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SYoxt Socrwry,’ vol. x. part ix. 1878. | 4
XI. A further Contribution to the Knowledge of the existing Ziphioid Whales. Genus
Mesoplodon. By WituiaM Henry Fiower, F7.RS., V.P.Z.S.
Received July 31st, 1877. Read November 6th, 1877.
{PLates LX XI.-LXXIILI.]
DURING the six years that have elapsed since I communicated to the Society a
memoir on the recent Ziphioid Whales (Trans. Zool. Soc. vol. viii. p. 203) very consi-
derable additions have been made to our knowledge of the group. Instead of being
so rare as was then supposed, since the attention of naturalists resident in our colonies
has been directed to the importance of losing no opportunity of securing such specimens
as accidents of wind and waves may cast upon their shores, it has been proved that in
the seas of the southern hemisphere these Whales exist in considerable numbers both as
species and as individuals, and that one species at least is gregarious, having been met
with in two instances in “ schools”’ of considerable numbers.
On the other hand, no remarkable deviations from the forms already known have
been met with; and all additional information as to their osteology has fully confirmed
the value of the division of the Ziphioid Whales into four distinct types or genera, at
that time indicated. With perhaps one exception (to be noticed further on), which
presents some signs of transition, all the known individuals lately discovered can be
arranged without any hesitation either as Hyperoodon, Berardius, Ziphius, or Meso-
plodon, as defined in the previous memoir.
It is in our knowledge of the animals of the last-named genus that the greatest
advances have been made of late; and it is to these that the present communication
will chiefly relate.
After examining all the available specimens and published descriptions, I have
arrived at the conclusion that evidence exists at present of six distinct specific modifica-
tions of this form, of which four inhabit the southern temperate seas. To these I shall
have to add two more, although in neither case on evidence so satisfactory as might be
desired. The synonymy and habitats of the hitherto known species are as follows :—
1. M. Bipens (Sowerby).
Physeter bidens, Sowerby, Brit. Miscellany, p. 1 (1804).
Delphinus (Heterodon) sowerbiensis, Blainville, Nouv. Dict. d’Hist. Nat. 2nd ed. tome ix. p. 177
(1817).
D. sowerbyi, Desmarest, Mammalogie, p. 521 (1822).
Delphinorhynchus micropterus, Cuvier, Régne Animal, t. i. p. 288 (1829) ; Dumortier, Mém. Acad.
Roy. Bruxelles, t. xi. (1839).
VOL. X.-—PART 1X. No. 2.— August 1st, 1878. 3 L
416 PROF. W. H. FLOWER ON THE GENUS MESOPLODON.
Mesoplodon sowerbiensis, Van Beneden, Mém. Acad. Belgique, coll. im 8vo, t. xvi. (1863).
M. sowerbensis, Gervais and Van Beneden, Ostéographie des Cétacés, p. 892 (1877).
Micropteron bidens, Malm, “ Hvaldjur i Sveriges Museer ar 1869,” K. Svenska Vetenskaps-Akad.
Handl. 1871.
Hab. Coast of Elginshire (Sowerby) ; Havre (Blatnville); Ostend (Dumortier); Sale-
nelles, Calvados (H. Deslongchamps); Brandon Bay, Ireland (Andrews); Norway
(Malm).
2. M. EUROP&US (Gervais).
Dioplodon europeus, Gervais, Zool. et Pal. francaises, le édit. t. 11. Explic. No. 40 (1850).
D. gervaisit, E. Deslongchamps, Bull. de la Société Linnéenne de Normandie, tome x. (1866).
A single specimen found floating in the sea at the entrance of the British Channel
about 1840. ‘The skull is now in the Museum at Caen. It should be mentioned that
some doubts have been thrown upon the supposed distinctive characters between this
and the last species.
3. M. DENSIROSTRIS (Blainville).
Delphinus densirostris, Blainville, Nouy. Dict. d’Histoire Nat. 2nd edit. tome ix. p. 178(1817).
Mesodiodon densirostris, Duvernoy, Annales des Sciences Naturelles, sér. 3, t.xv. p. 59 (1851).
Ziphius sechellensis, Gray, Zool. E. & T. p. 28 (1846) ; Krefft, P. Z. S. 1870, p. 426.
Dioplodon densirostris, Gervais, Zool. et Pal. frangaises, 1°¢ édit. t. ii. Explic. No. 40 (1850) ; Ann.
des Sciences Nat. sér. 3, t. xiv. p. 16 (1850).
Hab. Seychelles Islands (Duvernoy); South Africa (Mus. R. Coll. Surgeons); Lord
Howe’s Island (Arefft).
4. M. Layarpi (Gray).
Ziphius layardi, Gray, P. Z. S. 1865, p. 358; Owen, Crag Cetacea, p. 12, pl. i. (1870).
Ziphius (Dolichodon) layardi, Gray, Cat. Seals and Whales Brit. Mus. p. 353 (1866).
Dolichodon layardii, Hector, Trans. New-Zealand Inst. vol. v. p. 166, pl. iii. (1872).
Mesoplodon longirostris, Krefft, MS., M. giintheri, Krefft, MS., and Callidon giintheri, Gray,
Ann. Mag. Nat. Hist. ser. 4, vii. p. 368 (1871).
Dolichodon traversit, Gray, Trans. New-Zealand Inst. vol. vi. p. 96 (1874).
Mesoplodon floweri, Haast, P. Z.S. 1876, p. 478.
Hab. Cape of Good Hope (Layard) ; Near Sydney (Krefft) ; New Zealand (Haast) ;
Chatham Islands (Hector).
Two skulls, an adult and a young specimen, of this well-established species were
brought home by the naturalists of the ‘ Challenger,’ and will be described in detail by
Professor Turner in his Report on the Marine Mammalia collected during the expedition.
5. M. HEcTORI (Gray).
Berardius arnuxii, Hector, Trans. New-Zeal. Inst. vol. ii. p. 27 (1870).
Smaller Ziphioid Whale, Knox & Hector, T. New-Zeal. Inst. vol. iii. p. 125, pls. xiii., xiv. & xv. (1871).
PROF. W. H. FLOWER ON THE GENUS MESOPLODON, AIT
erardius hectori, Gray, Annals & Mag. N. H. ser. 4, vol. viii. p. 117 (1871).
Mesoplodon knoxi, Hector, Trans. New-Zeal. Inst. vol. v. p. 167 (1873).
Hab. Titai Bay, New Zealand. Very young; length 9! 3", 1866 (nox and Hector).
6. M. erayi, Haast, Proc. Zool. Soc. 1876, p. 7.
Oulodon grayt, Haast, P. Z. 8. 1876, p. 457°.
Hab. New Zealand.
These six species do not differ greatly in size, and present a close family or, rather,
generic resemblance ; but still there are certain cranial and dental characters by which
the skulls can readily be distinguished.
The most easily recdgnized of these are :—
a. In some species there is a deep lateral longitudinal groove at the base of the
rostrum, commencing posteriorly in a blind pit below the tubercle of the maxilla
situated in front of the anteorbital notch, and bounded above and below by sharply
defined prominent ridges, both formed by the maxilla. This groove characterizes
M. grayi and M. densirostris. No trace of this groove, but a prominent ridge instead,
which gives a wide base to the rostrum, as seen from above or below, exists in
WM. bidens, M. ewropeus, and M. hectori. The groove is slightly developed in 1. layardi,
which in this respect presents an intermediate condition.
b. The relative position of the foramina for the exit of the facial branches of the
second division of the fifth nerve appears to afford constant distinctive characters between
certain of the species. The principal foramina are in two pairs:—1, the larger or more
external, situated in the maxilla, and often double, corresponding to the infraorbital
foramen in man (Plate LXXI. mf); and, 2, the smaller and nearer the middle line,
situated in the preemaxilla (pf). The latter are rather in front of, or on the same level
with, the maxillary foramina in WV. bidens, M. europeus, M. hectori, and M. layardi, and
placed decidedly on a posterior level in MW. denstrostris and M. gray.
c. The constant presence of a row of small teeth in the upper jaw is said to dis-
tinguish WZ. grayi from the other species, though, as will be discussed presently, this
can scarcely be considered of generic importance.
d. The position of the large tooth in the lower jaw, whether close to the apex of the
' Tn the notice of this paper in the Proceedings of the Zoological Society, 1877, p. 684, for
Te IE Vane obo Oe present memoir,” read :—
7. MU. haasti, n. sp.
Hab. New Zealand.
Known only by a portion of a cranium in the Museum of the Royal College of Surgeons.
8. UM. australis, n. sp. = M. hectori (Gray). Hector, Trans. New-Zealand Inst. vol. vi. p. 86, and yol. yu.
p. 362.
Hab. New Zealand.
A complete skeleton, now in the British Museum, described in the present memoir.
418 PROF. W. H. FLOWER ON THE GENUS MESOPLODON.
mandible, as in J. hectori, or near the hinder part of the symphysis, as in all the
others, forms another good distinctive character.
e. Finally, the character and form of this tooth presents some important variations.
In MW. layardi, its apex is everted, and seated upon a flat strap-like base, which grows
upwards, backwards, and finally inwards, closing over the upper jaw and meeting its
fellow of the opposite side. In IZ. densirostris the apex is directed vertically, placed
on a very massive base, which is implanted in a greatly expanded alveolar margin of
the jaw, not found in any of the other species. In MJ. grayi the tooth is triangular
and compressed, and the apex is vertical at all ages. In I. hectori the apex of the
tooth in the young animal is directed forwards. In J. bidens the apex of the tooth is
at first directed backwards, but during growth becomes rotated forwards. In the only
specimen of WM. ewropeus known (an adult) the apex of the tooth is directed somewhat
forwards.
These characters may be thus placed in a synoptical form :—
Lateral Premaxillary | ve
basirostral foramen to Position of Other dental characters.
groove. maxillary. mandibular tooth.
Absent. Level. Close; to Vapexotg go. -useci yack wie neta. | M. hectori.
mandible.
Absent. Level. | DMamtabiatimemes)| bao saeomncedo ods cou M. europeus.
dibular sym-
physis.
Absent. Level. Near hinder edge | Apex of tooth at first di- | IW. bédens.
of mandibular rected backwards, and }
symphysis. during grow rotated
forwards.
Slight. Level. Ditto. Apex of tooth M. layardi.
mounted |
base, which increases Tx. |
length with age, and |
finally curves round the
maxilla. a |
Deep. Behind. Ditto. Apex of tooth vertical. Al- | M. densirostris. |
veolar portion of jaw , |
expanded. I
Deep. Behind. Ditto. Apex of tooth vertical. A | MW. grayi. |
row of small teeth in |
upper jaw.
Besides the above, there are many minor differences, more appreciable by a com-
parison of the specimens than by description, some of which will be pointed out in
the sequel.
The special materials upon which the present communication is based are :—
1. The skull of a young specimen of I. hectori, the type and at present only known
example of that species. It was taken in Titai Bay, Cook’s Strait, January 1866 ; and
the skull is figured by Dr. Hector in the Trans. New-Zeal. Inst. vol. ii. 1870, pls. xiv.
PROF. W. H. FLOWER ON THE GENUS MESOPLODON. 419
& xv. The length of the animal is stated by Dr. Knox to have been 9 feet 3 inches.
The skull is now in the British Museum.
2. The skeleton of a perfectly adult animal, attributed by Dr. Hector to the same
species, prepared from a specimen cast ashore in Lyall Bay, and described by
Dr. Hector in Trans. New-Zeal. Inst. vol. vii. p. 262, 1875. This is also in the
British Museum !.
3. The skeleton of an adolescent male JZ. grayi, stranded, with others, on the coast
near Saltwater Creek, about thirty miles north of Banks’s Peninsula, New Zealand.
4. The rostrum, mandible, and mandibular teeth of an old male animal, stranded
in December 1875 on the east coast of the North Island, attributed by Dr. v. Haast to
the same species.
The last two were presented to the Museum of the Royal College of Surgeons by
Dr. Julius y. Haast, F.R.S., of Christchurch, New Zealand.
Of the specific distinction or identity of the 2nd, 3rd, and 4th I am not perfectly
satisfied, as the materials at hand are unfortunately not sufficiently complete for the
purpose of arriving at a definite conclusion.
No. 3, though a perfect skeleton, is still immature. No. 2, though adult, is incom-
plete as regards the very important evidence furnished by the dentition. No. 4 is
only a fragment, and, although identified by Dr. v. Haast with his species WZ. grayt,
presents strongly marked differential characters which seem beyond the range of
individual variation. Under the circumstances, it is somewhat difficult to know what
course to pursue with reference to the names by which these specimens are to be
respectively distinguished; but on the whole it will lead to less confusion if I designate
them, provisionally at least, by specific appellations, bearing in mind that it is quite
possible that further information and more abundant materials may cause a modi-
fication of this view. I shall therefore, in the present memoir, speak of the British-
Museum skeleton from Lyall Bay as WZ. australis, and the fragmentary skull from the
east coast of the North Island as MW. haasti.
Skull.—The skull of animals of the genus Mesoplodon is very easily distinguished
from those of the other Ziphioids by the characters previously given (T. Z. 8. vol. viii.
p. 208). The ossification of the mesethmoid cartilage, and its coalescence with the
surrounding bones to form a solid rostrum, appear to be greatly dependent upon
the influence of age. It has not hitherto been found wanting in any thoroughly adult
example of any species of Mesoplodon or of Ziphius; on the other hand, it appears
never to occur either in Hyperoodon or Berardius.
* T am indebted to Dr. Giinther’s kindness for facilities in examining and comparing these specimens, which
were sent to the British Museum by Dr. Hector, F.R.S., of Wellington, New Zealand.
* Almost simultaneously with my memoir appeared an excellent description of the skull of M. sowerbyi, com-
pared with that of Zphius cavirostris, by Professor Turner (Transactions of the Royal Society of Edinburgh,
vol. xxvi.).
420 PROF. W. H. FLOWER ON THE GENUS MESOPLODON.
The tympanic bone of Mesoplodon is quite different from that of Ziphius', the groove
between the lobes at the posterior end being very well marked as in the true Dolphins,
whereas in Ziphius it is obsolete. In Hyperoodon also it is scarcely apparent, while in
Berardius it is nearly as well marked as in Mesoplodon. ‘Thus, by the form of this very
characteristic bone, Ziphius allies itself to Hyperoodon, and Mesoplodon to Berardius,
and the two former approach nearest to Physeter, and the latter to the less-modified
Dolphins.
The relationship between Berardius and Mesoplodon is undoubtedly close. One of
the specimens (JZ. hector’) now under consideration has been placed sometimes in one
genus and sometimes in the other; and it does certainly present some transitional cha-
racters; but as it is only known by the skull of a very young individual, it is scarcely
safe to decide its position, except provisionally, especially as it is not yet known in
which direction the alterations which must take place during the progress towards
maturity tend.
The animal stranded at Lyall Bay, which I have called UM. australis, is perfectly
adult, as shown not only by the union of the sutures at the base of the cranium,
but also by the condition of the other bones of the skeleton, the terminal epiphyses
of the bodies, even of the thoracic vertebre, being completely consolidated. The
mesorostral or mesethmoid cartilage is densely ossified throughout its whole length.
There is still much to be learned with regard to the mode of ossification of this cartilage.
All the specimens which I have had an opportunity of examining are either so young
that ossification has not commenced, and the trough of the vomer in the rostrum
proper is completely empty in the dried skull, or so old that the consolidation of the
cartilage and its union with the surrounding bone has been completed. But it must
be observed that, although the cartilage appears to be nothing more than a con-
tinuation forwards of the ordinary mesethmoid lamina or septum of the nose, the
ossification is not a simple extension forwards of that which occurs in all Cetacea (in
all Mammalia, in fact) in the hinder or internarial portion of the septum, but appears
to be an independent production, peculiar to the genera Mesoplodon, Ziphius, and certain
allied extinct forms. It is separated by an interval (which appears to diminish with age,
but of which traces can always be seen on the upper surface of the rostrum near its
base) from the true mesethmoid ossification. It differs from the latter in being
intensely hard and compact, whereas the mesethmoid is, especially at its anterior part,
somewhat spongy in texture. It differs also in showing strong indications of being
formed by a pair of lateral ossifications, united in the middle line, as the upper surface
in many parts and the anterior apex show a marked median groove. I think it will
be well therefore to adopt Professor Turner’s name of “ mesorostral” bone for this
solid bar forming the centre of the rostrum, restricting ‘ mesethmoid” to the part
lying between the nares and a short distance in front of them, which is ossified in the
* Figured by Prof. Turner, loc. cit. plate xxx.
PROF. W. H. FLOWER ON THE GENUS MESOPLODON. 421
young animal and in all other species of Cetavea. The median ossicle in the snout
of the Pig offers a similar case of an independent ossification in the anterior extremity
of the mesethmoid cartilage.
The skull, as seen from above (Plate LXXI. fig. 1), presents several characters by
which it can be readily distinguished from J. didens and the closely allied MW. ewropeus.
The principal of these is the sudden narrowing of its outline at the commencement
of the rostrum, due to the presence of the deep lateral groove mentioned above. In
the British species the rostrum is wider at the base, and passes more insensibly into the
cranium. The next point is the relative position of the premaxillary (pf) and the maxil-
lary (mf) infraorbital foramina—the former being behind the latter in the southern,
and on the same level in the northern species. The third is the form of the premaxille,
which are more expanded and flattened out (especially the right) immediately in front
of the narial apertures in I/. australis. In the conformation of the upper ends of
the premaxille and of the nasal bones the differences are but slight.
MM. layardi, as mentioned above, agrees with the northern species in the disposition
of the infraorbital foramina, and the absence of any marked lateral basirostral groove ;
and by these as well as by its very characteristic dental characters it is distinguished
from the one now under consideration. J. densirostris is much more closely allied to
our new species, having (judging by the figure in Van Beneden and Gervais’s Ostéo-
graphie, pl. xxv.) a very similar outline as seen from above. It differs, however,
independently of the mandibular and dental characters, in the far more massive rostrum,
which, although no longer, has exactly double the vertical height at the middle, as seen
in the profile view of the skull.
A marked difference is also seen on the lower surface of the skull. In I. densi-
rostris the palate-bones completely surround the anterior pointed ends of the pterygoids,
widely separating the latter from the maxillaries; while in I. australis the palatines
on the inferior surface of the skull lie altogether on the outer side of the pterygoids,
and do not eyen extend so far forwards as the pointed extremities of those bones,
which thus by their free end and inner side come into direct contact with the
maxillaries.
It remains only to compare this skull with the other New-Zealand forms—the
typical IZ. grayi of Haast, the rostrum which I propose to call M. haasti, and MW. hectorv.
With the former it has very near affinities—so much so, that from an examination
of the cranium alone I should scarcely think of placing them in different species.
It is needless to dwell upon differential characters which may be merely individual ;
but I may indicate the following as perhaps of more importance. a@. Size: MV. australis
would appear to be smaller than WZ. grayi. The two complete skulls are practically of the
same size; but that of the former is adult and that of the latter young, and the tympanic
bone of the latter is slightly larger than that of the former. Dr. Haast gives 17' 6"
as the length of an adult female of WV. grayi, whereas the adult MW. australis (sex not
429 PROF. W. H. FLOWER ON THE GENUS MESOPLODON.
stated) is given by Dr. Hector as 15! 6"; this difference, however, can scarcely be con-
sidered beyond the limits of individual variation. 6. The form of the tympanic bone
differs slightly, the inner side of the posterior edge (as seen from below) being more
bevelled and the inner posterior lobe less prominent in J. australis. ec. The lateral
groove at the base of the rostrum is very much deeper in JV. australis, running up-
wards into the maxillary bone, and forming a blind pit at its upper termination, of the
depth of nearly one inch, whereas in JZ. grayi this pit is quite shallow. It is possible,
however, that this may be a question of age.
Mesoplodon australis (adult). Mesoplodon grayi (young).
Mesoplodon haasti (adult). Mesoplodon hectori (young).
Outlines of section of middle of rostrum. Natural size.
While the section of the rostrum of both the above (figs. 1 & 2) agree very closely
in general outline and size, that of the supposed I. haasti (fig. 3) differs very greatly,
being altogether more compressed and wanting the lateral wing-like ridges near the
lower border ; its length is also considerably greater, being 223’, while that of the adult
M. australis is only 18". Making every allowance for individual variation, it scarcely
seems possible that a rostrum such as that shown in fig. 2 could change in the course
PROF. W. H. FLOWER ON THE GENUS MESOPLODON. 423,
of growth to that in fig. 3. If so, most of the determinations of the fossil species based
solely upon the form of the rostrum are quite valueless.
It should be mentioned that in this fragmentary rostrum the lateral groove and pit
at its base are deeper even than in MV. australis, and that it departs, therefore, in this
character, still further from (the young) M. grayi.
The cranium of MW. hectori, from Titai Bay, New Zealand, now in the British
Museum, presents a conformation quite distinct from that of either of the specimens
last under consideration. As is shown in the synoptical table at p. 418, it more nearly
resembles the northern species in the relation of the maxillary foramina, the absence of
the basirostral groove, and, it may be added, the correlated greater breadth of the base
of the rostrum. The skull is evidently that of a very young animal: the teeth are still
concealed beneath the gum; the symphysis of the lower jaw is not united; the meso-
rostral cartilage is entirely unossified; the basisphenoidal suture is open (as seen
from the interior of the cranium; for on the outer surface it is covered by the vomer) ;
and the bones generally are very spongy and greasy. On the other hand, the elements
of the occipital bone have all coalesced with each other and with the basisphenoid.
The principal difference between this cranium and that of MZ. bidens and, in fact, of all
others of the genus, is that the upper extremities of the preemaxille are less developed
laterally and less everted, and therefore approximate more to the form of Berardius ;
but unfortunately a large piece of one of these bones is broken off from the right side,
or that which is usually most strongly marked, and it is impossible to say how much of
the absence of characteristic eversion may be due to the immaturity of the specimen.
There is also a greater elevation of the longitudinal ridge on the upper surface of the
maxilla, immediately to the inner side of the anteorbital notch, than in any other
Mesoplodon. This ridge, it should be remarked, forms a strongly pronounced elevation
in Berardius, and is developed in Hyperoodon into the immense bony mass which forms
such a characteristic feature of the skull of that genus. The nasals, however, are com-
pletely sunk between the ends of the premaxille, whereas in Berardius they form
prominent masses rising to the vertex.
The palate-bones only appear on the palatal surface as narrow strips on the outside
of the pterygoids, not reaching as far forwards as they do. The vomer is visible in
the middle third of the inferior surface of the rostrum.
The mastoids and squamosals are quite free; and the former are largely developed,
forming the principal part of the great post-tympanic processes of the skull.
The tympanic is very slightly larger than that of UW. australis, and therefore equal
to that of IV. grayi, and presents a very close resemblance to both, but is nearer in the
form of the lobes at the posterior part to the former. I have not had an opportunity
of comparing it with the tympanic of J. bidens. As the tympanic bone of Berardius is
constructed on exactly the same type as that of MWesoplodon, this bone gives no assist-
ance in determining the supposed relationship of JZ. hectori to Berardius,
vou. X.—PaART Ix. No. 3.—August 1st, 1878. 3M
424 PROF. W. H. FLOWER ON THE GENUS MESOPLODON.
The principal dimensions of these crania are as follows :—
| l
M. australis, M. grayi, M. hectori,
| adult. young. | young.
| inches. | millim. inches. | millim. | inches. millim.
Extreme length of cranium ...........-.. 30:3 770 30°32 770} || 29:3 567
Length of rostrum (from the apex of the
| premaxilla to the middle of a line drawn | |
| between the anteorbital notches?) ...... 19:9 505 20:2 510 || 12:6 320
| From middle of hinder edge of palate (formed
| by the pterygoids) to apex of rostrum....| 23°8 605 24:5 | 622 | 17-4 442
| Greatest height of cranium from yertex to |
WeepLELy SOLUS eae eccrs cts causicte cases sxeilsvel ax 110 278 10-4 264 || 9:5 241
, Breadth of cranium across middle of superior
) . WAAC CMOS, 4 HobnG doa edo asoebe oT 11:2 283 10:5 268 9:2 235
, Breadth of cranium between zygomatic pro-
lMenxcessesolisquamosals etic: elke fees kc wale 7 298 111 282 || 10-2 259
| Breadth between anteorbital notches ...... TA 187 7:2 183 | is} 135
| Breadth of middle of rostrum ............ Tf 44 TL} 5} 40 15 37
| Breadth of occipital condyles ............ 3-9 99 37 | 95 3°6 92
| Premaxille, greatest width behind anterior |
MATOS vere ees eset esiare ee tae eve ee Rude eee Se 5-4 138 OU WSS Uae G 117
Preemaxille, least width opposite anterior Z
WP ATLATOS Gee u tM euenr git wan sstuckt duo sesaecavcey sche 4-4 111 40 | 103 4:2 107
| Premaxille, greatest width in front of an- |
| RASC TCRRESE Sp eet bo Boce OMe on on 6 ate 4-6 ulate Ara | 8 Lats! 4:5 114
| Width ofsanterior mares! 0... 150.6. 5. ' « 2:0 52 PPA — = G3} 2°0 51
| eensth\of tympanic bones... 2... k ee ig) 49 2:02 [ib 21 54
| Greatest breadth of tympanic bone ........ 13 32 14 | 385
| Mandible, length of ramus .............. | 26:0 660 19-1 484
| Mandible, length of symphysis............ | 84 214 5-9 150
| Mandible, greatest vertical height of ramus. . 4-1 105 33 83
The lower jaw appears to present no differentiating characters whereby the different
species of Mesoplodon can be distinguished, except as regards the implantation of the
teeth, which will be considered presently.
The thyro- and basihyals are present in the younger skeleton but not in the older
one. They are not united. In general form they resemble those of Berardius and the
ordinary Dolphins, presenting none of the peculiar enlargement and flattening cha-
racteristic of Physeter and Kogia and seen to a slight extent in Hyperoodon. The
stylo-hyals are preserved in both skeletons, and, allowing for difference of age, are
similar. They do not present the simple cylindrical characters of the same bones of
all the true Dolphins, and even of Physeter and Berardius, but are (especially those of
the older specimen) remarkably flattened and widened towards the lower end, and thus
approximate to the form which has hitherto been thought peculiar to Hyperoodon.
In M. australis these bones have a length of 5:6 = 142 millims., and a greatest
1 As the apex of the rostrum is stil] covered with the dried gum, this is not absolutely exact.
2 [have thus defined the commencement of the rostrum for the sake of uniformity with previous measure-
‘ments ; but, owing to the breadth of the maxilla in front of the anteorbital notches, the rostrum in Jesoplodon
appears to commence more than an inch in advance of the spot indicated.
PROF. W. H. FLOWER ON THE GENUS MESOPLODON. 425
breadth of 13 = 34 millims; in VW. grayi, a length of 5-3 = 135 millims. and a
breadth of 1-0 = 26 millims.
Dental characters.—All the Ziphioid Whales have previously been characterized as
having “ no functional teeth in the upper jaw. Teeth of mandible quite rudimentary
and concealed in the gum, with the exception of one or occasionally two pairs, which
may be largely developed, and project like tusks from the mouth, especially in the
male sex”?
A series of small rudimentary and concealed teeth has been shown to exist in the lower
jaw, behind the large functional teeth, in Mesoplodon bidens, by Gervais, in the upper
jaw of Ziphius cavirostris by Gervais, and in both the upper and lower jaw of a member
of the same genus, if not species, by Burmeister; and they may have been present, but
overlooked, in many other specimens, as such teeth, not being fixed in the bony
alveoli but only attached loosely in the gums, would invariably be lost by maceration.
The discovery by Dr. von Haast of the constant presence, in many individuals of dif-
ferent ages and sexes, of a row of small teeth of determinate number and definite
form, with their crowns projecting beyond the gum of the upper jaw, in a species of
Mesoplodon, is therefore of great interest. We have here the permanent retention
of a condition intermediate between that of the irregular, completely concealed, pro-
bably only temporary, and quite functionless teeth mentioned above and the normal
state of dentition of the true Dolphins; and it is especially interesting that this
should have been met with in a member of the genus otherwise least modified from the
Dolphins. I must, however, take leave to differ from Dr. Haast in thinking that it is
a difference of generic importance, especially as it is associated with no other structural
peculiarity, as far as is at present known’.
These teeth, from seventeen to nineteen in number on each side, are entirely uncon-
nected with the bone; and when the dense fibrous gum is removed there is no trace of a
socket or groove forthem. They have been preserved in the specimen under description,
the soft coverings of the rostrum having been allowed to dry upon it. They are nineteen
in number on each side, and occupy a space of four inches in the middle of the rostrum,
the anterior teeth being seven anda half inches from the apex. The first is very
minute ; but they gradually increase in size to about the sixth, then remain nearly equal,
but diminish again, though not to the same extent, in the posterior part of the series.
They are slender, conical, compressed from before backwards, and incurved at the apex,
with sharp-pointed enamelled crowns, resembling, in fact, those of the genus Delphinus
in miniature. ‘The apices of many are broken or blunted; whether from use or after
1 Trans. Zool. Soc. vol. viii. p. 204.
* Professor Geryais seems to be of the same opinion, as he says, “ Mais c’est particulicrement auprés du
Mesoplodon sowerbense qu’il doit prendre place, et dans la rangée de dents qu’il présente dans une partie de la
longueur de sa machoire supérieure, on ne trouyerait aucune bonne raison pour le séparer génériquement de ce
dernier, encore moins du Dolichodon” (Ostéographie des Cétacés, livr. xy. p. 518).
3M 2
426 PROF. W. F. FLOWER ON THE GENUS MESOPLODON.
death can scarcely be determined now; but Dr. v. Haast mentions that in his recent
specimens some of the teeth were “ evidently worn down from use.” The crowns of
the largest of these teeth are 4 millims. in length, projecting that distance above the
gum in its present dried state, though when recent they must have been covered to a
larger extent. Some teeth of an older individual (figured in P. Z. S. 1876, p. 10) are
somewhat larger. Their whole length, including crown and root, is 12 millims. or
half an inch, and their greatest thickness (in the part concealed by the gum) 23 millims.
There is no evidence of the presence of any such teeth in MW. australis or M. hectori;
and whether they were present or not in J/. haasti I am not able to say, as the bones
were cleaned before they came into my possession.
The genus Mesoplodon is characterized by possessing a single pair of compressed
mandibular teeth, which are usually situated at about one third of the distance from
the apex of the mandible to the condyles, or opposite the hinder edge of the symphysis.
This is, in fact, their situation in all known species, except J/. europeus and M. hectori;
and it is doubtful whether the first should be considered an exception ; for though the
teeth are situated further forward than in the others, relatively to the whole ramus of
the mandible, they have the same relation to the hinder border of the symphysis, which
appears (judging only by the published figure) to be shorter than in the other species.
The teeth of the different species appear to agree in their essential structure’, having
a small and pointed enamel-covered crown, composed of true dentine, which, instead of
surmounting a root of the ordinary character, is raised upon a solid mass of osteo-
dentine, the continuous growth of which greatly alters the form and general appear-
ance of the organ as age advances, as seen most strikingly in the case of MZ. layardi.
The changes that take place during growth in the mandibular teeth of M/. grayi are
well exemplified in the different specimens described by Dr. v. Haast. In the young
male skeleton the apex of the tooth projects but a quarter of an inch above the level
of the alveolar margin, and is quite unworn. During life it was probably altogether
concealed beneath the gum, though the animal had attained three fourths of its
full dimensions. ‘The apex is directed upwards and outwards. It hes in a special
expansion of the dental groove, which groove, it should be mentioned, continues to
exist, even in the aged animal, in a marked degree, along the whole upper border of
the dentary portion of the ramus. Although quite loose, it could not be removed to
examine its base until a portion of the bone had been cut away; yet in the old tooth,
which completely filled the socket, owing to the alteration of the form of the latter
this difficulty did not exist.
When removed the tooth has a triangular form; the apex, which is pointed and
somwhat everted, is covered with a thin layer of enamel, which extends to a greater
distance (7.e. 15 millims.) in front and behind than at the sides (7 millims.) The
1 See E. Ray Lankester, Trans. Microsc. Soc., new series, vol. xv. 1867, p. 55, and Trans. Zool, Soc. vol. viii.
p. 223.
PROF. W. H. FLOWER ON THE GENUS MESOPLODON. 427
height of the tooth at the middle is 1"°7, or 43 millims. The base is extended antero-
posteriorly to a length of 2!"1 or 53 millims., and has a rough, spongy, unfinished
and laminated edge, without any distinct pulp-cavity. The greatest lateral thickness
of the tooth is 6 millims. or 0!°25.
The old specimen (J. haasti of this communication) has a tooth very closely re-
sembling that of the adult JZ. grayi, judging of the latter from Dr. Haast’s description
and from photographs which he has sent me, and well illustrates the changes which have
taken place in passing to maturity. These are:—rounding of the apex, with almost
complete destruction of the enamel, except a small patch on the hinder part of the
tooth ; great wear of the whole anterior edge or shoulder of the tooth; and a great
increase of the supporting mass or bone, the growth of which is evidently completed,
as the lower edge is now firm and rounded, and has developed upon it a number of
short root-like processes. The height of the tooth is 5°3 = 84 millims., its greatest
antero-posterior breadth 3/"9=99 millims., its greatest lateral width 11 millims. or
nearly half an inch’.
This tooth is therefore of altogether a different form from that of the adult
M. bidens and europeus, in neither of which does the base enlarge in the antero-pos-
terior direction, but forms an oblong figure with parallel sides, the axis of which does
not coincide with the axis of the apex. Neither has it any resemblance to that of
M. layardi, the highly characteristic form of which has already been pointed out. In
MM. densirostris, though the exposed portion of the tooth is triangular and has its
apex placed nearly vertically, as in WZ. grayi and WM. haasti, the root when complete
assumes an oval form, contracting towards the base, and its vertical direction greatly
exceeds the antero-posterior. Most unfortunately, in the otherwise nearly perfect
adult skeleton of MW. australis, these most important and characteristic organs are
wanting. They were carried off, as Dr. Hector informs us, by some mischievous boys,
by whom the jaw-bone was broken into many pieces in extracting them; and thus no
satisfactory idea even of the form of the alveolus can be obtained. As, however, the
whole of the symphysis, with the anterior margin of the left alveolus, is preserved, it
can be demonstrated that the situation of the tooth in relation to the mandible was
quite the same as in WV. grayi and M. haasti, though (in accordance with the general
smaller size of the individual) the distance between the apex of the mandible and the
tooth is somewhat less than in the latter.
It is very probable that the fragment of a jaw of a very young Mesoplodon, found on
the beach at Kaikoura, and described and figured by Dr. Hector, may belong to this
1 Tt has long been suspected that the sexes of Mesoplodon differ greatly in the development of the teeth ;
but, owing to the want of materials, this has not been definitely proved. Of J. gray, however, Dr. Haast
informs us that in a full-grown female “‘ the mandibular tooth could scarcely be felt when passing the finger
over the gums; and its existence could scarcely have been ascertained in that way had I not known its
position” (P. Z. 8. 1876, p. 458).
428 PROF. W. H. FLOWER ON THE GENUS MESOPLODON.
species’. ‘The tooth is in a very early stage of development, being merely a hollow
cap of dentine, with a smooth and enamelled tip, and entirely embedded within the
alveolus. The formation of the base of osteo-dentine appears not to have com-
menced. It is compressed and triangular, smaller than the young specimen of MW. grayi
described above, and of different form, being narrower at the base, compared with the
height; but, as in that specimen, the apex was placed quite vertically. From so very
young a tooth it is not safe to draw conclusions as to the ultimate form the organ
would assume.
With regard to WV. hectori the same difficulty exists. The only specimen known is
so young as to have the teeth entirely embedded in the alveoli; but, as before men-
tioned, their situation at the anterior extremity of the mandible distinguishes this
from all other species. The apex, which is the anterior part of the tooth as it lies in
its socket, is exactly 1 inch from the anterior margin of the symphysis. The tooth is
very much compressed, being no where more than 5 millims. in breadth. ‘The base is
composed of two lamin, nearly in contact, with an open fissure between them. The
point is sharp, capped .with enamel to the extent of 8 millims. The height of the
tooth is 1-2 or 31 millims., the antero-posterior diameter of base 1-1 or 28 millims.
Vertebral Column.—the length of the vertebral column of the adult J/. australis, all
the bones being placed in apposition, is 150 inches. Allowing the same increase for
the intervertebral substance which was found in a common Porpoise, viz. as 115 is to
1002, we should obtain a length of 150 inches; the length of the skull (30 inches)
added to this gives a total length of 180 inches, or 15 feet, for the whole skeleton,
which corresponds very closely with Dr. Hector’s measurement of 15 feet 6 inches for
the animal when recent ®.
In the younger skeleton of the typical J/. grayi the vertebree measure 123 inches;
and by the same process the whole skeleton would be 14 feet 5 inches, rather more
than the length given by Dr. Haast, viz. 15 feet 8 inches; but of course the allowance
made above for the intervertebral substance is only approximative. Dr. Haast gives
the length of an adult female of the same species as 17 feet 6 inches?+.
The skeleton of J. australis has the following vertebral formula :—C. 7, D. 9, L. 11,
C. 20 = 47; that of IZ grayi has C. 7, D. 10, L. 11, C. 20 = 48. In both cases there
is probably one minute terminal vertebra wanting.
These numbers closely correspond to those of the few other complete skeletons of
Mesoplodon known, viz.:—that of IZ. bidens at Brussels, which has (according to Van
Beneden) C. 7, D.10, L. 10, C.19 = 46; that of the same species at Gothenburg,
which has C.7, D. 10, L. 9, C. 20 = 46 (Malm); and that of I. layardi, at the Canter-
1 Notice of a variation in the dentition of Mesoplodon hectori, Gray, Trans. N.Z. I. vol. vi. p. 86, pl. xv. a,
1874.
2 See Trans. Zool. Soc. vol. vi. p. 311. ’ Trans. New-Zeal. Inst. vol. vii. p. 262.
“ Proc. Zool. Soc. 1876, p. 457.
PROF. W. H. FLOWER ON THE GENUS MESOPLODON. 429
bury Museum, New Zealand, which has C. 7, D. 10, L. 10,C. 19 = 46 (Haast, P. Z. S.
1876, p. 481).
The cervical vertebre of the two skeletons are very much alike; indeed, considering
the difference of age and the great tendency to individual variation, it may be doubted
whether any differences that may justly be called specific can be detected between
them. In both the atlas and axis are completely united by the whole of the body and
arch, except for the spaces in the latter which constitute the foramina for the exit of
the second cervical nerves ; in both all the other vertebre are perfectly free’. This is
the minimum of vertebral union known in any Ziphioid, and is the same which occurs
in the specimens of J. bidens at Brussels and at Gothenburg. In I. layardi the
atlas and the second and third cervical vertebree are united into one large triangular
bone, the rest are all free®. In the skeleton of Berardius arnouai, in the Museum of
the College of Surgeons, the third is united by its body to the completely conjoined
first and second. In the fine skeleton of Ziphius cavirostris from Villa Franca, in the
Museum at Jena, the first, second, and third vertebrae are completely united by their
bodies, and the fourth is joined to them by the spine. In another specimen from
Corsica, described by Fischer, the six anterior vertebre are united. In the Gothenburg
Ziphius the first four are joined. In Hyperoodon the bodies of all seven are firmly
united together, and the spines of all except the seventh join to form a single elevated
conical mass.
The arch of the atlas bears in both specimens a complete foramen for the passage
of the cervical or suboccipital nerve, not a groove only, as in Berardius. In other
respects these bones bear a great resemblance to those of that genus, more than they
do to Ziphius and, @ fortiori, to Hyperoodon.
In WV. grayi the upper and lower transverse processes of the axis are united on the
left side, leaving only a small circular foramen between them at the base; but on the
right side in this specimen, and on both in JZ. australis, the processes are quite distinct,
and separated by a wide notch. In all the other vertebre, as far as the sixth, the
upper and lower processes are present and not united. They are developed in a nearly
corresponding degree in the two specimens, except that in the sixth vertebra the upper
process is very rudimentary in WV. australis, and the left lower process is developed to
nearly double the length of the right. ‘This is evidently an individual peculiarity.
These processes differ very considerably from the corresponding parts of Berardius,
as they are directed downwards, outwards, and in the main backwards, instead of for-
wards as in Berardius, in which it is the anterior portion of the process, and not the
principal or posterior part, which is chiefly developed *.
The arches and spines of the cervical vertebre are better developed generally than in
1 Dr. Haast tells me in a letter (Dec. 12, 1876) that in the skeleton of an adult female of JZ gray? the third
cervical vertebra is united to those before it.
2 Haast, P. Z.8. 1876, p. 482. * See Trans. Zool. Soc. vol. viil. p. 225.
430 PROF. W. H. FLOWER ON THE GENUS MESOPLODON.
Berardius. They are all completely united above, except the fifth of the young J. gray.
In the sixth, in both specimens, the spine is so well developed as to equal in height the
spinal canal over which it is placed. In the seventh, in MW. australis, it is considerably
higher than in J. grayi, and also much stouter at the base. It is a question, however,
how much the difference between the two may be due to age. ‘The seventh in both
has a large articular surface on the posterior edge of the side of the body for arti-
culation with the head of the first rib. At the lower end of this surface isa small
tubercle representing the inferior transverse process of the preceding vertebra.
Dimensions of the Cervical Vertebre.
M.australis, || MM. grayi. |
|
|
|
inches. | millim. | inches, | millim.
Antero-posterior length of the bodies of the seven vertebre in con-
acti G@nteriomsurtace)| wy. sete teeter ol ein che ret ears 51 130 || 49 | 124
Length of body of united first and second .................... 1:9 48 || 1:7 | 43)
LGU Or LOE POLG CMI Saacpenvadddaueascooubec saeduLded 3) 23 6 15 |
| Height from top of spine to lower edge of body of atlas .......... a1 WSO F456 | eblig
| rf 55 es third vertebra ..| 4:1 104 || 3°5 89 |
- a sixth vertebra . 5 55 | 140 BHO |) UAC |
| 3 5 3 seventh vertebra.| 7-1 | 180 || - 5-6 142 |
| Breadth between outer borders of articular surfaces of atlas ...... | 37 | 94 | 39 99
Breadth between ends of transverse processes of atlas............ | 56 | 142 | 55 | 140 |
}
It will be seen from the above that the only measurement in which /. grayi exceeds
M. australis is the width of the surfaces for articulation with the occipital condyles.
This, taken in conjunction with the immaturity of the specimen, is a tolerably sure
indication that it would, if adult, attain a larger size than the latter.
The thoracic vertebra of both skeletons present all the characteristic features of the
group, the most remarkable of which is the sudden transition of the articulation of the
ribs from the upper to the lower processes, by which the Physeteride are distinguished
from the Delphinide. The principal difference between the two is that in WZ. australis
there are but nine with nine pairs of ribs (as in Hyperoodon), whereas in MW. grayi there
are ten, as in all the other recorded examples of Mesoplodon'.
In the erect high spines of the thoracic vertebrae Mesoplodon resembles both Ziphius
and Hyperoodon, and differs in a marked manner from Berardius’.
1 It should be mentioned that, according to Hector’s original description (Trans. N.Z. Inst. vol. vii. p. 262),
the skeleton of WZ. australis had ten pairs of ribs; but the number now present with it is that which I have
stated above, and the vertebra which I have reckoned as the first lumbar bears no trace of an articulation for
arib at the end of the transverse processes. Moreover a close comparison shows that the rib which is wanting
would not correspond to the last of 1. grayi or the other ten-ribbed species, but rather to one from the middle
of the series; and yet there is no indication of the accidental loss of both vertebra and pair of ribs, which must
have been the ease if Dr. Hector’s enumeration was correct,
* See Trans. Zool. Soc. vol. viii. p. 226.
PROF. W. H. FLOWER ON THE GENUS MESOPLODON. 431
Articular surfaces of the arch, or zygapophyses, are developed in all the anterior
vertebra as far as to between the sixth and seventh in WV. australis, and as far as the
tenth (though in a very rudimentary condition) in UM. grayi. Metapophyses first
appear as distinct tubercles in both on the anterior edge of the transverse process of the
third vertebra, and gradually increase in size in passing backwards. Articular sur-
faces for the heads of the ribs on the hinder edge of the body occur from the first to
the seventh inclusive in both, for the tubercles of the ribs on the transverse processes
from the first to the seventh in J/. grayi, and to the seventh on the left side only in
M. australis. The last two (or the ninth and tenth in MW. grayi, and the eighth and
ninth in J. australis) have the rib attached to the end of the transverse process
springing from the side of the body of the vertebra. The eighth rib in WZ. grayi is
attached to the articular surfaces on the contiguous sides of the body of the seventh
and eighth vertebre, on a line with the last, and not to any transverse process; so in
this specimen seven ribs are attached in the ordinary way by upper and lower attach-
ments to body and transverse process, and three to the lower attachment only. In
M. australis six were attached by the former method, two by the latter, and the inter-
mediate one by the former method on the left, and the latter on the right side ; so that
the seventh vertebra and pair of ribs appear clearly to represent both seventh and
eighth of the other specimen combined in one, and so to account for the smaller
number altogether.
The lumbar vertebre are all very much alike in both skeletons. ‘Their bodies, as in
all the Ziphioids, are long compared with those of the true Dolphins, their length con-
siderably exceeding their transverse diameter. They are strongly carinate below. The
transverse processes are short, their length not exceeding the width of the body; their
bases arise from rather more than the anterior half of the sides of the body; they are
all directed forwards, and rounded at the free extremity. The spines are very long,
and nearly vertical, much compressed laterally, but broad from before backwards (ex-
ceeding half the length of the body), expanding and truncated at their upper extremity.
They all have broad, flattened, lamelliform metapophyses, projecting forwards from the
anterior edge of the lamina, at nearly the same level throughout the series.
The caudal vertebre in the two skeletons bear the closest resemblance in all essen-
tial features. ‘The one which I have reckoned as the first of this series differs from the
lumbar vertebre in having the keel rudimentary and divided posteriorly, where it
presents slight indications of the presence of chevron bones, probably in a rudimentary
condition, as they were not preserved in either skeleton. In the second and succeeding
vertebrae the median line of the inferior surface is channelled, and has a pair of lateral
longitudinal ridges terminating in front and behind in the articular surfaces for the
chevron bones. In both skeletons there are eleven of the series of compressed or
chevron-bone-bearing caudal vertebre; the twelfth is the transitional vertebra; and
there are thirteen of the depressed form, which lie within the laterally expanded portion
VOL. X.—Part 1x. No. 4.—August 1st, 1878. 3N
432 PROF. W. H. FLOWER ON THE GENUS MESOPLODON.
of the tail. As before mentioned, the appearance and size of the last vertebra present
leads to the belief that in both cases an additional minute terminal vertebra has
been lost.
The arches and spines gradually diminish from the first, the last being on the eleventh
in both skeletons; and in both the last trace of a transverse process is seen on the
ninth. As in Berardius, the transverse processes are not perforated, but grooved pos-
teriorly. In the eighth of JZ. grayi a vertical arterial canal passes through the base of
the rudimentary transverse process, which in the corresponding vertebra of J. australis
is grooved only; in the ninth and all succeeding vertebre of both skeletons the canals
pierce the sides of the bodies.
Of chevron bones there are nine present in the skeleton of J/. australis, and eight in
that of WZ. grayi. These evidently correspond, the ninth being absent in the latter.
The first (probably in reality the second) is small, and with the apex directed forwards.
It consists of two lamine, united in the middle line below, but not developing a keel. The
next has a considerable keel. The third is the largest, and in the adult skeleton has a
flattened rough surface at the apex. The fourth has a shorter but broader keel; and
from this they gradually diminish to the end. The hemal spines are longer proportion-
ally than in Berardius, corresponding in this respect with the neural spines.
The Sternum of the adult MW. australis (Plate LX-XIII. fig. 5) consists of four distinct
segments, connected only by cartilage, and, as is usual in the Ziphioids, not completely
ossified in the middle line, so that a notch is left at the anterior and posterior end of
each, forming a series of median foramina in the complete sternum. ‘That between the
third and fourth, however, is very small; that between the second and third is largest.
At the anterior extremity two processes, projecting forwards, curve inwards, and nearly
enclose another fenestra. The articular surfaces for the costal cartilages are well
marked. ‘The first is on the side of the first segment; the second between the first
and second segment; the third between the second and third; the fourth between
the third and fourth; the fifth on the side of the fourth segment. The terminal
piece is unsymmetrical, being longer on the right than on the left side. The extreme
length of the ossified sternum is 20 inches (508 millims.); the greatest breadth across
the first segment is 5°8 inches (148 millims.).
In WV. grayi the sternum (Plate LXXIII. fig. 2) presents all the characters of imma-
turity, the anterior and other processes being completely undeveloped. Otherwise it is
of much the same form as the former; but the part corresponding to the fourth
segment of that one is larger, and separated into two by a transverse suture, of which
but slight indications are to be seen at the same place in the sternum of VW. australis.
Its length is 17 inches =452 millims., its greatest breadth 5’-7=145 millims.
The Ribs.—In the skeleton of MW. australis there are nine, in that of IZ. grayi ten
PROF. W. H. FLOWER ON THE GENUS MESOPLODON. 433
pairs of ribs!. In the former the first to the seventh inclusive have distinct articular
surfaces on the head and the tubercle; the seventh on the left side is transitional ; the
remainder want the neck and head, articulating only by the tubercle to the extremity
of the transverse process of the vertebra.
In UW. grayi the first to seventh on both sides have distinct necks, and capitular and
tubercular articular surfaces; the three others, wanting the head and neck, articulate
only by the tubercle to the extremity of the transverse process. The general characters
of the ribs are, of course, more marked in the older specimen ; but otherwise they agree.
The first is very broad and flat; and in all the angle and a space beyond presents a
more or less strongly pronounced flattened surface on the outer side, with prominent
posterior edge. This character, scarcely seen in the first, gradually becomes more
marked to the last. None of the ribs materially increase in breadth at the lower
extremity.
The sternal ribs in both skeletons, as in all Physeteride, are not ossified.
Pectoral Limb.—Yhe scapula is perfectly delphinoid in form, more even than in
Berardius, and therefore further removed from Physeter than any other Ziphioid,
Hyperoodon being the nearest. The only differences between the two specimens are
such as might arise from want of ossification of the suprascapular cartilage, and of the
end of the coracoid process in the younger specimen. The humerus, and especially the
ulna and radius, present the simple elongated narrow form characteristic of the
Ziphioids, and by which they are easily distinguished from the Delphinide. Allowing
for difference of age, there is nothing by which the two individuals can be specifically
distinguished. The same can be said of the bones of the manus, as far as the available
materials serve for comparison. In JZ. grayi both are quite complete; and in MW. aus-
tralis one hand is nearly so.
This segment of the limb closely resembles that of Berardius?, except that the
singular union of scaphoid and lunar, and of cuneiform and unciform bones, noticed
in that animal, does not occur. The bones of the first row, the scaphoid, lunar,
cuneiform, and pisiform, are all distinct. In the second row the trapezoid and
magnum are united into a single bone, as in the skeleton of MZ. bidens in the Brussels
Museum®. The first digit has a single slender phalanx; the second has six, the third
the same number, the last being a minute nodule, scarcely larger than a pin’s head;
the fourth has three, and the second two phalanges. These are the complete number
of the adult skeleton.
* Dr. Haast mentions in a letter (dated March 8th, 1876) that one specimen of MW. grayi has ten, another
eleven pairs of ribs.
* Trans. Zool. Soe. vol. viii. pl. 28. fig. 10.
* In the specimen at Gothenburg, Malm figures these bones as distinct, loc. cit. pl. v. fig. 52.
3Nn 2
434 PROF. W. H. FLOWER ON THE GENUS MESOPLODON.
Dimensions of the Bones of the Pectoral Limb.
M, australis, M. gray?.
Scapula: height from middle of external margin of glenoid fossa in. millim. in. millim
Loumiddlevofysuperior border’ aoe eee eee cee ee 8:2=208 7-3=185
Length from anterior to posterior superior angle ............ 13°8=325 ks = 287
Length of pectoral limb from head of humerus to end of second digit 19:0=483 17:0=432
Humerus gl ons thw mea. eee Tees abe ee ete Els 5:3=1385 5:2=132
Bread thgatploweriend says tA gongs, heres eek Riese es 2-5= 64
Radius: mlene themem re tre tocra tik crore’. scat otra evon ae tte ee nseeete 6-°0=153 5:6=143
Ulnaj:lenothtancludingyolecranon 25: .0...222. ee.) c sande ones 6°5=166 5°9=150
Breadth of radius and ulna at lower end ...............+---+ 31= 78 | 3°0= 75
IWlsenothyoremanust semen: oe eee re eee ree ere 9:0=229 7-0=178!
Summary and Conclusion.
The evidence which has accumulated up to the present time is sufficient to
show that there are at present existing in many parts of the ocean several distinct,
though closely allied, species of Cetaceans, united by common characters, to which the
generic name Mesoplodon may be applied. The reasons for the choice of this name
in preference to many others which have been applied to members of the group have
been fully stated in a previous communication ?.
The nearest known ally to these animals is unquestionably Berardius, although
in some characters, especially the form of the spinous processes of the vertebra, they
rather resemble Ziphius and Hyperoodon.
With the three genera just named they form a natural group, allied on the one
hand to Physeter, and on the other to the true Dolphins; but their affinities are,
on the whole, nearest the former, which, indeed, may be regarded as a highly special-
ized form of the same type, Hyperoodon coming next to it, and Mesoplodon being the
most generalized.
Gervais, who founded the name Mesoplodon, making M. bidens or sowerbiensis the
type, did not include two species assigned to it above, viz. IW. densirostris and
M. ewropeus, but put them in another genus, Dioplodon, an arrangement to which he
adheres in his recently published ‘ Ostéographie des Cétacés.’ He has, however, never
clearly defined the distinction between the two groups; and it is exceedingly difficult
to discover any characters common to the two last-named species which is not also pos-
sessed by the former. This has been the opinion of most authors who have investigated
the subject subsequently, as Duvernoy, Fischer, E. Deslongchamps (who, however,
uses the name Dioplodon for the whole genus), and Van Beneden. A _ close
examination of the cranial characters of the known species does, however, reveal
* As these measurements were taken after drying, the disproportionate shortness of the manus of the second
specimen may in a great measure be accounted for by the shrinking of the cartilages, which occupy in it a far
more considerable proportion than in the older animal.
* Trans. Zool. Soc. vol. viii. p. 208.
PROF. W. H. FLOWER ON THE GENUS MESOPLODON. 435
Q
differences by which the group may be divided into two sections, not equivalent to
those of Gervais, but to which, if thought advisable, his names might be applied, as
his JMJesoplodon sowerbiensis may be taken as the type of one, and his Dioplodon
densirostris as the type of the other. To the first (JZesoplodon proper) belong MW. bidens,
M. europeus (if distinct), MW. layardi, and M. hectori; to the second (to which the
name Dioplodon may be given if it should appear that a generic term is necessary
to distinguish it), IW. densirostris, M. australis, M. grayt, and M. haasti. Certain cranial
characters, and the position of the large mandibular tooth, distinguish JZ. hectori, and
may possibly afford grounds for placing it in another section intermediate between
Mesoplodon and Berardius; but with the present limited knowledge of its structure,
derived only from the cranium of a very young individual, I think it preferable to
retain it provisionally in the position to which it is assigned above. The structure
of the vertebral column, when known, will throw light upon its relationship to
Berardius. The objection that the name Mesoplodon is not applicable to this species
is scarcely worth considering, as it is a case in which “the sooner a term becomes an
arbitrary sign the better” !.
The further subdivisions which have been propesed, as Dolichodon and Callidon, of
Gray, and Oulodon of Haast, appear to me unnecessary, though, with regard to the
latter, this may be a matter of opinion.
Tt has been assumed above that the different individuals assigned by various authors
to the IZ. bidens or sowerbiensis are really one species; for I have not had an opportunity
of making a critical comparison of them, though it would be desirable to do so as soon
as a sufficient number of specimens are collected in our Museums to afford materials for
thoroughly estimating all the variations to which they may be subject according to sex
and age. Perhaps more light may then be thrown upon the relation to that species of
the solitary example known of I. ewropeus, which is at present not a very satisfactorily
established species.
Of the southern species, MW. layardi, M. densirostris, and M. grayi are thoroughly
well established; complete skeletons of each are in existence, and have been more or
less perfectly described, though, unfortunately, no skeletons of either of the two former
have yet reached Europe. MV. australis is known by the skeleton, complete but for the
important exception of the mandibular teeth; further evidence of its distinctness from
M. grayi may perhaps be considered desirable. J/. haasti-is only known by a rostrum
and mandible. J/. hectori must be a totally distinct form, though at present very
imperfectly known.
The geographical distribution of the group has a very great interest in relation
to that of many other Australian groups, both of vertebrates and invertebrates.
Among the earliest known remains of Cetacea, in the Belgian and Suffolk Crags, Mesop-
lodon and closely allied forms are most abundant. Up to little more than ten years
* Owen, Trans. Geol. Soc, 1857, p. 55.
436 PROF. W. H. FLOWER ON THE GENUS MESOPLODON.
ago the few stray individuals of MW. didens occasionally stranded on the shores of North
Europe were supposed to be their sole survivors. Since that time it has been proved
that they are still numerous in species, and even in individuals (as many as twenty-five
of M. grayi having been stranded on one occasion on the Chatham Islands, and four
at another time on the New-Zealand coast, where it is sufficiently abundant and well
known to have obtained the local name of Cowfish!) in the seas which surround the
Australian continent, extending from the Cape of Good Hope on the one side to New
Zealand on the other, though beyond these limits no specimens have yet been met
with. It is the history of the Marsupial Mammals, of Ceratodus, of Terebratula, and
of numerous other forms.
As the food of these Cetaceans consists, as far as is known, mainly of Loligo and
other free-swimming Cephalopods, which frequent the open ocean, we need not be
surprised if it should be found that their range in space is tolerably wide, although,
with our present imperfect information, many of the species appear to be local.
The great advance that has been made in our knowledge of this interesting group
in so short a period should encourage those to whom it is due not to relax their efforts
to secure every specimen that may be brought within their reach.
Postscript.—Since this memoir was communicated to the Society I have received (Dec.
4th, 1877) the X Vth part of the great ‘ Ostéographie des Cétacés, Vivants et Fossiles,’
by Van Beneden and Gervais, in which the latter author describes and figures a com-
plete skeleton of Mesoplodon grayi, also obtained by Dr. v. Haast, of the same age and
sex as the specimen described above, and from the same locality. It appears to cor-
respond in every detail with that described in the present memoir.
DESCRIPTION OF THE PLATES.
PLATE LXXI.
Fig. 1. Upper surface of cranium of adult Mesoplodon australis, from the specimen
in British Museum.
Fig. 2. Upper surface of cranium of young Mesoplodon grayi, from the specimen in
the Museum of the Royal College of Surgeons.
Fig. 8. Upper surface of rostrum of adult Iesoplodon haasti, from the specimen in the
Museum of the Royal College of Surgeons.
* Haast, P. Z. 8. 1876, pp. 7 and 457.
PROF. W. H. FLOWER ON THE GENUS MESOPLODON. 437
Fig. 4. Upper surface of cranium of young Wesoplodon hectori, from the specimen in
the British Museum.
m.f, maxillary, p.f, premaxillary division of the foramen for the exit of the infra-
orbital branch of the second division of the fifth nerve.
All one fourth of the natural size.
PLATE LXXII.
Fig. 1. Side view of cranium of adult Mesoplodon australis, from the specimen in the
British Museum.
Fig. 2. Side view of cranium and mandible of young Mesoplodon grayi, from the speci-
men in the Museum of the Royal College of Surgeons.
Fig. 3. Side view of the rostrum and mandible of adult Mesoplodon haasti, from speci-
men in the Museum of the Royal College of Surgeons.
Fig. 4. Side view of the cranium and mandible of young Mesoplodon hectori, from the
specimen in the British Museum.
All one fourth of the natural size.
PLATE LXXIII.
Fig. 1. Side view of the articulated skeleton of the young Mesoplodon grayi, in the
Museum of the Royal College of Surgeons.
Fig. 2. Sternum of the same.
Fig. 3. Upper surface of the vertebral column of adult Mesoplodon australis, from the
specimen in the British Museum.
Fig. 4. Side view of the same.
Fig. 5. Sternum of the same.
[It was intended that these figures should all be reduced to the same scale, viz. 75;
but by some error figs. 1 and 2 are reduced to about 74, and figs. 3, 4, and 5 to between
g and 39, giving the appearance of greater relative size to the latter skeleton than
it really possesses. |
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