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[From the TRANSAcTIONS oF THE Zootoareat-Socrery oF Lonvon, vol. viii. part i. |

VI. On the recent Ziphioid Whales, with a Description of the Skeleton of Berardius
amouxi. By WituiaAM Henry Fiower, /.B.S., V.P.Z.S., Hunterian Professor
of Comparative Anatomy, and Conservator of the Museum of the Royal College
of Surgeons.

Read November 7th, 1871.
[Puates XXVII., XXVIII., XXIX.]

THE interest which attaches itself to the remarkable division of the Cetacea which
forms the subject of the present communication, is in some respects even greater than
that which belongs to all the other members of the order.

The Ziphioid Whales form a very compact group, closely united together by the
common possession of very definite structural characters, and as distinctly separated
from all other groups by equally definite characters.

With the singular exception of Hyperoodon rostratus (the structure and habits of
which species are as well known perhaps as those of any other Cetacean), no specimen
of the group had ever come under the notice of any naturalist up to the commencement
of the present century. Since that time, however, at irregular intervals, in various
and most distant parts of the world, solitary individuals have been caught or stranded,
now amounting to about thirty, which by some naturalists are referred to upwards of
a dozen distinct species, and to very nearly as many genera. No case is recorded of
more than one of these animals having been observed in one place at a time; and their
habits are almost absolutely unknown. Their very presence in the ocean seems to pass
unnoticed and unsuspected by voyagers, and even by those whose special occupation is
the pursuit and capture of various better known and more abundant cetaceans, until
one of the accidental occurrences just alluded to reveals the existence of forms of animal
life of considerable magnitude (for they range between fifteen and thirty feet in length),
and at least sufficiently numerous to maintain the continuity of the race.

This comparative rarity at the present epoch contrasts greatly with what once obtained
on the earth, especially in the period of the deposition of the Crag formations, and leads
to the belief that the existing Ziphioids are the survivors of an ancient family which
once played a far more important part than now among the Cetacean inhabitants of the
ocean, but which have been gradually replaced by other forms, and are themselves pro-
bably destined ere long to share the fate of their once numerous allies or progenitors.

These considerations are sufficient to lead to the endeavour to collect all available
information with regard to them, and to put it in a convenient form for the guidance
of those who may have opportunities to pursue their history further. Doubtless such

VOL. VIII.—PART 11. September, 1872. 2H

204 PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES.

opportunities will become more and more frequent as the attention of naturalists in
various parts of the world is directed to the importance of never omitting any chance of
observing and, if possible, securing the remains of every specimen of the group that
may come within his reach.

The Ziphioid’ Cetaceans belong to the great primary division or suborder of the
Odontocetes, or Toothed Whales. The following are their principal common charac-
teristics, as far as they can be defined in the absence of sufficient knowledge of the
structural characters of several forms :—

1. No functional teeth in the upper jaw.

2. Teeth of mandible quite rudimentary and concealed in the gum, with the excep-
tion of one, or occasionally two, pairs, which may be largely developed, and project like
tusks from the mouth, especially in the male sex.

3. Bones of the cranium raised so as to form an elevated prominence or crest behind
the nares.

4, Rostrum long and narrow.

5. Pterygoid bones very large and solid, produced backwards, meeting in the middle
line, and not involuted but simply hollowed on the outer surface.

6. A distinct bone in the orbit, segmented from the posterior part oe the malar, and
ee the homologue of the lachrymal.

. The tympano-periotic bone fixed to the cranium by a posterior, long, wedge-
shane (mastoid ?) process lying in a groove between the exoccipital and the squamosal.

8. Number of vertebra not exceeding 50; ribs not exceeding 10 pairs.

9. ‘Transverse processes of arches of dorsal vertebre ceasing abruptly near the end of
the series, and replaced by processes on the body at a much lower level, and which are
in a line (or serially homologous) with the lumbar transverse processes.

10. Spines of dorsal and lumbar vertebre very long, transverse processes short, bodies
of posterior lumbar and anterior caudal vertebree much elongated.

11. Sternal ribs permanently cartilaginous.

12. Pectoral limbs small, with a rounded extremity. The five digits all moderately
well developed.

13. Asmall subfalcate dorsal fin, situated considerably behind the middle of the back.

14. External respiratory aperture a single, transverse, median, subcrescentic opening.

15. A pair of longitudinal cutaneous furrows on the throat, nearly meeting in the
middle line in front, and diverging posteriorly.

By the majority of the above characters, and all the more important ones, as Nos. 1,

1 According to strict rules of priority, ‘“‘ Hyperoodontoid” would be the more correct term, as Hyperoodon
was the first genus of the group distinctly characterized; but as the name is erroneous in its signification, and
not hitherto used, I have thought it better to keep to the more generally adopted and less objectionable term
of “ Ziphioid” first applied by Gervais to the animals allied to the Cuvierian genus Ziphius. They constitute
Eschricht’s Rhynchoceti.

PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES. 205

3, 5, 7, 9, 11, and 12, they differ from the typical Dolphins and agree with the
Cachalots; and though they are separated from the latter by certain definite characters,
as Nos. 2 and 6, and others of less importance, there is no doubt that the group of
which Physeter is the type and that which includes Zphius and its immediate allies
are closely related, and should be considered subdivisions of one great division which
excludes all the other Odontocetes. Perhaps it is most convenient to treat them as
subfamilies of a common family Physeteride, taking its name from the earliest charac-
terized genus Physeter (Linn.), and, according to the convenient nomenclature rules of
the British Association Committee, to call one Physeterinw and the other Ziphiine’ ;
though some zoologists may prefer to raise both to the rank of families.

Among the restricted Delphinidz no genus is known which appears to form any true
transition towards the Physeteride; but the strange and aberrant genera Platanista,
Inia, and Pontoporia, which cannot be placed in either of the two principal families of
the Odontocetes, and yet have scarcely sufficient common characters to constitute a
group apart, do in some few respects fill up the otherwise wide intermediate space’.

The excessively confused synonymy of the genera and species of the Ziphiine, as
thus defined, is a cause of great difficulty in writing about this group, and makes one
almost hesitate to enter upon the subject, lest in endeavouring to clear up the confusion
the perplexity should be inadvertently increased, either by adding new synonyms or by
adopting and perpetuating ill-chosen and incorrect terms.

In a recent memoir on the group by Professor Owen’, the difficulty is disposed of
in a very summary manner by uniting all the known forms, both living and extinct
(with the exception of Hyperoodon), under the generic name of Ziphius. This pro-
ceeding at all events has the merit of running no risk of adding to the confusion
of nomenclature which has been caused by hasty or ill-defined generic subdivisions,
often founded on imperfect or fragmentary knowledge of the animal described. But,
however great our admiration may be for this strong-handed resistance to the system
of name-coining, which is fast rendering the study of zoology almost an impossibility,
it must not lead us to overlook well-marked structural characteristics by which certain
small groups of species are allied together and differentiated from others, whether we
call them genera or by any other term. The question now is, can the genus Zphius,
as understood by Owen, be divided into any such groups?

1 This is the arrangement adopted in the sketch of the classification of the Cetacea given in Trans. Zool. Soc.
vol. vi. p. 113.

* In the attempt at a natural classification of the Cetacea just referred to, I had provisionally grouped these
genera into a single family, between the Physeteride and the Delphinide; but the details of the anatomy of
Pontoporia, first made known in the valuable monograph of Dr. Burmeister, published in the ‘ Anales del
Museo Publico de Buenos Aires,’ vol. i. pp. 389-442 (1869), show that this family can scarcely be retained,
without at least considerable modification of the characters assigned to it. The only alternative seems to be to
make of each of these three genera a distinct family.

3 « British Fossil Cetacea from the Red Crag,” Paleeontographical Society’s vol. xxiii. (1870).

2H2

206 PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES.

It must be premised that as far as is at present known, putting aside the peculiar
rounded form of the head in Hyperoodon, the external characters of the various known
Ziphiine afford no grounds for generic subdivision. It is to the skeleton and the teeth
that we must look in examining whether the group is truly homogeneous or not; and it
is only very recently that complete skeletons of a sufficient number of individuals have
been known, to attempt a comparison between them. ‘The teeth have been relied upon
almost entirely; and I agree with Professor Owen that the trifling differences in the
situation of the developed teeth are not such as, unless accompanied by other more
important and constant characters, are sufficient for generic distinction; but, at the
same time, if such differences are constantly associated with others, they may be useful
guides to classification.

Hyprroopon (Lacépéde) appears to differ from all the other Ziphiine in the cha-
racters of the cervical vertebra, especially in their great antero-posterior compression,
the constant ankylosis of all seven, and the absence of inferior transverse processes
in the third, fourth, fifth, and sixth’; it has also only nine pairs of ribs, and but forty-
five vertebre in all’. In the more essential characters of the cranium it resembles one
of the other sections to be spoken of presently (M/esoplodon), with the superaddition of
the great maxillary crests; and in the dentition it resembles more nearly another
section (Ziphius).

Having separated Hyperoodon, the remaining known members of the subfamily agree
in having a comparatively well-developed cervical region, with certain of the posterior
vertebre (one, or usually more) permanently detached, and with distinct inferior
transverse processes as far as the sixth, in having almost always ten pairs of ribs*®
and at least forty-six vertebree, usually forty-eight or forty-nine. There are, however,
in the conformation of the skull and the form and situation of the teeth considerable
differences, by which they may be divided into three distinct sections, which appear to
me to be natural, and which are not, as far as is yet known, united by intermediate
forms; so I think that they may well be considered generic, though of course this is
a subject upon which the judgment of zoologist$ may differ. I can see no grounds
at present for any further subdivision.

These sections may be characterized as follows—though the distinctive peculiarities
are more readily appreciated by an inspection of a specimen than they can be expressed

in words.

1 The great differences between the cervical vertebre of Hyperoodon and Ziphius cavirostris (erroneously
called Hyperoodon gervaisii) were pointed out by Duvernoy (Annales des Sciences Naturelles, 1851, p. 24).

2 The animal described by Cope, apparently without personal examination, as Hyperoodon semijunctus (Proc.
Acad. Nat. Sc. Philadelphia, 1865, p. 280), and stated to be in the Charleston Museum, is evidently not a
Hyperoodon, but most probably a true Ziphius. It was but between twelve and thirteen feet long, has the
four posterior cervical vertebre free, has ten pairs of ribs, and two more vertebra than Hyperoodon.

3 The skeleton of Ziphius cavirostris at Jena has but nine pairs of ribs.

PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES. 207

Zipuius, Cuvier’. | ~27

Skull with the premaxille immediately in front and at the sides of the nares ex-
panded, hollowed, and with elevated lateral margins, the posterior ends rising to the
vertex and curving forwards, the right being considerably more developed than the left ;
the conjoined nasals forming a strongly pronounced asymmetrical eminence at the top
of the cranium, projecting forwards over the nares, flat above, most prominent and
rounded in the middle line in front, and separated by a notch on each side from the
premaxille. Anteorbital notch not distinct. Rostrum (seen from above) triangular,
gradually tapering from the base to the apex, upper and outer edges of maxille at
base of rostrum raised into low roughed tuberosities. Mesethmoid cartilage usually
densely ossified, and coalescing with the surrounding bones of the rostrum.

A single conical tooth of moderate size on each side of the mandible, close to the
anterior extremity, with its apex directed forwards and upwards.

The type of this genus is Ziphius cavirostris, Cuvier, founded on an imperfect skull
picked up in 1804 on the Mediterranean coast of France, near Fos, Bouches-du-Rhone.
and described and figured in the ‘ Ossemens Fossiles’*, under the impression that it was
that of an extinct species, a view which Gervais has clearly shown to have been
erroneous. This is Petrorhynchus mediterraneus of Gray’s Suppl. Cat. Seals and Whales
in British Museum, 1871, p. 98.

A second specimen was taken on the coast of Corsica: its external characters are
described and figured by Doumet in the ‘ Revue Zoologique,’ v. 1842, pp. 207, 208; and
its skeleton is preserved at Cette*.

A third specimen was stranded near Aresquiers, Hérault, South France, in 1850.
The skull, which is preserved in the Paris Museum, is described by Gervais (Annales
des Sciences Nat. 5° sér. tome xiv. 1850). This is the Hyperoodon gervaisit of Duvernoy
(Annales des Sc. Nat. 1851), Ziphius gervaisii of Fischer, and Epiodon desmarestii of
Gray’s Catalogue*.

A fourth is a skull is the Museum of Arcachon. It was found on the beach at
Lanton, Gironde, West France, in 1864, and is very carefully described and figured by
Fischer in the ‘ Nouvelles Archives du Muséum,’ tome iii. 1867, p. 42, pl. 4°.

5. A complete skeleton of a very old animal in the Anatomical Museum of the
University of Jena. This was obtained at Villa Franca in 1867, by Professor Haeckel,
but has not yet been described.

1 “Yappliquerai au genre dont elle [skull of Z. cavirostris] devient le premier type, le nom de Zphius,
employé par quelques auteurs du moyen Age (voyez Gesner, i. p. 209) pour un Cétacé quils n’ont point
déterminé.”—Cuvier, Ossemens Fossiles.

? This skull is also figured in Van Beneden and Gervais’s ‘ Ostéographie des Cétacés,’ pl. 21. fig. 7.

% Figured in Van Beneden and Gervais, op. cit. pl. 21. figs. 8, 9. 34

4 Figured in Van Beneden and Gervais, op. cit. pl. 21. figs. 1-6.

5 Figured in Van Beneden and Gervais, op. cit. pl. 21. fig. 6.

208 PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES.

6. A skeleton in the Museum at Gottenburg. From Gullmarsfjirden, on the Swedish
coast, north of Gottenburg, 1867 (vide Malm, “ Hvaldjur i Sveriges Museer ar 1869.”
Kong]. Svenska Vetenskaps Akademiens Handlingar, Band 2. no. 2. 1871).

7. A skull in the Anatomical Museum, Edinburgh, obtained from Shetland, 1871.

8. A skeleton in the Museum at Pisa from the Mediterranean. Professor Gervais
has informed me of this specimen, which has not been described.

9. In the Museum of the University of Louvain is a skull of an animal of this genus
brought from the Cape of Good Hope, of which an excellent description has been
published by Professor Van Beneden, under the name of Ziphius indicus (Mém. de
PAcad. Roy. de Belgique, coll. in-8vo, tome xvi. 1863").

10. A very similar skull in the British Museum, also from the Cape of Good Hope,
has been described by Gray (Proc. Zool. Soc. 1865, p. 524) under the name of Petro-
rhynchus capensis; and the same name is retained in the Suppl. Cat. Seals and Whales
(1871), p. 98, although its specific identity with the last-named previously described
specimen is admitted. It is further described by Owen (Crag Cetacea, Palzont. Soc.
NSKO> ps1) 7.

11. A complete specimen of a young male, 3°95 metres long, was taken near Buenos
Ayres in 1865, and is the subject of an elaborate memoir by Burmeister (Anales del
Museo publico de Buenos Aires, vol. i. p. 312, 1869), accompanied by detailed figures
of external characters, skeleton, and some of the viscera. The specimen was first
named in a preliminary notice (Ann. & Mag. Nat. Hist. 1866, xvii. p. 94) Ziphior-
rhynchus cryptodon, but described subsequently as Epiodon australis.

Until more abundant materials are obtained, and especially a complete knowledge of
the external characters or entire skeleton of several individuals, it is impossible to deter-
mine whether the differences that have been noticed in the above specimens are the
results of age, sex, individual peculiarity, or whether they denote specific distinctions.
For the present it may be advisable to admit Z. cavirostris (Nos. 1, 2, 3, 4, 5, 6, 7, and 8)
and Z. indicus (Nos. 9 and 10) as species; but with reference to No. 11, it is not impro-
bable that it is the young of one of the others. It should be remarked that Fischer,
after a careful comparison, arrives at the conclusion that No. 3 is specifically distinct
from Z. cavirostris, although not agreeing with Duvernoy’s opinion that it should be
placed in the genus /Hyperoodon.

MEsoPLopon®, Gervais.
Premaxille not greatly expanded and hollowed in front of the nares, rising suddenly

1 Figured in Van Beneden and Gervais’s ‘ Ostéographie des Cétacés,’ pl. 21. figs. 11-13.

2 Figured in Van Beneden and Gervais, op. cit. pl. 21. fig. 10.

* There is much difficulty in determining the most appropriate name for this genus. The earliest known
specimen was assigned by its discoverer Sowerby to Physeter, from which, however, it is clearly distinct. In the
classification of the Dolphins furnished by De Blainville to Desmarest’s article “ Dauphin” in the ‘ Nouveau

PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES. 209

on the sides of the nares to the vertex, where they are dilated laterally, the right one
especially, the outer edges curving backwards, their anterior surface arching forwards
above, overhanging the nares. Nasals lying, more or less sunken, in a hollow between
the upper ends of the premaxille; their anterior surface more or less concave, not
projecting so far forward as the upper part of the premaxille, and not separated on
each side from those bones by a distinct notch. Anteorbital notch not very distinct.
Rostrum long and narrow. No maxillary tuberosities. Mesethmoid generally ossified
in its entire length, and coalescing with the surrounding bones.

A much compressed pointed tooth in each ramus of the mandible, 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 males).

Dictionnaire d’Histoire Naturelle,’ 2nd edit. Paris, 1817, the subgenus Heterodon comprises eight species,
of which five (D. grenlandicus, chemnitzianus, edentatus, bidentatus, and butskode) are synonyms of Hyperoodon
rostratus, one (D. epiodon) an ill-described species from the Mediterranean, perhaps a true Ziphius, and two
(D. sowerbensis and D. densirostris), undoubtedly belong to the section at present under consideration, being
founded on the only specimens at that time known to naturalists. It is clear, therefore, that Blainville’s
Heterodon is equivalent to the present section, plus Hyperoodon; and the latter being removed, the name might
very well have been retained for the remainder, if it had not been previously in use for a genus of snakes.
Heterodon is employed in the same sense as by De Blainyille for a subgenus in Desmarest’s ‘ Mammalogie,’ pt. 2.
1822, and as a genus in Lesson’s ‘ Manuel de Mammalogie,’ 1827. The specimen taken at Havre in 1825,
apparently a female of Sowerby’s Dolphin, supposed by its first describer, De Blainville, to be of the same
species as the Dolphin described by Dale (now considered a Hyperoodon), was named by Cuvier Delphinus
micropterus, and forms the type of the genus Delphinorhynchus of F. Cuvier’s ‘ Histoire des Cétacés’ (1836),
being associated with several other Dolphins of very different structure and even belonging to different families.
But Delphinorhynchus had been previously used by Blainyille, in the article above cited, for a heterogeneous
group of Dolphins, among which none of the present genus appears; so that it is perfectly inadmissible. The
term Diodon, proposed by Lesson for the male, was already in general use for a genus of fish. Aodon (Lesson,
Compl. de Buffon), changed to Modus (Wagler, Syst. de Amph. 1830), likewise proposed for the female, being
positively erroneous in signification, have never been generally received. Wagner (Schreber, Supplement,
p- 352, 1846) constituted Micropterus as a subgenus of Delphinus, for the then known animals of the group,
uniting them into a single species, but overlooking the fact that the name had already been given to more than
one genus in the animal kingdom. Eschricht, however, adopted it in a generic sense (Nordische Wallthiere,
p- 50, 1849), altering the spelling to Micropteron, in which form it has been used by Huxley (Proc. Geol. Soc.
1864, p. 388). In 1850 Gervais (Annales des Sciences Naturelles, 3° sér. tom. xiv.) divided the group (as
defined above), though, as appears to me, on very insufficient grounds, into two genera, which he named
Mesoplodon and Dioplodon, Blainville’s Heterodon sowerbensis being the type of the one, and his H. denstrostris
the type of the other. In the following year Duvernoy, in a memoir in the same journal, reunited them,
bestowing the name of Mesodiodon on the wholé group. Subsequently Fischer (Nouy. Archives du Muséum,
iii. 1867, p. 67), not recognizing Gervais’s divisions, adopted his name Mesoplodon for the entire genus, in
which I have followed him. Owen, as above mentioned, includes this group, with all the rest of the subfamily,
except Hyperoodon, in the Cuvierian genus Ziphius (Crag Cetacea, Palsont. Soc. vol. xxiii.), while Gray
(Suppl. Cat. Seals and Whales in Brit. Mus. 1871) divides it into Zphius, Dolichodon, Neoziphius, and Dio-
plodon, which, with Berardius, constitute the family Ziphiide—the type of Cuvier’s Ziphius being placed,
under the name of Petrorhynchus mediterraneus, in a different family.

210 PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES.

The specimens of animals referable to this genus preserved in museums are more
numerous than those of Ziphius. ‘They include :—

1. An imperfect skull in the University Museum, Oxford, from the animal (a male
16 ft. long) obtained on the coast of Elginshire, figured and described by Sowerby (Brit.
Miscellany, p. 1, 1804) under the name of Physeter bidens, but to which the specific
name of sowerbiensis or sowerbyi has since been generally attached. It is called Del-
phinus (Heterodon) sowerbensis by Blainville (Nouveau Diction. d'Histoire Naturelle,
2nd ed. tome ix. 1817, p. 177), and D. sowerbyi by Desmarest (Mammalogie, 1822,
p- 521).

2. A skull in the Paris Museum, from a female specimen 15 feet long, stranded at
Havre, September 9th, 1825, described by Blainville (Nouv. Bulletin Sciences, Soc.
Philom. t. iv. 1825, p. 139) as the “ Dauphin de Dale,” by Cuvier (Régne Animal,
1829, t.i. p. 288) as Delphinus micropterus, by F. Cuvier (Hist. Nat. des Cétacés) as
Delphinorhynchus micropterus, and. afterwards by other authors under a variety of dif-
ferent names, but now generally considered to be specifically identical with the first
mentioned’.

3. A complete skeleton in the Brussels Museum from a young specimen stranded at
Ostend, August 31st, 1855, described by Dumortier (Mém. Acad. Royal. Bruxelles, 1839,
t. xii. tabb. 1-3) under the name of Delphinorhynchus micropterus. ‘The skeleton sub-
sequently described by Van Beneden as Mesoplodon sowerbiensis (Mém. Acad. Belgique,
coll. in-8vo, t. xvi. 1865)’.

4. A skull and part of skeleton in the Museum at Caen, from Sallenelles, Calvados,
north of France, 1825; described by E. Deslongchamps (Bulletin de la Scc. Linn. de
Normandie, tom. x. 1866) as Mesoplodon sowerbyensis?.

5. A mutilated skull in the Museum of the Irish Academy, Dublin, from an animal
15 feet long, stranded in 1864 in Brandon Bay, Kerry, Ireland (Andrews, Trans. Roy.
Trish Academy, vol. xxiv. 1869, Ziphius sowerbyi).

6. The skull of another specimen from the same locality, a male, about 17 feet long,
stranded May 31st, 1870 (Andrews, Proc. Roy. Irish Acad. ser. 2, vol. i. p. 49).

7. A mandible in the Museum at Christiania, from the Norway coast, figured and
described by Van Beneden (Bulletin de lAcad. Roy. de Belgique, t. xxii. 1866) as
Mesoplodon sowerbiensis.

8. A skull in the University Museum, Edinburgh, of unknown origin. (I am indebted
to Professor Turner for information of this specimen, which has not yet been described.)

9. A complete skeleton in the Gottenburg Museum, described by Malm (loc. cit.)
under the name of Micropteron bidens. From the coast of Norway, 1869.

All the above appear to belong to one species.

10. A skull in the Museum at Caen, from an animal caught in the entrance of the

? Figured by Van Beneden and Gervais, op. cit. pl. 26. figs. 5-8.
? Ibid. pl. 22, 5 Thid. pl. 26. figs. 1-4.

PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES. 211

Channel about 1840. Described by Gervais as D. europeus, by E. Deslongchamps
(Joc. cit.) as Dioplodon gervaisii’.

This appears to be at present a unique specimen.

11. In the British Museum is a skull received from the Cape of Good Hope, with
remarkably developed teeth in the lower jaw, passing upwards and backwards, and
finally curving inwards so as to meet over the rostrum. This has been named Ziphius
layardii by Gray (P. Z. 8. 1865, p. 358), and is figured and fully described by Owen
(Crag Cetacea, p. 12, pl. 1). The condition of the teeth suggest an individual pecu-
liarity; but Mr. E. Layard has in his possession a single tooth of another individual
(also from South Africa) having an exactly similar conformation.

12. The Australian Museum at Sydney has lately obtained a skeleton of an animal of
this group, stranded at Little Bay, about six miles from Sydney, which has not yet been
described; but, judging from the photograph sent by Mr. Krefft, it is closely allied to,
if not identical with the last. The teeth, however, are much less developed.

13. A skull in the Paris Museum, from the Seychelle Islands, has been figured and
described by Duvernoy (Annales des Sciences Naturelles, 1851) under the name of
Mesodiodon densirostris, being apparently identical with the rostrum, of unknown origin,
described by De Blainyille under that name (Nouv. Dict. d'Histoire Nat. 2nd edit.
tome ix. 1817). It has also received the specific name of seychellensis from Dr. Gray”.

14. A complete skeleton of an animal of the same species, obtained from Lord
Howe’s Island, is in the Australian Museum at Sydney. A brief description has been
given of it by Krefft (P. Z. S. 1870, p. 426), and an outline figure in Ann. and Mag. Nat.
Hist. vol. vi. 4th ser. 1870, p. 343.

The last two belong undoubtedly to a species distinct from any of the others, charac-
terized by the peculiar form of the ramus of the lower jaw, and of the very massive
tooth which it supports. It is to be hoped that further details of the structure of the
skeleton, especially of the cervical vertebrae (which appear to be different from those of
other members of the group), will be published before long.

15. In the Museum at Wellington, New Zealand, is a skull and some bones of an
animal 9 feet 3 inches long, figured and partially described by Dr. Hector in the Trans.
New-Zealand Institute, vol. ii. p. 27, and vol. ii. pls. 14 and 15. This has been named
Berardius hectori by Gray (Ann. and Mag. Nat. Hist. August 1871). The conformation
of the skull shows that it is a member of the present group; but the single compressed
tooth in the lower jaw is situated further forwards than in any other known species,
thus completing, with densirostris, sowerbyi, layardii, and europeus, the series of dif-
ferent positions in the side of the ramus occupied by the developed tooth, and proving
its little importance as a generic character.

16. In the Report of the Director of the Museum of Comparative Zoology at
Cambridge, U.S. A., for the year 1869, among the additions made to the collection by
' Figured by Van Beneden and Gervais, op. cit. pl. 24. * Thid. pl. 25. figs. 2, 3.

VOL. VIII.—ParT 11. September, 1872. 21

212 PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES.

donation, a skull of ‘‘ Mesoplodon sowerbyensis” is mentioned as presented by Stephen
C. Martin, but without any further details.

Berarpius, Duvernoy.

Upper ends of the preemaxille nearly symmetrical, moderately elevated, very slightly
expanded, and not curved forwards over the nares. Nasals broad, massive, and rounded,
of nearly equal size, forming the vertex of the skull, flattened in front, most prominent
in the middle line. Anteorbital notch distinct. Rostrum long and narrow. Mesethmoid
only partially ossified. Small rugous eminences on the outer edge of the upper surface
of the maxille at base of rostrum.

Two moderate-sized, compressed, pointed teeth on each side of the symphysis of the
mandible, with their apices directed forwards, the anterior being the larger of the two,
and directed forwards’.

History of Berardius arnouxi’.

The genus Berardius was founded by Duvernoy® upon a skull received at the Museum
of Paris in 1846, having been obtained from an animal stranded in Akaroa Harbour,
Banks Peninsula, New Zealand. The following brief description of the animal, and
the circumstances attending its capture, was supplied by M. Arnoux, surgeon to the
corvette ‘ Rhin,’ commanded by Captain Bérard, by whom the skull was presented to
the Museum.

‘“¢Cet animal vint échouer, sur la céte, dans le port d’Akaroa. ... . Des habitants
anglais, voisins de ce lieu, le tuérent 4 coups de lance... .. Ils en retirérent trois
barils de graisse. La corvette ‘le Rhin’ rentra dans le port d’Akaroa trois ou quatre
jours apres cet événement . . . . Je m’empressai d’aller voir les restes de cet animal, et
je m’emparai de la téte et d'un aileron... .. L’animal vivant avait 32 pieds anglais de
longueur totale; il était pourvu d’une nageoire dorsale assez étendue précédée d’une
bosse assez considérable. Sa couleur était entiérement noire, sauf une partie grisatre
claire vers les organes génitaux: c’était un male.” j

No portion of the animal except the skull was brought to the Museum. Duvernoy’s
description of this is accompanied by a figure of the upper surface of the cranium, of
the mandible, and a side view of one of the teeth*.

1 Tn the specimen in the Wellington Museum the second tooth does not seem to be developed. See Trans.
New-Zealand Institute, vol. iu. pls. 13 & 14, p.129. This, together with the circumstances mentioned with
respect to the teeth of different species of Mesoplodon, shows that the diagnoses of the genera of Ziphioids from
the teeth alone, relied upon by some authors, as Fischer, are insufficient.

? Spelt arnuwii by Duvernoy, arnouai and arnouxii by other authors.

5 “Mémoire sur les caractéres ostéologiques des genres nouveaux ou des espéces nouvelles de Cétacés vivants
ou fossiles.”” (Annales des Sciences Naturelles, 3° série, Zoologie, tome xv. 1851, pp. 1-71.)

* Loc. cit. pl. 1.

PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES. 213

In the great work on the osteology of the Cetacea in the course of publication by
Van Beneden and Gervais, figures on a larger scale are given of several views of the
same skull (pl. 23), but the description has not yet appeared.

Until the arrival of the subject of the present communication the Paris skull was the
only portion of Berardius contained in any European museum. The length of the
cranium is 595 inches.

2. The next indication of the existence of Berardius is thus recorded by Mr. F. J,
Knox :—

“In January 1862, a male specimen was embayed in the Porirua harbour, and was
captured by Mr. London, of which I was only able to make a rude sketch and take a
few of the measurements.”

The following are the dimensions given :—

ft. in.

“Total length pal
Greatest circumference: 375 3 8 > Le. 0
Tail, from tip to tip cies 40
Head.—Length of basal surface ; 4 4
FLEW) meet tere fick op SEE 2 2

Breadth across occiput Zi 0

Length of lower jaw 3 10

Length of pectoral extremity, free 3 0

“Produce of fine sperm oi], about 240 gallons; spermaceti, a considerable quantity
in the upper surface of the face’”’’.

Except a statement that in the recent state no vestige of teeth could be discovered,
no further description and no preparations were obtained from this specimen, which
can only be conjectured to be a Berardius from its locality and size. The extremely
slight sketch (pl. xiii. fig. 1) of the outline of this animal shows only the characters
common to the Ziphioid Whales, except that the pectoral fin is unusually long and
pointed. This, however, corresponds neither with Mr. Knox’s measurements nor with
the form of the manus of the present skeleton’.

3. No animal of the species was again observed until the subject which afforded the
skeleton to be described in the present memoir was stranded, of which event the fol-
lowing account was given by Dr. Julius Haast, F.R.S., the eminent geologist and
energetic curator of the Museum at Christchurch, Canterbury. As our knowledge of
this rare animal is so scanty, I think it desirable to reproduce Dr. Haast’s notice
entire :—

1 Transactions and Proceedings of the New-Zealand Institute, 1870, vol. iii. p. 126.

2 It may be noticed that further doubt is thrown on the accuracy of this sketch by its being attributed in
the letterpress to two quite different animals, viz. the 27-feet long Berardius, taken in 1862 in Porirua
Harbour (p. 126), and the 9-feet long Mesoplodon, taken in 1866 in Titai Bay (p. 125).

Bh

214 PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES.

“ Preliminary Notice of a Ziphioid Whale, probably Berardius arnuxii, stranded on the 16th of December, 1868,
on the Sea-beach, near New Brighton, Canterbury, New Zealand. By Jurivs Haast, Ph.D., PRS.

“ Towards the latter part of December, last year, it was stated that a whale had been stranded on the sea-
beach, near the mouth of the Avon. Unfortunately the notice reached me too late to enable me to see the
body in its fresh state ; and when I went to the sea-beach the blubber had been cut off nearly a week, and the
animal was already in such an advanced state of putrefaction that the external appearance was greatly
destroyed. Before entering into a description of its affinities and peculiarities, I may be allowed to offer a few
observations on its capture.

“Mr. William Walker, a fisherman, living near the mouth of the Avon, one mile and a half below New
Brighton, observed, on the 16th of December, early in the morning, that a huge animal was in the surf, making
the most strenuous efforts to return to deeper water. The fisherman had only a large sheath-knife with him,
with which he stabbed it several times, making it bleed very freely. ach time when the surf reached it, it
threw out a large quantity of water and sand from its blowers, like a fountain; at the same time it moved its
tail with such vehemence that it threw its captor several times when he came too near it. Seeing that he
could not manage the large animal by himself, he returned home to fetch a rope, a larger knife, and assistance.
After having, with some trouble, placed the rope round the tail, and fastened it securely to the stump of a tree
on the beach, he inflicted with the large knife some deep wounds, from which the blood ran copiously; but
the animal, notwithstanding this great loss of blood, still lived for fourteen hours. The fisherman also put a
large stick several times into its mouth, which, to use his own words, made the whale ‘ bellow like a bull.’

“A yery interesting fact may be deduced from the observations of Mrs. Walker, who accompanied her
husband on the second trip, She told her husband that each time he put the stick into the whale’s mouth, she
could see several large teeth in front of its lower jaw, which, however, were not observed by any body else,
and the existence of which was only revealed when the skull was cleaned, when, in front of the lower jaw, two
large triangular and movable teeth on each side became exposed. It thus seems that the Ziphioid Whales,
when defending themselves from their enemies, or attacking their prey, have the power to protrude these four
teeth at will. Such an hypothesis gains still more in probability when we consider the nature of the principal
food of the animal, which, judging from the contents of its stomach, seems to consist almost exclusively of the
common Sea-Spider, or Octapus—a cephalopod which, as in the northern hemisphere, does not seem to be very
numerous along the coast. In the stomach of the whale in question there was about half a bushel of the
horny beaks of this cephalopod, which were nearly all of the same size. It would be rather difficult for any
whale to obtain possession of such an agile animal as the Octopus, had not nature furnished the former with
the means of taking good hold of it. It is interesting that the allied genera Ziphius and Hyperoodon, of the
northern hemisphere, feed also on similar species of cuttlefish, as I learn from a paper of Dr. J. E. Gray, of the
British Museum (Proc. Zool. Soc. 1868, p. 422). Also the Sperm Whales are said to feed almost exclusively
on the same vivacious animal, which, by its agility and organization, is so well adapted to make great hayoe
amongst the smaller inhabitants of the sea. And, as Dr. Gray justly observes, it proves, at the same time,
that these cephalopods, although apparently of rare occurrence, must in many localities be very numerous, as it
would otherwise be impossible to understand how they could furnish those huge whales with sufficient food.

«© When I proceeded to the beach the animal was still lying in the surf, partly covered with sand, but still
intact. I measured its length exactly, and found it to be 30 feet 6 inches from the tip of the nose to the end
of the lobes of the tail. The colour of the whole animal was of a deep velvety black, with the exception of the
lower portion of the belly, which had a greyish colour. The tail was 6 feet 6 inches broad, and had the usnal
two faleate lobes. The pectoral fins were situated near the neck, a little aboye the middle of the body, and
were 17 inches broad and 19 inches long. They had a triangular form; and one of them was buried in the
sand when I saw the animal first. The dorsal fin was unfortunately destroyed when I first saw the whale, so
that I cannot describe its form and position from my own observations; but Mr, Walker told me that it was
small, had the usual faleate form, and was situated not far from the tail,

PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES. 215

‘«*T may here observe that, from the form of the skull and some other characteristics, it appears evident that
this whale is the Berardius arnuwxii of Duvernoy, of which a specimen was caught in 1846, in Akaroa harbour,
the skull of which, of the length of four feet, is at present in the Imperial Museum in Paris. The animal to
which it belonged is described as having been 32 feet long, and possessing a large dorsal fin, with a large boss
or hump in front of it. As putrefaction and the cutting off of the blubber had greatly changed the outlines of
the animal, I could not observe whether it possessed the larger boss in front. Mr. Walker did not speak of it
when he gave me a description of the animal as it appeared when captured. However, as the figure of the
skull, as given by Duvernoy in the ‘Annales des Sciences Naturelles,’ and copied into Dr. Gray’s British-
Museum ‘ Catalogue of Seals and Whales,’ is identical with that of our own specimen, I do not hesitate to state
that both belong to the same species. It also seems to me that this whale is very local, probably inhabiting
only the coast of New Zealand, and perhaps the regions south of it, because, as far as I can find, it has never
been observed elsewhere. It has without doubt not been met with on the coasts of Australia (or it would not
have passed unnoticed), as, amongst others, the energetic director of the Australian Museum, Gerard Krefft,
F.L.S., has not observed it. I may here state that the form of the skull is very peculiar, reminding one
strongly of that of a dolphin.

«There seems to be nothing known of this peculiar whale, except its external appearance and its skull; and
it is therefore a matter of congratulation to us that we shall be able to supply all the details of its osteological
characteristics, which are peculiar in many respects.

«<The specimen in our possession was evidently a young animal, because all the disk-like epiphyses of the
vertebre are still detached. The same is the case with the epiphyses of the limb-bones, which are not yet
united with them; also the sutures of the cranium are not yet obliterated. The beginning of coalescence is,
however, to be observed in the seven cervical vertebra, of which the first three are already anchylosed, the first
two completely, and the second and third only partially, as the neural arches and transverse processes are not
yet united into one bone. In the allied Hyperoodon all the cervical vertebre coalesce; and it is therefore
possible that when Berardius is in an adult state the same will take place, The Ziphius has six cervical
yertebre separate; and it will therefore be necessary to examine very carefully into the character of the
uncoalesced vertebre of our skeleton before giving a decided opinion upon the subject. It possesses ten dorsal
yertebre, in common with Ziphius sowerbiensis; the Hyperoodont Whales have nine, and the Dolphins thirteen
to fifteen. I haye not yet been able to count and examine the lumbar and caudal vertebra, as the animal was
in such a state of putrefaction that, after cleaning the bones as well as possible and leaving often a great
portion of the vertebral column together, we put them at once to macerate. We obtained only one of the
small pelvic bones, the other having probably been washed away by the surf; it might, however, owing to its
diminutive size and sticking loosely in the flesh, easily have been overlooked. As soon as the bones are clean,
so that I can examine them, I shall offer a few more observations upon the osteology of this remarkable
animal, for the complete skeleton of which the Canterbury Museum is indebted to the members of the Philo-
sophical Institute, without whose pecuniary assistance I should have been unable to secure it for the
Provincial collections.’—Proc. Phil. Institute of Canterbury, New Zealand, May 5, 1869; also Annals and
Magazine of Nat. History, October 1870, p. 348.

It is much to be regretted that no mention is made in this account of the sex of the
animal, especially as some of the allied forms are supposed to present considerable
sexual differences.

4, In January, 1870, a large Ziphioid Whale was stranded in Worser’s Bay, near the
entrance to Port Nicholson, and was captured. Its dimensions are thus given by
Mr. Knox :—

216 PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES.

ft. in.

Total length sie
From beak to nostrils 3 6
From nostrils to occipital foramen LG
Total length [of head ?] over vertex ait 5 5 0
Head.—Tip of beak to occipital foramen, basal sur pice ALO
Greatest height 2 0

Length of lower jaw . 3 10

Length of symphysis . 1g)
Breadth between condyles . 1 10
Breadth at symphysis eG}

The skull, the cervical vertebra, scapula, and imperfect pectoral limb of this animal
are preserved in the Museum at Wellington, and have been figured on a reduced scale
in the Transactions of the New-Zealand Institute, vol. ui. Although there was but a
single tooth on each side of the lower jaw, near the apex, the skull in form and size so
closely resembles that in the Paris Museum, that it is difficult to believe that they are
not specifically identical.

Though no complete description of the portions of this animal which were preserved
has yet been published, the following paragraph relating to it by Dr. James Hector,
F.R.S., is important:—“‘The preparation of the nose (figures 4@ & 46) shows that,
notwithstanding this is a full-sized animal, the tooth is still sheathed in the gum,
being imbedded in a tough cartilaginous sac, which adheres loosely in the socket of
the jaw, and is moved by a series of muscular bundles that elevate and depress it”?.

The dimensions of the skull are thus given by the same naturalist :—

inches.
Bensthiiofheady pereteds sd. Maio Seis ts side Cdule-medea ele cae peo EOE
hength! of. nose) uivia wie ts Law amas ake ee athe: ens oO)
ensthyofidentaligroove 44. 9.) 42:2 =p, Heth ee OLD)
Lens thyoflowerejaw slivers) vhtic sinew meee eaepeees aes oo
Wadthyofpmotehus Gs} ce! ye custets a2 r eyed cee elder ae eo
Widthyatiorbitsyis se. yon ayes wiemeehs 25 el ei ei eae Ae
Wadthat-blow-holes) 2... 4:4 din dee didek bag eter pede.
Wrdthjat nose.) o> oa pe +e) a ge ee eet
Heightrofjocciputec iil iced is Are al gene ee ti PLO

1 Trans. New-Zealand Institute, 1870, vol. iii. p. 129.

2 This is evidently a mistake, as it is far too large for the other dimensions. According to the figure it should
be 47:5.

3 The smaller specimen, 9 feet 3 inches in length, which was described in the 2nd vol. of the Trans. N.-Z,
Inst. (p. 27) under the name of Berardius arnuxii, and which has since been named by Dr. Gray Berardius
hectori (Ann. & Mag. Nat. Hist. August 1871), belongs, as above mentioned, to a different section of the
group.

PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES. 217

Description of the Skeleton of Berardius arnouxi.

I must return to No. 3 of the above list, of which the skeleton has been lately placed
among the fine series of Cetaceans in the Museum of the Royal College of Surgeons ;
which fortunate circumstance is due partly to the extremely liberal desire of Dr. Haast
that it should be made as available as possible for scientific examination, comparison,
and description, and partly to the generosity of Mr. Erasmus Wilson, F.R.S., a Member
of the Council of the College, in providing the means of adding it to the collection
without expense to the Institution.

The skeleton is complete, with the exception of one of the pelvic bones, and a few
of the phalanges. Although it may have attained nearly to its full size, the condition
of the bones shows that the animal was far from adult. The terminal epiphyses of the
bodies of the vertebra are separate throughout the thoracic, lumbar, and caudal regions,
though united to the rest of the bone in the neck-vertebree. The epiphyses of both
ends of the radius, the upper end of the humerus, and lower end of the ulna are free,
but that of the lower end of the humerus has partially coalesced with the shaft.

The length of the skeleton as now mounted is 29 feet in a straight line frem the tip of
the lower jaw to the end of the tail; but, notwithstanding the careful indications kindly
furnished by Dr, Haast, it is possible that the allowance made for the intervertebral
substance is not quite exact.

Viewing the skeleton as a whole (see Pl. XXVII.), the most striking feature is the
small size of the head, compared with the great length of the vertebral column, and
the massiveness of the individual vertebrae, especially of the lumbar and anterior caudal
regions. It presents a remarkable contrast to Physeter in this respect, though agreeing
generally with the other Ziphine.

Skull.—The cranium agrees so closely in form with the type specimen in the Paris
Museum, described and figured by Duvernoy, and subsequently by Gervais, that a
detailed description of its external characters will not be required. That specimen,
however, is one inch longer than the present one, and probably belongs to a fully adult
individual, The most prominent parts of the pterygoid bones are broken off, which
alters the contour of the lower margin in the figure; and the petro-tympanic bones are
wanting. ‘The present specimen is quite perfect; and as a longitudinal median vertical
section has been made through it, I am enabled to give for the first time this highly
characteristic view (Pl. XXVIII. fig. 7).

As compared with the other Ziphioids, the most remarkable features of the skull are
the almost perfect bilateral symmetry of the upper surface and the comparative sim-
plicity of the posterior ends of the premaxille, which do not curve forwards to overhang
the superior narial apertures as in the other members of the group.

The vertex is formed by the massive nasals (Va), prominent and rounded in the middle
line in front and above, and behind these by a small, but elevated, portion of the
united frontals (Zr), which at this spot, instead of being solid, are composed of several

218 PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES.

distinct irregular and freely moveable wedge-shaped pieces of bone of the nature of the
so-called ‘“‘ Wormian bones.”

The mesethmoid (JZ/) forms a strong ridge between the narrow nostrils, rising to a
level with the premaxille; its ossification becomes irregular and nodular anteriorly,
and extends as far forwards as 2 inches in front of the base of the rostrum (7. e. a line
drawn between the deepest part of the two anteorbital notches, and which is supposed
to mark off the rostrum from the cranium proper). In the Paris specimen the ossifica-
tion extends somewhat further, doubtless in consequence of the superior age of the
individual; but it is not likely that it ever attains to the remarkable extent and
solidity characteristic of some species of Ziphiine.

As in Hyperoodon and other allied forms (but not in Physeter) the bone which lies
in ordinary Dolphins beneath the anterior part of the orbital plate of the frontal, in
contact with the maxilla in front, the frontal behind, and the palatine on the inner
side, is divided by a distinct suture into two parts. ‘The anterior part gives origin to
the slender zygomatic arch, and undoubtedly corresponds to the malar of ordinary
mammals; while the posterior part appears to represent the lachrymal, as pointed out
by F. Cuvier’, Eschricht?, Van Beneden’, and others, though Duvernoy considered it a
prolongation of the orbito-sphenoid.

There are several differences in detail in the form and arrangement of these bones
between Berardius and Hyperoodon, one of the most important being that the orbit of
the former is considerably smaller than that of the latter, which, together with the
inferior size of the optic foramen, would indicate a smaller organ of vision.

The zygoma, like that of Hyperoodon, is broader and flatter, especially at its anterior
extremity, than in the ordinary Dolphins.

A most important and characteristic region of the base of the skull in the
Cetacea, as in other Mammals, is that surrounding the organ of hearing. Here
Berardius agrees with the other Ziphioids in showing affinity to Physeter* rather
than to the true Dolphins, both in the form of the tympanic bulla, and in the greater
fixedness by which it is attached to the skull. This is chiefly effected by a large
irregular wedge-shaped process (Pl. X XIX. figs. 1 & 2, m), which passes backwards and
outwards from the hinder edge of the portion of the tympanic which articulates with
the periotic, and lodges in a groove between the exoccipital and squamosal, reaching
the external border of the skull. This process so closely occupies the position of the
“mastoid” in ordinary Mammals, that it has very naturally received that name: but its
exact homology must be cleared up by a study of its development; for it differs from

1 Histoire Naturelle des Cétacés (1836), p. 76, pl. 7, 1.

2 Untersuchungen iiber die nordischen Wallthiere (1849) p. 44.

* Mem, sur une nouvelle espéce de Ziphius, Mém, de Acad, Royale de Belgique, coll, in-8vo, tome xvi.
1863, p. 14.

* And more remotely to the Whalebone Whales, as pointed out by Eschricht,

PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES. 219

the ordinary mastoid in being united to the tympanic instead of the periotic. To the
anterior and inner side the tympano-periotic is supported by a long slender process of
the squamosal, with which it comes into closer contact than in the true Dolphins’.

The tympano-periotic bones are slightly larger than those of Hyperoodon (Pl. X XIX.
fig. 6), but have much the same general form, with certain differences in detail, of which
the most important are:—in the tympanic the anterior (Eustachian) end (é) is more
prolonged, pointed, and spout-like; the groove between the posterior lobes is deeper ;
the posterior articular surface (a) for the periotic is larger and smoother; the periotic
is more elongated, the notch between its anterior (a./) and middle lobe (m./) is wider, the
anterior lobe is more prolonged and pointed in front, and the orifice of the meatus
auditorius internus (7.a.m) is rather smaller.

It is important to note that in every one of these points of difference, Hyperoodon
approaches nearer than Berardius to Physeter—a confirmation of the great taxonomic
value of the characters of this region of the skull.

On comparing the median section of the skull (Pl. XXVIII. fig. 7) with that of
Hyperoodon, the main difference is the smaller degree of elevation and of antero-
posterior compression in the great supracranial crest, and the greater extent, both in
yertical and antero-posterior direction, of the bony mass formed by the coalescence of
the presphenoid, mesethmoid, and frontals which lies in front of the cerebral cavity,
separating it from the nasal passages. In these, as in so many other respects,
Hyperoodon approaches much nearer to the Cachalot than does Berardius.

The hinder edge of the vomer, which is prolonged beneath the presphenoid and basi-
sphenoid, is much less massive in Berardius than in Hyperoodon ; but, generally speaking,
the conformation of the cranium and the relations of the bones to each other as seen in
this view are strikingly similar’.

The basioccipital and the basisphenoid have completely coalesced; but the fissure
between the latter and the presphenoid is open to the extent of more than a quarter of
an inch. ‘The floor of the cranial cavity is less curved from before backwards than in
Hyperoodon, and therefore much less than in Physeter; and the commencement of the
spinal canal is not directed upwards to the same extent. The pituitary fossa is very
indistinct ; but at some distance behind it there is a broad and deep groove on the upper
surface of the basioccipital.

The cerebral cavity presents much the same general form and size as in Hyperoodon
and the allied species, being high, very broad, and flattened from before backwards.
Its greatest breadth is 113 inches, its greatest vertical height 7 inches, its length
8 inches. The posterior upper part has a strong median projection, or osseous falar

1 See the description of this region in Hyperoodon by Eschricht (op. cit. p. 45), and in Zphius indicus by
Van Beneden.

2 The comparison was made with the skull of a Hyperoodon of corresponding age in the Museum of the
College of Surgeons ; it is figured in Trans. Zool. Soc. vol. vi. pl. 56. fig. 3.

VOL. VIII.—PART 11, September, 1872. 2k

220 PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES.

cerebri, which, in the figure, conceals the real height of the lateral parts of the cavity.
A well-marked ridge, more conspicuous than in Hyperoodon, commencing on the orbito-
sphenoid below, and extending upwards on the frontal almost to the vertex, divides the
anterior from the posterior cerebral fossa. The cerebellar fossa is relatively larger than
in Hyperoodon, and is separated from the posterior cerebral fossa by a strong ridge.
There is no olfactory fossa.

The periotic bone is excluded from the cerebral cavity by a distance of 54 inches.

The foramina which pierce the base of the cranium, as seen from within, are :—

1. A very small hole, $ inch from the middle line, and 3 inches in front of the suture
between the presphenoid and basisphenoid, and passing through the posterior lateral ex-
pansion of the mesethmoid which corresponds to the cribriform plate of other mammals,
to the nasal passage, may be an olfactory foramen. A similar foramen has been noticed
in Ziphius by Fischer, and exists on a larger scale in Physeter’.

2. The optic foramen is a rather small oval fissure (0°4 inch by 0°3 inch) perforating
the_orbito-sphenoid, near its hinder border, and soon joining the great orbital or
sphenoidal fissure. It is less than half the size of the corresponding opening in
Hyperoodon.

3. Immediately behind the sharp ridge formed by the hinder edge of the orbito-
sphenoid is the large funnel-shaped opening, compressed from side to side (about 1 inch
by 4 inch in diameter at the narrowest part), corresponding to the sphenoidal or
orbital fissure together with the foramen rotundum, as it transmits the nerves to the
orbit, as well as the middle division of the fifth nerve.

4, The foramen ovale, for the third division of the fifth, forms a distinct perforation
through the alisphenoid, about 0°3 inch in diameter. It is connected with the last by
a shallow groove.

5. Behind the orbito-sphenoid is a large infundibuliform depression, divided at the
bottom into an anterior smaller circular aperture (0°3 inch in diameter) and a posterior
larger oval opening (1 inch by 0-4 inch). The former is for the seventh nerve,
which it conducts to the meatus auditorius internus on the periotic; the latter for the
various nerves commonly known as the eighth pair; this is further divided near its
termination on the surface by a narrow bony septum.

6. Immediately behind the last, in the cerebellar fossa, and 1? inch from the middle
line, is the small condylar foramen (0-3 inch by 0-2 inch in width) for the hypoglossal
nerve. After a course of about four inches through the bone, it opens into a groove in
the hinder surface of the skull between the basioccipital and the exoccipital.

7. In the basisphenoid, 1 inch from the middle line, opposite the prominence in
front of the sella turcica (tuberculum sell) is the longitudinal narrow oval aperture
(0-4 inch by 0-2 inch) for the carotid artery. This canal opens externally in the
pterygoid close to its posterior border.

1 See “ Osteology of the Sperm Whale (Physeter macrocephalus),’” Trans. Zool. Soc. vol. vi. p. 316.

PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES. 221

In general arrangement these foramina correspond very closely with those of Hypero-
odon. In the Sperm-Whale the greater massiveness of the cranial walls, compared
with the brain-cavity, and the consequent greater distance that has to be traversed by
the nerve-canals, causes some alteration in the condition of the foramina, several (as in
the case of that for the seventh and eighth nerves) more completely coalescing into a
single aperture, at least at their cerebral ends, than in the smaller-headed Ziphioids.

Dimensions of the Skull.

inches.
Extreme length of cranium . . MRE Rees? Tas host kes OL
Length of rostrum (from the apex ai due premaxille to the middle at
a line drawn between the anteorbital notches) . . . . . . . . 36:2
Anterior end of vomer to anterior end of premaxille. . . .. . . 136
From middle of hinder edge of palate ee by pterygoid bones) to
end of premaxille . . . =) sy wants LOLS
Greatest height of skull, from a a ee to ae eeaet of ey eiiay 21
Greatest breadth, across postorbital processes of frontals . . . . . 27
Breadth across zygomatic processes of squamosals. . . . . . . . 264
Breadth between outer borders of suprafrontal processes of maxille . . 24°6
Breadthrotoccrpitalicondyles yn sate Lt ory le ote Site, ae AO
edn Or Tomien me ge Lig MONE B IA ONS RS) Eom a eg OE
Reig it olsioramenpMagnuUiaecr a aul armel. ae poy tsa) ete) iat cee ALO
Breadth of exoccipitals . . . wate cen “ee OaO
Breadth of base of rostrum (hecncen tron oF arial ieoneai loan
J SIRSAYG KAA Oe IOAN, OES BEE gee Awe he OS A oe ea GO
INasalibones;antero-nostenior length au cs wet ied ek tu on OLA
Nasalsbones, sreatest breadth of the two. <0. cus = seis oo ey not &
Anterior nares, greatest width of thetwo. . . . Sy cu a a ee amen
Breadth between outer borders of preemaxillee Ae cite MaLeS) sary ieee tet Oc O
Breadth between outer borders of Perey a at their widest part, in
front of the nares. . : th se Ores
Breadth between outer tories ai Cee at Witacte ae TOsiium’ 2 3:6
Mandible;——ene thot ramusy 900 nue goa bere tee cy fu tO
Dene chuotasym ply Sis itp tae et Veet in) en org ns LA
Vertical height of ramus at coronoid process ae 8:3
Apex of mandible projecting beyond apex of rostrum. . 2°7

The rami of the mandible are not ankylosed at the symphysis. They are more
massive in form and of a denser substance than in Hyperoodon, especially near the
symphysis, corresponding with the greater development of the teeth. A slight irre-
gular groove, with numerous minute vascular canals opening into it, extends along the

22

222 PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES.

upper margin from the apex for two thirds the length of the ramus, to opposite the
entrance of the wide infundibuliform dental canal on the inner side. Just before its
posterior termination this groove becomes wider and deeper than elsewhere ; behind it
the superior edge of the ramus rises into a sharp elevated crest, terminating behind
in the feebly marked coronoid process.

At the anterior extremity of the ramus the groove dilates into a large, oval, narrow,
alveolar cavity, 2°7 inches long, 1 inch wide and 2 inches deep. ‘Two and a half inches
behind this is another, smaller alveolus 1-8 long, 0-6 inch wide in front. and very narrow
behind. The floors of both these cavities slope backwards and upwards; and their
openings are directed forwards.

Teeth.—Kach of the above-mentioned alveolar depressions contains a tooth, which
very nearly fills the socket, and projects but slightly above the level of the upper
surface of the ramus of the jaw.

The anterior tooth of the left side, although loose in its alveolus, cannot be removed
from it as long as the bone is intact. The right tooth, however, was extracted without
much difficulty (see Pl. XXIX. fig. 7). It is compressed laterally to the form of a
nearly equilateral triangle, with a base, an apex, and an anterior and a posterior margin.
The base is 2°8 inches long, the anterior margin 2°8 inches, the posterior margin 5:2
inches, the height from the middle of the base to the apex 3 inches, the greatest thick-
ness between the lateral surfaces 0°8 inch. The pulp-cavity is completely closed below,
the base being rounded and rugose. The inner surface is concave, the outer one slightly
convex in both directions. Both surfaces are marked with irregular shallow furrows
and ridges running in a longitudinal direction, or rather radiating from apex to base.
The apex itself is conical, with a deep linear longitudinal groove on the middle of
its inner side; it appears to be formed of dentine, without any enamel covering, and
projects for a distance of 0°3 inch from the mass of cementum which covers the greater
part of the tooth. The extremity is somewhat polished, but presents no distinct signs
of wear.

The left tooth, as far as can be seen without removing it from the alveolus, exactly
resembles the right. The apex projects scarcely one inch above the level of the alveolar
border; so that before the gum was removed very little, if any, of the tooth could have
been exposed.

_ The second tooth (Pl. XXIX. fig. 8) fits very well into, and nearly fills its alveolus ;
but, owing to the form of its root, it is readily removed. It also is compressed and
triangular, but narrower from before backwards than the other. Its base is closed;
and its apex is formed, as in the other, of a small cone of dentine, emerging out of an
enveloping mass of cement; the anterior margin is thicker, more rounded and curved
than the posterior. The whole tooth lies very obliquely in its alveolus; so that the
posterior margin is nearly horizontal, and the apex projects forwards. Such a very
small portion is raised above the level of the alveolus, that we might naturally infer

PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES. 223

that in life the tooth was entirely concealed'. The height of the right tooth from
the middle of the base to the apex is 1:8 inch, the length of the base is 1-1 inch,
the length of the anterior margin is 1°5 inch, of the posterior margin 2 inches, the
greatest thickness 0-6 inch. ‘The left tooth is slightly larger than the right.

A longitudinal section in the antero-posterior direction having been made through
these teeth, their structure was seen to be very similar to that of MWesoplodon sowerbyi,
as described by Mr. E. Ray Lankester’. In the larger anterior tooth the only remains
of the pulp-cavity is a small irregular vacuity (p), an inch below the apex of the tooth,
and consequently more than an inch and a half from the base. ‘The true dentine (d) is
limited to the portion of the tooth above this spot, the large bulk of the tooth below
being composed of very coarse-looking osteo-dentine with numerous wavy fissures and
channels, having a general longitudinal direction; immediately around the pulp-cavity
a tissue (7) having a botryoidal or globular arrangement forms a transition from the last-
named structure to the true dentine. There is no enamel. The surface of the tooth,
except at the apex and base, is covered with a layer of cement (c), which nowhere
exceeds 5/5 inch in thickness. The constituent elements of the smaller tooth are
arranged in a precisely similar manner.

Hyoid bones.—The basihyal and the thyro-hyals are not yet united (Pl. XXVIII.
fig. 9). The former is more elongated transversely than in Hyperoodon, being 5°8 inches
in width, and 2:7 inches in antero-posterior length at the middle. The posterior
border is straight; the anterior border excavated in the middle, and with a roughened
prominence near each end for the attachment of the anterior cornua.

The thyro-hyals are each 9 inches long, and 2°5 inches in greatest diameter. Their
under surface is flattened. They are less wide in proportion to their length than in
Hyperoodon, and thus, as in so many other details of the skeleton, they show that Berar-
dius recedes further from Physeter than does that genus’, and consequently approaches
nearer to the ordinary Dolphins.

The stylo-hyals are 14 inches long, and 2°9 inches in greatest thickness, slightly
curved, and with three distinct surfaces bounded by three longitudinal ridges, the
broadest and flattest surface being in the concave side of the bone. Towards what
appears to be the upper end there is a neck-like constriction, surmounted by an
expanded and obliquely truncated head.

Vertebral Column.—The numbers of the vertebre are:—cervical 7, thoracic 10,

' See the remarkable observations upon the teeth of the animal when alive, recorded by Dr. Haast (p. 214),
and the mention of “ muscular bundles” by which they are moved by Dr. Hector (p. 216), which accord so
little with any thing hitherto known in mammalian anatomy, that further observations on this subject are
extremely desirable.

2 Transactions of the Royal Microscopical Society, new series, vol. xv. 1867, p. 55.

* The breadth and flatness of the thyro-hyals is eminently characteristic of both the genera of Physeterine,
Physeter and Kogia; and the affinity of the Ziphiine to them is in this respect only slightly marked, though
most so in Hyperoodon.

224 PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES.

lumbar 12, caudal 19; total 48. This appears to be very nearly the usual number in
Ziphius and Mesoplodon, and to exceed that of Hyperoodon by 3.

Though generally resembling that of the three allied genera, the column differs from
Mesoplodon sowerbyi (Van Beneden and Gervais, pl. 22) chiefly in the spines of the
anterior dorsal region being smaller, more pointed, and more recumbent—also in all
the other spines sloping more backwards, and being smaller in proportion to the body,
both in height and width from before backwards; thus in a middle lumbar vertebra,
the antero-posterior diameter of the spine in Sowerby’s Dolphin is about two thirds
the length of the body, while in Berardius it is little more than one third. 7. densi-
rostris and another as yet undetermined species in the Sydney Museum agree generally
with I. sowerbyi in these respects, judging from photographs of their skeletons sent by
Mr. Krefft. Ziphius australis appears in Burmeister’s figure to be rather intermediate
between Mesoplodon and Berardius in the form of the spines of the vertebre. In
Hyperoodon the spines are as long as in Mesoplodon and as slender as in Berardius.

Cervical Vertebre.—The vertebre of the neck (Pl. XX VIII.), especially those of the
posterior part of the region, are better developed than in most Dolphins, and, indeed
(except for the coalescence of the first three), bear a considerable resemblance to those
of the Beluga. Among the Ziphiine, as far as is yet known, Mesoplodon and Ziphius
both approach Berardius in the structure of this important region, while, as will be
seen, Hyperoodon offers the greatest possible contrast.

The entire length of the bodies of the seven cervical vertebree, when placed in con-
tact, is 10 inches. The atlas, axis, and the third vertebra are united firmly by their
bodies; and the first two are also united by the lamine of the neural arches, but the
axis and the third only by that portion of the arch corresponding to the zygopophyses,
the upper part being free. All the remaining vertebre are separate throughout. The
free ends of their bodies show traces of the epiphyses by which they have been com-
pleted ; and their surfaces appear so completely formed or finished, as it were, that it
does not appear probable that much, if any, further union would have taken place had
the animal attained a greater age.

In Hyperoodon not only is the whole length of the cervical region scarcely more than
half that of Berardius, but the bodies of all seven vertebre are firmly united together,
and the spines of all, except the seventh, join to form a single elevated conical mass ;
the vertebre, except the first two and the last, are evidently extremely compressed,
almost rudimentary in fact. In Ziphius australis, according to Burmeister, the first,
second, and third are united, and also the fourth and fifth to each other, though not to
the third, and the sixth and seventh are free. In the specimen of Ziphius cavirostris,
from Corsica, briefly described in Fischer’s memoir, the six anterior yertebre are stated
to be united and the seventh free. In Sowerby’s Mesoplodon at Brussels the first two
only are united, and all the others free. In I. densirostris, according to Krefft’, “the

1 PZ. 8. 1870, p. 426.

PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES. 220

first three cervical vertebrae are anchylosed, the next one is more or less free, and the
remaining three are anchylosed again.” In another specimen of the same genus lately
added to the Sydney Museum, the first, second, and third are said to be united and the
remainder all free (Krefft, MS.).

The articular surfaces on the atlas for the occiput (fig. 2) are considerably smaller than
in Hyperoodon, and do not coalesce at their inferior margins. Above the upper end of
each there is a deep groove for the suboccipital nerve, instead of a foramen as in
Hyperoodon.

The coalesced spines of the first and second vertebree are but moderately developed,
and slope backwards, overhanging the short pointed spine of the third (see fig. 1). The
transverse process of the atlas is very little developed, and placed low on the sides of
the bone; it does not unite at its extremity with that of the axis as in Hyperoodon.
The inferior surface of the conjoined bodies of the first three vertebrz has a backward
projecting compressed tubercle, wanting in Hyperoodon, and probably representing that
so well developed in the Narwhal and Beluga.

The axis has two very short transverse processes on each side, compressed from before
backwards, both placed on the side of the body, the lower one corresponding serially
with the transverse process of the atlas, and being of about the same length.

The third has two distinct transverse processes on each side, further apart than those
of the second, the upper one arising from the upper part of the body and root of the
arch; the lower one is longer and narrower, and directed downwards and backwards.

Each of the three following vertebre (figs. 3, 4, and 5) have two transverse processes,
the upper ones (diapophyses) arising from the pedicle of the arch, rather slender,
conical, and inclined downwards, in the sixth also somewhat forwards; they increase
in size from the fourth to the sixth. The lower processes ( parapophyses) arise from the
inferior outer angle of the body, are thick and massive; that of the fourth vertebra is
most compressed and longest, while that of the sixth, though scarcely extending laterally
beyond the body of the vertebra, is greatly developed downwards, forwards, and inwards
(passing beneath the body of the antecedent vertebra), being, in fact, little more than
a great development of the inner basal tubercle of the others, and answering to the
“inferior lamella” of the transverse process of the corresponding vertebra of the
Carnivora, Ungulata, &c.’,

‘The seventh vertebra (fig. 6) has only the upper transverse process from the arch, the
inferior projecting edge of the well-marked articular surface for the head of the first rib
taking the place of the inferior process.

The bodies of all these vertebree are broader than they are high. The arches of the
fourth and fifth are incomplete in the middle line above for a space of more than half
an inch. The arches of the sixth and seventh are complete, and incline backwards,
but without any distinct spinous process.

1 See ‘Introduction to the Osteology of Mammalia,’ p. 22, 1870.

226 PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES.

The neural arches thus differ greatly from those of Hyperoodon, which has one tall
conical upright spine springing from the conjoined arches of the six anterior vertebrae,
and a second slender but equally elevated and vertical spine, belonging to the seventh
vertebra alone. On the other hand they resemble generally those of Ziphius and
Mesoplodon, as far as their condition in these genera is at present known; for the
incompleteness of the upper part of the arch exists in the fourth and fifth vertebree in
Ziphius australis, and in the third, fourth, and fifth in Mesoplodon sowerbyi. Perhaps
even more strikingly does the presence of well-developed inferior transverse processes
to the third, fourth, fifth, and sixth cervical vertebre show the affinity of Berardius to
these two genera and its dissimilarity to Hyperoodon.

Dimensions of the Cervical Vertebre.

inches.
Antero-posterior length of bodies of all seven, lower surface . . . 10
Length of body of united first, second, and third . . ... . 346
cencthvofbodyrot fourths Wier Wet ean PEN essen ree eee ee
Meng throtibedytofetitt ly sey swear eters Met crete eeeme Maneneeir, eee te, seem CA
Wenothtombodyiot sixthis site sme tt pie) CANADA ease Osea ee mitae de 6;
Length of body of seventh. . . Sid big hea i
Height from top of spine to lower sine of ae ai seseti ee wag
Height from top of spine to lower edge of body of seventh . . . 85
Breadth between outer borders of articular surfaces of atlas . . . 8:7
Height: ofiarticular'surfacesiof atlas’ 0) W112, Pim re. et Pee 528
Breadth between tips of transverse processes of atlas . . . . . 115
Height of neural canal in arch of atlas . see Set i778)
Greatest breadth of neural canal in arch of atlas. . . . . . . 3:8
Height of neural’canalsin-arch?of seventh’... ees se ee ord
Greatest breadth of neural canal in arch of seventh. . . . . . 36
Breadthsofibodytot seventh Ors i 1.-.aatrtsnis. terra) Peet. marae
Heighthtot bodyorseventhy 1. 5a ees ones SEE TAD
Breadth between tips of transverse processes of on ite sei Phd S105)

Thoracic Vertebre.—TVhe bodies of the twelve thoracic vertebre increase gradually in
length (see table on p. 228, and Pl. XXVIII. fig. 1). The inferior surface of those at
the commencement of this region is broad, flat, and somewhat rough ; posteriorly they
gradually acquire a median keel.

The spines are moderately high, compressed, sloping much backwards; that of the
first is very little developed, and pointed; though tapering in the anterior vertebrae,
they gradually become more obtuse and truncated at the extremity as they approach the
posterior end of the series.

The zygapophyses are unusually well developed, continuing on the contiguous anterior

PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES. 227

and posterior edges of the arch until between the eighth and ninth dorsal vertebre
inclusive, but not developed between the ninth and tenth.

Metapophyses first appear as distinct tubercles on the transverse processes of the
third, and gradually increase in size and become more compressed, pointing forwards
and slightly upwards.

Articular surfaces for the heads of the ribs are developed only on the hinder edges of
the bodies, without any corresponding surface on the anterior edge of the next vertebra,
so that the head of the rib appears not to articulate directly with the body of the same
vertebra to which the tubercle-is attached, but only to the one in front of it. In the
first vertebra this surface is entirely on the side of the body, in the second at the
junction of the body and the arch, from the third to the seventh at the root of the
pedicle of the arch ; on and after the eighth it is absent altogether, and the rib is attached
only to the transverse process.

The transverse processes, in a line with the upper transverse processes (diapophyses) of
the cervical region, are short and thick, with large rounded articular extremities for the
tubercles of the ribs. In the seventh vertebra this process is small, and in the eighth
reduced to a mere low longitudinal ridge on the outside of the metapophysis, which
has here acquired a considerable size. In the ninth vertebra a large and massive process
springs from the upper part of the side of the body near the anterior edge, in a situation
corresponding to which no trace of a process exists on any of the vertebre in front. It
has a large articular surface at its extremity, looking somewhat backwards, for the
ninth rib. The tenth vertebra bears a corresponding process, but rather longer, more
depressed, wider from before backwards, situated rather lower on the side of the body,
and not quite so near its anterior edge. Its articular surface (for the tenth rib), also
directed obliquely backwards, is not so large as that of the ninth. ‘This process corre-
sponds serially with the transverse processes of the lumbar vertebre.

Berardius thus conforms to the type of the Physeteride in the transverse processes of
the dorsal vertebre not gradually sinking from the arch to the body, as in the true
Dolphins, but disappearing near the end of the series, and being replaced by a new
process; but it differs from Physeter, and exactly agrees with Mesoplodon’, in not
having both upper and lower processes developed simultaneously on several of the
vertebre. Hyperoodon approaches nearer to Physeter in this characteristic feature, as
its seventh thoracic vertebra has distinct upper and lower transverse processes, which in
some specimens completely unite at their extremities, so as to form a ring, to the outer
edge of which the rib is attached.

The twelve lumbar vertebre are very much alike. ‘Their bodies increase in size
towards the hinder end of the series, where they are remarkably elongated. Below

1 Tn the skeleton of Sowerby’s Mesoplodon in the Brussels Museum, the upper process continues as far as
the seventh vertebra, and the lower process commences abruptly on the eighth.
VOL. VIII.—PaRT 11. September, 1872. 21

228 PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES.

the transverse processes they are compressed or pinched in, as it were, at the middle of
each side; and the inferior surface has a well-marked median keel.

The arches arise nearer the anterior than the posterior end of the bodies. The spines
are long, compressed, of nearly equal antero-posterior breadth from base to apex, the
edges being approximately parallel, and roundly truncated above; they slope backwards
forming an angle of 45° with the long axis of the body. The anterior end of the arch,
below the spine, develops a conspicuous, broad, flattened metapophysis, the lower edge of
which carries another tubercle. The metapophyses remain at the same level throughout
the series, instead of gradually rising on the sides of the arch in the posterior lumbar
and caudal vertebrae, as in Physeter and Orca.

The caudal vertebre, reckoning from the first which bears a chevron bone at its
hinder border, are nineteen in number.

The first resembles those of the lumbar region; but it is distinguished from them by
wanting the median keel on the inferior surface, and by the pair of articular facets on
its hinder edge for the first chevron bone. Its spine is also shorter and considerably
broader than in the last lumbar vertebra.

The bodies gradually shorten, though retaining their vertical height as far as the
ninth; the tenth is much compressed; the eleventh is small, being the ‘“ transitional
vertebra.” The series of depressed vertebree, or those of the tail proper, begins at the
twelfth; but they are less flattened and less transversely extended than in the true
Dolphins.

The under surfaces of the bodies are deeply channelled in the middle line, and have
strongly marked tuberosities at each corner for the attachment of the chevron bones;
the anterior and posterior tuberosities of each side become united in the seventh and
succeeding vertebra, enclosing a foramen which gradually diminishes in size.

The spines gradually decrease to the tenth, after which they are no longer developed.
The transverse processes also gradually diminish, and cease to be apparent after the
eighth. The last vertebra is a small depressed triangular nodule.

Antero-posterior length of the Bodies of the Thoracic, Lumbar,
and Caudal Vertebre.

inches. inches. inches. inches.
Thoracic: first .... 2°3| Lumbar: second .. 7-0 | Lumbar: twelfth .. 9-8} Caudal: tenth .... 5-6
second .. 2:7 third! 2) 22) Y7-leii@audallt) firstiacwen = 9-7 eleventh .. 4:1
third) 22.4 (3:5 fourthe. sence second 9-5 twelfth.... 3°5
fourth .. 4-0 fifth 5 YD third 5°9:0 thirteenth .. 3-1
fifth .... 4:3 sixthae. tod fourth atone) fourteenth.. 2-9
sixth .... 4:6 seventh .. 8:7 fittheaeerss 81 fifteenth .. 2:7
seventh .. 5:0 eighth .. 9-1 sixth aes sixteenth .. 2:2
eighth .. 5:5 Minthy yaa4 19'S seventh .. 7-5 seventeenth 1-8
ninth.... 6:0 tenthiees ont eighthie cc vel eighteenth.. 1:5
tenthieyane eos eleventh.. 9-8 ninth . 66 nineteenth.. 0-9
| Lumbar: first .... 6:4 i

PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES. 229

As noticed by Van Beneden' in Mesoplodon sowerbyi, the bases of the transverse
processes of the hinder lumbar and caudal vertebre are not perforated by vertical
vascular canals as in most Dolphins, but only the bodies of the posterior caudal vertebre
(beginning at the tenth) have such perforations.

There are nine chevron bones. ‘The first consists of two pieces, not united in
the middle line. The fourth is the largest. Their principal characteristic is that
their spines are not particularly elongated downwards, but are large from before
backwards.

The Ribs.—There are ten pairs of ribs, tolerably stout, especially thickening at their
lower ends, though somewhat contracted rather above the middle.

The first is short and broad, articulating above by a well-developed tubercle with the
transverse process of the first thoracic vertebra, and by a short capitular process with
the body of the seventh cervical. It has a strongly pronounced angle.

The second to the sixth gradually increase in length, and diminish in thickness;
afterwards they gradually become shorter. As far as the seventh inclusive, they have
all distinct tubercles articulating with the transverse processes, and well-developed
capitular process articulating with the hinder edge of the body or root of the arch of
the vertebra in front. After the fourth the angle becomes obscure and is lost in the
general convexity of the upper surface of the bone. In the eighth rib the tubercle is
rudimentary, and does not reach the much diminished upper transverse process of the
vertebra ; the head articulates with the hinder edge of the base of the pedicle of the
seventh vertebra, and on the right side only it has a small irregular articulation (not
found in any other case) with the anterior edge of the pedicle of the arch of the eighth
vertebra.

The ninth rib wants the whole of that portion representing the head, neck, and
tubercle of the ribs in front, and has a truncated upper extremity, articulating to the
end of the large transverse process of the ninth thoracic vertebra. The tenth rib is
similarly articulated with the corresponding process of the tenth vertebra, and is nearly
as well developed as the one in front of it.

The greatest length of each rib, in a straight line, is—

inches. inches. inches.
Burst yee ee USB bah oS Ga dommon ee AD oo) BET oh thwarts 38
Seconds Mercator Xo}. Mtiotde nn cao one anon SY) Vi INiitddh "SS 6605 50 aso 32
pind is Pricey ate or SAP Neventiieis sot cler fel ar. Soe Renthy sags ciel stars 29
Mounties a ne 37

As in Physeter, Hyperoodon, and the other known Ziphioids, there are no ossified
sternal ribs. ;
The Sternum.—The sternum (Pl. XX VII. fig. 3) is remarkably long and narrow. It

1 Loe. cit. p. 42,
22

230 PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES.

resembles generally the form of the same part in Ziphius and Mesoplodon, but is pro-
portionally longer than in Hyperoodon, and still more than in Physeter. The inferior
surface is slightly convex from before backwards, and also from side to side.

It consists of five distinct segments, not connected together by bone. The anterior
segment is the largest, and has a shallow fossa in the middle line in front; the posterior
segment ends in a pair of narrow xiphoid processes, of which the right is slightly larger
than the left.

As in the other Ziphioid Cetaceans, development is less complete along the middle
line of the sternum than at the sides; not only are there median notches at each
extremity, but there are three large median fenestra, one between the first and second
segments, one between the second and third, and one between the third and fourth seg-
ments. The edges of these fenestre are bevelled and smooth; so that it does not appear
that ossification would have advanced further in this direction if the animal had lived
to be older.

Each side of the sternum shows six rough articular facets for sternal ribs :—the first
near the anterior end of the first segment, at its broadest part; the second at the
junction of the first and second segments; the third at the junction of the second and
third segments; the fourth at the junction of the third and fourth segments; the fifth
near the hinder end of the fourth segment; the sixth on the side of the fifth segment,
near its anterior end.

The entire length of the sternum in a straight line is 45 inches. The greatest breadth
of the first segment is 12°8 inches, of the second segment 10-9 inches, of the third
10 inches, of the fourth 9 inches, of the fifth 8-8 inches.

Pectoral Limb.—The pectoral limb, as in all other members of the group to which
Berardius belongs, is small in proportion to the size of the animal.

The scapula bears a considerable resemblance to that of Hyperoodon ; it is less narrow
and elongated than in Physeter, and more regularly triangular, and less fan-shaped
than in the ordinary Dolphins. The external surface is smooth and slightly concave,
with one vertical ridge near the hinder border. The acromion is large and flat, curving
regularly inwards, and somewhat upwards towards the extremity. The coracoid is
rather longer, and more slender and rounded than the acromion. The glenoid fossa is
irregularly oval, the outer edge more convex than the inner, and the anterior extremity
more pointed than the posterior.

The humerus and bones of the forearm are elongated, slender, and simple in
character; the latter more resemble the corresponding bones of a Balewnoptera than
of one of the Delphinide.

The humerus has a rounded head and single tuberosity, formed by a large and still
free epiphysis. The lower epiphysis includes little more than the articular surface,
and is united to the shaft, though the line of separation is distinct in many points.
The radial border of the bone presents, near the middle of its length, a low rounded

PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES. 231

protuberance, representing the deltoid ridge, with a shallow groove below it. Rather
higher than this, just below the commencement of the expansion for the head, on the
outer surface of the bone, is a roughened shallow depression. ‘The articular surface
for the ulna encroaches for a space of 14 inch on the corresponding margin of the
humerus.

The radius is a simple narrow flattened bone of almost equal breadth throughout.

The ulna has more pronounced characters, having a very well-developed, broad, and
flattened olecranon, with a strongly marked groove in the middle of the outer side of
its upper border, apparently for the passage of a tendon, and which I have not observed
in any other Cetacean. The bone is considerably contracted in its upper third, and
expands moderately towards its lower extremity.

Both radius and ulna have well-marked epiphyses at either extremity, including the
whole of the articular surfaces: that of the upper end of the ulna“is united to the
shaft ; but the other three are still separate.

The carpus (Pl. XXVIII. fig. 10) presents some unusual features in the mode of union
of its several elements with each other. All the bones which are met with in Hyperoodon
and the other Ziphioids appear to be present and well ossified. There is thus one bone
more (viz. the magnum) than in the ordinary Dolphins and in the Sperm-Whale.

The scaphoid (s) and the lunar (/) have almost completely united, though a groove
shows their original limits. ‘The cuneiform (c), which is the largest bone of the carpus,
is united at its outer edge with the unciform (w); and a long bone to the distal side of
which the second and third metacarpals are articulated, represents the magnum (m) and
the trapezoid (td) coalesced as in the skeleton of Mesoplodon sowerbyi at Brussels.
The bone, of the distal row, most to the radial side might well be considered the
trapezium; but it represents the bone which, in other Cetacea, has been considered,
with good grounds, to be the first metacarpal. Excluding this, the number of carpal
bones are reduced by ankylosis to four. The arrangement on the two sides is precisely
similar.

The second and third digits are of nearly equal length, the fourth (JV) not much
shorter; the fifth (V) is well developed and stands considerably apart from the others ;
the first (2) is very small and adpressed. The phalanges were artificially articulated
before they came under my observation; and some of the smaller terminal ones are
probably missing; but the numbers present are, including the metacarpals, in the first
digit 2, in the second 5, in the third 4, in the fourth 4, in the fifth 3.

Dimensions of the Bones of the Right Pectoral Limb.

inches.
Scapula.—Height from middle of superior border to middle of external

MArgiINOL Slenold Hossa ely. a. Tat! Si et ay ath ie cane
Length of anterior border, from anterior superior angle to
anterior margin of glenoid fossa . . . . . . . 15

232 PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES.
inches.
Scapula—tLength of posterior border from posterior superior angle to

posterior margin of glenoid fossa . . . . . . . . . 12
Breadth from anterior to posterior es angle’) =.) jut an aloes
Length of acromion . . . RAPS ; 8a NG
Vertical height of acromion at the narrowest or (ateai itsroot) 2°8
Vertical height at the broadest part (nearitsend) . . . . 41
Length or coracoid: processtt =!) EOP RE) ae fein ra HOPS
Length of glerioid fossa) S100) 2. ROSE cake ees iin GA
Breadth ‘ofigl énoid fossa‘! 7) 4 2 ROD ee Rae ees
Humerus.—Length. . . . RANA. “CURSE CARA BEL OES
Breadth at lower end RPO al Pe ee ee gi eS
Radius: —Extreme! lensth y 77" 226.1, VER Ene) TEI: ES DLO
Breadth af wpperiendhs posi" OS eR De ae oso
Breadth‘at-middle! 070.192 sia RiG | NEE Ge Pa Bato aE
Breadth*attlower nd ®. Sew Pas ENS See AP SOS ra eee
Ulna.—Extreme length, including olecranon . . . . . . . . . 127
Breadth at upper part, including olecranon . . . . . . . 56
Breadthtat marrowestaparts J) Saiimace oe) © eo teuekl = | ¢sSec ute ane ee,
Breadth atelower end J'eia, a. Le B/C Rae eal
Manus.—Length, allowing for loss of terminal Tee, abou Lie el a
ibenethrof firstametacarpale AR setes) Saks ence eects aap GO
Weng th of'second metacarpal iw giavey a ics Gent © eee iste OO
hengthof third’metacarpalls4”. se) ee oc0 cpus Mota jeer)
Itengthiofifourthsmetacarpal ue 8. lee) le. iat tee bat in ume OSO
Length of fifth metacarpal . . . . 3:2

Pelvic Bones.—The one pelvic bone sent with the seals is very light ae spongy
in texture, and is apparently not completely ossified at the extremities. Its length is
5:8 inches, and its greatest thickness 0:9 inch; it is thus small in relation to the
general size of the skeleton, and would indicate that the animal was a female, if the
same sexual proportions obtain in Berardius as in many other Cetaceans. It is of very
simple form, subcylindrical, a little compressed, and with a slight bend at one third
of the length from one end, causing a low obtuse angular prominence on one edge.
The characters and dimensions of the bone would probably have been somewhat
different if the age of the animal had been more advanced.

Conclusion—The special osteological characteristics of Berardius will be more fully
appreciated when our knowledge of the skeletons of the other Ziphioids, especially of
the genera Mesoplodon and Ziphius, is more perfect. We can hardly go wrong,
however, in affirming that, as far as the structure of the skeleton is concerned,
Berardius is a peculiar form of the group, and that it occupies one end of the series
of which- Hyperoodon forms the other, the remaining Ziphioids being in a certain

PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES. 233

sense middle forms, and that Berardius is, on the whole, the least modified or
specialized form, and approaches, therefore, nearer to the true Dolphins, while Hype-
roodon is the most specialized, being modified in the direction of the Physeterine or
Sperm- Whales.

The principal points still to be ascertained with reference to the skeleton of Berardius
are :—1. Whether in the perfectly adult animal any further ankylosis of the cervical
vertebre takes place. 2. What is the number of the phalanges of a perfectly com-
plete skeleton. 3. The form and size of the pelvic bones in the adult of both sexes.
4. Whether there is any difference in the development or form of the teeth in the two
sexes. 5. Whether there are any rudimentary, non-alveolar teeth concealed in the
gum of either the upper or the lower jaw.

Any one who may be fortunate enough to see the animal in a recent state should
observe whether the cutaneous furrows on the throat, noticed in all other Ziphioids
hitherto examined, are present; and it need scarcely be added that all observations on
the visceral anatomy as well as any facts bearing upon the geographical distribution or
habits of this rare and interesting Cetacean will be of great value in elucidating its
history and determining its true place in the natural system.

DESCRIPTION OF THE PLATES.

PLATE XXVII.

Fig. 1. Side view of the skeleton of Berardius arnouai.
Fig. 2. The skull and vertebral column of the same, seen from above.
Fig. 3. The sternum, seen from below.

All one sixteenth of the natural size.

PLATE XXVIII.

Fig. 1. Side view of the cervical, dorsal, and two anterior lumbar vertebre. The lines
from the Roman numerals below the figure point to the surfaces for the
attachment of the ribs.

Fig. 2. Anterior surface of the atlas.

Fig. 5. Anterior surface of the fourth cervical vertebra.
Fig. 4. Anterior surface of the fifth cervical vertebra.
Fig. 5. Anterior surface of the sixth cervical vertebra.
Fig. 6. Anterior surface of the seventh cervical vertebra.
Fig. 7. Vertical median section of the skull.

Mz, maxilla; PMza, premaxilla; Vo, vomer; M/E, mesethmoid, united with the frontal
above and the presphenoid below; Na, nasal; Fr, a number of loose wedge-shaped

234 PROFESSOR FLOWER ON THE RECENT ZIPHIOID WHALES.

fragments of bone, apparently dismemberments of the frontal; SO, supraoccipital ;
BO, basioccipital ; BS, basisphenoid ; PS, presphenoid ; Pt, pterygoid.
Fig. 8. Inner surface of the right ramus of the lower jaw.
Fig. 9. Hyoid bones. BH, basihyoid; 7H, thyro-hyoid.
Fig. 10. Dorsal surface of the left manus, some of the terminal phalanges missing.
R, radius; U, ulna; s, scaphoid; 7, lunar; ¢, cuneiform; td, trapezoid;
m, magnum ; wv, unciform; J to V, the five digits.

All one sixth of the natural size.

PLATE XXIX.

. United right periotic and tympanic bones of Berardius arnouat, outer surface.

. The same bones, under surface. :

The periotic bone, upper surface.

The periotic bone, under surface.

. The tympanic bone, upper surface.

. The united periotic and tympanic bones of Hyperoodon rostratus, outer surface.
From a specimen in the Museum of the University of Cambridge.

7
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Letters to all the above figures :—per, periotic; ty, tympanic; a, principal or posterior
articular surface between these bones; a’, broken surface where the two bones had
been united; e.a.m, external auditory meatus, closed in life by the membrana tympani ;
e.p.l, external posterior lobe of the tympanic; @.p./, internal posterior lobe of the
tympanic; e, groove for the Eustachian tube; m, posterior bony process corre-
sponding in its relations with the mastoid of other mammals, but united with the
tympanic; 7.a.m, internal auditory meatus; a.f, aqueduct of Fallopius; f.0, fenestra
ovalis; fir, fenestra rotunda; a@./, anterior lobe of the periotic; m./, its middle
lobe ; p.l, its posterior lobe.

Fig. 7. Median vertical antero-posterior section of the anterior tooth of the right side.

é, cementum; d, dentine; g, globular dentine; p, remains of pulp-cavity.

Fig. 8. Corresponding section of the posterior tooth of the same side.

All of the natural size.

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oS z c z ee z =e S =
wo S o = o S o = wy
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Sal = Se 3 = pe 2 - Fa
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z - = & : Zz Ni = cS Zz
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a ar e ; L = a a i
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