Bibliography.—Leopold Rius,Bibliografía crítica de las obras de Miguel de Cervantes Saavedra(Madrid, 1895-1905, 3 vols.);Obras completas(Madrid, 1863-1864, 12 vols.), edited by Juan Eugenio Hartzenbusch;Complete Works(Glasgow, 1901-1906, 8 vols. in progress), edited by James Fitzmaurice-Kelly;Don Quijote(Madrid, 1833-1839, 6 vols.), edited by Diego Clemencíu;Don Quixote(London, 1899-1900, 2 vols.), edited by James Fitzmaurice-Kelly and John Ormsby;Don Quijote(Madrid, 1905-1906, 2 vols. in progress), edited by Clemente Cortejón;Rinconete y Cortadillo(Sevilla, 1905), edited by Francisco Rodriguez Marín;Epístola á Mateo Vázquez(Madrid, 1905), edited by E[milio] C[otarelo]; Julián Apráiz,Estudio histórico-crítico sobre las Novelas ejemplares de Cervantes(Madrid, 1901); Francisco A. de Icaza,Las Novelas ejemplares de Cervantes(Madrid, 1901); Francisco Rodríguez Marín,El Loaysa de “El Celoso Extremeño”(Sevilla, 1901); Narciso Díaz de Escovar,Apuntes escénicos cervantinos(Madrid, 1905); Manuel José García,Estudio crítico acerca del entremés “El Vizcaino fingido”(Madrid, 1905); Alfred Morel-Fatio,L’Espagne de Don QuichotteinÉtudes sur l’Espagne(Paris, 1895, 2me série); Julio Puyol y Alonso,Estado social que refleja “El Quijote”(Madrid, 1905); James Fitzmaurice-Kelly,Cervantes in England(London, 1905); Raymond Foulché-Delbose,Étude sur “La tia fingida,”in theRevue hispanique(Paris, 1899), vol. vi. pp. 256-306; Benedetto Croce,Due illustrazioni al “Viage del Parnaso,”in theHomenaje á Menéndez y Pelayo(Madrid, 1899), vol. i. pp. 161-193; Paul Groussac,Une Énigme littéraire: le Don Quichotte d’Avellaneda(Paris, 1903); Alonso Fernández de Avellaneda,El ingenioso hidalgo Don Quixote de la Mancha(Barcelona, [1905]), edited by Marcelino Menéndez y Pelayo; Julio Cejador y Franca,La Lengua de Cervantes(Madrid, 1905, &c.); Martin Fernández de Navarrete,Vida de Miguel de Cervantes Saavedra(Madrid, 1819); Cristóbal Perez Pastor,Documentos Cervantinos hasta ahora inéditos(Madrid, 1897-1902, 2 vols.); Emilio Cotardo y Mori,Efemérides Cervantinas(Madrid, 1905); Francisco Rodríguez Marín,Cervantes estudió en Sevilla, 1564-1565(Seville, 1905).
Bibliography.—Leopold Rius,Bibliografía crítica de las obras de Miguel de Cervantes Saavedra(Madrid, 1895-1905, 3 vols.);Obras completas(Madrid, 1863-1864, 12 vols.), edited by Juan Eugenio Hartzenbusch;Complete Works(Glasgow, 1901-1906, 8 vols. in progress), edited by James Fitzmaurice-Kelly;Don Quijote(Madrid, 1833-1839, 6 vols.), edited by Diego Clemencíu;Don Quixote(London, 1899-1900, 2 vols.), edited by James Fitzmaurice-Kelly and John Ormsby;Don Quijote(Madrid, 1905-1906, 2 vols. in progress), edited by Clemente Cortejón;Rinconete y Cortadillo(Sevilla, 1905), edited by Francisco Rodriguez Marín;Epístola á Mateo Vázquez(Madrid, 1905), edited by E[milio] C[otarelo]; Julián Apráiz,Estudio histórico-crítico sobre las Novelas ejemplares de Cervantes(Madrid, 1901); Francisco A. de Icaza,Las Novelas ejemplares de Cervantes(Madrid, 1901); Francisco Rodríguez Marín,El Loaysa de “El Celoso Extremeño”(Sevilla, 1901); Narciso Díaz de Escovar,Apuntes escénicos cervantinos(Madrid, 1905); Manuel José García,Estudio crítico acerca del entremés “El Vizcaino fingido”(Madrid, 1905); Alfred Morel-Fatio,L’Espagne de Don QuichotteinÉtudes sur l’Espagne(Paris, 1895, 2me série); Julio Puyol y Alonso,Estado social que refleja “El Quijote”(Madrid, 1905); James Fitzmaurice-Kelly,Cervantes in England(London, 1905); Raymond Foulché-Delbose,Étude sur “La tia fingida,”in theRevue hispanique(Paris, 1899), vol. vi. pp. 256-306; Benedetto Croce,Due illustrazioni al “Viage del Parnaso,”in theHomenaje á Menéndez y Pelayo(Madrid, 1899), vol. i. pp. 161-193; Paul Groussac,Une Énigme littéraire: le Don Quichotte d’Avellaneda(Paris, 1903); Alonso Fernández de Avellaneda,El ingenioso hidalgo Don Quixote de la Mancha(Barcelona, [1905]), edited by Marcelino Menéndez y Pelayo; Julio Cejador y Franca,La Lengua de Cervantes(Madrid, 1905, &c.); Martin Fernández de Navarrete,Vida de Miguel de Cervantes Saavedra(Madrid, 1819); Cristóbal Perez Pastor,Documentos Cervantinos hasta ahora inéditos(Madrid, 1897-1902, 2 vols.); Emilio Cotardo y Mori,Efemérides Cervantinas(Madrid, 1905); Francisco Rodríguez Marín,Cervantes estudió en Sevilla, 1564-1565(Seville, 1905).
(J. F.-K.)
CERVERA, PASCUAL CERVERA Y TOPETE(1839-1909), Spanish admiral, was born at Medina Sidonia on the 18th of February 1839. He showed an early inclination for the sea, and his family sent him to the naval cadet school at the age of twelve. As a sub-lieutenant he took part in the naval operations on the coast of Morocco during the campaign of 1859-60. Then he was for some time engaged in operations in the Sulu Islands and the Philippines. Afterwards he was on the West Indian station during the early part of the first Cuban War (1868-78), returning to Spain in 1873 to serve on the Basque coast against the Carlists. He distinguished himself in defending the Carraca arsenal near Cadiz against the Federals in 1873. He won each step in his promotion up to flag-rank through his steadiness and brilliant conduct in action, and was awarded the crosses of the Orders of Military and Naval Merit, Isabella the Catholic, and St Hermengilde, besides several medals. Cervera had a great reputation for decision, unbending temper and honesty, before he was placed at the head of the Bilbao building-yards. This post he resigned after a few months in order to become minister of marine in 1892, in a cabinet presided over by Sagasta. He withdrew from the cabinet when he found that his colleagues, from political motives, declined to support him in making reforms and,on the other hand, unwisely cut down the naval estimates. When in 1898 the Spanish-American War (q.v.) broke out, he was chosen to command a squadron composed of four first-class cruisers, the “Maria Theresa,” his flagship, “Oquendo,” “Vizcaya,” and “Columbus,” and several destroyers. This ill-fated squadron only started upon its reckless cruise across the ocean after its gallant commander had repeatedly warned both the minister of marine and the prime minister, Sagasta, in despatches from Cadiz and from the Canary and Cape Verde Islands, that the ships were insufficiently provided with coal and ammunition. Some of them, indeed, even lacked proper guns. In compliance with the instructions of the government, Admiral Cervera made for the landlocked harbour of Santiago de Cuba, where he co-operated in the defence, landing some guns and a naval brigade. In spite of his energetic representations, Cervera received an order from Madrid, dictated by political considerations, to sally forth. It meant certain destruction. The gallant squadron met forces trebly superior to it, and was totally destroyed. The admiral, three of his captains, and 1800 sailors and marines were taken by the victors to Portsmouth, New Hampshire, U.S.A. After the war, Cervera and his captains were tried before the supreme naval and military court of the realm, which honourably acquitted them all. In 1901 he became vice-admiral, in 1902 was appointed chief of staff of the Spanish navy, and in 1903 was made life senator. He died at Puerto Real on the 3rd of April 1909.
CESAREVICH,or more properlyTsesarevich, the title of the heir-apparent to the Russian throne. The full official title isNasliednik Tsesarevich,i.e.“heir of Caesar,” and in Russia the heir to the throne is commonly called simplyNasliednik, the wordTsesarevichnever being used alone.Tsarevich, a form now much used in England, means simply any “king’s son”; it is an antiquated term now out of use in Russia, and was last borne as heir to the throne by the unfortunate Alexius, son of Peter the Great. The style of the wife of the tsesarevich isTsesarevna. The Cesarewitch handicap race at Newmarket, founded in 1839, was named after the prince who was afterwards Alexander II. of Russia, who paid a state visit to England that year.
CESARI, GIUSEPPE,called Il Cavaliere d’ Arpino (born in or about 1568 and created a “Cavaliere di Cristo” by Pope Clement VIII.), also named Il Giuseppino, an Italian painter, much encouraged at Rome and munificently rewarded. His father had been a native of Arpino, but Giuseppe himself was born in Rome. Cesari is stigmatized by Lanzi as not less the corrupter of taste in painting than Marino was in poetry; indeed, another of the nicknames of Cesari is “Il Marino de’ Pittori” (the pictorial Marino). There was spirit in Cesari’s heads of men and horses, and his frescoes in the Capitol (story of Romulus and Remus, &c.), which occupied him at intervals during forty years, are well coloured; but he drew the human form ill. His perspective is faulty, his extremities monotonous, and his chiaroscuro defective. He died in 1640, at the age of seventy-two, or perhaps of eighty, at Rome. Cesari ranks as the head of the “Idealists” of his period, as opposed to the “Naturalists,” of whom Michelangelo da Caravaggio was the leading champion,—the so-called “idealism” consisting more in reckless facility, and disregard of the common facts and common-sense of nature, than in anything to which so lofty a name could be properly accorded. He was a man of touchy and irascible character, and rose from penury to the height of opulence. His brother Bernardino assisted in many of his works.
CESAROTTI, MELCHIORE(1730-1808), Italian poet, was born at Padua in 1730, of a noble but impoverished family. At the university of his native place his literary progress procured for him at a very early age the chair of rhetoric, and in 1768 the professorship of Greek and Hebrew. On the invasion of Italy by the French, he gave his pen to their cause, received a pension, and was made knight of the iron crown by Napoleon I., to whom, in consequence, he addressed a bombastic and extravagantly flattering poem calledPronea. Cesarotti is best known as the translator of Homer and Ossian. Much praise cannot be given to his version of theIliad, for he has not scrupled to add, omit and modernize. Ossian, which he held to be the finest of poems, he has, on the other hand, considerably improved in translation; and the appearance of his version attracted much attention in Italy and France, and raised up many imitators of the Ossianic style. Cesarotti also produced a number of works in prose, including aCourse of Greek Literature, and essaysOn the Origin and Progress of the Poetic Art,On the Sources of the Pleasure derived from Tragedy,On the Philosophy of LanguageandOn the Philosophy of Taste, the last being a defence of his own great eccentricities in criticism. His weakness was a straining after novelty. His style is forcible, but full of Gallicisms.
A complete edition of his works, in 42 vols. 8vo. began to appear at Pisa in 1800, and was completed in 1813, after his death. SeeMemoirs, by Barbieri (Padua, 1810), andUn Filosofo delle lettere, by Alemanni (Turin, 1894).
A complete edition of his works, in 42 vols. 8vo. began to appear at Pisa in 1800, and was completed in 1813, after his death. SeeMemoirs, by Barbieri (Padua, 1810), andUn Filosofo delle lettere, by Alemanni (Turin, 1894).
CESENA(anc.Caesena), a town and episcopal see of Emilia, Italy, in the province of Forlì, 12 m. S.E. by rail from the town of Forlì, on the line between Bologna and Rimini, 144 ft. above sea-level. Pop. (1905) 12,245 (town); 43,468 (commune). The town is picturesquely situated at the foot of the slopes of the Apennines, and is crowned by a medieval fortress (Rocca), begun by the emperor Frederick I. (Barbarossa) probably, but altered and added to later. The cathedral has two fine marble altars by the Lombardi of Venice (or their school). The library, built for Domenico Malatesta in 1452 by Matteo Nuzio, is a fine early Renaissance building, and its internal arrangements, with the original desks to which the books are still chained, are especially well preserved (see J.W. Clark,The Care of Books, Cambridge, 1901, p. 199). In it are valuable MSS., many of which were used by Aldus Manutius. It also contains a picture gallery with a good “Presentation in the Temple” by Francesco Francia. There are some fine palaces in the town. Three-quarters of a mile south-east on the hill stands the handsome church of S. Maria del Monte, after the style of Bramante, with carved stalls of the 16th century. Wine, hemp and silk are the main articles of trade. About the ancient Caesena little is said in classical authors: it is mentioned as a station on the Via Aemilia and as a fortress in the wars of Theodoric and Narses. During the middle ages it was at first independent. In 1357 it was unsuccessfully defended by the wife of Francesco Ordelaffi, lord of Forlì, against the papal troops under Albornoz. In 1377 it was sacked by Cardinal Robert of Geneva (afterwards Clement VII., antipope). It was then held by the Malatesta of Rimini until 1465, when it came under the dominion of the church. Both Pius VI. (1717) and Pius VII. (1742) were born at Cesena.
(T. As.)
CESNOLA, LUIGI PALMA DI(1832-1904), Italian-American soldier and archaeologist, was born near Turin on the 29th of July 1832. Having served in the Austrian and Crimean Wars, in 1860 he went to New York, where he taught Italian and French and founded a military school for officers. He took part in the American Civil War as colonel of a cavalry regiment, and at Aldie (June 1863) was wounded and taken prisoner. He was released from Libby prison early in 1864, served in the Wilderness and Petersburg campaigns (1864-65) as a brigadier of cavalry, and at the close of the war was breveted brigadier-general. He was then appointed United States consul at Larnaca in Cyprus (1865-1877). During his stay in the island he carried on excavations, which resulted in the discovery of a large number of antiquities. The collection was purchased by the Metropolitan Museum of New York, and Cesnola became director in 1879. Doubt having been thrown by Gaston L. Feuerdant, in an article in the New YorkHerald(August 1880), upon the genuineness of his restorations, the matter was referred to a special committee, which pronounced in his favour.1He is the author ofCyprus, its ancient Cities, Tombs and Temples(1877), an interesting book of travel and of considerable service to the practical antiquary; and of aDescriptive Atlas of the Cesnola Collection of Cypriote Antiquities(3 vols., 1884-6). He died in New York on the 21st of November 1904. He was amember of several learned societies in Europe and America, and in 1897 he received a Congressional medal of honour for conspicuous military services.
His brother,Alessandro Palma di Cesnola, born in 1839, conducted excavations at Paphos (where he was U.S. vice-consul) and Salamis on behalf of the British government. The results of these are described inSalaminia(1882).
1For the Cesnola controversy see C.D. Cobham’sAttempt at a Bibliography of Cyprus(4th ed., 1900). See also articleCyprus.
1For the Cesnola controversy see C.D. Cobham’sAttempt at a Bibliography of Cyprus(4th ed., 1900). See also articleCyprus.
CESPEDES(in Ital.Cedaspe),PABLO DE(1538-1608), Spanish poet, painter, sculptor and architect, was born at Cordova, and was educated at Alcalá de Henares, where he studied theology and Oriental languages. On leaving the university, he went to Rome, where he became the pupil and friend of Federigo Zuccaro, under whose direction he studied particularly the works of Raphael and of Michelangelo. In 1560, while yet in Rome, proceedings were taken against him by the Inquisition at Valladolid on account of a letter which, found among the papers of the archbishop of Toledo, had been written by Cespedes during the preceding year, and in which he had spoken with great freedom against the holy office and the inquisitor-general, Fernando de Valdés. Cespedes remained in Rome at this critical moment, and he appears rightly to have treated the prosecution with derision. It is not known how he contrived to bring the proceedings to an end; he returned, however, to Spain a little before 1577, and in that year was installed in a prebend of the cathedral at Cordova, where he resided till his death. Pablo de Cespedes has been called the mostsavantof Spanish artists. According to his friend Francisco Pacheco, to whom posterity is indebted for the preservation of all of Cespedes’s verse that is extant, the school of Seville owes to him its introduction to the practice of chiaroscuro. He was a bold and correct draughtsman, a skilful anatomist, a master of colour and composition; and the influence he exerted to the advantage of early Spanish art was considerable. Cristobal de Vera, Juan de Peñalosa and Zambrano were among his pupils. His best picture is a Last Supper at Cordova, but there are good examples of his work at Seville and at Madrid. Cespedes was author of several opuscules in prose on subjects connected with his profession. Of his poem onThe Art of Paintingenough was preserved by Pacheco to enable us to form an opinion of the whole. It is esteemed the best didactic verse in Spanish; and it has been compared, not disadvantageously, with theGeorgics. It is written in strong and sonorous octaves, in the majestic declamatory vein of Fernando Herrera, and is not altogether so dull and lifeless as is most didactic verse. It contains a glowing eulogy of Michelangelo, and some excellent advice to young painters, insisting particularly on hard work and on the study of nature. The few fragments yet remaining, amounting in all to some six hundred lines, were first printed by Pacheco in his treatiseDel arte de la pintura, in 1649.
CÉSPEDES Y MENESES, GONZALO DE(1585?-1638), Spanish novelist, was born at Madrid about 1585. Nothing positive is known of him before the publication of his celebrated romance, thePoema trágico del Español Gerardo, y desengaño del amor lascivo(1615-1617); there is evidence that he had been sentenced to eight years at the galleys previous to the 1st of January 1620, and that the penalty had been remitted; but the nature of his offence is not stated. His treatment of political questions in theHistoria apologética en los sucesos del reyno de Aragón, y su ciudad de Zaragoza, años de 91 y 92(1622), having led to the confiscation of the book, Céspedes took up his residence at Saragossa and Lisbon. While in exile he issued a collection of short stories entitledHistorias peregrinas y exemplares(1623), the unfinished romanceVaria fortuna del soldado Píndaro(1626), and the first part of hisHistoria de Felipe IV.(1631), a fulsome eulogy which was rewarded by the author’s appointment as official historiographer to the Spanish king. Céspedes died on the 27th of January 1638. His novels, though written in a ponderous, affected style, display considerable imagination and insight into character. ThePoema trágicohas been utilized by Fletcher inThe Spanish Curateand inThe Maid of the Mill.
TheHistorias peregrinashas been reprinted (1906) with a valuable introduction by Sr. Cotarelo y Mori.
TheHistorias peregrinashas been reprinted (1906) with a valuable introduction by Sr. Cotarelo y Mori.
CESS(a shortened form of “assess”; the spelling is due to a mistaken connexion with “census”), a tax; a term formerly more particularly applied to local taxation, in which sense it still is used in Ireland; otherwise it has been superseded by “rate.” In India it is applied, with the qualifying word prefixed, to any taxation, such as “irrigation-cess” and the like, and in Scotland to the land-tax.
CESSIO BONORUM(Latin for a “surrender of goods”), in Roman law, a voluntary surrender of goods by a debtor to his creditors. It did not amount to a discharge unless the property ceded was sufficient for the purpose, but it secured the debtor from personal arrest. The creditors sold the goods in satisfaction,pro tanto, of their claims. The procedure ofcessio bonorumavoided infamy, and the debtor, though his after-acquired property might be proceeded against, could not be deprived of the bare necessaries of life. The main features of the Roman law ofcessio bonorumwere adopted in Scots law, and also in the French legal system. (See furtherBankruptcy.)
CESTI, MARC’ ANTONIO(1620?-1669?), Italian musical composer, was born at Florence about 1620. He was a pupil of Carissimi, and after holding a post somewhere in Florence asmaestro di cappellaentered the papal chapel in 1660. In 1666 he becameVice-Kapellmeisterat Vienna, and died at Venice in 1669. Cesti is known principally as a composer of operas, the most celebrated of which wereLa Dori(Venice, 1663) andIl Pomo d’ oro(Vienna, 1668). He was also a composer of chamber-cantatas, and his operas are notable for the pure and delicate style of their airs, more suited to the chamber than to the stage.
CESTIUS, LUCIUS,surnamed Pius, Latin rhetorician, flourished during the reign of Augustus. He was a native of Smyrna, a Greek by birth. According to Jerome, he was teaching Latin at Rome in the year 13B.C.He must have been living afterA.D.9, since we are told that he taunted the son of Quintilius Varus with his father’s defeat in the Teutoburgian forest (Seneca,Controv.i. 3, 10). Cestius was a man of great ability, but vain, quarrelsome and sarcastic. Before he left Asia, he was invited to dinner by Cicero’s son, then governor of the province. His host, being uncertain as to his identity, asked a slave who Cestius was; and on receiving the answer, “he is the man who said your father was illiterate,” ordered him to be flogged (Seneca,Suasoriae, vii. 13). As an orator in the schools Cestius enjoyed a great reputation, and was worshipped by his youthful pupils, one of whom imitated him so slavishly that he was nicknamed “my monkey” by his teacher (Seneca,Controv.ix. 3, 12). As a public orator, on the other hand, he was a failure. Although a Greek, he always used Latin in his declamations, and, although he was sometimes at a loss for Latin words, he never suffered from lack of ideas. Numerous specimens of his declamations will be found in the works of Seneca the rhetorician.
See the monographDe Lucio Cestio Pio, by F.G. Lindner (1858); T. Brzoska in Pauly-Wissowa’sRealencyclopadie, iii. 2 (1899); Teuffel-Schwabe,Hist, of Roman Lit.(Eng. tr.), § 268, 6; M. Schanz,Geschichte der romischen Litteratur, ii.
See the monographDe Lucio Cestio Pio, by F.G. Lindner (1858); T. Brzoska in Pauly-Wissowa’sRealencyclopadie, iii. 2 (1899); Teuffel-Schwabe,Hist, of Roman Lit.(Eng. tr.), § 268, 6; M. Schanz,Geschichte der romischen Litteratur, ii.
CESTUI, CESTUY,an Anglo-French word, meaning “that person,” which appears in the legal phrasescestui que trust,use, orvie. It is usually pronounced as “cetty.”Cestui que trustmeans literally “the person for whose benefit the trust” is created. Thecestui que trustis the person entitled to the equitable, as opposed to the legal, estate. Thus, if land be granted unto, and to the use of A. in trust for B., B. iscestui que trust, and A. trustee. The term, principally owing to its cumbersomeness, is being gradually superseded in modern law by that of “beneficiary.”Cestui que use(sometimescestui à que use) means “the person for whose benefit a use” is created (see TRUST).Cestui que vieis “the person for whose life” lands are held by another (seeRemainder).
CETACEA(from the Gr.κέτος, a whale), the name of the mammalian order represented by whales, dolphins, porpoises, &c. From their fish-like form, which is manifestly merely an adaptation to their purely aquatic life, these creatures are often regardedas fishes, although they are true mammals, with warm blood, and suckle their young.
The general form is essentially fish-like, the spindle-shaped body passing anteriorly into the head without any distinct neck, and posteriorly tapering gradually towards the extremity of the tail, which is provided with a pair of lateral, pointed expansions of skin supported by fibrous tissue, called “flukes,” forming a horizontal triangular propelling organ, notched behind in the middle line. The head is generally large, in some cases attaining more than one-third the entire length; and the mouth is wide, and bounded by stiff, immobile lips. The fore-limbs are reduced to flattened paddles, encased in a continuous skin, showing no external sign of division, and without trace of nails. There are no signs of hind-limbs visible externally. The surface of the skin is smooth and glistening, and devoid of hair, although in many species there are a few bristles in the neighbourhood of the mouth which may persist through life or be present only in the young state. Immediately beneath the skin is a thick layer of fat, held together by a mesh of tissue, constituting the “blubber,” which retains the heat of the body. In nearly all species a compressed dorsal fin is present. The eye is small, and not provided with a true lacrymal apparatus. The external ear is a minute aperture in the skin situated at a short distance behind the eye. The nostrils open separately or by a single crescentic aperture, near the vertex of the head.
The bones generally are spongy in texture, the cavities being filled with oil. In the vertebral column, the cervical region is short and immobile, and the vertebrae, always seven in number, are in many species more or less fused together into a solid mass. The odontoid process of the second cervical vertebra, when that bone is free, is usually very obtuse, or even obsolete. In a paper on the form and function of the cervical vertebrae published in theJenaische Zeitschriftfor 1905, Dr O. Reche points out that the shortening and soldering is most pronounced in species which, like the right-whales, live entirely on minute organisms, to capture which there is no necessity to turn the head at all. Accordingly we find that in these whales the whole seven cervical vertebrae are fused into an immovable solid mass, of which the compound elements, with the exception of the first and second, are but little thicker than plates. On the other hand, in the finner-whales, several of which live exclusively on fish, and thus require a certain amount of mobility in the head and neck, we find all the cervical vertebrae much thicker and entirely separate from one another. Among the dolphin group the narwhal and the white whale, or beluga, are distinguished from all other cetaceans by the great comparative length of their cervical vertebrae, all of which are completely free. In the case of the narwhal such an abnormal structure is easily accounted for, seeing that to use effectively the long tusk with which the male is armed a considerable amount of mobility in the neck is absolutely essential. The beluga, too, which is believed to feed on large and active fishes, would likewise seem to require mobility in the same region in order to effect their capture. On the other hand, the porpoise preys on herrings, pilchards and mackerel, which in their densely packed shoals must apparently fall an easy prey with but little exertion on the part of their captor, and we accordingly find all the neck-vertebrae very short, and at least six out of the seven coalesced into a solid immovable mass. None of the vertebrae are united to form a sacrum. The lumbar and caudal vertebrae are numerous and large, and, as their arches are not connected by articular processes (zygapophyses), they are capable of free motion in all directions. The caps, or epiphyses, at the end of the vertebral bodies are flattened disks, not uniting until after the animal has attained its full dimensions. There are largely developed chevron-bones on the under side of the tail, the presence of which indicates the distinction between caudal and lumbar vertebrae.In the skull, the brain-case is short, broad and high, almost spherical, in fact (fig. 1). The supra-occipital bone rises upwards and forwards from the foramen magnum, to meet the frontals at the vertex, completely excluding the parietals from the upper region; and the frontals are expanded laterally to form the roof of the orbits. The nasal aperture opens upwards, and has in front of it a more or less horizontally prolonged beak, formed of the maxillae, premaxillae, vomer, and mesethmoid cartilage, extending forwards to form the upper jaw or roof of the mouth.There are no clavicles. The humerus is freely movable on the scapula at the shoulder-joint, but beyond this the articulations of the limb are imperfect; the flattened ends of the bones coming in contact, with fibrous tissue interposed, allowing of scarcely any motion. The radius and ulna are distinct, and about equally developed, and much flattened, as are all the bones of the flippers. There are four, or more commonly five, digits, and the number of the phalanges of the second and third always exceeds the normal number in mammals, sometimes considerably; they present the exceptional character of having epiphyses at both ends. The pelvis is represented by a pair of small rod-like bones placed longitudinally, suspended below and at some distance from the vertebral column at the commencement of the tail. In some species, to the outer surface of these are fixed other small bones or cartilages, the rudiments of the hind-limb.Fig.1.—A Section of the Skull of a Black-Fish (Globicephalus melas).PMx, Premaxilla.Mx, Maxilla.ME, Ossified portion of the mesethmoid.an, Nostrils.Na, Nasal.IP, Inter-parietal.Fr, Frontal.Pa, Parietal.SO, Supra-occipital.ExO, Ex-occipital.BO, Basi-occipital.Sq, Squamosal.Per, Periotic.AS, Alisphenoid.PS, Presphenoid.Pt, Pterygoid.pn, Posterior nares.Pl, Palatine.Vo, Vomer.s, Symphysis of lower jaw.id, Inferior dental canal.cp, Coronoid process of lower jaw.cd, Condyle.a, Angle.sh, Stylo-hyal.bh, Basi-hyal.th, Thyro-hyal.Teeth are generally present, but exceedingly variable in number. In existing species, they are of simple, uniform character, with conical or compressed crowns and single roots, and are never preceded by milk-teeth. In the whalebone whales teeth are absent (except in the foetal condition), and the palate is provided with numerous transversely placed horny plates, forming the “whalebone.” Salivary glands are rudimentary or absent. The stomach is complex, and the intestine simple, and only in some species provided with a small caecum. The liver is little fissured, and there is no gall-bladder. The blood-vascular system is complicated by net-like expansions of both arteries and veins, orretia mirabilia, The larynx is of peculiar shape, the arytenoid cartilages and the epiglottis being elongated, and forming a tubular prolongation, which projects into the posterior nares, and when embraced by the soft palate forms a continuous passage between the nostrils and the trachea, or wind-pipe, in a more perfect manner. The brain is relatively large, round in form, with its surface divided into numerous and complex convolutions. The kidneys are deeply lobulated; the testes are abdominal; and there are no vesiculae seminales nor an os penis. The uterus is bicornuate; the placenta non-deciduate and diffuse. The two teats are placed in depressions on each side of the genital aperture. The ducts of the milk-glands are dilated during suckling into large reservoirs, into which the milk collects, and from which it is injected by the action of a muscle into the mouth of the young animal, so that sucking under water is greatly facilitated.
The bones generally are spongy in texture, the cavities being filled with oil. In the vertebral column, the cervical region is short and immobile, and the vertebrae, always seven in number, are in many species more or less fused together into a solid mass. The odontoid process of the second cervical vertebra, when that bone is free, is usually very obtuse, or even obsolete. In a paper on the form and function of the cervical vertebrae published in theJenaische Zeitschriftfor 1905, Dr O. Reche points out that the shortening and soldering is most pronounced in species which, like the right-whales, live entirely on minute organisms, to capture which there is no necessity to turn the head at all. Accordingly we find that in these whales the whole seven cervical vertebrae are fused into an immovable solid mass, of which the compound elements, with the exception of the first and second, are but little thicker than plates. On the other hand, in the finner-whales, several of which live exclusively on fish, and thus require a certain amount of mobility in the head and neck, we find all the cervical vertebrae much thicker and entirely separate from one another. Among the dolphin group the narwhal and the white whale, or beluga, are distinguished from all other cetaceans by the great comparative length of their cervical vertebrae, all of which are completely free. In the case of the narwhal such an abnormal structure is easily accounted for, seeing that to use effectively the long tusk with which the male is armed a considerable amount of mobility in the neck is absolutely essential. The beluga, too, which is believed to feed on large and active fishes, would likewise seem to require mobility in the same region in order to effect their capture. On the other hand, the porpoise preys on herrings, pilchards and mackerel, which in their densely packed shoals must apparently fall an easy prey with but little exertion on the part of their captor, and we accordingly find all the neck-vertebrae very short, and at least six out of the seven coalesced into a solid immovable mass. None of the vertebrae are united to form a sacrum. The lumbar and caudal vertebrae are numerous and large, and, as their arches are not connected by articular processes (zygapophyses), they are capable of free motion in all directions. The caps, or epiphyses, at the end of the vertebral bodies are flattened disks, not uniting until after the animal has attained its full dimensions. There are largely developed chevron-bones on the under side of the tail, the presence of which indicates the distinction between caudal and lumbar vertebrae.
In the skull, the brain-case is short, broad and high, almost spherical, in fact (fig. 1). The supra-occipital bone rises upwards and forwards from the foramen magnum, to meet the frontals at the vertex, completely excluding the parietals from the upper region; and the frontals are expanded laterally to form the roof of the orbits. The nasal aperture opens upwards, and has in front of it a more or less horizontally prolonged beak, formed of the maxillae, premaxillae, vomer, and mesethmoid cartilage, extending forwards to form the upper jaw or roof of the mouth.
There are no clavicles. The humerus is freely movable on the scapula at the shoulder-joint, but beyond this the articulations of the limb are imperfect; the flattened ends of the bones coming in contact, with fibrous tissue interposed, allowing of scarcely any motion. The radius and ulna are distinct, and about equally developed, and much flattened, as are all the bones of the flippers. There are four, or more commonly five, digits, and the number of the phalanges of the second and third always exceeds the normal number in mammals, sometimes considerably; they present the exceptional character of having epiphyses at both ends. The pelvis is represented by a pair of small rod-like bones placed longitudinally, suspended below and at some distance from the vertebral column at the commencement of the tail. In some species, to the outer surface of these are fixed other small bones or cartilages, the rudiments of the hind-limb.
PMx, Premaxilla.
Mx, Maxilla.
ME, Ossified portion of the mesethmoid.
an, Nostrils.
Na, Nasal.
IP, Inter-parietal.
Fr, Frontal.
Pa, Parietal.
SO, Supra-occipital.
ExO, Ex-occipital.
BO, Basi-occipital.
Sq, Squamosal.
Per, Periotic.
AS, Alisphenoid.
PS, Presphenoid.
Pt, Pterygoid.
pn, Posterior nares.
Pl, Palatine.
Vo, Vomer.
s, Symphysis of lower jaw.
id, Inferior dental canal.
cp, Coronoid process of lower jaw.
cd, Condyle.
a, Angle.
sh, Stylo-hyal.
bh, Basi-hyal.
th, Thyro-hyal.
Teeth are generally present, but exceedingly variable in number. In existing species, they are of simple, uniform character, with conical or compressed crowns and single roots, and are never preceded by milk-teeth. In the whalebone whales teeth are absent (except in the foetal condition), and the palate is provided with numerous transversely placed horny plates, forming the “whalebone.” Salivary glands are rudimentary or absent. The stomach is complex, and the intestine simple, and only in some species provided with a small caecum. The liver is little fissured, and there is no gall-bladder. The blood-vascular system is complicated by net-like expansions of both arteries and veins, orretia mirabilia, The larynx is of peculiar shape, the arytenoid cartilages and the epiglottis being elongated, and forming a tubular prolongation, which projects into the posterior nares, and when embraced by the soft palate forms a continuous passage between the nostrils and the trachea, or wind-pipe, in a more perfect manner. The brain is relatively large, round in form, with its surface divided into numerous and complex convolutions. The kidneys are deeply lobulated; the testes are abdominal; and there are no vesiculae seminales nor an os penis. The uterus is bicornuate; the placenta non-deciduate and diffuse. The two teats are placed in depressions on each side of the genital aperture. The ducts of the milk-glands are dilated during suckling into large reservoirs, into which the milk collects, and from which it is injected by the action of a muscle into the mouth of the young animal, so that sucking under water is greatly facilitated.
Whales and porpoises are found in all seas, and some dolphins and porpoises are inhabitants of the larger rivers of South America and Asia. Their organization necessitates their passing their life entirely in the water, as on land they are absolutely helpless. They have, however, to rise very frequently to the surface for the purpose of respiration; and, in relation to the upward and downward movement in the water thus necessitated, the principal instrument of motion, the tail, is expanded horizontally. The position of the nostril on the highest part of the head is important for this mode of life, as it is the only part of the body the exposureof which above the surface is absolutely necessary. Of numerous erroneous ideas connected with natural history, few are so widespread as that whales spout through their blow-holes water taken in at the mouth. But the “spouting,” or “blowing,” of whales is nothing more than the ordinary act of expiration, which, taking place at longer intervals than land-animals, is performed with a greater 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 charged with vapour in consequence of the respiratory changes. This rapidly condensing in the cold atmosphere in which the phenomenon is often observed, forms a column of steam or spray, which has been taken for water. It happens, however, especially when the surface of the ocean is agitated into waves, that the animal commences its expiratory puff before the orifice has cleared the top of the water, some of which may thus be driven upwards with the blast, tending to complete the illusion. From photographs of spouting rorquals, it appears that the height and volume of the “spout” of all the species is much less than was supposed to be the case by the older observers; even that of the huge “sulphur-bottom” (Balaenoptera sibbaldi) averaging only about 14 ft. in height, although it may occasionally reach 20 ft.
As regards their powers of hearing, the capacity of cetaceans for receiving (and acting upon) sound-waves is demonstrated by the practice of shouting on the part of the fishermen when engaged in driving a shoal of porpoises or black-fish into shallow water, for the purpose of frightening their intended victims. As regards the possession of a voice by cetaceans, it is stated that one species, the “buckelwal” of the Germans, utters during the breeding-season a prolonged scream, comparable to the scream of a steam-siren, and embracing the whole musical scale, from base to treble. In respect of anatomical considerations, it is true that the external ear is much reduced, the “pinna” being absent, and the tube or “meatus” of very small calibre. On the other hand, the internal auditory organs are developed on the plan of those of ordinary mammals, but display certain peculiar modifications (notably the remarkable shell-like form of the tympanic bone) for intensifying and strengthening the sound-waves as they are received from the water. It seems, therefore, perfectly evident that whales must hear when in the water. This inference is confirmed by the comparatively small development of the other sense-organs. The eye, for instance, is very small, and can be of little use even at the comparatively small depths to which whales are now believed to descend. Again, the sense of smell, judging from the rudimentary condition of the olfactory organs, must be in abeyance; and whales have no sense-organs comparable to the lateral-line-system of fishes. Consequently, it would seem that when below the surface of the water they must depend chiefly upon the sense of hearing. Probably this sense is so highly developed as to enable the animals, in the midst of the vibrations made by the screw-like movements of the tail, or flukes, to distinguish the sound (or the vibrations) made by the impact of water against rocks, even in a dead calm, and, in the case of piscivorous species, to recognize by the pulse in the water the presence of a shoal of fish. Failing this explanation, it is difficult to imagine how whales can find their way about in the semi-darkness, and avoid collisions with rocks and rock-bound coasts.
In the ChristianiaNyt Magazin for Naturvidenskaberne, vol. xxxviii., Dr G. Guldberg has published some observations on the body-temperature of the Cetacea, in which he shows how extremely imperfect is our knowledge of this subject. As he remarks, it is a matter of extreme difficulty to obtain the temperature of living cetaceans, although this has been taken in the case of a white-whale and a dolphin, which some years ago were kept in confinement in a pond in the United States. With the larger whales such a mode of procedure is, however, obviously quite impracticable, and we have, accordingly, to rely onpost-mortemobservations. The layer of blubber by which all cetaceans are protected from cold renders thepost-mortemrefrigeration of the blood a much slower process than in most mammals, so that such observations have a much higher value than might at first be supposed to be the case. Indeed, the blood-temperature of a specimen of Sibbald’s rorqual three days after death still stood at 34° C. The various observations that have been taken have afforded the following results in individual cases: Sperm-whale, 40° C.; Greenland right-whale, 38.8° C.; porpoise, 35.6° C.; liver of a second individual, 37.8° C.; common rorqual, 35.4° C.; dolphin, 35.6° C. The average blood-temperature of man is 37° C., and that of other mammals 39° C.; while that of birds is 42 C. The record of 40° C. in the case of the sperm-whale seems to indicate that at least some cetaceans have a relatively high temperature.
In the ChristianiaNyt Magazin for Naturvidenskaberne, vol. xxxviii., Dr G. Guldberg has published some observations on the body-temperature of the Cetacea, in which he shows how extremely imperfect is our knowledge of this subject. As he remarks, it is a matter of extreme difficulty to obtain the temperature of living cetaceans, although this has been taken in the case of a white-whale and a dolphin, which some years ago were kept in confinement in a pond in the United States. With the larger whales such a mode of procedure is, however, obviously quite impracticable, and we have, accordingly, to rely onpost-mortemobservations. The layer of blubber by which all cetaceans are protected from cold renders thepost-mortemrefrigeration of the blood a much slower process than in most mammals, so that such observations have a much higher value than might at first be supposed to be the case. Indeed, the blood-temperature of a specimen of Sibbald’s rorqual three days after death still stood at 34° C. The various observations that have been taken have afforded the following results in individual cases: Sperm-whale, 40° C.; Greenland right-whale, 38.8° C.; porpoise, 35.6° C.; liver of a second individual, 37.8° C.; common rorqual, 35.4° C.; dolphin, 35.6° C. The average blood-temperature of man is 37° C., and that of other mammals 39° C.; while that of birds is 42 C. The record of 40° C. in the case of the sperm-whale seems to indicate that at least some cetaceans have a relatively high temperature.
With the possible exception of one West African dolphin, all the Cetacea are predaceous, subsisting on living animal food of some kind. One kind alone (Orca) eats other warm-blooded animals, as seals, and even members of its own order, both large and small. Many feed on fish, others on small floating crustaceans, pteropods and jelly-fishes, while the principal staple of the food of many is constituted by cuttle-fishes and squids. In size cetaceans vary much, some of the smaller dolphins scarcely exceeding 4 ft. in length, while whales are the most colossal of all animals. It is true that many statements of their bulk are exaggerated, but the actual dimensions of the larger species exceed those of all other animals, not even excluding the extinct dinosaurian reptiles. With some exceptions, cetaceans are generally timid, inoffensive animals, active in their movements and affectionate in their disposition towards one another, especially the mother towards the young, of which there is usually but one, or at most two at a time. They are generally gregarious, swimming in herds or “schools,” sometimes amounting to many thousands in number; though some species are met with either singly or in pairs.
Commercially these animals are of importance on account of the oil yielded by the blubber of all of them; while whalebone, spermaceti and ambergris are still more valuable products yielded by certain species. Within the last few years whalebone has been sold in America for £2900 per ton, while it is also asserted that £3000 per ton has been paid for two and a quarter tons at Aberdeen, although there seems to be some degree of doubt attaching to the statement. Soon after the middle of the last century, the price of this commodity was as low as £150 per ton, but, according to Mr Frank Buckland, it suddenly leapt up to £620 with the introduction of “crinoline” into ladies’ costume, and it has apparently been on the rise ever since. Ambergris, which is very largely used in perfumery, is solely a product of the sperm-whale, and appears to be a kind of biliary calculus. It generally contains a number of the horny beaks of the cuttle-fishes and squids upon which these whales chiefly feed. Its market-price is subject to considerable variation, but from £3 to £4 per oz. is the usual average for samples of good quality. In 1898 a merchant in Mincing Lane was the owner of a lump of ambergris weighing 270 ℔, which was sold in Paris for about 85 s. per oz., or £18,360.
Whalebone Whales.—Existing Cetacea are divisible into two sections, or suborders, the relationships of which are by no means clearly apparent. The first section is that of the whalebone whales, or Mystacoceti, in which no functional teeth are developed, although there are tooth-germs during foetal life. The palate is furnished with plates of baleen or whalebone; the skull is symmetrical; and the nasal bones form a roof to the nasal passages, which are directed upwards and forwards. The maxilla is produced in front of, but not over, the orbital process of the frontal. The lacrymal is small and distinct from the jugal. The tympanic is welded with the periotic, which is attached to the base of the skull by two strong diverging processes. The olfactory organ is distinctly developed. The two halves of the lower jaw are arched outwards, their anterior ends meeting at an angle, and connected by fibrous tissue without any symphysis. All the ribs at their upper extremity articulate only with the transverse processes of the vertebrae; their capitular processes when present not articulating directly with the bodies of the vertebrae. The sternum is composed of a single piece, and articulates only with a single pair of ribs; and there are no ossified sternal ribs. External openings of nostrils distinct from each other, longitudinal. A short conical caecum.When in the foetal state these whales have numerous minute teeth lying in the dental groove of both upper and lower jaws. They are best developed about the middle of foetal life, after which they are absorbed, and no trace of them remains at the time of birth. The whalebone does not make its appearance until after birth; and consists of a series of flattened horny plates, between three and four hundred in number, on each side of the palate, with a bare interval along the middle line. The plates are placed transversely to the long axis of the palate, with short intervals between them. Each plate or blade is somewhat triangular in form, with the baseattached to the palate and the apex hanging downwards. The outer edge of the blade is hard and smooth, but the inner edge and apex fray out into long bristly fibres, so that the roof of the whale’s mouth looks as if covered with hair, as described by Aristotle. At the inner edge of each principal blade are two or three much smaller or subsidiary blades. The principal blades are longest near the middle of the series, and gradually diminish towards the front and back of the mouth. The horny plates grow from a fibrous and vascular matrix, which covers the palatal surface of the maxillae, and sends out plate-like processes, one of which penetrates the base of each blade. Moreover, the free edges of these processes are covered with long vascular thread-like papillae, one of which forms the central axis of each of the hair-like fibres mainly composing the blade. A transverse section of fresh whalebone shows that it is made up of numbers of these soft vascular papillae, circular in outline, and surrounded by concentrically arranged epidermic cells, the whole bound together by other epidermic cells, that constitute the smooth (so-called “enamel”) surface of the blade, which, disintegrating at the free edge, allows the individual fibres to become loose and assume a hair-like appearance.Whalebone really consists of modified papillae of the mucous membrane of the mouth, with an excessive and horny epithelial development. The blades are supported and bound together for a certain distance from their base, by a mass of less hardened epithelium, secreted by the surface of the palatal membrane or matrix of the whalebone in the intervals of the plate-like processes. This is the “gum” of the whalers. Whalebone varies much in colour in different species; in some it is almost jet black, in others slate colour, horn colour, yellow, or even creamy-white. In some descriptions the blades are variegated with longitudinal stripes of different hues. It differs also greatly in other respects, being short, thick, coarse, and stiff in some cases, and greatly elongated and highly elastic in those species in which it has attained its fullest development. Its function is to strain the water from the small marine molluscs, crustaceans, or fish upon which the whales subsist. In feeding, whales fill the immense mouth with water containing shoals of these small creatures, and then, on closing the jaws and raising the tongue, so as to diminish the cavity of the mouth, the water streams out through the narrow intervals between the hairy fringe of the whalebone blades, and escapes through the lips, leaving the living prey to be swallowed.Although sometimes divided into two families,BalaenidaeandBalaenopteridae, whalebone-whales are best included in a single family group under the former name. The typical members of this family are the so-called right-whales, forming the genusBalaena, in which there are no folds on the throat and chest, and no back-fin; while the cervical vertebrae are fused into a single mass. The flippers are short and broad, with five digits; the head is very large and the whalebone very long and narrow, highly elastic and black; while the scapula is high, with a distinct coracoid and coronoid process. This genus contains the well-known Greenland right-whale (B. mysticetus) of the Arctic seas, the whalebone and oil of which are so much valued in commerce, and also other whales, distinguished by having the head somewhat smaller in proportion to the body, with shorter whalebone and a larger number of vertebrae. These inhabit the temperate seas of both northern and southern hemispheres, and have been divided into species in accordance with their geographical distribution, such asB. biscayensisof the North Atlantic,B. japonicaof the North Pacific,B. australisof the South Atlantic, andB. antipodarumandnovae-zelandiaeof the South Pacific; but the differences between them are so small that they may probably be regarded as races of a single species, the black whale (B. australis). On the head these whales carry a peculiar structure which is known to whalers as the “bonnet.” This is a large horny excrescence, worn into hollows like a much-denuded piece of limestone rock, growing probably in the neighbourhood of the blow-hole. More than one theory has been suggested to account for its presence. One suggestion is that it indicates the descent of whales from rhinoceros-like mammals; another that this species of whale is in the habit of rubbing against rocks in order to free itself from barnacles, and thus produces a kind of corn—although why on the nose alone is not stated. Dr W.G. Ridewood, however, considers that the structure is due to the fact that the horny layers which are produced all over the skin are not shed on this particular spot.The pigmy whale (Neobalaena marginata) represents a genus agreeing with the right-whales in the absence of throat-flutings, and with the rorquals in the presence of a dorsal fin. The cervical vertebrae are united, and there are only 43 vertebrae altogether. The flippers are small, narrow, and with only four digits. The ribs remarkably expanded and flattened; the scapula low and broad, with completely developed acromion and coracoid processes. The whalebone is long, slender, elastic and white. The species which inhabits the South American, Australian and New Zealand seas is the smallest of the whalebone-whales, being not more than 20 ft. in length.In contrast to the preceding is the great grey whale (Rachianectes glaucus) of the North Pacific, which combines the relatively small head, elongated shape, and narrow flippers of the fin-whales, with the smooth throat and absence of a back-fin distinctive of the right-whales. The whalebone is shorter and coarser than in any other species. In the skeleton the cervical vertebrae are free, and the first two ribs on each side expanded and united to form a large bony shield. In the humpback-whale (Megaptera longimanaorboops) the head is of moderate size, the whalebone-plates are short and wide, and the cervical vertebrae free. The skin of the throat is fluted so as to form an expansible pouch; there is a low back-fin; and the flippers, which have four digits each, are extremely long, equalling about one-fourth the total length of the animal. The acromion and coracoid processes of the scapula are rudimentary. SeeHumpback-Whale.The right-whales are built for cruising slowly about in search of the shoals of small floating invertebrates which form their food, and are consequently broad in beam, with a float-shaped body and immovable neck. The humpback is of somewhat similar build, but with a smaller head, and probably attains considerable speed owing to the length of its flippers. The finners, or rorquals (Balaenoptera), which prey largely on fish, are built entirely for speed, and are the ocean greyhounds of the group. Their bodies are consequently long and attenuated, and their necks are partially mobile; while they are furnished with capacious pouches for storing their food. They chiefly differ from the humpback by the smaller head, long and slender build, small, narrow, and pointed flippers, each containing four digits, and the large acromion and coracoid processes to the low and broad scapula. Rorquals are found in almost every sea. Among them are the most gigantic of all animals,B. sibbaldi, which attains the length of 80 ft., and the smallB. rostrata, which does not exceed 30. There are certainly four distinct modifications of this genus, represented by the two just mentioned, and byB. musculusandB. borealis, all inhabitants of British seas, but the question whether almost identical forms found in the Indian, Southern and Pacific Oceans are to be regarded as specifically identical or as distinct awaits future researches, although some of these have already received distinct names. SeeRorqual.In the report on the zoology of the “Discovery” expedition, published in 1907 by the British Museum, E.A. Wilson describes a whale frequenting the fringe of the Antarctic ice which indicates a new generic type. Mainly black in colour, these whales measure about 20 or 30 ft. in length, and have a tall dorsal fin like that of a killer.Toothed Whales.—-The second suborder is represented by the toothed whales, or Odontoceti, in which there is no whalebone, and teeth, generally numerous, though sometimes reduced to a single pair, and occasionally wanting, are normally developed. Unlike that of the whalebone-whales, the upper surface of the skull is more or less unsymmetrical. The nasal bones are in the form of nodules or flattened plates, applied closely to the frontals, and not forming any part of the roof to the nasal passage, which is directed upwards and backwards. The olfactory organ is rudimentary or absent. Hinder end of the maxilla expanded and covering the greater part of the orbital plate of the frontal bone. Lacrymal bone either inseparable from the jugal, or, if distinct, large, and forming part of the roof of the orbit. Tympanic bone not welded with the periotic, which is usually only attached to the rest of the skull by ligament. Two halves of the lower jaw nearly straight, expanded in height posteriorly, with a wide funnel-shaped aperture to the dental canal, and coming in contact in front by a flat surface of variable length, but constituting a symphysis. Several of the anterior ribs with well-developed capitular processes, which articulate with the bodies of the vertebrae. Sternum almost always composed of several pieces, placed one behind the other, with which several pairs of ribs are connected by well-developed cartilaginous or ossified sternal ribs. External respiratory aperture single, the two nostrils uniting before they reach the surface, usually in the form of a transverse sub-crescentic valvular aperture, situated on the top of the head. Flippers with five digits, though the first and fifth are usually little developed. No caecum, except inPlatanista.The first family,Physeteridae, is typified by the sperm-whale, and characterized by the absence of functional teeth in the upper jaw; the lower teeth being various, and often much reduced in number. Bones of the skull raised so as to form an elevated prominence or crest behind the nostrils. Pterygoid bones thick, produced backwards, meeting in the middle line, and not involuted to form the outer wall of the post-palatine air-sinuses, but simply hollowed on their outer side. Transverse processes of the arches of the dorsal vertebrae, to which the tubercles of the ribs are attached, ceasing abruptly near the end of the series, and replaced by processes on the body at a lower level, and serially homologous anteriorly with the heads of the ribs, and posteriorly with the transverse processes of the lumbar vertebrae. Costal cartilages not ossified.The first group, orPhyseterinae, includes the sperm-whale itself and is characterized by the presence of a full series of lower teeth, which are set in a groove in place of sockets, the groove being imperfectly divided by partial septa, and the teeth held in place by the strong, fibrous gum. No distinct lacrymal bone. Skull strikingly asymmetrical in the region of the nasal apertures, in consequence of the left opening greatly exceeding the right in size.Fig.2.—Skull of Sperm-Whale (Physeter macrocephalus).In the sperm-whale (Physeter macrocephalus) the upper teeth are apparently of uncertain number, rudimentary and functionless, being embedded in the gum. Lower jaw with from 20 to 25 teeth on each side, stout, conical, recurved and pointed at the apexuntil they are worn, without enamel. Upper surface of the skull concave; its posterior and lateral edges raised into a very high and greatly compressed semicircular crest or wall (fig. 2). Zygomatic processes of jugal bones thick and massive. Muzzle greatly elongated, broad at the base, and gradually tapering to the apex. Lower jaw exceedingly long and narrow, the symphysis being more than half the length. Vertebrae: C 7, D 11, L 8, Ca 24; total 50. Atlas, or first vertebra, free; all the other cervical vertebrae united by their bodies and spines into a single mass. Eleventh pair of ribs rudimentary. Head about one-third the length of the body; very massive, high and truncated, and rather compressed in front; owing its huge size and form mainly to the accumulation of a mass of fatty tissue filling the large hollow on the upper surface of the skull and overlying the long muzzle. The single blow-hole is longitudinal, slightly S-shaped, and placed at the upper and anterior extremity of the head to the left side of the middle line. The opening of the mouth is on the under side of the head, considerably behind the end of the snout. Flippers short, broad and truncated. Dorsal fin represented by a low protuberance. SeeSperm-Whale.Fig.3.—Bottle-nose (Hyperoödon rostratus). From a specimen taken off the coast of Scotland, 1882.In the lesser or pigmy sperm-whale (Cogia breviceps) there may be a pair of rudimentary teeth in the upper jaw, while on each side of the lower jaw there are from 9 to 12 rather long, slender, pointed and curved teeth, with a coating of enamel. Upper surface of the skull concave, with thick, raised, posterior and lateral margins, massive and rounded at their anterior terminations above the orbits. Muzzle not longer than the cranial position of the skull, broad at the base, and rapidly tapering to the apex. Zygomatic process of the jugal rod-like. Lower jaw with symphysis less than half its length. Vertebrae: C 7, D 13 or 14, L and Ca 30; total 50 or 51. All the cervical vertebrae united by their bodies and arches. The head is about one-sixth of the length of the body, and obtusely pointed in front; the mouth small and placed far below the apex of the snout; the blow-hole crescentic, and placed obliquely on the crown of the head in advance of the eyes and to the left of the middle line; while the flippers are bluntly sickle-shaped, and the back-fin triangular. This species attains a length of from 9 to 13 ft.A second subfamily is represented by the bottle-noses and beaked whales, and known as theZiphiinae. In this group the lower teeth are rudimentary and concealed in the gum, except one, or rarely two, pairs which may be largely developed, especially in the male. There is a distinct lacrymal bone. Externally the mouth is produced into a slender rostrum or beak, from above which the rounded eminence formed by a cushion of fat resting on the cranium in front of the blow-hole rises somewhat abruptly. The blow-hole is single, crescentic and median, as in theDelphinidae. Flippers small, ovate, with five digits moderately well developed. A small obtuse dorsal fin situated considerably behind the middle of the back. Longitudinal grooves on each side of the skin of the throat, diverging posteriorly, and nearly meeting in front. In external characters and habits the whales of this group closely resemble each other. They appear to be almost exclusively feeders on cuttle-fishes, and occur either singly, in pairs, or in small herds. By their dental and osteological characters they are easily separated into four genera.In the first of these,Hyperoödon, or bottle-nose, there is a small conical pointed tooth at the apex of each half of the lower jaw, concealed by the gum during life. Skull with the upper ends of the premaxillae rising suddenly behind the nostrils to the vertex and expanded laterally, their outer edges curving backwards and their anterior surfaces arching forwards and overhanging the nostrils; the right larger than the left. Nasal bones lying in the hollow between the upper extremities of the premaxillae, strongly concave in the middle line and in front; their outer edges, especially that of the right, expanded over the front of the inner border of the maxilla. Very high longitudinal crests on the maxillae at the base of the beak, extending backwards almost to the nostrils, approaching each other in the middle line above; sometimes compressed and sometimes so massive that their inner edges come almost in contact. Preorbital notch distinct, and mesethmoid cartilage slightly ossified. Vertebrae: C 7, D 9, L 10, Ca 19; total 45. All the cervical vertebrae united. Upper surface of the head in front of the blow-hole very prominent and rounded, rising abruptly from above the small, distinct snout. Two species are known. SeeBottle-nose Whale.The typical representative of the beaked whales isZiphius cuvieri, in which there is a single conical tooth of moderate size on each side close to the anterior extremity of the lower jaw, directed forwards and upwards. Skull with the premaxillae immediately in front and at the sides of the nostrils expanded, hollowed, with elevated lateral margins, the posterior ends rising to the vertex and curving forwards, the right being considerably more developed than the left. The conjoint nasals form a pronounced symmetrical eminence at the top of the skull, projecting forwards over the nostrils, flat above, prominent and rounded in the middle line in front, and separated by a notch on each side from the premaxillae. Preorbital notch not distinct. Rostrum (seen from above) triangular, tapering from the base to the apex; upper and outer edges of maxillae at base of rostrum raised into low roughened tuberosities. Mesethmoid cartilage densely ossified in adult age, and coalescing with the surrounding bones of the rostrum. Vertebrae: C 7, D 10, L 10, Ca 22; total 49. The three anterior cervical vertebrae united, the rest free.Fig.4.—Sowerby’s Beaked Whale (Mesoplodon bidens).In the numerous species of the allied genusMesoplodonthere is a much-compressed and pointed tooth in each half of the lower jaw, variously situated, but generally at some distance behind the apex; its point directed upwards, and often somewhat backwards, occasionally developed to a great size. In the skull the region round the nostrils is as inHyperoödon, except that the nasals are narrow and more sunk between the upper ends of the premaxillae; like those ofHyperoödon, they are concave in the middle line in front and above. No maxillary tuberosities. Preorbital notch not very distinct. Rostrum long and narrow. Mesethmoid in the adult ossified in its entire length, and coalescing with the surrounding bones. Vertebrae: C 7, D 10, L 10 or 11, Ca 19 or 20; total 46 to 48. Two or three anterior cervicals united, the rest usually free.Fig.5.—Skull of a Beaked Whale (Mesoplodon densirostris).Though varying in form, the lower teeth of the different members of this genus agree in their essential structure, having a small and pointed enamel-covered crown, composed of dentine, which, instead of surmounting a root of the ordinary character, is raised upon a solid mass of osteo-dentine, the continuous growth of which greatly alters the form and general appearance of the tooth as age advances, as in the case ofM. layardi, where the long, narrow, flat, strap-like teeth, curving inwards at their extremities, meet over the rostrum, and interfere with the movements of the jaw. In one species (M. grayi) a row of minute, conical, pointed teeth, like those of ordinary Dolphins, 17 to 19 in number, is present even in the adults, on each side of the middle part of the upper jaw, but embedded by their roots only in the gum, and not in bony sockets. This, with the frequent presence of rudimentary teeth in other species of this genus, indicates that the beaked whales are derived from ancestral forms with teeth of normal character in both jaws. The species are distributed in both northern and southern hemispheres, but most frequent in the latter. Among them areM. bidens,M. europaeas,M. densirostris,M. layardi,M. grayiandM. hectori; but there is still much to be learned with regard to their charactersand distribution. This group was abundant in the Pliocene age, as attested by the frequency with which the imperishable long, cylindrical rostrum of the skull, of more than ivory denseness, is found among the rolled and waterworn animal remains which compose the “bone-bed” at the base of the Red Crag of Suffolk.Finally, in Arnoux’s beaked whale (Berardius arnouxi), of New Zealand, which grows to a length of 30 ft., there are two moderate-sized, compressed, pointed teeth, on each side of the symphysis of the lower jaw, with their summits directed forwards, the anterior being the larger of the two and close to the front of the jaw. Upper ends of the premaxillae nearly symmetrical, moderately elevated, slightly expanded, and not curved forward over the nostrils. Nasals broad, massive and rounded, of nearly equal size, forming the vertex of the skull, flattened in front, most prominent in the middle line. Preorbital notch distinct. Rostrum long and narrow. Mesethmoid partially ossified. Small rough eminences on the outer edge of the upper surface of the maxillae at base of rostrum. Vertebrae: C 7, D 10, L 12, Ca 19; total 48. The three anterior cervicals welded, the rest free and well developed. Apparently this whale has the power of thrusting its teeth up and down, exposing them to view when attacked.Fig.6.—The Susu, or Ganges Dolphin (Platanista gangetica).In a family by themselves—thePlatinistidae—are placed three cetaceans which differ from the members of the preceding and the following groups in the mode of articulation of the ribs with the vertebrae, as the tubercular and capitular articulations, distinct at the commencement of the series, gradually blend together, as in most mammals. The cervical vertebrae are all free. The lacrymal bone is not distinct from the jugal. The jaws are long and narrow, with numerous teeth in both; the symphysis of the lower one exceeding half its length. Externally the head is divided from the body by a slightly constricted neck. Pectoral limbs broad and truncated. Dorsal fin small or obsolete. In habits these dolphins are fluviatile or estuarine. In the Indian susu, or Ganges dolphin (Platanista gangetica), the teeth number about30⁄30on each side, are set near together, are rather large, cylindrical, and sharp-pointed in the young, but in old animals acquire a large laterally compressed base, which in the posterior part of the series becomes irregularly divided into roots. As the conical enamel-covered crown wears away, the teeth of the young and old animals have a totally different appearance. The beak and tooth-bearing portion of the lower jaw are so narrow that the teeth of the two sides are almost in contact. Maxillae supporting large, incurved, compressed bony crests, which overarch the nostrils and base of the rostrum, and almost meet in the middle line above. Orbits very small and eyes rudimentary, without crystalline lens. Blow-hole longitudinal, linear. Vertebrae: C 7, D 11, L 8, Ca 25; total 51. A small caecum. No pelvic bones. Dorsal fin represented by a low ridge.Fig.7.—River Plate Dolphin (Stenodelphis blainvillei).The second genus is represented byInia geoffroyi, of the Amazon, in which the teeth vary from 26 to 33 pairs in each jaw; those at the posterior part with a distinct tubercle at the inner side of the base of the crown. Vertebrae: C 7, D 13, L 3, Ca 18; total 41. Transverse processes of lumbar vertebrae very broad. Sternum short and broad, and consisting of a single segment only. Dorsal fin a mere ridge. The long cylindrical rostrum externally furnished with scattered, stout and crisp hairs. The third type isStenodelphis blainvillei, the River Plate dolphin, a small brown species (fig. 7), with from 50 to 60 pairs of teeth in each jaw, furnished with a cingulum at the base of the crown. Jaws very long and slender. Vertebrae: C 7, D 10, L 5, Ca 19; total 41. Transverse processes of the lumbar vertebrae extremely broad. Sternum elongated, composed of two segments, with four sternal ribs attached. Dorsal fin rather small, triangular, pointed. Blow-hole transverse. In several respects this species connects the two preceding ones with theDelphinidae(seeDolphin).Fig.8.—Upper surface of the Skull of male Narwhal (Monodon monoceros), with the whole of both teeth exposed by removal of the upper wall of their alveolar cavities.The last family of existing cetaceans is the above-mentionedDelphinidae, which includes the true dolphins, porpoises, grampuses and their relatives. As a rule there are numerous teeth in both jaws; and the pterygoid bones of the skull are short, thin and involuted to form with a process of the palate bone the outer wall of the post-palatine air-sinus. Symphysis of lower jaw short, or moderate, never exceeding one-third the length of the jaw. Lacrymal bone not distinct from the jugal. Transverse processes of the dorsal vertebrae gradually transferred from the arches to the bodies of the vertebrae without any sudden break, and becoming posteriorly continuous serially with the transverse processes of the lumbar vertebrae. Anterior ribs attached to the transverse process by the tubercle, and to the body of the vertebra by the head; the latter attachment lost in the posterior ribs. Sternal ribs ossified. The blow-hole is transverse, crescentic, with the horns of the crescent pointing forwards.First on the long list is the narwhal,Monodon monoceros, in which, apart from some irregular rudimentary teeth, the dentition is reduced to a single pair of teeth which lie horizontally in the maxilla, and in the female remain permanently concealed within the socket, so that this sex is practically toothless, while in the male (fig. 8), the right tooth usually remains similarly concealed while the left is immensely developed, attaining a length equal to more than half that of the entire animal, projecting horizontally from the head in the form of a cylindrical, or slightly tapering, pointed tusk, without enamel, and with the surface marked by spiral grooves and ridges, running in a sinistral direction. Vertebrae: C 7, D 11, L 6, Ca 26; total 50. Cervical region comparatively long, and all the vertebrae distinct, or with irregular unions towards the middle of the series, the atlas and axis being usually free. Flipper small, short and broad, with the second and third digits nearly equal, the fourth slightly shorter. No dorsal fin. SeeNarwhal.Closely allied is the beluga or white-whale (Delphinapterus leucas), of the Arctic seas, in which, however, there are from eight to ten pairs of teeth in each jaw, occupying the anterior three-fourths of the rostrum and corresponding portion of the lower jaw, rather small, conical, and pointed when unworn, but usually become obliquely truncated, separated by intervals considerably wider than the diameter of the tooth, and implanted obliquely, the crowns inclining forwards especially in the upper jaw. Skull rather narrow and elongated, depressed. Premaxillae convex in front of the nostrils. Rostrum about equal in length to the cranial portion of the skull, triangular, broad at the base, and gradually contracting towards the apex, where it is somewhat curved downwards. Vertebrae: C 7, D 11, L 9, Ca 23; total 50. Cervical vertebrae free. Flippers broad, short and rounded, all the digits being tolerably well developed, except the first. Anterior part of head rounded; no distinct snout. No dorsal fin, but a low ridge in its place. SeeBeluga.In all the remaining genera ofDelphinidaethe cervical region of the vertebral column is very short, and the first two, and usually more, of the vertebrae are firmly united. The common porpoise (Phocaena communis, orP. phocaena) is the typical representative of the first genus, in which the teeth vary from18⁄18to25⁄25, are small, and occupy nearly the whole length of the rostrum, with compressed, spade-shaped crowns, separated from the root by a constricted neck. Rostrum rather shorter than the cranium proper, broad at the base and tapering towards the apex. Premaxillae raised into tuberosities in front of the nostrils. The frontal bones form a somewhat square elevated protuberance in the middle line of the skull behind the nostrils, rising above the flattened nasals. Symphysis of lower jaw very short. Vertebrae: C 7, D 13, L 14, Ca 30; total 64. First to sixth cervical vertebrae and sometimes the seventh also, coalesced. Flippers of moderate size, oval, slightly sickle-shaped, with the second and third digits nearly equal in length, and the fourth and fifth well developed, but shorter. Head short, moderately rounded in front of the blow-hole. Dorsal fin near the middle of the back, triangular; its height considerably less than the length of the base; its anterior edge frequently furnished with one or more rows of conical horny tubercles.The porpoise, which is so common in British waters and the Atlantic, seldom enters the Mediterranean, and apparently never resides there. There is, however, a porpoise in the Black Sea, which, according to Dr O. Abel, is entitled to rank as a distinct species, with the name ofPhocaena relicta. This Black Sea porpoise is readily distinguished from the Atlantic species by the contour of the profile of the head, which, in place of forming a continuous curve from the muzzle to what represents the neck, has a marked prominence above the angle of the mouth, followed by an equally marked depression. The teeth are also different in form and number. The absence of porpoises from the Mediterranean is explained by Dr Abel on account of the greater saltness of that sea as compared with the ocean in general; his idea being that these cetaceans are near akin to fresh-water members of the group, and therefore unsuited to withstand an excessively saline medium. From the Taman Peninsula, on the north shore of the Black Sea, the same writer has described an extinct type of ancestral porpoise, under the name ofPalaeophocaena andrussowi. Another species is the wholly blackP. spinipennis, typically from South America. Black is also the hue of the Indian porpoise (Neophocaena phocaenoides), which wants a dorsal fin, and has eighteen pairs of teeth rather larger than those of the ordinary porpoise. (SeePorpoise.)Fig.9.—Beluga or White-Whale (Delphinapterus leucas). From a specimen taken in the river St Lawrence and exhibited in London, 1877.Next comes the Indo-Malay genusOrcella, in which the12⁄12to14⁄14, small, conical teeth are pointed, rather closely set, and occupy nearly the whole length of the rostrum. Skull sub-globular, high. Rostrum nearly equal in length to the cranial portion of the skull, tapering. Flippers of moderate size, not elongated, but somewhat pointed, with all the bones of the digits broader than long, except the first phalanges of the index and third fingers. Head globular in front. Dorsal fin rather small, placed behind the middle of the body. Two species, both of small size—O. brevirostris, from the Bay of Bengal, andO. fluminalis, from the Irrawaddy river, from 300 to 900 m. from the sea.In the grampus, or killer,Orca gladiator(orO. orca) the teeth form about twenty pairs, above and below, occupying nearly the whole length of the rostrum, very large and stout, with conical recurved crowns and large roots, expanded laterally and flattened, or rather hollowed, on the anterior and posterior surfaces. Rostrum about equal in length to the cranial part of the skull, broad and flattened above, rounded in front; premaxillae broad and rather concave in front of the nostrils, contracted at the middle of the rostrum, and expanding again towards the apex. Vertebrae: C 7, D 11-12, L 10, Ca 23; total 51 or 52; bodies of the first and second and sometimes the third cervical vertebrae united; the rest free. Flippers very large, ovate, nearly as broad as long, with all the phalanges and metacarpals broader than long. General form of body robust. Face short and rounded. Dorsal fin near the middle of the back, very high and pointed. SeeGrampus.Fig.10.—The Grampus or Killer (Orca gladiator).The lesser killer or black killer,Pseudorca crassidens, has its8-12⁄8-12teeth confined to the anterior half of the rostrum and corresponding part of the lower jaw; they are small, conical, curved and sharp-pointed when unworn, but sometimes deciduous in old age. Skull broad and depressed; with the rostrum and cranial portions about equal in length. Upper surface of rostrum broad and flat. Premaxillae concave in front of the nostrils, as wide at the middle of the rostrum as at the base, and nearly or completely concealing the maxillae in the anterior half of this region. Vertebrae: C 7, D II, L 12-14, Ca 28-29; total 58 or 59. Bodies of the anterior five or six cervical vertebrae united. Length of the bodies of the lumbar and anterior caudal vertebrae about equal to their width. Flippers very long and narrow, with the second digit the longest, and having as many as 12 or 13 phalanges, the third shorter (with 9 phalanges), the first, fourth and fifth very short. Fore part of the head round, in consequence of the great development of a cushion of fat, placed on the rostrum of the skull in front of the blow-hole. Dorsal fin low and triangular, the length of its base considerably exceeding its vertical height.Next comes the ca’ing whale, or black-fish (Globicephalus melas), with about ten pairs of upper and lower teeth. Cranial and dental characters generally like those ofOrca, except that the roots of the teeth are cylindrical. Vertebrae: C 7, D 10, L 9, Ca 24; total 50; first to sixth or seventh cervical vertebrae united; bodies of the lumbar vertebrae distinguished from those of the preceding genera by being more elongated, the length being to the width as 3 to 2. Flippers of moderate size, narrow and pointed. Dorsal fin situated near the middle of the back, of moderate size, and sickle-shaped. Head in front of the blow-hole high, and compressed anteriorly, the snout truncated. SeeCa’ing Whale.Risso’s dolphin,Grampus griseus, represents another genus, characterized by the absence of teeth in the upper and the small number of these in the lower jaw (3 to 7 on each side, and confined to the region of the symphysis). Vertebrae: C 7, D 12, L 19, Ca 30; total 68. General external characters much as inGlobicephalus, but the fore part of the head less rounded, and the flippers less elongated.G. griseusis about 13 ft. long, and remarkable for its great variability of colour. It has been found, though rarely, in the North Atlantic and Mediterranean.The common dolphin (Delphinus delphis) is the typical representative of a large group of relatively small species, some of which are wholly marine, while others are more or less completely fluviatile. They are divided into a number of genera, such asProdelphinus,Steno,Lagenorhynchus,Cephalorhynchus,Tursiops, &c., best distinguished from one another by the number and size of the teeth, the form and relations of the bones on the hinder part of the palate, the length of the beak and of the union of the two halves of the lower jaw, and the number of vertebrae. For the distinctive characters of these genera the reader may refer to one of the works mentioned below; and it must suffice to state that, collectively, all these dolphins are characterized by the following features. The teeth are numerous in both jaws, and more than20⁄20in number, occupying nearly the whole length of the rostrum, and small, close-set, conical, pointed and slightly curved. Rostrum more or less elongated, and pointed in front, usually considerably longer than the cranial portion of the skull. Vertebrae: C 7, D 12-14, L and Ca variable; total 51 to 90. Flippers of moderate size, narrow, pointed, somewhat sickle-shaped, with the first digit rudimentary, the second longest, third nearly equal, and the fourth and fifth extremely short. Externally the head shows a distinct beak or pointed snout, marked off from the antenasal fatty elevation by a V-shaped groove. Dorsal fin rather large, triangular or sickle-shaped, rarely wanting. A curiously marked brown and white species, perhaps referable toLagenorhynchusis found on the fringe of the Antarctic ice (see report on the zoology of the “Discovery,” published in 1907 by the British Museum). SeeDolphin.Extinct Cetacea.At present we are totally in the dark as to the origin of the whalebone-whales, not being even assured that they are derived from the same stock as the toothed whales. It is noteworthy, however, that some of the fossil representatives of the latter have nasal bones of a type recalling those of the former. Such fossil whalebone-whales as are known occur in Pliocene, and Miocene formations are either referable to existing genera, or to more or less nearly related extinct ones, such asPlesiocetus,HerpetocetusandCetotherium.The toothed whales, on the other hand, are very largely represented in a fossil state, reaching as low in the geological series as the upper Cretaceous. Many of these present much more generalized characters than their modern representatives, while others indicate apparently a transition towards the still more primitive zeuglodonts, which, as will be shown later, are themselves derived from the creodont Carnivora. In the Pliocene deposits of Belgium and England are preserved the teeth and other remains of a number of cetaceans, such asPhysodon,Encetus,Dinoziphius,Hoplocetus,BalaenodonandScaldicetus, more or less nearly related to the sperm-whale, but presenting several primitive characters. A complete skull of a member of this group from the Tertiary deposits of Patagonia, at first referred toPhysodon, but subsequently toScaldicetus, has a full series of enamelled teeth in the upper jaw; and it is probable that the same was the case in other forms. This entails either a modification of the definition of thePhyseteridaeas given above, or the creation of a separate family for these primitive sperm-whales. In other cases, however, as in the MioceneProphyseterandPlacoziphius, the anterior portion or the whole of the upper jaw had already become toothless; and these forms are regarded as indicating the descent of the sperm-whales from the under-mentionedSqualodon. The beaked whales, again, are believed to be independently descended from the latter type,Berardiusbeing traced into the MioceneMioziphius,AnoplonassaandPalaeoziphius, the last of which shows signs in its dentition of approximating to the complicated tooth-structure of the squalodonts.Another line of descent from the latter, apparently culminating in the modernPlatanistidae, is represented by the familyEurhinodelphidae, typified by the European MioceneEurhinodelphis, but also including the contemporary PatagonianArgyrocetusand the nearly allied EuropeanCyrtodelphis. All these were very long-beaked dolphins; and inArgyrocetus, at all events, the occipital condyles, instead of being closely pressed to the skull, are as prominent as in ordinary mammals, while the nasal bones, instead of forming mere rudimentary nodules, were squared and roofed over the hind part of the nasal chamber.In the MioceneSqualodon, representing the familySqualodontidae, the dentition is differentiated into incisors, canines and cheek-teeth, the hinder ones of the latter series having double roots and compressed crowns carrying serrations on the hinder edge; generally the dental formula has been given as i.3⁄3, c.1⁄1, p.4⁄4, m.7⁄7, the single-rooted cheek-teeth being regarded as premolars and those with double roots as molars. Dr Abel is, however, of opinion that the formula is better represented as i.3⁄3, c.1⁄1, p.8 or 9⁄9, m.3⁄2; the teeth reckoned as molars corresponding to those of the creodont Carnivora. The single-rooted cheek-teeth are regarded as due, not to the division of double-rooted ones, but to the fusion of the two roots of teeth of the latter type. InSqualodonthe nasal bones were of the modern nodular type, but in the Miocene PatagonianProsqualodonthey partially covered the nasal chamber.At present there is a gap between the most primitive squalodonts and the Eocene zeuglodonts (Zeuglodontidae), which are regarded by Messrs Max Weber, O. Abel and C.W. Andrews as the direct forerunners of the modern-toothed whales, forming the suborderArchaeoceti. It is, however, right to mention that some authorities refuse to admit the relation of the Archaeoceti to the whales.In the typical zeuglodonts the long and flat skull has large temporal fossae, a strong sagittal crest, a long beak formed mainly by the premaxillae (in place of the maxillae, as in modern whales), and long nasal bones covering over the nasal chamber, so that the nostrils opened about half-way down the beak. All the cervical vertebrae were free. Normally the dentition in the typical genusZeuglodon(which is common to the Eocene of North America and Egypt) is i.3⁄3, c.1⁄1, p.4⁄4, m.3⁄3; the cheek-teeth being two-rooted, with compressed pointed crowns, of which the fore-and-aft edges are coarsely serrated. In the EgyptianZeuglodon osiristhe number of the molars is, however, reduced to2⁄3, while some of the earlier cheek-teeth have become single-rooted, as in the squalodonts. The probable transitional form between the latter and the zeuglodonts is the smallMicrozeuglodon caucasicusdescribed by the present writer, from the Caucasus. As regards the origin of the zeuglodonts themselves, remains discovered in the Eocene formations of Egypt indicate a practically complete transition, so far at least as dental characters are concerned, from these whale-like creatures to the creodont Carnivora. In the earliest type,Protocetus, the skull is practically that of a zeuglodont, the snout being in fact more elongated than in some of the earliest representatives of the latter, although the nostrils are placed nearer the tip. The incisors are unknown, but the cheek-teeth are essentially those of a creodont, none of them having acquired the serrated edges distinctive of the typical zeuglodonts; and the hinder premolars and molars retaining the three roots of the creodonts. In the somewhat laterProzeuglodonthe skull is likewise essentially of the zeuglodont type, although the nostrils have shifted a little more backwards; as regards the cheek-teeth, which have acquired serrated crowns, the premolars at any rate retain the inner buttress supported by a distinct third root, so that they are precisely intermediate betweenProtocetusandZeuglodon. Yet another connecting form isEocetus, a very large animal from nearly the same horizon asProzeuglodon; its skull approaching that ofZeuglodonas regards the backward position of the nostrils, although the cheek-teeth are of the creodont type, having inner, or third, roots. It is noteworthy thatZeuglodonapparently occurs in the same beds as these intermediate types.It follows from the foregoing that if zeuglodonts are the ancestors of the true Cetacea—and the probability that they are so is very great—the latter are derived from primitive Carnivora, and not, as has been suggested, from herbivorous Ungulata. The idea that the zeuglodonts were provided with a bony armour does not appear to be supported by recent discoveries.Authorities.—The above article is based on that by Sir W.H. Flower in the 9th edition of this work. See also W.H. Flower, “On the Characters and Divisions of the Family Delphinidae,”Proc. Zool. Soc.(London, 1883); F.W. True, “Review of the Family Delphinidae,”Proc. U.S. Museum, No. 36 (1889); R. Lydekker, “Cetacean Skulls from Patagonia,”Palaeontol. Argentina, vol. ii:An. Mus. La Plata(1893); W. Dames, “Über Zeuglodonten aus Ägypten,”Paläontol. Abhandlungen, vol. i. (1894); F.E. Beddard,A Book of Whales(London, 1900); O. Abel, “Untersuchungen über die fossilen Platanistiden des Wiener Beckens,”Denks. k. Akad. Wiss. Wien., vol. lxviii. (1899); “Les Dauphins longirostres du Bolérien,”Mém. musée d’hist. nat. belgique(1901 and 1902); “Die phylogenetische Entwickelung des Cetaceengebisses und die systematische Stellung der Physeteriden,”Verhandl. deutsch. zool. Gesellschaft(1905); E. Fraas, “Neue Zeuglodonten aus dem unteren Mittelocean vom Mokattam bei Cairo,”Geol. und paläontol. Abhandl.ser. 2, vol. vi. (1904); C.W. Andrews, “Descriptive Catalogue of the Tertiary Vertebrata of the Fayum” (British Museum, 1906).
Whalebone Whales.—Existing Cetacea are divisible into two sections, or suborders, the relationships of which are by no means clearly apparent. The first section is that of the whalebone whales, or Mystacoceti, in which no functional teeth are developed, although there are tooth-germs during foetal life. The palate is furnished with plates of baleen or whalebone; the skull is symmetrical; and the nasal bones form a roof to the nasal passages, which are directed upwards and forwards. The maxilla is produced in front of, but not over, the orbital process of the frontal. The lacrymal is small and distinct from the jugal. The tympanic is welded with the periotic, which is attached to the base of the skull by two strong diverging processes. The olfactory organ is distinctly developed. The two halves of the lower jaw are arched outwards, their anterior ends meeting at an angle, and connected by fibrous tissue without any symphysis. All the ribs at their upper extremity articulate only with the transverse processes of the vertebrae; their capitular processes when present not articulating directly with the bodies of the vertebrae. The sternum is composed of a single piece, and articulates only with a single pair of ribs; and there are no ossified sternal ribs. External openings of nostrils distinct from each other, longitudinal. A short conical caecum.
When in the foetal state these whales have numerous minute teeth lying in the dental groove of both upper and lower jaws. They are best developed about the middle of foetal life, after which they are absorbed, and no trace of them remains at the time of birth. The whalebone does not make its appearance until after birth; and consists of a series of flattened horny plates, between three and four hundred in number, on each side of the palate, with a bare interval along the middle line. The plates are placed transversely to the long axis of the palate, with short intervals between them. Each plate or blade is somewhat triangular in form, with the baseattached to the palate and the apex hanging downwards. The outer edge of the blade is hard and smooth, but the inner edge and apex fray out into long bristly fibres, so that the roof of the whale’s mouth looks as if covered with hair, as described by Aristotle. At the inner edge of each principal blade are two or three much smaller or subsidiary blades. The principal blades are longest near the middle of the series, and gradually diminish towards the front and back of the mouth. The horny plates grow from a fibrous and vascular matrix, which covers the palatal surface of the maxillae, and sends out plate-like processes, one of which penetrates the base of each blade. Moreover, the free edges of these processes are covered with long vascular thread-like papillae, one of which forms the central axis of each of the hair-like fibres mainly composing the blade. A transverse section of fresh whalebone shows that it is made up of numbers of these soft vascular papillae, circular in outline, and surrounded by concentrically arranged epidermic cells, the whole bound together by other epidermic cells, that constitute the smooth (so-called “enamel”) surface of the blade, which, disintegrating at the free edge, allows the individual fibres to become loose and assume a hair-like appearance.
Whalebone really consists of modified papillae of the mucous membrane of the mouth, with an excessive and horny epithelial development. The blades are supported and bound together for a certain distance from their base, by a mass of less hardened epithelium, secreted by the surface of the palatal membrane or matrix of the whalebone in the intervals of the plate-like processes. This is the “gum” of the whalers. Whalebone varies much in colour in different species; in some it is almost jet black, in others slate colour, horn colour, yellow, or even creamy-white. In some descriptions the blades are variegated with longitudinal stripes of different hues. It differs also greatly in other respects, being short, thick, coarse, and stiff in some cases, and greatly elongated and highly elastic in those species in which it has attained its fullest development. Its function is to strain the water from the small marine molluscs, crustaceans, or fish upon which the whales subsist. In feeding, whales fill the immense mouth with water containing shoals of these small creatures, and then, on closing the jaws and raising the tongue, so as to diminish the cavity of the mouth, the water streams out through the narrow intervals between the hairy fringe of the whalebone blades, and escapes through the lips, leaving the living prey to be swallowed.
Although sometimes divided into two families,BalaenidaeandBalaenopteridae, whalebone-whales are best included in a single family group under the former name. The typical members of this family are the so-called right-whales, forming the genusBalaena, in which there are no folds on the throat and chest, and no back-fin; while the cervical vertebrae are fused into a single mass. The flippers are short and broad, with five digits; the head is very large and the whalebone very long and narrow, highly elastic and black; while the scapula is high, with a distinct coracoid and coronoid process. This genus contains the well-known Greenland right-whale (B. mysticetus) of the Arctic seas, the whalebone and oil of which are so much valued in commerce, and also other whales, distinguished by having the head somewhat smaller in proportion to the body, with shorter whalebone and a larger number of vertebrae. These inhabit the temperate seas of both northern and southern hemispheres, and have been divided into species in accordance with their geographical distribution, such asB. biscayensisof the North Atlantic,B. japonicaof the North Pacific,B. australisof the South Atlantic, andB. antipodarumandnovae-zelandiaeof the South Pacific; but the differences between them are so small that they may probably be regarded as races of a single species, the black whale (B. australis). On the head these whales carry a peculiar structure which is known to whalers as the “bonnet.” This is a large horny excrescence, worn into hollows like a much-denuded piece of limestone rock, growing probably in the neighbourhood of the blow-hole. More than one theory has been suggested to account for its presence. One suggestion is that it indicates the descent of whales from rhinoceros-like mammals; another that this species of whale is in the habit of rubbing against rocks in order to free itself from barnacles, and thus produces a kind of corn—although why on the nose alone is not stated. Dr W.G. Ridewood, however, considers that the structure is due to the fact that the horny layers which are produced all over the skin are not shed on this particular spot.
The pigmy whale (Neobalaena marginata) represents a genus agreeing with the right-whales in the absence of throat-flutings, and with the rorquals in the presence of a dorsal fin. The cervical vertebrae are united, and there are only 43 vertebrae altogether. The flippers are small, narrow, and with only four digits. The ribs remarkably expanded and flattened; the scapula low and broad, with completely developed acromion and coracoid processes. The whalebone is long, slender, elastic and white. The species which inhabits the South American, Australian and New Zealand seas is the smallest of the whalebone-whales, being not more than 20 ft. in length.
In contrast to the preceding is the great grey whale (Rachianectes glaucus) of the North Pacific, which combines the relatively small head, elongated shape, and narrow flippers of the fin-whales, with the smooth throat and absence of a back-fin distinctive of the right-whales. The whalebone is shorter and coarser than in any other species. In the skeleton the cervical vertebrae are free, and the first two ribs on each side expanded and united to form a large bony shield. In the humpback-whale (Megaptera longimanaorboops) the head is of moderate size, the whalebone-plates are short and wide, and the cervical vertebrae free. The skin of the throat is fluted so as to form an expansible pouch; there is a low back-fin; and the flippers, which have four digits each, are extremely long, equalling about one-fourth the total length of the animal. The acromion and coracoid processes of the scapula are rudimentary. SeeHumpback-Whale.
The right-whales are built for cruising slowly about in search of the shoals of small floating invertebrates which form their food, and are consequently broad in beam, with a float-shaped body and immovable neck. The humpback is of somewhat similar build, but with a smaller head, and probably attains considerable speed owing to the length of its flippers. The finners, or rorquals (Balaenoptera), which prey largely on fish, are built entirely for speed, and are the ocean greyhounds of the group. Their bodies are consequently long and attenuated, and their necks are partially mobile; while they are furnished with capacious pouches for storing their food. They chiefly differ from the humpback by the smaller head, long and slender build, small, narrow, and pointed flippers, each containing four digits, and the large acromion and coracoid processes to the low and broad scapula. Rorquals are found in almost every sea. Among them are the most gigantic of all animals,B. sibbaldi, which attains the length of 80 ft., and the smallB. rostrata, which does not exceed 30. There are certainly four distinct modifications of this genus, represented by the two just mentioned, and byB. musculusandB. borealis, all inhabitants of British seas, but the question whether almost identical forms found in the Indian, Southern and Pacific Oceans are to be regarded as specifically identical or as distinct awaits future researches, although some of these have already received distinct names. SeeRorqual.
In the report on the zoology of the “Discovery” expedition, published in 1907 by the British Museum, E.A. Wilson describes a whale frequenting the fringe of the Antarctic ice which indicates a new generic type. Mainly black in colour, these whales measure about 20 or 30 ft. in length, and have a tall dorsal fin like that of a killer.
Toothed Whales.—-The second suborder is represented by the toothed whales, or Odontoceti, in which there is no whalebone, and teeth, generally numerous, though sometimes reduced to a single pair, and occasionally wanting, are normally developed. Unlike that of the whalebone-whales, the upper surface of the skull is more or less unsymmetrical. The nasal bones are in the form of nodules or flattened plates, applied closely to the frontals, and not forming any part of the roof to the nasal passage, which is directed upwards and backwards. The olfactory organ is rudimentary or absent. Hinder end of the maxilla expanded and covering the greater part of the orbital plate of the frontal bone. Lacrymal bone either inseparable from the jugal, or, if distinct, large, and forming part of the roof of the orbit. Tympanic bone not welded with the periotic, which is usually only attached to the rest of the skull by ligament. Two halves of the lower jaw nearly straight, expanded in height posteriorly, with a wide funnel-shaped aperture to the dental canal, and coming in contact in front by a flat surface of variable length, but constituting a symphysis. Several of the anterior ribs with well-developed capitular processes, which articulate with the bodies of the vertebrae. Sternum almost always composed of several pieces, placed one behind the other, with which several pairs of ribs are connected by well-developed cartilaginous or ossified sternal ribs. External respiratory aperture single, the two nostrils uniting before they reach the surface, usually in the form of a transverse sub-crescentic valvular aperture, situated on the top of the head. Flippers with five digits, though the first and fifth are usually little developed. No caecum, except inPlatanista.
The first family,Physeteridae, is typified by the sperm-whale, and characterized by the absence of functional teeth in the upper jaw; the lower teeth being various, and often much reduced in number. Bones of the skull raised so as to form an elevated prominence or crest behind the nostrils. Pterygoid bones thick, produced backwards, meeting in the middle line, and not involuted to form the outer wall of the post-palatine air-sinuses, but simply hollowed on their outer side. Transverse processes of the arches of the dorsal vertebrae, to which the tubercles of the ribs are attached, ceasing abruptly near the end of the series, and replaced by processes on the body at a lower level, and serially homologous anteriorly with the heads of the ribs, and posteriorly with the transverse processes of the lumbar vertebrae. Costal cartilages not ossified.
The first group, orPhyseterinae, includes the sperm-whale itself and is characterized by the presence of a full series of lower teeth, which are set in a groove in place of sockets, the groove being imperfectly divided by partial septa, and the teeth held in place by the strong, fibrous gum. No distinct lacrymal bone. Skull strikingly asymmetrical in the region of the nasal apertures, in consequence of the left opening greatly exceeding the right in size.
In the sperm-whale (Physeter macrocephalus) the upper teeth are apparently of uncertain number, rudimentary and functionless, being embedded in the gum. Lower jaw with from 20 to 25 teeth on each side, stout, conical, recurved and pointed at the apexuntil they are worn, without enamel. Upper surface of the skull concave; its posterior and lateral edges raised into a very high and greatly compressed semicircular crest or wall (fig. 2). Zygomatic processes of jugal bones thick and massive. Muzzle greatly elongated, broad at the base, and gradually tapering to the apex. Lower jaw exceedingly long and narrow, the symphysis being more than half the length. Vertebrae: C 7, D 11, L 8, Ca 24; total 50. Atlas, or first vertebra, free; all the other cervical vertebrae united by their bodies and spines into a single mass. Eleventh pair of ribs rudimentary. Head about one-third the length of the body; very massive, high and truncated, and rather compressed in front; owing its huge size and form mainly to the accumulation of a mass of fatty tissue filling the large hollow on the upper surface of the skull and overlying the long muzzle. The single blow-hole is longitudinal, slightly S-shaped, and placed at the upper and anterior extremity of the head to the left side of the middle line. The opening of the mouth is on the under side of the head, considerably behind the end of the snout. Flippers short, broad and truncated. Dorsal fin represented by a low protuberance. SeeSperm-Whale.
In the lesser or pigmy sperm-whale (Cogia breviceps) there may be a pair of rudimentary teeth in the upper jaw, while on each side of the lower jaw there are from 9 to 12 rather long, slender, pointed and curved teeth, with a coating of enamel. Upper surface of the skull concave, with thick, raised, posterior and lateral margins, massive and rounded at their anterior terminations above the orbits. Muzzle not longer than the cranial position of the skull, broad at the base, and rapidly tapering to the apex. Zygomatic process of the jugal rod-like. Lower jaw with symphysis less than half its length. Vertebrae: C 7, D 13 or 14, L and Ca 30; total 50 or 51. All the cervical vertebrae united by their bodies and arches. The head is about one-sixth of the length of the body, and obtusely pointed in front; the mouth small and placed far below the apex of the snout; the blow-hole crescentic, and placed obliquely on the crown of the head in advance of the eyes and to the left of the middle line; while the flippers are bluntly sickle-shaped, and the back-fin triangular. This species attains a length of from 9 to 13 ft.
A second subfamily is represented by the bottle-noses and beaked whales, and known as theZiphiinae. In this group the lower teeth are rudimentary and concealed in the gum, except one, or rarely two, pairs which may be largely developed, especially in the male. There is a distinct lacrymal bone. Externally the mouth is produced into a slender rostrum or beak, from above which the rounded eminence formed by a cushion of fat resting on the cranium in front of the blow-hole rises somewhat abruptly. The blow-hole is single, crescentic and median, as in theDelphinidae. Flippers small, ovate, with five digits moderately well developed. A small obtuse dorsal fin situated considerably behind the middle of the back. Longitudinal grooves on each side of the skin of the throat, diverging posteriorly, and nearly meeting in front. In external characters and habits the whales of this group closely resemble each other. They appear to be almost exclusively feeders on cuttle-fishes, and occur either singly, in pairs, or in small herds. By their dental and osteological characters they are easily separated into four genera.
In the first of these,Hyperoödon, or bottle-nose, there is a small conical pointed tooth at the apex of each half of the lower jaw, concealed by the gum during life. Skull with the upper ends of the premaxillae rising suddenly behind the nostrils to the vertex and expanded laterally, their outer edges curving backwards and their anterior surfaces arching forwards and overhanging the nostrils; the right larger than the left. Nasal bones lying in the hollow between the upper extremities of the premaxillae, strongly concave in the middle line and in front; their outer edges, especially that of the right, expanded over the front of the inner border of the maxilla. Very high longitudinal crests on the maxillae at the base of the beak, extending backwards almost to the nostrils, approaching each other in the middle line above; sometimes compressed and sometimes so massive that their inner edges come almost in contact. Preorbital notch distinct, and mesethmoid cartilage slightly ossified. Vertebrae: C 7, D 9, L 10, Ca 19; total 45. All the cervical vertebrae united. Upper surface of the head in front of the blow-hole very prominent and rounded, rising abruptly from above the small, distinct snout. Two species are known. SeeBottle-nose Whale.
The typical representative of the beaked whales isZiphius cuvieri, in which there is a single conical tooth of moderate size on each side close to the anterior extremity of the lower jaw, directed forwards and upwards. Skull with the premaxillae immediately in front and at the sides of the nostrils expanded, hollowed, with elevated lateral margins, the posterior ends rising to the vertex and curving forwards, the right being considerably more developed than the left. The conjoint nasals form a pronounced symmetrical eminence at the top of the skull, projecting forwards over the nostrils, flat above, prominent and rounded in the middle line in front, and separated by a notch on each side from the premaxillae. Preorbital notch not distinct. Rostrum (seen from above) triangular, tapering from the base to the apex; upper and outer edges of maxillae at base of rostrum raised into low roughened tuberosities. Mesethmoid cartilage densely ossified in adult age, and coalescing with the surrounding bones of the rostrum. Vertebrae: C 7, D 10, L 10, Ca 22; total 49. The three anterior cervical vertebrae united, the rest free.
In the numerous species of the allied genusMesoplodonthere is a much-compressed and pointed tooth in each half of the lower jaw, variously situated, but generally at some distance behind the apex; its point directed upwards, and often somewhat backwards, occasionally developed to a great size. In the skull the region round the nostrils is as inHyperoödon, except that the nasals are narrow and more sunk between the upper ends of the premaxillae; like those ofHyperoödon, they are concave in the middle line in front and above. No maxillary tuberosities. Preorbital notch not very distinct. Rostrum long and narrow. Mesethmoid in the adult ossified in its entire length, and coalescing with the surrounding bones. Vertebrae: C 7, D 10, L 10 or 11, Ca 19 or 20; total 46 to 48. Two or three anterior cervicals united, the rest usually free.
Though varying in form, the lower teeth of the different members of this genus agree in their essential structure, having a small and pointed enamel-covered crown, composed of dentine, which, instead of surmounting a root of the ordinary character, is raised upon a solid mass of osteo-dentine, the continuous growth of which greatly alters the form and general appearance of the tooth as age advances, as in the case ofM. layardi, where the long, narrow, flat, strap-like teeth, curving inwards at their extremities, meet over the rostrum, and interfere with the movements of the jaw. In one species (M. grayi) a row of minute, conical, pointed teeth, like those of ordinary Dolphins, 17 to 19 in number, is present even in the adults, on each side of the middle part of the upper jaw, but embedded by their roots only in the gum, and not in bony sockets. This, with the frequent presence of rudimentary teeth in other species of this genus, indicates that the beaked whales are derived from ancestral forms with teeth of normal character in both jaws. The species are distributed in both northern and southern hemispheres, but most frequent in the latter. Among them areM. bidens,M. europaeas,M. densirostris,M. layardi,M. grayiandM. hectori; but there is still much to be learned with regard to their charactersand distribution. This group was abundant in the Pliocene age, as attested by the frequency with which the imperishable long, cylindrical rostrum of the skull, of more than ivory denseness, is found among the rolled and waterworn animal remains which compose the “bone-bed” at the base of the Red Crag of Suffolk.
Finally, in Arnoux’s beaked whale (Berardius arnouxi), of New Zealand, which grows to a length of 30 ft., there are two moderate-sized, compressed, pointed teeth, on each side of the symphysis of the lower jaw, with their summits directed forwards, the anterior being the larger of the two and close to the front of the jaw. Upper ends of the premaxillae nearly symmetrical, moderately elevated, slightly expanded, and not curved forward over the nostrils. Nasals broad, massive and rounded, of nearly equal size, forming the vertex of the skull, flattened in front, most prominent in the middle line. Preorbital notch distinct. Rostrum long and narrow. Mesethmoid partially ossified. Small rough eminences on the outer edge of the upper surface of the maxillae at base of rostrum. Vertebrae: C 7, D 10, L 12, Ca 19; total 48. The three anterior cervicals welded, the rest free and well developed. Apparently this whale has the power of thrusting its teeth up and down, exposing them to view when attacked.
In a family by themselves—thePlatinistidae—are placed three cetaceans which differ from the members of the preceding and the following groups in the mode of articulation of the ribs with the vertebrae, as the tubercular and capitular articulations, distinct at the commencement of the series, gradually blend together, as in most mammals. The cervical vertebrae are all free. The lacrymal bone is not distinct from the jugal. The jaws are long and narrow, with numerous teeth in both; the symphysis of the lower one exceeding half its length. Externally the head is divided from the body by a slightly constricted neck. Pectoral limbs broad and truncated. Dorsal fin small or obsolete. In habits these dolphins are fluviatile or estuarine. In the Indian susu, or Ganges dolphin (Platanista gangetica), the teeth number about30⁄30on each side, are set near together, are rather large, cylindrical, and sharp-pointed in the young, but in old animals acquire a large laterally compressed base, which in the posterior part of the series becomes irregularly divided into roots. As the conical enamel-covered crown wears away, the teeth of the young and old animals have a totally different appearance. The beak and tooth-bearing portion of the lower jaw are so narrow that the teeth of the two sides are almost in contact. Maxillae supporting large, incurved, compressed bony crests, which overarch the nostrils and base of the rostrum, and almost meet in the middle line above. Orbits very small and eyes rudimentary, without crystalline lens. Blow-hole longitudinal, linear. Vertebrae: C 7, D 11, L 8, Ca 25; total 51. A small caecum. No pelvic bones. Dorsal fin represented by a low ridge.
The second genus is represented byInia geoffroyi, of the Amazon, in which the teeth vary from 26 to 33 pairs in each jaw; those at the posterior part with a distinct tubercle at the inner side of the base of the crown. Vertebrae: C 7, D 13, L 3, Ca 18; total 41. Transverse processes of lumbar vertebrae very broad. Sternum short and broad, and consisting of a single segment only. Dorsal fin a mere ridge. The long cylindrical rostrum externally furnished with scattered, stout and crisp hairs. The third type isStenodelphis blainvillei, the River Plate dolphin, a small brown species (fig. 7), with from 50 to 60 pairs of teeth in each jaw, furnished with a cingulum at the base of the crown. Jaws very long and slender. Vertebrae: C 7, D 10, L 5, Ca 19; total 41. Transverse processes of the lumbar vertebrae extremely broad. Sternum elongated, composed of two segments, with four sternal ribs attached. Dorsal fin rather small, triangular, pointed. Blow-hole transverse. In several respects this species connects the two preceding ones with theDelphinidae(seeDolphin).
The last family of existing cetaceans is the above-mentionedDelphinidae, which includes the true dolphins, porpoises, grampuses and their relatives. As a rule there are numerous teeth in both jaws; and the pterygoid bones of the skull are short, thin and involuted to form with a process of the palate bone the outer wall of the post-palatine air-sinus. Symphysis of lower jaw short, or moderate, never exceeding one-third the length of the jaw. Lacrymal bone not distinct from the jugal. Transverse processes of the dorsal vertebrae gradually transferred from the arches to the bodies of the vertebrae without any sudden break, and becoming posteriorly continuous serially with the transverse processes of the lumbar vertebrae. Anterior ribs attached to the transverse process by the tubercle, and to the body of the vertebra by the head; the latter attachment lost in the posterior ribs. Sternal ribs ossified. The blow-hole is transverse, crescentic, with the horns of the crescent pointing forwards.
First on the long list is the narwhal,Monodon monoceros, in which, apart from some irregular rudimentary teeth, the dentition is reduced to a single pair of teeth which lie horizontally in the maxilla, and in the female remain permanently concealed within the socket, so that this sex is practically toothless, while in the male (fig. 8), the right tooth usually remains similarly concealed while the left is immensely developed, attaining a length equal to more than half that of the entire animal, projecting horizontally from the head in the form of a cylindrical, or slightly tapering, pointed tusk, without enamel, and with the surface marked by spiral grooves and ridges, running in a sinistral direction. Vertebrae: C 7, D 11, L 6, Ca 26; total 50. Cervical region comparatively long, and all the vertebrae distinct, or with irregular unions towards the middle of the series, the atlas and axis being usually free. Flipper small, short and broad, with the second and third digits nearly equal, the fourth slightly shorter. No dorsal fin. SeeNarwhal.
Closely allied is the beluga or white-whale (Delphinapterus leucas), of the Arctic seas, in which, however, there are from eight to ten pairs of teeth in each jaw, occupying the anterior three-fourths of the rostrum and corresponding portion of the lower jaw, rather small, conical, and pointed when unworn, but usually become obliquely truncated, separated by intervals considerably wider than the diameter of the tooth, and implanted obliquely, the crowns inclining forwards especially in the upper jaw. Skull rather narrow and elongated, depressed. Premaxillae convex in front of the nostrils. Rostrum about equal in length to the cranial portion of the skull, triangular, broad at the base, and gradually contracting towards the apex, where it is somewhat curved downwards. Vertebrae: C 7, D 11, L 9, Ca 23; total 50. Cervical vertebrae free. Flippers broad, short and rounded, all the digits being tolerably well developed, except the first. Anterior part of head rounded; no distinct snout. No dorsal fin, but a low ridge in its place. SeeBeluga.
In all the remaining genera ofDelphinidaethe cervical region of the vertebral column is very short, and the first two, and usually more, of the vertebrae are firmly united. The common porpoise (Phocaena communis, orP. phocaena) is the typical representative of the first genus, in which the teeth vary from18⁄18to25⁄25, are small, and occupy nearly the whole length of the rostrum, with compressed, spade-shaped crowns, separated from the root by a constricted neck. Rostrum rather shorter than the cranium proper, broad at the base and tapering towards the apex. Premaxillae raised into tuberosities in front of the nostrils. The frontal bones form a somewhat square elevated protuberance in the middle line of the skull behind the nostrils, rising above the flattened nasals. Symphysis of lower jaw very short. Vertebrae: C 7, D 13, L 14, Ca 30; total 64. First to sixth cervical vertebrae and sometimes the seventh also, coalesced. Flippers of moderate size, oval, slightly sickle-shaped, with the second and third digits nearly equal in length, and the fourth and fifth well developed, but shorter. Head short, moderately rounded in front of the blow-hole. Dorsal fin near the middle of the back, triangular; its height considerably less than the length of the base; its anterior edge frequently furnished with one or more rows of conical horny tubercles.
The porpoise, which is so common in British waters and the Atlantic, seldom enters the Mediterranean, and apparently never resides there. There is, however, a porpoise in the Black Sea, which, according to Dr O. Abel, is entitled to rank as a distinct species, with the name ofPhocaena relicta. This Black Sea porpoise is readily distinguished from the Atlantic species by the contour of the profile of the head, which, in place of forming a continuous curve from the muzzle to what represents the neck, has a marked prominence above the angle of the mouth, followed by an equally marked depression. The teeth are also different in form and number. The absence of porpoises from the Mediterranean is explained by Dr Abel on account of the greater saltness of that sea as compared with the ocean in general; his idea being that these cetaceans are near akin to fresh-water members of the group, and therefore unsuited to withstand an excessively saline medium. From the Taman Peninsula, on the north shore of the Black Sea, the same writer has described an extinct type of ancestral porpoise, under the name ofPalaeophocaena andrussowi. Another species is the wholly blackP. spinipennis, typically from South America. Black is also the hue of the Indian porpoise (Neophocaena phocaenoides), which wants a dorsal fin, and has eighteen pairs of teeth rather larger than those of the ordinary porpoise. (SeePorpoise.)
Next comes the Indo-Malay genusOrcella, in which the12⁄12to14⁄14, small, conical teeth are pointed, rather closely set, and occupy nearly the whole length of the rostrum. Skull sub-globular, high. Rostrum nearly equal in length to the cranial portion of the skull, tapering. Flippers of moderate size, not elongated, but somewhat pointed, with all the bones of the digits broader than long, except the first phalanges of the index and third fingers. Head globular in front. Dorsal fin rather small, placed behind the middle of the body. Two species, both of small size—O. brevirostris, from the Bay of Bengal, andO. fluminalis, from the Irrawaddy river, from 300 to 900 m. from the sea.
In the grampus, or killer,Orca gladiator(orO. orca) the teeth form about twenty pairs, above and below, occupying nearly the whole length of the rostrum, very large and stout, with conical recurved crowns and large roots, expanded laterally and flattened, or rather hollowed, on the anterior and posterior surfaces. Rostrum about equal in length to the cranial part of the skull, broad and flattened above, rounded in front; premaxillae broad and rather concave in front of the nostrils, contracted at the middle of the rostrum, and expanding again towards the apex. Vertebrae: C 7, D 11-12, L 10, Ca 23; total 51 or 52; bodies of the first and second and sometimes the third cervical vertebrae united; the rest free. Flippers very large, ovate, nearly as broad as long, with all the phalanges and metacarpals broader than long. General form of body robust. Face short and rounded. Dorsal fin near the middle of the back, very high and pointed. SeeGrampus.
The lesser killer or black killer,Pseudorca crassidens, has its8-12⁄8-12teeth confined to the anterior half of the rostrum and corresponding part of the lower jaw; they are small, conical, curved and sharp-pointed when unworn, but sometimes deciduous in old age. Skull broad and depressed; with the rostrum and cranial portions about equal in length. Upper surface of rostrum broad and flat. Premaxillae concave in front of the nostrils, as wide at the middle of the rostrum as at the base, and nearly or completely concealing the maxillae in the anterior half of this region. Vertebrae: C 7, D II, L 12-14, Ca 28-29; total 58 or 59. Bodies of the anterior five or six cervical vertebrae united. Length of the bodies of the lumbar and anterior caudal vertebrae about equal to their width. Flippers very long and narrow, with the second digit the longest, and having as many as 12 or 13 phalanges, the third shorter (with 9 phalanges), the first, fourth and fifth very short. Fore part of the head round, in consequence of the great development of a cushion of fat, placed on the rostrum of the skull in front of the blow-hole. Dorsal fin low and triangular, the length of its base considerably exceeding its vertical height.
Next comes the ca’ing whale, or black-fish (Globicephalus melas), with about ten pairs of upper and lower teeth. Cranial and dental characters generally like those ofOrca, except that the roots of the teeth are cylindrical. Vertebrae: C 7, D 10, L 9, Ca 24; total 50; first to sixth or seventh cervical vertebrae united; bodies of the lumbar vertebrae distinguished from those of the preceding genera by being more elongated, the length being to the width as 3 to 2. Flippers of moderate size, narrow and pointed. Dorsal fin situated near the middle of the back, of moderate size, and sickle-shaped. Head in front of the blow-hole high, and compressed anteriorly, the snout truncated. SeeCa’ing Whale.
Risso’s dolphin,Grampus griseus, represents another genus, characterized by the absence of teeth in the upper and the small number of these in the lower jaw (3 to 7 on each side, and confined to the region of the symphysis). Vertebrae: C 7, D 12, L 19, Ca 30; total 68. General external characters much as inGlobicephalus, but the fore part of the head less rounded, and the flippers less elongated.G. griseusis about 13 ft. long, and remarkable for its great variability of colour. It has been found, though rarely, in the North Atlantic and Mediterranean.
The common dolphin (Delphinus delphis) is the typical representative of a large group of relatively small species, some of which are wholly marine, while others are more or less completely fluviatile. They are divided into a number of genera, such asProdelphinus,Steno,Lagenorhynchus,Cephalorhynchus,Tursiops, &c., best distinguished from one another by the number and size of the teeth, the form and relations of the bones on the hinder part of the palate, the length of the beak and of the union of the two halves of the lower jaw, and the number of vertebrae. For the distinctive characters of these genera the reader may refer to one of the works mentioned below; and it must suffice to state that, collectively, all these dolphins are characterized by the following features. The teeth are numerous in both jaws, and more than20⁄20in number, occupying nearly the whole length of the rostrum, and small, close-set, conical, pointed and slightly curved. Rostrum more or less elongated, and pointed in front, usually considerably longer than the cranial portion of the skull. Vertebrae: C 7, D 12-14, L and Ca variable; total 51 to 90. Flippers of moderate size, narrow, pointed, somewhat sickle-shaped, with the first digit rudimentary, the second longest, third nearly equal, and the fourth and fifth extremely short. Externally the head shows a distinct beak or pointed snout, marked off from the antenasal fatty elevation by a V-shaped groove. Dorsal fin rather large, triangular or sickle-shaped, rarely wanting. A curiously marked brown and white species, perhaps referable toLagenorhynchusis found on the fringe of the Antarctic ice (see report on the zoology of the “Discovery,” published in 1907 by the British Museum). SeeDolphin.
Extinct Cetacea.
At present we are totally in the dark as to the origin of the whalebone-whales, not being even assured that they are derived from the same stock as the toothed whales. It is noteworthy, however, that some of the fossil representatives of the latter have nasal bones of a type recalling those of the former. Such fossil whalebone-whales as are known occur in Pliocene, and Miocene formations are either referable to existing genera, or to more or less nearly related extinct ones, such asPlesiocetus,HerpetocetusandCetotherium.
The toothed whales, on the other hand, are very largely represented in a fossil state, reaching as low in the geological series as the upper Cretaceous. Many of these present much more generalized characters than their modern representatives, while others indicate apparently a transition towards the still more primitive zeuglodonts, which, as will be shown later, are themselves derived from the creodont Carnivora. In the Pliocene deposits of Belgium and England are preserved the teeth and other remains of a number of cetaceans, such asPhysodon,Encetus,Dinoziphius,Hoplocetus,BalaenodonandScaldicetus, more or less nearly related to the sperm-whale, but presenting several primitive characters. A complete skull of a member of this group from the Tertiary deposits of Patagonia, at first referred toPhysodon, but subsequently toScaldicetus, has a full series of enamelled teeth in the upper jaw; and it is probable that the same was the case in other forms. This entails either a modification of the definition of thePhyseteridaeas given above, or the creation of a separate family for these primitive sperm-whales. In other cases, however, as in the MioceneProphyseterandPlacoziphius, the anterior portion or the whole of the upper jaw had already become toothless; and these forms are regarded as indicating the descent of the sperm-whales from the under-mentionedSqualodon. The beaked whales, again, are believed to be independently descended from the latter type,Berardiusbeing traced into the MioceneMioziphius,AnoplonassaandPalaeoziphius, the last of which shows signs in its dentition of approximating to the complicated tooth-structure of the squalodonts.
Another line of descent from the latter, apparently culminating in the modernPlatanistidae, is represented by the familyEurhinodelphidae, typified by the European MioceneEurhinodelphis, but also including the contemporary PatagonianArgyrocetusand the nearly allied EuropeanCyrtodelphis. All these were very long-beaked dolphins; and inArgyrocetus, at all events, the occipital condyles, instead of being closely pressed to the skull, are as prominent as in ordinary mammals, while the nasal bones, instead of forming mere rudimentary nodules, were squared and roofed over the hind part of the nasal chamber.
In the MioceneSqualodon, representing the familySqualodontidae, the dentition is differentiated into incisors, canines and cheek-teeth, the hinder ones of the latter series having double roots and compressed crowns carrying serrations on the hinder edge; generally the dental formula has been given as i.3⁄3, c.1⁄1, p.4⁄4, m.7⁄7, the single-rooted cheek-teeth being regarded as premolars and those with double roots as molars. Dr Abel is, however, of opinion that the formula is better represented as i.3⁄3, c.1⁄1, p.8 or 9⁄9, m.3⁄2; the teeth reckoned as molars corresponding to those of the creodont Carnivora. The single-rooted cheek-teeth are regarded as due, not to the division of double-rooted ones, but to the fusion of the two roots of teeth of the latter type. InSqualodonthe nasal bones were of the modern nodular type, but in the Miocene PatagonianProsqualodonthey partially covered the nasal chamber.
At present there is a gap between the most primitive squalodonts and the Eocene zeuglodonts (Zeuglodontidae), which are regarded by Messrs Max Weber, O. Abel and C.W. Andrews as the direct forerunners of the modern-toothed whales, forming the suborderArchaeoceti. It is, however, right to mention that some authorities refuse to admit the relation of the Archaeoceti to the whales.
In the typical zeuglodonts the long and flat skull has large temporal fossae, a strong sagittal crest, a long beak formed mainly by the premaxillae (in place of the maxillae, as in modern whales), and long nasal bones covering over the nasal chamber, so that the nostrils opened about half-way down the beak. All the cervical vertebrae were free. Normally the dentition in the typical genusZeuglodon(which is common to the Eocene of North America and Egypt) is i.3⁄3, c.1⁄1, p.4⁄4, m.3⁄3; the cheek-teeth being two-rooted, with compressed pointed crowns, of which the fore-and-aft edges are coarsely serrated. In the EgyptianZeuglodon osiristhe number of the molars is, however, reduced to2⁄3, while some of the earlier cheek-teeth have become single-rooted, as in the squalodonts. The probable transitional form between the latter and the zeuglodonts is the smallMicrozeuglodon caucasicusdescribed by the present writer, from the Caucasus. As regards the origin of the zeuglodonts themselves, remains discovered in the Eocene formations of Egypt indicate a practically complete transition, so far at least as dental characters are concerned, from these whale-like creatures to the creodont Carnivora. In the earliest type,Protocetus, the skull is practically that of a zeuglodont, the snout being in fact more elongated than in some of the earliest representatives of the latter, although the nostrils are placed nearer the tip. The incisors are unknown, but the cheek-teeth are essentially those of a creodont, none of them having acquired the serrated edges distinctive of the typical zeuglodonts; and the hinder premolars and molars retaining the three roots of the creodonts. In the somewhat laterProzeuglodonthe skull is likewise essentially of the zeuglodont type, although the nostrils have shifted a little more backwards; as regards the cheek-teeth, which have acquired serrated crowns, the premolars at any rate retain the inner buttress supported by a distinct third root, so that they are precisely intermediate betweenProtocetusandZeuglodon. Yet another connecting form isEocetus, a very large animal from nearly the same horizon asProzeuglodon; its skull approaching that ofZeuglodonas regards the backward position of the nostrils, although the cheek-teeth are of the creodont type, having inner, or third, roots. It is noteworthy thatZeuglodonapparently occurs in the same beds as these intermediate types.
It follows from the foregoing that if zeuglodonts are the ancestors of the true Cetacea—and the probability that they are so is very great—the latter are derived from primitive Carnivora, and not, as has been suggested, from herbivorous Ungulata. The idea that the zeuglodonts were provided with a bony armour does not appear to be supported by recent discoveries.
Authorities.—The above article is based on that by Sir W.H. Flower in the 9th edition of this work. See also W.H. Flower, “On the Characters and Divisions of the Family Delphinidae,”Proc. Zool. Soc.(London, 1883); F.W. True, “Review of the Family Delphinidae,”Proc. U.S. Museum, No. 36 (1889); R. Lydekker, “Cetacean Skulls from Patagonia,”Palaeontol. Argentina, vol. ii:An. Mus. La Plata(1893); W. Dames, “Über Zeuglodonten aus Ägypten,”Paläontol. Abhandlungen, vol. i. (1894); F.E. Beddard,A Book of Whales(London, 1900); O. Abel, “Untersuchungen über die fossilen Platanistiden des Wiener Beckens,”Denks. k. Akad. Wiss. Wien., vol. lxviii. (1899); “Les Dauphins longirostres du Bolérien,”Mém. musée d’hist. nat. belgique(1901 and 1902); “Die phylogenetische Entwickelung des Cetaceengebisses und die systematische Stellung der Physeteriden,”Verhandl. deutsch. zool. Gesellschaft(1905); E. Fraas, “Neue Zeuglodonten aus dem unteren Mittelocean vom Mokattam bei Cairo,”Geol. und paläontol. Abhandl.ser. 2, vol. vi. (1904); C.W. Andrews, “Descriptive Catalogue of the Tertiary Vertebrata of the Fayum” (British Museum, 1906).