Order Vi. Cetacea

Chapter XII. Cetacea—Whales and Dolphins

Aquatic Mammalia of fish-like form; tail expanded into horizontal flukes; a fatty dorsal "fin" present in most species; anterior limbs converted into fin-like paddles; posterior limbs only represented by skeletal rudiments. Hairy covering reduced to a few isolated hairs in the neighbourhood of the muzzle. Nostrils represented by the single or double blow-hole, nearly always situated far back upon the skull. Bones of loose texture and much impregnated with oil. The skull has a greatly-developed facial portion; supra-occipital bones meeting the frontal by overgrowing, or growing in between the parietals; bones surrounding the organ of hearing loosely attached to the skull, the tympanics of peculiar cowrie-shell form. Coronoid process of mandible absent, or very feebly developed. Teeth, when present, few or numerous, always of simple conical form, with at most traces of additional cusps (Inia); if absent their place taken by whalebone. Cervical vertebrae of short antero-posterior diameter, often more or less completely welded together into a single mass. Articulations between dorsal and other vertebrae feeble. Scapula peculiarly flattened; acromion strongly developed as a rule, but arising from a slightly-marked spine; coracoid process generally strongly developed. Phalanges of digits always more numerous than in other mammals. Clavicles absent. Stomach complex, consisting of at least four and often more chambers. Lungs simple and non-lobulated. Diaphragm obliquely set and very muscular. Brain much expanded transversely and well convoluted. Testes abdominal. Teats two, inguinal in position. Placenta diffuse and non-deciduate.

The Whales and Dolphins, which constitute this order, form an assemblage which is easily characterised by reason of the fact that their affinities to other groups of Mammalia are so doubtful that they furnish matter rather for speculation than for authoritative statement. Some hold that they resemble in certain points the Ungulata; while others again see in them the culminating term of a series which commences with such a form as the Otter, and of which the Seals and Sea-lions are intermediate stages. A third opinion is that the Whales have arisen from some low mammalian stock, too primitive to be assigned to any existing order of mammals. Palaeontology, as will be seen later, throws no light whatever upon their origin. This matter has already been referred to (see p. ) in considering the position of the Cetacea.

The Whales include the most gigantic of all the orders of vertebrated animals. No creature living or extinct is so large as the Sibbald's Rorqual, which attains to a length of some 85 feet, or perhaps even rather more. On the other hand we have what are by comparison minute forms. Apart from the possibly problematical Delphinus minutus, stated to be only 2 feet in length, we have as a minimum 3 or 4 feet. The size of the Cetacea has been subjected to much exaggeration. The first duty of a Whale, observed the late Sir William Flower, is to be large; and Natural Historians, in the recent as well as in the remote past, have not hesitated to put very round numbers upon the dimensions of the larger members of the order. We may perhaps pass over Pliny's "fish called balaena or whirlpool, which is so long and broad as to take up more in length and breadth than two acres of ground," and a number of analogous exaggerations, which gradually dwindled down to the dimensions just stated of the great Rorqual. M. Pouchet has made the ingenious suggestion that the statements of the ancients may have been nearer the truth than observations of to-day would have us believe; he pointed out justly that in former times Whales were not so relentlessly pursued as during the last century; the inference being that they may have lived to a greater age, and attained a more colossal bulk. The more modern exaggerations in the dimensions of the bigger Whales are probably due to the fact that measurements have been taken, not in a straight line from snout to tail, but along the bulging sides of the Cetacean, rendered even more convex than in nature by decomposition, and by pressure due to the immense tonnage of the creature.

The Cetacea are the most perfectly aquatic of all mammals; they never leave the waters which they inhabit. It is true that legends have represented them as pasturing upon the shore—Aelian spoke of Dolphins basking in the sun's rays upon the sand; and the "Devil Fish" of California, Rhachianectes (see p. ) has given rise to improbable stories—but they are apparently only legends. Indeed a stranded Whale cannot live long, for it is unable to breathe, the comparatively feeble breast being crushed by its own weight. In accordance with the purely aquatic habit, we find a modification of the outward form of the body (and as we shall see later of many of the internal organs), which renders the Cetacea externally unlike all other mammals. The form is fish-like, the fore-limbs are paddles, the tail is expanded into two horizontal flukes, which serve to propel the creature through the water.

Fig. 180.—Killer. Orca gladiator. × 1⁄40 (After True.)

The skin is smooth and shiny, so smooth and so shiny that it has often been compared to coach leather. But nevertheless they are not entirely without that most essential character of the class Mammalia, a coating of hair. The hairy covering is, however, reduced to the very smallest proportions; it is represented by a few hairs only—so few that they can be counted with ease—in the neighbourhood of the muzzle. These hairs are not present in all Whales; they are absent, for example, in the White Whale or Beluga. When present they are not furnished either with sebaceous glands or with muscular fibres, which are such universal concomitants of the hair follicles in the Mammalia generally. This appears to be conclusive evidence that the hairs, few as they are, are still undergoing degeneration. The need for a furry coat is removed by the presence of a thick coating of fat immediately underlying the skin. This is known as the blubber, and is the main incentive to the pursuit of Whales. It must not, however, be assumed without further argument that the hair is absent because its place is taken, as a mechanism for retaining the heat, by the blubber; for the Seal tribe possess both fur and blubber. Another conceivable explanation is quite at variance with such a view of economy. It may be noticed that among Ungulates there is a tendency to lose hair, particularly among more or less aquatic forms. Thus the Hippopotamus is almost naked (as is indeed the Walrus); the Rhinoceros, too, often a frequenter of marshy soil, is almost as denuded as is the Hippopotamus. It is not, however, settled that the Whales have anything to do with the Ungulata; otherwise an additional argument might be used, that is, the secular loss of hair in some members of this group. The Hairy Rhinoceros, Rh. tichorhinus, was, as its name denotes, a hairy beast; the Mammoth was equally so. The descendants, or at least the modern representatives of both these creatures, are but scantily clad with hairs.

A final reason for the naked character of the skin in existing Cetacea is closely connected with a feature in the organisation of three or four living species which must first be described.

Some years ago the late Dr. J. E. Gray of the British Museum described from the sea, off Margate, what he considered to be a new species of Porpoise, characterised by the presence on the dorsal fin of a row of stony tubercles. As a matter of fact it was subsequently shown that the Common Porpoise has the same structures, so that there was no need for a Margate species, Phocaena tuberculifera. Moreover, in the Indian Neomeris, a close ally of the Porpoise, a more abundant calcified covering of scales exists along the whole back of the animal. These plates, it has been discovered, are larger in the foetus, a fact which naturally points to their being an inheritance from the past, now undergoing retrogressive changes. Such a way of looking upon the facts is confirmed by the finding, many years ago, by the naturalist and physiologist Johannes Müller, of bony plates in connexion with the remains of a Zeuglodont Cetacean. It looks, therefore, very much as if the Eocene ancestors of the modern Cetacea had a skin studded with bony plates, as have the armadillos. This being the case, the disappearance of hair is not surprising. The room would be taken up by the calcified plates, and when the latter disappeared, as they have in the vast majority of existing Whales, the naked skin alone would be left.

Whales possess no externally-visible hind-limbs; rudiments of these appendages are present, which will be dealt with under the description of the principal features of the skeleton. But it has been discovered that in the Porpoise, external vestiges of hind-limbs do appear in the foetus, a fact which, be it observed, does away with the old view that the flukes of the Whale are the last term in the series of vanishing hind-limbs, of which the Seals, with their hind-limbs and tail bound up together, offer an intermediate step.

The tail is fish-like in form, but the flukes are horizontal instead of vertical as in fishes and Ichthyosaurus. This arrangement is no doubt associated with the need for rapid return to the surface waters after a prolonged immersion in search of food. A downward stroke, such as is given by the powerful and large tail flukes, would naturally bring about this result rapidly. The tail, moreover, is under all circumstances the swimming organ. Its motion has been stated to be slightly rotatory, like that of a screw, and it is the case that the two flukes are often alternate in shape like the flanges of a screw; one being convex upwards, the other convex downwards.

The fore-limbs are in the form of paddles, but they do not apparently serve as organs of locomotion so much as balancers. When a Whale is killed, it falls over on to one side, the office of the flippers being to maintain the proper position. It is believed, however, from the fact that the embryo often shows a relatively larger pectoral fin than that of the adult—the difference being due to a reduction in the adult of the number of phalanges—that the fin was once an organ of progression.

The pectoral fin of Whales exists in two forms. In the Toothed Whales it is shorter and rounder; in the Whalebone Whales longer and narrower. Structural differences accompany these outward dissimilarities. In the first-named group the humerus and the beginning of the radius and ulna are within the body, and do not form a part of the fin. In the Whalebone Whales, on the other hand, the fin contains all the bones of the fore-limb. Another remarkable contrast between the hand in the two groups of Whales is that while the Toothed Whales have five fingers, thus justifying the prevailing opinion that they are the more primitive of the two groups, the Whalebone Whales have only four fingers. Actually the Right Whale, Balaena, seems to have five fingers; and, indeed, the fact that it has, is often used to distinguish it from the Humpback, which has undoubtedly only four. But a careful consideration of the state of affairs which prevails in the foetus of Balaenoptera dispels this idea. Between what are apparently the second and third fingers, a rudimentary finger, consisting of four phalanges, appears. This is not produced, as is an additional finger found in the White Whale or Beluga, by a splitting of a finger. Accordingly the four-fingered condition of the Whalebone Whales is produced by the dropping out of a finger in the middle of the series,—a very remarkable fact. When fingers disappear, as, for instance, in the Horse, etc., it is at the two ends of the series that the digits vanish. If this view of Professor Kükenthal's be accepted, it follows that the presumed thumb of the Right Whale is what has been termed the prepollex.

The hand of the Whales, like those of some other aquatic creatures, e.g. the reptile Ichthyosaurus, has a larger number of phalanges than have terrestrial animals. The result of this is, of course, to increase the length of the fin and its utility as a paddle. It is commonly not all the fingers that have developed this great number of accessory phalanges. Rudimentary nails have been found upon the Cetacean hand; but in no case are they functionally developed. In the Manatees we have the disappearance of the nails still imperfectly accomplished. In M. latirostris there are nails; these have vanished, apart from possible traces to be seen with a microscope, in M. inunguis.

A very characteristic feature of certain Whales are the furrows to be seen on the throat. This is especially the case with the Rorquals, in which group the Humpback Whale, Megaptera, is to be included. The whales of these two genera (Balaenoptera and Megaptera) have a large number of the throat furrows—as many as sixty have been counted. Some other Whales have a smaller number; thus Rhachianectes has but two on each side, and the Physeteridae have not many more. These furrows are absent in very young embryos. It is thought by Professor Kükenthal that they allow of a wide opening of the mouth.

Fig. 181.—Dorsal surface of bones of right anterior limb of Round-headed Dolphin (Globicephalus melas). × 1⁄10. The shaded portions of the digits are cartilaginous. c, Cuneiform; H, humerus; l, lunar; R, radius; s, scaphoid; td, trapezoid or magnum; U, ulna; u, unciform; II-V, digits. (From Flower's Osteology.)

The blow-hole of Whales is, of course, the aperture of the nostrils, which are not so far back in the foetus as in the adult. By the characters of the nostrils the Toothed Whales can be distinguished from the Baleen Whales; in the latter the orifice is double, in the former single. In embryos of Dolphins, however, the two apertures are quite independent. The phenomena of spouting have often been misinterpreted. When the Whale breathes, the expired air rushes out through the nostrils. The water vapour in the breath condenses into drops of water in the cold Arctic regions where the phenomenon has been mainly observed. Hence the idea that water taken in at the mouth is expelled through the blow-hole. As the Whale approaches the surface to breathe, it may be that some of the water of the sea is driven upwards by the forcible expulsion of air from the lungs. But for the most part the water which is spouted is simply condensed breath.

Like some, but not all, other aquatic Mammalia the Whales have apparently no external ear. Indeed the opening of the ear is excessively small. In a huge Rorqual it will "admit a quill"; and although "a quill" is rather vague, we may fairly allow any sized quill without proving that the orifice of the auditory passage is anything but exceedingly minute. As a proof, added to so many, that the Whales are the progeny of terrestrial creatures, we have the occasional traces of external ears.

Fig. 182.—Left lower jaw of foetus of Balaenoptera rostrata. Inner aspect, natural size, showing teeth. (After Julin.)

Whalebone Whales never possess permanent teeth as well as the baleen; but in the foetus are more than traces of true teeth, which, however, never arrive at maturity. The whalebone itself is described later (p. ). That the Whalebone Whales possess teeth while in the foetal condition was discovered so long ago as 1807. It has since been confirmed by many observers. Not only is there one set of teeth developed in the foetal Balaenoptera but two, of which one comes to a greater maturity; the other, in fact, remaining at a very early stage of development. The more complete dentition belongs to the milk series, as is the case with the Toothed Whales. A very interesting conclusion with regard to the derivation of the simple conical teeth of Whales seems to follow from the development of these structures in Balaenoptera. There are in the young foetus fewer teeth than in the more advanced embryo. Now in the younger embryo some of the teeth are furnished with more than one cusp; they are bi- or even tri-conodont. As Sir R. Owen observed, the teeth—some of them—are literally double teeth. This is a suggestion of the more complicated teeth of the Zeuglodonts, and shows so far that the simple conical teeth of existing Whales (cf. however the Platanistidae) are not by any manner of means so primitive as their actual structure would undoubtedly lead one to believe. Further than this, the greater number of teeth in the older embryo coincided with the disappearance of these double teeth, which seem to split up into the simple conical teeth.

The Toothed Whales are not furnished with baleen, but with teeth only. These teeth are more or less numerous, their arrangement being of value in the classification of the group; a matter which is dealt with later.

In the Narwhal, whose dentition in the adult is reduced to the well-known tusk or tusks (properly developed only in the male), there is a complete foetal dentition. A very curious fact has been elucidated by Professor Kükenthal about the dentition of the Common Porpoise. It appears that in this Cetacean the two teeth corresponding to each other of the two dentitions may fuse into a single tooth, which has in consequence a double crown. It may be that this is the case with the Platanistid Inia, and that its diconodont teeth are not, therefore, a reminiscence of the comparatively complicated teeth of the ancient Zeuglodonts.

The internal organs of Whales which show the greatest peculiarities as compared with other mammals are the stomach, the lungs, and the diaphragm. Whales always possess a complicated stomach divided into many, but into a variable number of, chambers: there are as few as four in some, as many as fourteen in Ziphioids.

On account of its complication the stomach has been compared to that of Ruminants—it has even been alleged that Whales "ruminate"—but the comparison will not hold good. Nor, on the other hand, is there a very close resemblance to the equally-complicated stomach of the Sirenians.

The Rorqual has a stomach with as few compartments as any. The only Whale which appears to have fewer is Balaena mysticetus, where there are but three. In the Rorqual the oesophagus opens into a more or less globular sac; from the upper end of this, i.e. close to the entry of the oesophagus, arises the second chamber, long and narrowish; then follows an extremely short third sac, then a larger fourth, after which comes the dilated commencement of the small intestine. The latter might be regarded as a chamber of the stomach were it not for the fact that the ducts of the liver and the pancreas open into it. This represents one type of the Cetacean stomach, which seems to be found in all Whales except the Ziphioids. In the latter, the oesophagus opens into the first compartment as usual; but the second division of the stomach arises not close to the entrance of the oesophagus, but at the opposite end. It would seem, therefore, as if the first division of the stomach, found in most Whales, were missing in Ziphioids. This way of looking at the matter is confirmed by the fact that in Hyperoodon a remnant of the missing first stomach is found in the shape of a small diverticulum of the oesophagus just before it enters the stomach.

The essential difference between the Whale's and the Ruminant's stomach is this: in the latter the stomach is primarily divided into two portions, of which the first is non-digestive and is clothed with oesophageal epithelium. The second, the abomasum, is the digestive region. The first part is again divided into three compartments. In the Whales, on the other hand, it is the digestive part which is again subdivided, while if the first part is divided it is not markedly so as in the Ruminants.

The lungs are remarkable for their unlobulated character; in this they agree with the lungs of the Sirenia. The thoracic cavity in which they lie is barrel-shaped, and not, as is usual in terrestrial mammals, boat-shaped, i.e. narrower sternally than above. The alteration of the shape of the thoracic cavity is associated with the aquatic life; so at any rate the fact that it is also marked in Seals and even in the Otter seems to show. The Whales are also characterised by the great obliquity of the diaphragm, which is extremely muscular. In this character again we find an agreement with the Sirenia, and also with other aquatic mammals; it is not therefore a character of Whales so much as evidence of an adaptation to the aquatic life. The advantage is, it appears, in the increased capacity of the thoracic cavity, and the consequent greater possibilities of expansion of the lungs, which it must be remembered serve as hydrostatic as well as breathing organs.

Some of the internal arteries of Whales break up into retia mirabilia. Their kidneys are lobulated; whether this has anything to do with the aquatic life is not so clear. It also characterises the Sirenia, more or less, and the Otters; but, on the other hand, the terrestrial Bears show the same structure as do also some Ungulates. It must be borne in mind, too, that the kidneys of foetal Man are lobulated.

The liver is a compact organ not showing such lobulation as is common, but not universal, among mammals.

The bones of Whales have a somewhat loose structure, and are much impregnated with oil. In many features the skeleton of Whales is highly distinctive of the order.

Fig. 183.—Skeleton of Porpoise (Phocoena communis), acr, Acromion process of scapula; cerv, united cervical vertebrae; chev, chevron bones; cor, coracoid process; hu, humerus; hy, hyoid; jug, jugal; lumb.trans, lumbar transverse processes; max, maxilla; nas, nasal; orb, orbit; par, parietal; pelv, vestige of pelvis; per.ot, periotic; pr.max, premaxilla; rad, radius; rb, first rib; rb, twelfth rib; sc, scapula; s.occ, supra-occipital; st, sternum; uln, ulna; zyg, prezygapophysis. (From Parker and Haswell's Zoology.)

Fig. 184.—Under surface of the cranium of a young Caa'ing Whale (Globicephalus melas). × 1⁄5. AS, Alisphenoid; BO, basioccipital; cf, condylar foramen; ExO, exoccipital; Fr, supra-orbital process of frontal; gf, glenoid fossa of squamosal; Ma, body of malar; Mx, maxilla; OS, orbitosphenoid; Per, posterior (mastoid) process of periotic; Pl, palatine; PMx, premaxilla; Pt, pterygoid; Sq, squamosal; tg, deep groove on squamosal for meatus auditorius externus, leading to tympanic cavity; Ty, tympanic; Vo, vomer; ZM, zygomatic process of malar. (From Flower's Osteology.)

The brain case is small proportionately and rounded. The "face" is therefore long, and in some cases, especially among the fossil forms of Platanistidae, the rostrum is extraordinarily elongated. The asymmetry of the Whale's skull is one of its most remarkable features; this, however, is entirely limited to the Toothed Whales, and among them is more pronounced in some forms than in others. Thus the Platanistidae and many Ziphioids are not nearly so asymmetrical as the Dolphins and, especially, Physeter. This asymmetry affects particularly the premaxillae, the maxillae, and the nasals. The base of the skull is symmetrical. The Whale's skull has very long premaxillae which, however, do not, except in the extinct Zeuglodonts, bear any teeth. The nasal bones, whether symmetrical or the reverse, are very small in existing Whales, which arrangement, together with the long and broad maxillary bones, removes the anterior nostrils, the blow-hole, far backwards. The roof of the skull is not at all formed by the parietals externally. These bones form a portion of the side of the cranium, but are replaced or covered by the enormously-developed supra-occipital in the adult. Here again the Zeuglodonts are more typically Mammalian, for in them the parietals have a normal development and situation, rising even into a median crest as in so many quadrupeds. The bones related to the organ of hearing, the tympanis and petrous bones, are very solid and dense in structure. Moreover they are but loosely attached to surrounding bones, and are thus easily and frequently lost. Nearly the only mammals which resemble the Whales in the fact that the pterygoids sometimes meet in the middle line below are the Edentata (Anteater and Armadillo, see p. ). But in both groups this peculiarity is not universal.

Fig. 185.—A section of a skull of a young Caa'ing Whale (Globicephalus melas). × 1⁄5. a, Angle; an, anterior nares; AS, alisphenoid; bh, basihyal; BO, basioccipital; BS, basispnenoid; cd, condyle; cp, coronoid process; ExO, exoccipital; Fr, frontal; id, inferior dental canal; IP, interparietal; ME, ossified portion of the mesethmoid; Mx, maxilla; Na, nasal; Pa, parietal; Per, periotic; Pl, palatine; PMx, premaxilla; pn, posterior nares; PS, presphenoid; Pt, pterygoid; s, symphysis of mandible; sh, stylohyal; SO, supra-occipital; Sq, squamosal; th, thyrohyal; Vo, vomer. (From Flower's Osteology.)

The vertebral column is remarkable for the fact that more or fewer of the cervical vertebrae may be fused together into a short and compact mass. This is seen at its maximum in the genera Balaena and Neobalaena. The odontoid process of the second vertebra, though hardly at all marked, is nevertheless really present and developed from a bony centre of its own, as in other mammals. The dorsal and lumbar vertebrae are, of course, to be distinguished by the presence of ribs attached to the former; but as there is only a rudimentary pelvis, not attached to the vertebral column, no sacral region can be detected. The caudal vertebrae are to be recognised by the V-shaped chevron bones below.

Fig. 186.—Section through middle line of united cervical vertebrae of Greenland Right Whale (Balaena mysticetus). × 1⁄9. a, Articular surface for occipital condyle; e, epiphysis on posterior end of body of seventh cervical vertebra; sn, foramen in arch of atlas for first spinal nerve; 1, arch of atlas; 2, 3, 4, 5, 6, conjoined arches of the axis and four following vertebrae; 7, arch of seventh vertebra. (From Flower's Osteology.)

Fig. 187.—A, Sternum of Greenland Right Whale (Balaena mysticetus). × 1⁄15. B, Sternum of Common Rorqual or Fin Whale (Balaenoptera musculus). × 1⁄10. (From Flower's Osteology.)

The sternum in the Whale tribe is much more modified in the Whalebone Whales than in the Odontocetes. In the latter it is made up of several pieces, as in other mammals, which often, however, become coalesced. In the Mystacoceti this bone is a single piece, to which only one pair of ribs is attached, and its form is characteristic of the genus. It is heart-shaped, more or less, in Balaena, and somewhat cross- or T-shaped in the genus Balaenoptera. In the Odontocetes the ribs have, some of them, the normal attachment by capitulum and tuberculum. In the Mystacocetes the attachment, where it exists, is very loose, and the tuberculum alone is attached to its vertebra. This allows of the freer play of the ribs during respiration. The scapula has a very characteristic form in these animals. The acromion, where it exists, is placed near the anterior margin of the shoulder blade, and overlaps the generally long coracoid process. Clavicles are totally absent. The pelvis is very rudimentary, consisting merely of a single bonelet, to which are attached the rudiments (in some cases) of a femur, and, in Balaena (Fig. 188), of a tibia also.

Fig. 188.—Side view of bones of posterior extremity of Greenland Right Whale (Balaena mysticetus). × ⅛. i, Ischium; f, femur; t, accessory ossicle representing the tibia. (After Eschricht and Reinhardt) (from Flower's Osteology.)

Whales are to be divided into three great groups:—(1) the Whalebone Whales or Mystacoceti; (2) the Toothed Whales or Odontoceti; and (3) the entirely-extinct Archaeoceti or Zeuglodonts.