zoology—and especially its paleontological branch—while only a small section is devoted to Geology proper. While some of the zoological chapters treat of the natural history of particular groups or species of animals, others are more especially devoted to some of those problems connected with the evolution, development, and mutual relationships of animals which at the present day are attracting so much attention on the part of students of the science. Although necessarily somewhat technical in places, the Author hopes that the popular style, which he has ‘endeavoured to preserve throughout the work, will render it acceptable to those lovers of Nature who are too often repelled by the number of hard words they meet with in the books professedly written for their edification. HARPENDEN, March, 1894. XII. XIII. XIV. XV XVI XVII. XVIII. XIX. XX. XXII. CONTENTS. Elephants, Recent and Extinct Tusks and their Uses Moles and their Like Spiny Animals Parallelism in Development Toothed Whales and their Ancestry Whalebone and Whalebone Whales Runinants and their Distribution The Tallest Mammal Lemurs Armadillos and Aard-Varks The Oldest Mammals Crocodiles and Alligators The Oldest Fishes and their Fins Living Fossils The Extinction of Animals Protective Resemblance in Animus Sea-Urchins Nummulites and Mountains A Lump of Chalk and its Lessons A Flake of Flint and its IJIistory LIST OF ILLUSTRATIONS. African Elephant Skeleton of Mammoth Foot of Elephant Tooth of Mastodon Indian Elephant . Teeth of Elephants Skeleton of Mastodon Tusks of Babirusa Narwhal Skull of Chinese Water-Decr Skull and Tusk of Uintathere Cape Golden Mole Long-clawed Mole-Vole . Marsupial Mole . Common Porcupine Hedgehog Spiny Anteater Foot of dArtionyx Skull of Protoceras Bridled Dolphin Teeth of Killer Indian Porpoise . Pacific Black-Fish LIST OF ILLUSTRATIONS. Section of Skull of Greenland Whale Skeleton of Extinct Irish Deer Cheek Tooth of Antelope Bones of Foot of Ruminant Horns of Gazelle Skull of Sivathere Antlers of Red and Sambar Deer Giraffe Slender Loris Tarsier Three-banded Armadillo Pichiciago Six-banded Armadillo Ethiopian Aard-Vark Jaw of Stonesfield Mammal Jaw of American Jurassie Mammal Jaw of Triconodon Jaw of Plagiaulax Jaws and Teeth of Rat-Kangaroo Skull of South African Secondary Mamunal Side view of Skull of Crocodile Skull of Crocodile Skull of Extinet Crocodilian The Bichir An Extinct Ganoid Skeleton of Perch . Skeleton of Fin of Extinct Shark Skeleton of Extinct Shark Tooth of Extinct Lung-Fish LIST OF ILLUSTRATIONS, Shell of Pleurotomaria A Pentacrinid Skeleton of Northern Sea-Cow Leaf Butterfly Test of Common Sea-Urchin Test of Secondary Sea-Urchin Test of Paleozoic Sea- Urchin Test of Cretaceous Heart-Urelun A Nunmulite Flint Implement Cone of Flint xi PAGE, 154 158 168 178 186 189 190 191 197 210 212 LIFE AND ROCK: A COLLECTION OF ZOOLOGICAL AND GEOLOGICAL ESSAYS CHAPTER I. ELEPHANTS, RECENT AND EXTINCT. SSUREDLY of all the mammals now inhabiting this A earth elephants are those most justly entitled to the epithet “antediluvian,” since they remind us, far more vividly than any of their modern contemporaries, of the gigantic extinct mammals of various kinds which flourished in that latest epoch of geological history when man was but a comparatively new comer. A long acquaintance has, indeed, made us so familiar with the appearance of elephants that we are too apt to forget what altogether strange and uncouth creatures they really are. If, however, they had happened to be included among those animals which disappeared from the face of the earth before the historic period, and were known to us solely by their skeletons, there can be no doubt that they would be regarded as among the most remarkable of mammals. Moreover, if elephants were only known to us by their skeletons it would be more than doubtful if we should ever have attained a correct idea of their true form; since although the conformation of their jaws and teeth would clearly indicate that they must have had some very peculiar method of feeding, yet it would have required a very bold, not to say a very imaginative man to have conceived the idea that these creatures were furnished with that unique organ which we term the trunk or proboscis. At the present day, it need scarcely be said, there are but two living species of elephants, differing remarkably from one another not only in external characters, but also, as we shall notice later on, in the structure of their B 2 ELEPHANTS, RECENT AND EXTINCT. teeth; these two species being respectively confined to Africa, and to India and the adjacent regions. These two kinds of elephants are, however, merely the last survivors of a vast host of extinct forms, some of which were closely related to their living cousins; while others differed so markedly in the structure of their teeth as to have received the distinctive appellation of mastodons, although lephant. Dj 1.—Young African I] Fia. MASTODONS. 3 Fre. 2.—Skeleton of Mammoth in the St. Petersburg Museum. they are really nothing but very generalized elephants, These so-called mastodons carry us backwards to the middle portion of that_division of the Tertiary period of B2 A ELEPHANTS, RECENT AND EXTINCT. the earth’s history known as the Miocene; but when we have reached to that stage all below is dark as regards the elephantine pedigree. And it is, indeed, one of the most remarkable circumstances in paleontology that although we know that elephants belong to the great group of Hoofed or Ungulate Mammals, of which they form a well-marked division, yet we have practically no sort of knowledge of the many extinct forms which we presume must have connected them with Ungulates of a more ordinary type. Although the trunk and tusks of elephants form their most striking external features, yet it is not to these that the naturalist looks at first when inquiring into the true affinities and general structure of these animals, since these come under the category of specialized and acquired structures, which tell but little of an animal’s past history; he looks rather to the structure of the internal skeleton, which is always of especial value, as being that part of the organism which is usually alone preserved ina fossil state. Let us then first turn our attention to the skeleton of these animals, of which we may see examples in our larger museums. The most remarkable feature noticeable in such a skeleton is that the various long-bones of the limbs are placed almost directly one above another, so as to form nearly vertical columns of support for the body ; whereas in ordinary Ungulates, such as a horse or an ox, these bones are set very obliquely to one another. Moreover, as a similar vertical position of the limb-bones occurs in several old extinct Ungulates which are known to be of extremely primitive organization, we may take it that an elephant’s limbs are likewise of a primitive type. We have, however, further evidence in confirmation of this primitive structure. Thus elephants differ from all other living Ungulates in having five complete toes to all their feet (Fig. 3). Moreover, whereas in other living Ungulates (except the little hyrax) the bones of the wrist are not situated in vertical rows immediately over the metacarpal bones of the foot, but, on the contrary, cross and overlap one another, in elephants they have the former relation, with the single exception that the bone marked 1 overlaps the STRUCTURE OF FEET. 5 one lettered td to a certain extent. This difference may be illustrated by saying that if we were to take a hatchet and chop vertically up- wards between the third and fourth toes of an elephant’s foot there would be nothing to resist the passage of the blade till it reached the bones of the leg, whereas in all other living Ungulates, iy except the hyrax—the pig oS for instance — the blade could not pass through the wrist without cleaving solid bone. Again, where- as ordinary Ungulates Fie. 3.—Bones of the Left Fore Foot walk solely on the tips of of an Elephant, one-eighth natural their toes, and are thus size. The lettered bones are those «tgs : f the wrist or carpus, an termed digitigrade, while aioe ones the ae the bones of the toes them- }elow which are the bones of the selves are more or less toes. (After Osborn.) elongated, elephants walk on the soles of their feet im the so-called plantigrade fashion, and have very short toe-bones. Now, since all the large extinct Ungulates of the lower Eocene, or earliest Tertiary period, also have five-toed feet, very similar to, but still shorter and of even simpler structure than those of elephants, there can be no doubt as to the extremely primitive plan on which the entire limbs of the latter are constructed. As regards, therefore, its limbs and feet, an elephant may be said to be an essentially old-fashioned animal. If, however, we turn to their teeth we shall find that elephants are very far indeed from being of a primitive or old-fashioned type; the truth being that they are, on the contrary, very peculiar and specialized in this respect. The first and most obvious peculiarity in regard to their dentition is to be found in their tusks, which correspond to one of the pairs of upper front teeth in man, and 6 ELEPHANTS, RECENT AND EXTINCT. also to the single pair of such teeth in the Rodents (rats, hares, &c.). Moreover, these teeth, like the incisors of the Rodents, grow continuously throughout the life of the animal, owing to the cir- cumstance that the pulp-cavity at their base always remains open, and has a permanent con- nection with the soft structures of the gum. In our own teeth, on the contrary, the pulp-cavity closes at a cer- tain period, after which there is a total cessation of growth. These ever-growing tusks of the elephants are preceded in the young animal by a pair of small milk-tusks, with a closed pulp- cavity, which are shed at an early period of life. In both of the living species of elephant the tusks are con- fined to the upper jaw; but whereas they occur in both sexes in the African elephant, in the Indian species large permanent aS Fia. 4.—Last or Sixth Upper Molar Tooth of a Mastodon; half natural size. TUSKS AND MOLARS. 7 tusks are restricted as a rule to the male, and are not, indeed, invariably present in all individuals of that sex. In the female Indian elephant the tusks are usually very small, and soon shed. The primitive elephants, or masto- dons, frequently, however, had tusks in both the upper and lower jaws; and since these did not generally attain the huge dimensions which they reach in many true elephants, it is evident that in this respect the mastodons departed less from the ordinary type of mammals, where the front teeth are not greatly larger than the hinder ones, and those of the upper and lower jaws correspond with one another in size and number. Before leaving the subject of tusks, it may be mentioned that the ivory of which they are composed differs from the so-called ivory of other teeth in a manner which renders it always easy to determine whether a reputed ivory article is genuine. This peculiarity consists in the circumstance that a trans- verse section of a tusk exhibits a series of fine, decussating, curved lines radiating from the centre to the circumference, and forming curvilinear lozenges at their intersections. This remarkable structure is in fact precisely similar to the “engine-turning”’ on the back of a watch; and in an ivory knife-handle it should be distinctly visible at the butt. We turn now to the grinding or molar teeth, which present far more remarkable peculiarities. So peculiar, indeed, and unique are the structure and mode of suc- cession of the molar teeth of elephants, that they are often very imperfectly understood, even by those who have spent half their lives among these animals. For instance, we find the late veteran elephant-hunter, Sir 8. Baker, in his “Wild Beasts and their Ways,” making a statement in regard to elephants that ‘“ both the Indian and the African varieties have only four teeth,” whereas, as a matter of fact, every elephant has, in the course of its life, six molars on each side of both the upper and lower jaws. Instead, however, of all these teeth being in use at one and the same time, as are those of a cow or ahorse, in existing elephants there are never more than portions of two teeth in use at any one time, although, in the extinct 8 ELEPHANTS, RECENT AND EXTINCT. mastodons, portions of three may co-exist. In all elephants, the more anterior of these six teeth are smaller and of simpler structure than the hinder ones, and the whole series is protruded in an are of a circle from back to front, the larger hind teeth being pushed up and gradually coming into use as the small anterior ones are Fie. 5.—The Indian Elephant. MOLAR TEETH. 9 worn away and finally discarded. The hinder ones of these teeth (Fig. 6) are so large that while the front portion is being worn away, the back is still bedded in the gum. In the earlier elephants, or mastodons, these molar teeth are composed of a series of relatively low and widely separated transverse.ridges, more or less completely divided into inner and outer moieties, and with large open valleys between them. Moreover, in all the teeth, except the last, these ridges do not exceed four in number, although in the sixth tooth (Fig. 4) there may be as many as five or six. Such a molar tooth can be easily derived from the ordinary type of tooth presented by the molars of the pig, in which the crown carries four columns, severally placed at its corners. When a tooth like the one represented in Fig. 4 becomes worn down by use, the enamel coating each of the columns would be cut through, so as to expose a series of more or less trefoil-shaped surfaces of ivory, each surrounded by a ring of the hard enamel. And it will be obvious that a tooth thus constructed of alternations of substances of different degrees of hardness would act as an efficient millstone. Elephants do not appear, however, to have been by any means satisfied with this comparatively simple kind of tooth, for as we pass upwards in the geological scale we find that there has been a gradual increasing complexity in the structure of the molar teeth of these animals, this being due to a graduated increase in the height of their transverse ridges, accompanied by an increase in the number of the ridges themselves. There is, indeed, an almost perfect structural gradation, now known to exist between the mastodon tooth, repre- sented in Fig. 4, to the teeth of true elephants shown in Fig. 6. Both the latter examples are in a somewhat worn condition, but it will be readily seen that the lozenge- shaped surfaces of ivory, surrounded by enamel, in the tooth of the African elephant, correspond to the transverse ridges of the mastodon’s tooth. In the true elephants, however, the open valleys between these ridges (which have now assumed the form of tall, thin, and nearly parallel laminz) have been completely filled up by a third constituent of the tooth, known as the cement. The 10 ELEPHANTS, RECENT AND EXTINCT. grinding surface of the tooth of such an elephant con- sequently consists of a solid mass, made up of alternating vertical transverse layers of various substances, arranged Fic. 6.—Molar Teeth of Indian (a) and African (4) Elephant. In athe anterior half is worn, and the remainder unworn. Much reduced. (After Owen.) in the order of cement, enamel, ivory, enamel, cement; and since each of these constituents differs in hardness, it will be obvious that the millstone-like apparatus is now of a far more efficient type than it was in the mastodon. Moreover, since the crowns of the molars of the true elephants are very much taller than are those of the mastodons, it is evident that they will require a longer period of time before they become worn away, and that they will therefore allow a longer life to their owner. Tiven, however, among true elephants there is a con- siderable amount of difference in regard to the number and narrowness of the component plates of their molar teeth, and it will be seen from Fig. 6 that in this respect the African elephant departs far less widely from the mastodon than does its Indian cousin. Since the food of EVOLUTION. 11 the latter consists to a large extent of boughs and twigs, while that of the former is composed more of juicy leaves, fruits, and roots, the necessity of a more complicated masticating apparatus in the one than in the other is apparent. In modern elephants, the six molar teeth on either side of each jaw are all that are ever developed. This is, however, by no means the case with some of the earlier elephants, and with most of the mastodons. In these animals, when the second and third molars became worn out, they were succeeded vertically by much smaller teeth, from which we learn that the first three molars of the modern elephants really correspond to the milk-teeth of other mammals, which, as we all know, are succeeded vertically by some of the teeth of the permanent series. This succession of the teeth shows us, therefore, another point in which mastodons tend to connect modern elephants with ordinary Ungulates. We might go further and enter upon the consideration of some of the structural peculiarities presented by the soft parts of elephants. Enough has, however, been stated to show that while these animals have preserved a very ancient type of structure in their limbs, they have acquired a very special modification in the structure and mode of succession of their teeth. And it is highly probable that it is owing to this particular specialization that elephants have survived to our own day, while all the other plantigrade and five-toed primitive Ungulates have completely passed away; while it is almost certam that it is this feature alone which has enabled them to attain the gigantic bulk which forms one of their most striking features. In regard to their evolution, perhaps no group shows more clearly than that of the elephants how exceedingly important is the study of fossils to eluci- date the relations of existing animals. Had we only the two living species of elephants to deal with, we should never have had the least mkling of the manner in which they were related to other Ungulates, imperfect as our knowledge of the relationship still is. Moreover, from the distribution of these two species, it would have been 12 ELEPHANTS, RECENT AND EXTINCT. naturally inferred that elephants were creatures suited solely to tropical or sub-tropical climates. The discovery of frozen carcases of the mammoth—a species closely related to the Indian elephant—in the Siberian “tundras”’ shows, however, that this animal was suited to dwell in at least comparatively cold regions, although it is probable that the climate of Siberia was formerly not so rigorous as at present. To withstand the cold of these northern regions, the mammoth was protected by a coat of long reddish hair, beneath which was a shorter covering par- taking more of the nature of wool. Along the borders of the Arctic Ocean for hundreds of miles mammoth remains are met with in incredible quantities ; and it is still one of the puzzles of geology to account adequately and satisfactorily for the manner in which these creatures perished, and how their bodies were buried beneath the frozen soil before decomposition had begun its work, for it is hardly possible to believe that they lived in a climate so rigorous that their bodies would have been frozen on the surface of the ground immediately after death. Another rude shock to our common ideas of elephantine nature is afforded by the extinct elephants of Malta, which show us that gigantic size is not a necessary concomitant of the group; and that when the area in which a species dwelt was small, the size of the species itself was proportionately reduced. These little Maltese elephants were very closely allied to the livmg African species, but whereas “Jumbo’’ attained eleven feet in height, and wild specimens of the African elephant may be still larger, the smallest of the Maltese species was scarcely taller than a donkey. So small, indeed, are the bones and teeth of this species exhibited in the Natural History Museum, that it is sometimes difficult to convince people that they really belong to elephants at all. As regards their distribution, elephants and mastodons formerly roamed over the whole world with the exception of Australia; true elephants ranging over the whole of the Northern Hemisphere, while mastodons extended as far south in the New World as the confines of Patagonia. It isin the north-east of India, Burma, and the islands of DISTRIBUTION. 13. the Malayan region that the fossil elephants connecting the living species with the mastodons are alone found ; and it is thus probable that from these regions the true elephants migrated westward into Europe and Africa, while the mammoth in later times crossed from Asia into Alaska by way of Behring Strait. That the mammoth, which ranged from the Arctic regions to the Alps and Pyrenees, was a contemporary of the primeval hunters of Europe is now a well-established fact, but it appears that throughout the Old World mastodons had utterly died out before the advent of man. In the New World, however, the continuity between the old and the new fauna was more fully sustained, the Missouri mastodon having survived well into the human period, so that we have in this survival a good instance of the vast changes that have taken place in the fauna of the globe within what we may metaphorically call the memory of man, 14 TUSKS AND THEIR USES. CHAPTER II. TUSKS AND THEIR USES. Many mammals, such as the elephant, hippopotamus, and walrus, are furnished with one or two pairs of pointed conical or compressed teeth largely exceeding all the others in length, and to which the term “tusks” is usually applied. Lions and wolves likewise have two pairs of somewhat similarly enlarged teeth in the fore part of their jaws; and although these are proportionately smaller than in the animals above named, it will be obvious that in popular language it is difficult not to include them under the same general title. Using, then, the term “tusks” for all such enlarged simple teeth, it may be of interest to note the groups in which these attain their greatest development, and also endeavour to learn something regarding their uses. An especial interest attaches, indeed, to the subject, on account of the extreme beauty of many of these teeth, and also from the circumstance that it is these alone which yield the various descriptions of ivory. In a great number of instances such tusks comprise a pair in both the upper and lower jaws, which are situated immediately behind the front, or incisor teeth, and, from their marked development in the dog tribe, are scientifically designated canine teeth; the name “eye- teeth” being also not unfrequently applied to them in popular language. Tusks of this sort are characterized by the circumstance that the lower one on each side bites in front of the upper one ; while the latter is always the first tooth situated in the true upper jaw-bone, the incisors, or front teeth, heimg implanted in a more anteriorly situated bone known as the pre-maxilla. In any ordinary NATURE OF TUSKS. 15 carnivorous mammal, such as a lion or a wolf, the upper tusks are considerably larger than the lower, although neither project beyond the edges of the muzzle when the jaws are closed. If we examine such tusks in a dried skull we shall find that their roots (which, as in all tusks, Fie. 7.—Skeleton of Missouri Mastodon. are simple) are completely closed ; thus indicating that their growth ceased at a certain period of life. Moreover, we may notice that the upper and lower pairs of tusks do not abrade against one another to any marked extent, and that consequently their summits are only subject to the ordinary wear and tear necessarily undergone during use. In many other mammals the form and structure of these tusks is, however, very different. Take, for instance, a wild boar, in which the upper tusks are short and curved 16 TUSKS AND THEIR USES. upwards, while the larger and more slender lower pair abrade against their outer surfaces, and are thus worn to sharp, cutting edges. Obviously such tusks, if they were incapable of growth like those of the lion, would soon be worn away to mere stumps and become useless to their owners. To prevent this, the bases of the tusks remain permanently open (as shown in Fig. 10), and contain a soft pulp connected with the vascular structures of the jaw, in consequence of which the teeth continue to grow through- out life; their rate of growth thus keeping pace with that of the abrasion to which they are subject. Tusks may accordingly be divided into two classes, which we may designate hollow and solid. The open tusks referred to above are prevented from attaining any very great length by the abrasion of the lower against the upper pair, but in other cases, as in those of the remarkable pig of Celebes known as the babirusa (Fig. 8), no such abrasion takes place, and both pairs then attain enormous dimensions, projecting in this particular instance far above the upper surface of the head, and the down-curving points of the upper pair sometimes even penetrating the skull. A babirusa may in fact be compared to a wild boar in which one pair of tusks having been broken, the other cont‘nues to grow without any abrasion by wear; only that in these animals both pairs are thus developed. In the wild boar and its allies the tusks—more especially those of the 1ower jaw—are purely offensive and defensive weapons ; but it is hard indeed to imagine the use of those of the babirusa. Probably Mr. Wallace is right in regarding them as an ultra-development of organs originally useful, but which have now, from some reason or another, become of no functional advantage to their owner, and have thus, so to speak, run riot. The babirusa presents us with an instance of a mammal in which both pairs of tusks have acquired their enormous development owing to the cessation of the mutual attri- tion between those of the upper and lower jaws, character- izing all its allies. On the other hand, in the mastodons and elephants we have examples where the development ig normal, and either one or both pairs attain a vast size. TUSKS OF ELEPHANTS. 17 In both species of existing elephants, as in many of the mastodons (Fig. 7), only the upper pair of tusks is thus developed; but in other mastodons these organs were present in both jaws, the upper pair being, however, always much larger than the lower. In the pigs and babirusa the tusks, of which the upper pair are implanted in the true jaw-bone, correspond severally with those of Fie. 8.—Fore part of Skull and Tusks of Babirusa. the lion and the wolf, and are accordingly reckoned as canines. This, however, is not the case with those of the elephants and mastodons, which grow in great part from the pre-maxillary bone, and thus correspond with one pair of the front or incisor teeth of other mammals. Consequently, much as they resemble them in general appearance, the tusks of an elephant are not homologous with those of a pig or a babirusa. As we have entered Cc 18 TUSKS AND THEIR USES, in some detail into the structure of the tusks of tke elephants in the previous chapter, it will be unnecessary to recapitulate the facts here; although we may mention that in modern elephants these organs consist wholly of ivory, without any investing coat of enamel. The tusks of these animals, which belong to the hollow type, are the largest developments of dental structure to be met with in the whole animal kingdom. To a great extent they are weapons of defence and offence, but in the African species, where they are common to both sexes, they are also largely used in grubbing up roots and overturning trees ; while in an extinct species from India their length is so great that they must have quite ceased to be useful, and were probably an actual encumbrance. Another allied extinct animal, known as the dinothere, is unique in having a large pair of downwardly-bent tusks in the lower jaw and none in the upper; the use of these being very difficult to conjecture. A still more remarkable condition obtains in the hippopotamus, in which not only are the canine teeth developed into an enormous pair of hollow, ever-growing curved tusks in each jaw, but the central pair of lower front or incisor teeth are so enlarged as likewise to merit the title of tusks. These incisor-tusks—which thus cor- respond to the lower pair of the four-tusked mastodons— are hkewise of permanent growth, and project forwards from the front of the jaw in the form of two elongated cones. In thus possessing three pairs of tusks the hippopotamus is quite peculiar among animals. Largely employed for tearing up the erasses, on which these monsters feed, the tusks of the hippopotami are also most effectual offensive weapons. In the land carnivores, of which more anon, the tusks are always of the closed type; but in their aquatic ally, the walrus, we again meet with a huge pair of ever- growing tusks directed downward from the upper jaw. On comparing the head of a walrus with that of au elephant, most persons would say at once that the tusks of the two animals were homologous. In this, however, they would be wrong, since, as we learn from the con- TUSK OF NARWHAL. 19 dition in the young animal, those of the walrus are true canines, whereas, as we have seen, the tusks of the elephant are incisors. We have here, therefore, a well- marked instance of the parallel development of severally dissimilar structures to attain a marked general simi- larity. The tusks of the walrus, which in old animals attain a great length, are mainly employed in digging up molluses and crustaceans from the sand and shingle, and also, it is said, to enable their owners to clamber up on the ice. Although the whalebone-whales are entirely deficient in teeth, and many of the dolphins and their allies have these organs but poorly developed, there are two cetaceans which exhibit a most remarkable development of tusks. The first of these creatures is the well-known narwhal, of the Arctic seas, in which, as a rule, there is one huge spirally-twisted cylindrical tusk projecting from the left side of the upper jaw of the male, which continues to grow throughout life. Whether this solitary tusk is a canine or an incisor, is not very easy to determine ; but it is Fie. 9.—The Narwhal. (After True.) remarkable that its fellow of the opposite side generally remains concealed in the jaw-bone, like the kernel of a nut in its shell, while in the female both teeth are thus rudi- mentary. Occasionally, however, male narwhals are met with in which both the right and left tusks are developed ; and it is somewhat curious that in such cases the direction of the spiral in the two tusks is the same, instead of being, as in the horns of antelopes, opposite. Although narwhals have never been known to charge and pierce ships with their tusks, after the manner ‘of sword-fish, it is still uncertain whether these formidable weapons ( which may attain a length of from eight to nine feet) are normally c2 20 TUSKS AND THEIR USES. used for purposes of attack, or for procuring food. Doubtless, however, the narwhal’s tusk is of some use to its owner; but in another cetacean, known as Layuard’s mesoplodon, the tusks appear not only useless, but actually harmful. In the whale in question, which is a rare species from the southern seas, there is but one large strap-like tusk on each side of thie. middle of the lower jaw, both of these curving upwards and inwards over the snout, so as actually to prevent the mouth from opening to its full extent. The only possible use we could suggest of such a structure would be to prevent the creature dislocating its jaw by yawning; but as other animals manage to get on without such an arrangement, this is scarcely likely to be a solution of the problem. It is more probable, indeed, that we have here to do with another instance of ultra, or monstrous development. Other examples of hollow or permanently-growing tusks occur among the hoofed mam- mals, other than the pigs, in all of which these teeth are found only in the upper jaw, and are developed chiefly or solely in the males. Among recent forms these tusks attain their greatest development in the little musk- deer of the Himalaya, where they are frequently over three inches in length, and project considerably below the lower Fie. 10.—Extremity of the jaw. In form they are sabre- Skull of a young Chinese like, and recall the upper tusks Water-Deer, withthe base of the feline carnivores, only o ie ces being more slender, and grow- ae “ine permanently. Similar but smaller tusks are met with in the Chinese water-deer (Fig. 10), in the Indian muntjac, and the little deer-like animals known as chevrotains. The latter belonging to a totally distinct group from the others, it is evident that these scimitar-like tusks have been inde- pendently acquired in the two groups; while it is quite TUSKS OF UINTATHERES. 21 probable that those of the musk-deer and Chinese water- deer are likewise of separate origin. With the exception of the muntjac, in which they are very small, all these deer- like animals are devoid of antlers; and it is thus evident that their tusks have been developed in lieu of those weapons. It is true, indeed, that the males of some of the antlered deer have small tusks, but these are of no use for offensive purposes, and are evidently organs in process of degeneration. Moreover, in the hollow-horned ruminants, such as oxen and antelopes, where the horns are permanent and generally present in both sexes, not a vestige of tusks remains. Although the Asiatic rhinoceroses have procumbent tusks of considerable size in the lower jaw, none of the odd-toed hoofed mammals, such as horses and tapirs, have upper tusks of any size. In past times there were, however, in North America a group of somewhat allied creatures known as uintatheres, in which an enormous pair of upper tusks, somewhat like those of the musk-deer, was developed (Fig. 11). Like those of the latter, these tusks grew permanently, and they were also protected by a descending flange of the lower jaw, which was often deeper than in the figured example. As these animals had front teeth only in the lower jaw, we have here, therefore, another curious instance of the parallel development of similar conditions in totally unconnected groups. What use these creatures could have made of their tusks is, however, not very clear, as in the living condition they could have projected but little below the lower jaw, while they were too long to have been effectual when the mouth was open. These uintatheres were also noteworthy on account of having a number of bony projections on the top of the skull, which in life may have been sheathed in horn; and it is not a little remarkable that another extinct creature—Protoceras (Fig. 19)—belonging to the even-toed eroup of hoofed mammals had a skull with very similar bony projections and also similar upper tusks. Our last instance of animals furnished with per- manently-growing upper tusks is afforded by the reptilian class, in which the extinct South African creatures 22 TUSKS AND THEIR USES. described many years ago by Sir R. Owen, under the name of Dicynodon, are thus armed. In these reptiles, some of which attained gigantic dimensions, the jaws were mainly sheathed in horn like those of a turtle ; the single pair of tusks curving down- wards and forwards from near the middle of the upper one. Possibly these tusks were capable of being em- ployed when the mouth was open like those of lions or tigers; but otherwise it is difficult to see their use. Certain allied reptiles from the aforesaid formations, known as anomodonts, had a full series of teeth, with a large pain of tusks in the upper jaw, which may or may not have been rooted, but are exceedingly f like those of certain car- Fre. 11.—Extremity of the Skull Mivores. As these reptiles and Tusk of a Uintathere. are not the direct ancestors of mammals, we have thus evidence of the acquisition of tusks in two distinct classes. As regards solid tusks, or those in which the lower end is closed at a certain period coincident with the cessation of growth, almost the only mammals, save the rhinoceroses (where, as we have seen, there may be a forwardly-directed pair in the lower jaw) and some of the marsupials, in which they attain any marked development are the carni- vores. Among these, the maximum size of tusks at the present day is attained in the larger felines, such as the lion, tiger, and leopard. In these animals the tusks are, however, never so much elongated as to bar the front of the open mouth; while very frequently their tearing power is increased by the hinder cutting-edge being finely serrated. Their terrible effect in tearmg and GENERAL CONCLUSIONS. 23 rending the animals upon which these fearsome carnivores prey is too well known to need further mention. During the Tertiary period there existed certain carnivores nearly allied to the modern cats, but exhibiting a much greater development of the upper pair of tusks, with a corre- sponding reduction of those of the lower jaw. In these creatures, which are known as macherodonts, or sabre- toothed tigers, the upper tusks were greatly compressed and flattened, with serrations on one or both cutting-edges; their length in a species of the approximate size of a tiger being upwards of seven inches. As in the uintatheres, the anterior end of the lower jaw had a descending flange to protect the end of the tusk. Manifestly, such enormous weapons would completely bar the sides of the open mouth, and consequently they could not be used in the manner of the tusks of a lion or tiger. It is still more difficult to imagine that these animals could have struck with their tusks projecting from below the closed mouth ; and it would consequently seem that in this instance also the tusks attained a development which was harmful rather than advantageous—this conclusion being confirmed by the fact, for what it is worth, that the sabre-toothed tigers have become totally extinct, while their less specialized allies continue to flourish. Finally, as regards tusks in general, it appears that while in the land carnivores these are always canines, are present in both jaws, and have closed roots, in the other orders of mammals their development is apparently some- what capricious, while they are very frequently present in only one jaw—almost invariably the upper—and continue to grow permanently. Moreover, they are almost as frequently incisors as they are canines; so that tusks apparently similar may be in nowise homologous with one another. Never developed to any size in animals with large cranial appendages in the form of antlers or horns, tusks are frequently wanting in those lacking the latter. Primitively their use was undoubtedly as weapons of attack and defence, or to aid in procuring vegetable food ; but in many cases they have subsequently undergone a (frequently sexual) development beyond the needs of such 24 TUSKS AND THEIR USES. purposes, and are thus in this respect analogous to the antlers of many stags. In other instances, however—and this in all the groups in which they occur—they have undergone a still further semi-monstrous development, rendering them if not actually harmful, probably in some cases inconvenient to their owners. Lastly, the independent acquisition in closely allied or widely separated groups of mammals of tusks of very similar structure and appearance, shows how little reliance is to be placed on external characters as indicative of relationship. MOLES AND THEIR LIKE, 25 CHAPTER III. MOLES AND THEIR LIKE. Iv is probably well known to most of our readers that in the evolution of organized nature two great factors have constantly been working against each other—the one being the adherence to a particular type of structure, while the other is the adaptation to a special mode of life. The usual resultant of these two forces has been that, in any assemblage of animals specially adapted for a certain peculiar kind of existence, while internally its different members have preserved their essential structural peculiarities more or less intact, externally they have become so much like one another that it often requires the aid of the professed zoologist to point out their essential distinctness. Perhaps in no case is this adaptive similarity in external characters better displayed than among certain of the smaller mammals which have taken to a more or less completely subterranean burrowing existence, of which the common mole is the best known example. In the British Islands we have, indeed, only this one creature which has adopted this particular mode of life; and it is to this animal alone that the name “mole” properly belongs. Other parts of the world possess, however, several more or less closely allied animals to which the same name must clearly be also applied. If, however, we happen to have friends from the Cape, we may hear them applying the name “moles” to certain burrowing mammals from that district, which upon exami- nation would be found to differ essentially in structure both from the ordinary moles and from one another. Then, again, if we were to travelin Afghanistan or some of the neighbouring regions, we should meet with another mole-like burrowing animal to which we should likewise feel disposed to apply the same name, although it has not 26 MOLES AND THEIR LIKE. the most remote kinship with our English mole. Finally, the deserts of central South Australia are the dwelling- place of the “marsupial mole,” which, although mole-like in general form, differs from all the animals yet mentioned in belonging to the marsupial order. We thus arrive at the conclusion that in the popular sense the term “mole” now serves to indicate a number of widely different animals, whose sole or chief bond of union is to be found in their adaptation to a similar mode of life, and their consequent assumption of a more or less similar outward form. Hence, in order to avoid confusion, it will be necessary to prefix the epithet ‘“‘true”’ to those species which belong to the same family group as the “little gentleman in black velvet,” while the remainder must be designated by other distinctive epithets. It might have been thought that such an expanded application of the name “mole’’ was restricted to popular language. This, however, is not the case, as naturalists have found it convenient to adopt the names “ sand-mole,” ‘ golden mole,” “marsupial mole,” etc., as the distinctive titles of different members of this purely artificial assemblage of animals; and the reader will accordingly understand that when we speak of “moles and their like,” we merely refer to a similarity in habits, and a more or less marked external resemblance between the animals under consider- ation. The general bodily form of the common mole is so thoroughly well known and familiar, that the term “ mole- like”? has been introduced into zoological, if not into popular, literature as a definite descriptive epithet. Since it is perfectly obvious that this peculiar form is the one best adapted for the needs of the creature’s subterranean existence, no explanation is necessary why most of the other members of the assemblage have conformed more or less closely to the same type. We may especially notice the flat, tapering, and sharp-nosed head, passing backwards without any distinctly defined neck into the long and cylindrical body ; the comparative shortness of the limbs, and the immense strength of the front pair, which are placed close to the head, and have their feet expanded into TRUE MOLES. 27 broad, shovel-like organs. We shall also uot fail to observe the absence of any external conchs to the ears, and the rudimentary condition of the deeply-buried eyes. A long tail would also be useless to a burrowing animal, and we accordingly find this appendage reduced to very small dimensions ; while the close velvety hair is most admirably adapted to prevent any adhesion of earthy particles during the mole’s subterranean journeys. Equally well-marked adaptive peculiarities would also present themselves were we to undertake an examination of the mole’s skeleton. While the majority of the assemblage conform more or less closely to the true moles in appear- ance, there are others in which such resemblance is but slightly marked, if apparent at all, from which we may probably infer some minor differences in their mode of life. Proceeding to the consideration of the different groups of mole-like animals, it will be convenient to divide the mixed assemblage into insectivorous moles, rodent moles, and marsupial moles. The term “insectivorous moles,” it may be premised, does not primarily indicate carnivorous habits in the species thus designated, but merely refers to the fact that they are members of the order Insectivora. It would of course be impossible, not to say out of place, to attempt a definition of that order ; but it may be men- tioned that it includes small mammals, like shrews, moles, and hedgehogs, which differ from the rodents in not pos- sessing a pair of chisel-like teeth in the front of the jaws, and also have their molar teeth surmounted by a number of small sharp cusps. The insectivorous moles include not only our common mole (Talpa europea), but also many other species belong- ing to the same genus, as well as certain others which are referred to distinct genera, all of which, for our present purpose, may be collectively spoken of as true moles. Of these, two genera (Talpa and Scaptonyx) inhabit Europe and Asia, while the other three are North American ; Africa having no representative of the group. It may be well to mention that all the true moles have very broad naked hands, each furnished with five toes carrying lone flattened nails, in addition to which there is a sickle-like 28 MOLES AND THEIR LIKE. extra bone internally to the thumb. In burrowing, most of them throw up the well-known molehills at certain intervals from the tunnels driven in search of worms— their chief food. In addition to these trae moles, North America also possesses certain other species known as shrew-moles, which, while belonging to the same family (Zalpide), are distinguished by the absence of the sickle-like bone in the hand and the less expanded form of the bones of the upper part of the fore-limb. They are thus clearly seen to be less specialized creatures than our own mole, to which they closely approximate in general appearance. Although belonging to the insectivorous order, the mole- like creature represented in the accompanying figure indicates a totally different family group. If we were to examine the upper molar teeth of a common mole, we should find that they had broad crowns, carrying cusps arranged somewhat after the manner of the letter W. On the other hand, in the Cape golden mole (as the animal Fie. 12.—The Cape Golden Mole. here represented is termed) the corresponding teeth have triangular crowns carrying three cusps arranged in a V. f=) RODENT-MOLES. 99 Moreover, if we look at the fore-limb, we find instead of the five-fingered hand of the mole that there are but four digits, of which the lateral pair are small, while the two middle ones are enlarged and furnished with triangular claws of great power. As in the true moles, all external traces of ears and eyes are concealed by the fur; this latter, it may be added, having a peculiar golden-green metallic lustre, from which the name of the animal is derived. The golden moles, of which there are several species, are much smaller than our English mole, and are widely distributed in South Africa ; in which continent they, in conjunction with the under-mentioned sand-mole and its allies, take the place occupied in the northern hemisphere by the true moles. In tunnelling, the golden moles come so close to the surface as to leave a ridge marking their course. The true moles and the golden moles aftord us, therefore, an instance of two entirely distinct groups belonging to the same order having assumed a perfectly similar mode of life, and, con- sequently, having acquired a superficial general similarity in external appearance. With the rodent-moles, of which there is likewise more than a single group, we come to animals of a totally different order, which have assumed a mole-like form aud habits, and are popularly confounded with the true moles. In common with the other members of the order Rodentia, all these rodent-moles are characterized by the presence of a pair of powerful chisel-like incisor teeth in the front of each jaw, while their molars have broad and flattened crowns adapted for grinding. Moreover, instead of driving their tunnels in search of worms, these rodent-moles burrow for roots and bulbs. All of them have very small or rudimentary ears and eyes, large and powerful claws, and short tails. One of the best known of these rodent-moles is the great mole-rat (Spalax), ranging from south-eastern Europe to Persia and Egypt, in which the eyes are completely covered with skin; allied to which are the bamboo-rats (Rhizomys) of north-eastern Africa and Asia, distinguished by having minute uncovered eyes and small naked ear- 30 MOLES AND THEIR LIKE. conchs, and thus departing more widely from the mole type. In the ‘sandy soil of Eeypt the mole-rat constructs tunnels of great length in search of bulbs. In South Africa these forms are replac ‘ed by the huge sand-mole (Bathyergus), which attains a length of about ten inches; and also by certain smaller animals known as Georychus and Myose alops ; differing from the former by the absence of grooves in their incisor teeth. The sand-mole is commonly met with in the flats near the shore, while the smaller forms generally frequent land at a higher elevation. Sometimes, lowever, both are found together, and the country is then covered in all directions with hillocks precisely resembling those made by our English mole. Although the sand-mole has uncovered eyes, these are not bigger than the heads of large pins, and can have but little visual power. Still, however, their presence serves to indicate that these animals have not become so completely adapted to a subterranean life as is the common mole, and this is confirmed by the fact that if their burrows are opened, the sand-moles after a few minutes usually protrude their noses from the aperture with a view to discover the cause of the disturbance, whereas an ordinary mole would under similar circumstances remain below. All the foregoing belong to one family of rodents; but in addition to these certain members of the vole group (a sub-division of the Wuridw) have also taken to a subter- ranean burrowing life, with the assumption of a mole-like hodily form. These may be termed mole-voles, and range from Russia to central and northern Asia, where they are represented by the two genera Hllobius and Siphneus. ‘They all have mole-like heads and bodies, short limbs and tails, rudimental external ears, very minute eyes, and powerful fore-paws. In the Russian mole-vole (Ellobins) and the allied Quetta mole from Afghanistan the claws of the front paws are short; but, as shown in our figure, they become oreatly elongated in the members of the genus Siphneus. All of ‘bean agree with the ordinary Peg in the peculiar structure of shale molar teeth, which consist of a number of triangular prisms placed edge to edee; and all are described as driving subterranean tunnels and throwing RODENT-MOLES, 31 out at intervals heaps of earth precisely after the fashion of the common mole. The foregoing are the only rodents which have assumed amore or less distinctly marked mole-hke external form while retaining all the characteristic structural features of Fie. 13.—The Long-clawed Mole- Vole. the order to which they belong. There are, however, two other members of the same great order which, while having acquired mole-like habits, have not assumed a distinctly mole-like form. One of these is the tuco-tuco (Ctenomys) of South America, belonging to the same great family as the capivara and the coipu. This animal is rather smaller than a rat, with a relatively shorter tail, pale grey fur, and red incisor teeth. Its general form is also not unlike that of a rat, the limbs being of fair length, and the front paws not markedly enlarged, while the eves are of considerable size. The external conchs of the ears have, however, been greatly diminished in size. The tuco-tuco derives its name from its voice, which resounds day and night from its subterranean dwellings, and is compared by Mr. W. H. Hudson to the blows of u hammer on an anvil. 32 MOLES AND THEIR LIKE. It frequents loose and sandy soil, although occasionally found in moist heavy mould, through which it pierces its way as readily as the mole. Darwin, who states that the tuco-tuco is even more subterranean in its habits than the mole, was told by the Argentines that blind examples were often captured. This, however, is not the experience of Mr. Hudson, who lays stress on the relatively large size of the creature’s eyes. From the soft nature of the soil in which it tunnels, it is not difficult to understand why it has been unnecessary for the tuco-tuco to assume a mole-like bodily form; but the reason for the retention of fully- developed eyes—which we should have thought exceedingly prone to injury—is hard indeed to divine. The other rodents with mole-like habits are two tiny little creatures from the sandy districts of Somaliland, locally known by the name of farumfer, and scientifically as Heterocephalus. They are about the size of a mouse, with large heads, moderately long tails, long powerful fore-feet, no external ear-conchs, minute eyes, and the whole skin naked, save for a few sparse bristly hairs. About as ugly a creature as can well be conceived, the farumfer, if clothed with a thick coat of fur, would be not very unlike a rather long-tailed, long-limbed, and narrow- handed mole. For tunnelling beneath the hot sand of the Somali desert the naked skin of the farumfer is most admirably adapted; and as the creature is allied to the South African Georychus, it may be recarded as the member of this group most specially adapted fora subterranean existence. Mr. E. L. Phillips, who was the first to observe these curious rodents in the living state, writes that they threw up in certain districts groups of elevations in the sand which may be compared to miniature volcanic craters. When the animals are at work, the loose sand from their tunnels is brought to the bottom of the crater and sent with considerable force into the air with a succession of rapid jerks, the rodents themselves remaining concealed in the shelter of their burrows, from which they appear never to venture forth. The extensive assemblage of pouched or marsupial mammals contains groups corresponding to several of MARSUPIAL MOLE. 33 those of the placental or higher mammals. Thus, for instance, the kangaroos in Australia play the same réle as the deer and ruminants in other parts of the world, while the Tasmanian wolf takes the place of the ordinary wolf, the wombats act the part of the marmots, the phalangers of squirrels, and the bandicoots of the civets and weasels. Till recently, it was thought that the place of the moles (whether insectivorous or rodent) was unoccupied in the Antipodes, and that no marsupial had adapted itself to a tunnelling subterranean existence. Within the last few years it has, however, been discovered that the sandy deserts of south Central Australia are inhabited by a small burrowing creature belonging to the pouched group, which has been fitly termed the marsupial mole (Notoryctes) ; Fia. 14.—Under-surface of the Marsupial Mole, two-thirds natural size. (After Stirling.) and it is not a little remarkable that in general appearance this tiny animal is even more mole-like than are some of the above-mentioned burrowing rodents, thus showing how all-powerful is adaptation to environment, and of how little import is internal structure in modifying the external form of an animal. The general mole-like appearance of the Australian burrower will be apparent from the accom- panying figure; the most striking mole-like features being the elongated and depressed body passing imperceptibly into the bead, the absence of external ear-conchs, the rudimentary pin-like eyes, the small tail, and the short limbs, of which the front pair are armed with claws of D 34 MOLES AND THEIR LIKE. great power. The marsupial mole stands alone, however, in having the front of the muzzle protected by a leathery shield ; while its short and blunt tail is also covered with a peculiar naked leathery skin. In the fore limbs the structure of the feet recalls the golden moles rather than the true moles, the third and fourth toes being greatly enlarged at the expense of the others, and furnished with huge triangular claws of enormous digging power. In its pale, sandy-coloured hair, with a more or less golden tinge, the marsupial mole departs widely from our sable European friend; but it must be remembered that the difference in this respect is really not so great as it at first sight appears, seeing that cream-coloured varieties of the common mole are far from rare. In the Australian form the pale coloration is doubtless adapted to harmonize with the natural surroundings of its native desert, since it has been ascertained that the creature makes its appearance from time to time above ground. That the resemblance of the marsupial to the golden mole in the structure of the fore-paws is a purely adaptive one, there can be no reasonable doubt; but whether the identical structural conformation of the molar teeth of the two animals indicates any real genetic affinity, or is merely inherited from an old ancestral type which may have been common to many groups, is far less easy to answer. The marsupial mole, or ur-quamata, as it is termed by the aborigines, inhabits a very limited area lying about a thousand miles to the interior of Adelaide; and even there appears to be of extreme rarity. According to observations supplied to its describer, Dr. E. C. Stirling, the creature is generally found buried in the sand under tussocks of the so-called porcupine-grass (T'riodia), and its food appears to consist of insects and larve. The animals appear only to move about during warm, moist weather ; and as they are extremely susceptible to cold, it is probable that they lie in a more or less torpid condition during the winter months, when the surface of the ground is often white with frost. When on the move, the marsupial mole is said to enter the sand obliquely, and to travel for a few feet or yards beneath the surface, when it emerges, and after CONCLUSIONS. 35 traversing a short distance above ground, once more descends. As it seldom tunnels at a depth of more than two or three inches below the surface, its course may often be detected by a slight cracking or movement of the surface as the tunnel proceeds. Both in this respect, and in the nature of its food, the ur-quamata therefore exhibits a further analogy with the golden mole. In burrowing, the leathery shield of the head is said to be brought into play as a borer in the soft sand. As regards the advantages obtained by those mammals which have taken to a burrowing subterranean existence, it will be manifest that these are twofold. In the first place, the creatures are secure from all foes, except those which, like the weasel and the snake, are able to follow them into their underground labyrinths ; while, secondly, they tap a food-supply (whether animal or vegetable) inaccessible to most other animals. In the case of the mole at least, whose habitations are generally made in comparatively hard ground, the life must be an incessant round of labour; and to our thinking, at any rate, the existence of all these burrowing creatures must be so dull and monotonous as to leave no question as to whether it is worth living. 36 SPINY ANIMALS. CHAPTER IV. SPINY ANIMALS. In the preceding chapter it was shown how the adapta- tion to the necessities of a particular mode of life has produced a marked general external resemblance in certain burrowing mammals belonging to several more or less completely distinct groups. We now propose to point out the resemblances existing between certain other members of the same class of animals, owing to the assumption of a protective coat of spines. Although this resemblance is in some instances not so striking as among the creatures noticed in that chapter, yet it is quite sufficient to have obscured in popular estimation the real affinities of some of the spine-bearing mammals, as it is by no means uncommon to hear the hedgehog spoken of as the ‘British porcupine,’ while certain Madagascar spiny mammals are frequently alluded to as hedgehogs, and the Australian echidna is commonly alluded to as a porcupine. Moreover, the names “ sea- urchins’’ and “sea-hedgehogs,”’ applied to animals belonging to totally different classes, shows the important estimation held by spines in popular zoology. It is almost superfluous to add that the acquisition of the coat of spines in all the mammals here alluded to is solely for the pur- pose of protection : and how sufficient is this protection in most cases, is evident to all who have seen how the hedgehog, when rolled up, sets most dogs at defiance. Still, however, this panoply is by no means invariably proof against all attacks, as it appears to be well ascer- tained that leopards and pumas will kill and eat porcupines without the shghtest hesitation, and with a total disregard of their formidable spines, which may be found sticking in all parts of the bodies of the devourers. As we found to be the case with the mole-like mammals, all the spiny mammals belong to the lower orders of the class, their THEIR VARIOUS KINDS. 37 several representatives being distributed among the in- sectivores, rodents, and egg-laying groups, and the majority pertaining to the two former of these. In fact, in this respect an exact parallelism may be drawn between the mole-like and the spiny mammals, each assemblage having several representatives among the insectivores and rodents, while the former has a solitary marsupial type, and the latter two members among the egg-laying mammals. Some of these spiny mammals, such as the true porcupines and the echidnas, are burrowing creatures, Fig. 15.—The Common Porcupine.* and thus have a double means of defence against their enemies ; others, however, like the hedgehog, rely on their power of rolling themselves up into a ball, and thus pre- senting a chevauz-de-frise on all sides. Some again, like the tree-porcupines, are more or less completely arboreal in their habits; and the whole of them, like the mole-like mammals, indicate how urgent has been the need for the lowly-organized rodents, insectivores, and egg-laying * We are indebted to Messrs. F. Warne and Co. for the loan of the three figures illustrating this chapter. 38 SPINY ANIMALS, mammals to acquire some special means of protection in order to be able to hold their own among the higher forms. As our readers are doubtless aware, in mammals spines are nothing more than specially modified hairs, and in a poreupine the transition from a spine to an ordinary hair can be easily seen. There are many rodents in which a certain number of scattered spines are mingled with the fur of the back, but our remarks will be confined in the main to the forms in which the spines predominate suffi- ciently to render them the most striking feature in the external appearance of their possessors. ; Commencing with the rodents, our first representatives of the spiny mammals will be the true porcupines (Hystriz), which are such well-known creatures as to require but brief description. These animals conform, of course, to the ordinary rodent type in having a single pair of large chisel-like incisors in each jaw ; and their spines are most developed on the middle line of the head and back, the hinder part of the body, and on the short tail. Whereas, however, those on the body are solid throughout and pointed at each end, the spines at the extremity of the short tail are in the form of hollow quills inserted by narrow stalks. It is these hollow quills that make the loud rattling sound heard when a poreupine is walking ; and it appears to us not improbable that they may have given rise to the old legend of the porcupine ejecting its spines whev attacked, as such hollow quills might well have been thought to be receptacles for the ordinary spines. Although their owner is unable to voluntarily eject the latter, their pointed bases render them easily detached, and leopards which habitually feed on porcupines are found to be actually bristling with their quills. In attack- ing its foes, the porcupine rushes at them backwards, and thus gives full effect to its weapons. All the members of the typical genus are characterized by their large size, short tails, and highly convex skulls, and are confined to the warmer regions of the Old World. The brush-tailed porcupines (Atherura) from West and Central Africa and the Malayan region are, however, of much smaller size, and also distinguished by their much longer tails, which PORCUPINES AND SPINY MICE. 39 terminate in a brush of flattened spines, and are thus evidently less specialized creatures. America is tenanted by a group of porcupines easily distinguished from their Old World cousins by having the soles of their feet covered with rough tubercles, instead of being perfectly smooth, and also by their comparatively short spies being mingled with a number of long hairs, by which they may be partially concealed. The Canada porcupine (Hrethizon) ditters from all the other American species in having a short stumpy tail, and also in its non- arboreal habits ; its spines being almost hidden by the hairs. In parts of North America these porcupines are so abundant as to be a positive nuisance, and an enterprising engineer, with true American “cuteness,” hit upon the original idea of utilizing their bodies as fuel for his engine —apparently with the most satistactory results. The lighter-built tree-porcupines (Synetheres), which are mainly characteristic of the southern half of the American conti- nent, are easily distinguished by their long tails, which, as in so many South American mammals, are prehensile. These porcupines are thoroughly arboreal in their habits, and it is therefore easy to understand why their spines are so much shorter than those of their terrestrial Old World cousins, who have to rely solely on these weapons for their protection. In addition to the members of the porcupine family, there are several other groups of rodents which develop a more or less complete coating of spines. Among the most remarkable of these groups are the spiny mice (Acomys) of Syria and Eastern Africa, one of which, when it has its spines erected, is almost indistinguishable at the first glance from a diminutive hedghog. The spiny rat of Celebes (Echinothrix) is another member of the mouse family having the fur thickly intermingled with spines. In a third rodent family (Octodontide), nearly all the members of which are South American, there is also a genus (Hchinomys), taking its name from the number of flattened spines mingled with the fur of the back charac- terizing all its representatives. It will thus be obvious that even in a single mammalian order we have several 40 SPINY ANIMALS. instances where a protective coat of spines must have been acquired quite independently. te This independent origin is still more clearly indicated when we come to the consideration of the hedgehog and its Sr eer ar sce ea Nass Fic. 16.—The Hedgehog. allies, which bear precisely the same systematic relation- ship to the poreupines as 1s presented by the true moles to the mole-voles, as described in our last chapter. That is to say, whereas the hedgehogs and true moles belong to the insectivorous order, the porcupines and the mole-voles are herbivorous rodents. In spite, then, of the general similarity of appearance between a hedgehog and a porcu- pine, or, still better, a spiny mouse, we shall find, as already mentioned, that whereas the two latter have the ordinary chisel-like rodent teeth, the former has several narrow and somewhat irregularly-shaped teeth in the front of the jaws, while its back teeth are crowned with numerous sharp cusps, instead of having nearly smooth grinding surfaces. Accordingly, from the purely systematic point of view there is no justification for calling the HEDGEHOGS. 41 common hedgehog the “ British porcupine”; but, on the other hand, if we allow similarities in external appearance to be our guide in nomenclature, there are just as good grounds for applying the latter title to the hedgehog as there are for giving the names of golden or Cape mole, sand-mole, mole-vole, and marsupial mole to four of the creatures noticed in the last chapter. Our ancestors, to whom the hedgehog was commonly known as the “ urchin,” went, however, a step further than this, and, from the resemblance of its spine to those of the mammal, gave the name of sea-urchin to the Hehinus, a title which has stuck to it ever since. Systematic zoologists need not then wax so wroth as we have known them do when the name of “ British porcupine” is applied to the urchin, seeing that the analogies of nomenclature are sufficient to justify its use. As the sea-urchins come most dis- tinctly under the title of spiny animals, it may be mentioned here that, although the spines of the common British species are net unlike those of the hedgehog, yet their structure is totally different. Thus, whereas the spines of the mammal are of a horny nature, those of the invertebrate owe their solidity to the presence of carbonate of lime, and always break with the characteristic oblique fracture of the mineral calcite. Moreover, whereas the spines of the hedgehog are implanted in its skin, those of the sea-urchin are entirely external, being movably attached by their hollow bases to knobs on the surface of the shell or “test.” Whether our worthy ancestors believed that the land and sca-urchins were connected by ties similar to those which in their estimation affiliated barnacle-geese to barnacles, or how the name “urchin”? came also to indicate a child, we are quite unaware. In place of terminating in sharp points, by which they are but loosely attached to the skin, like those of the por- cupine, the spines of the hedgehog terminate interiorly in small knobs, which are placed beneath the skin, and may thus be compared to pins stuck through a piece of soft leather. Beneath the skin lies a layer of muscle known as the panniculus carnosus; and it is by the action of this 42, SPINY ANIMALS. muscle on their heads that the spines are raised from a recumbent to a vertical position when the creature rolls itself up into a ball—an action of which all porcupines are quite incapable. Not only does the hedgehog differ from the porcupine in this respect, but it is likewise peculiar in using its spines as a means of protection when throwing itself down a vertical bank or precipice, and by this means is able to accomplish a vertical descent of over a dozen feet without the sightest harm. As regards the develop- ment of its spiny armour, the hedgehog is perhaps the most highly specialized of all the spiny mammals, in spite of the inferior length of its spines as compared with those of the porcupine. Hedgehogs are now represented by about a score of species ranging over Europe, Africa, and a considerable portion of Asia; while the existing genus dates from the middle portion of the Miocene division of the Tertiary period. That the spines characterizing the existing forms have been independently developed within the limits of the group is pretty conclusively indicated by the close affinity of the hedgehogs to the long-tailed and spineless Malayan insectivores known as gymnuras; fossil types apparently indicating an almost complete transition from the gymnuras to the hedgehogs. In case anyone should suggest that the latter, and not the former, might be the ancestral stock, it may be mentioned that while the gymnuras have a generalized type of dentition and long tails, the hedgehogs have the teeth much reduced in number and specialized in character, and short tails. In Madagascar the place of the hedgehogs is taken by an entirely different group of insectivores known as the tenrecs, all of which have a certain number of spines mingled with the fur, at least in the young condition, and some of which are so hedgehog-like in general appearance that by the non-zoological observer they would certainly be regarded as members of the Erinaceidee. The tenrecs differ, however, from the hedgehogs pre- cisely in the same manner as the golden moles differ from the true moles. That is to say, whereas in the latter the crowns of the upper molar teeth are quadrangular, with their cusps arranged in a somewhat TENRECS. 43. W-like manner, in the former these teeth are triangular, with their cusps arranged in a V. There are five species of tenrecs, classed under three generic headings, and all characterized by the absence or small size of the tail. The largest, and at the same time the most generalized of all, is the common tenrec (Centetes ecaudatus), which attains a length of from twelve to sixteen inches, and is characterized by the absence of a tail, by the rows of spines on the back being shed in the adult state, and also by certain peculiar features in the dentition which appear to indicate relationship with the pouched mammals. The spiny tenrecs ( Hemicentetes) are much smaller avimals, of the size of moles,in which the longitudinal rows of spines on the back are retained throughout life. They have the same number of teeth as fully adult individuals of the common tenrec; but whereas in the latter there are four upper molars and two upper incisors, in the spiny tenrecs there are three of each of these teeth. In the loss of the last molar these tenrecs are evidently more specialized than the common species, but the presence of the third incisor shows that they are descended from a still more generalized type. Lastly, we have the hedgehog-tenrecs (Ericulus), in which the whole upper surface of the body, as well as the short tail, is thickly beset with spines, thus giving the hedgehog-like appearance from which the creatures derive theirname. The dentition is more reduced than in either of the upper groups, thus indicating the greater specialization of the genus; although the presence of a short tail indicates direct descent from a tailed ancestor. Although it is quite clear that the three genera of tenrecs are divergent branches from a common stock, yet it is not impossible that they may indicate the manner in which the complete coat of spines characterizing the third group has been gradually evolved. Against this view it may, however, be urged that if the common tenrec indicated the first commencement of the spiny coat, it would be more likely to find the spines in the adult rather than in the young. Be this as it may, the wide difference between the hedgehogs and the tenrecs, coupled with the affinity of the former to the gymnuras, leaves no doubt 44 SPINY ANIMALS. that the spines have been acquired independently in the two groups. As we found the last representative of the mole-like animals among the pouched mammals, so we observe that spiny animals are represented among the still lower egg- laying mammals by the spiny anteaters, or echidnas, of Australia and New Guinea. The two kinds of echidnas differ, however, externally from all the spiny animals hitherto mentioned in the production of the muzzle into a long, toothless tubular beak; and their spines are short, and in some cases largely concealed by the fur. Neither of them have the power of rolling the body into a ball; and, whereas the Australian echidna has five toes to each EVOLUTION OF SPINES. 45 foot, in the Papuan species the number is generally reduced to three. The wide structural differences separating the egg-laying mammals from all other members of their class render it almost unnecessary to observe that the spines of the echidnas are an entirely independent development. Like so many of the spined mammals, the echidnas have extremely short tails and thick bodies, with the neck indistinctly marked. We have thus decisive evidence that a more or less complete coat of spiny armour has been independently acquired in the following groups of mammals, viz., in the porcupines, mice, and octodonts among the rodents; in the hedgehogs and tenrecs among the inmsectivores; and in the echidnas among the ege- layers. From the perishable nature of these appendages we have, unfortunately, no evidence as to the existence of spines among fossil mammals; but from the foregoing considerations, we are strongly inclined to think they may be mainly characteristic of later epochs. In this connection it is interesting to notice that the spiny globe-fishes (Diodon, &c.), often termed ‘ sea- hedgehogs,” in which the spines are bony and therefore capable of preservation, do not date back below the Tertiary, and that spiny fishes are unknown in earlier epochs. Moreover, some—although by no means all—of the paleozoic sea-urchins appear to have had very minute spines. Hence it would rather seem as though the history of spines has been exactly the opposite of that of bony armour, which has tended gradually to disappear with the advance of time. Finally, we have to notice the general similarity in appearance of so many of the more specialized spiny mammals, due not only to their bristly coat, but likewise to the general shortness or absence of the tail, and the rounded, plump form of the whole body. The bearing of this independent development of spines in so many groups of mammals, together with the acquirement of a general external resemblance in the creatures thus clothed, on questions of wider import, forms the subject of the next chapter, in which we shall also have to take into considera- tion the conclusions reached in the previous one. 46 PARALLELISM IN DEVELOPMENT. CHAPTER V. PARALLELISM IN DEVELOPMENT. In the course of the three preceding chapters, it has been to a great extent our aim to show. that certain animals may resemble one another very closely in general external appearance, or may possess certain peculiar structural features in common, without being in any way intimately related; thus rendering it evident that such similarities of form or structure have been independently acquired, and are not inherited from a common ancestor. It has been shown, for instance, that mammals belonging to several distinct orders, or to different families of the same order, may assume such a marked external resemblance to the common mole as to be designated in popular language by the same general title; while in other cases a more or less striking approximation to the type of the ordinary hedge- hog has resulted from the independent development of very similar spines in totally distinct groups. Furthermore, it has been pomted out in the first of the three chapters that large tusks of very similar form may be independently developed in the jaws of totally different distinct groups of mammals; and even that certain extinct reptiles have acquired tusks which are almost indistineuishable from those of some of the carnivorous mammals, there being no direct relationship between the members of these groups. With regard to the external similarities of form in the above-mentioned instances, we have seen reason to believe that their inducing cause has been either similarity of habit or the need of protection; while in the case of the tusks the similarity may in certain instances be due to the necessity for efficient offensive weapons. Be their causes, however, what they may, it is evident that such resemblances among animals, which are, so to speak, accidental, indicate what may be termed a kind of parallel NATURE OF PARALLELISM. 47 development ; and as such parallelism in development, or shortly ‘‘ parallelism,” has recently attracted a good deal of attention among biologists, we propose in this chapter to present to our readers some of the more striking instances of this feature. In the instances above alluded to, the parallelism is either to a great extent shown in external characters, or in structures which are easily modified ; but, as we shall indicate in the sequel, in other cases it affects deeply-seated structures ; and its inducing cause is then very difficult to surmise on any of tbe ordinarily accepted doctrines of evolution. It will, more- over, be obvious that the acceptation of parallelism in development—and accepted to a certain extent it must undoubtedly be—throws a new difficulty in the inter- pretation of the affinities of animals, since before saying that an identity in some structural feature between the members of any two groups indicates their relationship, we have first of all to determine whether such similarity of structure is due to parallelism, or is inherited from a common ancestral type. As is so generally the case when any new theory is started, a host of enthusiastic writers have welcomed parallelism with acclamation, and have attempted to apply it im a number of instances where there is not at present sufficient evidence of its existence. We have been told, for instance, that the American monkeys have no relationship with their reputed cousins of the Old World; that whalebone whales are not allied to the sperm-whale and dolphins; that cats have no kinship with civets or other modern carnivores; and that the egg-laying mammals have been evolved from a reptilian or amphibian stock, quite independently of all other members of the mammalian class—their resemblances being solely due to parallelism. At present, we confess, we are totally unable to accept any of these conclusions, and we think it somewhat improbable that if the members of either of the above-mentioned pairs of groups had an independent origin, they would have presented such a similarity, both externally and internally, as we find to be the case. Still, however, it must be borne in mind that there is a considerable amount of evidence that the modern 48 PARALLELISM IN DEVELOPMENT. horses of the Old World and their extinct cousins of the New have been independently derived from earlier horse- like animals; and if this be confirmed, it will be one of the most remarkable known instances of parallelism, and will tend to show that many others may exist. Turning from these more or less problematical cases, we proceed to notice seriatim certain well-marked instances of parallel development as to the existence of which there can be no doubt; commencing with those displayed in external features, and then referring to such as are more deeply seated. As regards external resemblances, it is, of course—and more especially so far as the lower animals are concerned —not always easy to distinguish between parallelism and mimicry. The above-mentioned instances of mole-like and hedgehog-like animals belong, however, clearly to the former category. The external resemblances existing between swifts and swallows, which have no sort of relationship to one another, likewise comes under the same head. Another striking instance is to be found in the assumption of a snake-like form, accompanied by abortion of the limbs, by certain lizards, such as the familiar English “blind worm,” it being very doubtful, to our thinking, if this can be explained by mimicry. Still more remarkable is the external resemblance presented by whales and dolphins to fishes, and by the extinct fish- lizards (Ichthyosaurs) to both; such resemblances being clearly due to parallel development induced by the needs of adaptation to a purely aquatic life. If, moreover, it should prove that the whalebone whales have no connection with the dolphins, we should then havea far more remark- able instance of this feature, and one extending to internal structures as well as to external form. A similar remark will also apply to the case of the true seals and the eared seals, which it has been suggested may be of independent origin. The independent acquisition of wings by birds, bats, and the extinct flying dragons, or pterodactyles, may likewise certainly come under the heading of parallel development, so far as external form and the adaptation to a particular mode of life are concerned ; although here, IN TEETH. 49 as in the instance of whales and fishes, the structural features, by the aid of which the adaptation has been brought about, are very different in the respective groups. In respect to the teeth and dentition, many very well- marked instances of parallelism may be adduced. We have already referred to the similarity of the tusks in dif- ferent groups of mammals; while in a previous chapter we noticed the loss of upper front teeth, not only in the modern ruminants, but likewise in the peculiar even-toed ungulate known as Protoceras, of which the skull is figured on page 53. The rhinoceroses likewise show a gradual tendency to the loss of front teeth, resulting in the African forms in the disappearance of the whole series from both jaws. More remarkable, however, is the tendency to a complete loss of the whole of the teeth in certain groups of all the higher classes of vertebrates, as exemplified by all modern birds, by turtles and tortoises, as well as certain of the extinct flying dragons and fish-lizards among reptiles, and by the great anteater, the scaly anteater, and the echiduas, or spiny anteaters, among mammals ; in all of which teeth are completely lacking. Moreover, in many of these animals, such as birds, tortoises, and probably the toothless flying dragons, the beak-like jaws thus produced were sheathed in horn, thus showing a kind of double parallelism, viz., the loss of teeth coupled with the acquisition of a horny sheath to the jaws. Another instance of parallel development afforded by teeth relates to the gradual heightening of the crown and the production of a flat plane of wear not only among several distinct groups of hoofed mammals, such as the elephants, horses, and ruminants, but likewise among the rodents. It is evident that these high-crowned teeth have been independently evolved in the three great groups of hoofed mammals, of which the above-named creatures are typical representatives ; and it may be added that the molars of horses and ruminants present a further evidence of parallelism in their assumption of a more or less decidedly crescent-like (selenodont) structure, the essential peculiarities of which are described in the eighth chapter. This resemblance between the molars of horses and E 50 PARALLELISM IN DEVELOPMENT. ruminants is, however, comparatively remote, but in the latter croup and the camels these teeth are so alike as to require an expert to distinguish between them. Never- theless, there is good evidence to show that the camels and the ruminants, if not also the chevrotains, have acquired their crescent-like molar teeth quite independently of one another, and it therefore yet remains for those writers who explain evolution by some mode of what they are pleased to call natural selection to account adequately for the similarity thus existing between structures of such totally different origin, when they could have been made equally efficient if unlike. Passing from the consideration of teeth to that of limbs, we may mention the remarkable similarity displayed in the mode by which the lower segments of the limbs of the even-toed and odd-toed hoofed mammals have been gradu- ally elongated by the formation of a cannon-bone and the disappearance of either three or four of the lateral digits ; the cannon-bone in the horse consisting of but a single element carrying one digit, while in the ruminants it comprises two united elements supporting a pair of toes. This is clearly a case of parallelism in development attained by a shghtly different modification of principle. The parallelism does not, however, stop here, since an essentially similar type of cannon-bone has been produced in birds; only that im that group (with the exception of the ostrich) three long bones enter into its composition, which is further complicated by the addition of a bone from the ankle above. Sveing that in all these groups the parallelism has been arrived at by a different structural modification, the explanation of its mode of evolution is much less difficult than in the case of the molars of the camels and ruminants, where, as we have seen, the struc- ture is practically identical. Recent discoveries in North America have brought to light the existence of a kind of secondary parallelism among certain peculiar mammals which may be included among the hoofed or ungulate division of that class. In the even-toed group of that division, as exemplified by the pigs and ruminants, it is the third and fourth digits IN LIMBS. 51 which are symmetrical to one another, and tend to develop at the expense of the others ; while in the odd- toed group it is the third or middle digit which is symmetrical in itself and undergoes an ultra development. Now there are certain extinct creatures, which, while having claws instead of hoofs at the ends of their toes, yet are so closely related to the hoofed mammals that Fie. 18.—Front and side views of the Hind Foot of Artionyz. (After Osborn.) their separation therefrom is almost impossible. In one of these, which has long been known in Europe as the chalicothere, the third digit, as in the odd-toed hoofed mammals, is symmetrical in itself, and larger than either of the others; whereas in the newly-described animal known as Artionyx the third and fourth digits resemble those of the even-toed division of the hoofed mammals in being larger than the others and symmetrical to a line drawn between them. While, therefore, we have clearly a parallel development on different lines between the odd and even-toed hoofed animals im the development of a cannon-bone, these extinct clawed, hoof-like mammals (for want of a better expression) show a parallel parallel- ism (to coin another expression) which, if it had continued, might have resulted in the development of a two-clawed £2 52 PARALLELISM IN DEVELOPMENT. ruminant and a two-clawed horse. To our thinking, this is indeed one of the most curious phases of development yet discovered. In addition to the parallelism in the evolution of their molar teeth and limbs, the hoofed mammals likewise exhibit the same feature in regard to the second vertebra of the neck. As many of our readers are probably aware, the first, or atlas vertebra of the neck, turns with the head when the latter is moved sideways, the axis of rotation being formed by a process—the odontoid process —arising from the second or axis vertebra, and projecting into the central hollow of the atlas. Now in pigs and likewise all the primitive hoofed mammals, the so-called odontoid process is (as in ourselves) in the form of a flattened peg. On the other hand, in the ruminants, the modern horses, and the camels, which, as we have seen, represent three distinct phyla of the order, this peg has become modified into a spout-like half-cylinder, which must clearly have been separately evolved in each of these three groups. It is true that such a half-cylinder affords a far better basis of support for a heavy skull than does a mere peg; but the curious part of the matter is why these half-cylinders should be so exactly alike in the different groups, seeing that, as it would not be difficult to design some other structural modification by which the same end might have been attained, there is no necessity for their similarity. Our last example of parallelism will be drawn from two groups of extinct North American hoofed mammals, to which brief allusion has already been made in the chapter on “Tusks and their Uses” ; the one group being known as uintatheres, while the second is represented by a single species to which the name of Protoceras has been applied. Now, although the uintatheres have five-toed feet approxi- mating in structure to those of elephants, while in Proto- ceras each foot approximated to the ruminant type, in both groups the skull, as shown in the accompanying figure, was armed with several pairs of large, irregular, bony processes, which during life may have been sheathed in horn ; while in each case a pair of long tusks projected IN THE TAIL. 53 from the upper jaws, which were totally devoid of front or incisor teeth. Had such skulls been discovered without any indication as to the nature of the limbs with which they were associated, they would inevitably have been assigned to the same group of animals. The resemblance existing between thei, is, however, clearly due to parallel development, and we are thus shown another striking instance of caution necessary in endeavouring to determine the affinities of extinct animals from the evidence of in- complete remains. Fie. 19.—Skull of Protoceras. (After Osborn.) Our last instance of parallelism is alluded to in a later chapter, entitled ‘‘'The Oldest Fishes and their Fins,” in the course of which it is shown that while both the most ancient birds and the oldest fishes had long tapering tails with the joimts of tue backbone gradually diminishing in size, and each carrying either a pair of feathers or a pair of fin-rays, in all the modern representatives of the former group, and in a large section of the latter, the end of the vertebral column has been aborted into a composite bone, from which either the feathers or the rays of the tail diverge in a fan-like manner. In conclusion, we may say that although there is no very great difficulty in satisfactorily accounting for external parallelism obviously due to the necessity for adaptation to a particular mode of life, or in explaining those instances where a particular result has been brought about by 54 PARALLELISM IN DEVELOPMENT. different methods, yet when we find precisely similar structural modifications in different groups of animals which clearly cannot be traced to a common ancestry, and for which equally efficient substitutes could be readily suggested, we are fain to confess that the ordinarily accepted explanations of evolution appear to us altogether inadequate. Putting this aspect of the matter aside, our readers will, however, see from the imperfect sketch given above, what an important factor in evolution parallel development really is, and how largely it is likely in the future to modify our present views as to the mutual relationships of animated nature. TOOTHED WHALES AND THEIR ANCESTRY. 55 CHAPTER VI. TOOTHED WHALES AND THEIR ANCESTRY. Wiruiw the entire limits of the great mammalian class, there are, perhaps, no creatures which arouse a larger amount of interest—both among the general public and among naturalists—than those included under the names of whales, dolphins and porpoises, and collectively known as cetaceans. One reason for this universal interest is, doubtless, that among these denizens of the deep are com- prised the largest animals, not only of the present day, but likewise, so far as our information allows us to speak, of all epochs. Then, avain, the fact that such apparently fish-like creatures are really warm-blooded mammals, suckling their young in the manner distinctive of all other members of the class, and being under the necessity of coming to the surface at certain intervals for the purpose of breathing, cannot fail to strike even the most unobser- vant mind as being something quite beyond the ordinary. Moreover, the momentary glimpses which in general are all we obtain of these animals, and the halo of mystery which still to a great extent enshrouds their mode of life, are likewise important elements in generating the wide- spread interest they arouse. To the zoologist, cetaceans are indeed not only of prime importance as being the sole mammals which have assumed a purely fish-like form, and have become so thoroughly adapted to a completely pelagic life as to be unable to exist on land, but their study gives rise to many problems as to their origin and relationships, and the mode in which they attained their present con- dition. Into the consideration of the leading external features of cetaceans we need not enter very fully, merely pointing out that while the general contour of the body is fish- like, the tail-fin, or flukes, differs essentially from that of a fish in being horizontal instead of vertical; while 56 TOOTHED WHALES AND THEIR ANCESTRY. in place of the two sets of paired fins characterizing a fish, a whale has but a single pair of flippers representing the greatly modified fore limbs of other mammals. The hind limbs have, indeed, been completely lost externally, although more or less imperfect traces of them may still be detected deeply imbedded among the muscles of the body. In the great majority of the group the back is furnished with an upright fin, very similar in appearance to the unpaired back-fin of a fish. Whereas, however, such a back-fin is constantly present in fishes, in cetaceans it may be absent or present in different species of the same genus; while if we were to cut through such a fin we should find a total absence of the slender spine-like bones characterizing those appendages in a fish ; a similar condition also obtaining in the flukes. In marked contrast to the scaly armour of the majority of modern fishes, the skin of a cetacean is for the most part completely naked ; although the frequent presence, in the young state at least, of a few scattered bristles in the region of the mouth is of itself sufficient to indicate the derivation of these strangely modified creatures from more ordinary mammals. As regards their coloration, we may again, however, note a similarity to most pelagic fishes, in that while the upper parts are generally dark, the lower surface of the body is of a light hue; this arrangement being, of course, designed to render all these animals as inconspicuous as possible when viewed in the water either from above or from below. Although the flippers show no external indications of toes, and are unprovided with nails, yet their internal skeleton comprises the same elements as occur in the limbs of any ordinary mammal, and is thus quite different from that of a fish. This structural similarity is, however, to a certain degree obscured by the alteration in the form of the bones, and also by the circumstance that the number of joints in the skeleton of the individual toes is increased beyond the normal. As external ears would be mere useless incumbrances, these appendages are absent ; while the aperture of the ear itself is reduced to an extremely minute size. To prevent the ingress of water during the periods of submergence, the apertures of the nostrils, which CLASSIFICATION. 57 may be either double or single, can be completely closed at will, and are only opened at such times as the creatures come to the surface to breathe, when a column of water is generally thrown up by the rush of expired air let loose Fig. 20.—The Bridled Dolphin. (From True, Bull. U.S. Nat. Museum.) shortly before the head reaches the surface. In all their internal structures, as well as in the mode of production and nourishing of their young, cetaceans conform strictly to the ordinary mammalian type; and we accordingly see that their assumption of a fish-like form is, with the exception of the loss of the hind limbs and the modifi- cation of the front pair into flippers, mainly superficial. In departing from the fish-type in having the expansion of the tail-fin horizontal instead of vertical, the necessity of having an organ capable of bringing them rapidly to the surface has been the inducing cause; while in order to prevent their blood from being reduced below the proper temperature by the chill of ‘the surrounding water, the whole body is invested with a thick layer of oily fat, commonly known as the blubber, beneath the skin. Existing cetaceans are divisible into two great groups, distinguished as the whalebone whales and the toothed whales ; the latter group including sperm-whales, together 58 TOOTHED WHALES AND THEIR ANCESTRY. with the grampuses, porpoises, and dolphins. The most obvious distinction between these two groups, as the terms applied to them indicate, relates to the presence or absence in the adult condition of true teeth. Confining our attention in the present chapter to the toothed cetaceans, of which the dolphins and their allies are the least specialized representatives, we find that the two jaws may be in some cases provided with a full series of teeth, while in other forms the number of teeth may be reduced to a single pair, or even, as in the male narwhal (Fig. 9), to a solitary tusk. Whether, however, the teeth be many or few (and in the female narwhal there are none of any functional importance), the structure known as whalebone is never developed in the mouth; while all the members of the group are further distinguished from the whalebone whales by the circumstance that the nostrils invariably open by a single external aperture, which is very frequently in the form of a transverse crescentic slit, closed by an overhanging valve. In the latter respect these cetaceans are more specialized than are the whalebone whales; and as the presence of teeth in the former indicates that they could not have been Fria. 21.—The last six Upper Teeth of the Killer. (After Sir W. H. Flower.) derived from the latter, it is evident that the two groups are of extreme antiquity, and have undergone a parallel development. Till recently, it has indeed been considered that they were divergent branches from some common ancestral type; but, as mentioned in the chapter on ‘‘ Parallelism in Development,” it has been lately suggested NATURE OF TEETH. 59 that each may have had a totally distinct origin, although the evidence in favour of such a view is, at present at least, far from conclusive. In the structure of their teeth, the modern toothed whales differ very widely from the generality of mammals. In the first place, their teeth are always of the simple structure shown in Fig. 21, having conical or compressed crowns, and undivided roots; while, secondly, there is only one single series developed, the replacement of the anterior ones characterizing the majority of mammals being wanting. From this simple structure of their teeth it has been argued that these cetaceans are among the most primitive of all mammals; but, altogether apart from the conclusive evidence that all whales (as proved by their breathing air) are derived from land mammals, it has recently been shown by the researches of Dr. Kiikenthal, of Jena, that this view is quite untenable. By examining em- bryos of young cetaceans of this group, that observer has demonstrated that there are actually rudiments of a second series of teeth, which, although never coming to maturity, serve to show that there were once two complete sets, and that the permanent teeth correspond, in part at least, to the milk-teeth of other mammals; thus indicating that the present state of the cetacean dentition is a degraded one. Hitherto it has not, indeed, been shown by embryology that the teeth of this group were originally of a complex type (although in the case of the whalebone whales this has been demonstrated), but, fortunately, here paleontology comes to our aid. Thus in the middle of the Tertiary period there occur remains of what may be termed shark- toothed dolphins (squalodonts), in which the permanent teeth are differentiated into distinct series, corresponding to the incisors, canines, premolars, and molars of other mammals; while, for all we know to the contrary, there may also have been a regular replacement of the more anteriorly placed teeth. In these shark-toothed dolphins the molar teeth, instead of being of the simple structure of those represented in our illustration, were severally implanted in the jaws by two perfectly distinct roots ; while their large, laterally compressed, and somewhat fan- 60 TOOTHED WHALES AND THEIR ANCESTRY. shaped crowns were furnished with a number of cusps on their hinder cutting-edges. Indeed, these teeth much resemble the premolar teeth of a dog or the molars of a seal; and they obviously serve to indicate a transition from the modern toothed cetaceans towards ordinary mammals. This, however, is by no means all, since in a still earlier portion of the same division of the Tertiary period there occur other cetacean-like animals known as zeuglodonts, which have still more complicated teeth, and otherwise depart further from the modern cetacean type—so much so, indeed, that they have been regarded by some writers as more nearly allied to the seals. In our own opinion they are, however, undoubtedly primitive cetaceans, and thus serve, not only to connect the present group with other mammals, but also, in conjunction with the shark-toothed dolphins, to show that the simple teeth of the former are clearly pro- duced by degeneration from a complex type. As regards the particular group of land mammals from which whales were derived, it has been thought that their nearest allies are with the ancestors of the pig-like hoofed mammals. This, indeed, is the view of Sir W. H. Flower; but we confess that from the nature of the teeth of the two extinct groups above mentioned, coupled with certain resemblances of the skeleton of the zeuglodonts to those of seals, we are rather more inclined to look among flesh-eating land mammals for the lost ancestors. Still, however, it must be remembered that in the early Eocene period, which is probably the very latest epoch at which cetaceans could have originated, the distinction between carnivorous and hoofed mammals was but imperfect, so that, after all, the ancestral cetacean stock may well have been of an extremely generalized type. At present, however, we are almost completely in the dark in all that concerns this interesting subject. By far the largest of all the toothed whales is the gigantic sperm-whale, the typical representative of a family characterized by the absence of teeth in the upper jaw of the adult, while those of the lower jaw are very variable both as regards form and number. In the sperm-whale, of which the male attains a length of between fifty and sixty SPERM-WHALES, ETC. ~ 61 feet, the lower teeth vary in number from fifty to sixty on each side, and are characterized by their large size and pointed crowns, upon which there is not a trace of enamel. Another characteristic feature of this animal is the enormous size of the head, which terminates in an abruptly truncated muzzle of great depth, and in a cavity of which is contained a peculiar oily substance, yielding when refined the well-known spermaceti. An even more valuable product of this animal is ambergris, which, while accumu- lated as a concretion in the intestine, is generally found floating on the surface of the sea ; in appearance it is an amber-coloured substance, containing a number of the horny beaks of the squids on which the sperm-whale subsists. Omitting mention of the lesser sperm-whale—the only other member of the first division of the family—we pass on to mention the bottle-nosed and beaked whales, characterized by having all the lower teeth, with the exception of a single pair, rudimentary, and concealed in the gum. In the bottle-noses—so named from the extreme convexity of the crown of the head of the adult males, which rises suddenly above the short beak—there is but a single pair of teeth, situated in the front of the lower jaw, and even these are invisible during life. These whales, of which there is but a single well-defined species. (Hyperoédon rostratus), although not exceeding some thirty feet in length, are valuable on account of their oil, as well as from yielding spermaceti from their skulls. In contradistinction to the sperm-whale, they carry, in common with the beaked whales, a large back-fin. In the beaked whales, of which there are three existing genera, the skull is produced into a long beak, of which the upper half is formed by a solid bone of ivory-like density ; while in the lower jaw there are either one or two pairs of teeth, which, although variable in position, are generally of large size, Itis one of these whales (Mesoplodon layardi) which is alluded to in the chapter on ‘‘ Tusks and their Uses,” as being provided with teeth of such a size as actually to impede the free opening of the mouth. From their general avoidance of the neighbourhood of 62 TOOTHED WHALES AND THEIR ANCESTRY. coasts, and their apparently somewhat solitary habits, extremely little is known of the life-history of the beaked whales, of which the skeletons are but poorly represented in our museums. At the present day very rarely seen in the English seas, during the Pliocene period these whales must have been extremely numerous in the North Sea, since their fossilized beaks are amongst the most common vertebrate fossils obtained from the crags of Suffolk and Essex. The same deposits, together with others of corre- sponding age on the Belgian coasts, havealso yieldedremains of a number of extinct whales, more or less closely allied to the sperm-whale, thus indicating that the latter is the last survivor of a once numerous group. The teeth of many of these fossil sperm-whales differed, however, from those of their living cousins in having their crown capped with enamel. Fre, 22.—The Indian Porpoise. (From True, Bull. U.S. Nat. Museum.) In the general presence of numerous teeth in both the upper and the lower jaws, the dolphins and their allies the porpoises, killers, and white-whale differ from all the above-mentioned forms, and thus constitute a second family —the Pelphinide. The group is a numerous one, which is split up into a number of genera, some of which are by no means easy to distinguish. They may, however, be roughly ranged under two main divisions, in one of which the muzzle is short and rounded, as in the porpoises and black- fish, while in the other, as represented by the dolphins (Fig. 20), it is produced into a longer or shorter beak, of which the base is marked off from the main portion of the head by a distinct re-entering angle. The most aberrant member of the first group is the spotted Arctic narwhal, of which a brief notice, accompanied by a figure, was given PORPOISES AND KILLERS. 63 in the chapter on “‘ Tusks.” Allied to the narwhal is the beautiful white-whale (Beluwya), which is likewise a northern species, distinguished by its glistening white skin, the absence of any tusk, and the presence of numerous well-developed teeth in the fore part of the jaws ; neither of these species having a back-fin. Although the name “porpoise” is applied indiscriminately to several members of the family, it should properly be restricted to a few comparatively small-sized species characterized by the presence of some twenty-five small and flattened teeth with spade-shaped crowns on either side of each jaw. Porpoises are among the most common and familiar of all cetaceans, their rolling gambols being well known to all who have made a voyage; but whereas the common porpoise has a distinct back- fin, in the species represented in our illustration that appendage is lacking. Less familiar, on the other hand, are the much larger and handsomely coloured killers, or grampuses (Urea), differing by the great size of their teeth (Fig. 21), which are usually twelve in number on each side, and the great development of the back-fin. Fie. 23.—Pacific Black-Fish. (From True, Bull. U.S. Nat. Jus.) Attaining a length of about twenty-five feet, the killer derives its title from its rapacious habits; a single specimen having been known to swallow several whole seals in succession, while not unfrequently several individuals have been observed to combine their forces to attack and kill the larger members of the order. Killers may always be easily recognized while swimming near the surface by the 64 TOOTHED WHALES AND THEIR ANCESTRY. great height of their nearly vertical back-fins. Allied cetaceans with smaller teeth (Orcella) frequent the Bay of Bengal and ascend some distance up the Irrawadi. Another well-marked type is the black-fish (Fig. 23), characterized by its remarkably short and rounded head, the uniformly black hue of the skin, and by the eight or twelve small and conical teeth being confined to the front portion of the jaws. There are other less well-known representatives of this eroup which we have not space to notice; and we accordingly pass on to say a few words about dolphins—a term which should be restricted to those forms having a distinctly marked beak. Since, however, sailors will persist in speaking of dolphins indifferently, either as bottle-noses or porpoises, the inexperienced landsman must be on his guard not to confound them when thus spoken of either with the bottle-nosed whales or the true porpoises. Dolphins, which are divided into numerous genera, according to the number of their teeth, the length of the beak, and other characters, are all comparatively small species, seldom exceeding some ten feet in length ; and while the great majority are marine, a few ascend some of the larger tropical rivers, such as the Amazon. Fish of various kinds constitute their usual prey ; but one peculiar species recently described from the Cameruns district is believed to subsist on sea-weed. Of the better-known types, the common dolphin represents the genus Delphinus, the bottle-nosed dolphins constitute a distinct genus (Tursiops), while the long-beaked dolphins are separated as Steno. To the foregoing group of beaked dolphins the ordi- nary observer would, doubtless, be disposed to refer three peculiar species severally restricted to the larger rivers of India, the Amazon, and the mouth of the La Plata river, but as these differ more or less markedly from other dolphins in certain structural features they are referred to a distinct family. Moreover, since these peculiarities approximate to a more generalized type, while their fresh- water habits and scattered distribution indicate extreme antiquity, it is not improbable that these three dolphins. FRESH-WATER DOLPHINS. 05 are the most primitive of all existing cetaceans. At present we have, indeed, no evidence of fossil forms allied to the susu or Gangetic dolphin (Platanista); but in the older Tertiary deposits, both of the United States and Europe, there occur the remains of dolphins evidently nearly allied to the two existing South American species (Inia and Stenodelphis), and thus clearly proving the antiquity of those types. It may be added that it is probable that the ancestors of the cetaceans which first took to an aquatic life were inhabitants of fresh-water, and it is therefore only what we should expect that the most primitive of the existing representatives of the order were likewise of fluviatile habits. On the other hand, it must be confessed that the similarity in structure of the widely separated existing fresh-water dolphins is some- what difficult to account for on this hypothesis. Te 66 WHALEBONE AND WHALEBONE WHALEg. CHAPTER VII. WHALEBONE AND WHALEBONE WHALES. Sreine that the substance so-called has nothing in common with true bone, many zoological writers object to the use of the term ‘‘ whalebone”; and they have accordingly proposed the substitution of the word ‘‘ baleen,’’ which has been specially coined for the purpose. To our thinking, there is, however, no necessity for this substitution of a word of foreign origin for such a well-known English name, any more than there is for replacing the native term ‘ black Jead”’ by its foreign equivalent ‘‘ graphite.” Everybody knows what is meant by whalebone or black lead, while comparatively few are familiar with the terms “ baleen” and ‘graphite’; and as the two former are every bit as good as their foreign substitutes, we prefer to employ them. If, indeed, there should exist any persons so misguided as to imagine either that whalebone is equivalent to the bone of whales, or that black lead has any sort of affinity with load, we fear that the substitution of the terms ‘‘baleen”’ and “graphite” would not much aid in removing their ignorance. The substance which we accordingly take leave to call whalebone is one of the chief essential characteristics by which the whalebone whales are distinguished from the toothed whales forming the subject of the preceding chapter. As our readers are probably aware, this substance is attached to the upper surface of the mouth of the whale, from which it depends in the form of a series of parallel, narrow, elongated, triangular plates, placed transversely to the long axis of the mouth, with their external edges firm and straight, but the inner ones frayed out into a kind of fringe. The longest plates of whalebone are situated near the middle of the jaw, from which point the length of the plates gradually diminishes towards the two extremities, where ARRANGEMENT OF WHALEBONE. 67 they become very short. There is, however, whalebone and whalebone, and whereas in the Greenland whale the length of the longest plates varies from some ten to twelve feet, while the total number of plates in the series is about 380, in the great rorquals or fin-whales the length is only afew inches, while the number of plates is considerably less. To accommodate the enormous whalebone plates of the Greenland whale, the bones of the upper jaw are greatly arched upwards, while the slender lower jaw is bowed out- wards, thus leaving a large space both in the vertical and horizontal directions, the transverse diameter of the space being much wider below than above. When the mouth is Fia. 24.—Section through the Skull of the Greenland Whale, with the outline of the soft parts. he arrows indicate the position of the nasal passage and aperture. closed, the plates of whalebone are folded obliquely back- wards, with the front ones lying beneath those behind them ; but directly the jaws are opened, the elastic nature of this wonderful substance causes it to spring at once into a vertical position, and thus form a sieve-like wall on both sides of the mouth, the thin ends of the plates being pre- vented from pushing outwards by the stiff lower lip which F2 68 WHALEBONE AND WHALEBONE WHALES. overlaps them. By elevating its enormous fleshy tongue within the cavity thus formed, the whale causes the enclosed water to rush out between the plates, leaving such small creatures as it contained lying high and dry on the surface of the tongue ready for swallowing. In structure, whalebone (which, by the way, although black in the Greenland whale, is white in some of the other species) is of a horny nature, and grows from transverse ridges on the mucous membrane of the roof of the mouth ; being, in fact, nothing more than an ultra- development of the ridges on the palate of a cow, hardened and lengthened by an excessive growth of a horny super- ficial or epithelial layer. The whole of this stupendous horny growth takes place, however, after birth, young whales having smooth palates, with no trace of the horny plates. Although at birth young whalebone whales show no traces of the substance from which the group derives its name, they equally exhibit no evidence of the presence of teeth. If, however, their jaws be examined at a still earlier stage of development, it will be found that there are a number of small teeth lying within a groove beneath the gum on each side of both the upper and the lower jaws. Previous to birth these teeth become absorbed, and thus never cut the gum. Their presence in this transitory stage is, however, of the deepest interest to the evolutionist, since they unmistakably indicate the derivation of the whalebone whales from ancestors provided with a full series of functional teeth. This, however, is not the whole of the story these rudi- mentary structures have to tell. From the recent investigations of Dr. Kikenthal, it appears that in addition to the above-mentioned tooth-germs, the jaws of very young whales likewise exhibit traces of a still earlier deciduous series of milk-teeth; thus showing that the former correspond to the permanent series of other mammals. Accordingly, these tooth-germs do not represent the functional teeth of the toothed whales, which, as we have seen in the chapter on that group, correspond to the milk-teeth of ordinary mammals. Even more remarkable are certain observations relating to the structure of these CHARACTERISTICS. 69 tooth-germs. It has been shown, indeed, that the teeth in the latter part of the series, when first formed, consist of a number of adjacent cusps, and that as development proceeds these cusps become completely separated from each other so as to constitute distinct individual teeth of a simple conical form. This discovery is of the very highest import, since it serves to indicate how the numerous simple conical teeth of the dolphins and other toothed whales have probably been derived by the splitting and subdivision of originally complex cusped teeth more or less closely resembling those of the extinct zeuglodonts referred to in the last chapter. Being primarily distinguished from the toothed whales vy the total absence of teeth after birth and the presence of whalebone in the adult condition, the whalebone whales present certain other distinctive characters to which we may now briefly allude. In the first place, the whales of this group differ externally from all those furnished with teeth in that their nostrils open externally by two distinct longitudinal slit- like apertures ; while, if we cut into the head, we shall find that there is a distinct organ of smell, of which all traces have disappeared amcng the toothed whales. Moreover, instead of the skull being invariably unsymmetrical in the region of the nose, as it is in the latter group, it retains the normal symmetry ; while, instead of the mere nodules which in the toothed whales represent the nasals of other mammals, in the whalebone whales these bones are fairly well developed. Then again, the lower jaw of any member of the present group may always be distinguished from that of a toothed whale not only by the absence of teeth, but likewise by the circumstance that each of its branches is much bowed outwards in the middle, while their anterior extremities are connected together merely by ligamentous tissue, instead of by a bony symphysis of greater or lesser length. Many other points of difference between the two groups might be cited, but we have especially referred to those mentioned above, for the reason that while the presence of whalebone indicates that in one respect the 70 WHALEBONE AND WHALEBONE WHALES. members of this group are more specialized than their toothed cousins, in regard to the structure of the skull in the region of the nose and the retention of an organ of smell, as well as in the double apertures of the nostrils, they depart less widely from the ordinary mammalian type. And it is from this evidence that zoologists regard the two main groups of whales as being widely divergent branches from a common ancestral stock, if, indeed, they do not go so far as to consider that each has had a totally independent origin. With regard to the number of forms by which they are represented, the whalebone whales are far less numerous than the toothed group, the whole of them being included within a single family. What the group lacks in number is, however, amply made up by the great bodily size of its various representatives. Among the toothed whales, the sperm-whale alone attains gigantic dimensions; whereas in the present group there are several species, such as the Greenland whale, which nearly equal that enormous creature in bulk, while two of the rorquals far surpass it. This, however, is by no means all, for whereas the great majority of the dolphins and porpoises are relatively small cetaceans of less than twenty feet in length, the smallest member of the present group is the southern pigmy whale, which attains a length of twenty feet, while next to this comes the lesser rorqual, whose length is frequently close on thirty feet. Of the various genera of this group, the most specialized are the typical or right whales (Lalena), in which the black whalebone is far longer and more elastic than in any of the others, except the pigmy whale; while, in order to accommodate it in the mouth, the skull has the palate narrower and much more highly arched, and the two branches of the lower jaw more outwardly bowed than in the other members of the group. Ex- ternally, these whales—which are commercially of far greater value than the undermentioned rorquals—are characterized by the inordinately large dimensions of the head, by the smooth throat, the moderate length of the flippers, and the total want of a back-fin. So gigantic RIGHT WHALES. 71 indeed, is the size of the mouth in these whales that its capacity actually exceeds that of the whole of the other cavities of the body; and yet the size of the throat is so smallas almost to justify the common nautical saying that a herring will choke a whale. There are but two well- defined species of right whales, viz., the Greenland or Arctic whale (B. mysticetus), and the southern right whale (B. australis), the latter of which was nearly exterminated some centuries ago in the Atlantic by the Basque whalers, while the former is only too likely to share the same fate at the hands of their modern successors in the Arctic seas. Of the two, the Greenland whale is decidedly the more specialized, having a much larger head and longer whalebone, and is in this respect facile princeps among its tribe ; although, as it only measures from forty- five to fifty feet in length, it is inferior in point of size to the rorquals. The skeletons of both these whales are cha- racterized by the whole of the vertebre of the neck being welded together into a solid mass; and the same feature is exhibited by those of the pigmy whale (Neobalena) of the southern seas, which, as already mentioned, is a mere dwarf among giants, asitdoes not exceed some twenty feet in total length. Agreeing also with the right whales in its smooth throat, the pigmy whale differs by having a small hook-like fin on the back, while its long and elastic whalebone is white instead of black. Far larger than the last, the great grey whale of the Pacific (Rhachiancctes) forms a kind of connecting link between the right whales and the rorquals, having the smooth throat and finless back of the former, while its whalebone is even shorter and coarser than in the latter; the palate consequently is but little vaulted, and the entire head smaller in proportion to the body. The remaining whales of this groupare divided into hump- backs (Megaptera) and rorquals or finners (Balenoptera) ; both of which are characterized by the presence of a number of parallel groovings or flutings in the skin of the throat, as well as of a back-fin (whence their name of finners or fin-whales), and also by the shortness and coarseness of their whalebone, which is generally of a va WHALEBUNE AND WHALEBONE WHALES. yellowish colour. Their flukes are, moreover, less expanded than are those of the right whales ; while, as already said, their heads are relatively smaller and lower, with the cavity of the mouth much less vaulted. In their skeletons, the vertebre of the neck differ from those of the night whales in being longer and completely disconnected from one another; and in this respect the Pacific grey whale holds a position intermediate between the two groups. Hump-backs, of which there is but a single species, are specially characterized by the shortness and depth of the body, which behind the shoulder rises above the level of the back-fin, and by the exceeding elongation and slenderness of the flippers, which are about equal to one-fourth the total length of the animal. The female hump-back, which somewhat exceeds her partner in size, attains a length of from forty-five to fifty feet, or about the same as that of the Greenland whale. An enormous whale, believed to belong to this species, became some years ago entangled in the telegraph cable line off the coast of Baluchistan with three turns of the cable round its body. On the other hand, in the rorquals, the body is long and slender, while the flippers are small and pointed. Of the four well-established species of this genus, the blue or Sibbald’s whale (Bb. sibbaldi)—the “ sulphur-bottom ” of the American whalers—enjoys the proud distinction of being the largest of all known animals, whether living or extinct, attaining the enormous length of from eighty to eighty-five feet; while the common rorqual (B. musculus) comes in a good second with a length of from sixty-five to seventy feet. Both the others are, however, con- siderably smaller. As regards their distribution in time, whalebone whales have left their remains commonly enough in the Plio- cene strata of all parts of the world, and they likewise oceur in those of the Miocene period; but, although a single vertebra from the Eocene beds of Hampshire has been assigned to a member of this group, there is at present no decisive evidence that they had come into existence at such an early date. Since most of the Pliocene species are of smaller dimensions than their living repre- DESTRUCTION BY MAN. 73 sentatives, it appears that these marvellous creatures had only attained their maximum size shortly before the time when their very existence was to be threatened by the relentless hand of man. Formerly the only members of this group of whales which were thought worthy of general pursuit were the right whales; the shortness of their whalebone, coupled with their relatively small yield of oil, and their tre- mendous speed, rendering the rorquals scarcely worth the trouble and risk of hunting. Of the two right whales, the Greenland species, as being the more abundant, received the greatest share of attention ; and so relentless has been its pursuit, that it is now either well-nigh exterminated from many of its ancient haunts, or has retreated still farther north to regions almost impossible of access. As showing how the constant persecution in the Greenland seas has told upon the size of the comparatively few remaining individuals of this species, it may be mentioned that the eleven specimens killed there during the season 1890-91 yielded an average of less than 8 cwt. of whale- bone, whereas in five taken during the same season in Davis Strait the average yield was more than double this amount. In consequence of this diminution in the number and size of the Greenland whale, the value of whalebone has of late years gone up enormously; and whereas some time ago whalebone of over six feet in length sold at £1000 per ton, in 1892 it had reached the enormous price of upwards of £2800 per ton. The southern right whale yields a smaller quantity of rather shorter and less valuable bone, now selling at from £1600 to £1800 per ton; the quantity obtained from a well-grown example varying from 800 to 1200 pounds. The amount of oil produced by a whale of the same species averages from eight to fourteen tons, of which the present market value is about £28 per ton. If the unfortunate animals are not allowed some vespite, it is only too probable that the supply will before long cease altogether. As another result of this growing scarcity of the Green- land whale, attention has been directed to the previously despised rorquals and hump-backs; the employment of 74 WHALEBONE AND WHALEBONI WHALES. steam whaling vessels, and the use of explosive harpoons. having enabled the whalers easily to cope with the greater speed of these cetaceans. At Hammerfest, a special ‘‘ fishery” has, indeed, been established for the capture of rorquals, where the take of these animals is now large. The products of these whales are, however, nothing like the value of those of the Greenland species; and if the latter, together with the southern right whale, be so nearly exterminated as to render pursuit no longer profitable, the supply of long whalebone will absolutely come to an end. Postscript.—Since the foregoing was written, the author lias discovered that certain fossil toothed whales from Patagonia had their nasal bones nearly as well developed as in the whalebone whales ; this discovery removing one of the difficulties in regarding the latter as the descendants of the former, RUMINANTS AND THEIR DISTRIBUTION. ras CHAPTER VIII. RUMINANTS AND THEIR DISTRIBUTION. From early times we find the function of ruminating, or “chewing the cud,” recognized as a peculiarity of the group of mammals known in semi-popular language as ruminants. In ‘“ Deuteronomy,” for instance, the animals permitted for food are those that “ chew the cud and part the hoof” ; while the swine, “ which part the hoof but do not chew the cud,’ are forbidden. On the other hand, the camel, which chews the cud but has not paired hoofs, is in the forbidden list. In the permitted animals we thus have a recognition of the group of ruminants as represented by oxen, sheep, and deer; of which no better short definition can be given than that they chew the cud and have each foot furnished with a pair of hoofs symmetrical to a vertical line between them. The want of the paired hoofs in the camels, which are also cud-chewers, show, however, that these two characteristics will not hold good for the entire group. As we proceed, we sball find that there are structural features, common to the group, in addition to the peculiarity of rumination; but before going further, we may observe that the recognition of their paired hoofs, coupled with the absence of rumination, is an exact statement of the relationship of the swine to the true ruminants. The word “ruminant”? comes from the Latin rumen, which was applied both to the “cud” and to that part of the stomach in which the latter is contained previous to chewing. The Greeks had a word meruko or merukizo (from meruo, to revolve), to express this action of cud- chewing, and a derivative from the former was used by Aristotle to designate ruminants, who thus first distinguished the group by a definite name. This early recognition of the ruminants as a group is probably due to their importance to man, the Biblical record showing 76 RUMINANTS AND THEIR DISTRIBUTION. that they yielded the only mammalian food permitted to the Hebrew, and this pre-eminence as a source of food has scarcely decreased tothe present day. They are, moreover, Vic, 25.—skeleton of the Extinct Irish Deer—a typical Ruminant. now the dominant type of larger mammals, as witness the herds of bison which lately roamed over the American NATURE OF RUMINATION, Gils prairies, and the droves of antelopes on the African veldt. Commencing with the function of rumination, we may observe that it is a re-mastication of grass or other vegetable food, swallowed almost as soon as plucked, and transferred to a special receptacle in the stomach. From this it is regurgitated into the mouth by a reversed action of the muscles of the throat, and, after having undergone mastication,—or rumination—is transferred to the digesting part of the stomach. Now, it is evident that this compli- cated arrangement, so different from that of other animals, must be of some special advantage to the ruminants. As a matter of fact, these animals, like other large herbivores, are obliged to consume a vast quantity of food to obtain sufficient nutriment; and itis obvious that if this food had to be masticated as soon as plucked, the operation of feeding would be very protracted; but by the arrangement mentioned the requisite amount of food can be gathered within a comparatively short time, and the animals can then retire to ruminate in concealment. It is superfluous to comment on the advantage of this arrangement to creatures which, like many ruminants, have but little means of defending themselves against carnivorous foes; but we may mention that many still further increase this advantage by feeding only at dawn or evening, when they are far less conspicuous than in the mid-day glare. There is, moreover, evidence that when ruminants first appeared, this rapid feeding was of more importance than at the present day, since while many of the modern larger forms, like oxen, antelopes, and deer, are provided with formidable weapons in the shape of horns or antlers with which they can keep foes at bay, in earlier times such weapons were either absent or but feebly developed. Seeing, then, that the function of rumination is correlated with a special compartment of the stomach for the temporary reception of the freshly-gathered food, it would be expected that animals thus provided would also possess an efficient masticating arrangement for reducing their food to the condition in which it yields the fullest nutriment. Such, indeed, is the case, the grinding-teeth 78 RUMINANTS AND THEIR DISTRIBUTION. of ruminants being of a complex structure, unknown elsewhere ; the last three of these teeth in the upper Jaw (Fig. 26) being composed of four columns, of varying height, of which the two inner ones are crescent-shaped; such a type of tooth being termed selenodont. The lower grinding-teeth having their crescents directed the opposite way to those of the upper jaw, and both upper and lower teeth consisting of layers of different hardness, we can scarcely imagine a better masticating machine than is presented by the opposition of the two series of erinding - teeth of these animals. Bearing in mind this structure, the definition of cud-chewing, selenodont mammals will suffice to distinguish the ruminants from all other animals. When, however, we say that these characteristics dis- tinguish them from all other mammals it must be added that this refers only to those of the present day, for a transition can be traced through extinct forms to the more simply constructed orinding-teeth of the swine. We have already seen how the Mosaic law recognized the similarity in the structure of the hoofs of the ruminants and the swine, and it is curious that while under the Cuvierian system of zoology these two groups were widely sundered, modern paleontological researches have shown that they are really closely related, the want of the power of chewing the cud, with the correlated absence of the selenodont structure of the teeth, being the chief essential features in which the latter differ from the former. Here a curious problem is presented to those who put their faith im a mode of evolution dependent only upon so-called natural causes, in that it’is impossible to give any adequate explanation of what advantage would be the development of an incipient selenodont structure in the teeth of the early swine-like ungulates, or at what precise stage the function of chewing the cud, with the con- Fic. 26.—A left Upper Cheek-Tooth of an Antelope. STRUCTURE OF FEET. 79 comitant development of a separate compartment in the stomach, was superadded to the normal mode of feeding characteristic of the swine. Here we must say a few words as to the structure of the ruminant foot. The “cloven hoof” of ruminants and swine has become such a proverbial expression that the idea may still linger that this is due to the fission of a single hoof, like that of a horse. Nothing could, bowever, be further from the truth ; the two hoofs of a ruminant (Fig. 27) corresponding to the terminal joints of our own middle and ring-fingers (or the correspond- ing toes), which are the third and fourth of the typical series of five. The lateral or spurious hoofs (not shown in Fig. 27) of the ruminants represent our own index (second) and little (fifth) fingers, or toes. It is a further peculiarity of the true ruminants and camels that the two separate bones which in the swine connect the two large digits with the wrist or ankle are fused into a single cannon-bone (Fig. 27); the primary dual origin of which is indicated by the two distinct pulley- like surfaces at the lower end, which carry the bones of the digits. The peculiar little ruminants known as the chevrotains—of which more anon—retain, however, evidences of their kinship with the swine, in that some of them have the two elements of the front cannon-bone —or metacarpals as they are then called—quite separate from one another. Indeed, in the same Fic. 27.—Bones of the Hind Foot of a Rumi- nant. The letters 1 dicate the lower bones of the ankle. Osborn.) (After 80 RUMINANTS AND THEIR DISTRIBUTION. manner as we may trace a transition from the selenodont tecth of the ruminants to the bunodont ones of the swine, we may mark how the two-toed and cannon-boned ruminants passed into swine-like animals with four toes supported by as many separate metacarpal bones. Having now mentioned the leading characters of a modern ruminant, as distinct from other mammals, we may refer toa peculiarity, which, although by no means characteristic of all, is a striking one, and one sharply differentiating the group from all others. This is the tendency to the development of appendages on the skull, arranged ina pair at right angles to its longer axis, and taking the form either of solid branching antlers, as in the deer, or of hollow sheaths of horn covering bony cores on the skull, as in the oxen and antelopes. Passing to the consideration of the various kinds of cud- chewing mammals, we find that the true ruminants, or those with hoofs, no upper front teeth, and a cannon-bonc in both limbs, arrange themselves in several minor groups. The most important to man are the “ hollow-horned ruminants,” such as oxen, sheep, goats, and antelopes, all of which are characterized by the presence of horns, at least in the males. The vanety of form assumed by the horns renders this group one of the most attractive of all animals; and we have but to recall the curved and smooth horns of the oxen, the equally massive but wrinkled ones of the wild sheep, those of the ibex with their knotted points and scimitar-hke backward sweep, the spear-like form of those of the gemsbok, and the spiral twist of those of the kudu and eland, to realize the variety of contour assumed by these appendages. The oxen (including bison and buffaloes) are, with the exception of the American bison, Old World types, and were formerly abundant in Europe, where, however, they are now only represented by the bison preserved in the forests of Lithuania and the Caucasus, and by the half- wild cattle of Chillingham and some other British parks, which have been thought to be the direct descendants of the British wild ox, or aurochs, of Csesar’s time, but are more probably derived from ancient domesticated breeds SHEEP, GOATS, AND ANTELOPES. 81 which have reverted to a nearly wild state. True wild oxen now exist only in India and the adjacent regions, while wild buffalo occur both in India and Africa. Equally characteristic of the Old World are wild sheep and goats, the “big-horn” being an outlying North American type. Both groups are essentially mountain animals, the head-quarters of the former being the high- lands of Central Asia, while on the southern flanks of the same mountain-barrier the latter are more abundant. Both are also represented in the mountains of Europe; but in peninsular India there is but the wild goat of the Nilgiries, while in the whole of Africa we have only the wild sheep of Barbary and the ibex of Abyssinia. This absence of sheep and goats from Africa may, perhaps, be due to the fact that these animals are of comparatively late origin, and were probably poorly represented at the time when the other ruminants entered that continent from the north. The musk-ox of Arctic America is an aberrant form allied to the sheep. The antelopes have a distribution nearly the reverse of that of the sheep and goats, the great majority being restricted to Africa, where there are probably fully ninety species, against about a score in all the rest of the world, except Arabia and Syria, of which the fauna is allied to that of Africa. Indeed, the only typical antelopes found beyond these regions are the black-buck, the nilgai, the four-horned antelope of India, the saiga of Tartary, the chiru of Tibet, and several members of the widely dis- tributed gazelles. The rings marking the horns of the latter (Fig. 28) and many other antelopes are very distinc- tive of the group, although by no means universal. The European chamois, the goat-antelopes of India and China, andthe Rocky Mountain goat of America, serve to connect the typical antelopes with the goats, and it is these alone which represent the group in Europe, to the eastward of India, and in North America. Seeing that in Tertiary times, antelopes of African types occurred in southern Europe and India, it is difficult to determine why the group should have so dwindled or disappeared there ; although we can readily account for their extraordinary G 82 RUMINANTS AND THEIR DISTRIBUTION. development when they once obtained an entry into Africa, on account of the immense area open to them, in which there was no competition by any other ruminants except butfaloes and giraffes. To the zoologist Africa is indeed a country characterized by the number of animals living there which have disappeared from other regions ; and there is no better instance of this survival than the giraffe, a ruminant that, as regards its cranial appendages, stands midway between the hollow- horned group and the deer.* Weare all familiar with the ungainly and yet beautiful form of the giraffe ; but it is probably less well known that giraffes once roamed over Greece, Persia, India, and China, where, as in Africa at the present day, they were accom- panied by ostriches and hippopotami. And here again we are confronted by the problem how to account for the disappearance from regionsapparently exactly suited to their habits, of all these animals. The giraffe is, how- p14 98 Horns of ever, not only the sole survivor of Gazelle. (From several extinct species of its own Giinther.) kind, but it likewise represents a lost group of ruminants, intermediate between the horned and antlered Old World types. The head-quarters of this group was India, where, among other forms, occurs the gigantic sivathere, rivalling the elephant in bulk, and characterized by its two pairs of horns (Fig. 29), of which the hindmost were branching and _antler-hke, although apparently never shed, and probably covered during life with skin and hair. If our attention has been turned to Africa as the head- quarters of antelopes and giraffes, it must be directed to * See the following chapter. DEER. 83 other regions when we come to the deer, since, with the exception of the Barbary stag, there is no representative of the group in all that continent. With few exceptions, deer are characterized by the antlers of the males, the reindeer alone having these appendages in both sexes. They are the only true ruminants found in South America, where many of the species have comparatively simple antlers, and thus show affinity with the early fossil types, some of which were antlerless. Allied species range through North America, but it is not till the north of that continent is reached that we find in the wapiti a representative of our own red deer. The red deer group extends through Europe Fie. 29.—Skull of Sivathere, from the Pliocene of India. and a large part of Central Asia, but in India and the Malayan region it is replaced by the rusine deer, like the sambar, in which the antlers (Fig. 30, a) lack the bez-tine of the red deer (ibid, b). Other marked varieties of antler are exhibited by the elk, the fallow deer, and the reindeer; but none of these approach those of the extinct Trish deer (Fig. 25), which may have an eleven feet span from tip to @2 84 RUMINANTS AND THEIR DISTRIBUTION. tip. It is noteworthy that in a few small deer in which the males have no antlers, they are compensated by having long tusks in the upper Jaw. The tiny orien- tal chevrotains, and the larger African water- chevrotain form a group quite dis- tinct from all the above, and are in some respects related to the swine. None of them have ant- lers, and the African — species is the only living ruminant in which the two elements of the front cannon- bone remain separate, thus affording another instance of the Fre. 30.—Antlers of Red (A) and Sambar (B) survival of primi- Deer. a brow-, 6 bez-, c trez-tine, d e sur- tive forms in royals. (After Blanford.) Africa. Lastly, we have the group of camels and llamas, which differ from other ruminants in that their feet form cushion- like pads, while their upper jaws possess front teeth. According to the latest researches it 1s considered probable that this group has diverged from primitive swine-like animals quite independently of the true ruminants, an inference which is very remarkable, showing that selenodont teeth, a complex stomach, the function of rumination, and the single cannon-bone, have been acquired quite inde- pendently in the two groups. The present distribution of camels and llamas is remarkable, the former being CAMELS AND LLAMAS. 85 confined to Africa and Asia, and the latter to South America. Here, however, geology comes to our aid, for in former times camel-like ruminants were abundant in North America, while the fossil camels of India show certain resemblances to the llamas, and we can thus understand how the present distribution of the two sections of the group has come about. With the possible exception of some herds of the Bactrian species in Central Asia, which are, however, probably descended from individuals escaped from captivity, wild camels are now unknown, and we cannot even determine the original habitat of the single-humped species. Thus ends our brief survey of the chief groups of living ruminants and their distribution. Did space permit, we might go on to refer to their extreme importance to man, both as sources of food and of clothing, and as beasts of draught and burden, but having reached our limits, we trust that we may have aroused in our readers an interest in these highly specialized animals which may induce some of them to devote further consideration to the subject. 86 THE TALLEST MAMMAL. CHAPTER IX. THE TALLEST MAMMAL. Comparep with their extinct allies of earlier periods of the earth’s history, it may be laid down as a general rule that the large animals of the present day are decidedly inferior in point of size. During the later portion of the Tertiary period, for instance, before the incoming of the glacial epoch, when mammals appear to have attained their maximum development, there lived elephants alongside of which ordinary individuals of the existing species would have looked almost dwarfs, while the cave-bear and the cave- hyzna attained considerably larger dimensions than their living representatives, and some of the sabre-toothed tigers must have been decidedly larger than the biggest African lion or Bengal tiger. Again, the remains of red deer, bison, and wild oxen, disinterred from the cavern and other super- ticial deposits of this country, indicate animals far superior in size to their degenerate descendants of the present day ; while some of the extinct pigs from the Siwalik Hills of northern India might be compared in stature to a tapir rather than to an ordinary wild boar. The same story is told by reptiles, the giant tortoise of the Siwalik Hills, in spite of its dimensions having been considerably exaggerated, greatly exceeding in size the largest living giant tortoises of either the Mascarene or the Galapagos Islands. The Siwalik rocks have also yielded the remains of a long- snouted crocodile, allied to the garial of the Ganges, which probably measured from fifty to sixty feet in length, whereas it is very doubtful if any existing member of the order exceeds half the former of these dimensions. If, moreover, we took into account totally extinct types, such as the megatheres and mylodons of South America, and contrasted them with their nearest living allies—in this instance the sloths and anteaters—the discrepancy in size would be still more marked, but such a comparison would scarcely be analogous to the above. Fig. 31.—The Giraffe. 88 THE TALLEST MAMMAL. To every rule there is, however, an exception, and there are a few groups of living large mammals whose existing members appear never to have been surpassed in size by their fossil relatives. Foremost among these are the whales, which, as we have seen in a previous chapter, now appear to include the largest members of the order which have ever existed. The so-called white, or square-mouthed, rhinoceros of South Africa seems also to be fully equal in size to any of its extinct ancestors; and the same is certainly true of the giraffe, which may even exceed all its predecessors in this respect. Whether, however, the fossil giraffes, of which more anon, were or were not the equals in height of the largest individuals of the living species, there is no question but that the latter is by far the tallest of all living mammals, and that it was only rivalled in this respect among extinct forms by its aforesaid ancestors. Moreover, if we exclude creatures like some of the gigantic dinosaurian reptiles of the Secondary epoch, which, so to speak, gained an unfair advantage as regards height by sitting up on their hind legs in a kangaroo-like manner, and limit our comparison to such as walk on all four feet in the good old-fashioned way, we shall find that giraffes are not only the tallest mammals, but likewise the tallest of all animals that have ever existed. In the great majority of animals that have managed to exceed all their kin in height, the increment in stature has been arrived at by lengthening the hind limbs alone, and thus making them the sole or chief support of the body. In some of these cases, as among the living kangaroos and the extinct dinosaurs, the body was raised into a more or less nearly vertical position, and the required height attained without any marked elongation of the neck. In birds, on the other hand, like the ostrich, the body is carried in nearly the same horizontal position as in a quadruped, but both the hind legs and the neck have been elongated. The giraffe, however, has attained its towering stature without any such important departure from the general structure characterizing its nearest allies, and thus preserves all the essential features of an ordinary quad- ruped. Belonging, as we have had occasion to mention STRUCTURE AND HEIGHT. 89 in the preceding chapter, to the great group of ruminant ungulates, among which it is the sole living representative of a separate family, the giraffe owes its height mainly to the enormous elongation of two of the bones of the legs, coupled with a corresponding lengthening of the vertebre of the neck. As in all its kindred, the lower segment of each leg of this animal forms a cannon-bone, the nature of which has been explained in the same chapter; and in the fore limb it is the bone below the wrist (com- monly termed the knee), and the radius above the latter, which have undergone an elongation so extraordinary as to make them quite unlike, as regards proportion, the corresponding elements in the skeleton of a ruminant, such as an ox, although retaining precisely the same structure. Similarly, in the hinder limb, it is the cannon- bone below the ankle joint or hock, and the tibia or shin- bone above, which have been thus elongated. To anyone unacquainted with their anatomy, it might well appear that a giraffe and a hippopotamus would differ greatly in regard to the number of vertebre in their necks; but, nevertheless, both conform in this respect to the ordinary mammalian type, possessing only seven of such segments. Whereas, however, those of the latter animal are very broad and short, in the giraffe they are extremely long and slender, attaining in full-grown individuals a length of some ten inches. This remarkable adherence to one numerical type in the neck vertebre is, indeed, a very curious feature among mammals; the extreme contrasts in respect of form being exhibited by those of the Green- land whale, in which each vertebra is shortened to a broad disc-like shape, and the giraffe, where it is equally narrow and elongated. As regards the height attained by the male of the tallest of quadrupeds, there is, unfortunately, a lack of accurate information, and since it is probable that the majority of those now living are inferior in size to the largest individuals which existed when the species was far more numerous than at present, it is to be feared that this deficiency in our knowledge is not very likely to be remedied. By some writers the height of the male giraffe 20 THE TALLEST MAMMAL. is given at sixteen feet, and that of the female at fourteen feet, but this is certainly below the reality. For instance, Mr. H. A. Bryden states that a female he shot in southern Africa measured nearly seventeen feet to the summits of the horns, while a male measured within less than half an inch of nineteen feet; Sir S. Baker, whose experiences are derived from the north-eastern portion of the continent, also asserting that a male will reach as much as nineteen feet. From the evidence of a very large, though badly preserved specimen in the Natural History Museum, it may also be inferred that fine males certainly reach the imposing height of over eighteen feet. Although this towering stature is the most obvious external feature of the giraffe, it is not one which would of itself justify the naturalist in classing the animal as the representative of a family apart from other ruminants ; and we must accordingly inquire on what grounds such separation is made. On the whole, the most distinctive structural peculiarity of the giraffe is to be found in the nature of its horns. These, as mentioned in the preceding chapter, are quite unlike those of any other living ruminant, and take the form of a pair of upright bony projections arising from the summit of the head in both sexes, and completely covered during life with skin. In the immature condition separate from the skull, these horns become in the adult firmly attached to the latter; while below them, in the middle of the forehead, is another lower and broader protuberance, sometimes spoken of as a third horn. Obviously, these horns—for want of a better name—are quite unlike the true horns of the oxen and antelopes, or the antlers of the deer; and this essential difference in their structure is alone quite sufficient to justify the reference of the giraffe to a family all by itself. When, however, we come to inquire whether the creature is more nearly akin to the deer or to the hollow-horned ruminants (as the oxen, antelopes, and their allies are termed), we have a task of considerable difficulty. Relying mainly on the structure of its skull, and its low-crowned grinding teeth, which are invested with EXTERNAL FORM. 91 a peculiar rugose enamel having much the appearance of the skin of the common black slug, some naturalists speak of the giraffe as a greatly modified deer. A certain justification for this view is, indeed, to be found in the circumstance that the liver of the giraffe, like that of the deer, is usually devoid of a gall-bladder. Occasionally, however, that appendage, which is so characteristic of the hollow-horned ruminants, makes its appearance in the giraffe, thus showing that no great importance can be attached to it one way or another. On the other hand, in certain parts of its soft anatomy, the creature under con- sideration comes very much closer to the antelopes and their kin than to the deer. It would appear, therefore, on the whole, that the giraffe occupies a position midway between the deer on the one hand and the antelopes on the other ; while as neither of these three groups can be regarded as the direct descendant of either of the other two, it is clear that we must regard all three as divergent branches from some ancient common stock. In general appearance, the giraffe is too well known to require description, but attention may be directed to a few of its more striking external peculiarities. One remarkable feature is the total lack of the small lateral or spurious hoofs, which are present in the great majority of ruminants, and attain relatively large dimensions in the reindeer and musk-deer. Indeed, the only other members of the whole group iv which these hoofs are absent are certain antelopes; but this absence cannot be taken as an indication of any affinity between the latter and the giraffe, since it is most probably the result of independent development. Equally noticeable are the large size and prominence of the liquid eyes, and the great length of the extensile tongue; the former being obviously designed to give the creature the greatest possible range of vision, while the extensibility of the latter enhances the capability of reaching the foliage of tall trees afforded by the lengthened limbs and neck. In comparison with the slenderness of the neck, the head of the giraffe appears of relatively large size; but this bulk, which is probably necessary to the proper working of the long tongue, is 92 THE TALLEST MAMMAL. compensated by the extreme lightness and porous structure of the bones of the skull. Lastly, we may note that the long tail, terminating in a large tuft of black hairs, is a feature unlike any of the deer, although recalling certain of the antelopes. Somewhat stiff and ungainly in its motions—the small number of vertebre not admitting of the graceful arching of the neck characterizing the swan and ostrich—the giraffe is in all parts of its organization admirably adapted to a life on open plains dotted over with tall trees, upon which it can browse without fear of competition by any other living creature. Its wide range of vision affords it timely warning of the approach of foes; from the effect of sand-storms it is protected by the power of automatically closing its nostrils; while its capacity of existing for months at a time without drinking renders it suited to inhabit waterless districts like the northern part of the great Kalahari desert. And here we may mention in passing that the camel has gained a reputation for being adapted for a desert-life above all its allies which is not altogether deserved. It is true, indeed, that a camel can and does make long desert journeys, but these can only be maintained during such time as the supply of water in its specially constructed stomach holds out, and when this fails there is not an animal that sooner knocks up alto- gether than the so-called ‘ship of the desert.” Did their bodily conformation and general habits adinit of their being so employed, there can indeed be little doubt that the giraffe and some of the larger African antelopes, which are likewise independent of water, would form far more useful and satisfactory beasts of burden for desert travelling than the, to our mind, somewhat over-rated camel. Returning from this digression, it must be mentioned that when we speak of the giraffe being independent of water, we by no means intend to imply that it never drinks. On the contrary, during the summer this ruminant, when opportunity offers, will drink long and frequently; but it is certain that for more than half the year, in many parts of southern Africa at least, it never takes water at all. In certain districts, as in the northern DISTRIBUTION. 93 Kalahari, this abstinence is, from the nature of the country, involuntary ; but according to Mr. Bryden, the giraffes living in the neighbourhood of the Botletli river—their only source of water—never drink therefrom throughout the spring and winter months. When a giraffe does drink, unless it wades into the stream, it is compelled to straddle its fore-legs far apart in order to bring down its lips to the required level, and the same ungainly attitude is perforce assumed on the rare occasions when it grazes. There is yet one other point to be mentioned in con- nection with the adaptation of the giraffe to its surroundings before passing on, and this relates to its coloration. When seen within the enclosures of a menagerie— where, by the way, their pallid hue gives but a faint idea of the deep chestnut tinge of the dark blotches on the coat of a wild male—the dappled hide of a giraffe appears conspicuous in the extreme. We are told, however, that among the tall kameel-dhorn trees, or giraffe-mimosas, on which they almost exclusively feed, giraffes are the most inconspicuous of all animals; their mottled coats harmonizing so exactly with the weather-beaten stems and with the splashes of light and shade thrown on the ground by the sun shining through the leaves, that at a com- paratively short distance even the Bushman or Kaffr is frequently at a total loss to distinguish trees from giraffes, or giraffes from trees. At the present day, it is hardly necessary to mention, the single species of giraffe is exclusively confined to Africa, not even ranging into Syria, where so many other species of animals otherwise characteristic of that continent are found. This restricted distribution was, however, by no means always characteristic of the genus; for during the Pliocene period extinct species of these beautiful animals roamed over certain parts of southern Europe and Asia. The first of these extinct giraffes was discovered by Falconer and Cautley many years ago in that marvellous mausoleum of fossil animals, the Siwalik Hills of north-eastern India ; remains of the same species bcing subsequently brought to light in the equivalent deposits of Perim Island, in the Gulf of Cambay, and likewiseinthe Punjab. A second 94, THE TALLEST MAMMAL. species has also left its remains in the newer Tertiary rocks of Pikermi, near Athens ; while those of a third have been disinterred in China, and a fourth in Persia. It was, indeed, believed for a long time that France also was once the home of a member of the genus, but the specimen on which the determination was based is now known to be a jaw-bone belonging to the existing species. Although we are, unfortunately, unacquainted with the geology of the greater part of Africa, the foregoing evidence points strongly to the conclusion that giraffes (together with ostriches, hippopotami, and certain peculiar antelopes) are comparatively recent emigrants into that continent from the north-east ; but,as we have previously had occasion to mention, the reason why all these animals have totally died out in their ancient homes is still one of the darkest of enigmas. Unknown in the countries to the north of the Sahara, as well as in the great forest regions of the west, which are unsuitable to its habits, the giraffe at the present day ranges from the north Kalahari and northern Bechuana- land in the south, through such portions of eastern and central Africa as are suited to its mode of life, to the southern Sudan in the north. Unhappily, however, this noble animal is almost daily diminishing in numbers throughout a large area of southern and eastern Africa, and its distributional area as steadily shrinking. Whether it was ever found to the south of the Orange river and in the Cape Colony may be a moot point, although, according to Mr. Bryden, there are traditions that it once occurred there. Apart from this, it is definitely known that about the year 1813 these animals were met with only a little to the north of the last-named river; while as late as 1886 they were still common throughout the Transvaal, and more especially near the junction of the Marico with the Limpopo river. Now their last refuges in these districts are the extreme eastern border of the Transvaal (where only a few remain), and the district lying to the north of Bechuanaland and known as Khama’s ‘country, or Bawangwato, together with the northern Kalahari. Even here, however, their existence ig ITS GROWING SCARCITY. 95 threatened, as there isa proposal to put down tube-wells in the waterless Kalahari, which, if successfully accomplished, will open up the one great remaining stronghold of the animal to the merciless hunter. Unless, therefore, efficient and prompt measures are taken for its protection, there is but too much reason to fear that the giraffe will ere long be practically exterminated from this part of Africa ; although, fortunately, it has a prospect of surviving for many years to come in the Sudan and Kordofan. The great majority of the giraffes killed at the present day in southern Africa are shot solely for the sake of their skins, which are now, owing to the practical extermination of rhinoceroses south of the Zambesi, and the ever-increasing scarcity of the hippopotamus, used in the manufacture of the formidable South African whips known as jaimboks. The value of a skin usually varies, according to size and quality, from £2 10s. to £4, although they have been known to fetch £5 apiece; and it is for the sake of such paltry sums that one of the noblest and most strange of mammals stands in imminent danger of extermination ! We may conclude this notice by mentioning that although the giraffe was familiar to the Romans of the time of the empire, by whom it was known as the camelopard, it appears to have been almost completely lost sight of in Europe in later times till the closing decades of the eighteenth century, although a single example is stated to have been exhibited alive in Florence some four centuries ago. With that exception, it seems to have been generally regarded as a fabulous animal until one was shot near the Orange river in 1777 by an Englishman, and another by the French naturalist, Le Vaillant, in 1784. From that time onwards our knowledge of the animal and its habits gradually increased, although it was not till the spring of 1886 that four living specimens from the Sudan were brought alive to London, where some of their descendants lived continuously till 1892, since which date the species has been unrepresented in the Regent’s Park. 96 LEMURS. CHAPTER X. LEMURS. In a previous chapter we have had occasion to refer to Africa as an archaic kind of land containing types of mammalian life which, while formerly widely spread over the Old World, are now restricted to that continent. If this preservation of ancient types be highly characteristic of continental Africa, still more markedly is it so of the large island of Madagascar, lying off its eastern coast. In Africa itself many of the ancient types are more or less closely allied to other living mammals, and most of them belong to orders which are abundantly represented in other parts of the world. Very different is, however, the case with Madagascar, of which the great peculiarity is that it has preserved to us a whole fauna of those remark- able animals known as lemurs, which are represented else- where only in Africa and the Oriental region, and there by a comparatively small number of species belonging to genera totally distinct from those found in Madagascar. To put the matter more clearly, it may be stated that out of a total of thirteen genera of living lemurs no less than eight are absolutely confined to the island of Madagascar, while the remaining five are distributed over Africa, India, and the Malayan islands; two out of the five being African, one Indian, and two Malayan. The disproportion is, however, not even adequately expressed by the above statement, since, while most of the Malagasy genera are represented by a considerable number of species, of those found in other regions only one has more than two species, while two out of the other four have but a single species each. Lemurs are, indeed, in every sense the characteristic mammals of Madagascar, being far more common in its woods and coppices than are squirrels in those of this country. So common are they said to be in certain parts of the island that, according to the French traveller, THEIR DISTRIBUTION. 97 M. Grandidier, it is impossible to beat through a single copse without turning out at least one of these strange creatures. In order to arrive at the true reason of the present dis- tribution of any group of animals, it is always necessary to consult the records of geology; and it appears from these that while lemurs were unknown both in Europe and North America during the Pliocene and Miocene divisions of the Tertiary period, when we reach the upper part of the Eocene epoch we find their remains occurring in company with those of the extinct anoplotheres and paleotheres, or other allied animals, in both the eastern and western continents. It is, however, hardly necessary to observe that the whole of these early lemurs belonged to genera which are quite distinct from any of those living at the present day, although one of them appears to have nearly been allied to the African group. The discovery of these extinct lemurs, which is but com- paratively modern, at once reveals the fact that this group of animals is a very ancient one, and one formerly widely spread over the globe, and represented by at least one species in our own island. Not many years ago it was sought to explain the present peculiar distribution of lemurs by the supposition that a large island or continent formerly existed in the Indian Ocean; and the name Lemuria was suggested, appropriately enough, for this hypothetical land. From this presumed ancestral home it was considered that the lemurs had spread on all sides, some to find a refuge in the Malayan islands, others in the forests of Ceylon and southern India, but the larger number in Madagascar and Africa. Unfortunately, how- ever, for a very pretty theory, the geologists had a word to say on the matter; and this word was to the effect that Lemuria could not possibly have existed at the time its presence was required for the needs of the theory. With our fuller knowledge of fossil lemurs any such hypothetical land is, however, quite unnecessary to explain the present distribution of the group. At or about the time these animals existed in Europe there is little doubt that they were also spread over Africa, which there is H 98 LEMURS. good evidence to show was formerly connected by land with Madagascar. We do not, indeed, yet know how the ancient lemurs of Europe got into Africa, but when once there it is certain that they were ultimately cut off from Europe by a sea which stretched from the Atlantic to the Bay of Bengal in upper Eocene times. For some time afterwards, during which Africa and Madagascar still had a free communication, the large mammals characteristic of Miocene Europe were unknown in the lands to the south of this great sea, where lemurs and other lowly animals flourished in security. During some portion of this period Madagascar must have become separated from Africa, while an upheaval of land once more brought Africa into connection with Europe and Asia, and thus allowed it to be overrun by the great hoofed and carnivorous mammals which had hitherto existed only to the north of the dividing sea. This incursion of large quadrupeds at once put a final stop to the supremacy of the lemurs in Africa, where only a few species have since managed to survive by the aid of their nocturnal habits. In Madagascar, however, where there are still no large quadrupeds, and where the only carnivores are certain civet-like animals, the lemurs have continued to flourish in full exuberance, and the existing state of that island thus offers to our view a picture of what must have been the condition of Africa previous to the advent of its present fauna from the north. The few lemurs now inhabiting the Indian and Malayan region are, doubtless, also survivors from the original central home of the group, which have found safety in the dense forests of the regions they inhabit. A good deal more might be said on the subject of the past and present distribution of lemurs, but by this time the reader will probably be impatient to know something of the characteristics of the animals thus designated by naturalists. Of course comparatively little can be said on this subject in an essay of the present length, and it unfortunately happens that the lemurs, as a whole, do not present any very strongly marked single external feature by which they can be distinguished at a glance from all other mammals. It is, however, only with some of the CHARACTERISTICS. 99 lower monkeys of the New World that they could possibly be confounded by observant persons (although we have heard of some members of the family being mistaken for sloths) ; but the distinct geographical distribution of the two groups renders it improbable that any confusion is likely to arise between them. The nearest relations of the lemurs are undoubtedly the monkeys, and most naturalists in this country are now agreed in regarding the former as EN \ \ Fia. 32.—The Slender Loris, in waking and sleeping postures, with figures of the arm and leg. (From Sir J. E. Tennent.)* representing a primitive group of the order (Primates) which includes the latter. Lemurs may always be dis- tinguished from monkeys and apes by certain features in their skulls, as well as by several peculiarities in their * We are indebted to Messrs. Longmans for the use of this figure. H2 100 LEMURS. internal structure ; but as these require a certain amount of anatomical knowledge for their proper comprehension, we must ask our readers to take it on trust that such differences do exist. Externally, lemurs differ from ordi- nary monkeys by their more or less fox-like and immobile countenances, but since the marmosets of South America (which are a lowly type of monkey) resemble them in this respect, this character does not afford an absolute distinction between the two groups. All lemurs are further characterized by having the second toe terminating in a long pointed claw, as shown in the right hand lower figure of our first illustration, whereas in ordinary monkeys the same toe has a flattened nail. Unfortunately, how- ever, the marmosets have also a pointed claw on the toe in question, so that this character does not afford an absolute point of distinction between the two. On examining the upper teeth of the lemurs it will, however, be found that, except in the aye-aye of Madagascar, the first pair of incisor or front teeth are separated from one another in the middle line by a distinct gap, whereas in all monkeys they are, as in ourselves, in contact. Now as the aye-aye differs from all monkeys in having its first upper incisors of the chisel-like form characteristic of rodents (rats, beavers, &c.), the upper front teeth of the lemurs will serve to distinguish them absolutely from the whole of the monkeys. In appearance the various kinds of lemurs differ greatly from one another, some of them looking not unlike monkeys; while others, as the one represented in Fig. 32, are characterized by their long and slender limbs, enormous eyes, and general ghostly form. Then, again, while some of them are furnished with long tails, otbers are destitute of these appendages; and the common cat-lemur of Madagascar is distinguished from all the rest by the bold alternating rings of black and white with which the tail is ornamented. The last-named species is further peculiar in living chiefly among rocks, whereas the others are arboreal and mainly nocturnal in their habits. It is from these nocturnal habits, coupled with the large eyes, ghostly appearance, and stealthy movements, characteristic of HABITS, ETC. 101 many of the species, that Linnzus was induced to propose the name of lemurs (from the Latin term for evil spirits) for this group of animals—a name which, in the absence of any vernacular title, has been adopted as their ordinary designation. None of the lemurs are of large size, the length of the head and body in the largest species being only about two feet, and some of them are not larger than rats. They are all excellent climbers, and the majority spend the day sleep- ing either in the hole of a tree, in a specially constructed nest, or rolled up in a ball after the manner shown in our first illustration. Their food consists of leaves and fruits, birds and the eggs, reptiles and insects, and occasionally honey or sugar-cane ; and most of them spend the whole of their time im trees, rarely, if ever, descending to the ground. Some of the larger species inhabiting Madagascar are, however, an exception in this respect, as well as in their diurnal habits, and they may sometimes be observed in numbers jumping across the plains from wood to wood in their own peculiar fashion, when it is necessary to seek fresh food. From the structure of their brains and other parts of their organization, it is evident that the lemurs hold a very low place in the mammalian class, although their near relatives, the monkeys and apes, occupy the highest position. It is probable, indeed, that the modern lemurs are the descendants of the ancient ancestral stock from which monkeys have originated, and since they them- selves are also nearly related to the so-called insectivores (shrews, moles, hedgehogs, &c.), while the latter may have been directly descended from marsupials (opossums, &c.\, we see how close is the connection between the very highest and the very lowest representatives of the mammalian class. The low position of the lemurs in the zoological scale is in harmony with their antiquity and their peculiar geographical distribution, and it is note- worthy that both in Madagascar and in Africa lemurs are accompanied by certain insectivores of a very low degree of organization, and unlike those found in any other part of the world. The limits of this essay render our notice of the various 102 LEMURS. kinds of lemurs necessarily very brief, and our chief attention will accordingly be directed to some of the more peculiar and interesting forms. Among the Madagascar lemurs are included a group known as indris, or, in the vernacular, sifakas, and containing the largest of all these animals. These sifakas are distinguished by the circum- stance that all the toes of the foot, except the first, are united together at their bases; and they are further characterized by their parti-coloured fur, in which white, black, and various shades of brown and orange predo- minate. Most of them have long tails, but in one species this appendage is represented by a mere stump. They live in smail parties in the woods of Madagascar, and feed entirely upon vegetable substances. By the aid of their powerful limbs they are able to take enormous flying leaps —sometimes as much as thirty feet in length—from tree to tree, and when passing from one clump of trees to another on the ground they hop on their hind limbs, with their arms raised above their heads, in a series of ’short jamps, when they are said to present the most ludicrous and grotesque appearance. They are largely diurnal in their habits, although sleeping during the heat of the day. The true lemurs, which are likewise confined to Mada- gascar and some of the adjacent Comoro islands, differ from the sifakas in having thirty-six in place of thirty teeth, by their perfectly free toes, and their less elongated hind limbs. They all have long tails, and the best known species is the above-mentioned cat, or ring-tailed lemur, easily recognized by the feature from which it derives its second name. This animal, which is often exhibited in menageries, is about the size of a domestic cat, and is peculiar in frequenting rocks rather than trees, in at least certain districtsin Madagascar. The black lemur, in which the male is black and the female red, is a nearly allied species ; and there are also several others. The females of these lemurs have a peculiar habit of carrying their young clinging to the under surface of their body, with the head on one side and the tail on the other; but this strange position is only maintained for a certain time, after which GALAGOS AND SLOW-LEMURS. 103 the young creature mounts upon its parent’s back, where it remains until able to shift for itself. A third group is formed by the galagos, which differ from the true lemurs in having two of the bones of the ankle greatly elongated, so as to make this segment of the limb much longer than ordinary. They all have long and bushy tails, and are mostly of small size, some of them being even smaller than a rat. The galagos are divided into two groups—one confined to Madagascar and the other to Africa. The former, or dormouse-lemurs, are peculiar in that during the hot dry season several of the species undergo a kind of hibernation, coiled up in the hollow of some tree, and in order to prepare themselves for such a protracted fast they accumulate on their bodies a large store of fat. The true African galagos, in addition to other features, differ from the last in that their large ears are capable of being partially folded up, somewhat after the fashion obtaining in the common long-eared bat. Like the dormouse-lemurs, they are purely nocturnal, and when on the ground they hop after the manner of kangaroos, the elongation of the bones of the ankle doubtless sub- serving this kind of movement. Widely different from all the above are the curious slow- lemurs of Asia and Africa, of which an Asiatic species is represented in Fig. 32. These lemurs are characterized by the index finger of the hand being either very short or rudimentary, and likewise by their tail being similarly abbreviated. The Asiatic representatives of this group, of which there are two genera and about four species, have the usual three joints to the index finger, which is, however, extremely short, and no trace of a tail. It was one of these lemurs which doubtless suggested to Linnzus his name for the whole group, as their movements are slow and deliberate in the extreme, their eyes large, and their habits completely nocturnal. The Asiatic forms are known by the name of loris. The common loris, with some allied species, extends from Burma through the Malayan region to Siam and Cochin China, and is a solitary animal inhabiting the depths of the forests. The strange and weird little animal represented in our first illustration is 104 LEMURS. an inhabitant of southern India, and is commonly known as the slender Joris. It differs from the common loris by its much larger eyes, which are separated from one another only by a very thin partition, as well as by its slender body and limbs; and in consequence of these and other points of difference is referred by naturalists to a distinct genus, of which it is the sole representative. This beauti- ful little creature is about the size of a squirrel, and is of a yellowish-brown colour. It occurs in the forests, usually in pairs. Sir E. Tennent observes that “the naturally slow motion of its limbs enables the loris to approach its prey so stealthily that it seizes birds before they can be alarmed by its presenre. During the day, the one which I kept was usually asleep in the strange position repre- sented in the figure; its perch firmly grasped with both hands, its back curved into a ball of soft fur, and its head hidden deep between its legs. The singularly large and intense eyes of the loris have attracted the attention of the Singhalese, who capture the creature for the purpose of extracting them as charms and love-potions, and they are said to effect this by holding the little animal to the fire till the eyeballs burst.” I once brought a pair of these little creatures from Madras to Calcutta, and during the voyage they lived chiefly on plantains and bread-and-milk. In West Africa the slow lemurs are represented by the potto and the smaller awantibo, both of which differ from the loris by the reduction of the index finger to a mere stump, and also by the presence of a distinct tail, which is of greater length in the former than in the latter species. These animals resemble their Asiatic cousins in their habits, but are even more deliberate in their movements ; and while the potto appears to be not uncommon, the awantibo is of extreme rarity. The whole of the preceding species are included by naturalists in a single tamily, but the two following representatives of the group are so different from all the others that each is made the type of a distinct family. The first of these two aberrant creatures is the weird tarsier, of the forests of Sumatra, Borneo, Celebes, and some of the Philippine islands. This animal derives its name from TARSIER, 105 its greatly elongated ankle (tarsus), and is rather smaller than an ordinary squirrel, with large ears, enormous eyes, and a long tufted tail. Dr. Guillemard, who when in Celebes was fortunate enough to obtain a living tarsier, writes that ‘these little creatures, which are arboreal Fie. 33.—The Tarsier. (From Guillemard’s ‘“ Cruise of the Marchesa.” * and of nocturnal habits, are about the size of a small rat, and are covered with remarkably thick woolly fur, which is very short. The tail is long and covered with hair at * Messrs. Murray have kindly lent this figure. 106 LEMURS. the root and tip, while the middle portion of it is nearly bare. The eyes are enormous, and indeed seem, together with the equally large ears, to constitute the greater part of the face, for the nose and jaw are very small, and the latter is set on, like that of a pug dog, almost at a right angle. The hind limb at once attracts attention from the great length of the tarsal bones, and the hand is equally noticeable for its length, the curious claws with which it is provided, and the extraordinary disc-shaped pulps on the palmar surface of the fingers which probably enable the animal to retain its bold in almost any position. This weird-looking little creature we were unable to keep long in captivity, for we could not get it to eat the cockroaches which were almost the only food with which we could supply it.” Our brief account of the lemurs appropriately closes with the strangest of them all—the now well-known aye- aye of Madagascar. This remarkable animal, whose systematic position was long a puzzle to naturalists, was discovered as far back as 1780, but for eighty years after that was only known in Europe by the single specimen then obtained. The aye-aye differs from all other lemurs in having but eighteen teeth, but its most marked peculiarity is to be found in the circumstance that the single pair of front or incisor teeth in each jaw have chisel- shaped crowns like those of rats and beavers, and grow continuously throughout the life of their owner. Another peculiarity occurs in the extremely long and attenuated third finger of the hand, which surpasses all the others in length, although the whole hand is remarkably elongated. It also differs from other lemurs in that all the toes of the foot, with the exception of the first, have pointed claws, and thus resemble the second toe of the ordinary kinds. In size the aye-aye may be compared to a cat, and it has a rounded and somewhat cat-like head, with a short face, and large naked ears. The tail is long and bushy, and the general colour of the fur dark brown. The aye-aye represents the extreme development of the lemur type, and it is evident that its peculiarities of structure are correlated with equally well-marked traits of AYE-AYE. 107 habit. Unfortunately, however, from the nocturnal habits and rarity of the creature itself, as well as from the absence of a sufficient number of competent observers in its native haunts, we are by no means so well acquainted with the habits of this strange creature as is desirable. It appears, however, that the aye-ayes live either in pairs or alone in the bamboo forests of Madagascar, and that they are in the habit of constructing ball-like nests of leaves placed in the forks of trees, to which they retire for their diurnal slumber. The strong incisor teeth are certainly used for ripping up the hard external cases of the sugar-canes, on which these animals delight to feed; and it is said that they are like- wise employed to tear away the bark and wood from the trunks of trees, and thus expose the burrows of wood-eating larve, which are then extracted by the aid of the thin middle finger. If any of our readers who have friends living in Madagascar can induce them to try and obtain more particulars in regard to the aye-aye, they will be conferring a real benefit on science. 108 ARMADILLOS AND AARD-VARKS. CHAPTER XI. ARMADILLOS AND AARD-VARKS. Or the four animals represented in the figures accom- panying the present chapter, three are sufficiently alike to suggest to the ordinary observer their relationship to one another, but the fourth is so utterly different, that it is difficult to point out any important character it has in common with the tbree others; nevertheless, naturalists generally regard all these four strange creatures as belonging to a single order of mammals, for which the name of Edentata is adopted. The signification of the term Edentata being toothless, the unsophisticated student would naturally be led to suppose that all the animals so named were utterly devoid of those useful but troublesome appendages. This, however, is far from being the case, the majority of the members of the group (among which are those figured here) having a considerable number of teeth. THE GROUP OF EDENTATES. 109 Still there is one feature in connection with the dentition exhibited by the whole of these so-called edentates; and this is, that teeth in the front of the jaws, corresponding to the incisors of other mammals, are totally absent. Instead, therefore, of being described as edentates, or toothless mammals, these creatures ought rather to have been named aprotodonts, or incisorless mammals. After all, however, it is not much consequence what is the proper meaning of a name, so long as we know the sense in which it is used, and there is accordingly no real objection to the employment of the term edentates, which bas obtained the almost universal sanction of zoologists. In addition to this total absence of front teeth, the edentates are further characterized by the circumstance that all their teeth (when they possess any) show no trace of the hard layer of enamel which is so characteristic and essential a constituent of those of other mammals; these teeth at no period of life forming roots, but continually growing from below. Moreover, in nearly all the eden- tates there is never any set of milk-teeth developed, although, unfortunately, this cannot be taken as a charac- teristic of the order, sinc2 such teeth occur in one of the armadillos, and also in the animal represented in our fourth figure. Premising that the edentates are quite distinct from the marsupials and egg-laying mammals, we may say, then, that the only features by which they can be collectively characterized are the want of front teeth, and the absence of enamel on those of the cheek series, while in certain rare instances they may be utterly devoid of teeth. Such characters, it must be confessed, are by po means of first importance. The mammals thus associated by these negative charac- teristics, are now chiefly confined to the southern hemi- sphere, and include the sloths, anteaters, and armadillos of South America, the pangolins, or scaly anteaters of south-eastern Asia and Africa, and the aard-varks of Africa; the true anteaters and pangolins being those in which teeth are wanting. In past times they were also represented by the gigantic megathere, and a number of 110 ARMADILLOS AND AARD-VARKS. other ailied extinct forms ranging throughout America, which in some respects serve to connect the sloths with the anteaters. This marked restriction of the existing edentates in the southern hemisphere, and their especial abundance in South America, at once stamps them as a very lowly group of animals, there being a well-marked tendency for the preservation of the humbler forms of life in the southern contents and islands of the globe. There is, indeed, a question whether the pangolins, and more especially the aard-varks of the Old World, have any real kinship with the more typical American edentates, but apart from this possibility of the artificial nature of the eroup as at present constituted there can be no doubt but that all its members are what may be called degraded types—that is to say, that instead of having advanced in the struggle for existence they have lost some of the attributes of the higher animals ; evidence of this degrada- tion being afforded by the indications above mentioned of their having been formerly provided with two sets of teeth. In saying that the edentates are lowly and degraded examples of the mammalian type we by no means intend, however, to imply that they are not admirably adapted to their own particular modes of life, or that they have not developed special structures unknown among the higher mammals. In truth, precisely the contrary is the case, since these creatures have taken to modes of life unlike those of the majority of the larger mammals, and have more or less specially modified their structure in accordance with such peculiar habits, so that they are thoroughly and perfectly suited to their environment. And we may add that it is doubtless due to these peculiarities of structure and habits that they have been enabled to survive and hold their own against the competition of the higher forms. Thus, in the first place, with the exception of a few armadillos, the whole of the edentates are strictly noc- turnal; and while the sloths spend the whole of their lives among the branches of the dense forests of South America, some of the others have taken to a burrowing subterranean life. Moreover, the armadillos and the pangolins have ANT-EATING. 111 acquired a special protection from their foes in the shape of a bony or scaly armour, which is a perfectly unique feature in the whole mammalian class. Another peculiarity of the group is that no less than three distinct sections of its members—namely, the anteaters of South America, the pangolins of southern Asia and Africa, and the African aard-varks—have taken to feed mainly or exclusively on termites, or so-called white ants. This practice obviously gives them an advantage in the struggle of existence, since with the exception of the marsupial banded anteater and the egg-laying spiny anteater of Australia (with which, of course, they do not come into competition), no other mammals are in the habit of subsisting exclusively on those insects. And we may notice here that of the three groups of termite-eating edentates, two—namely, the pan- golins and the anteaters—are those which have entirely lost their teeth ; while in the aard-varks those organs are retained. As teeth are obviously of no sort of use to animals subsisting on such a diet, we may regard the two former groups as those most specially modified for their particular mode of existence ; and it may thus be suggested that they have taken to termite-eating for a longer period than the aard-varks. A similar observation also applies to the banded and spiny anteaters of the antipodes, the former retaining a number of minute teeth, while in the latter they have completely disappeared. Needless to say, all termite-eating mammals, whether they be edentates, marsupials, or egg-layers, have extremely long, narrow, and extensile tongues with which to pick up their insect- food; but the presence of such an organ does not, of course, imply any mutual affinities between the possessors thereof, and is merely an instance of the similarity of organs arising from adaptation to a similar mode of existence. The tongue of the aard-varks is, however, far less elongated and extensile than that of the pangolins aud true auteaters; and, therefore, tends to confirm our suggestion as to the relative duration of time since the ancestors of these creatures severally took to termite- eating. Another instance of adaptation displayed by all the edentates, except the arboreal sloths, is to be found in 112 ARMADILLOS AND AARD-VARKS. the powerful and generally elongated claws or nails with which their feet are armed, such claws being obviously necessary for a fossorial subterranean existence. The aard-varks, as will be seen from our fourth figure, have, however, much shorter and blunter claws than any other member of the group; and this leads me to hazard the suggestion that, in addition to having taken at a com- paratively late period to termite-eating, these animals have not been accustomed to a subterranean life for so long a time as their reputed kindred. Having said thus much as to edentates in general, we must turn to the special consideration of the creatures whose names form the title of this chapter. The armadillos, as their name (a Spanish one) implies, are distinguished by the solid armour with which their heads and backs are protected; and it is doubtless the peculiar appearance presented by these animals to which we owe the expression ‘“ hog-in-armour.” In all the armadillo family the armour takes the form of a series of thicker or thinner bony plates imbedded in the skin covering the head and back, and overlain by horny scales ; while the under parts of the body and jimbs are hairy, and in many species a larger or smaller number of stiff hairs protrude from between the joints of the armour. This bony armour is a perfectly unique feature among existing mammals; and since each plate is ornamented with a more or less elaborate sculptured pattern, such armour when cleaned by maceration forms a most beautiful object. In the true armadillos, as the one represented in Fig. 34, the shield of armour covering the head is quite distinct from that of the body ; while the latter is divided into three distinct portions, namely, a large solid shield covering the fore-quarters, and separated by a larger or smaller number of free movable bands occupying the middle of the body from a nearly similar shield protecting the hinder portion of the animal. In our first figured example the number of the movable bands is only three, but they may vary from six to nme (Fig. 36) up to as many as twelve or thirteen in other species. In one extinct armadillo there were, however, no solid shields, the whole body being THE PICHICIAGO. 113 covered by a series of thirty-two movable bands. The latter species evidently, therefore, leads on to the rare and beautiful little creature represented in Fig. 85, which rejoices in the name of pichiciago. In this tiny animal, which is only about five inches in length, and has a pink- coloured armour above, and long silky white hair below, the armour of the head and body forms a continuous shield of horny plates underlain by very thin plates of bone, and is attached only to the middle line of the back, so that the lateral portions form a kind of cloak loosely overhanging the hairy sides of the body. The hinder end of this cloak is abruptly truncated, and beneath it the hind-quarters of the animal are protected by a solid bony shield, through a notch in the centre of which protrudes Fie. 35.—The Pichiciago. the small cylindrical tail. The pichiciago is found on sandy plains only in the western portions of the Argentine pampas. It will be seen from our illustrations that this creature also differs from the true armadillos in the I 114 ARMADILLOS AND AARD-VARKS. absence of the large external ears which form such a characteristic feature in the physiognomy of the latter. Reverting to the true armadillos, we find that the majority of the species protect themselves from attack by squatting on the ground, and tucking their hmbs within the shelter of the edges of the armour of the body, while the plated head is drawn as close as possible to the front shield. On the other hand, the species represented in Fig. 34:has the power of rolling itself up into a complete ball, like the pill-millipedes of our own country, the wedge- shaped head and tail fitting most perfectly side by side into the deep notches of the front and hind shields. Thus coiled up, the three-banded armadillo is safe from most animals except man. Trusting in this immunity from attack, this armadillo, together with two other species inhabiting the Argentine, has become almost exclusively diurnal in its habits. These diurnal habits, as Mr. W. H. Hudson, in -his charming work, ‘The Naturalist in La Plata,” suggests, may also have had the advantage of avoiding any encounters with the larger animals of prey, which are mostly nocturnal, and some of which may have been able to break through the protecting armour, more especially in the species which lack the power of rolling themselves up. Whatever advantage may have formerly accrued from these nocturnal habits before the appearance of man on the scene, is, however, now completely lost in cultivated districts, where these species stand a good chance of being completely exterminated by the hand of man. On the other hand, the six-banded peludo, or hairy armadillo (Fig. 36), of the Argentine, which differs from its cousins in preferring an omnivorous diet to one of insects, is a far wiser beast in its generation. This creature, according to Mr. Hudson, adapts itself to the conditions under which it exists, and thus stands a good chance of surviving when its fully-armoured relatives perish. “Where nocturnal carnivores are its enemies,” writes the observer mentioned, “it is diurnal; but where man appears as a chief persecutor, it becomes nocturnal. It is much hunted for its flesh, dogs being trained for the purpose; yet it 115 ARMADILLOS. Fig. 36.—Six-Banded Armadillo. i | 116 ARMADILLOS AND AARD-VARKS. actually becomes more abundant as population increases in any district.” As I have witnessed, beneath any decomposing carcase lying in the Argentine pampas the burrow of a peludo is almost sure to be found ; and it is not a little remarkable that the flesh of a creature which has such unpleasant tastes in the matter of diet, should be so eagerly sought after as an article of human consumption. Before taking leave of the peludo we must not omit to mention two other peculiar habits which are recorded of it by Mr. Hudson, since these also mark it as a creature far above the generality of its kind in point of intelligence. The first of these peculiarities is the ingenious way the creature catches mice, by approaching them with extreme caution, raising itself on its hind-quarters, and then suddenly proceeding to “sit down” on the unfortunate rodents, which become entrapped under the projecting edges of its armour. The sharp edges of the armour are also brought into requisition when this armadillo attacks a snake preparatory to devouring it; the snake being pressed close to the ground beneath the edges of the bony plates, and literally sawn to death by means of a back- wards-and-forwards motion of the body of its assailant. ‘he largest of living armadillos is one which inhabits the moist forests of Brazil and Surinam, and has a length of about thirty-six inches, exclusive of the unusually long tail, which is some twenty inches in length. These dimen- sions were, however, vastly exceeded by some extinct armadillo-like animals, of which the remains are found in the caverns of Brazil. The most gigantic of these creatures, which flourished during the Pleistocene epoch— the period par eacellence of giant mammals—is estimated to have been nearly equal in size to a rhinoceros, and has been named the chlamydothere. The armour appears to have been very like that of the true armadillos, but the bony plates measured as much as five and six inches in length, in place of little more than an inch. The teeth differed, however, from the simple conical ones of the modern armadillos, and more nearly resembled the verti- cally fluted ones characteristic of the extinct glyptodonts. Unfortunately, space does not admit of further reference AARD-VARES. 117 to the gigantic creatures from the Pleistocene of. South America, to which the latter name has been applied, all of which are distinguished from the armadillos by the armour of the body being welded into a single solid dome- like shell. Passing on to the animals whose name comes second in the title of this chapter, we have first of all to mention that the designations by which these creatures are com- monly known exhibit that remarkable want of originality in nomenclature which appears to be characteristic of Europeans when they are brougut for the first time into contact with hitherto unknown animals. Thus, whereas the Dutch Boers of South Africa applied to the creatures in question the title of ‘“aard-vark’’ (meaning “ earth- pig”), the English colonists of the Cape commonly speak of them as the ant-bear. Now, if there is any one parti- cular animal which the aard-vark (as we must perforce term the creature) is unlike, it is a bear ; while its resem- blance to a pig is only of the most distant kind. Still, however, as in the case of the order to which it belongs, we must be content to designate the animal by the name by which it is most commonly known. In appearance, aard-varks, of which there are two species, are decidedly ugly creatures, having thick ungainly bodies, a long pointed snout, enormous erect ears, and a thick cylindrical and tapering tail, nearly as long as the body. The skin is eitber almost naked, or thinly covered with bristle-like hairs. The fore-feet have but five toes, which are armed with broad and strong nails, as are the five toes of the hind limb. As we have already mentioned, almost the only feature which the aard-vark bas in com- mon with the armadillos is the absence of front teeth, and its cheek teeth are quite unlike the simple ones of the latter, as, indeed, they are dissimilar to those of any other mammals. In the first place, they are preceded by a functionless series of milk-teeth (a feature found else- where among edentates only in one species of armadillo), while in the second place the premolars are unlike the molars. The latter are composed of «a number of closely packed denticules, each furnished with a central pulp 118 ARMADILLOS AND AARD-VARKS, (From Selater, Proc. Zool. Soc.) 7.—The Ethiopian Aard-Vark, 9 € Fie. AARD-VARKS. 119 cavity, and by their close approximation forming polygonal prisms, so that a cross-section of one of these teeth presents the appearance of a pavement. No dental struc- ture among mammals is at all comparable to this, although there is some approximation to it among certain fishes. Of the two living species of aard-vark, one is confined to South Africa, while the other (represented in our figure) inhabits part of Egypt and other districts in the north- western portion of the same continent. A third species oceurs fossil in the Pliocene deposits of the Isle of Samos, but with this exception the paleontological record is silent as to the past history of these strange creatures, as to whose origin and relationship to the other animals we are at present utterly in the dark. Indeed, the aard-vark is placed among the edentate mammals chiefly because zoologists do not know where else to put it, and they take that group as a kind of refuge for the destitute. Were it not that the burdening of zoological science with new names is from all points of view to be deprecated, there is, indeed, much justification for regarding these animals as the sole representatives of a distinct order; but, although im some ways such a new departure would be convenient, I do not know that in others it would be of any great advantage. But in including them provisionally among the edentates we have to recollect that their affinities with other members of that group—not even excepting the pangolins—must be extremely remote. Aard-varks lead what would seem to us a very dull and monotonous kind of Jife, passing the whole of the day curled up in their deep burrows, which are generally ex- cavated hard by the tall pyramidal hills made by the termites, and only issuing forth at night to dig in the mounds for their favourite insect-food. Nota great many years ago it used to be said at the Cape that wherever a clump of termite hills was to be seen there an aard-vark’s burrow might be pretty confidently expected. Unfortu- nately, however, this is no longer the case, and the aard- vark of that district runs a good chance of being extermi- nated at no very distant date. This deplorable result is being brought about by the 120 ARMADILLOS AND AARD-VARKS. incessant pursuit of these animals by the natives for the sake of their hides and flesh, and also by their being dug out by Europeans for so-called sport. Their flesh is said to be excellent, and is compared to superior pork ; while the value of each hide is about fifteen shillings. This threatened extermination is a very short-sighted policy on the part of the South African farmers, to whom the aard- vark is a valuable ally, not only on account of the enormous number of termites it consumes, but likewise from the circumstance that while it is engaged in digging for these insect-pests it covers with loose earth a quantity of the seeds of grass and other pastoral herbage which would otherwise perish during the hot season. Although there is no likelihood at present of the Ethiopian aard- vark sharing the threatened fate of its southern cousin, yet the extermination of the latter would be a sad loss to zoological science, and we therefore wish every success to a movement which we hear has been set going by the Cape Farmers’ Association for the protection of this most strange and curious creature ere it be too late. Addendum.—Quite recently some remains have been discovered in the upper Eocene, or Oligocene strata of France, which are considered to indicate the existence at that period of small ancestral types of aard-varks. THE OLDEST MAMMALS. 121 CHAPTER XII. THE OLDEST MAMMALS. Up to the year 1818 it was a generally received axiom of geology that mammals were totally unknown before the Tertiary period; and that period was consequently designated the age of mammals—a name, by the way, which is still perfectly 2 appropriate, if taken to imply that these animals then, and then only, became the dominant inhabitants of the world. In that year, however, the illustrious Cuvier, during a visit to the museum at Oxford, was shown two minute jaws, carrying a number of cusped teeth, which had been obtained in the neighbouring quarries of Stonesfield, from the rock known as the Stonesfield slate, belonging to the lower part of the great Jurassic, or Oolitic, system. After careful examination, the French anatomist pronounced confidently that these two tiny little jaws, neither of which exceeded an inch in length, were those of mammals, and he further suggested that they would prove to belong to a species of opossum. Although this opmion was given in the year 1818, it does not appear that it was published till the year 1825, when the second edition of the fifth volume of the immortal “Ossements Fosiles”’ saw the light. In publishing this epoch-making notice of the occurrence of mammals in the Secondary period, Cuvier, with the usual caution of naturalists, was careful to add the proviso that everything depended on whether the specimens he saw had really been obtained from the Stonesfield slate. Unfortunately, there does not appear to be any record stating by whom, or at what date, these original specimens—now forming some of the most valued treasures of the Oxford Museum —were obtained from the Stonesfield slate; but that they did come from that formation is perfectly certain. Indeed, other specimens have been subsequently obtained from the same beds, showing certain characteristic Stonesfield 122 THE OLDEST MAMMALS. shells imbedded in the fragments of rock in which the mammalian jaws are contained. Here we may mention that the Stonesfield slate is the equivalent of the lower portion of tbe great, or Bath, oolite—a deposit which is separated from the underlying lias by the beds known as the inferior oolite. Conse- quently, the mammial-yield- ing beds are separated from the rocks of the Tertiary period, not only by the immense series of Cretaceous deposits (chalk, gault, greensands, and Wealden), but likewise by a large thickness of those belonging to the Jurassic system, such as the Purbeck and Portland oolites, the Kimeridge clay, the coral-rag, and the Oxford clay. Needless to say, no sooner was the existence of mammals in the Stonesfield slate announced than it was received with a howl of incredulity. First of all it was attempted to show that the specimens themselves did not come from Stonesfield ; and no sooner was this objection knocked on the head than doubts were raised as to the Jurassic age of the Stonestield slate itself. These, however, were equally soon disposed of, and the only thing then remaining was to dispute the mammalian nature of the fossils. This task was undertaken by the French naturalist, De Blainville, who attempted to show that the mammalian character of the specimens was not proved by the double roots of their molar teeth. In the course of the argument, great stress was laid on the circumstance that two-rooted molars were found in a gigantic animal from the Eocene of the United States, then known as Bast/osawrus, and regarded as a reptile. De Blainville was, however, here treading on very dangerous ground, for it subsequently turned out that Basilosaurus itself was really a mammal, which is now generally placed among the whales, under the name of Fra. 38.—One half of the lower Jaw of a Stonesfield Mammal. Twice natural size. The restoration of the front teeth is conjectural. STONESFIELD SPECIES. 123 Zeuglodon. The correctness of Cuvier’s original determi- nation was thus in the end triumphantly sustained, and the existence of Jurassic mammals became henceforth an established fact in geology, although the suggestion that these fossils belonged to opossums was, of course, unfounded. Passing on to the consideration of the specimens them- selves, we find that the great peculiarity of the jaws of these Stonesfield mammals (for one of which De Bilainville proposed the name of amphithere) is the excessive number of their cheek-teeth, a feature now paralleled only in the little banded anteater of Australia. This multiplicity of teeth is well shown in the jaw represented in Fig. 38, which is preserved in the museum at York, and shows upwards of nine cheek-teeth still remaiming whereas in practically all existing mammals with complex teeth, except the banded anteater, the number does not exceed seven. Other jaws were, however, subsequently discovered in Stonesfield, in which the number of cheek-teeth was considerably less; but one of these later specimens (described as the phascolothere) revealed the important fact that there were four pairs of front or incisor teeth in the lower jaw. Now since it is only among the pouched mammals, or marsupials, that more than three pairs of incisor teeth are found, while the banded anteater, with its numerous cheek-teeth, is a member of the same group, it became a very natural conclusion that the Stonesfield mammals were likewise marsupials. Support was lent to this conclusion by the circumstance that, with the exception of the egg-laying mammals, or monotremes, the marsupials are the lowest of all living mammals. And indirectly some further support to this view is afforded by the fact that Australia still retains other forms of animal life allied to those which were living in Europe during the period of the Stonesfield slate. For instance, it is in the Australian seas alone that there still survives the solitary representative of the beautiful genus of bivalve shells known as T’rigonia, which were so especially abundant in the oolites; while it is also there alone that swims the Port Jackson shurk, whose 124 THE OLDEST MAMMALS. mouth is armed with a pavement of crushing teeth, recalling those of many of its Jurassic forerunners. Moreover, in the presence of numerous cycads among its flora, Australia again recalls the Jurassic epoch of Europe ; and it has accordingly been suggested that modern Australia may be regarded as a kind of direct survival from Jurassic times. Before, however, we can say any- thing more as to the affinities of the Stonesfield mammals, we must turn our attention to subsequent discoveries of mammalian remains in other formations. The first of these discoveries was made in the year 1847, by Professor Plieninger, of Stuttgart, who obtained certain minute teeth from a bone-bed near that city belonging to the upper part of the Triassic period, which were declared to be mammalian, and for the owner of which the name Microlestes was proposed. Now, as the trias lies below the lias, the existence of mammalian Jife was by this discovery carried back at one bound very nearly to the commence- ment of the Secondary period. Subsequently, mammalian remains were obtained from the trias of Somerset, which proved to belong to the same genus as those from Stuttgart, while others of a different type were found in the equivalent deposits of North America. Fre. 39.—Lower Jaw of an American Jurassic Mammal ; twice natural size. (After Marsh.) A little later, the year 1854 was made memorable by the first discovery of mammalian remains in the freshwater Purbeck strata of Dorsetshire, belonging to the very top of the Jurassic system; from which formation in subsequent AMERICAN FORMS. 125 years a vast number of such remains were obtained, through the energy of the Rev. P. B. Brodie and Mr. Beccles. All these specimens were obtained from a single bed, and many of them indicated forms more or less closely allied to those from Stonesfield and Stuttgart. It was thus shown, once for all, that mammatian life must have been locally abundant throughout the Jurassic period. This conclusion was subsequently amplified by the discovery in the upper Jurassic rocks of North America of a whole Fie. 40.— Lower Jaw of Triconodon ; twice natural size. (After Marsh.) host of small mammals very closely allied to those from Dorsetshire, a large number of which have been described by Prof. O. C. Marsh, some of whose figures are here reproduced. Many of these small Jurassic mammals (Figs. 39 and 40) were evidently carnivorous, and such carnivorous forms exhibited two distinct types of dentition. In one of them (Fig. 39) there was a numerous series of cheek-teeth behind the tusk, or canine (a), each of which carried three cusps arranged in a triangle; while in the other type (Fig. 40) the cheek-teeth were fewer in number, and had the three cusps on their crowns ranged in the same line. From this peculiarity the animal to which the second type of jaw belonged was appropriately named Triconodon. The first type corresponds to the amphithere of the Stones- field slate, while the second is more like the phascolothere of the same formation. In Fig. 40 it will be observed that there is a pecu- liar groove (g) running along the inside of the jaw, and 126 THE OLDEST MAMMALS. since a similar groove is found among existing mammals only in the banded anteater and certain other carnivorous marsupials, we have pretty conclusive evidence that Tricon- odon and its allies were really marsupials. There can also be but little doubt that the species of the amphitherian type (I avoid mentioning the numerous genera of these animals) are likewise members of the same order. It is, however, quite possible that some of the Jurassic mammals of Dorsetshire and North America may be more nearly allied to that primitive group of mammals known as Insectivores, among which are included the mole, the shrew, and the hedgehog of Europe, as well as the more generalized tenrec of Madagascar, and many other peculiar creatures. All these insectivores are of a very low grade of organization, and the result of modern researches is to show that their connection with the marsupials is very close indeed. Hence it is highly likely that some of the Jurassic mammals may have been the actual connecting links between the marsupials and the insectivores ; and it is worthy of mention here that while marsupials at the present day linger on only in Australia and America, some of the most primitive types of insectivores are preserved to us in Madagascar, which is another refuge for animals of a low grade of organization. There is yet another type of mammal found in the English and American Jurassics, to which the Micro- lestes of the trias also appears to belong, which has given rise to Fie. 41.—Lower Jaw of Plagiaulax ; natural size and enlarged. (After Marsh.) a vast amount of discussion among the paleontologists. These re- markable mammals are mostly of very minute size, and were long known only by their lower jaws, of which a specimen is represented in Fig. 41, from which it will HERBIVOROUS TYPES. 127 be seen at a glance that the dentition is quite different from that of either of the carnivorous types figured above. The lower teeth comprise a single large incisor (a), behind which were either three or four tall premolar teeth with cutting edges, and marked on the sides with a number of oblique grooves, from which the name Plagiaulax was taken. When unworn, these grooves extended along the whole outer surface of the teeth, but when the teeth had been long in use (as in our figure) the groovings became worn away from the sides. Behind these four premolars are two smaller molar teeth, with the summits of their crowns warked by a single longitudinal groove bounded by prominent ridges. Now it was argued at first that this very peculiar type of dentition indicated carnivorous habits in the owners thereof; but it was subsequently pointed out that the existing rat-kangaroos of Australia (of which the front of the skull is shown in Fig. 42) presented a some- what similar type of tooth-structure. Thus, the last pre- mnolar tooth (p.m.) of the rat-kanga- roo has a cutting- crown marked with a number of parallel grooves ; while each half of the lower jaw terminates in a single large incisor not unlike that of the Jurassic Plagiau- lax. Hence it was argued—and, in our opinion, argued rightly—that as the living form is herbivorous, the same must have been the case with the extinct one. When, however, it was also urged that the rat-kangaroo and Plagiaulax were closely allied animals, important differences between the two were overlooked. For instance, as will be apparent from the figures, while in the former there was but one grooved tooth, in which the grooves are vertical, in the latter there were usually three or four Fie. 42.—Jaws and Teeth of Rat-Kangaroo. 128 THE OLDEST MAMMALS, such teeth in which the grooves are oblique. Moreover, whereas the recent form was provided with four molar teeth (m 1—m 4), the fossil had but two such teeth ; while the form of these teeth was quite unlike in the two. Hence, when we add that there are other important differ- ences between them (into the consideration of which it would be difficult to enter here), it will be apparent that the view of regarding Plagiaulax as a near ally of the rat- kangaroo was the result of attaching too much importance to resemblances, and overlooking differences. Indeed, such resemblances as do exist between the two may be regarded merely as a well-marked instance of the phenomenon of parallelism, treated of in an earlier chapter. Taking it, then, as proved that Plagiaulax is not a near ally of the rat-kangaroo, we have to consider whether it can be affiliated to any other group of existing mammals. Before doing so, we have, however, to mention that there are certain ghee Secondary mammals allied to Plagiaulaz, in which the whole of the cheek-teeth are hke the true molars of the latter. We have already stated that in Plagiaulux the lower molars have a median longitudinal groove, and it may be added that the ridges bordering such grooves are surmounted by a number of small tubercles. In the upper jaw, if we may judge by some allied genera, the molars had three such tubercular ridges, separated by two grooves. Similar molars occur in the skull represented in Fig. 43, which is that of a mammal discovered a few years ago in the Secondary rocks of South Africa, and apie by Sir R. Owen Tritylodon ; but it will be noticed that there is no trace of the cutting and obliquely-grooved premolar teeth of Pla- giaulax, the premolars being like the molars. Detached molars of similar type have been found in the Trias of Stuttgart, and others occur in the Stonesfield slate. Moreover, in Dorsetshire and North America, there are certain nearly allied mammals (Bolodon) in which the upper molars have only two, in place of three, longitudinal ridges of tubercles. These forms, if other. proofs were wanting, clearly show, indeed, that the resemblance between Plagiaulaa and the rat-kangaroo is not a genetic CRETACEOUS TYPES. 129 one. When, however, the molar teeth of the type in which there are but two longitudinal rows of tubercles are compared with the transitory teeth of the Australian duck- bill, a certain resemblance can be detected between the two, which seems sufficient to indicate that in Plagiaulax and Tritylodon we have to do in all probability with ancient types of egg-laying mammals. Till within the last few years the Cretaceous period formed a complete gap as regards the history of mammals; and seeing that in Europe, with the excep- tion of the Wealden, the rocks of this system are mainly of marine origin, while some of them, like the chalk, were laid down in seas of considerable depth, this absence of mammalian remains is not to be wondered at. In the United States the condition of things is, however, very different. There the upper- most Cretaceous rocks are Pie, 40 Vader pat ok the of fresh-water origin, and gkull of a South African Secondary constitute a series known Mammal. Two-thirds natural size. as the Laramie, which is in intimate connection with the lower part of the Tertiary, and has yielded the extraordinary horned dinosaurs. From these Laramie Cretaceous rocks Prof. Marsh has succeeded in obtaining a quantity of teeth of mammals, although these are, unfortunately, mostly found detached. These teeth indicate mammals closely allied to Plagiaulaz of the Jurassic, and also others of a carnivorous type related to Amphitherium, or some of the many-molared carnivorous forms from the Dorsetshire Purbeck. Mammals of the Triconodont type—that is, those with the three cusps of the molars in a straight line—seem, however, by this time K 130 THE OLDEST MAMMALS. to have totally disappeared. At a still later date a single tooth of the Plagiaulaz type, as well as one allied to Bolodon, have been described from the English Wealden, indicating that at that epoch the Purbeck mammals still survived in Europe, and leading to the hope that future researches will yield us further evidence of the European mammalian fauna of the Wealden. The present state of our knowledge, therefore, shows that from nearly the lowest beds of the Secondary period till the close of that vast epoch, there existed a numerous fauna of small mammals distributed over a large portion of the globe, and displaying a remarkable persistence of nearly similar type. It is further evident that such of these mammals as exhibit a carnivorous type of dentition appear to be allied to the more primitive of the existing marsupials, although some of them may be more nearly related to the almost equally low insectivores. On the other hand, those which exhibit what appears to be an herbivorous modification of dental structure, if they are related to any living forms, appear to have an affinity with the modern egg-laying mammals of Australia. Now the latter, together with the marsupials and insectivores, being the lowest representatives of mammalian life at present existing, are precisely such mammals as we should natu- rally have expected to have been foreshadowed by more or less nearly allied forms in the Secondary rocks; and, therefore, in this respect, theoretical paleontology is, so far as our present knowledge goes, precisely in accord with actual facts. That the few Triassic mammals at present known were the earliest representatives of the class cannot, however, be admitted for a moment, and we must accordingly look either to the lower Triassic rocks, or to those of the underlying Permian (forming the top of the Paleozoic series), for the discovery of such primitive types. Should such ever be discovered, it is to be confidently expected that they will exhibit such a combination of characters common to mammals, and certain extinct reptiles and amphibians, that it will be very hard to say under what class they will have to be ranked. TERTIARY MAMMALS. 131 Seeing that throughout the whole of the Secondary period, with the possible exception of a few lowly insecti- vores, there is no evidence of the existence of any mammals belonging to the higher placental type (under which are included all living representatives of the class save the marsupial and egg-laying groups), the reader will naturally inquire when such higher forms first made their appear- ance. We answer, with the first dawn of the Tertiary period ; for in the very lowest Eocene strata both of France and the United States there are found, side by side with small mammals allied to Plagiawlar and the marsupials of the Jurassic and Cretaceous, others, which, though still of small size, were evidently placentals. And it is very remarkable that this first definite appearance of the higher forms of mammalian life should, so far as we know, have been contemporaneous with the disappearance of so many gigantic types of extinct reptiles, such as the dinosaurs, the fish-lizards, and the plesiosaurs, which seem to have reached the end of their term of existence at or about the close of the Secondary period. Most of these early Tertiary mammals had molar teeth carrying three cusps arranged in a triangle, like their marsupial forerunners of the Secondary epoch, from which, indeed, they were probably derived; and at this comparatively early epoch the orders of mammals were but very imperfectly differentiated from one another, it being frequently difficult to decide which were carnivores and which were ungulates.