A STUDY OF THE GENUS BERARD/US DUVERNOY*CHARLES McCANN
8 Kiwi Street, Heretaunga, Upper Hutt, New Zealand

ABSTRACTThe taxonomy and distribution of the two known species of Berardius,
B. arnouxi Duvernoy and B. bairdi Stejneger are reviewed, and some anatomicaland biological aspects of the two species are considered.

INTRODUCTION

The members of the genus Berardius or Porpoise Whales, in spite of their large size,
are little known. Although one of its two closely allied species, B. bairdi Stejneger,
was known to and fished for by Japanese whalers very early in historic times, it was
not scientifically recognised until 1883.
The genus Berardius was founded by Duvernoy in 1851 on a skull from Akaroa,
Banks Peninsula, New Zealand. The specimen was taken to France by M. Arnoux,
who was at the time surgeon to the French corvette Rhin commanded by Captain
Berard. These two gentlemen are commemorated in the generic and specificnames of the type species, Berardius arnouxi Duvernoy.
Although more than a hundred years have elapsed since the original description of B. arnouxi, barely forty specimens have been reported in literature. Bysome mischance, the ' o ' was omitted from Arnoux in the original description.
Since then, some cetologists have retained the original spelling, arnuxi, while others
have amended it to arnouxi. As the original spelling was almost certainly a 'lapsus
calami ', the amended spelling should be adopted.
Flower (1872) gave the first detailed description of the general anatomy ofBerardius from a specimen then in the Museum of the Royal College of Surgeons.
This specimen which had stranded at New Brighton, Canterbury, New Zealand,on 16th December, 1868 had been sent to London by Sir Julius von Haast. Little
has since been added.
The two known species are geographically isolated: B. arnouxi Duvernoy occurs only in the Southern Hemisphere, in the Southern Pacific and Southern
Atlantic Oceans; whilest, B. bairdi Stjneger is confined to the Northern Pacific
Ocean, concentrating seasonally near Japan while strays are occasionally found* This paper was compiled some years ago while the author was still employed as the Vertebrate Zoologist
at the DOMINION MUSEUM, Wellington (now the NATIONAL MUSEUM) under the Directorship of Dr. R. A. Falla (now Sir Robert A. Falla), since retired. Dr. R. K. Dell, is the present Directorof the National Museum.
The specimen registration numbers quoted in the paper are referred to by the originals with theprefix D. M. or Dom. Mus.
Sci. Rep. Whales Res. Inst.,

No. 27, 1975, 111-137.

C. McCANN

112

along the western shores of North America.
Recent papers by Omura et al. (1955) and others have discussed the biology
and whaling of B. bairdi, but no such information is available for B. arnouxi. McCann (1962), in a revision of the genus Mesoplodon discovered that the calves of B.
arnouxi had until then been identified as Mesoplodon hectori (Gray). Hector (1870)
had originally identified the type of M. hectori (Gray) correctly as the young ofBerardius arnouxi Duvernoy, but Gray (1871) declared it a new species of Berardius.
Turner (1872) cautiously transferred B. hectori Gray to Mesoplodon where it hadremained ever since. McCann's discovery not only places the genus Mesoplodon ina more satisfactory systematic position, but indicates that the breeding grounds ofBerardius arnouxi are in the vicinity of the New Zealand Archipelago.
GENUS BERARDIUS DUVERNOY (PLATES I-II)
1851. Berardius

1872.

Mesoplodon

1922.

PaikeaDuvernoy, Ann. Sci. Nat. (3) 15: 52. Type Berardius
arnouxi Duvernoy. Type locality: Akaroa, New Zealand.
Type in Paris Museum.
Turner, Trans. Roy. Soc. Edin. 26: 778. (Berardius
hectori (Gray))
Oliver, Proc. Zool. Soc. Lond. p. 575. Type Mesoplodon
hectori (Gray), Type Brit. Mus. 1677b. 76. 2. 16. 3.
DESCRIPTION

Large whales attaining 7.8 to 12.7 metres (26 to 42 ft) in length. Head large, buffer-like, without nuchal depression but a pronounced rostrum or beak; upper jaw
subacute anteriorly; lower jaw longer than the upper, extremity rounded; twopairs of strongly compressed, functional teeth near the extremity, the anterior larger
than the posterior; teeth erupted in both sexes on reaching maturity; occasionally
only a single (anterior) tooth present in each ramus; normally no teeth in maxillae.
Eye of moderate size. Blowhole median, crescentic. A pair of diverging gulargrooves forming an inverted V-shape, the anterior arms not in contact; occasionallyan additional pair of shorter grooves may appear; no thoracic nor abdominal
pleats. Body fusiform, laterally compressed posteriorly only. Flippers moderately
broad and rounded distally; flukes large, shallowly falcate, without median notch
although occasionally a shallow indentation may appear mesially. Dorsal fin nearer
the caudal, triangular, slightly falcate or represented by a short obtuse elevation.*
SIZE

B. bairdi is larger than B. arnouxi; the former reaches 12.6 metres in length (42 ft),
whereas the latter has not been known to exceed 9.9 metres (33 ft).

Curiously,* Perhaps the result of injury or attack by shark.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

STUDY OF BERARD/US DUVERNOY

113

although B. bairdi is the larger and bulkier of the two species, it has a relatively
smaller head than B. arnouxi (True, 1910: 67).
SEXUAL DIFFERENCES

Females of B. bairdi are larger than males, but in B. arnouxi both sexes appear to beof very similar size. As the mandibular teeth erupt in both sexes at maturity, theexternal sexual differences are not as well marked as other genera of the family inwhich the teeth erupt in the male alone.
GESTATION

The few breeding records available, suggest that the gestation period is approximately ten months and calving takes place during the late spring or early summer.
ANATOMICAL OBSERVATIONSThe skull (Pls. III-VI)
The skull of Berardius differs from those of other ziphioid genera in its greater
symmetry and simplicity. Its elements are less firmly interdigitated than in othergenera. The mesorostral groove remains unossified throughout life, i.e. the mesethmoid
cartilage does not ossify partially nor wholly as in ,(iphius and Mesoplodon. The
cranium is relatively more depressed in adults than in the rest of the family, being
broader than deep (Pl. VI).
Flower (1872) drew attention to the imperfect closure of the suture between
the basisphenoid and presphenoid bones; the suture being, at times, open 12.5 mm(1 in) or more. This is quite normal, not only in Berardius but also in other members of the Ziphiidae. In Berardius, the suture between the supraoccipital and thefrontal bones is also imperfectly closed, but in the rest of the family there is a more
complete fusion of these elements. The basioccipital and the basisphenoid elements
fuse completely, so that, in all the genera, the line of fusion is not clearly discernible.
In dorsal view (Pl. III, fig. 2), the posterior cranial region is semilunate, theantorbital tubercles forming the ' horns ' of the moon. The rostrum is somewhat
lanceolate, arising from within the antorbital notches and is considerably shorter
than the mandibles. In side view (Pl. III, fig. 3 ), the dorsal profile is almost straight;
the lower profile rising gently from the pterygoids towards the end of the rostrum.
The maxillae are much shorter than the premaxillae, the latter alone forming theextremity. In ventral view (Pl. III, fig. 1) a small portion of the vomer is visible
between the pterygoids and palatines. For a short distance anteriorly, the approximation of the margins of the maxillae obscures the vomer, which reappears once
more as a linear-lanceolate bony wedge between the maxillae and premaxillae(Pl. III, Vo), but ends considerably short of the rostrum extremity. The premaxillae approximate (but do not fuse) in front of the vomer. An open canalSci. Rep. Whales Res. Inst.,
No. 27, 1975.

114

C. McCANN

forms a ridge on the ventral aspect of each premaxilla. From the anterior 'segment ' of the vomer, there is a marked difference in the transverse level between the
maxillae and premaxillae, the latter being nearer the midline, resulting in a certain
amount of flatness in this region.
No basirostral grooves are present, but there is a blind ' pit ' beneath the maxillae just in front of the malar bones (Pl. VII, MP). The position of this pit corresponds approximately to the commencement of the basirostral groove in Mesoplodon
grayi Haast, the species in which the basirostral grooves are best developed in thefamily. There is no maxillary notch or tubercle as in Mesoplodon, but a maxillarycrest is present (PL. 7, MXC).
In Berardous the premaxillae are subequal posteriorly, the right being only
very slightly wider than the left in the narial region. Their extremities are nodular
and are not turned outward as in Mesoplodon nor do they overhang the narial opening in the adult. The large bulbous nasal bones overtop the ends of the premaxillae slightly and over-hang the narial opening, supported below by a wide
'bracket' formed by the posterior wall of the vomer, (Pl. V, fig. 3, Vo). In the
newly-born animal the nasals do not attain the full size and are consequently below
the level of the extremities of the premaxillae. Two large ' blind pits ' (BP) on thewall of the vomer, just below the nasals (one on either side of the septum) arecharacteristic of Berardius and readily distinguish it from any other genus of Ziphiidae. These pits are clearly visible in both old and young (Pl. V, fig. 3, BP).
The mesethmoid is completely fused with the presphenoid posteriorly. Within the narial cavity the mesethmoid is sandwiched and fused between the recurved
walls of the vomer; together they form the narial septum. Anteriorly the mesethmoid emerges from between the vomerine walls and enters the mesorostral groove
as a free projection for a short distance, becoming thicker and roughened along its
dorsal surface. It is free from the vomer.
The walls of the narial cavity are composed posteriorly and mesially (septum)
largely of the vomer; the lateral faces largely of the pterygoid, the premaxillae andsmall wedges of the maxillae and palatines. A large foranien, situated on the internal lateral face of the nares leads by a canal, laterally, to a large foramen in theorbital region.

Foramina (Pl. III): In dorsal view, the skull shows several large formina, some ofwhich are useful accessory diagnostic features. Anteriorly, there are two premaxillary foramina, one in each premaxilla (PMXF); two large maxillary foramina, one (sometimes dual or multiple) in each maxilla (MXF). Both pairs offoramina are in front of the narial cavity; in Berardius they are approximately on
the same transverse level although slight variations in level may occur. At theposterior edge of the maxillary crest or occasionally slightly further back, there isa large foramen connected by a canal with a foramen in the orbital region. This
post-crest for amen (PCF) (as I propose to call it) is peculiar to Berardius and absent inother ziphioids, even in Hyperoodon ampullatus which has a greatly exaggerated maxillary crest. Still further back, on the maxillary plate, there is another large foramen
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

STUDY OF BERARD/US DUVERNOY

115(PMF) (and occasionally an additional one or two); a canal connects this foramen
to another in the orbital region.

Antorbitalforamina (Pls. VI, VIII): In ventro-lateral view, the skull of Berardius has
three large foramina in front of the large infundibulliform optic foramen (OPF).
These three large foramina are not readily seen in any of the other ziphioid genera
as they are concealed within the apparently single 'antorbital foramen' behinda ridge formed by the palatine bone. The anteriormost foramen (Pl. VII, fig. I)

Fig. 1. Arrangement of antorbital foramina (diagramatic).
figures. del. C. McCann.

Lettering as in otherleads by a wide canal to the maxillary foramen (MXF) on the dorsal surface. Occasionally, immediately above this foramen (1), a small foramen perforates thelachrymal bone (L) and passes under it. A short distance behind is another large
foramen (Pl. VII, fig. 2) which perforates the lachrymal bone posteriorly; a large
vessel leads from this foramen to the dorsal surface of the skull where it appears as
the post-maxillary foramen (PMF) (or foramina). Lastly, somewhat below and alittle behind the last mentioned foramen and partially below the optic foramen(OPF) is another large foramen (Pl. VII, fig. 3). From this foramen, a large canalpasses to a large foramen on the lateral wall of the narial cavity (NF) and anotherbackwards through the foramen lacerum (Pl. X, fig. 1) into the brain cavity. Thisis the passage of the 5th cranial nerve.
From within the maxillary foramen (MXF) several secondary canals diverge(text-fig. I): one large division from the maxillary contributes to the formation of
the premaxillary foramen (PMXF); anteriorly a canal passes through the matrixof the maxilla towards the extremity of the rostrum (I term this the median rostral
canal (MRC)); posteriorly, three similar canals pass backwards under the premaxilla towards the vertex, emerging laterally under the posterior extremity of thepremaxilla. Other smaller canals from within the maxillary foramen arise to appear dorsally on or near the base of the maxillary crest. Finally, a large canalSci. Rep. Whales Res. Inst.,
No. 27, 1975.

116

C. McCANN

from within the maxillary foramen emerges as the maxillary crest foramen (MCF)
towards the rear of the crest itself.
From within the premaxillary foramen, a canal is given off anteriorly leading
to the extremity of the rostrum. This canal, which I propose to call the upper rostral
canal (URC), is sandwiched between the premaxilla and the maxilla, particularly
proximally.
Attention has already been drawn to the canals leading from within the large
posterior foramen to the narial cavity and the brain case. Anteriorly, a small canalpasses through the maxtrix of the maxill towards the end of the rostrum; I propose
to call this the lower rostral canal (LRC). These three rostral canals are readily seen
in a cross section through the rostrum.
(In .?,iphius, the lower rostral canal, instead of passing through the matrix ofthe maxilla, passes to the exterior at the anterior edge of the palatine bone andcontinues in a shallow groove to the end of the rostrum underside).
The optic foramen is large and canals lead from it into the cranial cavity, tothe internal optic foramen, and through the foramen lacerum.
Behind the optic foramen (OPF), the external foramen ovale (FO) is situatedbeneath the falciform process (FP) of the squamosal surrounding the tympanic
bulla. A little further to the rear, and at a slightly lower level, is the external auditory foramen. The exterior exit of the carotid artery is somewhat irregular inposition and size; it is situated at the suture between the pterygoid and the alisphenoid.

Brain case (Pl. X, fig. 1): The brain cavity is amply described by Flower (1872).
Relatively, it appears to be larger than in other members of the family. A
well-marked ridge, formed by the orbito-sphenoid and continued upwards by thefrontal, is present in Berardius. This ridge is developed to a lesser degree in Hyperoodon, but absent in both ,Ziphius and Mesoplodon.
The base of the brain case appears to be flatter in Berardius than in .<:,iphius.
The sella tursica and tuberculum sella are both better developed in ,Ziphius (female)than in Berardius, and the walls of the cerebellum fossa appear to be more rugose.
In sagittal section the internal cranial foramina are readily visible. Fromanterior to posterior, they are:
1) A small foramen high up on the wall of the presphenoid close to the midline, entering the nares by a canal. Flower (1872) suggests this may be the rudiments of the olfactory foramen.
2) The large optic foramen perforating the presphenoid.
3) The large sphenoidal fissure (Pl. VII, fig. 1) and the foramen rotundum
slightly behind the optic foramen (Pl. VI, OPF), but obscured by the orbito-sphenoidal ridge. These transport the nerves to the orbit and the middle division of
the 5th cranial nerve.
4) A small foramen for the carotid artery (Pl. IX, ea). On the floor of thebasisphenoid, on a transverse level with the foramen ovale. The canal to the exterior appears as a foramen near the posterior border of the pterygoid.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

STUDY OF BERARDIUS DUVERNOY

1175) The large foramen ovale further to the rear of the sphenoidal fissure andpartially obscured by a ridge of bone. In Berardius the sulcus leading from the foramen ovale to the sphenoidal fissure is broad and shallow, but in :(,iphius andMesoplodon it is more canalised.
6) A foramen for the seventh nerve to the rear of the foramen ovale. This
enters the funnel of the auditory meatus and exits through the auditory foramen.
7) The condylar foramen, a small foramen a short distance behind the auditory foramen. This perforates the cerebellar fossa, but has its external opening
in the fissure formed by the basioccipital crest and the exoccipital.
8) The auditory foramen, clearly visible to the rear.

Mandibles (Pl. III, fig. 3)
The mandibles in Berardius are much longer than the rostrum and permanently
expose the large anterior teeth. The distal half of each ramus is comparatively
narrow and the upper and lower profiles in this region are almost parallel; theproximal half expands rapidly to form the posterior half. There is no defined' gonus ' in Berardius as in some other ziphioids. The symphysial union is relatively short and the two halves do not anchylose even in old age as in the rest of thefamily. The mental foramen may be single or multiple and is somewhat erratically situated.
Mandibular teeth (Pl. XII)
Unlike other genera of Ziphiidae, Berardius erupts two pairs of functional teeth
in both sexes, the anterior pair being usually larger than the posterior. They arestrongly compressed laterally. In this, both Berardius and Mesoplodon agree; anddiffer from the rest of the family in which the teeth are circular or oval in crosssection.
Viewed laterally, both pairs are roughly triangular, the base being the shortest
side, while the anterior edge is frequently slightly longer than the posterior. The
lateral faces may be flat or variously undulate with numerous furrows and striae
arising along the base and converging towards the apex. Along the antero-posterioredge, the axis is bent outwards slightly. The tip is an acute, enamelled point surrounded by a ' collar ' of dentine. In old animals the teeth are usually so severely
abraded, that their tops are flat, almost level with the gum.
Occasionally, only a single tooth (anterior) is present in each ramus.
Vertebrae:
The formula of the vertebrae in the two species is as follows:
Berardius arnouxi: C 7 (3+4); Th 10; L 12 (13); Ca. 19=48 (49).
Berardius bairdi: C. 7. (3+4); Th. 11; L. 12. ea. 16+=46+.
True (1910) observed that the skeletons of both species of Berardius are very
similar in most details, but there are some specific differences particularly in thenumber of thoracic vertebrae and of ribs.** Hale (1962) records a specimen of B. arnouxi with eleven ribs on one side.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

C. MaCANN

118

Hector (1878) recorded thirteen lumbar vertebrae in a specimen of B. arnouxi,
which is still in the collection of the National Museum, Wellington. Wellington(Slet Mus. 183) and has unmistakably thirteen lumbar vertebrae. However, twelve
lumbar vertebrae appears to be the normal number in B. arnouxi as shown by otherspecimens in the National Museum, Wellington. True (1910) remarks that the" discrepancy here shown cannot be accounted for at present, but at all events, noneof the formulae of B. arnouxi correspond to that of B. bairdi."
In the number of thoracic vertebrae, the two species differ specifically. B.

Fig. 2. Berardius bairdi Stejneger. Right scapular (714x 525 mm.) (after True, 1910,
pl. 33, fig. 2).

arnouxi has ten and B. bairdi, eleven.
bers of vertebrae respectively.

The numbers of ribs correspond to the num-

Cervical vertebrae: The three anterior cervical vertebrae are completely anchylosed
in the adult to form a single mass in both species. A newly born specimen of B.
arnouxi (Dom. Mus. 614) has only the first two vertebrae completely fused.
Caudal vertebrae: The number of vertebrae in the caudal region appears to varyslightly (eighteen to nineteen appears to be normal). The small terminal elements may easily be lost or dissolved during maeeration.
Chevron bones: There are normally nine chevron bones, the first usually of two
irregular pieces, not united mesially. The fourth and fifth are the largest and bestdeveloped.
Sternum:
The sternum is like that of .Z,iphius and Mesoplodon, but naturally larger. It is
normally composed of five elements, the fifth frequently anchylosed to the fourthand bifid posteriorly.
Scapulae: (Pl. XIII)
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

STUDY OF BERARD/US DUVERN OY

119Apart from size, the shape of the scapulae is remarkably different in the two
species. The acromion and coracoid processes are contrasted; there is a greater
space between the two processes in B. arnouxi than in B. bairdi, and in bairdi theacromion almost completely overlaps the dorsal margin of the coracoid (text-fig. 2).
SCHOOLING
Omura and his colleagues (1955) state that Berardius bairdi forms schools of ten or
more individuals, males appearing in greater numbers than females.
There is some evidence that the breeding school could be dominated by asingle master bull. Linear weals and other wounds occur on the bodies of males,
particularly old bulls, indicating that there is considerable rivalry among the males-the females seldom exhibit such scarring. Adult females of Berardius probably
add their quota of scarring on the bulls for unwanted attentions.
MALE RIVALRY
Pike (1953) referring to scars in Berardius bairdi offers another explanation of scarring"Numerous white scratch marks appear on the back and flanks of both specimens(Pl. II, Ill). The scratch marks when found on males of other Ziphiidae andDelphinidae, have been attributed to the teeth of rival males. This explanation
seems unlikely to be applicable in this species whose teeth are either buried in thegums [young animals] or are so blunt and protrude so slightly. A more plausible
explanation for all species is that the scratches are caused by the sharp ' beaks ' of
the squids on which they feed. Similar scratches are found on the heads of sperm
whales."This explanation overlooks the fact that the teeth are extremely sharp when
first erupted and only become blunt with age, and that scarring by cephalopods
would not be restricted to males, nor would it occur indiscriminately all over the
body. The present author believes the scars are the result of male rivalry, a viewalso held by Omura, Fujino and Kimura (1955). Over several years, I have examined numerous carcases of Ziphiidae and other whales for scarring. I find thatin male Z,iphius linear scars, which are numerous on back and flanks, less common
on the body, appear as single weals or exactly parallel weals, ranging from a few
centimetres to one metre or more in length. They are produced by one or a pairof teeth ripping along the body, the distance between the parallel scars corresponding to the distance between the mandibular teeth at the extremity of the mandible.
Females rarely exhibit linear scarring.
There are also round, oval or elliptical white scars*, particularly by the vent.

* Recently, my suspicions that the elliptical scars, so commonly met with on the bodies of ziphioids,
particularly in the region of the vent, were probably caused by some epizoic crustation were confirmed,
when I asked Dr. J.C. Yaldwyn, the Assistant Director of the National Museum to produce the largest' copepod ' he was aware of in the museum ' collection. This he did and I immediately recognised itas the epizoan responsible for the scars. It proved to be the female of Livoneca ravnaudi (M. Edwards):CYMOTHOIDAE. A paper on the subject will appear shortly under our joint names.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

120

C. McCANN

In ]l,fesoplodon, linear scars are very common on old males, but they are neverexactly parallel as in Ziphius. As the mandibular teeth are nearer the posterior
union of the symphysis in Mesoplodon (except in M. mirus True) both teeth cannot
be brought into play at the same time. (M. mirus could produce scars similar to
those in -?,,iphius). Circular, oval or elliptical scars also appear on the body as in
Ziphius.
In Berardius there are three types of linear weals: (a) a single rip probably
made by a single anterior tooth; (b) parallel line or scars (Pl. II, fig. 5) made by
the two anterior teeth; the distance between the weals being approximately thatbetween these teeth; and, (c) parallel linear scars, one larger and deeper than theother, but the width narrower than to the distance between the two anterior teeth.
These could be produced by both teeth of one ramus making contact at the sametime. All three forms of linear scars have been seen on one old male (Pl. II).
There were also the usual circular, oval and elliptical scars (Pl. II, fig. 5).
The circular, oval or elliptical scars referred to probably result from the attack or
attachment of suctorial animals. Pike (1951) and Nemoto (1955) discuss andillustrate scars made by cyclostomes, particularly the Sea Lamprey (Entosphenus
tridentatus), a species confined to the Northern Pacific Ocean. In the whales I
have examined the scars showed what appeared to be the impressions left by thetransverse lamellae of the discs of sucker-fish. Prolonged attachment could wellbring about permanent discolouration of the skin. Concentrations of scars areinvariably around the vent, which suggests that the fish congregate in the area
to feed on excrement. Squid have been known to scar sperm whales in much thesame manner.
Parasitic crustaceans could also be responsible for some of the scarring, somescars suggesting the erstwhile presence of barnacles, e.g. Coronula. However, whilethese dermal barnacles are commonly epizoic on baleen whles, they have not, as
far as I am aware, been observed on ziphioids. Circular scars could also result
from 'jab ' wounds caused by the tip of an opponent's rostrum.
SYNONYMYBERARDIUS ARNOUX! Duvernoy: The Southern Porpoise Whale
1851.
Berardius Arnuxii
Duvernoy, Ann. Sc. Nat. (3) 15: 51. (Typelocality: Akaroa, Banks Peninsula, New
Zealand. Type in Paris Mus.).
Gray, Cat. Seals & Whales Brit. Mus. p. 348.
Berardius Arnuxi
1866.
Owen, Monogr, Palaeontogr. Soc. p. 31, fig. 11.
.:{,iphius (Berardius)
1870.
Arnouxii
Hector, in Knox, Trans. N. Zeal. Inst. 2: 27.
1870.
Berardius Arnouxi
Hector, Ann. Mag. Nat. Hist. (4) 5: 222. Trans.
1870b. Berardius Arnouxi
N. Zeal. Inst.
Haast, Trans. N. Zeal Inst 2: 190.
1870a. Berardius Arnouxi
Haast, Ann. Mag. Nat. Hist. (4) 6: 348. (Repeat
1870b. Berardius Arnouxiof previous article)
Sci. Rep. Whales Res. Inst.,

No. 27, 1975.

STUDY OF BERARDIUS DUVERNOY

1871.

' :(,iphidae '1871.

Berardius Hectori

1872.
1872.

Mesoplodon Hectori
Berardius Arnouxi

1873.

Mesoplodon J;noxi

1873.
1874.
1876.
1878.
1878.

Mesoplodon Hectori
Mesoplodon Hectori
Berardius arnouxi
Mesoplodon hectori
Mesoplodon hectori

1880.

Berardius arnouxi

1893.
1900.
1900.
1910.
1920.

Mesoplodon hectori
Mesoplodon hectori
Berardius arnouxi
Berardius arnouxi
Berardius arnuxi

1922.
1922.

Berardius arnouxi
Paikea hectori

1924.
1939.
1950.
1957.

Mesoplodon hectori
Berardius arnuxi
Mesoplodon hectori
Berardius arnouxi

1962a.
1962b.
1962.

Berardius arnouxi
Berardius arnouxi
Berardius arnuxi

1974.

Berardius arnouxi

121

Knox & Hector, Trans. N. Zeal. Inst. 3: 125, pls.
13-17.
Gray, Ann. Mag. Nat. Hist, (4) 8: 115-117.
(Type locality: Titahi Bay, N. Zeal., Type Brit.
Mus. 1677b. 76. 2. 16. 3.)
Turner, Trans. Roy. Soc. Edin. 26: 778.
Flower, Trans. Zool. Soc. Lond. 8: 203-234, pls.
27-29.
(part.) Hector in Knox, Trans. N. Zeal. Inst. 5:167 (Part type, Titahi Bay specimen, Brit. Mus.
1677b. 76. 2. 16. 3)
Hector, Trans. N. Zeal. Inst. 5: 170.
Gray, Trans. N. Zeal. Inst. 6: 96-97.
Buller, Trans. N. Zeal. Inst. 8: 407.
Hector, Trans. N. Zeal. Inst. 10: 338, p. 16.
Flower, Trans. Zool. Soc. Lond. 10: 416, pls. 71
and 72 fig. 4.
Van Beneden & Gervais, Osteogr. Cet. pp. 377,
515, Atlas pls. 21-23.
Forbes, Proc. Zool. Soc. Lond. p. 277.
Beddard, Book of Whales, Lond. p. 220.
Beddard, Book of Whales, Lond. p. 228-233.
True, Bull. U.S.N.M. 73: 77.
Marelli, An. Mus. Nac. Secc. Zool. Buenos
Aires. 30: 411-444, pls. 1-5.
Oliver, N. Zeal. Journ. Sci. Tech. 5: 135.
Oliver, Proc. Zool. Soc. Lond. p. 575. (Type,Brit. Mus. 1677b. 76. 2. 16. 3).
Harmer, Proc. Zool. Soc. Lond. p. 542.
Hale, Ree. S. Austr. Mus. 19 (4); 5.
Fraser, Proc. Linn. Soc. Lond. 162: 50, pls. 3, 4.
Taylor, Proc. Zool. Soc. Lond. 129 (3): 325-332,
pl. 3, figs. 1-4.
McCann, Tuatara 10 (1): 13-18, pls. 1, 2.
McCann, Ree. Dom. Mus. 4 (9): 83-94 pls. 1-5.
Hale, Ree. S. Austr. Mus. 14 (2): 231-243, pls.
5, 6, text fig. 1.
McCann, Sci. Rep. Whales Res. Inst., Japan,
No. 26, 145-155, Figs. 1-23, pls. 1-8.
DESCRIPTION

General external characters as in the genus
Teeth (Pl. XII): The anterior mandibular teeth are normally larger than theposterior. Viewed laterally both are roughly triangular in shape, the base being
the shortest side, and the anterior frequently longer than the posterior. The
lateral faces may be flat or undulate with numerous furrows or. striae arising along
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

122

C. McCANN

the base and converging towards the tip. Viewed antero-posteriorly, the vertricalaxis is inclined slightly outwards in its upper portion. The apex is formed of anenamelled point of about 10 to 12 mm surrounded by a 'collar' of dentine. On
the inner face of the enamelled cusp there may be one or two vertical furrows. In
old animals the tooth is worn down to a flat-top level with the gum.
Size: Berardius arnouxi may attain a length of 9.6 m (32 ft). The difference insize between the sexes does not appear as pronounced as in B. bairdi.
Measurements in the flesh, ad. male. Dom. Mus. 1,433 (Col. C. McCann)
Total length 8.50 m; snout to anterior edge of dorsal 6.150 m; Snout togenital 5.60 m; genital to anal 0.30 m; length of penis 0.90 m; circum. of penis
at base 0.23 m; snout to eye 1.50 m; snout to flipper 1.77 m; gape 0.59 m; snout
to blowhole 1.10 m; blowhole 0.13 m; Beak 0.551 m; throat groove 0.55 m; anal
opening 0.23 m; Fluke across 2.444 m.
Neonatals so far recorded do not exceed 3.0 m (11 ft) but the records could
be of animals prematurely born when the true neonatal length be slightly greater.
Colour: " 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." (Haast,1870).
" The colour was black with a purple hue, except a narrow band along the
belly which was grey. The muzzle, flippers and tail lobes were intensely black."(Hector, 1878: 338).
An old male from Pukerua Bay (Pls. I-11), although decomposing, clearly
showed a dirty white dorsal surface. This colour extended from the ' crest ' of thehead to shortly behind the dorsal fin. Parts of the dorsal fin and some surrounding areas were even whiter. The rest of the body was brownish black (Pl.
I and 11, fig. 1). As most observers describe the dorsaum is black, it is possiblethat in this individual the dorsum may have bleached during exposure, after deathand before stranding.

DISTRIBUTION
Berardius arnouxi is apparently confined to the South Pacific and South Atlantic
Oceans, between 30°S and Antarctica. Odd specimens have stranded along theeastern shores of New Zealand as far north as 37°S, but these strandings have beenlater than the normal period when this whale visits the New Zealand area.
Specimens: (See text-fig. 3)
New Zealand Waters: Otago Peninsula, 1840 (Otago Mus. Al4.37 mandibles);Stewart Island (Otago Mus. A24.69, skull), 2 teeth (Dom. Mus. 523); Mason Bay,
Stewart Island, February, 1929 (Dom. Mus. 523); Mason Bay, Stewart Island,
February, 1929 (Dom. Mus. 416, skeleton); Centre Island, Foveaux Strait (Dom.
Mus. 528, worn tooth); Chatham Islands (Otago Mus. A339); Coast near WanSci. Rep. Whales Res. Inst.,
No. 27, 1975.

STUDY OF BERARDIUS DUVERNOY

123

ganui, before 1920 (Wang. Mus. A339); Coast near Wanganui, before 1920 (Wang.
Mus. skeleton); Worser Bay, Wellington, January 1870 (Dom. Mus. 7, skull), 12th
January, 1877 (Dom. Mus. 183, skull); 30th January, 1876 (Dom. Mus. 239, 2
na

178"

17"

17?

Three Kings Is.

NEW ZEALAND-SHOWING-

50

25

O

50

100

1'jJ M.tes

13:1(J)EJ3:S:WO!:E~==:;;10J=~~2CD KILOMETRES

75 Miles to an inch.

g

1870~WELLINGTON'/.

ll!~~~+-~~~~t--~~--t~t\-'-:.--~/'.r--__,,&~~~1-~~~~-f-~~---1~

CHATHAMS

x

17U

Fig. 3.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

17?

174"

17"

Strandings of Berardius arnouxi Duv. in New Zealand.

178'

C. McCANN

124

teeth); Saltwater Creek, north of Banks Peninsula, 27th December, 1873 (Dom.
Mus. 1402, skeleton); New Brighton Beach, Christchurch, 16th December, 1868
(Roy. Coll. Surg., skeleton); Akaroa, Banks Peninsula, 1846 (Type, Paris Mus.),(Brit. Mus. 96.11.20.1, male); South Tapu, Thames Coast, 13th March, 1944
(Auck. Mus.); Te Horo Beach Wellington District, 13th January, 1931, male,
8,400 mm (28 ft); Otaki Beach, Wellington Dist, 4th January, 1934, female, 8,
100 mm (28 ft.), (Dom. Mus. 528, 4 teeth);* Pukerua Bay, Wellington District, 18
th February 1960, old male, 8,850 mm (29-ft), (Dom. Mus. 1433, 2 teeth); **Pencarrow, Wellington Harbour, male 9,000 mm (30 ft, male), female 6,000 (20 ft),ft), (Dom. Mus. 526, 2 teeth); Lyall Bay, Wellington, 1874 (Buller); East Cape,(Dom. Mus. 527, 1 tooth); Titahi Bay, Wellington District, January, 1866, male
2,725 mm (9-ft), (Britt Mus. 1677b 76. 2. 16. 3, Type of B. hectori Gray); Plimmerton, Wellington District, 25th January, 1937, 3,200 mm (10 ft, 8 in.), (Dom. Mus.
614, skeleton); 'New Zealand', (Brit. Mus.), (U.S.N.M. 21511).
Australian waters: Port Lorne, S. Australia, 27th December 1935, 8, 700 mm(29 ft), (S. Austr. Mus. M. 5012)
Antarctic waters: Taylor, April 1955 (Photographs).
Argentine waters: Arroyo del Pescado, Los Talas, 29. 5. 1918, female, 7,800 mm(26 ft), (Mus. Nac. Buenos Aires).
South Atlantic waters: South Shetland Islands, female (Brit. Mus. 1934. 6. 15.
l); Falkland Islands (Brit. Mus. 1949. 8. 19. l); La Plata, 29th May, 1918, female,(Mus. Nac., Buenos Aires).
Of thirty specimens recorded from New Zealand waters, the majority stranded
during the late spring or early summer (December to February-March). Two
neonatals which may have been premature births are recorded as stranding inJanuary 1866 and 1937 (as Mesoplodon hectori (Gray)). The presence of adults andneonatals during the same period of the year suggests that Berardius arnouxi visits the
New Zealand area to breed. On the other hand, photographs of Berardius arnouxi
locked in the ice in April 1955 (Taylor, 1957) show that they return to the Antarctic
region to feed. These observations point to a seasonal north and south migration.
The single recorded Australian specimen, a gravid female was found in thesame period in which Berardius visits New Zealand waters.
The number of Berardius arnouxi visiting New Zealand waters annually is unknown, particularly as their visits do not coincide with the whaling season (Mayto August), and they are not at present of commercial value. Any information we
have is derived from chance strandings.
In the South Atlantic, there appears to be a similar pattern of movement presents itself, but the records are too few for definite conclusions. A single neonatal
has been recorded from one of the smaller islands of the Falklands group (Fraser,1950), but its exact time of stranding is unknown.
As with Mesoplodon, species of Berardius appear to calve near the larger archi-* Porirua Harbour, Wellington District,January 1862, male, 8100 (27 ft), (Fide Knox, 1871: 125)** The sex of these two specimens is in doubt as they were reported and the teeth of the adult collected bya layman.

They may have been a female and calf.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

STUDY OF BERARD/US DUVERNOY

125

pelagos of the world in both hemispheres. B. arnouxi in the New Zealand and South
Atlantic archipelagos and B- bairdi around the Japanese Archipelago.
FOOD

Haast (1870) notes that the stomach contents of one animal contained about halfa bushel of the horny beaks of cephalopods (squids). Haast mentions ' Octopus ',but this is hardly likely, as the octopus is an inshore animal, whereas squids are
pelagic and often occur in vast shoals.
ANATOMICAL OBSERVATIONSCranium (Pls. III, IV): The cranium of the adult has been well described and figured by Duvernoy (1851) and Flower (1872).
The skull of Berardius exhibits almost perfect symmetry (Flower 1872). The
same symmetry appears in the newly-born young. The large bulbous nasals area prominent feature of the vertex in the adult, but the nasals are not so conspicuous in the newly born (Pl. IV, fig. 1). The extremities of the premaxillae rise above
the nasals and are slightly curved forwards, not so recumbent as in the adult, nor dothey overhang the narial cavity as in some species of Mesoplodon. Further, they do
not twist outwards as asymmetrically as in Mesoplodon; they are almost equal insize, the right premaxilla being very slightly larger than the left.
The recurved walls of the vomer, with the mesethmoid bone sandwiched between them contribute to the formation of the narial septum, the mesethmoid bone
eventually emerging anteriorly into the mesorostral groove as a free projection
between the margins of the premaxillae. The ossified projection of the mesethmoid
terminates as an irregular, rough or nodular projection, extending as far forwards
as a transverse line drawn through the middle of the maxillary crest. In the flesh,
the mesethmoid bone is continuous with the mesethmoid cartilage and fills themesorostral groove. In Berardius the mesorostral groove is permanently open andis never partially nor wholly filled by bone as in some species of Mesoplodon. Posteriorly, the vomer forms a broad ' bracket ' beneath the large nasals with a large
blind pit (BP) on either side of the septum. The shape of the ' bracket ' and thepresence of the blind pits are distinctive of Berardius.
Anteriorly, the vomer extends about two-thirds the length of the rostrum
forming a trough for the mesorostral cartilage. Ventrally, much of the anterior
end of the vomer is visible, wedged between the premaxillae. This condition is
also found in the newly-born young (Pl. IV, fig. 2, Vo).
In dorsal view, the rostrum is somewhat spearshaped. No distinct maxillaryturbercle is present but there is a maxillary crest. In side view, its dorsal profileis straight; the lower gradually descends from its anterior extremity towards thepterygoids. In ventral view, the broad maxillary prolongations are seen embracing the premaxillary extremities with the vomer wedged between. In transverse
section, the rostrum is considerably broader than deep near the middle.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

126

C. McCANN

The adult skull is broader than high, but in the newly-born the height is greaterthan the breadth.

Foramina (Pl. IX, fig. 1): The premaxillary and maxillary foramina are on approximately the same transverse level. The premaxillary is very much smaller
than the maxillary which frequently has secondary foramina around it. The postcrest foramen and the secondary foramina are not pronounced in the neonatalskull.
Antorbital region (Pl. IX, fig. 2): The antorbital tubercle is exaggerated by thepresence of a deep antorbital notch. In dorsal view, the tubercle is seen to beformed by the maxilla and a small portion of the malar bone, with the anterior
extremity of the frontal, a short distance behind forming the orbit. In side view,
the broad anterior end of the frontal is shortly behind the end of the maxilla andthe anterior projection of the malar bone. At the lower corner of the frontal, anarrow portion of the lachrymal bone is visible but not on a level with the lateral
margin of the frontal. Ventrally, the malar bone alone appears to form the tubercle; the lachrymal bone falls short of the lateral margin of the frontal. Thereis a triangular wedge between the frontal and the malar.
The antorbital notch is margined by the maxilla alone.
In the newly-born, the arrangement is somewhat different. In dorsal view,the malar borders the antorbital notch; the lachrymal is just visible in front of theanterior extremity of the frontal. The orbital portion of the frontal is not laterally
expanded as in the adult, but forms a narrow margin to the maxilla behind thetubercle. In side view, the large ' cuboidal ' end of the lachrymal abuts the anterior end of the frontal and a very small portion of the malar is also visible. Ventrally, the lachrymal and malar bones predominate in the formation of the tubercle
and malar bones predominate in the formation of the tubercle to the exclusion of
the anterior end of the frontal and maxilla.
Mandibles: The extremity of the mandibles protrudes considerably beyond therostrum, so much so that the anterior teeth are outside the mouth when it is closed.
The distal half of the mandibles is comparatively narrow and the upper and lower
profiles are almost parallel. The proximal half expands rapidly to form the posterior
portion of the jaw. The symphysial union is comparatively short and the two
halves appear never to fuse together as in some other genera. The mental foramen
may be single or multiple and somewhat irregularly situated on either side, even in
the same individual.
Eyes: The relative size of the eye ' sockets,' in relation to the length of the skull,is smaller than in either Ziphius or Mesoplodon. The jugal process is accordingly
comparatively short.
Ear Bones (Pl. XI):

The bulla and the periotic of an adult measure 73 by 48 mm
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

STUDY OF BERARD/US DUVERNOY

127

and 80 by 45 mm respectively (Dom. Mus. 1406). In the newly-born the same bones
in the same order measure 54 by - - mm, periotic not stated (Brit. Mus. 1677b. 76.
2. 16. 3); 51 by - - mm, periotic not stated (Brit. Mus. 1949. 8. 19. l); 51.5 by
36 mm and 56 by 32 mm (Dom. Mus. 614).
Scapula (Pl. XIII): In B. arnouxi the vertebral border is more or less straight, notconves as in B. bairdi; both the anterior and posterior borders are more verticalthan in B. bairdi. The anterior border is almost straight, whereas in B. bairdi it isconvex dorsally, and slightly concave ventrally; the posterior border is less concave than in B. bairdi. The anterior spinous fossa is relatively narrower than in
B. bairdi.
The acromion is almost level with a vertical dropped from the anterior angle;
the upper and lower profiles are, at first, almost parallel but gradually diverge towards the extremity; in some specimens the extremity is decidedly wider. In B.
bairdi, the acromion falls short of a vertical dropped from the anterior angle andthe lower profile turns abruptly upwards producing an almost semicircular anterior
margin, which gives the entire process a somewhat circular or spiral border.
The coracoid and acromion processes are subequal in length whereas in B.
bairdi the former exceeds the latter. In dorsal view the two processes are divergent,but in B. bairdi there is very little space between the two. The coracoid is narrow
with both upper and lower profiles slightly concave; in B. bairdi the process is broadand slightly curved upwards with the upper profile concave, the lower convex.

SYNONYMYBERARDIUS BAIRD/ Stejneger: Japanese Porpoise Whale
1883. Berardius bairdi
Stejneger, Proc. U.S.N.M. 6: 75-77. (Type locality:
Bering Island; type U.S.N.M. 20992).
Malm, Bihang K. Svenska Akad. Handl. 8 (4): 109.
1883. Berardius vegae
Stejneger, Journ. Amer. Geogr. Soc. N.Y. 18: 317-328.
1886. Berardius bairdi
Beddard, Book of Whales, Lond. p. 233.
1900. Berardius bairdi
True, Bull. U.S.N.M. 73: 60, pls. 26-33. 34, fig. 7, 35,
1910. Berardius bairdi
fig. 7; 36, fig. 7; 39; 42.
Kellogg, Journ. Mamm. 12: 73-77.
1931. Berardius bairdi
Matsuura, Zool. Mag. Zool. Inst. Fae. Sci. Jap. Univ.
1942. Berardius bairdi
54: 466-473.
Matsuura Marine Mamm. (Kaiju). 298 pp.
1943. Berardius bairdi
Scheffer & Slipp, Amer. Midl. Nat. 39 (2): 266.
1948. Berardius bairdi
Scheffer, Pacific Sci. 3: 353, fig. 1.
1949. Berardius bairdi
Pike, Journ. Mamm. 34: 98-104, pl. 1.
1953. Berardius bairdi
Slipp & Wilke, Journ. Mamm. 34: 105-113, pls. 1, 2.
1953. Berardius bairdi
Omura, Fujino & Kimura, Sci. Rep. Whales Res. Inst.
1953. Berardius bairdi
J ap. 10: 89-132 (per errorum baildi)
Omura, Sci. Rep. Whales Res. Inst. Jap. 13: 213-214,
1958. Berardius bairdi
figs. 1, 2.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

C. McCANN

128

DESCRIPTION (PL. XIV FIG. 1)
General external characters as in the genus. There is no marked difference in thesize and shape of the mandibular teeth in either sex; (they are erupted in both sexes
alike on reaching maturity (Omura, 1955). Pike (1953) examined an 8.7 m (29 ft)
females which he regarded as both physically and sexually immature, and found theteeth were " completely buried." His observations support those of Omura (1955 ).
The largest teeth on record are: (anterior) vertical height 89; antero-posterior
width 103 mm; lateral diameter--; (posterior) vertical height 60 mm; anteroposterior width 37 mm; lateral width ? for a male of 9.9 m (33 ft); (interior) vertical height 93 mm; antero-posterior width 72 mm; lateral diameter?; (posterior)vertical height 55 mm; antero-posterior width 33; lateral width ? ; for a 8. 7 m(29 ft) female (Pike, 1953).

Size:

B. bairdi is up to 3.0 m (10 ft) longer than B. arnouxi.

female measured 12. 6 m (42 ft).
than females.

The largest known
Males are about 2.0 m (approx. 7 ft) smaller

Colour:Pike ( 1953) describes a male and female as follows: "In these specimens
the body is black on the head, back, flukes and flippers. The undersurface is of aslightly lighter shade, with white patches at the umbilicus and some white and grey
markings. The male has a conspicuous d~amond-shaped white patch at the umbilicus and some white around the genital opening. The female is lighter incolour and the skin was covered by abrasions caused while the whale was being
towed into the station. Numerous white scratch marks appear on the back andflanks of both specimens (Pls. I, II, V)."
Omura et al. (1955) state: "According to our observations, however, color ofwhole body is pure black or somewhat lighter, even on the under surface of the
body is pure black or somewhat lighter, even on the under surface of the body. It
is not certain, but most whalers say that there are two groups of Berardius, one being
black and another slatish. As there are many white scratch marks (Pls. I-II) on
the skin of Berardius, especially in old bull, the colour looks more lighter than it isin such whales. There are white patches in three regions of the ventral side of thebody, i.e. on the throat, between the flippers and at the umbilicus (Pls. II-IV,
Pl. III, fig. 2). Every Berardius seems to have the white patch on the throat or
between the flippers, there are considerably individual differences."
DISTRIBUTION

Berardius bairdi is restricted to the Northern Pacific Ocean; concentrating in Japanese waters during the breeding season in spring and summer. Occasional strayindividuals or small schools are found off the North American coast.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

STUDY OF BERARDIUS DUVERNOY

129

Specimens: Bering Island (Type, U.S.N.M. 20992); St. George Island, ad. female(U.S.N.M. 49726); Centerville, California, ad. male (?), (U.S.N.M. 49725); St.
George Island, young male, ( U.S.N.M. 49727); Bering Island ( ?), (Grebnitzki ?)( U.S.N.M.); about 10 miles off Kains Island, Quatsino Sound, 5th July, 1950, ad.
male, 9.9 m (33 ft), (Pike, 1953); 20 miles E.S.E. of Cape St. James, 9th August,
1951, imm. female, 8.7 m (29 ft), (Pike, 1953); one mile south of Ocean City,
Washington, July 1950, 16.5 m (35 ft), (Slipp and Wilke, 1953); Japan, Nat. Sci.
Mus., 10.8 m (36 ft); Santa Cruz, California, June, 1925 (Mus. Calif. Acad. Sci.
skull);COMMERCIAL WHALING
In summer and autumn, B. bairdi concentrates in Japanese waters north of the 30°N
parallel. The Japanese whaling season is from April to November (Omura et al,
1955: 93 ), reaching its peak in the summer months of July and August. Omura
and colleagues found the main concentration in the southern part of the range off
the east coast of Japan proper, the whales moving in northwards, according to thewhalers, as the season advances. After breeding, they apparently leave for feeding
grounds in more northern latitudes.
Whaling commences in the northernmost area in the Okhotsk Sea earlier thanelsewhere and has a second peak in autumn but it is supposed that there is a different
population or community in the Okhotsk Sea from elsewhere.
The small numbers taken along the western shores of Japan may be due tofew schools passing west through the Soya and Tsugaru Straits.
The American season is from April to September or October, but as Berardius
is a sporadic visitor it is less important than in Japan (Scheffer and Slipp, 1948:
256).
BREEDING
Omura et al (1955, fig. 28) extrapolate length records to arrive at the following" the pairing of Berardius takes place in February and the parturition in December
in most individuals, length of gestation being about 10 months." Omura (1958)
however, records an anomalous 12 mm long embryo taken in August, and True(1910) records a newly-born calf discovered in June.
The neonatal length, in most cetaceans, is approximately one-third the lengthof the parent. Accordingly, in Berardius bairdi which can grow to about 12.6 m(42 ft), the newly-born calf should approximate 4.2 m (14 ft). Omura et al (1955)
estimated the neonatal length as 4.5 m (15 ft). The largest foetus they record wasin September and measured 4.2 m (14 ft). Stejneger (True, 1910: 64) records
a newly-born calfof 4.8 m (16 ft) washed up on Bering Island on June 5, 1883, withthe remains of the umbilicus still adhering.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

130

C. McCANN

FOOD

Pike ( 1953) records the stomach contents of a young female as: " about two gallons
of small rockfish and squid remains."
PARASITES

Omura et al. (1955) record Conchoderma sp. attached to the teeth (Pl. XIV, 2). Thisbarnacle has been recorded on the teeth of Hyperoodon ampullatus and Physeter catodon(Wolff, 1960), and I have found it on a tooth of Mesoplodon grayi. A species of Conchoderma is also found commonly on Megaptera nodosa, never on the animal itself but
attached to the barnacle Coronula, so commonly found on the Humpback. Unlike
Coronula, Conchoderma must have a hard substrate to attach itself to.
Pike (1953) recorded nematodes in the stomach of a specimen of B. bairdi.
ACKNOWLEDGMENTS

The author wishes to thank Dr. R. A. Falla, Director, Dominion Museum, Wellington, for the opportunity of undertaking the work; Dr. R.R. Forster, Director,
Otago Museum, Dunedin, for the loan of material; Dr. R. K. Dell and Mr. B. Hamlin for valuable comments and suggestions on the manuscript; Mr. P. M. Hedgland,
Photographer, Dominion Museum, for the excellent anatomical photographs; Dr.
H. Omura, Whales Research Institute, Japan, for permission to reproduce thephotographs of Berardius bairdi; and Miss M. Hall, Librarian, Dominion Museumat the time, for help in obtaining literature.
REFERENCESBEDDARD, F. E., 1900. A book of whales. London, 320pp. 40 illustrations: 228-233.
BENEDEN, P. J. VAN, and F.L.P. GERVAIS, 1868-1880. Osteogr. cet. osteographie des cetaces vivants et fossiles.
Paris, pp. 377, 515, Atlas, pls. XXI-XXIII.
BULLER, W. L., 1876. Presidential address 7th August, 1875; Trans. N. Zeal. Inst., 8: 407.
CARCELLES, A., 1948. Los cetaceos en las aquas argentinas. Mus. Arg. cien. Nat. (Pub. Extra), 238: 6.
DuvERNOY, G. L., 1851. Memoire sur les caracteres osteologiques des genus nouveaux ou des especes nouvelles de oetaces vivant ou fossiles. Ann. Sci. Nat., 3(15): 1-71.
FLOWER, W. H., 1872. On the recent ziphioid whales, with a description of the skeleton of Berardius arnouxi.
Trans. Z,ool. Soc. Lond., 8(3): 203-234, pls. XXVII-XXIX.
FLOWER, W. H., 1878. A further contribution to the knowledge of the existing ziphioid whales: genus
Mesoplodon. Ibid., 10(11): 416, pls. LXXI-LXXII fig. 4 (M. hectori).
FORBES, H. 0., 1893. Observations on the development of the rostrum in the cetacean genus Mesoplodon,
with remarks on some of the species. Proc. ,(,ool. Soc. Lond., 216-236, pls. XII-XV. (M. hectori).
FRASER, F. C., 1950. Note on a skull ofHector's beaked whale, Mesoplodon hectori (Gray) from the Falkland
Islands. Proc. Linn. Soc. Lond., 162: 50-52, pls. III-IV, figs. 1-5.
GRAY, J.E., 1866. Cat. Seals, Whales and dolphins in the British Museum, 384 pp.
GRAY, J.E., 1871. Notes on the Berardius of New Zealand. Ann. Mag. Nat. Hist., (4)8: 115-117.
GRAY,]. E., 1874. List of seals, whales and dolphins of New Zealand. Trans. N. ,(,ea[. Inst., 6: 96-97.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

STUDY OF BERARD/US DUVERNOY131

HAAST, J. voN, 1870. Preliminary notice of a ziphid whale, probably Berardius arnuxi, stranded on the 16th
December, 1868 on the beach near New Brighton, Canterbury. Ibid., 2: 190-192.
HAAsT,j. VON, 1870. Ditto. Ann. Mag. Nat. Hist., (4)6: 348 (Repeat of previous article).
HALE, H. M., 1939. Rare whales in South Australia. S. Austr. Nat., 19(4): 5-6, photo.
HALE, H. M., 1962. Occurrence of the whale Berardius arnuxi in Southern Australia. Ree. S. Austr. Mus.,14(2): 231-243, pls. V-VI, fig. 1.
HARMER, S. F., 1924. On Mesoplodon and other beaked whales. Proc. Zool. Soc. Lond., 541-587.
HECTOR,j., 1870. List of the bones of Seals and whales in the Colonial Museum, Wellington, New Zealand.
Ann. Mag. Nat. Hist., (4)5: 220-224.
HECTOR, J., 1872. On the whales and dolphins of New Zealand seas. Trans. N. Zeal. Inst., 5: 155-170,
pls. I-VI.
HECTOR,j., 1878. Notes on the whales of New Zealand Seas. Ibid., 10: 331-343, pls. XVI-XVII.
KELLOGG, R., 1931. Whaling statistics for the Pacific Coast of North America. Journ, Mamm., 12: 73-77.
KNOX, F.J., and J. HECTOR, 1871. Observations on the Ziphiidae, a family of cetacea. Trans. N. Zeal.
Inst., 3: 125-129.
MALM, A. W., 1883. Skelettdelar af hval, insamlade under expeditionear med Vega 1878-1880. Bihang.
K. Svenska Akad. Hand!., 8(4): 109.
MATSUURA, Y., 1942. On the northern Porpoise whale, Berardius bairdi Stejneger, in waters near Boshu.
Zoo[. Mag., Zoo!. Inst., Fae. Sci. Tokyo Imp. Univ., 54: 466-473.
MATSUURA, Y., 1943. Berardius bairdi Stejneger ranges the Sea of Okhotsk. Ibid, 55: 237.
MATSUURA, Y., 1943. Marine mammals (Kaiju). Tokyo, 298 pp.
MARELLI, C. A., 1920. Revision osteologica de Berardius arnouxii Duv. An. Mus. Univ. Nac. Buenos Aires.,30: 411-444, pls. I-V.
McCANN, C., 1962. The taxonomic status of the beaked whale, Mesoplodon hectori (Gray)-Cetacea. Ree.
Dom. Mus., 4(0).
NEMOTO, T., 1955. White scars on Whales. (I) Lamprey Marks. Sci. Rep. Whales Res. Inst., 10: 69-77,
figs. 1-10.
NISHIMURA, S., and M. NISHIWAKI, 1964. (Berardius bairdi Stejn.). Pub!. Seto Mar. Biol. Lab., Art. 25, pl.
XXI, fig. 1 (only).
NISHIWAKI, M., and N. 0GURO, 1971. Baird's beaked whales caught on the coast of Japan in recent 10
years. Sci. Rep. Whales Res. Inst., 23: 111-122. pls. 1-11.
0HSUMI, S., 1964. Comparison of maturity and accumulation rate of corpora albicantia between left andright ovaries in cetacea. Ibid., 18: 123-147, pl. I.
OLIVER, W.R.B., 1922a. The whales and dolphins of New Zealand. N. Zeal. Journ. of Sci. & Tech., 5(3):129-141.
OLIVER, W.R.B., 1922b. A review of the cetacea of New Zealand seas. Proc. Z,ool. Soc. Land., 557-585, pls.
I-IV.
OMURA, H.K. FUJINO, and S., KIMURA, 1955. Beaked whale Berardius bairdi of Japan, with notes on Z,iphiuscavirostris. Sci. Rep. Whales Res. Inst. Japan., 10: 89-132, pls. 1-X, figs. 1-30.
OMURA, H., 1958. Note on embryo of Baird's beaked whale. Ibid., 13: 213-214, figs. 1-2.
OWEN, R., 1870. Monograph on the British fossil cetacea from the Red Crag. Monogr. Palaeontogr. Soc., 140, pl. 1-V.
PIKE, G. C., 1953. Two records of Berardius bairdi from the coast of British Columbia. Journ. Mamm., 34:
98-104, pl. I.
SLIPP, J. W., and F. WILKE, 1953. The Beaked Whale Berardius on the Washington Coast. Ibid., 34: 105113, pls. 1-11.
SCHEFFER, V. B., andJ. W. SLIPP, 1948. The whales and dolphins of Washington State with a key to thecetaceans of the West Coast of North America. Amer. Mid!. Nat., 39(2): 257-337.
SCHEFFER, V. B., 1949. Notes on three whales from the Aleutian Islands. Pacific Sci., 3: 353, fig. 1.
STEJNEGER, L., 1883. Notes on the natural history of the Commander Islands including descriptions of
new cetaceans. Proc. U.S.N.M., 6: 75-77.
STEJNEGER, L., 1886. On the extermination of the great Northern Sea-cow (Rytina) a Reply to Professor
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

132

C. McCANN

A. E. Nordenskioeld. Journ, Amer. Geog. Soc. N.Y., 18: 317-328.
An unusual record of three species of whales being restricted to pools in Antarctic
sea-ice. Proc. :(,ool. Soc. Lond., 129(3): 325-332, pls. I-VII.
TRUE, F. W., 1910.
An account of the beaked whales of the family ziphiidae in the collection of the UnitedStates National Museum, with remarks on some specimens in other American Museums. Bull. U.S.
N.M., 73: 1-89. pls. I-ILi.
TURNER, W., 1872. On the Occurrence of :(,iphius cavirostris in the South Shetland Seas, and a cpmparison ofits skull with that ofSowerby's whale (Mesoplodon Sowerbyi). Trans. Roy. Soc. Edin., 26: 759-778, Append.
779-780.
WOLFF, T., year.
Rankefodderne Conchoderma og Coronula pa hvaler. Saertryk af 'Flora og Fauna', 66(1):1--8, 6 photos.
TAYLOR, R.H.F., 1957.

Sci. Rep. Whales Res. Inst.,

No. 27, 1975.

133

STUDY OF BERARDIUS DUVERNOYAPPENDIX I.
A

2
3
4
5
6

B

1392
894
1080
494 !
684
748

7

8

150

c

D

E

F

G

H

I

J

1420
870+
1040
560
720
700
445
169

1240
820+
970+
555
623
675
380
166
108
205
199
87
235
350
78
215
156

1193
780
918
455
615
610
405
158
90
201
195
75
210
370
65
194
137
1065
262
202

1262
800+
1000+
570
710
710
430

1220
780+
900+
518
660
640
410
172
92
192
183
90
200
340
65
186
133
1055
250
213

1200
785
910
475
620
620
395
160
95
180
195
90
202
350
82
192
128
1050
280
193

1270
865
1040
530
625
660

1174
800

9

10
11 21012 10213
14
15
16 21317
18 1236
19 29420 222

211
219
92
240
390
71 ·

1245
350
230

MEASUREMENTS OF SKULLS

218
168
1240
327
210

184

95
208
221
82
215
415
73

194
166
1158
287
212

APPENDIX II.
A

1378
2 880
3
4

5

666

6
7

8

188

B

c

1524
960
1185
563
766
750

1423
925
1130
544
682
675

207

197

215
235
110

195
217
98

165

85
261
193
1334
292

82
228
171
1289
295
230

D

1062(?)
578%
720+
530
520

E
1474
1025
1187
575
716

740
223

155
80

493
577

584
378
149

189

91
180
380
63

80

72
186

1011
245

MEASUREMENTS OF

G

H

I

1440
908
1146
569
693
732
447
208

1343
870
865
525
657
660
397
194

1438
923
1122
582
757
748
445
206

1421

204
238
98

203
239

215
246
102

194
230
98

86
246
180
1334

72
216
173
1190

78
221
170

241

212

222

F

1170
571

686

201

9

10 181
11 238
12
96
13
14
84
15
16 24017 16818 1282
19 145+20 223271

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

187

197
239

83

JOO

83
195
142
883+
145+
175

235
178
1360
310
245

71

94

238
163
1300
276
234

134

C. McCANN

OF BERARDIUS ARNOUX! (MILLIMETERS)
K

L

M

N

1350
903
1078
525
662

1372
919
1097
533
625
671

587E
343
443E
235
228
258

150

152

37

203
69

208
74

49

62
188

61
191

90

1225
303
204

1245+
310
211

481
155
83

505+
280+
383+
252
207
238
140
46
37
82
116
49
84
197
38
93
59
440
125
92

0
601
358
469
240
228
242

p
1260
765
648(?)

35

700
435
168

50

200
218
98

91
498
161
91

220
154
1155
290
230

Type Paris Mus.
Otago Mus. Al4.31
c. Otago Mus. A24.69
D. Otago Mus.
E. Dom. Mus. 7
F. Eom. Mus. 416
G. Dom. Mus. 143
H. Dom. Mus. 1402
Mus. Nac. Bs. AiresI.
J. S.R.N.M. 21511
K. Flower 1868 R.C. 3
L. True
M. Brit. Mus. 1677G, 76.2.16.3
N. Dom. Mus. 614
o. Brit. Mus. 1949-8.A.l
P. S. Aust. Mus. M.5012
A.
B.

BERARDIUS BAIRD! (MILLIMETERS)

A.
B.
C.
D.
E.
F.
G.
H.
I.

Type U.S.N.M. 30992
U.S.N.M. 49736? U.S.N.M. 49725
U.S.N.M. 49727
Bering Is!. U.S.N.M.
33 ft. (Pike, 1953)
29 ft. (Pike, 1953)? 34 ft. 5 in. (Slipp & Wilke, 1953)
36 ft. (Omura, Fujino, & Kimura, 1955)

Key to the numbers, 1 to 20, in the appendix tablesof measurements.
1. Total length of skull; 2. Length. of rostrum;
3. Tip of rostrum to posteror median margin ofpterygoids ; 4. Height of vertex to pterygoids, vertical ; 5. Breadth between centre of orbits ; 6.
Breadth between zygomatic processes ; 7. Breadth at
maxillary notches ; 8. Breadth of rostrum at middle ;JO. Greatest breadth of premaxillaries, proximally ;11. Greatest breadth of premaxillaries in front ofanterior nares; 12. Greatest breadth of anteriornares; 13. Length of temporal fossa; ]4. Breadth
between temporal fossae; 15. Breadth of foramen
magnum; 16. Width of occipital condyles; 17.
Height of occipital condyle; 18. Length of mandible; 19. Length of symphysis; 20. Greatest depthof mandible.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

STUDY OF BERARD/US DUVERNOY135

EXPLANATION OF PLATESPLATE I
Berardins arnouxi Duvernoy stranded in Pukerua Bay, New Zealand.

Photos E. Pain.
PLATE II
Old male of Berardius arnouxi Duvernoy.
Fig. 1. Stranded animal; Fig. 2. Head showing frontal bas, beak and gular grooves;
Fig. 3 Blowhole and scarring; Fig. 4. genital and anal area; Fig. 5. Dorsal fin, newand old scars and wounds; note the parallel scars and possible shark teeth marks(left lower corner).
Photos McCann.
PLATE III
Skull of adult Berardius arnouxi Duvernoy.
(See index to lettering, page 137).
Photos P. M. Hedgland.

Courtesy National Museum.

PLATE IV
Skull of neonatal Berardius arnouxi Duvernoy.
(See index to lettering, page 137).
Photos P. M. Hedgland.

Courtesy National Museum

PLATE V
Sagittal section of adult Berardius arnouxi Duvernoy.
Figs. 1. and 2. general aspect. Fig. 3. Narial region; note' blind pit' (BP) in vomer.
(See index to lettering, page 137)
Courtesy National Museum
Photos P. M. Hedgland.
PLATE VI
Posterior aspect of Skulls of Berardius arnouxi Duvernoy.
Fig. I. neonatal; Fig. 2. adult.
(See index to lettering, page 137)
Courtesy National Museum
Photos C. Hale Fig. I. & P. M. Hedgland Fig. 2.
PLATE VII
Antorbital region of adult Berardius arnouxi Duvernoy.
(See index to lettering, page 137)
Photos P. M. Hedgland.

Courtesy National Museum

PLATE VIII
Antorbital and aural region of neonatal Berardius arnouxi Duvernoy.
Fig. I. Antorbital foramina and optic foramen; note only foramen No. 2 is visibleat this stage. Fig. 2. aural area.
(See index to lettering, page 137).
Courtesy National Museum
Photos P. M. Hedgland.
PLATE IX
Formation of Antorbital tubercle and associated bones in Berardius arnouxi
Duvernoy (adult).
Fig. I. Dorsal aspect; Fig. 2. ventral aspect; Fig. 3. orbital region.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

136

C. McCANN(See index to lettering, page 122).
Photos P. M. Hedgland.

Courtesy National Museum·

PLATE X
Adult Berardius arnouxi Duvernoy.
Fig. I. Brain cavity; Fig. 2. Aural region.
(See index to lettering, page 122).
Photos P. M. HedglandCourtesy National Museum

PLATE XI
Adult and neonatal tympanic bones contrasted.
A-adult; B-neonatal.
(See index to lettering, page 122).
Photos P. M. HedglandCourtesy National Museum

PLATE XII
Teeth of Berardius arnouxi Duvernoy.
Upper set, inner faces; centre, outer faces of same set; lower, antero-posterior faces.
Courtesy National Museum
Photos P. M. Hedgland
PLATE XIII
Scapulae of Berardius arnouxi Duvernoy.
Fig. 1. Dorsal aspect of adult and neonatal.
Fig. 2. Ventral aspect of same pair.
Photos P. M. Hedgland.

Courtesy National Museum

PLATE XIV
Berardius bairdi Stejneger.

Fig. I. Adult animal; Fig. 2. Extremity of rostrum; note Conchoderena attached toteeth; Fig. 3. Teeth of adult and of a 2,700 mm foetus; top row, anterior, bottomrow, posterior.
PhotosCourtesy Dr. H. Omura.
PLATE XV
Adult skull of Berardius bairdi Stejneger.
(See index to lettering, page 122).
PhotosCourtesy Dr. H. Omura.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

STUDY OF BERARD/US DUVERNOY137

KEY TO LETTERING AND FIGURES ON PLATES
Ac-Acromion.
ALS(AS)-Alisphenoid bone.
AN-Antorbital notch.
AOF-Antorbital foramen.
AT-Antorbital tubercle.
AU(auf)-Auditory foramen.
B(ty)-Tympanic bulla.
BOC-Basioccipital crest.
BP-' Blind pit' in posterior wall ofvomer (nares).
BS-Basisphenoid bone.
Ca-Carotid foramen.
CE-Condylar foramen.
Cor-Coracoid.
EO-Exoccipital bone.
ET(e)-Eustacian tube.
F-Frontal bone.
FC-Falx cerebri.
FM-Foramen magnum.
FO(fo)-Foramen ovale.
FP-Falciform process.
FS(fl)-Sphenoidal fissure.
J-Jugal process.
L-Lachrymal bone.
LRC-Lower rostral canal.
M-Malar bone (jugal).
MAS-Mastoid process.
MCF-Post maxillary crest foramen.
MD-Mandible.
ME(MES)-Mesethmoid bone.
MF-Mandibular foramen.
MP-Maxillary pit.
MRC-Median Rostral canal.
MRG-Mesorostral groove.
MX-Maxilla.
MXC-Maxillary crest.
MXF-Maxillary foramen.
N-Nasal bone.
NF-Narial foramen.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

OBS-Orbito-sphenoid bone.
QC-Occipital condyle.
OF(OPF)-Optic foramen.
OS-Orbitosphenoid.
Pa-Parietal.
Pal.-Rid.-Palatine ridge.
PCF-Post crest foramen.
PE(per)-Periotic bone.
PL(Pl)-Palatine bone.
PMF-Post maxillary foramen.
PMX-Premaxillary.
PMXF-Premaxillary foramen.
PS-Presphenoid.
PT-Pterygoid bone.
S-Nasal septum.
SO-Supraoccipital bone.
SQ-Squamosal bone.
Ty(B)-Tympanic bulla.
Vo.-Vomer.
URC-Upper rostral canal.
Ear bones:
a-Posterior articular surface between bones.
a'-Anterior articular surface between bones.
af-Aquaduct of Fallopius.
al-Anterior lobe of periotic bone.
e-Eustacean tube.
earn-External auditory meatus.
epl-Exterior posterior lobe of tympanic.
fo-Fenestra ovalis.
fr.-Fenestra rotunda.
ipl-Internal posterior lobe of tympanic.
m-Mastoid process.
per-periotic bone.
pi-Posterior lobe of periotic.
ty-Tympanic bulla.
Numbers: Nos. I, 2 and 3 refer to foramina in the
antorbital region.

I
EI
PLAT

.McCAl'¥N
C

,
.
t
s
n
s.I
sRe
p.Whale
i.Re
c
S

.
5
7
9
,1
7
.2
o
N

＠

4

fL6l LZ・oN
・

S
'
!
y"
a
f
M ・r
1
1
v
7
a
1ロHs
1・

し 叩

一 −
・

l
l
Ji
百/

4X凶 d
ゴ）（~ぜ ’，
／

JXW
:

ω−
d

＠

)
:
NNV::PW"I宜上V寸dI
I

V
PLATE I

.McCANN
C

,
.
t
s
1
1
./
s
sRe
.Whalepi.Re
cSλ1975.
No.2

ムV寸d
l
八：

G'OJ{
f
f'
日 6
口S
y.
a
f
!
M・
1
1
v
7
a
ys
a
s
ザ・
z
"1s

N~V:)O ~吋＇：）PLATE VJ.McCA~N
C

＠

l.
s
1
1
.l
s
sRe
e
l
ha
ペ
I
.I
p
.Re
i
c
S

.
5
,197
7
Xo.2

.V1d
l
l.
.
八 '
1
I

「
OA
fZど5・
6
L
§
.
y・ns
a
l
II≪
J
f
'
l
l
/
3
If
:
J
'
3
f
r・
n
r
1
・

~'.\J V:):>J'J

)
:
.

l
l
I
しATE ¥'》
l.M cCA~N
C

.
t
s
n
I
s.
sRe
e
l
a
.Wh
p
.Rt
i
c
S
.
5
,197
7
λ匂.2

I
.
I
.
XI'

γ
υ
;・
a
"YL61・

・
S
.
J
!
U"
1
/
1
I・
A
/
l
J
I
/
J
U'
J
s
ザ・
＂SI
／
・

可・0
Ol
>
:
:
:
V
N
!
¥
.
'

C
.McCAN:¥'

PLAT" X

・
t
.~

o
c

S
r
i.
Re
/
J
.W
h
a
l
e
sRe
s
.1
1
1
.
<
I
.,
No.27
.1
97
.
'
i
.

＠

g

i
'
l
r
J

a・，＇.＼.
I・
ι
f
:
̲
≪
S
J
!
u"
1
/
i
If・
J
/
l
J
I
/
J
fS
l
J
¥
•"/HI/ .

v
d

、
ぜ
令

e

』1uJ」wd1u:L1いいんふ!ulu~:iム~wJ，ムムいいl叫
y

＠
、
3
1
J<
g

,9

叫が

d
l
Vr
L
.
J
IX/

~.＇－： VOO附・0

C 恥1cCAN NI
PLATE XI

i
.
r

総

a
,̲.
}・
.
!
f
・
−
.￡

J五.
̲
.

E

＠

＠

/.
s
n
.I
s
sRe
e
l
a
1.Wh
/
口.Re
S
.
975
,1
7
.2
o
X

5

v
,
o
;・
a
'

61
・ど

S
"
N"
a
f
t
lj・
f
(
l
j叫 V
l
J
"
'
f"
l
l
S
/
"
'

＠

＠

！......
＇
.
＼

.
. .ー

: LV1cl
1X ・L
11~NV:):>Y\l

)
:
"

¥'
" XI
.
PしA TI

:
'
i
.
N
A
C.McC

,
.
l
.
r
1
1
. Rec1
円
l
n
t
l
'
l
1.l
/
.Rr
i
c
S
.
5
7
9
,1
7
o.2
'
¥
.

。・

¥"
aC
1・
6
.
t
;
s
・

S
J
!
H"
a
f
t
，1・fJY'"/Vlf;H ・
叩
.

＠

＠

d
VI
.
.
J
I
.¥X :

~NV ::PJぺ＂：）