On the development and structure of the whale

Part I. On The Development Of The Dolphin

GUSTAV GULDBERG, m d. AND FRIDTJOF NANSEN, ph. d

BERGEN

PRINTED BY JOHN GRIEG

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Contents

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Preface . V

Historical Survey of the knowledge regarding the development of the whale . 1

Preface

It is now about four years since my friend, Dr. Fridtjof Nansen asked me if I would join him in a work on the line collection of Whale embryos contained in the Bergen Museum, and which, to a great extent, Dr. Nansen himself, when curator of that museum, assisted by the curator, James Grieg, had brought together. As the committee of management were favorably disposed towards Dr. Nansen's proposal, I was quite willing to enter into so interesting a labour, the more so from my having formerly been greatly interested in Cetacea.

In the autumn of 1890, some small embryos of Lagenorhyn cbus acutus were sent to us in Christiania, where Dr. Nansen was also residing, having been made fellow of the University after his Greenland expedition, and keeper of the zootomical collection in the Royal Norwegian University of Christiania.

In the summer of 1891, I spent some time in Bergen in order to become better acquainted with the large collection of embryos and foetuses belonging to the museum there; and I would here offer my sincerest thanks to Dr. Danielssen1) and the officers of the museum, Dr. Brunchorst, Dr. Appellof and James A. Grieg, for the friendliness and good will there shown me.

During the winter of 1891 — 92, Dr. Nansen and I worked together in the laboratory of the Anatomical Institution, studying more particularly the small embryos of Lagenorhyn chus acut us, Gray. Later on, we had only occasional meetings for consultation about the work, as Dr. Nansen, being occupied with preparations for his North Pole Expedition, left it entirely to me. Dr. Nansen, as is well-known started on his Polar Expedition in June, 1893.

Owing to the numerous duties of my office, and to other calls upon my time, the completion of this work has been delayed for about two years, as I did not resume it until August, 1893. In the mean time, the material had been increased by the addition of a few interesting small embryos of Phocsena communis which I received from Greenland, through the University’s Zoological Museum. In these I was able to show the existence of rudimentary hind-limbs. I spoke of this discovery and demonstrated it at the Anatomical Congress in Strasburg, on May 14th, 1894.

In addition to the species Lagenorhynchus acutus, L. albirostris, Phocsena communis and Orca gladiator, I have examined several embryos of Globicephalus me! as and Delphinapterus leu cas, and to some extent Delphinus del phis. With the last three species, however, I have not dealt in this treatise, as its publication would thereby have been still further delayed.

Our special thanks are due to the trustees of the Bergen Museum for the liberality with which the work has been got up, and to Miss Jessie Muir in Christiania for the care and labour bestowed upon its translation.

Christiania, July, 1 894. G. GULDBERG.

J) Since writing these lines, I have received the sad intelligence of Dr. Danielssen’s death.

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HISTORICAL SURVEY OF THE KNOWLEDGE REGARDING THE DEVELOPEMENT OF THE WHALE

Historical Survey.

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X rom olden times until the beginning of this century, not only has the natural history of the Cetaceans belonged, as is well known, to the least familiar sections of zoology, but the accounts of this remarkable group of aquatic Mammals have been perhaps more than usually mixed up with additions of the imagination. Although Oetology, like so many other branches of biology, has grown considerably during the present century, we find its essential features already indicated in the by no means unimportant literature of the last century. John Hunter’s must he considered as a fundamental work in this direction.

The older cetological literature presents many difficulties, and its value is in a great measure diminished on account of the great inaccuracy in the descriptions of the different Cetacean species, and the consequent difficulty and often impossibility in recognising them, whereby much confusion and many mistakes have arisen. This is, of course, very excusable, not to say natural, as both methods and material have been defective ; but on the other hand, we have here a striking example of how misleading and disadvantageous is the desire in certain writers to cut up the genera — I had almost said, in a wholesale manner, — into a number of species, thus manufacturing a diagnosis on the basis of defective material, without regard to differences of age and sex, or to variations, or only founded on anatomical (osteological) descriptions. It is not, therefore, until the later decades that a reliable specific definition — that first basis of all profound knowledge of any group of animals — has obtained a solid foundation in cetology; whereby also, it has become possible to identify several species which have been known and described in former centuries on the basis of numerous observations.

As was the case with so many other orders of the Mammal class, it was long before the Cetacea found recorders of the history of its developement, not through any want of interest in it, but because there was no material to be obtained. When I have named Eschricht, Anderson, Sir William Turner, and in later years Kiikenthal, I have named the most important contributors. It would certainly be interesting if a complete literary survey of the history of the developement of the Cetacea and their biology from the earliest times could be made ; for many a useful fact lies buried among the superfluous and useless matter which occupies so many pages in the old folios. I think, however, that the result, with regard to zoology, would scarcely he proportionate to the labour.

In here giving a short introductory survey of what is known about the history of the developement of the Cetacea, I ask that it may be criticised indulgently, as so much will remain incomplete and defective, the reason of this being partly that I have not had a sufficient quantity of literature at my disposal, and partly that I have not had the wished-for opportunity of working up the whole in the manner I had originally intended.

In the developmental history of the Cetacea, as in so many other fields of natural science, we find interesting

facts recorded by that wonderful genius of ancient times, Aristotle, in which the general views of the philosopher, and the

sober inquiry into details of the naturalist are combined in an exceptional manner.

In the 4th chapter of his first book, in Avhich the propagation of animals is generally treated, Aristotle states

that the Cetacea give birth to living young ones. In his second book (chap. 13 in the German translation, chap. 9 in the

French), it is stated that dolphins have mammee, and that these are not situated on the upper part of the body but by the regio inguinalis; they have not, however, as in quadrupeds protruding nipples, but a kind of opening on each side out of which the milk flows, the young one sucking as it swims by the side of its mother, as it has already sometimes been clearly observed to do. In the 6th book (chap. 11 in the French translation, chap. 12 [66 — 70] in the German) the propagation of the Cetacea is more fully treated of. We subjoin the following: ,, The Dolphin, the Pliakena (Physeter?) and other Whales which have no gills, but a spouting-tube, give birth] to living young ones, as do also the sawfish and the „bus“ (le boeuf marin) ; for in none of these animals are eggs found, but from the first, germs (embryos) whence the young emerge hy developement, as is the case with man and the viviparous quadrupeds. The Dolphin generally gives birth to 1 young one, sometimes 2 ; the Whalebone Whale to at most 2, which is most frequently the case, seldom 1 ; as with the Dolphin, so is it with the Porpoise (Phocsena) which has the appearance of a small dolphin and lives in Pontus, but is distinguishable from the Dolphin by its smaller size and greater breadth of back : its colour is dark blue. Many consider the Phocsena to be a species of Dolphin. [All animals that have a spouting-tube, breathe and draw in air, for they have lungs. The Dolphin has

also been observed when asleep to hold its snout above water, and to snore in its sleep.] The Dolphin and the Phocsena

have milk and give suck to their young; they alst take their young into themselves as long as they are small. Young Dolphins grow quickly, for they attain to their full size in 10 years: they are pregnant for 10 months. The Dolphin does not bear at any other season than the summer. At the period ot the dog-star, it hides itself for about 30 days. It accompanies its young for a long time and takes great care of them. It lives for a long time, and it is certain that many attain to the age of 25 and even 30 years; for fishermen have cut the tails off certain Dolphins, and let them go again, so that their age could be ascertained. “

Other ancient authors, who mention the Cetacea, have not been able to add anything of lasting value about these animals which was not known from Aristotle.

Galen (Kuhn’s edition, Leipz. 1821) states that Whales, Seals and Dolphins breathe air (HI. p. 444) and that Whales .... have their eyes „on their long neck“ (ibid. p. 631). He further states that the flesh is hard and unpleasant to eat, but may be eaten, if necessity requires, salted and spiced (VI, p. 728 and 737); also that eating whale’s flesh occasions ’’sanguis melancholichus11 (VIII. p. 13). *) Pliny, Oppian, Paulus iEgineta and Varro include the Cetacea in their description of other large marine animals, such as the Tunny, the Shark and others. Pliny has made himself

conspicuous by the fables with which he has adorned the natural history of the Whale. iElianus and Plutarch also

describe certain features of its natural history.

There are certain Arabian writers in the middle ages who are of interest to the history of cetology, as they mention some of the whale species among other wonders of the ocean.

In Abbe Renaudot’s translation in 1718, of an old Arabian work: ’’Anciennes Relations des Indes et de la

Chine de deux Voyageurs Mahometans qui y afferent dans le IX e. Siecle de notre Ere,“ revised and criticised by M. Renaud, a large fish is stated to be found in the Indian Ocean, of which the size and width of the jaws is described as enormous. Ships dread this fish greatly, and bells are rung to frighten it away. Much is said, too, about ambergris, which is cast up by the sea on to the coasts, also that it grows from the bottom of the sea like a plant. The word al-oual occurs as the name of a large fish. According to Renaud’s critical remarks, the large fish in question must be a Cachelot (Physeter macr ocephalus, L.).

In Edrisi’s Geography from the year 1154 (translated from Arabic to French, by P. Amedee Jaubert, Paris

1834 it is stated that in the Gulf of Oman, in Indian Herkend, there is a kind of fish that is about 200 feet long. ’’It is

called el-waly; it is white.11 The translator interprets this as ”la baleine11 (in which case it is Eubalsena australis) and thinks that the words ’whale1 and ’Waff have come from the Arabic, In Dimichqui’s Cosmography (from the latter half of the 13th century) ambergris is stated to occur in the Indian Ocean, and fables are related about its origin and effect; the stranding is also recorded of a large fish — probably a cachelot — on which 30C people lived for a whole month. A fuller description of certain whale species is found in El-Kazwini’s Cosmography. A ’’well-known genus of fish,11 100 feet in length is here mentioned, which, by the aid of harpoons, is drawn to land and cut in pieces with axes, while from ’’the

brain, a quantity of oil" is extracted. The Dolphin receives high praise as a ’’blessed animal11 that helps those who are

in danger of drowning. Two other kinds are also described, called ’’Ketos11 and ’’Kata11, i. e. Whales, of which the first destroys ships, and the second is so large that bridges may be built of its ribs. „If leprosy be spread over with its fat, it disappears with God the Almighty’s help.11 The same writer also mentions the taking of whales — al iblenah (= la baleine) — on the island of Irlandah; its young are hunted to procure ”suvl“ (fat and lean). It is related that the young are born in the month Ilul (i. e. September) and are hunted from October to January, and the mode of capture is described as very adventurous. It is evidently the well-known mode of whale fishery practised by the Biscayans. In other respects, the accounts

of these Arabian writers are highly embellished, and any fresh items of information about the Cetacea will scarcely be found

in their literature.2)

In the time of the Renaissance, we find the famous Conrad Gesner as a distinguished author also in Cetology.

For tlie quotations from Galen I have to thank my honoured colleague Prof. Stenersen.

-) The above accounts of the Cetacea from Arabian writers have been obtained through my colleague, Prof. A. Seippel, to whom I would here offer my sincere thanks.

This learned man. who was so well versed in classical literature, quotes and criticises the accounts of these animal forms given by older authors. In this way he has numerous quotations from Aristotle, Oppian. Pliny, iElianus, etc. for which reason his great work will retain its full value from a historical point of view, and as an original source. This edition also contains extracts from Be Ion (Petrus Belonius) and Rondelet (Guilielmus Rondeletius). The animals mentioned in his work, are. as is well known, arranged in alphabetical order, and most of them are also figured. Under the heading Delphinus, a gravid uterus with the embryo is figured. The Dolphin’s anatomy is mentioned, also that the females usually give birth to one, occasionally two young ones, after 10 months gestation. The birth takes place in the summer. The foetus is inclosed, in the uterus, in the chorion, the amnion, and the allantois: ’’sanguine nutritur per venas umbilicales, spiritual per arterias

from Oppian is quoted in support of this (p. 383). Under the same heading a number of stories is related about the habits, disposition and intelligence of the Dolphin.

Among older writers in Northern literature, who have treated of Whales to any great length, Peder Clausson Friis (b. 1545 at Ekersund, Norway; d. 1614) claims a special mention. In his account ”Om Fische,“ he gives an extract from the Royal Mirror (Speculum regale), in which the different species of Whales are enumerated and generally briefly described. The sperma of the Whale is moreover mentioned under "Huall-Raff”, ’’which on account of its fat and the air which it contains, floats on the top of the water like froth.1' In his account of a whale-fishery, he relates that ’’when whales are at play, several may often run after one mother-whale." He describes a large Whale — probably a fin-backed whale, Balaenoptera musculus — which was stranded in August 1532 at Thinemudt (Tynemouth?) (England). Olaus Magnus repeats the fact of the stranding of this whale. Olaus Worm (Museum Wormianum, 1656) translates into Latin the account of the different species of whales repeated by Peder Claus on Friis from the Royal Mirror1). Nothing further is related concerning the embryos of whales. These accounts of Whales and their habits have probably arisen in a great measure from a gradual collection of the experiences of the fishing population of the coast.

Thomas Bartholin describes the anatomy of the Porpoise, cinatome tursionis, which may presumably be identified with our ordinary Phocsena communis. In the presence of the Dano-Norwegian king, Frederick III and other spectators, he dissected a female Dolphin and its foetus. He says: "The internal organs do not greatly differ from those in the human being11 ; and he mentions the differences in his description. In the left part of the uterus he found a male foetus, with its head turned towards the mouth of the uterus (capite ad os uteri verso) : the right part of the uterus was empty. He describes the umbilical cord with prominent quadruple vessels, and its division and disappearance in the sides of the placenta. His remarks on the placenta, amnion and allantois are worthy of notice: "Placenta exilis concreto sanguine similis, parietibus chorii agglutinatin’. Tenuis amnios. Allantoides humores serosum continet, farciminibus inclusum.11

In his anatomical description of the embryo, Bartholin pays special attention to the foetal openings and passages, such as the communications between the v. umbilicalis and the v. hepatica. the open urachus, the small supra-renal bodies, the testes with their connections, hidden in the abdomen, and the connection of the membrum with the cartilaginous” hyoid-like11 bone; the position of the heart beneath the ensiform process is moreover mentioned, and its main fcetal communications.

The works of Major (De anatome phocaence . . . Ephemer, medico-physicarum annus tertius) and E. Tyson (The anatomy of a porpess, London, 1680, 40) 2) also treat of the anatomy of the Porpoise.

Martens describes several species of whale, especially the Greenland Whale and makes a few remarks on the Nordcaper. With regard to the breeding of the Greenland Whale (pp. 103, 104) he only mentions that it gives birth to 1 or 2 young ones, and the author knows nothing about the duration of pregnancy. His attempt to make spermaceti from the semen of the whale (which at that time was a general supposition), strikes us as rather comical. Klein gives an account of the anatomy of the Phocsena, with special mention of certain muscles, and makes a systematic survey of the different Cetacea. He took the holes on the embryo’s upper lip, left by the hairs, to be organs of smell.

Johan Anderson repeats from the ’’unanimous assertions of Greenland whalers, “ stories of the pairing of Greenland whales in a perpendicular position in the water, both ’’sinking on to their broad, flat-lying tail-fins,11 with their heads pointing upwards, and embracing each other with their pectorals (swimmers). But at the same time he quotes P. Dudley’s account (Philos. Trans, no. 387, Art. 2), which states that the female lies upon her back and bends her tail backwards, while the male passes over her. They breed every other year, and the gravidity is said to last 9 or 10 months. A foetus no more than 17 inches in length, says Anderson, has the form of the species, and is white. They are 20 feet long when they are born. The young one sucks the maternal breast for 1 year. There are illustrations of the Narwhal, and among them of a foetus, which, however, is unrecognisable.

Briss on’s ’’Regnum animale11 contains only a system, in which the Cetacea form one particular class separate from Fishes; while Linnaeus, in his ”Sy sterna Naturae1-', allows the Cetacea to belong definitively to Mammals.

1) My attention was directed to the above literary particulars by Prof. Gustav Storm, to whom I tender my sincere thanks.

£) These two authors I know only from extracts.

Erik Pontoppidan, bishop of Bergen, among other things mentions (20 and 21 Bd. II. p. 228) the breeding of whales, repeating the statement of the perpendicular position of the animals in the water while pairing; that the new-born animal is from 8 to 10 feet long; that it sucks the breast of which the nipples are near the anus; and that the mother shelters the young one, when it is tired, beneath her great pectoral.

Gunnerus, bishop of Trondhjem, gives a long account of the Phocaena (22. II. 1765, pp. 258 — 269) and the Orca (22. IV, 1768, pp. 99 — 112) hut says nothing about their embryos or breeding. The contributions of these learned divines towards the natural history of the Cetacea is of considerable interest.

We find several observations in the work by Otho Fabriciu s, which is of such importance to Arctic fauna. He quotes in his ’’Fauna gronlandica“ remarks on the breeding and the young of certain Whales. Thus, he says of the pairing of B a Le n a mysticetus: ’’C’orpore erecto, capite supra aquam prominente.44 It brings forth its young, generally only 1, in April. ’’The Balaena hoops (i. e. Megaptera boops) gives birth to only one young one, but certainly not every year.41 The young one follows the mother until the next one is born. The Phocaena gives birth to one young one, which keeps with the mother the first year. "Si gravida occiditur, ex utero cauda pulli prominens conspici solet, forsitan matre in angustiis parere conante.41 Of the Delphinus albicans (i. e. Delphinapterus leucas, Pall.) it is related that it gives birth to only one young one which even in the fcetal state becomes light-coloured, and when newly born is of a bluish colour, afterwards becoming gradually whiter, observations which have only lately been confirmed.

J. G. Schneider’s critical collection, from his own and older times, of accounts of the natural history of the whale, is also important to the advance of Cetology in the last century. He relates the usual story about the pairing of the Greenland whale, adding also Dudley’s version, which has an appearance of greater probability. Pairing takes place only every other year, and only one young one is born. A foetus is mentioned that was 17 inches long, and of a white colour; the new-born animal is black and generally 20 ft. long. Pregnancy lasts from 9 to 10 months, and when the mother gives suck to her young, she lies on her side in the water. He gives full accounts of the other whalebone Whales that were

known in the last century, critically sifting their literature. He also gives a survey of the then known toothed whales, among

which the Cachelot takes a conspicuous place. Minor anatomical remarks are added; and he gives a representation of the common Porpoise, based upon personal observation. Schneider’s work gives valuable information for the study of the cetological literature of the 18th century.

While John Hunter, in his fundamental work on the organisation and habits of whales, enlarged the knowledge of these ocean-mammals, and brought a stronger light to bear upon them, it did not fall to his lot to make observations on

their embryos and developement. On the other hand, he treats fully of the anatomy of the organs of generation.

In the beginning of the present century we find, as an eminent cetologist, Lacepede, who describes and figures, in his well-known ’'Histoire Naturelle des Cetaces44, a number of Cetacean forms; we are, as is well known, indebted to him for the separation of the genera Balaena and Balsenoptera. He does not, however, state anything new about the breeding and developement of whales, though he describes in vivid colours and with the pen of a Buffon, the habits of these animals.

Sonnini, a contemporary of Lacepede but not of the same importance, who continued Leclerc de [Buffon’s great ’’Histoire Naturelle'4, describes the Cetacea, the different species known at that time, their different anatomical relations, especially of B alien a mysticetus, its habits, propagation, etc. He gives attractive descriptions in fine language from the

best sources of that period, though he also produces much that is false. A much greater interest in Cetology seems, however,

from this time to have been awakened in the prolific French zoologists, and has borne good fruit in the cetological literature now to be mentioned.

The accounts of the Caaing-whale, and its capture in particular, given by J. Landt (24) and H. Chr. Lyngbye (26, pp. 204 — 232) are of subordinate interest. Lyngbye mentions that young ones are often found in the Caaing-whale when cut up; but they differ very considerably in size, large embryos having been found at all times of the year, from which he infers that the Caaing-whale breeds at any season of the year. It gives birth to only one young one.

A cetological writer of some mark, William Scoresby, describes, both in his ’’Account of the Arctic Regions44 and his ’’.Journal of a Voyage to the Northern Whale-fishery,44 etc., the appearance and habits of several Arctic Whales, and among others gives a full description of the Greenland Whale, with which, from abundant experience, he was best

acquainted. ’’The pairing,44 he says ’’has often been observed in the latter end of the summer, and the young one is supposed

to be born in February or March.44 It is added that at the end of April 1811 a young one with umbilical cord was captured. "The young one at the time of parturition is said to be at least 10, if not 14 feet in length. It goes under the protection of its mother for probably a year or more, or until by the evolution of the whalebone, it is enabled to procure its own nourishment.'4 He also gives an excellent drawing and description of the Narwhal, and describes several Balsenopteridse, Globicephalus and Beluga. Scoresby’s work lias still a great interest for Cetology, notwithstanding that he was the originator of the well-known mistake about the Nordcaper, a mistake which was long maintained under G. Cuvier’s authority.

Lesson, whose name is renowned in Cetology, describes in his ’’Histoire Naturelle des Cetachs44 both the herbivorous

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Cetacea and the whales proper, their habits and anatomy, as far as they were then known, being shortly treated of. Concerning their breeding he says (p. 177): ’‘Les femelles des dauphins re^oivent dans l’accouplement les males en les serrant entre leurs nageoires. On dit cpie la gestation est de dix mois, et que la conception a lieu dans l’automne; on assure aussi qu’elles ne font qu’un petit ou deux a chaque portee, et que la mere surveille avec sollicitude leurs mouvements, les fagonne ou les habitue a la natation . . . . Ce n’est qu’en se penchant sur le cote que les jeunes dauphins saisissent le ] lamelon du sein de leur nourrice, et qu'ils y puisent un lait onctueux, de couleurs bleuatres, mais tres-nourrissant.“

In the well-known ’’Medicinische Zoologie“ by Brandt and Ratzeburg, where Physeter, Baltena and several other Cetacean genera are described with special reference to their outward form and skeleton, and to the products they yield which are used in medicine, nothing new is found about the gravidity or the embryo of these animals.

Georges Cuvier has also rendered great services to Cetology, notwithstanding the mistakes he has made on

certain points, e. g. the geographical distribution of the Greenland whale. With his critical ability and thoroughness he brings order into the numerous descriptions and the confusion of names, searching for literary sources, and procuring osteological signs for the identification of the species. His works are therefore of considerable value with regard both to the systematism and anatomy of these animals; but it did not fall to his lot to contribute any fresh facts about their breeding and developement.

Nor does F. Cuvier, in his well-known "Histoire Naturelle des Cetaces,“ seem able to extend the knowledge in

these particulars, for he says (p. XXIV seq.): ”On ignore d’accouplement des Cetaces souffleurs. Personne jusqu’a present

n’en a ete temoin. L’opinion la plus probable, c’est qu’ils s’unissent couches tous deux sur le cote .... On ignore aussi la duree de la gestation que quelques-uns portent a dix mois pour la baleine, dont le jeune, en naissant, aurait vingt pieds de longueur. “ He then enters into the subject of suckling. After describing the different Cetacean forms, — certain of the descriptions being undoubtedly based partly on autopsy, — he gives in conclusion a good, though scarcely complete list of the literature on this subject.

In the first half of the present century, we find a number of cetological treatises, some of them descriptions of stranded specimens, others, anatomical pamphlets. Inquiry seems to be trying to make up for the previous neglect of these giant organisms of the world. While there have appeared as we have seen, large collections of accounts of the true and the herbivorous Cetacea in French zoological literature, we find on the English side a number of special descriptions following John Hunter. Contributions to Cetology have also been made by German naturalists, the Phocsena communis having been taken as the model, and in the main furnishing material for special descriptions.

We find an eminent explorer in the field of Cetology in Rapp, who in his well-known little book, ’’Die Cetaceen zoologisch-anatomisch dargestellt“ 1837, has recorded the outlines of the anatomy of these animals, based upon careful observation and thorough study, while besides the addition of fresh information, he critically sifts the already existing material. He had not, however, an opportunity of becoming acquainted with Cetacean embryonic forms. He states that he found in the umbilical cord of a Phocsena 2 arteries and 2 veins,- and that the urachus seems to be open until the birth of the young one. ’'Ueber die Fbtushiillen der Cetaceen feblt es an Beobachtungen,“ he says, quoting from E. Home the statement that the chorion takes the place of the placenta, as in the sow and the mare.

Nor has Karl Ernst von Baer, in his famous work "Ueber Entwicklungsgeschichte der Thiere,“ much to say about the developement of the Cetacean type, for he states that he knows nothing about the ovum of the Cetacea, but that he believes, from the remarks made about it in literature, that no placenta is formed, and that the ovum shows conditions similar to those in the Pachydermata. He has only had the opportunity of seeing one embryo 9 inches in length, — a narwhal, whose navel string had only a single umbilical vein, a fact which he emphasized in contradiction to Rudolph’s observation of 2 umbilical veins in Phocsena.