Sir William Flower

Next at any rate to the study of the various races of the human species (which he took up seriously later on in his career), the group of mammals to which Flower devoted special attention, and which attracted his greatest interest, was undoubtedly that of the Cetacea, or whales, dolphins, porpoises, etc. At the time when he set himself seriously to study these aquatic and fish-like mammals, the zoology of the group was certainly in a most confused and unsatisfactory state; partly, no doubt, owing to the comparative rarity of complete specimens in our museums, and the consequent difficulty of instituting accurate comparisons, and partly to the reckless prodigality with which names had been given to imperfect or insufficiently characterised specimens by some of his predecessors and early contemporaries, and the needless multiplication of generic terms. It was consequently at this time almost impossible to be sure which was the right name for even many of the commoner species; while in the case of the rarer kinds, the confusion was almost hopeless. When Flower left the subject—which he only did when his working days were over—it was in great measure thoroughly in order, although of course much was left for future workers to fill in. Unhappily, his views on the nomenclature of the group have not been accepted by all his followers; so that a fresh and totally unnecessary source of confusion has been introduced of late years into a subject which had already sufficient difficulties of its own.

In regard to the discrimination of species, Flower took a view almost the reverse of that held by some of his predecessors and colleagues; and, as he says himself, he may have consequently erred in a direction the very opposite of theirs. “As species have not generally been recognised as such,” he wrote in the British Museum List of 1885, “unless presenting constant distinguishing characters capable of definition, it is probable that, in the imperfect state of knowledge of many forms, some may have been grouped together which a fuller acquaintance with all parts of their structure, external and internal, will show to be distinct.”

Apart from his explaining to popular audiences that whales were mammals and not fishes, Flower emphasised three points very strongly in regard to the organisation and physiology of these animals. First of all, he pointed out that, as a rule, they do not “spout” water from their “blowholes.” “The ‘spouting,’ or more properly the ‘blowing’ of the whale,” he wrote, “is nothing more than the ordinary act of expiration, which, taking place at larger intervals than in land animals, is performed with a greater amount of emphasis. The moment the animal rises to the surface it forcibly expels from its lungs the air taken in at the last inspiration, which is of course highly charged with watery vapour in consequence of the natural respiratory changes. This, rapidly condensing in the cold atmosphere in which the phenomena is generally observed, forms a column of steam or spray, which has been erroneously taken for water.”

Secondly, he drew attention to the importance of the rudiments of hind-limbs which occur in many whales as affording decisive evidence of the descent of the group from land mammals. And thirdly, he emphasised the marked distinction between baleen, or whalebone, whales (Mystacoceti), and toothed whales and dolphins (Odontoceti); although he appears never to have gone so far in this direction as some modern naturalists, who are of opinion that these two groups have originated independently of one another from separate types of land mammals.

Another point to which Flower devoted a considerable share of attention was the dimensions attained by the larger species of whales. Previously, there is no doubt that very great exaggeration had been current in this respect, and that such things as 150-feet whales are unknown. With his excessive caution, and determination to be on the safe side, it is however probable that in some instances—notably the Greenland right-whale and the sperm-whale—Flower somewhat under-estimated the maximum dimensions.

At what date Flower first began to study whales seriously, it is not easy to ascertain. From the fact of his contributing three papers on this subject to the Zoological Society’s Proceedings for 1864, it may, however, be inferred that by that time he had devoted no inconsiderable amount of attention to the group. In the first of those he described a specimen of a lesser fin-whale, then recently stranded on the Norfolk coast; while in a second, and much more important communication, he gave notes on the skeletons of whales preserved in the museums of Holland and Belgium which he had recently visited. Two of these he described as indicating apparently new species; although their right to distinction was not maintained. In the same year he described two skulls of grampuses from Tasmania, which were regarded as representing a new species, under the name of Orca meridionalis; a further note on these being added in the Society’s Proceedings for 1865, when the species was transferred to the genus Pseudorca. Later still it was found that the supposed species was inseparable from the typical P. crassidens; named by Owen many years previously on the evidence of a skeleton from the Lincolnshire Fens. In another note published the same year in the same journal he showed that one of the whales named by him in 1864 was identical with the one now known as Balænoptera sibbaldi; while a second paper described a specimen of the fin-whale commonly known as B. musculus. A further note on the synonymy of B. sibbaldi appeared in the Proceedings for 1868.

Reverting to earlier publications, in 1866 the Royal Society of London issued a volume containing translations by Flower of certain very important memoirs on Cetacea by Professors Eschricht, Reinhardt, and Lilljeborg. As these were written in a language understood by comparatively few Englishmen, the translation was a distinct benefit to “cetology” in this country.

Between the years 1869 and 1878 inclusive, six very important memoirs on whales (including in that term porpoises, dolphins, etc.) from Flower’s pen appeared in the Transactions of the Zoological Society of London. The first of these, which was published in the year first mentioned, was devoted to the description of the skeleton of the very interesting and then little-known South American freshwater or estuarine dolphins, Inia and Pontoporia. In the course of this memoir it was demonstrated that, in spite of the wide distance between their habitats, these dolphins and the freshwater dolphin of the Ganges and certain other Indian rivers, Platanista gangetica, collectively form a distinct family group—the Platanistidæ, which exhibits many very generalised features.

In the second memoir of this series, which appeared in 1869, Flower treated in an exhaustive manner of the osteology of the sperm-whale, or cachalot. “The fine skeleton of a young male which he procured for the Museum of the Royal College of Surgeons,” writes Professor M’Intosh in his obituary notice of Sir William, “formed the basis of this important paper, and enabled him to add to and correct much which had been written on this subject. The description of its huge cranium as a large, pointed slipper, with a high heel-piece and the front trodden down, the hollow limited behind by the occipital crest, continued laterally into the elevated ridges of the broadly expanded maxillæ, which rose from the median line to the edge of the skull, instead of falling away, as in most Cetaceans, must be familiar to all students of the group. In this vast cavity lies the ‘head-matter,’ composed of almost pure spermaceti.”

It was further demonstrated that the available evidence pointed to the existence of only a single species of true cachalot; the small adult jaws not unfrequently seen in collections being apparently those of females, which are known to be far inferior in size to the old bulls.

It may be added, in connection with sperm-whales, that the abrupt termination of the muzzle, shown (in a somewhat modified degree) in the model of the old bull, set up under Sir William’s direction in the Whale Room at the Natural History Museum, has been said by certain modern naturalists to be incorrect. Inquiries instituted at the present writer’s suggestion at the New Bedford Cachalot-whaling Station have, however, proved that the abruptness is under-estimated rather than exaggerated in the restoration.

This brief reference to the Whale Room at the museum, and Flower’s work in superintending the construction of models of several of the larger members of the group, must, it may be further added, suffice in this place, seeing that fuller mention of the subject has been already made in an earlier chapter.

The third memoir of the series in the Zoological Society’s Transactions treats of the Chinese white dolphin (Delphinus, or Prodelphinus, sinensis), and was published in 1872. In the following year appeared one on Risso’s dolphin, Grampus griseus, in which the author directed attention to certain variable markings always seen on the skin of this species. These, it has been subsequently shown, are produced by the claws in the suckers of the cuttlefish which forms the food of this species.

The two remaining memoirs in the Transactions, which appeared respectively in 1873 and 1878, were devoted to that difficult, and at the time imperfectly known group, termed ziphioid, or beaked whales. In the first of the two attention was concentrated on the aberrant and rare form known as Berardius arnuxi; while the second was exclusively devoted to the much more abundant types included under the generic title Mesoplodon, in allusion to the single pair of lower teeth near the middle of the sides of the lower jaw, which forms the single dental armature of the cetaceans of this genus. The beaked whales, it should be added, had been previously discussed by Flower in a preliminary paper published in the Zoological Society’s Proceedings for 1871 and 1876, and likewise in an article communicated in 1872 to Nature.

Special interest attaches to a paper by Flower published in the Transactions of the Royal Geological Society of Cornwall for 1872, and also in the Annals and Magazine of Natural History for the same year, on the bones of a whale dug up at Petuan, in Cornwall, sometime previously to 1829, and now preserved in the museum of the above-named Society. The whale represented by these remains was made the type of the new genus and species Eschrichtius robustus, by the late Dr. J. E. Gray. That it was a member of the group of whalebone-whales, and that it could not be identified with either of the genera then known, namely Balæna, Balænoptera, and Megaptera, was fully demonstrated by Flower, who also showed that it agreed with the two latter in having the neck-vertebræ free.

“The interesting question,” he added, “remains, whether this species still exists in our seas; if extinct, it must have become so at a comparatively recent period, certainly long after Cornwall was inhabited by man. The negative evidence of no specimen having been met with by naturalists in a living or recent state, is hardly conclusive as to its non-existence, as our knowledge of this group of animals is lamentably deficient. We are acquainted with many species, even of very large size, only through isolated individuals, and the discovery of others new to science is by no means an infrequent or unlooked-for occurrence at the present time.”

In the opinion of the present writer, it is quite probable that this whale may be identical with the grey whale of the Pacific, described many years subsequently by the late Professor Cope as Rhachianectes glaucus, in which event that name will have to give place to Eschrichtius robustus.

In the year 1879, and for some time after, Flower directed his attention more especially to the dolphins and porpoises, which collectively constitute the family Delphinidæ of naturalists, and he published a series of papers on this group in the Proceedings of the Zoological Society. In the volume for 1879 there appeared, for instance, one paper on the common dolphin (Delphinus delphis); a second on the bottle-nosed dolphin, now known as Tursiops tursio; and a third on the skull of the white whale, or beluga (Delphinapterus leucas). Of far greater importance was, however, the appearance in 1883 of a paper in the same serial on the generic characters of the family Delphinidæ as a whole. Special attention was directed in this communication to the value of the pterygoid bones, on the under surface of the skull, in the classification of the family; and characters were formulated which enabled the various genera to be identified, wholly or in part, by this part of the skull. Flower’s classification of the Delphinidæ has, with some slight modifications, been very generally accepted by later naturalists. Some time after the publication of this paper the present writer pointed out to the author that two of the generic names employed by him were barred by previous use in a different sense; and in a note subsequently published in the Proceedings, these were accordingly replaced.

Flower was, however, by no means forgetful of his earlier love for the cachalot and beaked whales (Physeteridæ); and in 1883 and again in 1884 he published papers in the Proceedings on their near relatives the bottle-nosed whales (not to be confounded with the bottle-nosed dolphins) of the genus Hyperöodon. In these investigations he was much indebted, as on several previous occasions, to the observations of Captain Gray, a well-known whaler. As regards the common bottle-nose (H. rostratus), Sir William succeeded in demonstrating that the great differences which had long been noticed in the skull were due to distinctions either of sex or age; the old males developing huge maxillary crests—with a broad and flattened front surface—of which there is scarcely any trace in the younger members of the same sex, or in females of all ages. In consequence of this difference in the skull, the head of the old bull bottle-nose is easily recognisable by the abrupt and prominent elevation of the forehead immediately behind the base of the beak. Flower was also able to show that bottle-noses yield true spermaceti, especially in the head; a fact which does not appear to have been previously known to zoologists, although it may have been to whalers. At the present day there is a considerable trade in bottle-nose sperm-oil and spermaceti; these being often blended with the products of the cachalot, from which they are distinguishable by their specific gravity. In his 1882 paper Flower described a water-worn bottle-nose skull from Australia, which he regarded as indicating a second species of the genus—Hyperöodon planifrons. The correctness of this determination has been demonstrated by complete skeletons of the same whale from the South American seas.

The last two papers on Cetacea by Sir William in the Proceedings of the Zoological Society refer to the occurrence of examples of Rudolphi’s rorqual (Balænoptera borealis) on the English coasts. In the one paper he described a specimen stranded on the Essex shore in 1883, and in the other an example captured in the Thames four years later.

As regards other contributions to our knowledge of the Cetacea, Sir William in 1883 delivered before the Royal Institution a lecture on “Whales, Past and Present,” which is reproduced in the Proceedings of that body for the same year. A second lecture, “On Whales and Whaling,” was delivered before the Royal Colonial Institute for 1885, and is published in the Journal of the Institute for that year. The article “Whale,” for the ninth edition of the Encyclopædia Britannica, is also the work of Flower; it is reproduced, almost as it stands, in the Study of Mammals.

The year 1885 saw the publication of the “List of the Specimens of Cetacea in the Zoological Department of the British Museum,” a small, but nevertheless valuable work, from which an extract has already been made. Even when this was written, the museum contained skulls or skeletons of nearly all the more important and well-established representatives of the order, the only notable deficiency being the large whalebone whale from the North Pacific commonly known as the grey whale, and scientifically termed Rhachianectes glaucus. It was not many years before this gap was filled by the acquisition of a complete skeleton of the species in question.

In concluding this brief notice of the work accomplished by Flower on the Cetacea, an extract may be made to illustrate his views with regard to the ancestry and origin of the group:—

“The origin of the Cetacea,” he wrote, “is at present involved in much obscurity. They present no signs of closer affinity to any of the lower classes of vertebrates than do many other members of their own class. Indeed in all that essentially distinguishes a mammal from the oviparous vertebrates, whether in the osseous, nervous, reproductive, or any other system, they are as truly mammalian as any other group. Any supposed marks of inferiority, as absence of limb-structure, of hairy covering, of lachrymal apparatus, etc., are obviously modifications (or degradations, as they may be termed) in adaptation to their special mode of life. The characters of the teeth of Zeuglodon and other extinct forms, and also of the fœtal Mystacocetes, clearly indicate that they have been derived from mammals in which the heterodont type of dentition was fully established. The steps by which a land mammal may have been modified into a purely aquatic one are indicated by the stages which still survive among the Carnivora in the Otariidæ and in the true seals. A further change in the same direction would produce an animal somewhat resembling a dolphin; and it has been thought that this may have been the route by which the Cetacean form has been developed. There are, however, great difficulties in the way of this view. Thus if the hind-limbs had ever been developed into the very efficient aquatic propelling organs they present in the seals, it is not easy to imagine how they could have become completely atrophied and their function transferred to the tail. So that, from this point of view, it is more likely that whales were derived from animals with long tails, which were used in swimming, eventually with such effect that the hind-limbs became no longer necessary. The powerful tail, with its lateral cutaneous flanges, of an American species of otter (Lutra brasiliensis) may give an idea of this member in the primitive Cetaceans. But the structure of the Cetacea is, in so many essential characters, so unlike that of the Carnivora, that the probabilities are against these orders being nearly related. Even in the skull of the Zeuglodon, which has been cited as presenting a great resemblance to that of a seal, quite as many likenesses may be traced to one of the primitive Pig-like Ungulates (except in the purely adaptive character of the form of the teeth) while the elongated larynx, complex stomach, simple liver, reproductive organs, both male and female, and fœtal membranes of the existing Cetacea, are far more like those of that group than of the Carnivora. Indeed, it appears probable that the old popular idea which affixed the name of ‘Sea-Hog’ to the porpoise, contains a larger element of truth than the speculations of many accomplished zoologists of modern times. The fact that Platanista, which, as mentioned above, appears to retain more of the primitive characteristics of the group than any other existing form, and also the distantly related Inia from South America, are both at the present day exclusively fluviatile, may point to the freshwater origin of the whole group, in which case their otherwise rather inexplicable absence from the seas of the Cretaceous period would be accounted for.

“On the other hand, it should be observed that the teeth of the Zeuglodonts approximate more to a carnivorous than to an ungulate type.”

This difficulty with regard to the teeth is indeed one which it is impossible to disregard, since it is scarcely credible that grinding teeth such as characterise herbivorous mammals of all descriptions could ever have been modified into the teeth of whales, either living or extinct. There is, moreover, the unmistakable resemblance presented by the cheek-teeth of the aforesaid extinct zeuglodons to those of Carnivora. Both these facts seem to point to the derivation of toothed whales, at any rate, from flesh-eating rather than herbivorous mammals; although they have certainly no relationship with the eared seals.

Since the foregoing passage was written it has been practically demonstrated that the toothed whales, at any rate, are the descendants of primitive Carnivora. Professor E. Fraas, of Stuttgart, and Dr. C. W. Andrews, of the British Museum, have, for instance, shown that the zeuglodons are derived from the Eocene group of Carnivora known as Creodontia; while there is every reason for regarding the zeuglodons themselves as the ancestors of modern toothed whales.