B4F47I1

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180
A. B. VAN DEINSE AND G. C. A. JUNGE
richtius robustus 1). It is of interest to note that Gray in his Catalogue
(1866) enumerates Balaena gibbosa as well as Eschrichtius robustus. It
was impossible for him to know at that time that they could be identical,
for the only fact which was known of Dudley's scrag whale was the
external form.
The third phase starts with Cope's description of the California gray
whale. In 1868 (p. 159) Cope mentioned "that he had the opportunity
of examining a portion of the scrag whale of Dudley, Balaena gibbosa
of Erxleben, and ascertained that it represented a genus not previously
known". The genus was called Agaphelus. He continues: "A second
species of the genus was to be found in the "gray whale" of the coasts of
California". This species was named Agaphelus glaucus. In the same year
(pp. 221-227) he gives a more detailed account of these two species. He
stated now that the specimen he called Agaphelus gibbosus was cast
ashore on Long Beach, New Jersey, in the autumn of 1866 (two years
previously), and that the most important portions for its identification
were lost. From the description of the bones, especially the mandibular
ramus, the lumbar and caudal vertebrae, which according to Cope resembled
those of Balaenoptera rostrata more than those of any other whale and
from the description of the colour of the pectoral fins, which were white
at the base and black at the end, it seems that Cope was misled and that
this whale was a Balaenoptera acuto-rostrala. In 1884 (p. 1124) Cope
came also to this conclusion saying: "The Agaphelus gibbosus must be
withdrawn from the list of authentic species. The bones which I referred
to it are probably those of Balaenoptera rostrata. The characters of the
animal in the flesh were given me by persons whom I supposed to be
trustworthy, but who may have been mistaken". Cope who had not seen
the stranded whale himself was informed of the external form by some
people, who stated that the dorsal fin was lacking and that the whale had
no plicae on the gular and thoracic regions. Because the animal must have
been dead already some time before stranding, we cannot attach too much
value to these statements, but it is interesting that Cope immediately
combined Dudley's description of the scrag whale with the Pacific whale,
which he called Agaphelus glaucus and about whose external form he was
informed by Dall.
1) Allen (1916, p. 176) placed Balaenoptera robusta Lilljeborg into the synonymy
of Balaenoptera physalus (L.) saying (l.c., p. 179): "Fossil remains, now considered
indistinguishable from B. physalus, have been found in Sweden, and formed in 1860
the basis of Lilljeborg's Balaenoptera robusta." It will be clear after our discussion
of these bones that this cannot be upheld, the differences between Lilljeborg's
specimen and Balaenoptera physalus are too striking.
THE CALIFORNIA GRAY WIIALE IN THE ATLANTIC
181
In 1884 Cope thought that the specimen he originally described as
Agaphelus gibbosus in reality was a Balaenoptera acuto-rostrata, but before
that time in 1869 (p. 15) he had established already a new genus, Rhachia-
nectes, for the California gray whale especially on account of the different
form of the scapula between his Agaphelus gibbosus (= Balaenoptera
acuto-rostrata) and Agaphelus glaucus. The form of the scapula in the
first was identical with Balaenoptera, in the latter more like that of Balacna.
On the previous page (p. 14) he says in a synopsis that the difference
should be the absence of an acromion in Rhachianectes. It is not clear
how he came to this statement, for the California gray whale possesses a
well developed acromion. When we bear in mind that Cope compared the
bones of a California gray whale and a Balaenoptera acuto-rostrata, it will
be clear that Agaphelus glaucus remains as the only representative of the
genus Agaphelus and that Rhachianectes is merely a synonym. It is remark-
able that Cope overlooked this consequence in 1884 and that the name
Rhachianectes instead of Agaphelus could hold till now.
We believe that the recent finds of the skull-fragments and bones in
the Netherlands, which parts seem not to be different in any essential
point from those of "Rhachianectes glaucus" make it possible to combine
the three phases and to state that Dudley's scrag whale (1725), Balaena
gibbosa Erxleben (1777), is the same as Balaenoptera robusta Lilljeborg
(1861), Eschrichtius robustus (Lilljeborg) (1865), Agaphelus glaucus
Cope (1868) and Rhachianectes glaucus Cope (1869). After this it is clear
that the name of this whale must be changed into Eschrichtius gibbosus
(Erxleben). We must emphasize that with the bones we have now at hand
we cannot give any argument that the Pacific and Atlantic representatives
of Eschrichtius should be specifically different.
The fact that one species occurs in the northern Pacific as well as in
the northern Atlantic is also mentioned for Crustacea, Mollusca, Echino-
dermata and Pisces (Berg, 1934, PP. 391-409 and Schmidt, 1928,
[AD] pp. 1019-1020).
The question whether the populations of the California gray whale in
the Atlantic and Pacific Ocean were perhaps connected along the American
north coast or (and) the European-Siberian coast cannot be answered yet.
Material from these coasts is totally lacking till now. The skull-fragments
[AD] found during the Vega expedition (1878-1880) are found too near the
Bering Strait to be of any importance for this question.

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178
A. B. VAN DEINSE AND G. C. A. JUNGE
Temminckia, 2, 1937
Scrag whale
NOMENCLATURE AND DISCUSSION OF LITERATURE
The skull-fragments and other bones all point to the fact that the whale
now commonly called Rhachianectes glaucus has not always been restricted
to the Pacific, but has occurred in the Atlantic too. This at first sight
astonishing find made necessary an ample study of the literature and we
will show that it is possible now to give an explanation to some facts which
were imcomprehensible till now. First of all we believe that this find is
the clue to understand what Dudley (1725) meant with the mysterious
scrag whale. About the whales described by Dudley, True wrote in his
splendid work on the whalebone whales of the Atlantic (1904, p. 37):
"All of these are recognizable and have been assigned to their proper
places generically, except the scrag whale, which is, and always has been,
a stumbling block to cetology."
Dudley's description of the scrag whale is the following (1725, p. 258):
"The Scrag whale is near a kin to the Fin-back, but, instead of a Fin
upon his Back, the Ridge of the Afterpart of his Back is scragged with
half a Dozen Knobs or Nuckles; he is nearest the Right Whale in Figure
and for Quantity of Oil, his Bone is white, but won't split."
The absence of a dorsal fin and the crenulations on the dorsal ridge
of the peduncle fully agrees with what is known of Rhachianectes glaucus.
Andrews (1914, p. 258) mentions 6-14 crenulations on the peduncle in
this species. Also the remark that the bone (whalebone) is white and
"won't split" points to this species. Andrews says that the colour of the
laminae in this species is yellowish white or yellow and Scammon describes
it as "light brown or nearly white" (Andrews, 1914, p. 255). About the
structure of the plates Andrews remarks that they are very thick and heavy.
We believe, that, now it is known that Rhachianectes also inhabited the
Atlantic, Dudley's diagnosis is understandable without difficulties. The
scrag whale was said to occur near New England 1).
The scrag whale, Knotenfisch (German), Knobbelvisch (Dutch) is found
in nearly all older literature on the whales of the Atlantic, but nearly all
authors derived their knowledge at second hand and compiled Dudley's
description: Klein (1741), Anderson (1746), Brisson (1756), Cranz (1765),
Boddaert (1772), Müller (1773). Linneaus did not mention this whale in
the 10th and 12th edition of his Systema Naturae.
1) According to Allen (1016, p. 116) the fishermen of the New England coast apply
to-day the name scrag whale to small specimens of Eubalaena glacialis (Bonn.).
Judging from Dudley's description, however, it is certain that he did not design
this species.
baleine
"Recent and older finds of
THE CALIFORNIA GRAY WHALE IN THE ATLANTIC
179
Erxleben in 1777 (p. 610) gave it a Latin name for the first time,
grite
Balaena gibbosa, with diagnosis: "Figura Mysticeti, at gibbi in dorso sex,
Alautique N., laminaeque corneae albae. Species subobscura". He quoted most of the
18 ficcle, carlier authors in the synonymy. Much news was not added to the know-
ledge of this whale by Erxleben either and it seems that he too derived
his knowledge from others. Gmelin mentioned in 1789 (p. 225) Balacna
gibbosa. Under this name the scrag whale and the humpback (Megaptera)
are combined under a and b. His references also are of second hand.
It seems of less importance to enumerate all the later authors, who
mentioned Balaena gibbosa in their papers, the most important are Lace-
pède (1804), Desmarest (1822), G. Cuvier (1823), F. Cuvier (1836),
Brandt and Ratzeburg (1829), Gray (1846, 1850). Most of them do not
give much news and many of them combined the scrag whale and the
humpback under Balacna gibbosa. F. Cuvier thought it possible that Dud-
ley's scrag whale was a rorqual, which had been mutilated, Dudley's note
he considered to be insignificant (1836, p. 355). This view was criticized
already by Gray (1846, p. 16; 1850, p. 19), who thought the scrag whale
to be "a Balaena, probably well known formerly". He also criticized the
lumping together of the scrag whale and the humpbacked whale in one
genus as was done by many authors and said that they did not understand
Dudley's description of the belly of the latter, "being reeved', that is,
plaited". He established the genus Megaptera for the humpbacked whales
(1846, p. 16).
Though much was written about the scrag whale till then, hardly any-
thing was known about this species. A second phase in the history of this
whale starts independently from the former in 1860. In this year Lilljeborg
read a paper in Copenhagen about the find of a subfossil whale near
Gräsö in Sweden and he named it Balaenoptera robusta. In 1862 he gave
a detailed survey in Swedish and repeated it in English in 1867 with good
plates of many of the bones. In 1864 Gray (p. 350) referred this whale
to the genus Megaptera and established the subgenus Eschrichtius for it.
In 1865 he received a cervical vertebra from the Babbacombe bay, England,
and Lilljeborg, who compared it with the cervical vertebrae of his
Balaenoptera robusta, considered them identical. Gray (1865, p. 42) raised
Eschrichtius to generic rank now and thought this genus was related with
Megaptera. He mentions as especially characteristic for this genus the
very low processus coronoideus of the lower jaw, the small bodies of the
cervical vertebrae, which are suborbicular-quadrangular and possess a wide
spinal canal. In his paper of 1867 Lilljeborg too called this whale Esch-

[PAGE BREAK]

Statut
Inconnu
BALEINES À BEC 113
Population
Inconnue
Menaces Inconnues
IDENTIFICATION
.coloration foncée et claire
bande blanc crème
bec assez court
⚫ tête relativement plate
• melon petit mais bien visible
dimorphisme sexuel très marqué
⚫petits groupes en formation serrée
⚫léthargique et lent
⚫ comportement aérien réduit
probablement pas
d'échancrure
médiane
nageoire dorsale
basse, triangulaire
(variable)
bord de fuite
rectiligne ou
légèrement falciforme
base large
⚫ corps couvert de
cicatrices, surtout
derrière le bord de la bande
NAGEOIRE CAUDALE
licrée brun-noir ou
chocolat
tronçon caudal
⚫ assez étroit
Nouveau-ne Inconnu
Multe Em 55m
MÅLE
(RECONSTITUTION)
EAUX PROFONDES DU PACIFIQUE ORIENTAL TROPICAL
EN GENERAL AU LARGE
Poids à la naissance
Poids adulte
Inconnu
Inconnu
AIRE DE RÉPARTITION
La carte indique la zone où l'animal a été le
plus souvent signalé, mais les observations
sont si peu nombreuses qu'elles ne reflètent
pas nécessairement la distribution réelle de
l'espèce, qui est l'une des plus fréquemment
signalees, concernant le genre Mesoplodon,
dans les eaux tropicales du grand large du
Pacifique oriental. Elle a été vue à ce jour au
large des côtes du Guatemala, du Salvador,
du Mexique, du Costa Rica, du Nicaragua,
de Panama, de Colombie, de l'Equateur et
du Pérou, essentiellement dans les eaux
profondes, toujours dans des zones assez
chaudes, d'environ 27 'C. On ne sait pas
quel est son statut, mais le nombre des
animaux signalés indique qu'elle n'est pas
particulièrement rare pour une baleine à bec.
Régime alimentaire

[PAGE BREAK]

L
CEI LONATURE
BALEINES
DAUPHINS
ET
MARSOUINS
MARK CARWARDINE
Illustrations de
MARTIN CAMM
Conseiller
DR PETER EVANS
(Sea Watch Foundation, Université d'Oxford, R.U.)
MASON WEINRICH
(Cetacean Research Unit, Massachussets, E.U)
BORDAS
1996

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GOSSE (Philip Henry) The Romance of Natural History
CLondon, Nisbet, 1860)
366
THE GREAT UNKNOWN.
rally from fifty to three hundred fathoms' depth of water
within a boat's length of the shore. How could a carcase
or a skeleton be cast up here, even if it floated?
But, secondly, as to facts. Is it true, that of all the
larger oceanic animals we find the carcases or skeletons
cast up on the shore? Is it true even of the Cetacea,
whose blubber-covered bodies invariably ensure their
floating, and whose bones are so saturated with oil that
they are but little heavier than water?
In September, 1825, a cetacean was stranded on the
French coast which was previously unknown to natural-
ists. It was so fortunate as to fall under the examina-
tion of so eminent an zoologist as De Blainville; and hence
its anatomy was well investigated. It has become cele-
brated as the Toothless Whale of Havre (Aodon Dalei).
Yet no other example of this species is on record; and,
but for this accident, a whale inhabiting the British
Channel would be quite unrecognised.
Of another whale (Diodon Sowerbyi), likewise British,
our entire knowledge rests on a single individual which
was cast on shore on the Elgin coast, and was seen and
described by the naturalist Sowerby.
There is a species of sperm-whale (Physeter tursio)
affirmed to be frequently seen about the Shetland Islands;
a vast creature of sixty feet in length, and readily dis-
tinguishable from all other Cetacea by its lofty dorsal,
and, according to old Sibbald, by other remarkable pe-
culiarities in its anatomy. Yet no specimen of this huge
creature has fallen under modern scientific observation;
GOSSE'S DELPHINORHYNCHUS.
367
and zoologists are not yet agreed among themselves,
whether the High-finned Cachalot is a myth or a reality!
M. Rafinesque Smaltz, a Sicilian naturalist, described a
Cetacean which, he said, he had seen in the Mediterranean,
possessing two dorsals. The character was so abnormal
that his statement was not received; but the eminent
zoologists attached to one of the French exploring expedi-
tions, MM. Quoy and Gaimard,--saw a school of cetacea
around their ship in the South Pacific, having this extra-
ordinary character,-the supernumerary fin being placed
on the back of the head. Here is the evidence of com-
petent naturalists to the existence of a most remarkable
whale, no carcase of which, no skeleton, has ever been
recognised.
The last example I shall adduce is from my own ex-
perience. During my voyage to Jamaica, when in lat.
19° N., and long. from 46° to 48° W., the ship was sur-
rounded for seventeen continuous hours with a troop of
whales, of a species which is certainly undescribed I
had ample opportunity for examination, and found that
it was a Delphinorhynchus, thirty feet in length, black
above and white beneath, with the swimming paws white
on the upper surface, and isolated by the surrounding
black of the upper parts,-a very remarkable character.
This could not have been the Toothless Whale of Havre;
and there is no other with which it can be confounded.
Here, then, is a whale of large size, occurring in greut
numbers in the North Atlantic, which on no other occa-
sion has fallen under scientific observation.

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420
MARINE MAMMAL SCIENCE, VOL. 8, NO. 4, 1992
LITERATURE CITED
BOYD, I. L., AND C. D. Duck. 1991. Mass changes and metabolism in territorial male
Antarctic fur seals (Arctocephalus gazella). Physiological Zoology 64:375-392.
COSTA, D. P., B. J. LE BOEUF, A. C. HUNTLEY AND C. L. ORTIZ. 1986. The energetics
of lactation in the northern elephant seal, Mirounga angustirostris. Journal of Zoology,
London 209:21-33.
JOHNSON, R. R., AND D. J. FARRELL. 1988. The prediction of body composition in
poultry by estimation in vivo of total body water with tritiated water and deuterium
oxide. British Journal of Nutrition 59:109-124.
ØEN, E. O. 1982. A new darting gun for the capture of wild animals. Nordic Veterinary
Medicine 34:39-43.
ØEN, E. O. 1990. Trials of chemical immobilization of minke whales with etorphine
hydrochloride 1989. Reports of the International Whaling Commission (SC/42/
NHMi18).
891.
REILLY, J. J., AND M. A. FEDAK. 1990. Measurement of the body composition of living
grey seals by hydrogen isotope dilution. Journal of Applied Physiology 69:885-
RUMPLER, W. V., M. E. ALLEN, D. E. ULLREY, R. D. EARLE, S. M. SCHMITT AND T. M.
COOLEY. 1987. Body composition of white-tailed deer estimated by deuterium
oxide dilution. Canadian Journal of Zoology 65:204–208.
VILJOEN, J., S. E. COETZEE AND H. H. MEISSNER. 1988. The in vivo prediction of body
composition in Boer goat does by means of tritiated water space technique. South
African Journal of Animal Science 18:63-67.
CHRISTIAN LYDERSEN, University of Oslo, Department of Biology, Division of
General Physiology, P.O. Box 1051 Blindern, N-0316 Oslo 3, Norway; DAVID
GRIFFITHS, Norwegian College of Veterinary Medicine, Department of Anatomy,
[AD] P.O. Box 8146 dep., N-0033 Oslo 1, Norway; and IAN GJERTZ AND ØYSTEIN
WIG, Norwegian Polar Research Institute, P.O. Box 158, N-1330 Oslo Luf-
thavn, Norway. Received October 15, 1991. Accepted January 12, 1992.
MARINE MAMMAL SCIENCE, 8(4):420-425 (October 1992)
1992 by the Society for Marine Mammalogy
FIRST RECORD OF THE PYGMY BEAKED WHALE
MESOPLODON PERUVIANUS IN THE
NORTH PACIFIC
Two specimens of the genus Mesoplodon were found in the Bahía de La Paz,
Baja California Sur, México. The first one was found on 13 January 1990, on
a beach to the side of the road from La Paz City to San Juan de la Costa
(24°24'N, 110°38'W). The carcass was rotten, the length was estimated at 3.4
m, and the skull was collected and deposited in Aurioles' personal collection in
La Paz, B.C.S. (DAG 160). On 19 April 1990, an almost complete skeleton
of a second specimen was collected from a sandy beach on the west coast of the
Isla Espiritu Santo (24°25'N, 110°25'W). The length of this animal was cal-
Strictement réservé à l'usage privó
(Loi n° 57293 du 11.3.57, art. 41,0)

[PAGE BREAK]

421
ial male
392.
ergetics
Coology,
ition in
uterium
terinary
orphine
C/42/
f living
9:885-
T. M.
iterium
of body
South
ion of
DAVID
tomy,
YSTEIN
2.
E
Luf-
Paz,
0, on
Costa
t 3.4
on in
leton
of the
Scal-
NOTES
culated as 3.3 m. The skeleton was deposited in the Museo de Zoología of
the Universidad Autónoma de Baja California Sur, La Paz, B.C.S. (UABCS-
MM-053).
Because of the distal fusion of the premaxilla and maxilla, the degree of
filling of the mesorostral canal, and the fusion of the vertebral epiphysis to the
centra of the vertebrae, it was concluded that both animals were physically
mature.
Based on the length of the animals and their particular skull characteristics
such as tooth size and shape, tooth position, and synvertex configuration (Fig.
1), among others, they were initially thought to be a new species, and later
determined to be Mesoplodon peruvianus, the species described recently by Reyes
et al. (1991).
These records are the first of M. peruvianus for the North Pacific, and, in
fact, the first outside of the Peruvian coasts, where there is information on ten
specimens, including the holotype, which have either stranded or been captured
between 11°12'S and 15°19'S from 1976 to 1989 (Reyes et al. 1991). The
only other mesoplodont species recorded for the Mexican Pacific is M. ginkgodens
from a skull found on the Playa Mal Arrimo near the mouth of the Laguna
Ojo de Liebre, Baja California Sur, in December 1980 (Leatherwood et al.
1982).
Besides M. peruvianus, 6 of the 13 Mesoplodon species are present in the
North Pacific Ocean (Leatherwood et al. 1982, Mead 1989), with the possibility
of one more undescribed species (Pitman et al. 1987, 1988), for the Eastern
Tropical Pacific (ETP) (Table 1).
yr
There was almost no information about the ziphiids from the ETP until the
last 15 when important data were obtained by trained observers during marine
mammal survey cruises for the Southwest Fisheries Center, and by observers on
tuna purse seiners during their fishing activities (Perrin et al. 1985, Pitman et
al. 1988). Information on ziphiids has also been obtained from the long term
Marine Mammal Research Program of the Universidad Nacional Autónoma de
México (UNAM), in the Bahía de Banderas, Nayarit on the mainland coast of
México since 1981; from the Marine Mammal Research Program of the Uni-
versidad Autónoma de Baja California Sur (UABCS), in the Bahía de la Paz,
Península de Baja California since 1987; and from the work on stranded marine
mammals in the Bahía de La Paz during the last 12 yr by one of us (DAG).
As a result of these investigations, there have been 143 positive and 43
probable sightings of mesoplodonts of at least three different species in the ETP
(Pitman et al. 1988, in litt. 8 August 1991): (1) Mesoplodon densirostris (4
positive sightings). (2) Mesoplodon sp. "A" (24 positive sightings). This un-
identified species was described by Pitman et al. (1987). The animals reach a
maximum estimated length of 5.0-5.5 m, larger than the average for Mesoplodon.
The species appears to be sexually dimorphic, with a very distinctive color pattern
for the presumed adult male, with a broad white swath that originates in the
head area and runs postero-ventrally down both sides. The resulting chevron
pattern contrasts sharply with the otherwise black body. (3) Mesoplodon grayi
(2 positive sightings). This species has an extremely long, narrow rostrum. The

[PAGE BREAK]

Jorg
422
MARINE MAMMAL SCIENCE, VOL. 8, NO. 4, 1992
Figure 1. Skull photographs in antero-dorsal view of the two specimens collected.
Left, specimen UABCS-MM-046 collected on 19 April 1990. Right, specimen DAG-
060 collected on 13 January 1990.

[PAGE BREAK]

:0
NOTES
45
32
Table 1. Records of Mesoplodon spp. in the North Pacific.
Species
M. stejnegeri
M. carlhubbsi
Stranded animals Captured animals
Positive
sightings
M. ginkgodens
M. hectori
M. densirostris
M. peruvianus
M. grayi
M. sp.
25
"A"
423
9.9
ut
of
pio
[D
YST
12.
[1]
Pa
0, d
Cost
letofollected.
t 3.
on i
of th
s cal
2 DAG-
Sources: Leatherwood et al. 1982; Pitman et al. 1987, 1988; Aguayo et al. 1988;
Mead et al. 1988; Mead 1989; R. L. Pitman (in litt. 8 August 1991).
lower jaw contrasts markedly with the dark gray upper rostrum and has a
distinctive pale patch just behind the eye. Based on the descriptions above, we
can assume M. peruvianus is different from these three species.
Owing to the small size and not very distinctive pigmentation of M.
peruvianus, we can expect the field identification to be very difficult (particularly
prior to the species description), therefore several of the remaining (113) positive
sightings of Mesoplodon sp. could have been M. peruvianus.
In Bahía de Banderas, México, researchers from the UNAM have had several
encounters with mesoplodonts and, based on size and coloration, they found at
least three different species (Aguayo et al. 1988): (1) Mesoplodon sp. "A"
described before, (2) tentatively M. ginkgodens, and (3) tentatively M. hectori.
One of us (JUR) had the opportunity to observe the latter animals on 13 July
1986. Two individuals came to the surface very close to our two small boats,
and we could aproach them to within 4-6 m for three to four minutes. The
blow was inconspicuous, their length was between 3.0 and 3.5 m (slightly less
than half the length of the boats), their coloration was dark brown on the back
and sides and lighter below, and no teeth or scars were seen. Taking into account
the difficulty of positive identification of these animals at sea we can now,
tentatively, consider them as M. peruvianus. Thus we know of four possible
sightings of this species in the South Pacific (Reyes et al. 1991), and one positive
sighting in the North Pacific off southern Mexico at 10°58'N, 98°15'W of three
animals, two adults and a calf (R. L. Pitman, in litt. 8 August 1991).
Based on the above information we propose the Eastern Tropical Pacific as
the distribution area for M. peruvianus; the paucity of reports on sightings and
strandings in this area can be attributed to the lack of effort. The first reports
appeared after initiation of an active stranding investigation program in this
area. This is in contrast to the coast of California and to a lesser degree to Baja
California coasts, where the extensive survey efforts allow a conclusion of the
absence of the species based on an absence of reports. Thus, we propose a
northern limit to the species range corresponding to the north boundary of the
ETP, close to 25°N, on the west coast of Baja California Sur. If the hypothesis

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424
MARINE MAMMAL SCIENCE, VOL. 8, NO. 4, 1992
of the distribution in the ETP is correct, the mesoplodonts found on the Peruvian
coast should be close to the limit of their southern range (15°S).
This distribution is similar to the one proposed for Mesoplodon sp. "A" (Pitman
et al. 1987, 1988; Mead et al. 1988), and the distribution of the tropical
dolphins (Stenella spp. and Delphinus) in the ETP (Perrin et al. 1985). Based
on the geographical distribution of the M. peruvianus records, and their probable
foraging in mid and deep waters, it seems they have a distribution similar to
the striped and the common dolphins, which occur in equatorial and subtropical
waters with relatively large seasonal changes in the surface temperature, depth
of the thermocline, and upwelling.
ACKNOWLEDGMENTS
We thank the staff of the Marine Mammal Program of the School for Field Studies,
especially Dr. Gary Miller for finding information on the UABCS-MM-053 specimen.
Dr. D. Wartzok, the editor, reviewed the manuscript and corrected the English. Dr. J.
suggestions.
Mead, Dr. R. Brownell and R. Pitman critically reviewed the manuscript and made
LITERATURE CITED
AGUAYO L., A., D. AURIOLES G., J. URBAN R., M. SALINAS Z., O. VIDAL AND L. T.
FINDLEY, 1988. Beaked whales in Mexican waters. Paper SC/40/SM/13 presented
to the Scientific Committee of the IWC Annual Meeting, May 1988.
LEATHER WOOD, S., R. R. REEVES, W. F. PERRIN AND W. E. EVANS. 1982. Whales,
dolphins and porpoises of the eastern North Pacific and adjacent waters: a guide to
their identification. NOAA Technical Report, NMFS Circular 444. 245 pp.
MEAD, J. G., J. E. HEYNING AND R. L. BROWNELL, JR. 1988. Distribution and ex-
ploitation of beaked whales in the Northern Hemisphere. Paper SC/40/SM/21
presented to the Scientific Committee of the IWC Annual Meeting, May 1988.
MEAD, J. G. 1989. Beaked whales of the genus Mesoplodon. Pages 349-430 in S. H.
Academic Press, N.Y.
Ridgway and R. J. Harrison, eds. Handbook of marine mammals, volume 4.
PERRIN, W. F., M. D. SCOTT, G. J. WALKER AND V. L. CASS. 1985. Review of
geographical stocks of tropical dolphins (Stenella spp. and Delphinus delphis) in the
NMFS 28. 28 pp.
eastern North Pacific. U.S. Department of Commerce, NOAA Technical Report
PITMAN, R. L., A. AGUAYO L. AND J. URBAN R. 1987. Observations of an unidentified
3:345-352.
beaked whale (Mesoplodon sp.) in the eastern tropical Pacific. Marine Mammal Science
PITMAN, R. L., D. W. K. Au, M. SCOTT AND J. M. COTTON. 1988. Observations of
beaked whales (Ziphiidae) from the eastern tropical Pacific Ocean. Paper SC/40/
1988.
SM/14 presented to the Scientific Committee of the IWC Annual Meeting, May
REYES, J. C., J. G. MEAD AND K. VAN WEAREBEEK, 1991. A new species of beaked
Science 7:1-24.
whale Mesoplodon peruvianus sp. n. (Cetacea: Zhipiidae) from Peru. Marine Mammal
JORGE URBÁN-RAMÍREZ, Departamento de Biología Marina, Universidad Au-
tónoma de Baja California Sur. Ap. Post. 19-B, La Paz, B.C.S. 23081 México;

[PAGE BREAK]

PERSPECTIVES SCIENTIFIQUES
Un cousin
des dauphins
Le mésoplodon nain est la
treizième espèce du genre.
In 1976, le zoologue James Mead de
la Smithonian Institution, découvre
les restes d'un cétacé inconnu sur un
marché au poisson du Pérou. Un travail
de longue haleine sera nécessaire jusqu'à
ce qu'une nouvelle espèce soit officielle-
ment enregistrée sous le nom de méso-
plodon nain (Mesoplodon peruvianus
Reyes, Mead & van Waerebeek, 1991).
James Mead et ses collègues du Cen-
tre péruvien d'étude des cétacés mettent
près de 15 ans pour collecter un nombre
suffisant de spécimens, pris dans des
filets de pêche ou échoués sur des
plages: l'animal semble rare et les zones
où il vit sont peu fréquentées par les
cétologues. Une dizaine de spécimens
seulement a été récupérée, et il a fallu
attendre 1985 pour en obtenir un
complet. Le Musée océanographique de
La Rochelle présente actuellement une
exposition autour du crâne de l'un de ces
animaux qu'il a acquis pour enrichir ses
collections. Ce spécimen a été capturé en
janvier 1992 au large de la localité de
Pucusana au Pérou.
Les mésoplodons sont des odonto-
cètes, des cétacés à dents, membres de
la famille des ziphiidés. Les odontocètes
les plus connus sont les dauphins (famille
des delphinidés), les cachalots (famille
des physétéridés), et les marsouins (fa-
mille des phocénidés). La première
espèce de mésoplodon fut répertoriée
en 1804. Il s'agit du «mésoplodon de
Sowerby» qui vit principalement en Mer
du Nord. Les autres ont été découvertes
tout au long du XIXe et du XXe siècle. Les
deux dernières l'avaient été en 1958 et
1963. Le mésoplodon nain porte à treize
le nombre d'espèces du genre, une sorte
de record: les autres genres de la famille
des ziphiidés ne comptent qu'une ou
deux espèces chacun.
Les mésoplodons ne possèdent
qu'une paire de dents, implantée sur la
mâchoire inférieure. Leur aileron dorsal
est situé très en arrière, et leur queue ne
porte pas d'encoche centrale. Le méso-
plodon nain est plus petit que les autres
espèces. Le plus grand spécimen est un
mâle de 3,72 mètres de long alors que les
autres mésoplodons ont des tailles
comprises entre 4 et 6 mètres. La posi-
tion de ses dents et d'autres caractères
crâniens le différencient également. Son
dos est de couleur gris brun: la pigmen-
tation s'éclaircit sur les flancs pour deve-
nir presque blanche sur le ventre.
L'enregistrement d'une nouvelle
espèce est plus fréquent pour des petits
animaux comme les rongeurs que pour
des mammifères de grande taille Le peu
d'explorations menées dans le Pacifique
tropical sud où vit le mésoplodon nain
explique sans doute qu'il n'ait pas été
identifié plus tôt. Selon Daniel Robineau,
du Muséum national d'histoire naturelle,
la découverte d'une nouvelle espèce au
sein d'un groupe restreint comme les
cétacés, qui en compte environ 80, mon-
tre que notre connaissance de cette
faune n'est pas exhaustive: «L'image
très populaire des dauphins laisse penser
au public que l'on connaît tous les céta-
cés. Nombre d'espèces actuelles, que
l'on trouve dans différentes régions du
Globe, présentent des formes locales encore
mal connues dont certaines seront sans
doute considérées comme des espèces
distinctes. Par ailleurs la découverte d'es-
pèces entièrement nouvelles n'est pas à
exclure il existe encore de nombreuses
zones océaniques peu explorées. >>
Un mésoplodon marin, capturé accidentellement par des pêcheurs péruviens.
La physique
dans un plan
Élasticité et ruptures d'un
Cristal à deux dimensions.
L
es physiciens aiment à simplifier la
nature pour dépister les phénomènes
nouveaux. Si les cristaux ont habituelle-
ment trois dimensions, l'étude des carac-
téristiques à deux dimensions permet
d'isoler des phénomènes intéressants,
de valider les théories qui s'y rapportent
et d'imaginer ensuite des applications
techniques.
Les cristaux à deux dimensions sont
constitués par des molécules confinées
à la surface de l'eau. Une partie hydro-
phile de ces molécules «amphiphiles>>
réside dans l'eau, et une autre partie
hydrophobe se maintient hors de l'eau.
Nous avons utilisé des molécules d'acide
stéarique modifié dont les deux extrémi-
tés sont inégalement hydrophobes.
Selon la température et la pression (bidi-
mensionnelle), les molécules sont dans
les états gazeux, liquide ou solide.
Dans les phases gazeuse et liquide,
les deux groupes hydrophiles sont dans
l'eau. Dans la phase solide, la molécule
est redressée, et seul le groupe acide est
dans l'eau. Ainsi la solidification corres-
pond à un changement notable de l'aire
du film (d'un facteur presque égal à trois);
en comprimant lentement le film liquide,
on fait croître des cristaux dans la phase
liquide Une propriété optique permet de
suivre la progression de la phase solide :
celle-ci émet une fluorescence jaune
quand elle est éclairée en lumière bleue,
tandis que les phases liquide et gazeuse
n'émettent presque pas.
Lorsque la compression du film est
lente, les monocristaux qui croissent
dans la phase liquide sont des aiguilles
rectangulaires d'une molécule d'épais-
seur, de plus d'un millimètre de longueur
et de quelques centièmes de millimètre
de largeur. Pour mesurer la flexibilité de
ces bandes cristallines lors d'une défor-
mation dans le plan de l'eau, on utilise un
dispositif constitué de deux aiguilles en
fibre de verre fixes et d'une aiguille mobile
et flexible, qui pousse le cristal bidimen-
sionnel en son milieu (voir la figure). On
mesure au microscope le fléchissement
des aiguilles de verre pour connaître la
force appliquée et le fléchissement du
cristal sous l'action de cette force.
Après l'élimination de nombreux arte-
facts expérimentaux dus aux forces capil-
POUR LA SCIENCE N°201 JUILLET 1994
25
25

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MARINE MAMMAL SCIENCE, 3(4):345-352 (October 1987)
©1987 by the Society for Marine Mammalogy
ra novaeangliae
eds. Handbook
Academic Press,
ey, ed. Marine
ch Press, Seat-
tober 2, 1986
arch 23, 1987
OBSERVATIONS OF AN UNIDENTIFIED
BEAKED WHALE (MESOPLODON SP.) IN THE
EASTERN TROPICAL PACIFIC
ROBERT L. PITMAN
Southwest Fisheries Center,
[AD] P.O. Box 271,
La Jolla, California 92038
ANELIO AGUAYO L.
JORGE URBAN R.
Facultad de Ciencias, UNAM,
Apartado Postal 70-572, C.P. 04510,
México, D.F., México
ABSTRACT
Multiple sightings of a distinctive but unidentified species of beaked whale
have been made in the eastern tropical Pacific. The unidentified whale has two
color morphs: a conspicuously marked black and white form (judged to be
larger), and a uniformly gray-brown form. Maximum length estimates have
been 5-5.5 m. Other features include a relatively flat head, with a small, distinct
melon; a moderately long beak; and a low, wide-based, triangular dorsal fin. On
most animals the trailing edge of the dorsal fin is only slightly falcate and often
appears straight. On the black and white morph, a broad white or cream-colored
swathe originates immediately posterior to the dorsal surface of the head and
runs posterio-ventrally on either side of the animal. The prevalence of scarring
on the black and white animals suggests sexual dimorphism and that these
larger, more conspicuously marked animals are adult males, while the smaller,
browner, unscarred animals are females and young. Possibilities for identification
include: 1) a well-marked race of a known Mesoplodon sp., 2) Mesoplodon
(Indopacetus) pacificus or 3) an undescribed species.
Key words: Mesoplodon, unidentified beaked whale, eastern tropical Pacific.
In this paper we report on multiple sightings of a distinctive but as yet
unidentified species of beaked whale from the eastern tropical Pacific (ETP). A
morph of this whale that we assume is the adult male (see below) is conspicuously
description of a
marked and eminently identifiable in the field. Despite this, it does not fit the
any known species of ziphiid. We provide here a field description
of the whale, identify its range and comment on its identity.
345
[A
th
O
b
V

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PITMAN ET AL: UNIDENTIFIED BEAKED WHALE
347
[A
th
I
a
S
Figure 2.
b
Photographs of the unidentified beaked whale from Bahia de Banderas,
Jalisco, Mexico. The animal shown in 2a and 2b, foreground, is presumably an adult
male; notice the scratches behind the white area. Photos by P. Hernández and J. L. López.
The distinctive dorsal fin is low, wide-based and triangular. On most animals
the trailing edge of the fin is only slightly falcate and often appears straight.
Most of the dorsal surface of the black and white form, including the dorsal
fin, is dark and has been variously described as blackish, black/brown, chocolate
brown or dark olive brown. This is similar to the range in color variation noted
for short-finned pilot whales (Globicephala macrorhynchus) in the eastern Pacific.
Pilot whales in the field are generally described as jet black, but in the ETP
they often appear greenish brown. Whether these perceived differences are the
result of lighting effects, as seems likely, or some actual physical differences, is
not known. For the time being, the unidentified whale is best described as black
and white.
m
b

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346
30 N
MARINE MAMMAL SCIENCE, VOL. 3, NO. 4, 1987
Hawallan
Islands
⚫ Subtropical
20°
Seasonally
Tropical
10°
Tropical.
Equatorial
10°
20°S
160°
150°
Subtropical
140°
130°
120°
110°
Mexico
100°
O Sightings
Tentative sightings
Survey vessel noon
positions (500 10x1)
South
America
90°
60°
70°
Figure 1.
Sightings of the unidentified beaked whale in the eastern tropical Pacific
relative to surface water types (after Wyrtki 1967).
SIGHTINGS
Figure 1 shows 24 positive and 8 tentative sightings of the unidentified whale.
All except one were made by trained observers during marine mammal survey
cruises for the Southwest Fisheries Center in the central and eastern tropical
Pacific (see Perrin et al. 1983). Noon positions of survey vessels during these
cruises are also plotted in Figure 1 and indicate the extent of survey coverage.
The remaining sighting (Fig. 2) was photographed during a marine mammal
identification course taught by two of us (AAL and JUR) at Facultad de Ciencias,
Universidad Nacional Autonoma de Mexico (UNAM).
Figure
Jalisco, 1.
male; no
DESCRIPTION
The unidentified whale has two distinct color morphs: a conspicuously marked
black and white form and a relatively nondescript morph that appears to be
uniformly gray-brown (Figs. 2 and 3). When they have been seen together, the
black and white animal has been judged by several observers to be the larger
of the two. Estimates of maximum length have most often been between
approximately 5-5.5 m, making it an average-to-slightly-large-sized Mesoplodon
(see Mead 1984).
The head is relatively flat, as in most mesoplodonts, with a small but distinct
melon. The beak is moderately long, appearing similar in size to that of M.
densirostris. On one occasion an observer reported that the mandible was paler
than the upper part of the beak on one of the black and white animals.
The
the trail
Most
fin, is d
brown
for shor
Pilot w
they of
result d
not knea
and w

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PITMAN ET AL.: UNIDENTIFIED BEAKED WHALE
351
forms a unique biotic province in the eastern Pacific (Reid 1962, Wyrtki 1967,
McGowan 1972, Au and Perryman 1985). The range of the unidentified whale
is also nearly identical to the range of the eastern spinner dolphin (Stenella
longirostris), a pantropical species with a well-marked form endemic to the
eastern tropical Pacific (Perrin et al. 1985). From the evidence above, it seems
likely that a distinct, possibly endemic, species of Mesoplodon also occurs in the
eastern tropical Pacific.
Possibilities for identification of the unidentified whale include the following:
1) It may be a well-marked race of a known species of Mesoplodon.
2) Mesoplodon (Indopacetus) pacificus is currently the rarest extant species of
whale known and has never been identified in the flesh. It is known from
only two skulls: one from Queensland, Australia, and one from Somalia,
northeast Africa. It is possible, however, that these records represent
extralimital strays from a localized population inhabiting a little-explored
area, such as the eastern tropical Pacific.
3) It may be an undescribed species.
The identity of the whale will remain unknown until a stranded specimen is
collected.
ACKNOWLEDGMENTS
We would like to thank Jim Cotton, Gary Friedrichsen and Scott Sinclair for their
capable assistance in the field. David Au, Jim Mead, John Heyning and especially William
Perrin improved the manuscript with their comments. We thank P. Hernández and J
L. López for use of their photographs, Henry Orr for drafting Figure 1, and we gratefully
acknowledge the talents of Lou Silva in preparing Figure 3.
LITERATURE CITED
Au, D. W. K., AND W. L. PERRYMAN 1985. Dolphin habitats in the eastern tropical
Pacific. Fishery Bulletin, U.S. 83:623-643.
BAKER, A. N. 1983. Whales and dolphins of New Zealand; an identification guide.
Victoria University Press, Wellington. 133 pp.
BARBER, R. T., AND F. P. CHAVEZ. 1983. Biological consequences of El Niño. Science
[AD] 222:1203-1210.
GASKIN, D. E. 1982. The ecology of whales and dolphins, Heinemen, London and
Exeter, New Hampshire. 459 pp.
HEYNING, J. E. 1984. Functional morphology involved in intraspecific fighting of the
[AD] beaked whale, Mesoplodon carlhubbst. Canadian Journal of Zoology 62:1645-1654.
LEATHER WOOD, S., R. R. REEVES AND L. FOSTER. 1983. The Sierra Club handbook of
whales and dolphins. Sierra Club Books, San Francisco. 302 pp.
MCGOWAN, J. A. 1972. The nature of oceanic ecosystems Pages 9-28 in C. B. Miller,
ed. The biology of the oceanic Pacific. Proceedings of the 33rd Annual Biology
Colloquium, Oregon State University Press, Corvallis, Oregon.
MEAD, J. G. 1981. First records of Mesoplodon hectori (Ziphiidae) from the northern
hemisphere and a description of the adult male. Journal of Mammalogy 62:430-
432.
MEAD, J. G. 1984. Survey of reproductive data for the beaked whales (Ziphiidae).
Pages 91-96 in W. F. Perrin et al., eds. Cetacean reproduction: estimating parameters

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350
MARINE MAMMAL SCIENCE, VOL 3, NO. 4, 1987
only to sightings with adult males present. For 16 sightings where color morph
information was obtained for all of the animals present, 5 were lone males, 6
were male/female (or possibly male/juvenile) pairs, and 5 involved single males
with 2 or more female/juveniles, including one sighting with an estimated 8
animals.
At sea, this whale behaves similarly to other Mesoplodon species. It is usually
seen rolling over slowly at the surface or traveling at a moderate pace and in
fairly tight groups when more than one animal is present. It generally does
not engage in any sort of "playful" activity, e.g., breaching, tail-slapping, or
spy-hopping, though on one occasion a lone male was seen to breach clear of
the water three times in a row. Like other members of the genus, its behavior
repertoire, at least when it is at the surface, appears to be very limited and
stereotyped. Under normal circumstances there is no visible blow.
IDENTITY OF THE WHALE
In considering the possibilities for identification, two genera of beaked whales,
Berardius and Hyperoodon, can be eliminated because of excessive size (approx-
imately 12 and 7 m, respectively). Ziphius cavirostris is tan or whitish with a
stubby beak and is a species we commonly identify in the eastern tropical Pacific
(Pitman, unpublished observations). Tasmacetus shepherdi is probably too large
also (maximum length at least 7 m, Mead 1984) and its distribution appears
to be circumpolar in southern oceans (Gaskin 1982). The relatively small size
(5-5.5 m), distinct beak, and small but discernable melon lead us to identify
the whale with the genus Mesoplodon.
Currently there are 12 recognized living species of Mesoplodon. From descrip-
tions and photographs of stranded animals, the external pigmentation patterns
of most of these have gradually become known (e.g., Baker 1983, Leatherwood
et al. 1983), and none matches the description of the unidentified whale. (A
possible exception to this, M. (Indopacetus) pacificus, is discussed below.)
Nearly all of the information on Mesoplodon distribution has come from
stranded specimens. Gaskin (1982) stated that the genus "seems to be in a
process of prolific speciation, possibly as a result of population isolation events
which began in the Pliocene and were completed in the Pleistocene." Given the
large number of species in this open ocean group, it is not surprising that the
distributions of individual species appear to be limited to distinct water masses
(Mead 1981, Mead et al. 1982). Of the species known to inhabit tropical or
warm temperate waters, M. europaeus is known only from the Atlantic, M.
gingkodens inhabits the western Pacific (though apparently strays to the eastern
Pacific), and M. densirostris is pantropical. One of us (RLP) has identified M.
densirostris several times in the eastern tropical Pacific but only west of the area
where the unidentified whale occurs. To date, there has not been any Mesoplodon
specimen material collected from within the area that appears to be the normal
range of the unidentified whale (J. Mead, personal communication).
Tropical surface water and the apparent range of the unidentified whale (Fig.
1) both correspond to a distinct and highly productive oceanographic area that
forms a uni
McGowan
is also near
longirostris)
eastern trof
A
likely that
eastern troph
Possibili
1) It m
2) Mesi
wha
only
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3) It r
The id
collected.
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Perrin ima
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MEAD,
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348
MARINE MAMMAL SCIENCE, VOL. 3, NO. 4, 1987
On the black and white morph, a broad white or cream-colored swathe
originates immediately posterior to the dorsal surface of the head and runs
posterio-ventrally on either side of the animal. Anteriorly, the swathe is rather
broad, maybe 60-90 cm, but appears to taper out toward the trailing end. This
swathe has been seen to form a very large and contrasting chevron when the
animal is viewed head on (and probably from above also). When seen at a
distance, the posterior edge of this chevron forms a fairly distinct, contrasting
border against the dark color of the rest of the posterio-dorsal surface of the
whale. On closer inspection however, this border is seen to be broken up by
what appears to be white dappling on a dark background. A possible clue
concerning the ontogeny of the chevron pattern came from a sighting of an
animal, apparently a young adult, that had the contrasting white chevron pattern
described above, but it was formed entirely of individual white spots. The white
spotting may become confluent as the animal matures, eventually forming the
characteristic chevron pattern. The head and beak area of the black and white
animal is darker than the chevron but lighter than the back and usually appears
reddish brown or tan (Fig. 2).
Whenever the black and white animal has been seen at relatively close range,
scratches and scarring have been noted, particularly along the trailing edge of
the chevron (see Fig. 2). No scars have been seen on the lighter brown animals.
In most species of Mesoplodon, only adult males have teeth that erupt from the
surface of the gum (usually one on either side of the lower jaw). These teeth
generally protrude as tusks and, because of intraspecific fighting, males scar each
other up considerably at times. Consequently, adult males are often discernable
from females and young by the presence of extensive scarring (Heyning 1984).
The prevalence of scarring on the black and white animals suggests that the
unidentified beaked whale is sexually dimorphic and that the larger, more
conspicuously marked black and white animals are adult males, while the smaller,
browner, unscarred animals are females and young.
The second, ostensibly female, color form appears uniform gray/brown or
bronze-colored. No conspicuous color pattern, markings or scarring have been
noticed on this form. The dorsal fin of the brown animal is also low, wide-
based and triangular, although it tends to be slightly more falcate than in the
males. The brown animals are relatively nondescript and separating them from
other mesoplodonts in the field may not be possible. In fact, all of the positive
sightings shown in Figure 1 were based on the presence of at least one readily
identifiable adult male of the species.
Mead et al. (1982) described a similar sexual dimorphism with respect to
external color patterns in M. carlhubbsi. In that species, the adult male has a
rostrum and a "distinctive white patch centered on the
"brilliant white'
dorsal prominence of the melon," while the remainder of the body is dark gray
or black. Female and subadult M. carlhubbsi show an incipient version of this
pattern but are more uniformly gray-brown.
Figure 3 is an artist's rendition of both morphs of the unidentified whale
based on photographs (Fig. 2), and descriptions and sketches made during field
observations. The ventral color patterns shown are inferential, as that part of
S
S
O
Figure 3.
2 and illustr
male, while b
juveniles (set
and the ven
y
e
the animal
has not be
lodont hea
species.
No eru
specifically
do, they e
carlhubbs
DISTRIBU
The n
eastern t
Wyrtki
warm w
≥27°C
were as
sighting
anomal
(Barber
BIOLOC
Her
2 or 3 =
A
th

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PITMAN ET AL.: UNIDENTIFIED BEAKED WHALE
349
Figure 3.
An artist's depiction of the unidentified beaked whale drawn from Figure
2 and illustrations and descriptions made in the field. The larger animal is a presumed
male, while the smaller animal (not to scale) shows the pattern of presumed female/
juveniles (see text). We have seen little more of this animal than is visible in Figure 2
and the ventral patterns shown here are inferred. Illustration by Lou Silva.
the animal has never been seen. Likewise, the shape of the beak and line
has not been adequately examined; the illustration shows a generalized mesop-
lodont head and rostrum of about the right proportions for the unidentified
species.
gape
No erupted teeth have been seen on this species either, though we have
specifically looked for them. We suspect if they exist, and they almost certainly
do, they are probably not as large or conspicuous as in M. layardii or even M.
carlhubbsi.
A
[2
S
S
b
DISTRIBUTION
The normal range of the unidentified whale appears to be restricted to the
eastern tropical Pacific (Fig. 1). Nearly all of our sightings were within an area
Wyrtki (1967) characterized as tropical surface water. This is an area of very
warm water and recorded sea surface temperatures for 20 of our sightings were
≥27°C (80°F). The two sightings in Figure 1 that appear south of this range
were associated with southerly intrusions of warm water. The southernmost
sighting was made in February 1983 when a major El Niño event was shifting
anomalously warm water and associated organisms south along the coast of Peru
(Barber and Chavez 1983, Velez et al. 1984).
BIOLOGICAL OBSERVATIONS
Herd size for the 24 positive sightings ranged from 1 to 8, with groups of
2 or 3 being the most common. As noted above however, these results pertain
a

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352
MARINE MAMMAL SCIENCE, VOL. 3, NO. 4, 1987
for stock assessment and management. Reports of the International Whaling Com-
mission, Special Issue 6.
MEAD, J. G., W. A. WALKER AND W. J. HOUCK. 1982. Biological observations on
Mesoplodon carlhubbsi (Cetacea. Ziphiidae). Smithsonian Contributions to Zoology
344. 25 pp.
PERRIN, W. F., M. D. SCOTT, G. J. WALKER, F. M. RALSTON AND D. W. K. Au. 1983.
Distribution of four dolphins (Stenella spp. and Delphinus delphis) in the eastern
tropical Pacific, with an annotated catalogue of data sources. NOAA Technical
Memorandum NMFS-SWFC-38. 65
PP.
PERRIN, W. F., M. D. SCOTT, G. J. WALKER AND V. L. CASS. 1985. Review of
geographical stocks of tropical dolphins (Stenella spp. and Delphinus delphis) in the
eastern tropical Pacific. NOAA Technical Report NMFS 28. 28 pp.
REID, J. L., JR. 1962. On circulation, phosphate-phosphorus content, and zooplankton
volumes in the upper part of the Pacific. Limnology and Oceanography 7:287-306.
VELEZ, J., J. ZEBALLOS, AND M. MENDEZ. 1984. Effects of the 1982-83 El Niño on
the fishes and crustaceans off Peru. Tropical Ocean-Atmosphere Newsletter 28:
10-12.
WYRTKI, K. 1967. Circulation and water masses in the eastern equatorial Pacific Ocean.
International Journal of Oceanology and Limnology 1:117-147.
MARINE MA
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[PAGE BREAK]

MARINE MAMMAL SCIENCE, 3(4):345-352 (October 1987)
1987 by the Society for Marine Mammalogy
OBSERVATIONS OF AN UNIDENTIFIED
BEAKED WHALE (MESOPLODON SP.) IN THE
EASTERN TROPICAL PACIFIC
ROBERT L. PITMAN
Southwest Fisheries Center,
[AD] P.O. Box 271,
La Jolla, California 92038
ANELIO AGUAYO L.
JORGY URN R.
Facultad de Ciencias, UNAM,
Apartado Postal 70 572, CP. 04510,
México, DF, México
ABSTRACT
Multiple sightings of a distinctive but unidentified species of beaked whale
have been made in the eastern tropical Pacific. The unidentified whale has two
color morphs: a conspicuously marked black and white form (judged to be
larger), and a uniformly gray-brown form Maximum length estimates have
been 5-5.5 m. Other features include a relatively flat head, with a small, distinct
melon; a moderately long beak; and a low, wide-based, triangular dorsal fin. On
most animals the trailing edge of the dorsal fin is only slightly falcate and often
appears straight. On the black and white morph, a broad white or cream-colored
swathe originates immediately posterior to the dorsal surface of the head and
runs posterio-ventrally on either side of the animal. The prevalence of scarring
on the black and white animals suggests sexual dimorphism and that these
larger, more conspicuously marked animals are adult males, while the smaller,
browner, unscarred animals are females and young. Possibilities for identification
include: 1) a well-marked race of a known Mesoplodon sp., 2) Mesoplodon
(Indopacetus) pacificus or 3) an undescribed species.
Key words: Mesoplodon, unidentified beaked whale, eastern tropical Pacific.
In this paper we report on multiple sightings of a distinctive but as yet
unidentified species of beaked whale from the eastern tropical Pacific (ETP). A
morph of this whale that we assume is the adult male (see below) is conspicuously
marked and eminently identifiable in the field. Despite this, it does not fit the
description of any known species of ziphiid. We provide here a field description
of the whale, identify its range and comment on its identity.
345

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346
MARINE MAMMAL SCIENCE, VOL. 3, NO. 4, 1987
30° N
Hawallan
Islands
⚫ Subtropical
20°
Seasonally
Tropical
10°
Tropical.
Equatorial
10°
20°S
160°
150°
Subtropical
140°
130°
120°
110°
Mexico
100°
O Sightings
Tentative sightings
Survey vessel noon
positions (soo text)
90°
80°
South
America
70°
Figure 1. Sightings of the unidentified beaked whale in the eastern tropical Pacific
relative to surface water types (after Wyrtki 1967).
PITMAN ET AL.: UNIDENTIFIED BEAKED WHALE
347
a
SIGHTINGS
Figure 1 shows 24 positive and 8 tentative sightings of the unidentified whale.
All except one were made by trained observers during marine mammal survey
cruises for the Southwest Fisheries Center in the central and eastern tropical
Pacific (see Perrin et al. 1983). Noon positions of survey vessels during these
cruises are also plotted in Figure 1 and indicate the extent of survey coverage.
The remaining sighting (Fig. 2) was photographed during a marine mammal
identification course taught by two of us (AAL and JUR) at Facultad de Ciencias,
Universidad Nacional Autonoma de Mexico (UNAM).
Figure 2.
b
Photographs of the unidentified beaked whale from Bahia de Banderas,
Jalisco, Mexico. The animal shown in 2a and 2b, foreground, is presumably an adult
male; notice the scratches behind the white area. Photos by P. Hernández and J. L. López.
DESCRIPTION
The unidentified whale has two distinct color morphs: a conspicuously marked
black and white form and a relatively nondescript morph that appears to be
uniformly gray-brown (Figs. 2 and 3). When they have been seen together, the
black and white animal has been judged by several observers to be the larger
of the two. Estimates of maximum length have most often been between
approximately 5-5.5 m, making it an average-to-slightly-large-sized Mesoplodon
(see Mead 1984).
The head is relatively flat, as in most mesoplodonts, with a small but distinct
melon. The beak is moderately long, appearing similar in size to that of M.
densirostris. On one occasion an observer reported that the mandible was paler
than the upper part of the beak on one of the black and white animals.
The distinctive dorsal fin is low, wide-based and triangular. On most animals
the trailing edge of the fin is only slightly falcate and often appears straight.
Most of the dorsal surface of the black and white form, including the dorsal
fin, is dark and has been variously described as blackish, black/brown, chocolate
brown or dark olive brown. This is similar to the range in color variation noted
for short-finned pilot whales (Globicephala macrorhynchus) in the eastern Pacific.
Pilot whales in the field are generally described as jet black, but in the ETP
they often
appear greenish brown. Whether these perceived differences are the
result of lighting effects, as seems likely, or some actual physical differences, is
not known. For the time being, the unidentified whale is best described as black
and white.

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348
MARINE MAMMAL SCIENCE, VOL. 3, NO. 4, 1987
On the black and white morph, a broad white or cream-colored swathe
originates immediately posterior to the dorsal surface of the head and runs
posterio-ventrally on either side of the animal. Anteriorly, the swathe is rather
broad, maybe 60-90 cm, but appears to taper out toward the trailing end. This
swathe has been seen to form a very large and contrasting chevron when the
animal is viewed head on (and probably from above also). When seen at a
distance, the posterior edge of this chevron forms a fairly distinct, contrasting
border against the dark color of the rest of the posterio-dorsal surface of the
whale. On closer inspection however, this border is seen to be broken up by
what appears to be white dappling on a dark background. A possible clue
of the chevron pattern came from a sighting of an
concerning the ontogeny
animal, apparently a young adult, that had the contrasting white chevron patter
described above, but it was formed entirely of individual white spots. The whi
spotting may become confluent as the animal matures, eventually forming th
characteristic chevron pattern. The head and beak area of the black and whi
animal is darker than the chevron but lighter than the back and usually appears
reddish brown or tan (Fig. 2).
Whenever the black and white animal has been seen at relatively close range,
scratches and scarring have been noted, particularly along the trailing edge of
the chevron (see Fig. 2). No scars have been seen on the lighter brown animals.
In most species of Mesoplodon, only adult males have teeth that erupt from the
surface of the gum (usually one on either side of the lower jaw). These teeth
generally protrude as tusks and, because of intraspecific fighting, males scar each
other up considerably at times. Consequently, adult males are often discernable
from females and young by the presence of extensive scarring (Heyning 1984).
The prevalence of scarring on the black and white animals suggests that the
unidentified beaked whale is sexually dimorphic and that the larger, more
conspicuously marked black and white animals are adult males, while the smaller,
browner, unscarred animals are females and young.
The second, ostensibly female, color form appears uniform gray/brown or
bronze-colored. No conspicuous color pattern, markings or scarring have been
noticed on this form. The dorsal fin of the brown animal is also low, wide-
based and triangular, although it tends to be slightly more falcate than in the
males. The brown animals are relatively nondescript and separating them from
other mesoplodonts in the field may not be possible. In fact, all of the positive
sightings shown in Figure 1 were based on the presence of at least one readily
identifiable adult male of the species.
Mead et al. (1982) described a similar sexual dimorphism with respect to
external color patterns in M. carlhubbsi. In that species, the adult male has a
"brilliant white' rostrum and a "distinctive white patch centered on the
dorsal prominence of the melon," while the remainder of the body is dark gray
or black. Female and subadult M. carlhubbsi show an incipient version of this
pattern but are more uniformly gray-brown.
Figure 3 is an artist's rendition of both morphs of the unidentified whale
based on photographs (Fig. 2), and descriptions and sketches made during field
observations. The ventral color patterns shown are inferential, as that part
of
Figure 3.
PITMAN ET AL.: UNIDENTIFIED BEAKED WHALE
349
An artist's depiction of the unidentified beaked whale drawn from Figure
2 and illustrations and descriptions made in the field. The larger animal is a presumed
male, while the smaller animal (not to scale) shows the pattern of presumed female/
juveniles (see text). We have seen little more of this animal than is visible in Figure 2
and the ventral patterns shown here are inferred. Illustration by Lou Silva.
the animal has never been seen. Likewise, the shape of the beak and gape line
has not been adequately examined; the illustration shows a generalized mesop-
lodont head and rostrum of about the right proportions for the unidentified
species.
No erupted teeth have been seen on this species either, though we have
specifically looked for them. We suspect if they exist, and they almost certainly
do, they are probably not as large or conspicuous as in M. layardii or even M.
carlhubbsi.
DISTRIBUTION
The normal range of the unidentified whale appears to be restricted to the
eastern tropical Pacific (Fig. 1). Nearly all of our sightings were within an area
Wyrtki (1967) characterized as tropical surface water. This is an area of very
warm water and recorded sea surface temperatures for 20 of our sightings were
≥27°C (80°F). The two sightings in Figure 1 that appear south of this range
were associated with southerly intrusions of warm water. The southernmost
sighting was made in February 1983 when a major El Niño event was shifting
anomalously warm water and associated organisms south along the coast of Peru
(Barber and Chavez 1983, Velez et al. 1984).
BIOLOGICAL OBSERVATIONS
Herd size for the 24 positive sightings ranged from 1 to 8, with groups of
2 or 3 being the most common. As noted above however, these results pertain

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350
MARINE MAMMAL SCIENCE, VOL. 3, NO. 4, 1987
only to sightings with adult males present. For 16 sightings where color morph
information was obtained for all of the animals present, 5 were lone males, 6
were male/female (or possibly male/juvenile) pairs, and 5 involved single males
with 2 or more female/juveniles, including one sighting with an estimated 8
animals.
At sea, this whale behaves similarly to other Mesoplodon species. It is usually
seen rolling over slowly at the surface or traveling at a moderate pace and in
fairly tight groups when more than one animal is present. It generally does
not engage in any sort of "playful" activity, e.g., breaching, tail-slapping, or
spy-hopping, though on one occasion a lone male was seen to breach clear of
the water three times in a row. Like other members of the genus, its Lavior
repertoire, at least when it is at the surface, appears to be very limand
stereotyped. Under normal circumstances there is no visible blow.
IDENTITY OF THE WHALE
In considering the possibilities for identification, two genera of beaked whales,
Berardius and Hyperoodon, can be eliminated because of excessive size (approx-
imately 12 and 7 m, respectively). Ziphius cavirostris is tan or whitish with a
stubby beak and is a species we commonly identify in the eastern tropical Pacific
(Pitman, unpublished observations). Tasmacetus shepherdi is probably too large
also (maximum length at least 7 m, Mead 1984) and its distribution appears
to be circumpolar in southern oceans (Gaskin 1982). The relatively small size
(5-5.5 m), distinct beak, and small but discernable melon lead us to identify
the whale with the genus Mesoplodon.
Currently there are 12 recognized living species of Mesoplodon. From descrip-
tions and photographs of stranded animals, the external pigmentation patterns
of most of these have gradually become known (e.g., Baker 1983, Leatherwood
et al. 1983), and none matches the description of the unidentified whale. (A
possible exception to this, M. (Indopacetus) pacificus, is discussed below.)
Nearly all of the information on Mesoplodon distribution has come from
stranded specimens. Gaskin (1982) stated that the genus "seems to be in a
process of prolific speciation, possibly as a result of population isolation events
which began in the Pliocene and were completed in the Pleistocene." Given the
large number of species in this open ocean group, it is not surprising that the
distributions of individual species appear to be limited to distinct water masses
(Mead 1981, Mead et al. 1982). Of the species known to inhabit tropical or
warm temperate waters, M. europaeus is known only from the Atlantic, M.
gingkodens inhabits the western Pacific (though apparently strays to the eastern
Pacific), and M. densirostris is pantropical. One of us (RLP) has identified M.
densirostris several times in the eastern tropical Pacific but only west of the area
where the unidentified whale occurs. To date, there has not been any Mesoplodon
specimen material collected from within the area that appears to be the normal
range of the unidentified whale (J. Mead, personal communication).
Tropical surface water and the apparent range of the unidentified whale (Fig.
1) both correspond to a distinct and highly productive oceanographic area that
PITMAN ET AL. UNIDENTIFIED BEAKED WHALE
351
forms a unique biotic province in the eastern Pacific (Reid 1962, Wyrtki 1967,
McGowan 1972, Au and Perryman 1985). The range of the unidentified whale
is also nearly identical to the range of the eastern spinner dolphin (Stenella
longirostris), a pantropical species with a well-marked form endemic to the
eastern tropical Pacific (Perrin et al. 1985). From the evidence above, it seems
likely that a distinct, possibly endemic, species of Mesoplodon also occurs in the
eastern tropical Pacific.
Possibilities for identification of the unidentified whale include the following:
1) It may
be a well-marked race of a known species of Mesoplodon.
2) Mesoplodon (Indopacetus) pacificus is currently the rarest extant species of
whale known and has never been identified in the flesh. It is known from
only two skulls: one from Queensland, Australia, and one from Somalia,
northeast Africa. It is possible, however, that these records represent
extralimital strays from a localized population inhabiting a little-explored
area, such as the eastern tropical Pacific.
3) It may be an undescribed species.
The identity of the whale will remain unknown until a stranded specimen is
collected.
ACKNOWLEDGMENTS
We would like to thank Jim Cotton, Gary Friedrichsen and Scott Sinclair for their
capable assistance in the field. David Au, Jim Mead, John Heyning and especially William
Perrin improved the manuscript with their comments. We thank P. Hernández and J.
L. López for use of their photographs, Henry Orr for drafting Figure 1, and we gratefully
acknowledge the talents of Lou Silva in preparing Figure 3.
LITERATURE CITED
Au, D. W. K., AND W. L. PERRYMAN 1985. Dolphin habitats in the eastern tropical
Pacific. Fishery Bulletin, U.S. 83:623-643.
BAKER, A. N. 1983. Whales and dolphins of New Zealand; an identification guide.
Victoria University Press, Wellington. 133 pp.
BARBER, R. T., AND F. P. CHAVEZ. 1983. Biological consequences of El Niño. Science
[AD] 222;1203-1210.
GASKIN, D. E. 1982. The ecology of whales and dolphins, Heinemen, London and
Exeter, New Hampshire. 459 pp.
HEYNING, J. E. 1984. Functional morphology involved in intraspecific fighting of the
[AD] beaked whale, Mesoplodon carlhubbsi. Canadian Journal of Zoology 62:1645-1654.
LEATHERWOOD, S., R. R. REEVES AND L. FOSTER. 1983. The Sierra Club handbook of
whales and dolphins. Sierra Club Books, San Francisco. 302 pp.
MCGOWAN, J. A. 1972. The nature of oceanic ecosystems. Pages 9-28 in C. B. Miller,
ed. The biology of the oceanic Pacific. Proceedings of the 33rd Annual Biology
Colloquium, Oregon State University Press, Corvallis, Oregon.
MEAD, J. G. 1981. First records of Mesoplodon hectori (Ziphiidae) from the northern
hemisphere and a description of the adult male. Journal of Mammalogy 62:430-
432.
MEAD, J. G. 1984. Survey of reproductive data for the beaked whales (Ziphiidae).
Pages 91-96 in W. F. Perrin et al., eds. Cetacean reproduction: estimating parameters

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352
MARINE MAMMAL SCIENCE, VOL. 3, NO. 4, 1987
for stock assessment and management. Reports of the International Whaling Com-
mission, Special Issue 6.
MEAD, J. G., W. A. WALKER AND W. J. HOUCK. 1982. Biological bservations on
Mesoplodon carlhubbsi (Cetacea: Ziphiidae). Smithsonian Contributors to Zoology
344. 25 PP-
Au. 1983.
PERRIN, W. F., M. D. SCOTT, G. J. WALKER, F. M. RALSTON AND D.
Distribution of four dolphins (Stenella spp. and Delphinus delp in the eastern
tropical Pacific, with an annotated catalogue of data sources. NA Technical
Memorandum NMFS-SWFC-38. 65 pp.
PERRIN, W. F., M. D. SCOTT, G. J. WALKER AND V. L. CASS. 1985 Review of
geographical stocks of tropical dolphins (Stenella spp. and Delphinus delphis) in the
eastern tropical Pacific. NOAA Technical Report NMFS 28. 28 pp.
REID, J. L., JR. 1962. On circulation, phosphate-phosphorus content, and zooplankton
volumes in the upper part of the Pacific. Limnology and Oceanography 7:287–306.
VELEZ, J., J. ZEBALLOS, AND M. MENDEZ. 1984. Effects of the 1982-83 El Niño on
the fishes and crustaceans off Peru. Tropical Ocean-Atmosphere Newsletter 28:
10-12.
WYRTKI, K. 1967. Circulation and water masses in the eastern equatorial Pacific Ocean.
International Journal of Oceanology and Limnology 1.117-147.
Received: March 7, 1986
Accepted: May 11, 1987

[PAGE BREAK]

[17 decembre 1892, détart de
Bransfield, ocean antarctique, su
le Balaena
FROM EDINBURGH TO THE ANTARCTIC
209
Just after killing the seal there was a shout amongst
the men forward, 'A Uni! A Uni!'-the whalers' term
for a Narwhale. Several men said they saw their
horns.
de unicorn.
évidemment
The crow's-nest
cask, about five
was sent aloft to-day.
It is a
feet deep, paint-
ed white, with
iron clamps that
clasp on to the
main-topgallant
mast. In the
bottom there is
a trap-door. To
get into the nest
you climb up a
Jacob's ladder-
wooden ratlins
rigged on two
backstays that run from the top-gallant mast-head to the
cross-trees; these run through the bottom of the tub. You
climb up these and shove the trap open with your head,
and when you are right into the tub you let the trap shut
below you, and stand on it, and enjoy the extensive view.
If you prefer it, you can sit on a shelf-seat fixed in the back
of the tub-a sheltered, quiet place, far removed from the
troubles of the little world below: round the top of the
tub there is a small iron balustrade, on which a screen
runs, so as to shelter the watcher from the wind.
The boats were all lowered from the skids to the
MUSÉE OCEANOGRAPHIQUE
DE MONACO

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MARINE MAMMAL SCIENCE, 18(3):577-608 (July 2002)
©2002 by the Society for Marine Mammalogy
A NEW SPECIES OF BEAKED WHALE
MESOPLODON PERRINI SP. N.
(CETACEA: ZIPHIIDAE) DISCOVERED
THROUGH PHYLOGENETIC ANALYSES OF
MITOCHONDRIAL DNA SEQUENCES
MEREL L. DALEBOUT
School of Biological Sciences, University of Auckland,
Private Bag 92019, Auckland 1000, New Zealand
E-mail: m.dalebout@auckland.ac.nz
JAMES G. MEAD
National Museum of Natural History, Mailstop NHB 108,
Smithsonian Institution, Washington, DC 20560, U.S.A.
C. SCOTT BAKER'
School of Biological Sciences, University of Auckland,
Private Bag 92019, Auckland 1000, New Zealand
ALAN N. BAKER
Science & Research Unit, Department of Conservation,
P. O. Box 10-420, Wellington, New Zealand
ANTON L. VAN HELDEN
Museum of New Zealand Te Papa Tongarewa,
P. O. Box 467, Wellington, New Zealand
ABSTRACT
Mesoplodon perrini, a new species of beaked whale is described on the basis
of five animals stranded on the coast of California (between 32°55'N,
117°15'W and 36°37'N, 121°55'W) from May 1975 to September 1997.
Four of these animals were initially identified as Hector's beaked whales M.
bectori based on cranial morphology (Mead 1981). A fifth specimen was ini-
tially identified as a neonate Cuvier's beaked whale Ziphius cavirostris based
on external features. These specimens were first recognized as representatives
of an undescribed species through phylogenetic analysis of mitochondrial (mt)
DNA control region and cytochrome b sequence data. Although similar mor-
phologically, the genetic data do not support a close evolutionary relationship
between M. perrini and M. hectori. Instead, these data suggest a possible sister-
'Corresponding author; e-mail: cs.baker@auckland.ac.nz.
577

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578
MARINE MAMMAL SCIENCE, VOL. 18, NO. 3, 2002
species relationship with the lesser beaked whale M. peruvianus. Sightings of
two small beaked whales off California in the 1970s which were tentatively
identified as M. hectori are also likely to be M. perrini. We suggest that M.
hectori is confined to the Southern Hemisphere, while M. perrini is known to
date only from the North Pacific.
Key words: molecular genetics and systematics, morphology, external appear-
ance, natural history, distribution, Mesoplodon perrini, Perrin's beaked whale.
Beaked whales (Cetacea: Ziphiidae) are the least known of all cetacean fam-
ilies. In terms of species diversity, they are second only to oceanic dolphins
(Family Delphinidae), with 20 species currently recognized. Beaked whales are
rarely observed at sea due to their preference for deep ocean waters and elusive
habits. Most species are known from only a small number of stranded speci-
mens, and several have never been seen alive. Of the twelve cetacean species
described in the last 100 years, eight have been ziphiids, primarily of the
genus Mesoplodon. In the closing decade of the 20th century, two new beaked
whales were discovered; the lesser beaked whale M. peruvianus (Reyes et al.
1991), and Bahamonde's beaked whale M. bahamondi (Reyes et al. 1995),
although the latter is now recognized as synonymous with M. traversii (Gray,
1874) (van Helden et al. 2002).
Here we document the occurrence and characteristics of a previously un-
described species of Mesoplodon beaked whale in the North Pacific Ocean. In
the mid to late 1970s, four beaked whales stranded within 85 km of each
other along the southern coast of California (Table 1). These animals were
identified as Hector's beaked whales M. hectori, the first and only records of
this species from the Northern Hemisphere (Mead 1981). In 1997 a database
of mitochondrial (mt) DNA control region reference sequences was compiled
to assist in beaked whale species identification (Dalebout et al. 1998, Henshaw
et al. 1997). All specimens in this reference database were validated through
examination by experts in cetacean morphology and the collection of diag-
nostic skeletal material or photographic records (Dizon et al. 2000). Among
the specimens incorporated in these analyses was one of the "M. hectori" from
California (USNM504259; Henshaw et al. 1997). A phylogenetic review of
the database, which at this time consisted of reference sequences from 16 of
the 20 described species, including Southern Hemisphere M. hectori, suggested
that the California specimen was not of this species nor any other species in
the database (Dalebout et al. 1998).
To further investigate this anomaly, DNA was extracted from the remaining
three specimens from California described by Mead (1981). Phylogenetic anal-
yses of mtDNA control region and cytochrome b sequences from these spec-
imens, using a complete reference database which now includes all 20 de-
scribed beaked whale species, confirmed that all four were of the same species,
yet did not represent M. hectori nor any other known ziphiid species (Dalebout
2002). Instead, these results suggested that the California specimens repre-
sented an undescribed species of beaked whale. Here we present a formal
description of this new species, and include a description of a fifth specimen,

[PAGE BREAK]

Table 1. Specimens of Mesoplodon perrini from California, in order by stranding date. Total length in cm.
DALEBOUT ET AL: NEW SPECIES OF BEAKED WHALE
579
Museum number Field number
Date found
USNM504259
USNM504260
USNM504853
LAM088901
JRH 052
TMMC-C75
1975 May 22
1975 May 28
1978 Sept 9
1979 Dec 27
Locality
Camp Pendleton
Coordinates
Camp Pendleton
Carlsbad
Total
length
33°15'N, 117°26'W 210
33°16'N, 117°26'W 443
33°07'N, 117°20'W 390
Sex
M
F
M
1997 Sept 18
Torrey Pines State Reserve
Monterey
32°55'N, 117°15'W
36°37'N, 121°55'W
245
224
M
M

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580
MARINE MAMMAL SCIENCE, VOL. 18, NO. 3, 2002
which stranded in Monterey, California, in 1997, and was initially identified
as a neonate Cuvier's beaked whale Ziphius cavirostris from external morphol-
ogy.
METHODS
Material Examined
Five specimens of the undescribed species from California were examined
(Table 1). Phylogenetic analyses of mtDNA control region and cytochrome b
ziphiid species.
sequences were used to compare these specimens to all 20 previously described
Museums and institutions holding specimens of the new species are as fol-
lows: Los Angeles County Museum of Natural History, California (LAM) and
the National Museum of Natural History, Smithsonian Institution, Washing-
ton, DC (USNM). Tissue samples are held by: the School of Biological Sci-
ences, University of Auckland, New Zealand (AUNZ); Southwest Fisheries
Science Centre, La Jolla, CA (SWFSC); and, The Marine Mammal Centre,
Sausalito, CA (TMMC).
Morphological comparisons were made to 12 specimens of M. hectori held
(NMNZ), 11 =
by the following museums: Museum of New Zealand Te Papa Tongarewa
7; The Australian Museum, Sydney (AMS), " 1; Tasmanian
Museum and Art Gallery (TAM), n 1; the South Australian Museum (SAM),
n = 1; and, Museo Acatushún de Aves y Mamíferos Australes (MAAMA),
Tierra del Fuego, n = 2. Further information on specimens of M. hectori ex-
amined can be obtained from JGM or AVH.
DNA Extraction and Sequencing
DNA was extracted from the teeth and cartilage on the museum-held ma-
terial of the four specimens (USNM504259, USNM504260, USNM504853,
LAM088901) described by Mead (1981), using the silica-based method of
Höss and Pääbo (1993), as modified by Matisoo-Smith et al. (1997), and
techniques described in Pichler et al. (2001). Museum material from
USNM504259 was included to confirm that the original soft tissue sample
(SWFSC-24976) used by Henshaw et al. (1997) had indeed been derived from
this specimen. (Soft tissue samples were not available for the other three spec-
imens). Total genomic DNA was extracted from soft tissue samples obtained
from the Monterey specimen (TMMC-C75) using standard methods (Sam-
brook et al. 1989), as modified by Baker et al. (1993). Samples were stored in
salt-saturated, dimethylsulphoxide (DMSO) solution prior to analysis.
Segments of the 5' end of the mtDNA control region and 5' end of the
cytochrome b gene were amplified and sequenced from all five specimens, and
aligned to the sequences already in the beaked whale reference database, as
described in Dalebout (2002).

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DALEBOUT ET AL: NEW SPECIES OF BEAKED WHALE
Phylogenetic Analyses
581
Phylogenetic relationships among the California specimens and the 20
known beaked whale species were reconstructed from sequence data from both
mtDNA loci using maximum likelihood (ML) methods (consensus length of
alignments, control region, 437 bp; cytochrome b, 384 bp; Dalebout 2002).
Baird's beaked whale, Berardius bairdii, which likely represents the most basal
extant species in this family (e.g., Dalebout et al. 1998) was used as an out-
group.
To further investigate sister-species relationships among a subset of Meso-
plodon species, including the California specimens, the control region and cy-
tochrome b sequences were combined to increase phylogenetic signal (consen-
sus length of alignment, 821 bp). A partition-homogeneity test (Farris et al.
1995) indicated that these loci were congruent (branch and bound search,
1,000 replicates). Although the individuals representing each species differed
between the two loci in some cases, this was not considered a problem as
intraspecific variation was generally much lower than interspecific variation
(see discussion below). Starting parameters for ML reconstruction were esti-
mated from an initial neighbour-joining tree built using general-time-revers-
ible (GTR) distances. For ML analyses, the heuristic search option, with ran-
dom sequence addition (100 replicates), and sub-tree pruning-regrafting
branch swapping, was used. The statistical consistency of groupings was eval-
uated by 200 ML bootstrap resamplings of the data. Bremer support was
calculated using Tree Rot v.2a (Sorensen 1999), based on one of the three most
parsimonious trees (= ML tree) obtained through an exhaustive parsimony
search. All phylogenetic analyses were conducted using the program PAUP*
4.0 beta 6 (Swofford 1999).
RESULTS
DNA Sequence Data
For the mtDNA control region, fragments ranging in length from 245 bp
to 434 bp were sequenced successfully from the five California specimens. For
the mtDNA cytochrome b, fragments ranging in length from 276 bp to 384
bp were sequenced successfully. For those specimens represented by hard tissue
(i.e., tooth or cartilage; USNM504260, USNM504853, LAM088901), only
shorter fragments were obtained, as expected from DNA extractions from such
material (Höss and Pääbo 1993). Comparison of the mtDNA control region
sequence published by Henshaw et al. (1997) (Genbank Accession No.
U70466) with that obtained from museum-held material from USNM504259
confirmed that they were identical, and as such likely derived from the same
specimen. All previously unpublished sequences have been deposited in Gen-
bank (Accession No.'s: AF441254-AF441263).

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MARINE MAMMAL SCIENCE, VOL. 18, NO. 3, 2002
Phylogenetic Analyses
Phylogenetic analyses of sequence data from the mtDNA control region
(Fig. 1) and cytochrome b (not shown) confirmed that all five California spec-
imens were of the same species, and distinct from all other 20 known species
of beaked whale. Although higher-level relationships were not well resolved
by these rapidly evolving mtDNA loci (i.e., bootstrap scores for most internal
nodes, <50%), all species-specific groupings were supported by high bootstrap
scores (>80%). MtDNA control region sequences representing the southern
bottlenose whale Hyperoodon planifrons were the exception to this trend.
To further investigate the genetic distinctiveness of the California animals,
and exclude the possibility that they represent only a highly divergent, geo-
graphic subdivision of M. hector, we compared the combined control region
and cytochrome b sequences (821 bp) to a subset of potential sister-taxa (Fig.
1, gray box). While a monophyletic grouping of the California animals and
M. hectori would argue for a single-species classification despite a deep diver-
gence (Lento et al. 1997, Wayne et al. 1990), the results indicated instead that
the California specimens were more closely related to at least four other species
of Mesoplodon, than to the morphologically similar M. hectori (bootstrap score,
83%; Bremer support = 7; Fig. 2). No synapomorphies were found to unite
the California specimens and M. hectori exclusive of other beaked whales. These
results allowed us to reject the hypothesis of a deep intraspecific divergence,
and argue instead that these specimens represent a previously unrecognized
species of beaked whale.
Although higher-level relationships among this subset of Mesoplodon species
were not fully resolved (Fig. 2), there was some support for a sister-species
relationship between the California specimens and the lesser beaked whale M.
peruvianus (bootstrap score, 69%). There was also low-level support for a clade
consisting of these two species, plus Gray's beaked whale M. grayi, (bootstrap
score, 55%). Phylogenetic reconstructions based on nuclear sequences support
a similar pattern of relationships among these species (Dalebout 2002),
Intra- and Interspecific Genetic Divergence
Over the 245 bp fragment of the mtDNA control region covered by se-
quence data from all five specimens, all shared the same haplotype (Fig. 3).
Analysis of the 280 bp fragment of the cytochrome b covered by all five
specimens revealed two variable sites (one synonymous third position trans-
version and one non-synonymous first position transition) defining three
unique haplotypes (Fig. 4). The adult female (USNM504260), and two of the
three calves (USNM504259 and TMMC-C75) share the same haplotype at this
locus, while the adult male (USNM504853) and the remaining calf
(LAM088901) were both unique. The adult female and the calf,
USNM504259, both stranded at Camp Pendleton in the same week of May
1975 (Table 1).
Comparisons of intra- and interspecific pairwise sequence divergence for all

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DALEBOUT ET AL: NEW SPECIES OF BEAKED WHALE
583
20 beaked whale species confirmed that the California specimens follow a
similar pattern to other ziphiids (Fig. 5a, b). Over the 437 bp control region
alignment, intraspecific variation (using two representatives per species) was
found to be generally less than 2%, while interspecific variation was generally
greater than 4%. Similar trends were found in a previous analysis, which
compared intra- and interspecific genetic divergence among nine described
beaked whale species (Dalebout et al. 1998). The new species differs from all
other beaked whales by an average of 8.55% over this fragment. Over the 384
bp mtDNA cytochrome b alignment, intraspecific variation was found to be
generally less than 1.5%, while interspecific variation was found to be gen-
erally greater than 3% for 20 described beaked whale species. The new species
differs from all other beaked whale species by an average of 15.24% over this
fragment. See Dalebout (2002) for discussion regarding comparative levels of
divergence at the mtDNA cytochrome b versus the control region among the
Ziphiidae.
DESCRIPTION
Order Cetacea Brisson, 1762
Family Ziphiidae Gray, 1865
Mesoplodon perrini sp. n.
HOLOTYPE
Adult male (USNM504853); skull, mandible, and postcranial skeleton, at
the National Museum of Natural History, Smithsonian Institution, Washing-
ton, DC. This specimen was found on 9 September 1978, by G. Carsten, and
collected two days later by J.G.M.
TYPE LOCALITY
Carlsbad, California (33°07'N, 117°20'W), United States of America.
PARATYPES
Male calf (USNM504259); fragmented cranium and postcranial skeleton, at
the Smithsonian National Museum of Natural History, Washington, DC, col-
lected by W. F. Perrin.
Adult female (USNM504260); skull, mandible, and postcranial skeleton, at
the Smithsonian National Museum of Natural History, Washington, DC, col-
lected by W. F. Perrin.
Male calf (LAM088901); skull, mandible, at the Los Angeles County Mu-
seum of Natural History, collected by J. R. Henderson (JRH 052).
Male calf (TMMC-C75); skull, mandible, and postcranial skeleton, at the
Los Angeles County Museum of Natural History, collected by M. Haulena.

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584
MARINE MAMMAL SCIENCE, VOL. 18, NO 3, 2002
85
100
100
H
95
Sowerby's beaked whale
Mesoplodon bidens
True's beaked whale
Mesoplodon mirus
100 spade-toothed whale
Mesoplodon traversii (=M. bahamondi)
ginkgo-toothed beaked whale
Mesoplodon ginkgodens
Mesoplodon carlhubbsi
densebeaked whale
93
95
Hubbs' beaked whale
97
94
99
Gray's beaked whale
99
Mesoplodon densirostris
Stejneger's beaked whale
Mesoplodon stejnegeri
Mesoplodon grayi
lesser beaked whale
Mesoplodon peruvianus
100 Perrin's beaked whale
Mesoplodon perrini sp. n.
100
straptoothed whale
58
99
Mesoplodon layardii
Longman's beaked whale
Indopacetus pacificus
Andrews' beaked whale
Mesoplodon bowdoini
84 Hector's beaked whale
83
100
98
Mesoplodon hectori
Gervais' beaked whale
Mesoplodon europaeus
93 northern bottlenose whale
Hyperoodon ampullatus
southern bottlenose whale
Hyperoodon planifrons
Shepherd's beaked whale
Tasmacetus shepherdi
Cuvier's beaked whale
96 Arnoux's beaked whale
Berardius arnuxii
Baird's beaked whale
Berardius bairdii
Ziphius cavirostris
nucleotide divergence per lineage (GTR correction)
12%
8%
4%
0%
Treelength=357
Consistency index (CI)=0.401
Retention index (RI)=0.712
Figure 1. Phylogenetic relationships among the 20 described species of beaked
whales (Ziphiidae) from maximum-likelihood analyses, based on 437 bp of mitochon-
drial DNA control region. Numbers above internal nodes indicate bootstrap values
50%. All described species represented by two reference specimens where possible.
Arrows highlight respective positions of Mesoplodon perrini sp. n. and M. hectori, the
species as which several specimens of this new species were initially described. Gray
box indicates subset of taxa, with addition of M. hectori, used in further analyses. Note

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DALEBOUT ET AL.: NEW SPECIES OF BEAKED WHALE
585
ETYMOLOGY
The specific name, perrini, was chosen as a tribute to the American cetacean
biologist, Dr. W. F. Perrin, for his role in the collection of two of the known
specimens of this species, and his ongoing contribution to marine mammal
science and conservation. We propose this species be known by the common
name, Perrin's beaked whale.
DIAGNOSIS
Molecular Characters
M. perrini can be differentiated from all other species of Mesoplodon beaked
whales based on molecular genetic characters, as demonstrated by phylogenetic
analyses of mtDNA control region and cytochrome b sequences (Fig. 1-5).
Over the 434 bp control region segment, M. perrini is distinguished from M.
bectori by 26 diagnostic sites (5.99% pairwise sequence divergence), including
two insertion-deletions (indels), and from M. peruvianus, its likely sister-spe-
cies, by 16 diagnostic sites (3.46%), including one indel (Fig. 3). Over the
384 bp cytochrome b segment, M. perrini is distinguished from M. hectori by
37 diagnostic sites (9.64%), including four first position and three second
position substitutions, and from M. peruvianus by 48 diagnostic sites (12.50%),
including six first position and four second position substitutions (Fig. 4). In
comparisons including all ziphiid species, M. perrini is distinguished by one
diagnostic site (sensu Davis and Nixon, 1992) at the control region (position
111 = A; Fig. 3), and one diagnostic site at the cytochrome b (position 182
[2nd]T; Fig. 4), given a mean of two diagnostic sites per species for both
fragments. Note that high levels of homoplasy were observed at these mtDNA
loci due to the rapid rate of accumulation of mutations and the large number
of species to be differentiated (Sanderson and Donoghue 1989).
Morphological Characters
The following characters of the mandibles, teeth, and skull are, when com-
bined, diagnostic for M. perrini:
(1) Short mandibular symphysis (19%-23% mandible length).
(2) Convex profile to anterior part of mandible over the length of the sym-
physis.
that M. bahamondi (Reyes et al. 1995) was recently recognized as synonymous with M.
traversii, a species described by J. E. Gray in 1874 (van Helden et al. 2002). The
aligned sequence files used in these analyses are available electronically from http://
www.sbs.auckland.ac.nz/research-groups/ecology_and_evolution/molecular_ecol_evo l_lab
or http://www.dna-surveillance.auckland.ac.nz.

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100/40
BbaSW4965/X92541
BbaBC9220
MdeNZ01/NZ02
100/18
MdeNHMUK/SW4010
Mesoplodon densirostris
densebeaked whale
100/12
Mgr05/29
MgrI1/H04
Mesoplodon grayi
Gray's beaked whale
55/0
100/16 MpeJCR1926
MpeLAM95654/U13141
USNM504259
83/7 69/0
USNM504260
TMMC-C75
96/12
USNM504853
LAM088901
Mesoplodon peruvianus
lesser beaked whale
Mesoplodon perrini sp. n.
Perrin's beaked whale
MstSW10402/SW4962
100/15
MstSW9491/SW6481
Mesoplodon stejnegeri
Stejneger's beaked whale
MheNZ02/TAS
98/16 MheNMNZ2173/SAM16387
Mesoplodon hectori
Hector's beaked whale
Berardius bairdii
Baird's beaked whale
nucleotide divergence per lineage (GTR correction)
14%
Figure 2.
7%
0%
Treelength=289
Consistency index (CI)=0.653
Retention index (RI)=0.789
Phylogenetic relationships among Mesoplodon perrini, M. hectori, and sub-
set of related Mesoplodon beaked whales, based on maximum likelihood (ML) analyses,
using combined mitochondrial DNA control region and cytochrome b sequences (821
bp). Numbers adjacent to internal nodes are ML bootstrap values ≥50%/Bremer sup-
port indices. The figures adjacent to the branch termini show the diagnostic size, shape,
and position of the teeth in the lower jaw of the adult male of each species. The circle
indicates the position of the diminutive tooth in M. peruvianus. The arrows draw
attention to the morphological similarity between M. perrini and M. hectori.

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DALEBOUT ET AL. NEW SPECIES OF BEAKED WHALE
587
(3) Sub-terminal narrow teeth, up to 64 mm long, 47 mm wide and 12 mm
broad, with smooth anterior margins and a 60°-70° terminal angle.
(4) Narrow triangular subvertex.
(5) Narrow, reverse V-shaped space between right and left nasals.
(6) Narrow premaxillaries adjacent to the antorbital notches.
(7) Small, but distinct basirostral groove.
(8) Antorbital notches and prominences formed by the maxilla.
(9) Margins of the posteromedial portion of the maxillaries angled sharply
laterally.
Osteology and Dentition
MORPHOLOGICAL DESCRIPTION
Cranial and mandibular measurements are shown in Table 2, 3, respectively.
The premaxillary crest is relatively narrow and conservative in shape (Fig. 6),
similar to that of M. hectori and M. peruvianus. The cranium of the holotype
is not greatly inflated, adding to the narrow triangular appearance of the
synvertex, which is constricted laterally at the confluence of the maxillaries
with the supraocciptals. The margins of the posteromedial portion of the max-
illaries (posterior to the synvertex) are angled sharply laterally, more so on the
right side, and the sides of the cranium are steep. There are moderately formed
maxillary crests above the orbits. The rostrum is relatively short (82%-93%
zygomatic width [ZW]) for a Mesoplodon, and the mesorostral canal is fully
ossified in the adult male. The mandibles have a short symphysis, the ventral
profile of which is convex. The remaining ventral profile of the mandible is
concave and then convex again near the posterior end. As in other species of
this
genus, the teeth are reduced to a single, laterally compressed pair in the
mandible. The teeth are set close to the apex of the jaw, and are thus com-
pletely anterior to the posterior end of the symphysis. In the adult male ho-
lotype specimen, the anterior edge of the teeth is 23 mm from the tip of the
lower jaw in life, with 33 mm of the tooth exposed above the gumline. The
shape of the exposed portion of the tooth is a rough isosceles triangle, but
with a smoothly convex anterior margin. The teeth have a sharp terminal angle
of between 60° and 70°, and splay outwards from the perpendicular by an
angle of approximately 15° (Fig. 7, 8e). The teeth of the adult female
(USNM504260) are sharply triangular, and are not erupted.
Vertebral counts, based on the four type specimens are as follows: cervicals
7 (1 and 2 fused), thoracic 9-10, lumbars 11-13, total 46-50. The phalangeal
formula (based on two specimens) is I-2, II-6, III-7, IV-5, V-3 or 4.
COMPARISONS WITH MESOPLODON HECTORI
From the original misidentification of four of the five specimens of M. perrini
known to date (Mead 1981, Mead and Baker 1987), it is clear that this species
resembles M. hectori (Gray, 1871) morphologically. Further examination of the

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10
20
30
40
50
60
TMMC-C75
USNM504259
USNM504260 ?????????? ?????????? ?????????? ?????????? ?????????? ??.
USNM504853 ?????????? ?????????? ?????????? ?????????? ?????????? ??.
LAM088901 ?????????? ?????????? ?????????? ?????????? ?????????? ??.
MheNMNZ2173
GAAAAAGTCT TGTTATAGAA TCACCATAAC CTTACAGTAC TACGTCAGTA TTGAAAAAGA
.CT. .A..G.....
MpeLAM95654
70
80
..C... .C.
..T.
A..
90
100
110
120
TMMC-C75 AATACCCTAC AGTACATTTA CTGTATTAAT CATACAGACA CATTCA-CTG AGCGCTAATA
USNM504259
USNM504260
USNM504853
LAM088901
MheNMNZ2173
MpeLAM95 654
T.T.
TMMC-C75
USNM504259
USNM504260
USNM504853
LAM088901
MheNMN22173
MpeLAM95 654
TMMC-C75
USNM504259
USNM504260
USNM504853
LAM088901
MheNMNZ2173
MpeLAM95 654
130
140
150
.A.
..CC..C..A GA.A.
..CC..C..A
160
170
180
TATAGCGTCT CTCTAAAAGT GTATGTATAT ATATACTATG TATAACTGTG CATTCATTTA
190
TC.G..C.
200
GT
210
220
230
240
TTTTCACTAC GGAGAGTTAA AGCTCGTAAT TAATTTTTTT TATTTTACAT AAGTACATAA
250
260
270
280
290
300
TTTGCATTAT TCGTACATGT GCCCGTTCCA TTAGATCACG AGCTTAATCA CCATGCCGCG
TMMC-C75
USNM504259
USNM504260
USNM504853
LAM088901
MheNMNZ2173
..A
MpеLAM95 654
??
310
320
3.30
340
350
360
TMMC-C75
TGAAACCAGC AACCCGCTTG GCAGGGATCC CTCTTCTCGC ACCGGGCCCA TCGATCGTGG
USNM504259
USNM504260 ?????????? ?????????? ?????????? ?????????? ?????????? ??????????
USNM504853
?????????? ?????????? ?????????? ?????????? ?????????? ??????????
LAM088901
MheNMNZ2173
MpeLAM95 654
370
380
390
400
410
420
TMMC-C75 GGGTAGCTAA TATTGCCTTT TATAAGACAT CTGGTTCTTA CTTCAGGACC ATCTTAACTT
USNM504259
?????? ?????????? ?????????? ?????????? ?????????? ??????????
USNM504260 ?????????? ?????????? ?????????? ?????????? ?????????? ??????????
USNM504853
?????????? ?????????? ?????????? ?????????? ?????????? ??????????
LAM088901
MheNMNZ2173
MpeLAM95 654
..T..
TMMC-075
430
AAAATCGCCC ACTC
USNM504259 ?????????? ????
USNM504260 ?????????72 ????
USNM504853
LAM088901
?????????? 7777
MheNMNZ2173
MpeLAM95 654

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DALEBOUT ET AL. NEW SPECIES OF BEAKED WHALE
589
mandibles, teeth and skulls of M. perrini and M. hectori, including new material
from the latter species held in Southern Hemisphere museum collections, show
distinctive differences between the two species. These differences are docu-
mented below.
Mandibles and Teeth
The mandibles of M. perrini have a short symphysis (19.3%-23.2% of man-
dible length) compared to M. hectori (25.9% -33.8% of mandible length). The
ventral lateral outline of the rami is convex over the length of the symphysis
in M. perrini, whereas in M. hectori this outline is concave (Fig. 9). The entire
ventral margin of the M. hectori mandible consists therefore of two concave
areas, giving the rami a more slender appearance. This characteristic mandible
shape in M. hectori was noted by Harmer (1924).
In specimens of M. perrini, the teeth are situated slightly posterior (1-2
cm) to the tip of the mandibles. In M. hectori, this is only the case in juvenile
and sub-adult specimens; the fully adult male teeth are situated at the very
tip of the mandibles. In M. perrini, the alveoli are only slightly expanded to
take the teeth, which reach 12 mm in breadth in adult males, giving the jaw
an smoothly attenuating tip. In M. hectori, the adult male teeth are broader
(<17.5 mm) and the alveoli are consequently expanded, such that the tip of
the jaw is swollen.
There are small differences in the teeth of these species. The teeth in M.
perrini have, in profile, a smoothly convex anterior margin, whereas those of
M. hectori have a margin with three flattish areas between the denticle and
root (Fig. 10). In subadult specimens of M. hectori, these "flats" are more
defined and appear as steps in the margin. In situ, the teeth of M. perrini are
more erect, with a shorter posterior margin that those of M. hectori. The angle
formed by the denticle is 60°-65° in M. perrini and 85°-90° in M. hectori.
The female teeth of M. perrini are thin and sharply triangular with straight
or slightly concave margins, while those of M. hectori have the same flat areas
in the anterior margin as do the males of that species.
Cranial Morphology
The main difference in cranial morphology between these two species is in
the shape of the neurocranium and synvertex. In M. perrini, the synvertex
Figure 3. Aligned sequences for Mesoplodon perrini over 434 bp of mitochondrial
DNA control region. Identity to reference sequence, TMMC-C75, indicated by dots.
All five specimens of M. perrini share same haplotype over this fragment. Sequences
from M. hectori MheNMNZ2173 (underlined) and M. peruvianus MpeLAM95654 in-
cluded for comparison. Position 1 of alignment corresponds to position 15891 of fin
whale, Balaenoptera physalus, mtDNA genome (Arnason et al. 1991). Diagnostic nu-
cleotide position distinguishing M. perrini from all 20 previously described species of
beaked whales at this locus highlighted in gray

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10
20
30
40
50
ATG ATC AAC ATT CGA AAA ACA CAC CCA TTA ATA AAA ATT ATC AAC AAC GCA TTT
TMMC-C75
USNM504259 ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ???
??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ???
USNM504260
777 277 277 727 222 ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ???
USNM504853
777 777 777 777 777 777 777 277 227 222 227 277 ??? ??? ??? ??? ??? ???
LAM088901
MheSAM16387
.C...
Mpe-013141
....C.
60
..T...
C..
C.G
70
80
..C
..C G.T..T..T
T
90
100
..C
TMMC-C75
ATT GAT CTT CCC ACT CCA TCA AAC ATC TCC TCA TGA TGA AAC TTT GGT TCA TTA
USNM504259 ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??.
USNM504260 ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??....
USNM504853 777 27? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ???
LAM088901 777 777 ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??? ??.
MheSAM16387
Mpe-U13141 ..C
110
...T.C..G... T
..T
..T
120
130
140
150
160
TMMC-C75 CTC GGT CTC TGC CTA ATC ATA CAA ATT CTT ACA GGC TTA TTC CTA GCA ATA CAC
USNM504259
USNM504260
USNM504853
LAM088901
MheSAM16387
Mpe-U13141
??? ?? ??? ??? ?..
TMMC-C75
USNM504259
USNM504260
USNM504853
LAM088901
MheSAM16387
Mpe-U13141
TMMC-C75
USNM504259
USNM504260
USNM504853
LAM088901
MheSAM16387
Mpe-U13141
TMMC-C75
USNM504259
USNM504260
USNM504853
LAM088901
Mhe SAM16387
Mpe-U13141
170
180
190
200
210
TAC ACA CCA GAC ACT ACA ATA GCC TTT TCA TCA GTT GCA CAC ATC TGC CGA GAC
220
::::
230
240
250
..T
260
270
GTT AAC TAT GGT TGA ATT ATC CGA TAC CTA CAT GCA AAT GGA GCT TCC ATA TTT
280
290
300
310
320
TTT ATC TGC CTT TAC GCA CAC ATT GGA CGC GGT CTA TAT TAC GGC TCT TAT ATT
330
340
350
360
370
TTT CAA AAA ACA TGA AAT ATT GGA GTA ATT TTA CTC TTC ATA GTT ATA GCT ACT
TMMC-C75
USNM504259
USNM504260
USNM504853
LAM088901
G..
MheSAM16387 ..C
Mpe-U13141
380
TMMC-C75
GCA TTT
USNM504259
USNM504260
USNM504853
LAM088901
MheSAM16387
Mpe-U13141
Figure 4. Aligned sequences for Mesoplodon perrini over 384 bp of mitochondrial
DNA cytochrome b. Identity to reference sequence, TMMC-C75, indicated by dots.
Two variable sites, defining three unique haplotypes, were found among the five spec-
imens of M. perrini. Sequences from M. hectori MheSAM16387 (underlined) and M.
peruvianus Mpe-U13141 are included for comparison. Position 1 of alignment corre-
sponds to position 14613 of fin whale mtDNA genome (Arnason et al. 1991). Diag-

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DALEBOUT ET AL: NEW SPECIES OF BEAKED WHALE
591
narrows upwards, and the upper part of the skull is triangular in frontal view.
In M. hectori, the synvertex is flatter, giving the upper part of the skull a
rounded box-like shape. In M. perrini, the maxillaries below and behind the
synvertex on each side of the neurocranium are steep-sided and not greatly
inflated, whereas in M. hectori, the neurocranium is inflated and the maxillaries
are prominent in frontal view on each side of the synvertex. Also, in M. perrini
the margins of the posteromedial portion of the maxillaries at the uppermost
level are sharply angled laterally on each side (90° from the longitudinal me-
dian line on the right side, and 110° on the left). In M. hectori, these angles
are usually gentle (130°-140°) although one specimen of M. hectori from South
Africa (PEM 1511/15; Ross 1970) was found to have right side maxillary with
a 90° angle. Therefore, this feature may not be a reliable distinguishing char-
acter for M. perrini.
Associated with the synvertex is the space between the right and left nasals.
This is narrow and reverse V-shaped in M. perrini, but wide with parallel sides
in M. hectori. The width of this space can be expressed as the least distance
between the anterior prominences of the synvertex, which in M. perrini is
2.1%-6.7% ZW, and in M. hectori, 8.3%-18.5%.
The premaxillaries of M. perrini are noticeably narrower than M. hectori. At
the position of the antorbital notches, the width of M. perrini is 34.5%-35.4%
(relative to the distance between the apices of the antorbital notches) compared
to 43.3%-53.3% for M. hectori. At the position of the superior nares, the
width of M. perrini is 35.2% -37.2% (relative to ZW) compared to 46.0%-
49.0% for M. hectori.
There are a number of other, readily visible but less quantifiable features
which separate the skulls of these two species, the most significant of which
are:
(1) In adult males, the mesorostral groove is fully ossified in M. perrini, but
unossified in M. hectori (unusual in Mesoplodon, but the largest mature crania
(695 mm and 677 mm CBL) of both sexes of M. hectori show little ossification).
(2) M. perrini has a basirostral groove extending anterior to the maxillary
prominences, while in M. hectori this groove is very short in adults and does
not extend forward of the prominences,
(3) In M. perrini, the antorbital notch is formed by the maxillary overlaying
the jugal, and the antorbital prominence is formed by the maxillary; in M.
hectori the antorbital notch is formed by the jugal, and the antorbital promi-
nence is formed by the lacrimal.
(4) Moderately formed maxillary crests are present above the orbit in M.
perrini, but not in M. hectori.
(5) In lateral view, the rostrum of M. perrini is deep through to mid-length
nostic nucleotide position distinguishing M. perrini from all 20 previously described
species of beaked whales at this locus highlighted in gray.

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frequency
b 80
70
70
60
60
50
50
40
30
20
20
10
frequency
592
a
90
80
88
70
70
60
50
40
30
20
10
MARINE MAMMAL SCIENCE, VOL 18, NO. 3, 2002
Mesoplodon perrini sp. n.
versus all other
beaked whale species
(average, 8.55%)
intra-specific
inter-specific
pairwise sequence divergence (GTR-corrected)
mtDNA control region
low-level divergence
between Arnoux's & Baird's
beaked whales, B. arnuxii
& B. bairdii
8%.
%6
10%.
11%.
12%
13%-
14%.
15%
intra-specific
inter-specific
pairwise sequence divergence (GTR-corrected)
mtDNA cytochrome b
%91
17%
%81
Mesoplodon perrini sp. n.
versus all other
beaked whale species
(average, 15.24%)
19%
20%
21%

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DALEBOUT ET AL: NEW SPECIES OF BEAKED WHALE
593
and thereafter tapers to the tip. In M. hectori, the outline is smoothly concave
from the pterygoids to the rostrum tip.
Although Harmer (1924) suggested that the structure of the thin part of
the mesethmoid which forms the nasal septum was diagnostic for M. hectori,
our observations suggest that this is a variable character. No significant dif-
ferences were found in this feature among the specimens of M. hectori and M.
perrini examined. The presence of a dorsal sinus also varies between individuals,
and in both species, the septum rises above the level of the premaxillae in the
region of the anterior nares.
PREVIOUS ATTRIBUTIONS TO MESOPLODON HECTORI IN THE LITERATURE
A number of publications, in addition to those already mentioned (Mead
1981, Mead and Baker 1987, Henshaw et al. 1997), have erroneously used
data from specimens of M. perrini to represent M. hectori. In Mead (1984),
reproductive data from specimens of M. perrini was used for M. hectori. Mead
(1989) presented drawings of the cranium, mandible, and a tooth, as well as
photographs of the external appearance of the holotype of M. perrini
(USNM504853), as M. hectori. One of these photographs was used by Baker
(1990, 1999). Some of these photographs also appeared in Reeves and Leath-
erwood (1994). Jefferson et al. (1993) included some of the figures presented
in Mead (1989). In Mead (1993), information on the stomach morphology of
M. perrini was used for M. hectori. The artist's impression of M. hectori in
Carwardine (1995) was based on photographs of the holotype of M. perrini.
Messenger and McQuire (1998) used the mtDNA control region sequence
from the calf, USNM504259 (Henshaw et al. 1997) to represent M. hectori in
their phylogenetic analyses. Note that we have asked that this Genbank entry
be corrected to reflect the true species identity of this specimen. It is possible
that there are further publications that we are not aware of which have also
made the mistake of using specimens of M. perrini to represent M. hectori. As
no adult male specimens of M. bectori were known until relatively recently,
images of the holotype of M. perrini have unfortunately been used widely to
represent this species.
External Appearance
OTHER CHARACTERISTICS
The overall body shape of M. perrini is typical of Mesoplodon beaked whales,
with a relatively small head, a long thorax and abdomen, a deep peduncle,
Figure 5. Frequency distribution of pairwise sequence divergence within and be-
tween the 20 previously known species of beaked whales, based on: (a) 437 bp align-
ment of mtDNA control region and (b) 384 bp alignment of mtDNA cytochrome b.
Two representatives used per species where possible. GTR distance correction used to
adjust for multiple substitutions. All intraspecific (open bars) and interspecific distances
(shaded bars) are shown.

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MARINE MAMMAL SCIENCE, VOL 18, NO. 3, 2002
Table 2.
Cranial measurements for Mesoplodon perrini in mm. Methods of taking
measurements follow Moore (1963) taken on right hand side where possible. No cranial
measurements taken for USNM504259, as skull was crushed. Measurements currently
not available from TMMC-C75. E, estimated length.
Museum number
Measurement
number
USNM504853
USNM504260
LAM088901
563
594
424
511
554
365
377
414
263
412
446
264
446
483
314
501
541
371
310
334
221
262
261
203
271
282
208
10
271
276
207
11
191
208
162
12
226
232
178
13
99
97
76
14
46
37
34
15
66
61
51
16
54
43
38
17
25
27
19
18
50
59E
35
19
20
21
24
25
26
27
28
29
30
31
33
34
35
36
37
38
39
222222222-23eneran
10
16
19
15
23
25
14
105
110
88
41
40
28
96
97
77
96
98
73
159
160
108
103
115
67
61
62
45
32
36
18
31
42
25
42
39
32
32
29
23
48
40
31
259
267
186
168
185
137
74
69
64
41
43
38
81
88
78
311
346
246
40
284
317
216
41
236
259
179
42
416
452
43
449
44
90
80

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DALEBOUT ET AL: NEW SPECIES OF BEAKED WHALE
Table 2.
Continued.
Museum number
USNM
LAM
504260
088901
70
54
30
male/calf
Measurement
USNM
number
504853
45
64
46
595
sex/age class
male/adult
205
female/adult
greatest
Definitions of cranial measurements (numbers in parentheses refer to Moore 1963):
1 = condylobasal length (1), 2 = tip rostrum to posterior extension maxillary plate
(7); 3 tip rostrum to anterior margin superior nares (8); 4 tip rostrum to anterior
point maxillary crest (9); 5 tip rostrum to posterior extension premaxilla on lateral
tip of right premaxillary crest (11); 6 = tip rostrum to posterior extension temporal
fossa (10); 7 tip rostrum to apices of antorbital notches (2); 8 breadth skull across
orbital centres (19); 9 = breadth skull across postorbital process frontals (17); 10 =
breadth skull across zygomatic processes squamosals (18), 11 = least breadth skull
across posterior margins temporal fossae (20); 12 = greatest breadth skull across ex-
occipitals (25); 13 = greatest span occipital condyles (21); 14 greatest width of an
occipital condyle (22); 15 greatest length of an occipital condyle (23); 16 =
breadth foramen magnum (24); 17 = greatest length of right nasal on vertex (15); 18
length nasal suture (16); 19 = extension right premaxilla posterior to right nasal
on vertex (28); 20 greatest breadth nasals on vertex (26), 21 = least distance between
anterior prominences of the synvertex (27); 22 = greatest span premaxillary crests (29);
23 = greatest transverse width of superior nares (37); 24 = least width premaxillae
where narrow opposite superior nares (30); 25 greatest width premaxillae anterior
to position of previous (31); 26 = width rostrum in apices of antorbital notches (33);
27 = width rostrum in apices of prominential notches (34); 28 = least distance be-
tween main maxillary foramina (41); 29 = least distance between premaxillary foramina
(42); 30 = distance posterior margin of left maxillary foramina to anterior margin
maxillary prominence (43); 31 = width rostrum at mid-length of rostrum (35); 32 =
width premaxillae at mid-length of rostrum (32); 33 depth rostrum at mid-length
rostrum (36), 34 height of skull (39); 35 = external cranial height; 36 = greatest
length of temporal fossa (13); 37 = width of temporal fossa (40); 38 = length of orbit
taken from mid-point of frontals (14); 39 tip rostrum to posterior extension of
maxilla between pterygoids (6); 40 = tip rostrum to anterior extension of pterygoid
sinus (12); 41 = tip rostrum to most anterior extension of pterygoids (5); 42
rostrum to posterior margin of pterygoid mid-line (3); 43 = tip rostrum to posterior
extension of wing of pterygoid (4); 44 length of vomer visible at surface of palate
(44); 45 width between pterygoid notches (38); 46 = amount added to rostrum
because of breakage (45).
tip
and short tail (Fig. 8a-e). The rostrum is relatively short compared to all other
species in the genus, except M. hectori and M. peruvianus. In calves, the rostrum
appears to be shorter and stubbier than in adults. The blowhole is broad and
crescent-shaped, with anterior-pointing tips. The melon forms a small bulge,
and the mouthline is straight. Throat grooves are present. External measure-
ments are shown in Table 4.
The adult male (USNM504853) was dark gray dorsally grading to white
ventrally. The ventral side of the tail flukes was pale gray with converging
striations and there was a white patch around the umbilicus. The adult female

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MARINE MAMMAL SCIENCE, VOL. 18, NO. 3, 2002
Table 3. Mandibular measurements for Mesoplodon perrini. Measurements follow
methods of Moore (1963), taken on right hand side where possible. No measurements
available for USNM504259 as mandible has been lost. Measurements currently not
available from TMMC-C75. E, estimated length.
Museum number
Measurement
USNM
USNM
number
504853
504260
LAM
088901
47
516
486
356
48
387
379
289
49
455
402
314
50
120
108
69
51
97
52
37E
21
53
35
37
25
54
47
25
55
109
56
16
19
57
52
64
58
47
59
12
60
n/a
33
n/a
61
9.9 g³
sex/age class
male/adult
37.0 gh
female/adult
male/calf
right tooth, dry weight.
b right tooth, wet weight.
Definitions of measurements (numbers in parentheses refer to Moore 1963): 47 =
mandibular length (1); 48 = length from posterior extension of symphysis to condyles
(6); 49 = length posterior margin of alveolus to condyles (7); 50 = greatest length of
symphysis (2); 51 greatest height of mandible at coronoid processes (3); 52 = outside
height of mandible at midlength of alveolus (4); 53 = inside height of mandible at
midlength of alveolus (5); 54 = length of alveolus (8); 55 = width of alveolus (9); 56
tip of mandible to alveolus (10); 57 = greatest tooth length (11); 58 greatest
tooth width (12); 59 greatest tooth breadth (13); 60 = height of crown of tooth;
61 = tooth weight.
(USNM504260) was too decomposed to allow information on color pattern to
be collected. Calves (USNM504259, LAM088901 and TMMC-C75) are light
to dark gray dorsally and white ventrally (Fig. 8a-d). The lower jaw and throat
regions are white. A dark gray region extends from the corner of the mouth
and encompasses the eye and the rostrum, forming an extended mask. The
flippers are medium to dark gray dorsally and white ventrally. There is a
lighter-colored patch on the anterodistal portion. The flukes are dark gray
dorsally and medium to light gray ventrally. The ventral surface includes a
pattern of white striations that converge posteromedially.
Ontogeny and Reproduction
Both adult specimens (USNM504260, female, and USNM504853, male)
were considered physically mature based on the fusion of the thoracic epiphyses

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a
b
DALEBOUT ET AL. NEW SPECIES OF BEAKED WHALE
597
Figure 6. Skull photographs, USNM504853, holotype: (a) lateral view, (b) dorsal
view. Scale pattern in (b) = 5 cm, divided into 50-mm squares, and pertains to both
figures.
to the centra and the disappearance of the epiphyseal suture (Mead 1981).
While the mandibular rami of the male were fused at the symphysis, those of
the female were not. The teeth of the male were fully erupted (Fig. 7), and
the mesorostral canal completely filled, suggesting that it was sexually mature.
The testis weights were; 84.5 g (L), and 115.5 g (R). The adult female ap-
peared to have been dead for about two weeks when discovered (Mead 1981),
and was too decomposed for the reproductive organs to be recognizable. It is
likely that this animal was the mother of the calf (USNM504259), which had
been found in approximately the same location a week earlier (Table 1). Both

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a
b

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DALEBOUT ET AL: NEW SPECIES OF BEAKED WHALE
599
animals share the same haplotype at the mtDNA control region and cyto-
chrome b (Fig. 3, 4), as expected for a cow-calf pair. Although there has been
little opportunity to date for calibration and standardization in the aging of
Mesoplodon species, thin tooth sections were prepared from the right tooth of
both adult specimens, and the cemental growth layer groups (GLG's) counted,
giving an approximate age of nine years each, although the male appeared the
more mature of the two from cranial features (Mead 1981).
The 245-cm calf (LAM088901) had a squid eye lens in its stomach, which
suggests that it had been weaned. There are no data on stomach contents for
the other two calves (USNM504259 and TMMC-C75), but the fimbriated
edge on the tongue of the former animal suggests that it was still suckling
(Fig. 8d). The Monterey calf (TMMC-C75) had small, immature testes, which
were elliptical in shape and weighed approximately 1.5 g each.
NATURAL HISTORY
Food Habits
Only two of the animals had stomach contents. The stomach of a third
(TMMC-C75) was collected, but is currently not available for examination. A
squid eye lens, not identifiable to species, was found in the stomach of the
second calf (LAM088901). Two lower beaks of the squid, Octopoteuthis deletron,
and a fragment of an unidentifiable vertebrate were found in the stomach of
the adult female (USNM504260; Mead 1981). We assume that like many
other beaked whales, this species mainly eats pelagic squid.
Behavior
The adult male (USNM504853) bore a number of white, linear scars on its
postcranial flanks which were probably inflicted by the teeth of other males
of the species. However, it is noted that the scars on the adult male appear to
have been made with a single tooth, rather than with two teeth simultaneously,
as might be expected in species with apical teeth (e.g., Heyning 1984).
Parasites
Three soft-stalked barnacles, Conchoderma auritum, were found on the teeth
of the adult male (USNM504853), and a number of cysts of the cestode,
Phyllobothrium sp., were found encased in the blubber. Several oval scars (ap-
proximately 3 x 5 cm) were found on the flanks. The Monterey calf (TMMC-
C75) bore three similar scars, in various stages of healing. Such scars are likely
due to cookie-cutter shark attacks (Isistius spp.; Jones 1971). This calf was
Figure 7. Teeth of adult male (USNM504853) in situ: (a) oblique lateral view; (b)
anterior view. Scale bars = 30 mm. Photograph credit: J. G. Mead.

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MARINE MAMMAL SCIENCE, VOL. 18, NO. 3, 2002
b
d
e
Figure 8.
External photographs. (a) left lateral view of calf (TMMC-C75), scale bar
= 30 cm; (b) ventral view of calf (TMMC-C75), scale bar 30 cm; (c) right lateral
view of calf (TMMC-C75), scale bar = 30 cm; (d) right lateral view of head of calf
= 20 cm; (e) left lateral view of head of adult male
(USNM504259), scale bar
(USNM504853), scale bar = 24 cm. Note that striped pattern on right lateral side of
calf (TMMC-C75) is the imprint caused by bed of utility truck on which animal was
lying. Photograph credits: (a-c) M. Haulena; (d, e) G. Carsten and J. G. Mead.

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C
DALEBOUT ET AL: NEW SPECIES OF BEAKED WHALE
601
M. hectori
M. hectori
50mm
Figure 9.
M. perrini
Lateral view of mandibular ramus of Mesoplodon hectori (a, NMNZ2173
adult male; b, NMNZ614 juvenile male) and M. perrini (c, USNM504853 adult male).
M. perrini
40mm
M. hectori
Figure 10. Adult male tooth of Mesoplodon perrini (USNM504853) and M. hectori
(NMNZ2173). Right tooth of each species is shown, with anterior margin facing left.

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602
External measurements for Mesoplodon perrini. Due to extent of decomposition, few measurements were available from adult female
Table 4.
(USNM504260).
Museum or field number:
MARINE MAMMAL SCIENCE, VOL. 18, NO. 3, 2002
USNM504853
Male/adult
USNM504260
Female/adult
USNM504259
Male/calf
LAM088901
Male/calf
TMMC-C75
Male/calf
Sex/age class:
Measurement:
cm
cm
cm
cm
cm
Total length
390.0
100.0
443.0
100.0
210.0
100.0
245.0
100.0
223.5
100.0
Snout to center of blowhole
35.1
9.0
22.7
10.8
27.0
11.0
18.5
8.3
Snout to center of eye
Snout to angle of mouth
Snout to ear
Snout to anterior insertion of flipper
Snout to center of umbilicus
Snout to genital slit (center)
Snout to anus
42.5
10.9
25.3
12.0
28.5
11.6
25.5
11.4
24.0
6.2
19.5
9.3
15.2
6.2
14.0
6.3
50.2 12.9
29.6
14.1
33.7
13.8
72.0 18.5
48.4
23.0
51.7
21.1
47.0
21.0
150.0 38.5
123.4
50.4
105.5
47.2
254.0 65.1
140.0
66.7
162.0
66.1
142.0
63.5
285.0 73.1
159.0
75.7
184.8 75.4
160.1
71.6
Snout to tip of dorsal fin
240.0 61.5
274.0
61.9
140.0
66.7
162.9
66.5
146.0
65.3
Girth at anus
Girth at eye
Girth at axilla
Maximum girth
Girth midway anus to fluke notch
120.0 30.8
84.5
34.5
72.5
32.4
196.0 50.3
127.0
51.8
107.0
47.9
230.0 59.0
119.5
53.5
130.0 33.3
36.7
70.0
31.3
66.0 16.9
48.0
21.5
Height at same place as above
30.6
7.8
24.0
10.7
Thickness at same place as above
9.0
2.3
9.5
4.3
Projection lower/upper jaw
1.6
0.4
0.6
0.3
0.6
0.2
2.0
0.9
Length of eye opening
3.2
4.0
1.8
0.8
Center of eye to ear
9.0
2.3
5.4
2.6
7.0
2.9
Center of eye to angle of mouth
18.8
4.8
6.0
2.9
17.1
7.0
12.5
5.6
Blowhole width
9.9 2.5
4.8
2.3
5.8
2.4
5.7
2.6
Length of throat grooves
20.0
5.1
9.7
4.6
19.7
8.0
14.0
6.3
693

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Museum or field number: USNM504853
Table 4.
Continued.
USNM504260
USNM504259
Male/adult
Female/adult
Male/calf
LAM088901
Male/calf
TMMC-C75
Male/calf
Sex/age class:
Measurement:
cm
cm
cm
cm
cm
Flipper length, anterior
46.0
11.8
40.0
9.0
28.0
13.3
31.5
12.9
30.5
13.6
Flipper length, posterior
Flipper width, maximum
Fluke width
Fluke depth
29.0
7.4
32.0
7.2
19.0
9.0
20.1
8.2
24.0
10.7
13.2
3.4
7.6
3.6
9.0
3.7
8.5
3.8
103.0
26.4
89.0
20.1
46.7
22.2
52.0
21.2
58.0
26.0
32.0
8.2
18.0
8.6
20.8
8.5
19.0
8.5
Depth of fluke notch
NA
1.9
0.9
NA
1.2
0.5
Dorsal fin height
19.0
4.9
10.2
4.9
13.6
5.6
9.0
4.0
Length dorsal fin base
50.0
12.8
23.0
11.0
18.0
8.1
603

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MARINE MAMMAL SCIENCE, VOL. 18, NO. 3, 2002
severely emaciated, with minimal body fat and atrophied muscles. No parasites
were found in gross necropsy, but a histological examination of abscesses in
the periumbilical region revealed that they contained degenerating cestodes
(likely Phyllobothrium spp., possibly Phyllobothrium delphini) and other foreign
material. Structures resembling parasitic granulomas were found within the
gastrointestinal tract and one of the lymph nodes (necropsy performed by F.
Gulland, University of California at Davis; details courtesy of M. Haulena).
Distribution
This species is known from five specimens found beachcast along the Cal-
ifornian coast between Torrey Pines State Reserve, just north of San Diego
(32°55'N, 117°15'W) and Fisherman's Wharf, Monterey (36°37'N,
121°55'W). Although this stranding pattern is suggestive of an eastern North
Pacific distribution, there are too few records to date to draw any bounds on
this. Little can be concluded from the presence of cookie-cutter shark scars on
the Monterey calf. Isistius spp. are limited in their northern distribution, at
least in surface waters (Nakano and Tabuchi 1990), but the occurrence of such
scars on cetaceans is not (Jones 1971). This suggests either that these cetaceans
are migratory and pass through the territory of Isistius spp., or that the dis-
tribution of Isistius spp. extends farther north in deeper waters and they attack
cetaceans when they dive. Given the habitat preferences of other ziphiids, we
assume that M. perrini is found primarily in oceanic waters, over 1,000 m in
depth.
DISCUSSION
According to the International Code of Zoological Nomenclature (ICZN),
formal classification of M. perrini ideally requires genetic validation of the
holotype of M. hectori, as four of the five known specimens of M. perrini were
described previously as this species (Mead 1981). The holotype of M. hectori
is held by the British Museum (BM(NH) 1677/76.2.16.3), and consists of the
skull, mandible, scapulae, hyoids, cervical vertebrae and flippers of a juvenile
male collected in Titahi Bay, Wellington, New Zealand, in the 19th century
(Flower 1878; Gray 1871). We have attempted, but not been successful in
extracting and amplifying native DNA from the M. hectori holotype. This may
be due to the coat of varnish on this specimen, the acidic components of which
can degrade the already low levels of endogenous DNA contained in skeletal
material (Cooper 1994). In the absence of genetic data from the holotype of
M. hectori, validated specimens from New Zealand and Australia have been
used to represent this M. hectori in these analyses.
Given the increasing use of genetic information as a universal character in
species identification, systematics, and biodiversity assessment (Bisby 2000,
Wilson 2000), the designation of genetic voucher material should be consid-
ered for all taxa. It has been suggested that mtDNA typing of holotype spec-
imens should become part of standard museum protocol (Dalebout 2002), but

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DALEBOUT ET AL. NEW SPECIES OF BEAKED WHALE
605
it is recognised both that: (1) not all specimens will retain sufficient native
DNA for analysis due to a combination of age, the nature of the material
available (e.g., bone, tooth, pelts, feathers), and museum preparation and stor-
age methods and (2) not all specimens will be amenable to current DNA
extraction techniques (e.g., Wayne et al. 1999). In such cases, genetic material
should be obtained from other validated specimens of the species in question
(e.g., Dizon et al. 2000). Genetic voucher material, in the form of DNA se-
quences from a suite of loci, could be archived both at the institution holding
the holotype (or alternative validated specimen from which these data were
derived), as well as in the international genetic database, Genbank. When
describing new species, a genetic description should also be included wherever
possible.
A recent analysis of discovery trends has suggested that at least 40 species
of large marine animals still remain to be described (Paxton 1998). If so, it
is likely that several new cetacean species will be among them, including at
least one new form of Bryde's whale, Balaenoptera edeni sp. (e.g., Baker et al.
1996, Yoshida and Kato 1999), and possibly further species of ziphiids. The
current discovery represents the second new species of Mesoplodon discovered
in the last decade. Both were dependent on the opportunistic collection of
beachcast specimens and victims of fisheries bycatch (Reyes et al. 1991, this
paper). Yet, there is no reason to assume that all cetacean species can be
encountered in this way. Some may be distributed in areas far from shore or
shore-bound currents, where human presence is still minimal. To ensure that
such species do not go undocumented, we recommend that biopsy samples be
collected wherever possible from animals encountered on sighting cruises. In
addition to traditional morphological information, the collection of tissue sam-
ples from stranded and incidentally caught animals should also become stan-
dard procedure.
ACKNOWLEDGMENTS
We are extremely grateful to all those who gave us access to specimens used to establish
the beaked whale DNA reference database that has been so valuable for these and other
discoveries (see http://www.sbs. auckland.ac.nz/research-groups/ecology_and_evolution/
molecular ecol_evo_l_lab for a full list) For the work concerning M. perrini, we would
especially like to thank the following people and institutions for collection and access
to specimens, laboratory support, valuable discussion and other aid: field staff of the
New Zealand Department of Conservation; A. E. Dizon, M. Henshaw, and K. Rob-
ertson, NOAA SWFSC; J. E. Heyning, Los Angeles County Museum of Natural
History; P. Jenkins, A. Warlow, R. Sabin, Natural History Museum (BMNH), London;
S. J. O'Brien, N. Yuhki, Laboratory of Genomic Diversity, National Cancer Institute
(also M. Culver, V. David, G. M. Lento, J. Martenson, B. Murphy, M. Raymond, J.
Pecon-Slattery, and G. K. Pei); G. Carsten, J. M. Coe, J. R. Henderson, D. B. Holts,
K. LeVeille, W. F. Perrin, W. A. Walker, M. Haulena, NOAA SWFSC and The Marine
Mammal Centre; C. W. Potter, Smithsonian National Museum of Natural History; D.
J. Saul, A. G. Rodrigo, and V. Ward, University of Auckland; K. Van Waerebeek,
CEPEC, Peru. For access to specimens of M. bectori, we would like to thank C. Kemper,
SAM, P. Sabine, TAM, and N. Prosser-Goodall, MAAMA. We would also like to thank
our two reviewers, F. Cipriano and G. J. B. Ross, for their very useful comments on

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606
MARINE MAMMAL SCIENCE, VOL. 18, NO. 3, 2002
the manuscript. MLD was supported by a William Georgetti Scholarship, a University
of Auckland Doctoral Scholarship, and a Smithsonian Institution Short-Term Visitor's
Fellowship. This project was supported by grants to C. S. Baker from the University
of Auckland, the New Zealand Lottery Board, the New Zealand Marsden Fund, and
the International Fund for Animal Welfare.
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Received: 08 July 2001
Accepted: 14 January 2002

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NOTE INTORNO ALLA DISTRIBUZIONE
DELLA
FAUNA VERTEBRATA
NELL'OCEANO
prese durante un viaggio intorno al Globo
1865-68
dai Professore
ENRICO HILLYER GIGLIOLI
MEMBRO DELLA SOCIETÀ GEOGRAFICA ITALIANA, DELLA ZOOLOGICA DI LONDRA
DELL' ENTOMOLOGICA DI FIRENZE ECC.
Con una Carta.
Two- finned whalebone whate
75-76
FIRENZE,
STABILIMENTO DI GIUSEPPE CIVELLI
Via Panicale, 39.
1870.

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tri
l'At-
806;
mo-
il più
anco
ogici
gran-
balc-
assa-
ora,
il 10
mo
pure
pas-
vista
pinna
iso-
dei
1 go-
janco
oleva
inna
edere
ferire
com-
uper-
vidi.
grossi
pro-
que-
olari
nbini
come
Balaenoptera. Sp.?
Una grossa specie, la quale per quanto ho potuto giudicare vedendola
imperfettamente per alcuni minuti, non sembrava diversa dalla B. mu-
sculus dell'Atlantico, presentando come quella la pinna dorsale molto in-
dietro; la parte esposta del corpo era di un bruno-nero, non fece vedere
la tesla.
Fu incontrata il 1° settembre 1866, allo sbocco del Golfo di Yeddo.
Non meno di quattro balenottere e due balene sono state descritte c
baltezzate in tutte le regole, da soli disegni cinesi e giapponesi, sono
perciò specie più che dubbiosc. I Giapponesi danno ai grossi cetacci il
nome generico di kugira.
BALAENOPTERA? Sp. e gen. nov.
Il dopo pranzo del 4 settembre 1867 nel Pacifico, (lat. 28' 34' S.; long,
88° 10' O. Gr.); mentre stavo pescando minuti organismi pelagici alla
scala reale, fui interrotto da un grande getto d'acqua che venne quasi a
spruzzarmi, e vidi emergere dal mare la schiena grigio-verdastra di una bella
balenottera, sul culmine della quale si vedevano due pinne ben marcate,
separate da uno spazio di circa due metri. Queste pinne erano grandi, c
ben sviluppate relativamente alle dimensioni dell'animale, che poteva avere
un 18 metri di lunghezza. La testa era proporzionata, il muso largo e ar-
rotondato, le mascelle quasi uguali, quella inferiore però più sporgente;
gli occhi, come in tutti i cetacei, erano piccoli, le aperture delle narici
situate sulla sommità della testa, e poste l'una vicino all'altra, davano pas-
saggio al getto d'acqua misto ad aria, che aveva richiamate la mia atten-
zione; l'animale muoveva intanto la mascella inferiore, come se masticasse
qualcosa. La coda era relativamente grande, come pure le pinne pettorali;
e mentre l'animale si voltava sul fianco destro per tuffarsi sott'acqua, vidi
essere il ventre di color bianco puro, la pelle del petto mi sembrò liscia
e senza rughe. In questa occasione ebbi la rara opportunità di vedere
chiaramente tutti i movimenti di quel grosso cetaceo, che nuotava di poco
al di sotto della superficie, la coda è il vero motore, agisce precisamente
come una potente clice, le pinne pettorali erano poco usate, ed allora aiu-
tavano a cambiare la direzione, facendo da limoni supplementari.
Questa balenottera girò intorno alla Magenta fregando i suoi fianchi
contro il bordo, ed ebbi tutto l'agio di esaminarla bene per oltre un quarto
l'ora, mentre si stava ammainando una lancia onde vederla più da vicino.
Intorno ad essa erano molti pesci, non potci determinarne la specie.
Alcuni dei nostri vollero cercare di darle la caccia, ma non avevamo
i mezzi per questo, malgrado la buona volontà del nostromo, il quale andò
a cercare la sua migliore fiocina, ed una sessantina di metri di buona
func, ci voleva altro.
Scendemmo nella lancia, ma l'animale spaventato dal rumore non tardo
a sprofondarsi, e non comparve più.
l'er quanto io mi sappia, questa è la prima balenoltera osservata con

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due pinne sul dorso, e mi credo in dovere di considerarla non solo sic-
come specie nuova, ma come il tipo di un nuovo genere, pel quale pro-
pongo il nome di AMPHIPTERA; questo genero sarebbe per ora caratterizzato
dalla presenza di due pinne dorsali: augurando che presto qualche na-
turalista sia abbastanza fortdnato per avere a sua disposizione il cadavere
di uno di esse, onde dare al mondo scientifico una descrizione più com-
pleta di un cetaceo così interessante. La specie tipo, e per ora unica,
porterebbe il nome di AMPHIPTERA PACIFICA.
Non è improbabile che si trovi pure sulle coste meridionali dell' Au-
stralia, e su quelle della N. Zelanda.
Come abbiamo detto, non si conoscono altre balenottere munite di due
pinne mediane, è bensì vero che Rafinesque descrisse molti anni fa, un
delfino con due pinne dorsali, l'Oxypterus Mongitori, dei mari della Sici-
lia; ma d'allora in poi mai più visto. Ed i naturalisti Quoy e Gaimard che
accompagnarono Freycinet nel suo viaggio di circumnavigazione (1817-20),
descrissero, per averla veduta un momento a volo d'uccello, nel Pacifico
tra le isole Sandwich e l'Australia, un'altra specie più grande, il Delphi-
(nus (Oxypterus) rhinoceros, cosi chiamata per avere una pinna od una
prominenza curva indietro posta sull'occipite; anch'essa non fu più rive-
duta.
BALAENA.
I caratteri sono quasi tutti gli stessi di quelli che definiscono il genere
precedente, le balene si distinguono però per la mancanza della pinna
dorsale, per avere la testa relativamente molto più grossa, la bocca piu
grande, a margini sinuosi, la mascella inferiore più grande e più profonda
della superiore, i fanoni più sviluppati; il corpo più tozzo, e lo strato
adiposo soltocutaneo assai più spesso.
Sono le specie di questo genere che vengono pescate con maggior
profitto.
Balaena australis. Desmoul.
Questa specie ha la testa più proporzionata che non la B. mysticetus,
che rappresenta nei mari australi, ove ha una distribuzione molto estesa,
trovandosi in certe stagioni nei paraggi del Capo di Buona Speranza, in-
torno alla Nuova Zelanda, alla terra di Kerguelen, all'Australia meridio-
nale, ed alle terre più meridionali dell' America.
È stata in questi ultimi anni oggetto di una caccia assai attiva, resa
facile per l'abitudine che aveva di avvicinare le coste all'epoca della ri-
produzione; da Hobart Town (Tasmania) partivono molti bastimenti per
quella caccia; ora, il numero di questi cetacei è assai diminuito ina è
tuttora molto più considerevole che non quello della balena vera, nei mari
artici.
Dal 1830 al 1831 non meno di 91 bastimenti inglesi, montati da 2050
marinai, e rappresentando una portata di 30,083 tonnellate, pescavano la
balena nei mari del Sud, mentre dal 1840 al 1843 non se ne contavano

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THE BOOK OF
WHALES

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A.570
WRITTEN AND ILLUSTRATED BY
RICHARD ELLIS
ALFRED A KNOPF NEW YORK
1980
USEF
ROGEN QUE U

[PAGE BREAK]

said that calling an animal a whale does not necessarily
make it one.
Cetaceans are found in all the oceans of the world,
from deep within the polar ice to the tropics. There
are freshwater dolphins, restricted to certain rivers;
and one species, the whitefin dolphin, is found only in
Tung Ting Lake and its adjacent waters in China. The
killer whale is probably the most widely distributed of
all cetaceans, found in polar, temperate, and tropical
oceans all over the world. Many species are migratory,
moving from one area to another in clearly defined
routes, whereas in other species the movements and
distribution are poorly known. In recent years certain
species have turned up in unexpected locations; for
example, Fraser's dolphin, Lagenodelphis hosei, which
was previously known only from a skeleton found on
a beach in Borneo, has now appeared in the eastern
tropical Pacific, off Taiwan, off Australia, and in the
Caribbean.
There is even the possibility of still-undiscovered
cetaceans. Some of the beaked whales are known only
from single skeletons, and there may be Mesoplodons
whose remains have not yet washed ashore and been
discovered by knowledgeable cetologists. On a re-
mote beach in Tierra del Fuego, R. N. P. Goodall
(1978) discovered the collected remains of some four-
teen species of cetaceans and added substantially to
our knowledge of the range, habits, and osteology of
many of them. In 1964, during a whale-marking and
observation cruise off the coast of Chile, a number of
cetologists saw and photographed many of the ex-
pected species, including blue, fin, sei, sperm, and
smaller cetaceans. (As whalers will attest, the west
coast of South America is a particularly productive
area for their efforts.) On November 24 and 27, how-
ever, "there were eight sightings . of small whales
which we could not identify" (Clarke et al. 1978). The
investigators chased and photographed these odon-
tocete, single-blowhole, beakless whales, which had
light-colored heads and dark bodies, were in a school
of fifteen to twenty animals, and were about 20 feet
(6.0 meters) long. (Only Risso's dolphin, Grampus gri-
seus, fits this general description, but it is not known
to exceed a length of 13 feet.) To demonstrate the
exciting state of basic cetological research, I quote the
observer's concluding paragraph in full:
We can only conclude that these whales seen off Chile in
1964 may be a new species, possibly of the genus Pseudorca
or Grampus. It is clearly important to obtain specimens as
soon as opportunity affords. The animal agrees well with the
"undescribed whale" reported and figured by Wilson (1905,
P. 472; 1907, p. 4 and Whales Plate I) as 20-30 feet long,
black above but with some white around the mouth or chin,
and characterized by a high dorsal fin "erect, pointed and
sabre-shaped"; several were seen in the Ross Sea during the
[AD] British National Antarctic Expedition (1901-1904), but they
have never been reported since.
way:
In 1967 a sport-fishing boat captain saw eighteen to
twenty small cetaceans near the island of Tobago in
the northeastern Caribbean. He described them in
"The heads back to the dorsal fin and abdo-
this
mens were white, and the backs brownish gray. On the
larger ones, the dorsal fin was close to ca. 0.6m (2 feet)
high and hooked slightly backward" (Erdman et al.
1973). These whales, which were about 16 to 18 feet
(4.9 to 5.5 meters) long, might very well be the same
kind of mysterious animal observed by Clarke off
Chile.
gray
Some species exist in uncountable numbers, roam-
ing the open oceans far offshore, but others, closer to
human habitation, shipping, and industry, are all too
easily counted-and killed. The California whale,
which was known in historical times from the western
Pacific and the eastern Atlantic, exists now as a relict
population of some 15,000 animals that pass close to
the shoreline of western North America. The Indus
River susu, Plantanista minor, another primitive spe-
cies, is probably the most endangered of all cetaceans,
having been reduced to some five hundred-odd ani-
mals in the Indus River while hunting continues.
The cetaceans are perhaps the most highly special-
ized of all mammals. They are more or less fishlike in
shape and, like the fishes, have a caudal fin used for
propulsion and pectoral fins used for steering. The
caudal fin of a cetacean, commonly known as the
flukes, functions in an up-and-down movement,
whereas the fishes, whose tails lie in the vertical plane,
use a side-to-side motion. (In defining the whale, Mel-
ville wrote that it "is a spouting fish with a horizontal
tail.") Most cetaceans have a dorsal fin, which is be-
lieved to help stabilize the animal in the water. Neither
the caudal fin nor the dorsal fin has any bony internal
support structure, but both are composed solely of
connective tissue. Although some species have a num-
ber of hairs on the head, for the most part the mam-
malian hair in cetaceans has been replaced in function
by a layer of fat or blubber, which serves to insulate
the animal against the cold of its warmth-absorbing
habitat. Like all other mammals, whales and dolphins
give birth to live young and nurse them by means of