Content uploaded by Ismael Mijan
Author content
All content in this area was uploaded by Ismael Mijan
Content may be subject to copyright.
105
GEODIVERSITAS • 2013 • 35 (1)
© Publications Scientiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com
Bianucci G., Miján I, Lambert O., Post K. & Mateus O. 2013. — Bizarre fossil beaked whales
(Odontoceti, Ziphiidae) shed from the Atlantic Ocean oor off the Iberian Peninsula.
Geodiversitas 35 (1): 105-153. http://dx.doi.org/10.5252/g2013n1a6
ABSTRACT
Forty partial fossil skulls belonging to beaked whales (Cetacea, Odontoceti,
Ziphiidae) were collected by trawling and long-line fishing on Neogene (prob-
ably Late Early to Middle Miocene) layers of the Atlantic floor off the coasts
of Portugal and Spain (Asturias and Galicia). e systematic study of the most
diagnostic Iberian specimens, those preserving the rostrum and the dorsal part
of the cranium, led to the recognition of two new genera (Globicetus n. gen.
Giovanni BIANUCCI
Università di Pisa, Dipartimento di Scienze della Terra,
via S. Maria, 53, I-56126 Pisa (Italy)
bianucci@dst.unipi.it
Ismael MIJÁN
Sociedade Galega de Historia Natural, Apdo. 356, S-15480 Ferrol (Spain)
ismaelmijan@edu.xunta.es
Olivier LAMBERT
Muséum national d’Histoire naturelle, Département Histoire de la Terre,
57 rue Cuvier, F-75231 Paris cedex 05 (France)
and Institut royal des Sciences naturelles de Belgique,
Département de Paléontologie,
29 rue Vautier, B-1000 Brussels (Belgium)
olivier.lambert@naturalsciences.be
Klaas POST
Natuurhistorisch Museum Rotterdam,
P.O. Box 23452, NL-3001 Rotterdam (The Netherlands)
klaaspost@fishcon.nl
Octávio MATEUS
Universidade Nova de Lisboa, CICEGe, Faculdade de Ciências e Tecnologia,
FCT, 2829-516 Caparica (Portugal)
and Museu da Lourinhã,
Rua João Luís de Moura, 95, P-2530 Lourinhã (Portugal)
omateus@fct.unl.pt
Bizarre fossil beaked whales (Odontoceti,
Ziphiidae) fished from the Atlantic Ocean
floor off the Iberian Peninsula
106 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
KEY WORDS
Cetacea,
Odontoceti,
Ziphiidae,
Neogene,
Miocene,
Portugal,
Spain,
phylogeny,
skull morphology,
new genera,
new species.
and Imocetus n. gen.) and four new species (Choneziphius leidyi n. sp., G. hibe-
rus n. gen., n. sp., I. piscatus n. gen., n. sp., and Tusciziphius atlanticus n. sp.).
Based on the matrix of a previous work, the phylogenetic analysis places all the
new taxa in the subfamily Ziphiinae Gray, 1850. More fragmentary specimens
are tentatively referred to the genera Caviziphius Bianucci & Post, 2005 and
Ziphirostrum du Bus, 1868. Among these new ziphiids, extremely bizarre skull
morphologies are observed. In G. hiberus n. gen., n. sp. the proximal portion
of the rostrum bears a voluminous premaxillary spheroid. In T. atlanticus n. sp.
a medial premaxillary bulge is present on the rostrum; together with asymmet-
ric rostral maxillary eminences at the rostrum base, this bulge displays various
degrees of elevation in different specimens, which may be interpreted as sexual
dimorphism. Specimens of I. piscatus n. gen., n. sp. bear two sets of even crests:
spur-like rostral maxillary crests and longitudinal maxillary crests laterally bor-
dering a wide and long facial basin. A preliminary macroscopic observation of
these elements indicates very dense bones, with a compactness comparable with
that of cetacean ear bones. Questioning their function, the high medial rostral
elements (the premaxillary spheroid of G. hiberus n. gen., n. sp. and the medial
bulge of T. atlanticus n. sp.) remind the huge rostral maxillary crests of adult
males of the extant Hyperoodon ampullatus (Forster, 1770). In the latter, the
crests are very likely related to head-butting. However, they are made of much
more spongy bone than in the fossil taxa studied here, and therefore possibly
better mechanically suited for facing impacts. Other interpretations of these
unusual bone specializations, related to deep-diving (ballast) and echolocation
(sound reflection), fail to explain the diversity of shapes and the hypothetical
sexual dimorphism observed in at least part of the taxa. e spur-like rostral
maxillary crests and long maxillary crests limiting the large facial basin in
I. piscatus n. gen., n. sp. and the excrescences on the maxilla at the rostrum
base in Choneziphius spp. are instead interpreted as areas of origin for rostral
and facial muscles, acting on the nasal passages, blowhole, and melon. From a
palaeobiogeographic point of view, the newly described taxa further emphasize
the differences in the North Atlantic (including Iberian Peninsula) and South
African Neogene ziphiid faunal lists. Even if the stratigraphic context is poorly
understood, leaving open the question of the geological age for most of the
dredged specimens, these differences in the composition of cold to temperate
northern and southern hemisphere fossil ziphiid faunas may be explained by a
warm-water equatorial barrier.
RÉSUMÉ
Étranges baleines à bec fossiles (Odontoceti, Ziphiidae) pêchées sur le fond de l’Océan
Atlantique au large de la péninsule ibérique.
Quarante crânes partiels fossiles de baleines à bec (Cetacea, Odontoceti, Ziphii-
dae), pêchés au chalut et à la palangre sur des couches du Néogène (probable-
ment fin du Miocène inférieur à Miocène moyen) du fond de l’Atlantique au
large des côtes du Portugal et d’Espagne (Asturies et Galice), sont signalés.
L’étude sytématique des spécimens ibériques les plus diagnostiques, ceux dont
le rostre et la partie faciale sont préservés, a permis la reconnaissance de deux
nouveaux genres (Globicetus n. gen. et Imocetus n. gen.) et de quatre nouvelles
espèces (Choneziphius leidyi n. sp., G. hiberus n. gen., n. sp., I. piscatus n. gen.,
n. sp. et Tusciziphius atlanticus n. sp.). Sur la base de la matrice d’un travail
précédent, l’analyse phylogénique positionne l’ensemble des nouveaux taxons
107
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
INTRODUCTION
Including at least 21 extant species (Dalebout et al.
2002), beaked whales (Ziphiidae Gray, 1850) are the
second most species-rich modern cetacean family,
following the Delphinidae Gray, 1821. In addition
to dental reduction, these deep-diving, generally
teuthophagous animals are characterized by various
skull specializations, some of these specializations
being sexually dimorphic, that place them as one
of the most peculiar odontocete groups and lead
to contrasted functional interpretations (Heyning
1984; Buffrénil et al. 2000; MacLeod 2002; Lam-
bert et al. 2011). For a long time the fossil record
of ziphiids was scarce compared to its high present
diversity. Even now, the number of fossil species
dans la sous-famille Ziphiinae Gray, 1850. Des spécimens plus fragmentaires
sont provisoirement attribués aux genres Caviziphius Bianucci & Post, 2005
et Ziphirostrum du Bus, 1868. Parmi ces nouveaux ziphiidés, des morpholo-
gies extrêmement bizarres sont observées. Chez G. hiberus n. gen., n. sp., la
partie proximale du rostre porte une volumineuse sphère prémaxillaire. Une
crête médiale prémaxillaire est présente sur le rostre de T. atlanticus n. sp.;
cette crête, de même que des éminences rostrales maxillaires asymmétriques,
montre différents degrés d’élévation au sein de l’espèce, peut-être en lien avec
du dimorphisme sexuel. Les spécimens d’I. piscatus n. gen., n. sp. portent deux
paires de crêtes : des crêtes rostrales maxillaires en forme d’éperon et des crêtes
maxillaires longitudinales bordant latéralement un long et large bassin facial.
L’observation macroscopique préliminaire de ces éléments indique un os très
dense, avec une compacité comparable à celle des os de l’oreille des cétacés.
Au niveau de leur fonction potentielle, les éléments médians du rostre (sphère
prémaxillaire de G. hiberus n. gen., n. sp. et crête médiane prémaxillaire de
T. atlanticus n. sp.) rappellent les énormes crêtes maxillaires rostrales des mâles
adultes de l’espèce moderne Hyperoodon ampullatus (Forster, 1770). Chez ce
dernier, les crêtes sont très probablement utilisées lors de combats par coups de
tête. Cependant, elles sont constituées d’un os beaucoup plus spongieux que
chez les taxons fossiles étudiés ici, et donc peut-être plus aptes mécaniquement
à subir des impacts. D’autres interprétations des ces spécialisations osseuses
inhabituelles, liées aux plongées profondes (ballast) et à l’écholocalisation
(réflexion des sons), échouent à expliquer la diversité des formes et le possible
dimorphisme sexuel observé chez une partie des espèces. Les crêtes rostrales
maxillaires en forme d’éperon et les longues crêtes maxillaires limitant le grand
bassin facial d’I. piscatus n. gen., n. sp. et les excroissances sur le maxillaire à la
base du rostre de Choneziphius spp. sont, elles, interprétées comme des régions
d’origine pour des muscles rostraux et faciaux, agissant sur les conduits nasaux,
l’évent et le melon. D’un point de vue paléobiogéographique, les nouveaux
taxons confirment les différences de contenu des listes fauniques de ziphiidés
néogènes de l’Atlantique Nord (incluant la péninsule ibérique) et de l’Afrique du
Sud. Malgré le contexte stratigraphique peu précis, laissant ouverte la question
de l’âge géologique de la plupart des spécimens pêchés sur le fond de la mer,
ces différences dans la composition des faunes de ziphiidés fossiles des régions
froides à tempérées des hémisphères nord et sud pourraient être expliquées par
la présence d’une barrière d’eau équatoriale chaude.
MOTS CLÉS
Cetacea,
Odontoceti,
Ziphiidae,
Néogène,
Miocène,
Portugal,
Espagne,
phylogénie,
morphologie crânienne,
genres nouveaux,
espèces nouvelles.
108 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
based on specimens from inland deposits remains
small (e.g., Muizon 1984; Bianucci et al. 1994,
2010; Lambert & Louwye 2006; Lambert et al.
2009; Bianucci et al. 2010). However, specimens
recovered from the bottom of oceans proved to be
an essential source of information. Fossils, generally
isolated rostra, have been reported from the South
Pacific Ocean (Fordyce & Cullen 1979; Miyazaki &
Hasegawa 1992), Indian Ocean (Robineau 1973),
and Sea of Japan (Horikawa et al. 1987; Tazaki et al.
1987). Whitmore et al. (1986) also mentioned sev-
eral ziphiid fossils from other deep ocean sites and
analyzed in more detail the nature of these strange
and strongly phosphatized skull elements.
Later Bianucci et al. (2007, 2008) reported eight
new fossil ziphiid genera and ten new species, based
on better preserved cranial material recovered from
trawling activities on the ocean bottom along the
South African coasts, at depths of as much as 1000 m.
Here again, the relationships with phosphorite de-
posits were emphasized, allowing some argument
about the still problematic dating of the specimens
and the high local productivity (Bianucci et al. 2007).
Following a preliminary note (Miján 2007),
the present article describes a new large sample of
well preserved fossil ziphiid skulls, some of them
displaying unusual morphologies, recovered from
the Atlantic Ocean floor off Portugal and Spain
(Asturias and Galicia).
To our knowledge, the present article, together with
the Miján (2007) note, represent the first scientific
report of fossil ziphiids from Iberian Peninsula. Zbysze-
wski (1954) referred to the new species Palaeo ziphius
melidensis an incomplete mandible (MG5450) col-
lected in the Tortonian of Melides (Southwest Por-
tugal). However, the genus Palaeoziphius Abel, 1905,
primarily used for the species P. scaldensis (du Bus,
1872), based on another isolated mandible, has been
placed in Odontoceti incertae sedis by Lambert (2005).
Judging from Zbyszewski’s illustrations (Zbyszewski
1954: pls 1, 2), the mandible from Melides does not
exhibit any ziphiid character; consequently P. meliden-
sis must also be referred to Odontoceti incertae sedis.
e only reliable previous fossil ziphiid record from
Portugal (but not from Iberian Peninsula) originates
from the Archipelago of Azores and was referred to
Mesopodon sp. by Estevens & Ávila (2007).
ABBREVIATIONS
IEO Instituto Español de Oceanografia, Gijón,
Spain;
IGF Museo di Geologia e Paleontologia
dell’Università di Firenze, Florence, Italy;
MG Museu Geológico, Lisboa, Portugal;
MHNUSC Museo de Historia Natural Luis Iglesias,
Universidad de Santiago de Compostela,
Santiago de Compostela, Spain;
ML Museu da Lourinhã, Lourinhã, Portugal;
NMB Natuurhistorisch Museum Boekenberg,
Antwerp;
NMR Natuurhistorish Museum Rotterdam;
SGHN Museo da Natureza da Sociedade Galega
de Historial Natural, Ferrol, Spain.
MATERIAL AND METHODS
SPECIMENS AND LOCALITIES
We examined 40 partial ziphiid skulls recovered
from the sea floor off the coasts of Asturias, Gali-
cia, and Portugal during fishing activities based on
bottom set long-line and bottom trawl (Fig. 1).
All the specimens were collected on the borders
of the continental platform, most of them at a
depth ranging between 500 and 1000 m. Most
of the specimens were kept in private collections
for years; the location is precise in some cases, but
more approximate in others, for two reasons: some
were collected before the Global Positioning System
(GPS) was widely used and fishermen are often
reluctant to reveal their fishing spots. e fossils
collected off the Asturias coast are from the Can-
yon of Avilés and other imprecise localities along
the platform. e fossils collected off Galicia are
from several localities, among which As Paredes,
A Selva, and Cortada fishing grounds. e spec-
imens from Portugal were found in deep water
off central Portugal (Lourinhã, Peniche), south of
Nazaré Canyon; the latter extends about 210 km
westward from the coast and reaches depths near
5000 m in its distal part (Tyler et al. 2009). All
the fossils examined are now kept in IEO, ML,
MHNUSC, and SGHN.
All the fossils are strongly phosphoritized; some
of them are even partly included in a phosphatic
conglomerate. Fossilization and associated sediments
are actually similar for fossil ziphiids trawled off the
South African coasts (Bianucci et al. 2007).
109
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
As commonly observed for the phosphoritized
fossil beaked whale remains recovered from the sea
floor (Fordyce & Cullen 1979; Whitmore et al. 1986;
Horikawa et al. 1987; Tazaki et al. 1987; Bianucci
et al. 2007), all Iberian skulls are incomplete. e
preserved bones are the ones that are more compact
in the living ziphiids: bones from the rostrum and
in some cases from the facial area of the cranium,
including the vertex (Lambert et al. 2011). e
more spongy and/or delicate bones forming the
posteroventral portion of the braincase are gener-
ally not preserved.
Following Bianucci et al. (2007), we decided to
use only those specimens that include at least the
more diagnostic dorsal surface of the cranium,
including the vertex, for the description of new
taxa. More fragmentary fossils (e.g., isolated rostra)
originating from the same geographic area are listed
in the referred material only if their morphological
features fully overlap those of the more complete
SPAIN
Lisboa
Lourinhã
Peniche
PORTUGAL
ASTURIAS
BAY OF BISCAY
1
2
3
4
5
6
44°N
200
200
200
1000
1000
1000
2000
2000
2000
4000
4000
4000
3000
3000
3000
8°W 4°
42°
40°
7
8
9
Gijón
Ortegal Spur
Nazare Canyon
GALICIA
Coruña
100 km
Ferrol
FRANCE
EUROPE
Iberian
peninsula
Atlantic
Ocean
Mediterranean sea
Fig.1. — Map of Atlantic Iberian coast showing localities where fossil ziphiid skulls have been recovered by bottom set long-line and
trawling: 1, Asturias continental border, imprecise location; 2, Avilés Canyon, Asturias coast; 3, As Paredes shing ground (44°10’N,
8°20’W), depth 250-600 m; 4, A Selva shing ground (44°10’N, 8°40’W), depth 350-600 m; 5, Cortada shing ground (43°36’N, 9°0’W),
depth 400-800 m; 6, oceanographic research at 1500 m depth (42°27’N 11°59’W); 7, south of Nazaré Canyon, imprecise location
(c.39°18’N, 9°47’W); 8, escarapote shing ground (42°10’N, 09°26’W), depth 680-750 m; 9, 20 miles from Touriñán Cape (42°50’N,
9°40’W), depth 1500 m.
110 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
specimens. Eighteen of the 40 skulls are considered
as diagnostic. All these specimens are described for
the first time in this article, with the exception of
SGHN MA0632 and SGHN MA0644, which
were previously tentatively referred to the genus
Hyperoodon Lacépède, 1804 (Miján 2007).
SYSTEMATICS
e systematic classification used in the following
section is based on the phylogenetic analysis pub-
lished by Bianucci et al. (2010), here confirmed
with the addition of new taxa (see phylogenetic
paragraph). Most of the described material is dem-
onstrated to belong to the subfamily Ziphiinae Gray,
1850, which is redefined in this article. Following
Bianucci et al. (2010), this subfamily excludes
Beneziphius Lambert, 2005, Messapicetus Bianucci,
Landini & Varola, 1992 and Ziphirostrum du Bus,
1868, three genera forming, possibly together with
Aporotus du Bus, 1868, a more basal clade of the
ziphiid phylogenetic tree (“Messapicetus clade”).
SYSTEMATIC PALAEONTOLOGY
Order CETACEA Brisson, 1762
Suborder ODONTOCETI Flower, 1867
Family ZIPHIIDAE Gray, 1850
Subfamily ZIPHIINAE Gray, 1850
TYPE GENUS. — Ziphius Cuvier, 1823.
OTHER GENERA INCLUDED. — Choneziphius Duvernoy,
1851, Globicetus n. gen., Imocetus n. gen., Izikoziphius
Bianucci, Lambert & Post, 2007, Tusciziphius Bianucci,
1997, and possibly Caviziphius Bianucci & Post, 2005.
EMENDED DIAGNOSIS. — With the exception of Iziko-
ziphius and Ziphius, members of the subfamily Ziphiinae
differ from all other Ziphiidae in the dorsal closure of
the mesorostral groove by medial sutural contact of the
premaxillae extending posteriorly until the bony nares.
With the exception of Choneziphius and Imocetus n. gen.,
they further differ from all other Ziphiidae in having very
elongated nasals with the anterior tip of nasals located
anterior to the premaxillary crests (ratio between length
of medial suture of nasals and maximum width of na-
sals > 1.1). ey further differ from all other Ziphiidae,
with the exception of the clade formed by Africanacetus
Bianucci, Lambert & Post, 2007, Hyperoodon, Ihlen-
gesi Bianucci, Lambert & Post, 2007, and Mesoplodon
Gervais, 1850, in having the ascending process of the
premaxilla concave in lateral view, with the posterodorsal
portion partly overhanging the bony nares (apart from
Choneziphius planirostris (Cuvier, 1823), with bony na-
res still visible in dorsal view). ey further differ from
all other Ziphiidae, except Beneziphius, Messapicetus,
and Ziphirostrum, in having the left premaxillary crest
anterolaterally directed.
Genus Choneziphius Duvernoy, 1851
T
YPE
SPECIES
. — Choneziphius planirostris from southern
North Sea Basin, probably Late Miocene (Lambert 2005).
OTHER SPECIES INCLUDED. — Choneziphius leidyi n. sp.
EMENDED DIAGNOSIS. — Choneziphius differs from all
other ziphiid genera in the mesorostral groove dorsally
closed at the level of the antorbital notches by the joined
medial margins of the premaxillary sac fossae, forming
a prominent ridge posteriorly shifted to the left, and
separating deeply concave anterior portions of the pre-
maxillary sac fossae.
It also differs from the other ziphiine genera in the maxilla
covered at the rostrum base with prominent excrescen-
cies. Moreover it differs from Ziphius and Izikoziphius
in the medial fusion of the premaxillae dorsally closing
the mesorostral groove; from Globicetus n. gen., Imoce-
tus n. gen., and Tusciziphius in lacking an extremely
ossified trapezoidal vertex with the anterior part of the
nasals contacting the premaxillary crests; from Imo-
cetus n. gen. in lacking a wide facial depression, a rostral
maxillary spur-shaped crest, and in the location of the
premaxillary foramen (not posterior to the level of the
antorbital notch); from Globicetus n. gen. in lacking a
large spherical medial premaxillary prominence at the
rostrum base; from the possible ziphiine Caviziphius in
shallower premaxillary sac fossae and in the more slender
and lower right premaxillary crest.
DISCUSSION
e fossil record of Choneziphius is primarily based
on several partial skulls and rostra referred to the
type species C. planirostris. ese fossils have been
collected in sediments of North Sea, probably dated
from the late Miocene (Belgium, Netherlands,
and UK; see Lambert [2005] for bibliography and
review). Lankester (1870) described C. packardi
Lankester, 1870 based on an incomplete rostrum
from Suffolk (UK). Leidy (1876, 1877) described
111
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
C. liops Leidy, 1876 and C. trachops Leidy, 1876,
based on fragmentary rostra that are now lost, from
the Phosphate Beds of South Carolina (USA).
While reviewing the genus Choneziphius, Lam-
bert (2005) considered C. packardi as based on
too fragmentary material to allow specific or even
generic determination; he considered C. liops as a
possibly valid species (rostrum shorter and with
anterior narrowing stronger than in C. planiro-
stris), and C. trachops as possibly conspecific with
C. planirostris. According to Lambert (2005),
Proroziphius macrops Leidy, 1876 and probably
P. chonops Leidy, 1876, both based on fragmentary
and unfortunately lost skulls from the Phosphate
Beds of South Carolina, should be included in
the genus Choneziphius. Whitmore & Kaltenbach
(2008) considered C. trachops as a valid taxon and
assigned to this species a large rostrum collected
from reworked sediments at the Lee Creek Mine,
North Carolina. Although the above mentioned taxa
show the apomorphies of the genus Choneziphius
(at least on illustrations), we restrict these species,
based on too fragmentary material, to their holo-
types and consider them as incertae sedis.
Choneziphius leidyi n. sp.
(Figs 2-5; Table 1)
HOLOTYPE. — SGHN MA0633, partial skull including
rostrum, facial area and vertex.
REFERRED SPECIMENS. — SGHN MA0640, partial skull
including rostrum, facial area and vertex, Escarapote
fishing ground, depth of approximately 685 m, off the
Galician coast, 42°08’N, 09°26’W; SGHN MA0641,
partial skull including posterior portion of rostrum,
part of facial area and vertex, A Selva fishing ground,
depth of approximately 500 m, off the Galician coast,
44°10’N, 08°40’W; SGHN MA0937, partial skull in-
cluding rostrum and facial area, A Selva fishing ground,
depth of approximately 500 m, off the Galician coast,
44°10’N, 08°40’W; ML 533, partial skull including
rostrum and facial area, south of Nazaré Canyon, off
the Portuguese coast, exact locality unknown but likely
around 39°18’N, 9°47’W; ML 1366, fragment of skull
including the left dorsal surface of the cranium with
the left premaxillary crest, south of Nazaré Canyon off
the Portuguese coast, exact locality unknown, but likely
around 39°18’N, 9°47’W.
E
TYMOLOGY
. — In honour of the American palaeontolo-
gist Joseph Leidy (1823-1891), who described several
Choneziphius-like fossil ziphiids from the Mid Atlantic
Coastal Plain of the USA in 1876 and 1877.
TYPE LOCALITY. — A Selva fishing ground, depth of
approximately 500 m, off the Galician coast, 44°10’N,
08°40’W.
DIAGNOSIS. — Large species of Choneziphius differing
from C. planirostris in: longer and more pointed rostrum
with apex constructed of premaxillae alone; longer dorsal
opening of the mesorostral groove at the apex of the ros-
trum; premaxillary foramina located distinctly anterior
to level of prominental notch; lower maxillary crest on
the supraorbital process; shallower and wider premaxil-
lary sac fossae; less asymmetric premaxillary sac fossae;
higher vertex overhanging the bony nares.
DESCRIPTION
e rostrum, although longer than in Choneziphius
planirostris, exhibits the same massive appearance;
in both species it is relatively narrow with a subcy-
lindrical anterior half portion. As in C. planirostris,
the facial area is wider than long. Differing from
C. planirostris, the bony nares are not visible in
dorsal view, being hidden by the overhanging and
anteriorly projected vertex.
Premaxilla
On the complete rostra of the holotype and SGHN
MA0640, the apex of the rostrum is formed by
the premaxillae only, contrary to C. planirostris
where maxilla and premaxilla both reach the apex
of the rostrum. Anteriorly, the premaxillae are not
fused dorsally, leaving the narrow tunnel-shaped
mesorostral groove open for a length of at least
80 mm. Such a dorsal exposure of the mesoros-
tral groove is rarely present in the large sample of
C. planirostris, and, if present, never longer than
50 mm. For most of the rostrum length, the thick
premaxillae are firmly fused at midline, with a
suture remaining visible until the bony nares (as
in C. planirostris).
As in C. planirostris, the fused premaxillae form
a prominent ridge at the rostrum base, posteriorly
shifted to the left and separating the deeply excavated
anterior portions of the premaxillary sac fossae (the
main character defining the genus Choneziphius).
Each premaxillary sac fossa contains a premaxillary
112 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
foramen at its anteriormost point, located well an-
terior to the level of the prominental notch of the
maxilla (Fig. 2). In C. planirostris, the premaxillary
foramen is located at, or just posterior to, the level
of the prominental notch. An additional foramen
is observed on the medial margin of the left pre-
maxillary sac fossa of SGHN MA0640.
e right premaxillary sac fossa is distinctly wider
than the left (ratio between maximum width of left
and right fossae between 0.70 and 0.76, n = 3), and
vertical foramina left premaxillary crest
frontal
nasal
frontal
maxillary crest
antorbital notch
maxillary tubercle
A
B
prominental notch
left premaxillary sac fossa
right premaxillary sac fossa
right premaxillary crest
dorsal infraorbital foramen
dorsal infraorbital foramen
premaxillary foramen
premaxilla
maxilla
maxilla
ecrescence on maxilla
prominent ridge formed by fused premaxillae
Fig.2. — Skull of Choneziphius leidyi n.sp. (SGHN MA0633, holotype): A, dorsal view; B, corresponding line drawing. Tight parallel
lines indicate a break surface; more widely spaced parallel lines indicate supercial wear. Scale bar: 10cm.
113
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
the asymmetry is therefore less pronounced than in
C. planirostris (ratio between 0.48 and 0.65, n = 15).
e anterior part of the vertex, including the
ascending process of the premaxilla and the pre-
maxillary crest, is overhanging the premaxillary sac
fossae and the bony nares. is condition contrasts
with C. planirostris, in which in lateral view the
ascending process forms an angle of ≤ 90° with the
horizontal plane of the skull, whereas the angle is
> 90° in C. leidyi n. sp. e fairly slender premaxil-
lary crest is anterolateraly directed. e right crest
is distinctly larger than the left, as in C. planirostris.
Maxilla
From a roughly vertical orientation on the anterior
half of the rostrum, the lateral surface of the maxilla
progressively shifts to a subhorizontal dorsal surface
bordered by an acute lateral margin and the thick
premaxilla medially. In this part, the dorsal side
of the maxilla is covered with multiple, marked
excrescences. Rostra of C. planirostris bear similar
excrescences in the same area, usually less prominent
than in the known specimens of C. leidyi n. sp.
On the lateral surface of the maxilla, a marked alveolar
groove is visible which sharply slopes down from its
right premaxillary crest
A
B
left premaxilla
maxillary crest
maxilla-palatine suture mark
vestigial alveolar groove
right premaxilla
excrescences on maxilla
maxilla
maxilla
premaxilla
premaxillary bulge
frontal
jugal
lacrimal
Fig.3. — Skull of Choneziphius leidyi n.sp. (SGHN MA0633, holotype); A, lateral view; B, corresponding line drawing. Cross-hatching
indicates the presence of a concretion. Scale bar: 10cm.
114 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
uttermost point on the maxilla until approximately
⁹⁄₅ of the rostrum length. Within the groove SGHN
MA0640 shows 12-13 very shallow alveoli probably
corresponding to vestigial teeth. A few specimens
of C. planirostris also show traces of shallow alveoli.
From the prominent maxillary tubercle, the maxilla
forms a maxillary crest on the supraorbital process,
with a roughly antero-posterior direction. is crest
is less prominent than in C. planirostris, a feature
especially noticeable in anterior view.
Each maxilla is pierced by two dorsal infraorbital
foramina, one just behind the prominental notch,
and the other lateral to the vertex.
Nasal
e dorsal surface of the nasals is strongly worn in
each specimen, but their outline is clearly discern-
able thanks to the conspicuous sutures with the
premaxillary crests, frontals, and mesethmoid, with
a condition similar to C. planirostris. In the latter
the nasals are rarely preserved, and if preserved
they show a more spongy aspect than surrounding
bones. is is maybe the reason why they are easily
damaged and lost in Choneziphius.
Frontal
Frontals are heavily worn on the vertex of the
holotype and other referred specimens. However,
judging from the short distance between the ascend-
ing processes of the maxillae, they were originally
transversally narrower than the nasals, a condition
similar to C. planirostris. e supraorbital process
of the frontal is anteriorly bordered by the lacrimal
and the maxilla.
Table 1. — Measurements (in mm) on the skulls of Choneziphius leidyi n.sp. from the Atlantic Ocean oor off the Iberian Peninsula.
Abbreviations: e, estimate; +, nearly complete; –, no data.
SGHN
MA0633
(holotype)
SGHN
MA0640
SGHN
MA0641
ML
533
Rostrum length from level of antorbital notch +503 +495 –408
Rostrum length from level of prominental notch +488 +482 – –
Distance from apex of rostrum to bony nares +588 +585 – –
Length premaxillary portion of rostrum –+67 – –
Height of rostrum at anterior end of maxilla –42 – –
Width of rostrum at mid-length 82 74 –94
Width of premaxillae at mid-length of rostrum 50 e50 – –
Height of rostrum at mid-length 82 71 – –
Width of rostrum base at prominental notch 195 – – –
Width of rostrum base at antorbital notch 252 – – 205
Width of premaxillae at rostrum base (antorbital) 87 98 – –
Minimum distance between maxillae near rostrum base 37 –25 –
Distance rostrum base – anterior apex of palatine e115 e109 – –
Preorbital width of skull 354 – – –
Longitudinal distance right premaxillary foramen – rostrum base (antorbital) 52 49 – –
Longitudinal distance left premaxillary foramen – rostrum base (antorbital) 63 52 – –
Width of premaxillary sac fossae 172 168 158 150
Width of right premaxillary sac fossa 96 95 e82 92
Width of left premaxillary sac fossa 69 67 64 58
Width of bony nares 73 80 e81 –
Minimum width of right ascending process of the premaxilla 54 48 +45 –
Width of premaxillary crests 178 170 +155 –
Width of right premaxillary crest 65 71 +55 –
Width of left premaxillary crest 43 45 e48 –
Minimum distance between premaxillary crests 83 e74 e60 –
Maximum width of nasals e89 e84 e72 –
Minimum posterior distance between maxillae –86 80 –
115
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
Vomer
On the midline of the ventral side of the rostrum
of the holotype, a narrow exposure of vomer is vis-
ible over a length of 166 mm, from a level 181 mm
posterior to the apex of the rostrum.
Palatine
e palatine is only partially preserved in the holotype.
e anteriormost point of the maxilla-palatine suture
is 115 mm anterior to the level of the antorbital notch.
REMARKS
e most striking differences between C. leidyi
n. sp. and C. planirostris are the general size and
the rostrum length. erefore one has to wonder
whether these differences (and the other differences)
could be related to ontogeny and/or sexual dimor-
phism. Measurements of all available specimens
of C. planirostris (all from the North Sea) show a
mean rostrum length of 359 mm (n = 27, min =
297 mm, max = 416 mm), whereas C. leidyi n. sp.
Fig.4. — Skull of Choneziphius leidyi n. sp. (SGHN MA0633, holotype): A, anterior view; B, corresponding line drawing; C, ventral
view. Cross-hatching indicates the presence of a concretion. Scale bar: 10cm.
maxilla
maxilla
premaxilla
vomer
palatine
maxillary tubercle
maxillary tubercle
rostral tunnel
dorsal infraorbital
foramen
maxillary crest
ascending
process of
premaxilla
A
C
B
mesethmoid
bony
nares
left premaxillary
crest
prominent
ridge
on fused
premaxillae
maxilla
frontal
jugal lacrimal
excrescences
on maxilla
prominental notch
antorbital notch
premaxilla
vestigial alveolar groove
116 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
(from the Atlantic coast off Galicia) reports 499
mm (n = 2, min 495 mm, max 503 mm). It seems
obvious that the large North Sea sample cannot be
seen as a sexual dimorphic variant of the much larger
Galician specimens. Indeed, variation within the
North Sea sample (from very slender specimens to
more robust specimens, see Lambert [2005: fig. 21])
shows all the aspects of sexual and/or ontogenetic
variation within a same species.
Genus Tusciziphius Bianucci, 1997
T
YPE
SPECIES
. — Tusciziphius crispus Bianucci, 1997, from
Tuscany (Italy), early Pliocene, calcareous nannofossil
zone MNN14-15 (Bianucci et al. 2001).
O
THER
SPECIES
INCLUDED
. — Tusciziphius atlanticus n. sp.
EMENDED DIAGNOSIS. — Tusciziphius differs from all
other ziphiines except Imocetus n. gen. and Globicetus
n. gen. in having an extremely ossified trapezoidal ver-
tex in which the anterior part of the nasals contact the
premaxillary crests. It differs from Globicetus n. gen.
and Imocetus n. gen. in the extreme widening and an-
terior projection of the right premaxillary crest, and in
the lesser posterior constriction of the vertex; it further
differs from Globicetus n. gen. in lacking a large spher-
ical medial premaxillary rostral prominence; it further
differs from Imocetus in lacking a wide facial depression,
a rostral maxillary spur-shaped crest, and in having the
premaxillary foramen not located posterior to the level of
the antorbital notch. Among the other ziphiine genera it
further differs from Ziphius and Izikoziphius in the medial
fusion of the premaxillae closing the mesorostral groove;
it further differs from the possible ziphiine Caviziphius in
the shallower excavation of the premaxillary sac fossae.
Tusciziphius atlanticus n. sp.
(Figs 6-9; Table 2)
HOLOTYPE. — SGHN MA0926, a partial skull includ-
ing rostrum, facial area, and vertex.
PARATYPE. — NMR 9991-3020, a partial skull includ-
ing rostrum, facial area, and vertex, originally referred
to Tusciziphius crispus (see Post et al. 2008). Morgan
River, Beaufort County, South Carolina, USA, between
32°26’50”N, 80°35’57”W and 32°27’09”N, 80°28’44”W.
Found reworked on the bottom of a river, it has been
proposed to originate from late Miocene-Pliocene layers
(Post et al. 2008).
R
EFERRED
SPECIMENS
. — SGHN MA0632, a partial skull
including rostrum, facial area, and vertex, As Paredes
fishing ground, depth of approximately 470 m, off the
Galician coast, 44°07’N, 08°07’W; SGHN MA0644, a
partial skull including rostrum, facial area, and vertex,
A Selva fishing ground, depth of approximately 500 m, off
the Galician coast, 44°10’N, 08°40’W; SGHN MA0914,
a partial skull including rostrum, facial area, and vertex,
A Selva fishing ground, depth of approximately 500 m,
off the Galician coast, 44°10’N, 08°40’W; ML1365, a
right facial area including right side of the vertex, south
of Nazaré Canyon off the Portuguese coast, exact locality
unknown, but likely around 39°18’N, 9°47’W.
ETYMOLOGY. — From the Atlantic Ocean distribution of
the species (South Carolina, east coast USA and Iberian
Atlantic coast, western Europe).
T
YPE
LOCALITY
. — Cortada fishing ground, depth of
approximately 600 m, off the Galician coast, 43°30’N,
09°25’W.
DIAGNOSIS. — Tusciziphius atlanticus n. sp. differs from
all other ziphiids in the prominent medial rostral bulge
formed by the fused premaxillae, which elevation varies
individually and is probably related to sexual dimorphism
(this feature cannot be observed in T. crispus, of which
the rostrum is unknown; in Aporotus the elevated pre-
maxillae are not fused in a single bulge).
It differs from T. crispus in having the right premaxil-
lary sac fossa almost completely (except the posterior
portion) filled by compact bone forming a semicircular
shelf and in having the excavation for the premaxillary
sacs restricted to the posterior portion of both right and
left premaxillary sac fossae.
DESCRIPTION
e holotype, the paratype, and the referred speci-
mens of Tusciziphius atlanticus n. sp., share with
T. crispus the similar size, the large and asymmetrical
premaxilary sac fossae, and the vertex architecture
with the extreme widening and anterior projection of
the right premaxillary crest. All the Iberian specimens
referred to this species exhibit, when preserved, a
moderately elongated rostrum, suggesting that the
apparently short rostrum of the paratype may be
an artefact due to restoration of the broken apex
with plaster (Post et al. 2008).
A prominent medial elevation on the rostrum,
formed by the fusion of premaxillae over the meso-
rostral groove, is visible in three of the four speci-
mens with a preserved rostrum. In the paratype
NMR 9991-3020 and in SGHN MA0914, the
117
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
FIG. 5. — A-C, Skull of Choneziphius leidyi n.sp. (SGHN MA0640); A, dorsal view; B, lateral view; C, detail of the rostrum in lateral
view showing the vestigial alveolar groove; D, C. leidyi n. sp. (SGHN MA0641), partial skull in dorsal view; E, C. leidyi n.sp. (SGHN
ML533), partial skull in dorsal view. Scale bar: 10 cm.
additional foramen
premaxillary foramen phosphorite concretion
medial fusion of
premaxillae
prominent ridge
formed by fused premaxillae
A
B
E
CD
concave ascending
process of premaxilla
phosphorite concretion
maxillary crest
prominental notch
maxillary tubercle
antorbital notch
excrescences on maxilla
deep excavation on anterior
portion of premaxillary sac fossae dorsal exposure of the mesorostral
goove due to worn premaxillae
anterior end
of maxilla vertex overhanging
the bony nares
118 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
rostral bulge is present but less developed. is
character cannot be observed in T. crispus because
the rostrum is not preserved on the holotype and
only preserved specimen.
Premaxilla
Due to the apical erosion of the rostrum, more or
less pronounced in all specimens, it is not possible
to evaluate the length of the portion of the rostrum
formed by the premaxillae alone. In fact, this region
is partly preserved (5 mm) only in SGHN MA0914.
In all the specimens, the preserved portion of the
rostrum exhibits thick premaxillae with a medial
suture, dorsally closing the mesorostral groove. is
closure starts from the preserved apical portion of
the rostrum in all specimens except in the holo-
type, where the first 65 mm of the groove are still
dorsally open. e fused premaxillae are massive
and protuberant on the rostrum; they form a bulge,
with a varying position and height, on which the
medial premaxillary suture is completely obliterated.
e development of the medial premaxillary
bulge extends from the apex of the rostrum to
the level of the antorbital notch in the holotype,
SGHN MA0632 and SGHN MA0914, and to
the level of the anterior palatine suture in SGHN
MA0644. e elevation of the bulge increases
anteroposteriorly progressively in the holotype,
and more abruptly in SGHN MA0644 and SGHN
MA0632. e maximum height of the bulge above
the maxilla is 90 mm in SGHN MA0632, 67 mm
in SGHN MA0644, and 58 mm in the holotype.
In the paratype and in SGHN M0914, only a
small dome, respectively 27 and 33 mm above
the maxilla, is present in the posterior part of the
rostrum, just anterior to the antorbital notches.
Table 2. — Measurements (in mm) on the skulls of Tusciziphius atlanticus n.sp. from South Carolina (USA) and the Atlantic Ocean
oor off the Iberian Peninsula. Abbreviations: e, estimate; +, nearly complete; –, no data.
NMR
9991–
3020
SGHN
MA0632
SGHN
MA0644
SGHN
MA0914
SGHN
MA0926
(holotype)
Rostrum length –––+425 +440
Length of premaxillary portion of rostrum –––+5 –
Distance from apex of rostrum to bony nares ––––+525
Width of rostrum at mid-length –––63 64
Height of rostrum at mid-length –––89 82
Width of premaxillae at mid-length of rostrum –––26 50
Width of rostrum base at antorbital notch –––e150 230
Width of premaxillae at rostrum base 86 – – 68 62
Minimum distance between maxillae near rostrum base – – e51 –62
Distance rostrum base – anterior apex of palatine e130 – – e170 190
Preorbital width of skull –––e350 320
Postorbital width of skull +372 ––––
Width of premaxillary sac fossae 173 e171 142 170 150
Width of right premaxillary sac fossa 108 106 92 105 96
Width of left premaxillary sac fossa 51 61 46 58 41
Width of bony nares 69 e73 53 70 66
Minimum width of right ascending process of premaxilla 68 e60 60 – –
Width of premaxillary crests e184 188 155 –180
Width of right premaxillary crest +101 e87 e76 –101
Width of left premaxillary crest 51 33 29 48 44
Minimum distance between premaxillary crests e39 –e58 42 61
Maximum width of nasals –––55 58
Maximum width of right nasal e27 – – 29 34
Maximum width of left nasal –––27 24
Lenght of right nasal 82 e67 50 72 88
Length of medial suture of nasals 68 – – 62 71
Minimum posterior distance between maxillae 84 e105 –88 76
119
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
Macroscopic observation of transverse sections
along the medial bulge of SGHN MA0632 reveal
a high compacity of the bone tissue and the pres-
ence of a series of growth layers, a feature already
noted in the pachyosteosclerotic rostrum of several
other ziphiid taxa (Lambert 2005; Buffrénil &
left premaxillary
crest posterior excavation of the premaxillary sac fossae
concave left premaxillary sac fossae
antorbital notch
sulci
maxilla
maxilla
nasals
dorsal infraorbital foramina
filled right premaxillary sac fossa
filled right premaxillary
sac fossa
dorsal infraorbital foramen
right premaxillary crest
vestigial alveolar groove
maxilla-palatine suture marks
rostral maxillary
eminence
rostral maxillary
eminence
premaxilla
left premaxillary crest
bulge on fused premaxillae
bulge on fused premaxillae
open mesorostral groove
at the apex of rostrum
longitudinal depression
posterior excavation
of the premaxillary
sac fossae
A
B
C
D
right premaxillary
crest
Fig.6. — Skull of Tusciziphius atlanticus, n.sp. (SGHN MA0926, holotype): A, dorsal view; B, corresponding line drawing; C, lateral
view; D, detail of the vertex and premaxillary sac fossae in anterior view. Parallel lines indicate a break surface. Scale bar: 10cm.
120 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
Lambert 2011; Lambert et al. 2011). Posterior to
the premaxillary bulge, there is a low medial shelf
laterally delimited by two shallow depressions and
posteriorly margined by the premaxillae sac fossae.
e premaxillary sac fossae are strongly asym-
metric in all the specimens of T. atlanticus n. sp.
(ratio between the left and right width ranging
from 0.43 to 0.57) and in T. crispus (0.44). How-
ever, in all the specimens of T. atlanticus n. sp. the
anterior part of the right premaxillary sac fossa is
completely filled by compact bone, forming a thick
semicircular shelf. e filling is absent in the left
premaxillary fossa, which is deeply concave in all
specimens. Instead, in T. cripsus both premaxillary
sac fossae are excavated. e deep concavity of the
premaxillary sac fossae is likely a derived condition
shared with Caviziphius, Choneziphius, Globicetus
n. gen., and Imocetus n. gen., whereas the semicir-
cular shelf that partially fills the right premaxillary
fossa in T. atlanticus n. sp. may be homologous
with the rectangular premaxillary shelf of Globi-
cetus n. gen. (see below). Due to the presence of
the anterior shelf, the posterior portion of the right
premaxillary sac fossa displays an abrupt anterior
slope. Interestingly, a similar step is present at the
same level in the fully concave left premaxillary sac
A
left premaxillary crest
bulge on fused premaxillae
dorsal right premaxillary sac fossa
filled right premaxillary sac fossa
dorsal infraorbital foramen
right premaxillary crest
posterior excavation of the premaxillary sac fossae
concave left premaxillary fac fossa
sulci
rostral maxillary eminence
maxilla
depression
maxilla
premaxilla
nasal
nasal
B
Fig.7. — Skull of Tusciziphius atlanticus, n.sp. (SGHN MA0632): A, dorsal view; B, corresponding line drawing. Cross-hatching indi-
cates the presence of a concretion; tight parallel lines indicate a break surface. Scale bar: 10cm.
121
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
fossa. No premaxillary foramen is observed at the
anterior end of the premaxillary sac fossae. Only
one foramen is visible near the medial margin of
the left fossa in the holotype, in the paratype, and
SGHN MA0914, absent in SGHN MA0632 and
SGHN MA0644. Similar to T. crispus, the ascend-
F
ig
.8. — Skull of Tusciziphius atlanticus n.sp. (SGHN MA0632): A, lateral view; B, corresponding line drawing (the arrows indicate the level
of the transverse section); C, transverse section through the rostrum; D, detail of C showing the growth layers; E,anterior view; F,corre-
sponding line drawing. Cross-hatching indicates the presence of a concretion; tight parallel lines indicate a break surface. Scale bar: 10cm.
A
right premaxillary crest
right premaxillary crest left premaxillary
crest
growth layer marks
bulge on fused
premaxillae
bulge on fused
premaxillae
premaxilla
premaxilla
premaxilla
phosphorite
concretion
maxilla
foramen
mesorostral tunnel
maxilla
nasals
vestigial alveolar groove
maxilla-palatine suture marks
palatine fragment
anterior margin
of filled right
premaxillary
sac fossa
frontal
sulci
vomer
dorsal infraorbital
foramen
dorsal infraorbital
foramina
left rostal
maxillary
eminence
rostal
maxillary
eminence
filled right premaxillary
sac fossa
B
EF
CD
122 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
ing process of the premaxilla of T. atlanticus n. sp.
exhibits a strong transverse constriction. Its poste-
rior portion is anteriorly curved, overhanging the
premaxillary sac fossa and bony nares. e right
premaxillary crest of T. atlanticus n. sp. shows the
extreme transverse widening typical for Tusciziphius.
Moreover, as in T. crispus, the right premaxillary
crest is more anteriorly projected than the left. For
this character, Tusciziphius clearly differs from the
closely related Globicetus n. gen., in which both
crests have approximately the same anterior extent.
is difference may be related to the different direc-
tion of the right premaxillary crest (anterolateral
in Tusciziphius and more transversal in Globicetus
n. gen.). Finally, as in Globicetus n. gen., Imocetus
n. gen., and especially Caviziphius, the right pre-
maxillary crest is considerably larger and especially
higher than the left, a feature best seen in anterior
view. Moreover, due to the fact that the right pre-
maxillary crest is considerably higher than the left
one, the dorsal surface of the right nasal is more
medially inclined than that of the left nasal.
Maxilla
e distal half part of the rostrum is narrow and
strongly transversally compressed. Consequently
the dorsal surface of the maxilla is nearly vertical
and almost invisible in dorsal view. In the proximal
half portion, the lateral inclination of the maxilla
decreases progressively, with a wider portion visible
in dorsal view.
At the rostrum base, just medial to the right an-
torbital notch, the holotype and SGHN MA0644
develop a high and voluminous rostral maxillary
eminence, slightly medially curved. SGHN MA0632
lacks a portion of the right maxilla that probably
included, judging from the shape and the position
of the broken surface, a similar eminence. On the
left side of the holotype, SGHN MA0644, and
SGHN MA0632, the maxilla exhibits a similar
but lower rostral maxillary eminence. In the para-
type and SGHN MA0914, no prominent rostral
maxillary eminence is present, only some irregular
excrescences. On both sides of skulls bearing ros-
tral maxillary eminences, a shallow longitudinal
depression is margined laterally by the eminence,
slightly overhanging the depression, and medially
by the low medial premaxillary shelf. One to three
dorsal infraobital foramina pierce the maxilla near
the rostrum base, medial and/or posterior to the
rostral maxillary eminence (when the eminence is
present). From these foramina, several sulci run
anteriorly and posteriorly. e vestigial alveolar
groove is a narrow sulcus, with no visible alveoli.
Nasal
e shape of the nasals, as the general architecture
of the vertex, is rather stable in T. atlanticus n. sp.
and T. crispus. e sutures of the nasals are gener-
ally hard to detect due to the extreme ossification
and fusion of the vertex bones. e nasals are
anteroposteriorly elongated, with lateral margins
parallel or faintly convergent (but not as much as
in Globicetus n. gen.). As in Globicetus n. gen. and
Imocetus n. gen., the lateral margin of the nasal is in
contact with the premaxillary crest for all its extent
and the dorsal surface of the joined nasals forms a
shallow depression between the premaxillary crests.
Frontal
e frontals are visible on the vertex of several
specimens. ey are wider than in Globicetus n. gen.
and Imocetus n. gen., related to the lesser transverse
constriction of the posterior part of the vertex.
Vomer
e vomer is not visible dorsally due to the complete
closure of the mesorostral groove. It is visible only
ventrally between the premaxillae and the maxil-
lae along the mid-line of the rostrum, and anterior
to the choanae due to the non-preservation of the
palatine in that area.
Palatine
e palatine is partially preserved only in SGHN
MA0914. e maxilla-palatine suture extends about
150 mm anterior to the antorbital notch, the level
of the abrupt widening of the rostrum.
REMARKS
e previous assignation of the South Carolina
paratype of Tusciziphius atlanticus n. sp. to the Ital-
ian species T. crispus was made at a time when only
one skull was known (Post et al. 2008), preventing
123
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
any evaluation of the intraspecific or interspecific
character for the variation at the level of the pre-
maxillary sac fossae. e observation of the same
filling of the right premaxillary sac fossa in all the
examined skulls from the Atlantic Ocean floor off
the Iberian Peninsula suggests that this character is
valid for the definition of a new species, grouping the
South Carolina specimen with the Iberian specimens.
Additional specimens of T. crispus from Italy could
confirm this interpretation in the future.
Aexcavated left premaxillary sac fossa
filled with phosphorite concretion
bulge on fused premaxillae
bulge on fused
premaxillae
right rostral
maxillary eminence
right rostral
maxillary
eminence
right premaxillary
crest
high step
worn anterior portion
of rostrum
worn anterior portion of rostrum
worn protuberance on fused premaxillae
worn anterior portion of rostrum
worn lateral portion
of right premaxillary crest
right rostral
maxillary eminence
bulge on fused
premaxillae
D
E
B
C
Fig.9. — A-C, skull of Tusciziphius atlanticus, n.sp. (SGHN MA0644); A, dorsal view; B, lateral view; C, anterior view. D, E,skull of
T.atlanticus, n.sp. (SGHN MA0914); D, dorsal view; E, lateral view. Parallel lines indicate supercial wear. Scale bar: 10cm.
124 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
Genus Globicetus n. gen.
TYPE AND ONLY SPECIES. — Globicetus hiberus n. gen.,
n. sp., by present designation.
E
TYMOLOGY
. — From Latin “globus”, for the large spher-
ical medial premaxillary prominence on the rostrum, and
from Latin “cetus”, whale. Gender masculine.
DIAGNOSIS. — Same as for the species.
Globicetus hiberus n. sp.
(Figs 10-13; Table 3)
H
OLOTYPE
. — ML 1361, partial skull including rostrum,
facial area and vertex.
REFERRED SPECIMENS. — MHNUSC 3958, partial skull
including facial area and vertex, 20 miles from Touriñán
Cape, off the Galician coast, depth of 1500 m, 42°50’N,
9°40’W; IEO DR26 026, partial skull including rostrum
and anterior portion of facial area, off the Galician coast,
depth of approximately 1500 m, 42°27’N, 11°59’W.
ETYMOLOGY. — From Latin “hiberus”, Iberian, for the
geographical origin of the holotype and referred specimens.
T
YPE
LOCALITY
. — South of Nazaré Canyon off the
Portuguese coast, exact locality unknown, but likely
around 39°18’N, 9°47’W.
DIAGNOSIS. — Globicetus hiberus n. gen., n. sp. differs
from all other ziphiids in the large spherical medial ros-
tral prominence formed by the fused premaxillae, in the
large prominence of the right premaxilla anterior to the
right premaxillary sac fossa, and in the barely marked
antorbital notch, related to the important widening of
the rostrum base.
Among the other ziphiine genera it further differs from
Ziphius and Izikoziphius in the medial fusion of the pre-
maxillae closing the mesorostral groove; it shares with
Imocetus n. gen. and Tusciziphius the anterior part of the
nasal contacting the premaxillary crest and the extreme
ossification and fusion of the vertex elements, but it differs
from Imocetus n. gen. in lacking a wide facial depression,
rostral maxillary spur-shaped crest, and in having the
premaxillary foramina not located posterior to the level
of the antorbital notch; it differs from Tusciziphius in the
less transversally expanded vertex (lower width between the
premaxillary crests, and lower distance between maxillae
posterior to the vertex), and in the posterolateral direction
of the right premaxillary crest. It further differs from the
possible ziphiine Caviziphius in shallower excavation of
the premaxillary sac fossae.
DESCRIPTION
e skull is slightly smaller than in the largest speci-
mens of Imocetus piscatus n. gen., n. sp. (see below),
with a postorbital width estimated at 372 mm in
the holotype. e rostrum is elongated and its
base is wide. e posterior half of the rostrum is
characterized by the extreme thickening of the
premaxillae, forming a large spherical prominence
followed towards the right premaxillary sac fossa
by a high shelf (see below). e facial area is short
with the low and wide vertex overhanging it, hid-
ing the bony nares and most of the premaxillary
sac fossae in dorsal view.
Premaxilla
On the roughly complete rostrum of the holotype
the premaxilla is slightly longer apically (20 mm)
than the maxilla. On the anterior half of the massive
and subcylindrical rostrum, the mesorostral groove
is dorsally closed by the thick premaxillae, displaying
a medial sutural contact. In dorsal view, the poste-
rior half of the rostrum is partly covered by a large,
roughly spherical, element made by the joined pre-
maxillae. Preserved on the holotype and IEO DR26
026, this unusual prominence has a maximum width
of 150 mm and 141 mm respectively in these two
specimens and a maximum height above the maxilla
of 135 mm and 92 mm. It is slightly asymmetrical,
higher on the right side of the holotype and longer
on the right side of IEO DR26 026. It is made, at
least superficially, by compact bone, covered with
narrow and shallow anastomosed sulci likely related
to vascularization. On the holotype, the anterior and
anterolateral margins of the spheroid do not contact
the underlying premaxilla and maxilla, leaving an
open space of about 10 mm, whereas in IEO DR26
026 a high medial pad of bone joins the spheroid
to the dorsal surface of the rostrum. Posteriorly, the
spheroid is followed by a thick shelf predominantly
constituted by the right premaxilla, whose surface
is similarly compact and covered with sulci. Rec-
tangular in dorsal view in the holotype, this shelf is
distinctly narrower distally in IEO DR26 026 and
in the anteriorly incomplete skull MHNUSC 3958.
In lateral view the dorsal margin of the shelf of the
latter raises forwards similarly to the more complete
specimens, suggesting the presence of a similar large
125
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
spheroid in MHNUSC 3958. e abrupt posterior
margin of the shelf corresponds to the anterior limit
of the right premaxillary sac fossa, which is consid-
erably wider than the left fossa. Only a narrow and
much lower longitudinal crest is located anterior to
the left premaxillary sac fossa of the holotype and
MHNUSC 3958. Both fossae are short anteriorly,
more than in Tusciziphius, nearly completely dorsally
overhung by the vertex. e presence of the massive
shelf in Globicetus n. gen., associated with shorter
premaxillary sac fossae, might be interpreted as an
overgrowth of the thickened anterior portion of the
right premaxillary sac fossa observed in T. atlanticus
n. sp. e surface of the premaxillary sac fossae is
strongly concave and no premaxillary foramen could
be detected on the bottom of any of the fossae. Only
one foramen is observed on the medial margin of
the left fossa of the three specimens, similar to the
condition in Choneziphius leidyi n. sp., Imocetus
n. gen., and T. atlanticus n. sp.
A
B
left premaxillary
crest additional foramen
left premaxillary sac fossa
dorsal infraorbital foramina
spherical premaxillary prominence
maxilla
premaxilla
maxilla
dorsal infraorbital foramen
right premaxillary sac fossa
frontal
antiorbital notch
sulci
premaxilary shelf
medial suture between premaxillae
nasal
right
premaxillary crest
F
ig
.10. — Skull of Globicetus hiberus n.gen., n.sp. (ML 1361, holotype): A, dorsal view; B, corresponding line drawing. Scale bar: 10cm.
126 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
e ascending process of each premaxilla is strongly
constricted and short. On the low vertex, the over-
hanging right premaxillary crest is much wider
than the left, reaching laterally a level beyond the
lateral margin of the corresponding premaxillary
sac fossa. e top of the vertex is made by the right
premaxilla, much higher than the left. e anterior
margin of the right premaxillary crests is postero-
laterally directed, whereas the anterior margin of
the left premaxillary crest is roughly anterolaterally
directed. In Tusciziphius both crests are usually
anterolaterally directed.
Maxilla
Apically invisible in dorsal view, the maxilla only
slightly widens along the distal half of the rostrum,
with a somewhat medially convex maxilla-premax-
illa suture. From the level of the large spheroid
it sends a thin lateral plate whose lateral margin
reaches the preorbital process in a nearly rectilin-
ear line, forming a wide rostrum base. Differing
from T. atlanticus n. sp. and Imocetus n. gen., the
antorbital notch is therefore barely individualized;
a wide subhorizontal surface margins the premaxil-
lary shelf on both sides of the rostrum base. is
surface is pierced by several dorsal infraorbital
foramina, three on the right side and two on the
left side of the holotype, one less on the right side
of MHNUSC 3958, and one less on each side
of IEO DR26 026. From these foramina, several
sulci are sent anteriorly and anterolaterally. A large
longitudinal sulcus passes between the spheroid
of the premaxillae and the maxilla, exiting on the
anterior margin of the spheroid.
On the supraorbital area, the maxilla is thin,
lacking any maxillary crest contrary to part of
the specimens of T. atlanticus n. sp. and Imocetus
n. gen. Posterior to the nasals on the vertex, left and
right maxillae are close to each other, more than
in Tusciziphius, with a minimal distance between
the maxillae lower than the width of the nasals. At
this level the medial margin of the right maxilla is
Table 3. — Measurements (in mm) on the skulls of Globicetus hiberus n.gen., n.sp. from the Atlantic Ocean oor off the Iberian Pen-
insula. Abbreviations: e, estimate; +, nearly complete; –, no data.
ML 1361
(holotype) MLI 3958 IEO DR 26026
Rostrum length 530 –552
Length of premaxillary portion of rostrum 20 – –
Distance from apex of rostrum to bony nares 665 –e650
Width of rostrum at mid-length 99 –93
Height of rostrum at mid-length 98 –99
Width of rostrum base at antorbital notch 265 e285 e233
Width of premaxillae at rostrum base 104 – –
Minimum distance between maxillae near rostrum base –80 –
Distance rostrum base – anterior apex of palatine 227 –315
Preorbital width of skull –+335 –
Postorbital width of skull e372 –+305
Longitudinal distance right premaxillary foramen – rostrum base 78 –72
Width of premaxillary sac fossae 170 +165 –
Width of right premaxillary sac fossa 100.5 83 –
Width of left premaxillary sac fossa 62.5 e74 –
Width of bony nares 79 82 –
Minimum width of right ascending process of the premaxilla 44 42 –
Minimum width of left ascending process of the premaxilla 23 e17 –
Width of premaxillary crests 185 170 –
Width of right premaxillary crest 106 84 –
Width of left premaxillary crest 70 51 –
Maximum width of nasals 69.5 68 –
Maximum width of right nasal –47 –
Maximum width of left nasal –28 –
Length of medial suture of nasals 68 e62 –
127
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
more erected than the margin of the left maxilla;
the latter is the continuation of the depressed dorsal
surface of the nasals.
On the ventrolateral surface of the rostrum, the
remnant of alveolar groove does not contain indi-
vidualized alveoli.
A
maxilla
frontal
foramen
dorsal infraorbital
foramen
spherical premaxillary
prominence
lacrimal
lacrimal
vomer
maxilla
nasals
frontal
palatine-maxilla suture
vestigial alveolar groove
maxilla
maxilla
maxilla
right premaxillary crest
right premaxillary crest
left premaxillary
crest
spherical premaxillary prominence
premaxilla
premaxilla
premaxillary shelf
B
CD
Fig.11. — Skull of Globicetus hiberus n.gen., n.sp. (ML 1361, holotype): A, lateral view; B, corresponding line drawing; C, antero-
dorsal view; D, corresponding line drawing. Scale bar: 10cm.
128 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
A
C
choana
frontal
lacrimal
antorbital notch
vomer
vomer
palatine-maxilla suture
vestigial alveolar groove
premaxilla
maxilla
spherical premaxillary prominence
premaxillary shelf
cylindrical anterior part of rostrum
with fused premaxillae
frontal
bony
nares
nasals
left premaxillary
crest
right premaxillary
crest
ascending process of
right premaxilla
ventral infraorbital
foramen
B
Fig.12. — Skull of Globicetus hiberus n.gen., n.sp. (ML 1361, holotype): A, ventral view; B, detail of the facial area in anterior view;
C, anterolateral view; D, corresponding line drawing. Scale bar: 10cm.
129
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
AB
D
C
F
E
left premaxillary
crest left premaxillary
sac fossa concave ascending
process of premaxilla
premaxillary shelf
left premaxillary crest
right premaxillary
sac fossa
right premaxillary
crest
ascending process of
right premaxilla
medial pad of
premaxillae
right premaxillary crest
nasal
maxilla
premaxilla
premaxillary shelf
medial pad of premaxillae
spherical premaxillary
prominence
concave left and
right premaxillary
sac fossae
premaxillary shelf
additional foramen
Fig.13. — A-C, partial skull of Globicetus hiberus n.gen., n.sp. (MHNUSC 3958): A, dorsal view; B, lateral view; C, detail of the
vertex and premaxillary sac fossae in anterior view; D-F partial skull of G. hiberus n.gen., n.sp. (IEO DR26 026); D, dorsal view;
E,lateral view; F, detail of the spherical premaxillary prominence in anterior view. Parallel lines indicate a break surface. Scale
bar: 10cm.
130 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
Nasal
Excluded from the premaxillary crest, the nasal is
considerably narrower anteriorly than posteriorly.
e dorsal surface of the joined nasals forms a
depression between the premaxillary crests, as in
Imocetus n. gen. and Tusciziphius. e anterior tip
of the nasals does not reach a level anterior to the
premaxillary crests, differing from Izikoziphius and
Ziphius. e medial suture is distinctly shifted to the
left side compared to the sagittal plane of the skull.
Frontal
Only parts of the frontal are preserved on the heav-
ily worn supraorbital area. Frontals are lost on the
vertex; they apparently originally formed an an-
teromedial projection between nasals.
Vomer
Hidden dorsally by the development of the pre-
maxillae, the vomer is only visible ventrally be-
tween the premaxillae and between the maxillae.
At the rostrum base, its ventral exposure between
the maxillae of the holotype is likely due to partial
wear of the latter and loss of the palatines.
Palatine
Most parts of the palatine are likely lost in the three
specimens. A large depression with a distinct outline
marks the original anterior extent of the palatine on
the rostrum, far anterior from the antorbital notch.
e palatine was longer in IEO DR26 026, reach-
ing a level 315 mm anterior to the notch.
Lacrimal
e lacrimal is only partly preserved ventral to the
maxilla on the preorbital process of the holotype.
Its ventral exposure appears rather narrow but its
original extension cannot be estimated due to the
bad preservation of the ventral surface of the skull.
REMARKS
A series of arguments support the interpretation of the
conspicuous spheroid at the rostrum base of Globicetus
hiberus n. gen., n. sp. as a non-pathological element.
First, it is present with a very similar outer shape
in two of the described specimens. it has also been
observed by us in additional undescribed skulls kept
in private collections, and its presence is very likely
in the third described specimen. In addition, from a
morphological point of view, it is nearly symmetri-
cal, with smooth surfaces contrasting with different
kinds of pathological bone growths. Furthermore,
canals for vascularization/innervation at the base of
the spheroid are not interrupted. Finally no fracture
or pathological bone tissue has been detected ventral
to the spheroid in any of the described specimens.
From a systematic point of view, a dorsomedial
sutural contact between the premaxillae is observed
on the rostrum of part of the ziphiines (Choneziphius,
Imocetus n. gen., and Tusciziphius) and members of
the “Messapicetus clade”. e general morphology
of the low and wide, trapezoidal, extremely ossified,
and strongly asymmetric vertex, overhanging the
bony nares and the premaxillary sac fossae, closely
resembles Tusciziphius and, in a lesser extent, Imoce-
tus n. gen. A superficially similar morphology is also
observed in the hyperoodontine Hyperoodon, but in
this case the nasal is deeply thrusted in the premaxil-
lary crest and the left premaxillary crest is distinctly
directed posterolaterally. e development of a high
medial prominence of the joined premaxillae on the
rostrum is similarly observed in some specimens of
Tusciziphius atlanticus n. sp. and in a fragmentary
skull referred here to aff. Caviziphius sp. (see below).
In none of these specimens the prominence displays
a spherical volume shape. Additional differences
with Tusciziphius are: large prominence of the right
premaxilla anterior to the right premaxillary sac
fossa; barely marked antorbital notch, related to the
important widening of the rostrum base; less trans-
versally expanded vertex (lower width between the
premaxillary crests and lower distance between the
maxillae posterior to the vertex); and posterolateral
direction of the right premaxillary crest.
Genus Imocetus n. gen.
TYPE AND ONLY SPECIES. — Imocetus piscatus n. sp., by
present designation.
ETYMOLOGY. — From Latin “imum”, bottom, because
it was trawled on the sea floor, and from Latin “cetus”,
whale. Gender masculine.
DIAGNOSIS. — Same as for the species.
131
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
Imocetus piscatus n. sp.
(Figs 14-17; Table 4)
H
OLOTYPE
. — ML 1358, partial skull including rostrum,
facial area and vertex.
REFERRED SPECIMENS. — ML 1359, partial skull in-
cluding rostrum and anterior portion of facial area and
ML 1360, partial skull including rostrum and anterior
portion of facial area, south of Nazaré Canyon off the
Portuguese coast, exact locality unknown, but likely
around 39°18’N, 9°47’W.
ETYMOLOGY. — From Latin “piscatus”, fished, because
the holotype and the referred specimens were collected
at sea by fishermen.
T
YPE
LOCALITY
. — South of Nazaré Canyon off the
Portuguese coast, exact locality unknown, but likely
around 39°18’N, 9°47’W.
DIAGNOSIS. — Imocetus piscatus n. gen., n. sp. differs
from all other ziphiids in a wide and anteroposteriorly
elongated facial depression laterally margined by acute
longitudinal maxillary crests, a rostral maxillary crest
forming a posterodorsally directed spur, and premaxil-
lary foramen located very posterior to the level of the
antorbital notch. Among other ziphiines it further
differs from Izikoziphius and Ziphius in the medial fu-
sion of the premaxillae closing the mesorostral groove;
it shares with Globicetus n. gen. and Tusciziphius the
anterior part of the nasal contacting the premaxillary
crest, and the extreme ossification and fusion of the
vertex elements, but it differs from Globicetus n. gen.
in lacking a large spherical medial premaxillary rostral
prominence and in the presence of a distinct antorbital
notch. It differs from Tusciziphius in the less transversally
expanded vertex (lower width between the premaxillary
crests and lower distance between the maxillae poste-
rior to the vertex), and in the posterolateral direction
of the right premaxillary crest. It further differs from
the possible ziphiine Caviziphius in the less excavated
premaxillary sac fossae.
DESCRIPTION
e skull is large for a fossil ziphiid; based on the
width of the rostrum it is close to the size of the
modern Mesoplodon layardii. At the apex, the tapered
rostrum is cylindrical, whereas it is wider at its base.
e anteroposterior length of the rostrum is less
than half the estimated condylobasal length. e
rostrum is proportionally longer in the holotype,
in which the antorbital notch is more posteriorly
located than in ML 1359 and ML 1360. e ros-
Table 4. — Measurements (in mm) on the skulls of Imocetus piscatus n.gen., n.sp. from the Atlantic Ocean oor off the Iberian Pen-
insula. Abbreviations: e, estimate; +, nearly complete; –, no data.
ML 1358
(holotype) ML 1359 ML 1360
Rostrum length as preserved 495 382 326
Distance from apex of rostrum to bony nares –+392 –
Width of rostrum 100 mm from apex 68 +58 –
Width of premaxillae 100 mm from apex –46 –
Width of rostrum 200 mm from apex 99 101 –
Width of premaxillae 200 mm from apex –51 –
Width of rostrum 300 mm from apex 149 145 –
Width of premaxillae 300 mm from apex –52 –
Width of rostrum 400 mm from apex 212 – –
Width of rostrum base at antorbital notch 256 199 165
Width of premaxillae at antorbital notch –63 –
Distance rostrum base – apex of palatine-pterygoid suture 158 85 110
Maximum distance between lateral margins of rostral maxillary crests 198 205 162
Minimum distance between maxillary crests (on neurocranium) 186 – –
Distance rostrum base – anterior end of premaxillary sac fossa 211 367 –
Width of right premaxillary sac fossa e64 – –
Width of left premaxillary sac fossa e60 – –
Width of bony nares 76 – –
Minimum width of right ascending process of the premaxilla 29 – –
Width of premaxillary crests +139 – –
Maximum width of nasals 70 – –
Minimum posterior distance between maxillae 51 – –
132 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
trum base is also considerably wider in the holotype.
e facial area, between the antorbital notch and
the vertex, is very long. e vertex is proportionally
low, overhanging the bony nares. e supraorbital
area is only partly preserved and the basicranium
is completely missing.
Premaxilla
On the anterior portion of the rostrum, the half-
cylindrical robust premaxillae display a sutural
contact above the mesorostral groove. eir me-
dial suture is sinuous on the holotype and more
rectilinear in ML 1359 (not preserved in ML
1360). 195 mm posterior to the apex of the ros-
trum of the holotype, an artificial medial opening
between the superficial layers of the premaxillae
exposes bone that we interpret as deeper regions
of the premaxillae. Alternatively this element
could correspond to the thickened vomer, but
we prefer the first hypothesis taking into account
the anteriorly open mesorostral groove. Consid-
ering the preserved parts, the premaxillae seem
considerably thicker on the rostrum of ML 1359
than in the holotype, and even more than in ML
A
B
bony nares additional foramen
frontal
nasal
maxilla
dorsal infraorbital
foramen
right premaxillary
crest right premaxillary sac fossa
platform on maxilla
maxillary crest frontal
antorbital notch
rostral maxillary crest
right premaxilla
left premaxilla
maxilla
dorsal infraorbital foramina
premaxillary
foramen ?
broken surface
left premaxillary
sac fossa
left premaxillary
crest
Fig.14. — Skull of Imocetus piscatus n.gen., n.sp. (ML 1358, holotype): A, dorsal view; B, corresponding line drawing. Parallel lines
indicate a break surface. Scale bar: 10cm.
133
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
1360. From the level of the antorbital notch, the
premaxillae descend in a depression between the
more prominent maxillae, until the premaxillary
sac fossae. is depression seems homologous
to the prenarial basin described in Beneziphius,
Messapicetus, and Ziphirostrum (Lambert 2005;
Bianucci et al. 2010). e premaxillary sac fos-
sae, only well preserved on the holotype, are even
more depressed compared to the maxillae, with
a distinctly concave surface. e posteromedial
portion of the right fossa is slightly elevated.
e left fossa is lower than the right, and nar-
rower. However, the asymmetry at this level is
less pronounced than in Caviziphius, Globicetus
Fig.15. — Skull of Imocetus piscatus n.gen., n.sp. (ML 1358, holotype): A, lateral view; B, corresponding line drawing; C, ventral
view. Scale bar: 10cm.
A
B
maxilla
maxilla
maxillary crest
rostral maxillary crest
premaxillary
maxilla
vestigial alveolar groove
palatine-maxilla suture
palatine
frontal
frontal
choana
antorbital notch
vestigial alveolar groove
pterygoid-palatine sutural surface
vomer
ventral infraorbital
foramen
vomer
right premaxillary
crest
C
134 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
n. gen., and Tusciziphius. If present (area partly
covered with phosphorite concretions), right and
left premaxillary foramina are close to each other
and much more distant from the level of the an-
torbital notch than in any other known ziphiid,
even more than in Hyperoodon. is condition
stresses the unusual elongation of the facial area
between antorbital notches and bony nares in
Imocetus n. gen. An additional foramen is present
on the medial margin of the left premaxillary sac
fossa of the holotype and ML 1360, in the same
position as in Globicetus n. gen., Choneziphius
leidyi n. sp., and Tusciziphius atlanticus n. sp.
e ascending process of the premaxilla is short
and erected. Its upper part overhangs the corre-
sponding premaxillary sac fossa. On the vertex,
the premaxillary crests are highly asymmetric:
the right crest is more voluminous, distinctly
higher and more anteriorly bulging. Similar to
Globicetus n. gen., the right premaxillary crest is
posterolaterally directed whereas the left crest is
anterolarally directed.
Maxilla
In dorsal view the maxilla is visible along the
premaxilla on the whole length of the rostrum.
On the second third of the rostrum, the lateral
margin is acute. is margin only thickens con-
siderably before the antorbital notch, in relation
with the development of the rostral maxillary crest
located medial to the notch. Compared to the
holotype, this spur-like posterodorsally projecting
AB
right premaxillary crest
nasals dorsal depression
left premaxillary crest
maxillary crest
maxilla
premaxilla
frontal
dorsal infraorbital
foramina
right maxillary crest
right rostral
maxillary crest
large medial basin
between the maxillary crests
left maxillary crest
left rostral maxillary crest
antorbital notch
rostral maxillary
crest
prenarial basin
platform on
maxilla
C
Fig.16. — Skull of Imocetus piscatus n.gen., n.sp. (ML 1358, holotype): A, anterior view; B, corresponding line drawing; C, antero-
lateral view. Parallel lines indicate a break surface. Scale bar: 10cm.
135
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
crest is more laterally directed, larger, and more
posteriorly located compared to the antorbital
notch in ML 1359 and ML 1360. e crests are
asymmetric in the holotype and ML 1359; in the
former the left crest is slightly higher, whereas in
the latter the incomplete right crest was originally
more robust than the left crest. e antorbital
notch is deep and narrow in the three specimens,
medial to the robust and long preorbital process.
e notch is more slit-like in the holotype and
V-shaped in the two other skulls. From the bot-
tom of the notch, the maxilla forms an acute
longitudinal crest on the supraorbital area of
the skull. is crest is probably not homologous
to the huge rostral crest observed medial to the
antorbital notch in adults of Hyperoodon spp.
(see Mead & Fordyce 2009 for terminology).
is maxillary crest is rectilinear and posteriorly
diverging in ML 1359 and ML 1360, whereas
it is medially convex in dorsal view and slightly
overhangs the medial area of the maxilla in the
holotype. Right and left crests limit a vast and
depressed facial area, where dorsal infraorbital
foramina are present along the prenarial basin
(two on each side of the holotype, one on each
side of ML 1359 and ML 1360). In addition to
these large foramina, the surface of the maxilla
is covered with shallow and narrow anastomo-
sed sulci and tiny foramina. Between the dorsal
infraorbital foramina and the premaxillary sac
fossa, the maxilla is distinctly thickened, form-
ing a platform with a convex surface, much wider
on the right side (and also more elevated on the
right side of the holotype and ML 1359). e
lateral flank of the maxillary crest is a wide and
slightly concave surface with a steep slope. Due to
the non-preserved lateral part of the supraorbital
process, in lateral view the crest of the holotype
displays a high triangular section.
No alveoli could be detected on the heavily post-
mortem worn alveolar groove. In relation with the
anterior shift of the preorbital process and antorbital
notch compared to other ziphiids, the position of
the ventral infraorbital foramen is strongly modi-
fied. is foramen is about 200 mm posterior to
the antorbital notch in the three specimens, whereas
A
premaxillary
sac fossae
frontal
rostral maxillary crest
maxilla
premaxilla
antorbital notch
rostral maxillary crest
maxilla premaxilla
antorbital notch
frontal
maxillary crest
additional foramen
premaxillary
sac fossae
maxillary crest
prenarial basin
platform
on maxilla
B
Fig.17. — A, skull of Imocetus piscatus n.gen., n.sp. (ML 1359), dorsal view; B, skull of I. piscatus n.gen., n.sp. (ML 1360), dorsal
view. Parallel lines indicate a break surface. Scale bar: 10cm.
136 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
it is a short distance from the notch in other ziphi-
ids. e foramen is followed anteriorly by a deep
and long groove, edging the pterygoid sinus fossa
laterally until the antorbital notch. e location
of the ventral infraorbital foramen also gives a
clue about the level of the non-preserved orbit.
Indeed, in other ziphiids and other odontocetes
the frontal groove and optic canal are posterior to
the infraorbital foramen.
Nasal
e nasals are wide anteriorly. Each of them oc-
cupies a considerable portion of the corresponding
premaxillary crest and the nasal-premaxilla suture
reaches the anterior surface of the crest, a condition
more similar to Hyperoodon. e part of the nasal
thrusted in the premaxillary crest is more compact
than the smoothly depressed medial area. e dif-
ference of compactness only partly explains this
internasal fossa, extending posteriorly on the left
frontal and maxilla. e premaxilla-nasal suture being
hard to distinguish in this very ossified vertex, an
alternative interpretation would be a more medial
suture more similar to the condition in Globicetus
n. gen. and Tusciziphius (hatched line in Fig. 14B).
Frontal
Only fragments of the frontals are preserved posterior
to the nasals on the vertex. e frontal sends a short
anterior projection between the nasals. In lateral
view, the frontal is the main element of the robust
anterior part of the preorbital process, covered by a
thin sheet of maxilla. More posteriorly the bone is
incomplete laterally; the whole orbit area is lost and
the extent of the temporal fossa cannot be assessed.
Vomer
Our interpretation of the rostral bones suggests that the
vomer does not fill the mesorostral groove of Imocetus
n. gen., dorsally covered by the joined premaxillae.
Palatine
e palatine is preserved on its rostral portion in
the three specimens. e palatine-maxilla suture is
visible at some levels, but the best seen structure is
the surface of suture with the lost pterygoid, marked
by grooves, and limiting a large depression corre-
sponding to the vast anterior part of the pterygoid
sinus fossa. e palatine-pterygoid suture extends
far anterior to the antorbital notch in the three
specimens, even more in the holotype, which is
characterized by a more posteriorly located notch.
Lacrimal
Fragments of the lacrimal are preserved on the
holotype, but its original outline cannot be pre
-
cisely drawn.
REMARKS
e significant morphological variations between
the holotype and the more fragmentary referred
specimens is interpreted here as due to intraspecific
variation (possibly related to sexual dimorphism, see
below); pending the discovery of new more com-
plete specimens, we choose to maintain ML 1359
and ML 1360 in the same species as the holotype.
Despite similarities of Imocetus n. gen. with some
hyperoodontines (nasal probably included in the pre-
maxillary crest and reaching the anteromedial margin
of the crest; excavation of the dorsomedial surface
of the nasal; right premaxillary crest posterolater-
ally directed) and more specifically with Hyperoodon
(low vertex overhanging the premaxillary sac fossae;
premaxillary foramen posterior to the level of the
antorbital notch), this new genus is interpreted here
as a member of the subfamily Ziphiinae. Indeed,
it shares with all ziphiines, except Izikoziphius and
Ziphius, the medial fusion of the thickened premax-
illae dorsally closing the mesorostral groove, with
Globicetus n. gen. and Tusciziphius the extreme ossi-
fication and fusion of the vertex elements, and with
Choneziphius, Globicetus n. gen., and T. atlanticus
n. sp. the deep premaxillary sac fossae.
Family ZIPHIIDAE
Incertae sedis
Genus Caviziphius Bianucci & Post, 2005
TYPE AND ONLY SPECIES. — Caviziphius altirostris Bia-
nucci & Post, 2005, known from a single specimen from
Steendorp (Belgium), probably late Miocene (Bianucci &
Post 2005).
137
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
aff. Caviziphius sp.
(Fig. 18)
REFERRED SPECIMEN. — SGHN MA0920, partial skull
including rostrum and right part of facial area, Cortada
fishing ground, off the Galician coast, depth of approxi-
mately 400-800 m, 43°36’N, 9°0’W.
DESCRIPTION
e original shape of this badly preserved frag-
mentary skull might have been partially modified
by an intense wear. e sutures between the bones
are also almost completely obliterated. Nevertheless
some typical features of the premaxilla and maxilla
(medial rostral premaxillary bulge, strongly asym-
metric premaxillary sac fossae, and prominent right
rostral maxillary eminence) are conspicuous and
allow a relevant comparison with other ziphiids.
Premaxilla
In dorsal view, the premaxillae are medially su-
tured for all their rostral length except for the
apical 55 mm portion, where the premaxillae
abruptly diverge and leave the mesorostral groove
dorsally open. From the anteriormost point of
their junction, the height of the premaxillae
increases progressively, reaching an elevation of
78 mm above the maxilla at 186 mm from the
anterior margin of the right premaxillary sac
fossa, forming a protuberant bulge. Posteriorly,
the height of the premaxillae decreases abruptly,
generating a clear step on the dorsal outline of
the rostrum seen in lateral view. On the whole,
this premaxillary bulge is similar to the bulge
observed in some skulls of Tusciziphius atlanticus
n. sp. (holotype, SGHN MA0632, and SGHN
MA0644) even if its posterior margin is distinctly
more anterior. In fact, the distance from the
posterior margin of the bulge and the anterior
margin of the premaxillary sac fossa is 186 mm
in SGHN MA0920, whereas it varies from 81 to
128 mm in T. atlanticus n. sp. A similar anterior
premaxillary bulge is present in a partial skull
A
longitudinal depressions
maxilla
bulge on fused premaxillae
rostral maxillary eminence
ascending process
of right premaxilla
maxilla
concave left
and right
premaxillary
sac fossae
B
Fig.18. — Skull of aff. Caviziphiussp. (SGHN MA0920): A, dorsal view; B, lateral view. Parallel lines indicate a break surface. Scale
bar: 10cm.
138 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
from the Neogene of Antwerp, Belgium (NMB
002), referred by Lambert (2005: fig. 27A-C)
to Ziphiidae aff. Eboroziphius.
Between the bulge and the premaxillary sac
fossae, the premaxillae form a shallow medial
elevation laterally delimited by two longitudi-
nal depressions. A similar architecture is pre-
sent in T. atlanticus n. sp. (even if this area is
anteroposteriorly shorter), NMB 002, Eboro-
ziphius coelops Leidy, 1876, and Caviziphius
altirostris. e incompletely preserved premax-
illary sac fossae are strongly asymmetric (ratio
between maximum width of left and right fossae
approximately 0.5) and deeply excavated, even
more than in T. crispus. e fossae are separated
by a narrow septum corresponding to the medial
overlap of the premaxillae. For these characters,
the premaxillary sac fossae are similar to those
of NMB 002, Caviziphius, E. coelops, and Pely-
corhamphus pertortus Cope, 1895 (see Lambert
2005: fig. 28). It is important to outline that
the holotypes and only referred specimens of
E. coelops and P. pertortus are fragmentary and
considerably worn; consequently we restrict
the genera to their type-species and the type-
species to their holotypes, and consider them
as incertae sedis.
Only the incomplete ascending portion of
the right premaxilla is preserved. It abruptly
rises from the level of the premaxillary sac fossa;
consequently, in lateral view, the outline of the
anterior margin of the vertex displays a deep
concavity, followed ventrally by a semicircular
deep excavation corresponding to the premaxil-
lary sac fossa. A similar lateral profile is observed
in Caviziphius.
Although no premaxillary crest is preserved,
the thin broken surface at the posterior end of
the right ascending process suggests that the
right crest was not massive and wide as seen in
Globicetus n. gen. and Tusciziphius. Nevertheless,
the right premaxillary crest of Caviziphius, even
if not completely preserved, seems to have been
more robust than in SGHN MA0920, judging
from the wider break surface on the ascending
process of the right premaxilla of the holotype
and only referred specimen.
Maxilla
Due to the incompleteness and the strong ero-
sion, the maxilla does not show any significant
features, with the exception of a prominent right
rostral maxillary eminence. Roughly located at
the rostrum base, this semicircular crest is tilted
medially. A similar crest is present in several skulls
of T. atlanticus n. sp. and in NMB 002.
REMARKS
SGHN MA0920 shares with some skulls of Tus-
ciziphius atlanticus n. sp. and the fragmentary
skull NMB 002 the medial bulge on the fused
premaxillae, the strongly asymmetric premaxillary
sac fossae, and the prominent right rostral maxil-
lary eminence. Considering the deep excavation
of both premaxillary sac fossae, the anterior loca-
tion of the premaxillary bulge, and the diverging
premaxillae near the anterior end of the bulge,
SGHN MA0920 is more similar to NMB 002 than
to T. atlanticus n. sp. SGHN MA0920 also shares
with Caviziphius the deeply excavated asymmetric
premaxillary sac fossae, the longitudinal depressions
that laterally margin the shallow medial elevation
of the sutured premaxillae at the rostrum base,
and the abrupt elevation of the ascending process
of the right premaxilla. Unfortunately the ante-
rior part of the rostrum and the antorbital area
of the maxillae are not preserved in the holotype
and only referred specimen of Caviziphius alti-
rostris (see Bianucci & Post 2005); consequently
it is not possible to establish if the premaxillary
bulge and the prominent right rostral maxillary
eminence are also present in the latter. Neverthe-
less, considering that the holotype of C. altirostris
and NMB 002 are nearly identical for the parts
preserved in both specimens and show similar
dimensions, it is likely that both these incomplete
skulls belong to the same species. Furthermore
they were collected in the same area (Antwerp).
If this hypothesis is confirmed with future dis-
coveries, C. altirostris will be redefined with the
combination of the characters of the holotype
and NMB 002. e only significant differences
between SGHN MA0920 and these two skulls
from Antwerp are the smaller size and probably
the thinner right premaxillary crest.
139
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
“Messapicetus clade”
Genus Ziphirostrum du Bus, 1868
T
YPE
SPECIES
. — Ziphirostrum marginatum du Bus, 1868,
from Antwerp (Belgium), late Miocene (Lambert 2005).
O
THER
SPECIES
INCLUDED
. — Ziphirostrum recurvus (du
Bus, 1968) and Z. turniense du Bus, 1868.
aff. Ziphirostrum sp.
(Fig. 19; Table 5)
REFERRED SPECIMEN. — SGHN MA0936, partial skull
including rostrum and left part of facial area, A Selva
fishing ground, depth of approximately 500 m, off the
Galician coast, 44°10’N, 08°40’W.
DESCRIPTION
e rostrum of this medium-size ziphiid is narrow
and elongated, with size and proportions close to
Ziphirostrum turniense (see Lambert 2005).
Premaxilla
e premaxillae are distinctly swollen on the rostrum,
contacting each other dorsomedially above the hollow
mesorostral groove for most of the rostral length. e
nearly fused medial suture is asymmetric, shifted to
the right side on the posterior half of the rostrum. A
similar asymmetry has been noted, but on the other
side, on an isolated ziphiid rostrum from the Mio-
cene of Maryland, USA (Lambert et al. 2010). In
lateral view, the maximum height and width of the
premaxilla is more anterior than in Z. marginatum,
more similar to Z. turniense. From mid-length of the
rostrum, the premaxilla narrows considerably, and a
medial separation appears 110 mm anterior to the level
of the antorbital notch, with a progressive descent of
the premaxilla in the prenarial basin, a feature absent
in Choneziphius. e extent and depth of the basin
is again more similar to Z. turniense, shallower than
in Z. marginatum. Less anteriorly located than in the
latter, the premaxillary foramen is on the floor of the
basin, slightly anterior to the prominental notch. e
partly preserved surface of the left premaxillary sac
fossa is transversly convex, as in Ziphirostrum, differing
from the concave surface in Choneziphius and related
taxa. e ascent towards the vertex is not abrupt.
Maxilla
Even if the anterior part of the maxilla-premaxilla
suture is difficult to detect, the anterior end of the
maxilla is located 50-60 mm from the apex of the
rostrum. Barely visible in dorsal view for the anterior
half of the rostrum, the maxilla considerably widens
towards the prominental notch, forming an elongated
triangular surface. e posterior part of this surface,
along the prenarial basin, displays a steep slope, with
an elevated and thin lateral margin, more similar to
Z. turniense. Considerably wider on the right side
than on the left, the triangular surface is covered
with numerous and high excrescences, a character
found in Choneziphius, but also in Beneziphius and
an isolated ziphiid rostrum from the Neogene of the
North Sea (see Lambert 2005: fig. 26). e promi-
nental notch and maxillary tubercle are conspicu-
ous; this area is not well preserved in any specimen
of Z. turniense. No marked maxillary crest extends
posteriorly from the maxillary tubercle, differing
from Choneziphius.
e alveolar groove is vestigial, with shallow re-
mains of alveoli still visible, a condition observed in
several specimens of Z. marginatum and Z. turniense.
Palatine
e palatine is preserved at the rostrum base, with
a maxilla-palatine suture easy to distinguish. e
rounded anterior end of the palatine is 130 mm
anterior to the antorbital notch. An abrupt step in
the surface of the palatine indicates the suture with
the lost pterygoid.
REMARKS
Except for the development of excrescences on
the dorsal surface of the maxilla on the posterior
half of the rostrum and the distinct asymmetry of
the premaxillae on the rostrum, this specimen is
similar to the two specimens from the Neogene of
the North Sea referred to Ziphirostrum turniense.
e low diagnostic value of the excrescences on the
maxilla has previously been demonstrated (Lambert
2005) and the development of the premaxillae on
the rostrum is known to vary within one species.
e main features differentiating Z. turniense from
the better-known Z. marginatum are observed here:
maximum width and height of the premaxillae at
140 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
mid-length of the rostrum, shallower prenarial
basin, with dorsal exposure of the maxillae wider
and more steeply sloping along the basin. e third
species of the genus, Z. recurvus, is characterized
by a more elevated rostrum with a complete filling
of the mesorostral groove by the vomer, a feature
lacking here. Because this specimen originates from
a remote area, and because the vertex is lacking, as
in specimens of Z. turniense and Z. recurvus, we
prefer to maintain the attribution Ziphiidae aff.
Ziphirostrum sp., pending the discovery of more
complete specimens.
PHYLOGENY
To explore the phylogenetic relationships of the new
ziphiids described here (Choneziphius leidyi n. sp.,
Imocetus piscatus n. gen., n. sp., Globicetus hiberus
n. gen., n. sp., and Tusciziphius atlanticus n. sp.), we
included the new taxa in the matrix of 29 morpho-
logical characters published by Bianucci et al. (2010)
and undertook a similar cladistic analysis, using the
same outgroups (the squalodontid Squalodon and
the eurhinodelphinid Eurhinodelphis). e only
change in the matrix is the addition of a new state
(3) for the unordered character 10 (premaxillary
crest direction): left crest anterolaterally directed
and right crest posterolaterally directed, a condition
observed in Globicetus n. gen. and Imocetus n. gen.
e coding of characters for the Iberian new taxa
is given in the Appendix 1.
e cladistic analysis was achieved with the software
PAUP (version 4.0b10; Swofford 2001), using the
Branch-and-bound algorithm with the homoplastic
characters down-weighted using the default value of
3 for the constant k of the Goloboff method (1993).
e analysis produced 875 equally parsimonious trees,
with tree length 94, Goloboff fit-20.98, Consistency
Index (CI) 0.53 and Retention Index (RI) 0.72. e
consensus tree (Fig. 20) displays the same general
topology as in Bianucci et al. (2010). All the Iberian
ziphiids are placed inside the Ziphiinae. Choneziphi-
us leidyi n. sp. appears as sister taxon of C. planirostris,
the species coded in the previous analysis for the genus
Choneziphius. e unresolved relationships between
T. atlanticus n. sp. and T. crispus are probably due to
the incompleteness of the holotype and only referred
specimen of T. crispus. Nevertheless, both species are
more basal than the Choneziphius + Izikoziphius +
Ziphius clade, similar to the position of T. crispus in
Bianucci et al. (2010). Globicetus n. gen. and Imocetus
n. gen. have a more basal position inside the Ziphiinae,
redefined here (see the emended diagnosis above) with
the inclusion of these two new genera.
INTRASPECIFIC VARIATION
Significant morphological variation, considered
as possibly related to sexual dimorphism and/or
ontogeny, is observed in the samples of skulls re-
ferred to Tusciziphius atlanticus n. sp. and Imocetus
piscatus n. gen., n. sp.
Table 5. — Measurements (in mm) on the skull of aff. Ziphirostrumsp. SGHN MA0936 from the Atlantic Ocean oor off the Iberian
Peninsula. Abbreviation: e, estimate.
SGHN MA0936
Rostrum length 557
Distance from apex of rostrum to bony nares e635
Width of rostrum at mid-length 71
Width of premaxillae at mid-length of rostrum 49
Height of rostrum at mid-length 90
Width of rostrum base at prominental notch e120
Width of rostrum base at antorbital notch e208
Distance rostrum base – anterior apex of palatine 130
Preorbital width of skull e314
Longitudinal distance left premaxillary foramen-rostrum base 40
Width of left premaxillary sac fossa 49
141
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
e intraspecific variation observed at the
level of the medial premaxillary bulge within the
species T. atlanticus n. sp. is very conspicuous,
the most significant found until now in a fos-
sil ziphiid considering the range of sizes for the
concerned element. Interestingly, the develop-
ment and extension of the medial premaxillary
bulge shows a correlation with the development
of rostral maxillary eminences. Indeed, the speci-
mens that display a high medial premaxillary
bulge (holotype, SGHN MA0644, and SGHN
MA0632) also bear rostral maxillary eminences,
more developed on the right side. Such a varia-
tion could be related to sexual dimorphism. In
several extant ziphiids, a high intraspecific vari-
ation related to sex and age has been reported:
filling of the mesorostral groove by the vomer
more pronounced in adult males of Mesoplodon
Aexcrescences on maxilla
frontal
maxilla
foramina
maxilla
maxilla
premaxilla
frontal
vomer palatine
palatine
premaxilla
anterior end
of maxilla
vestigial alveolar
groove
frontal
vomer
vomer
vestial alveolar groove
antorbital notch
maxilary tubercle
prominental notch
maxilla
premaxilla
lacrimal
ventral infraorbital foramen
dorsal infraorbital
foramina
premaxillary foramen
prenarial basin
left premaxillary
sac fossa
B
C
Fig.19. — Skull of aff. Ziphirostrumsp. (SGHN MA0936): A, dorsal view; B, ventral view; C, lateral view. Parallel lines indicate a break
surface. Scale bar: 10cm.
142 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
spp. and Ziphius cavirostris Cuvier, 1823, further
development of the prenarial basin in adult males
of Z. cavirostris, much higher rostral maxillary
crests in adult males of Hyperoodon ampullatus
(Forster, 1770), and larger mandibular tusks
in adult males of many extant ziphiids, and
possibly fossil ziphiids (e.g., Heyning 1984,
1989a, b; Mead 1989a, b; MacLeod & Her-
man 2004; Lambert et al. 2010). Based on the
development of the medial premaxillary bulge
and the rostral maxillary eminences in T. at-
lanticus n. sp., we propose that the holotype,
SGHN MA0644, SGHN MA0632, and SGHN
MA0926 are males, whereas the paratype and
SGHN MA0914 are interpreted as females or
immature males, by analogy with extant ziphiids,
especially H. ampullatus. In adult males of the
latter, further development of the rostral maxil-
lary crests starts at the onset of sexual maturity,
leading to a larger size, massive proportions,
and a flattened anterodorsal surface (Hardy
2005). Similarly the different degrees of devel-
opment of the medial premaxillary bulge and
of the rostral maxillary eminences observed in
T. atlanticus n. sp. might be explained by sexual
dimorphism and maturity.
In I. piscatus n. gen., n. sp., the significant
morphological variation observed between the
holotype and the more fragmentarily known
referred specimens is also related to bony struc-
tures influenced by high sexual dimorphism in
several modern ziphiids. Indeed, the most striking
difference, namely the length of the preorbital
process and the related position of the antorbital
notch, might be linked to the development of
the maxillary crest on the supraorbital process.
is crest forms a lateral boundary to a wide
facial depression seemingly analogous (but not
homologous) to the wide prenarial basin of
adult male Z. cavirostris (see Heyning 1989a;
Cranford et al. 2008). Nevertheless, a larger
sample for I. piscatus n. gen., n. sp. would be
necessary to give a firmer interpretation for this
observed variation.
FUNCTIONAL ANATOMY
From a functional point of view, bony crests,
depressions, and prominences in the facial area
of modern odontocetes are often demonstrated
to be related to soft anatomy elements of the
forehead (facial and rostral muscles, blowhole,
melon, nasal passages, nasal sacs, and other
structures associated with echolocation; see e.g.,
F
ig
.20. — Consensus tree of 875 equally parsimonious clad-
ograms showing the relationships of the Iberian fossil ziphiids
(inbold) with other fossil and extant ziphiid genera. Tree length
94, Goloboff t –20.98, CI 0.53 and RI 0.72. See Bianucci et al.
(2011) for the description of characters and matrix, and Appendix1
for the coding of characters of taxa not included in that previous
analysis. †, strictly fossil taxa.
Squalodon †
Eurhinodelphis †
Xhosacetus †
Hyperoodontinae Ziphiinae
Ziphiidae
Berardiinae Messapicetus clade
Africanacetus †
Ihlengesi †
Izikoziphius †
Choneziphius †
planisostris †
C. Leidy n. sp. †
Tusciziphius
crispus †
T. atlanticus n. sp. †
Globicetus n. gen. †
Imocetus n. gen. †
Microberardius †
Archaeoziphius †
Nazcacetus †
Messapicetus †
Beneziphius †
Ziphirostrum †
Tasmacetus
Berardius
Ziphius
Pterocetus †
Indopacetus
Mesoplodon
Hyperoodon
143
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
Cranford et al. 2008; Huggenberger et al. 2009).
For example, the deep prenarial basin of adult
males of Ziphius cavirostris contains a fat body
usually identified as the melon (Heyning 1989a),
but it has been recently differentiated from the
melon as the anterior spermaceti organ (Cran-
ford et al. 2008). Low maxillary crests in the
supraorbital region of odontocetes correspond
to areas of origin for facial muscles (Mead 1975;
Heyning 1989a; Mead & Fordyce 2009), acting
on the nasal passages, blowhole, and melon. In
addition, the lateral and medial surfaces of the
huge rostral maxillary crests of Hyperoodon spp.
are areas of origin for several facial and rostral
muscles (Schenkkan 1973), even if this is much
likely not the unique function of the crests (see
Mead 1989a; Gowans & Rendell 1999; Hardy
2005). e premaxillary eminences anterolateral
to the bony nares of Phocoena bring the overlying
premaxillary sac fossae, potential sound reflectors,
closer to the acoustic pathway (Huggenberger
et al. 2009).
ROSTRAL PREMAXILLARY THICKENING
Several functions have been proposed for the
varied conditions of pachyosteosclerotic rostral
bones in extinct and extant ziphiids (Heyning
1984; Buffrénil & Casinos 1995; Zioupos et al.
1997; MacLeod 2002). Until now, no single func-
tional interpretation explains the whole diversity
of morphologies observed (Buffrénil & Lambert
2011; Lambert et al. 2011). e spherical promi-
nence of Globicetus n. gen. and the medial bulge
of Tusciziphius atlanticus n. sp. are certainly some
of the most bizarre rostral elements described to
date for odontocetes, and the question of their
potential function as well as the question of their
influence on, or link with, the echolocation system
are both difficult to answer.
In lateral view, the anterior margin of the sphe-
roid in Globicetus n. gen. and of the bulge in
T. atlanticus n. sp. occupies a position roughly
similar to the anterior surface of the elevated
rostral maxillary crests of extant Hyperoodon. In
addition to being areas of origin for facial muscles
(Heyning 1989a), the crests of H. ampullatus are
thought to be used as weapons during head-butting
encounters between adult males; additionally, they
might also provide a protection for the soft tis-
sues located posterior and between them, mostly
the melon, during impacts (Gowans & Rendell
1999; Hardy 2005).
A similar function could be proposed in Glo-
bicetus n. gen. and T. atlanticus n. sp. For the
latter, it would be corroborated by the sexual di-
morphism interpretation given above. However,
contrasting with the spongy aspect of the bone in
Hyperoodon (Hardy 2005; Lambert et al. 2011),
the superficial layers of the spheroid of Globicetus
n. gen. are made of very compact bone. Similarly,
high compactness has been detected through
preliminary macroscopic observations of trans-
verse sections of the bulge in T. atlanticus n. sp.,
suggesting very different mechanical properties
for this element (see discussion for Mesoplodon
densirostris Blainville, 1817 in Buffrénil et al.
2000). Another type of function might be related
to the deep-diving habit of extant ziphiids. In-
deed, this voluminous element, made of compact
bone, distinctly increases the weight of the skull,
particularly in Globicetus n. gen. Such a feature
has been proposed to help maintaining a verti-
cal position in the water during descents towards
feeding areas in other ziphiids (Buffrénil & Ca-
sinos 1995; Zioupos et al. 1997), but ecological
data on extant ziphiids do not explain for now
the observed sexual dimorphism. A combina-
tion of functions might likely better reflect the
diversity of rostrum forms observed (Buffrénil
et al. 2000; Lambert et al. 2011). It is clear that
additional analyses will be necessary to continue
the discussion of the potential function(s) of the
spheroid and of the bulge. e examination of
the inner bone organization, through Computed
Tomography scanning (CT scan) or ground sec-
tions of more fragmentary specimens, will bring
additional data about the compactness, mechanical
properties, and growth process of these unusual
bony structures.
In addition to the question of its function, the
spheroid of Globicetus n. gen., and in a lesser extent
the narrower medial bulge of T. atlanticus n. sp.,
must obviously be considered in the framework
of echolocation. Indeed, in odontocetes the echo-
144 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
location sounds are thought to be produced in
the area of the forehead roughly vertical to the
bony nares, and transmitted forwards via a low-
density pathway including the melon (Cranford
et al. 1996, 2008). If the lateral view of the skull
of Globicetus is compared to a CT scan of the
head of the extant Ziphius (see Cranford et al.
2008: fig. 6), the spheroid of Globicetus n. gen.
is only slightly more anterior than the posi-
tion of the melon in Ziphius, and it is nearly as
high. erefore, there is only little space for the
melon in Globicetus n. gen., and this soft tissue
was certainly located more posterodorsally than
in other ziphiids. In Hyperoodon, the melon has
been described as elongated, situated between
the rostral maxillary crests (Schenkkan 1973),
a condition impossible in Globicetus n. gen. In
the latter, the only way for the transmission of
sounds is dorsal to the spheroid, which must be
considered as an unsurpassable obstacle, an acous-
tic reflector, considering its high compactness
(acoustic impedance mismatch with surround-
ing soft tissues, including the phonic lips area,
where the sounds are produced, and the melon,
through which the sounds are transmitted). is
implies that the sounds were produced at a level
high enough above the level of the dorsal surface
of the spheroid.
ROSTRAL MAXILLARY CREST
For the rostral maxillary eminences and crests
of Tusciziphius atlanticus n. sp. and Imocetus
n. gen., n. sp., a similar functional explanation
might be proposed: protection of forehead soft
tissues facing more lateral impacts. However, the
supraorbital region and rostrum base of odon-
tocetes is mainly an area of origin for facial and
rostral muscles, acting on the air sac system, the
blowhole, the nasal plugs, and the melon (Heyn-
ing 1989a). e development of high crests might
provide surfaces for the attachment of muscles,
with a different direction of action. Considering
the posterodorsal direction of the pointed spur-
like rostral maxillary crest in Imocetus n. gen.,
muscles originating there reached a relatively
posterior region of the forehead, possibly the
posterior part of the melon. In various odon-
tocetes, rostral muscles have been proposed to
modulate the shape of the melon, influencing
therefore the shape of the sound beam (Mead
1975; Au 1993; Huggenberger et al. 2009). In
T. atlanticus n. sp., the asymmetry of the crests
(right crest considerably larger than left crest)
would mirror the asymmetry of the forehead soft
tissues observed in extant ziphiids, as well as in
other odontocetes (Heyning 1989a; Cranford
et al. 1996). A similar argument was proposed
to explain the asymmetry in the development of
excrescencies on both sides of the rostrum base
in Choneziphius planirostris, presumably for the
attachment of rostral muscles (Lambert 2005).
FACIAL BASIN AND MAXILLARY CREST
By comparison with the odontocetes displaying
the most developed facial basin (supracranial
basin in the sperm whale Physeter and prenarial
basin in adult males of Ziphius), the large facial
depression of Imocetus n. gen. probably contained
the main portion of the fat bodies of the forehead
(anterior spermaceti organ and/or melon). e
unusual length of the facial area, linked to the
derived anterior shift of the preorbital process,
might be an indication of an enlarged fat body.
In modern odontocetes, melon and spermaceti
organ are both considered as low density prefer-
ential acoustic pathways, for the transmission and
shaping of echolocation sound beams (reviewed in
Cranford et al. 1996; Cranford 1999). e lateral
flank of the long and high maxillary crest on the
supraorbital process of Imocetus was probably an
important area of origin for facial muscles. In the
modern Mesoplodon carlhubbsi Moore, 1963, Hey
-
ning (1989a: figs 6-8) describes two longitudinal
ridges on the supraorbital process: the lateral ridge,
ending anteriorly as the antorbital tubercle, and
the maxillary ridge, ending anteriorly as the maxil-
lary prominence (or maxillary tubercle). Possibly
corresponding to the supraorbital crest of other
odontocetes, the lateral ridge is the site of origin
for the pars anteroexternus of the m. maxillonasola-
bialis, whereas the maxillary ridge (= maxillary
crest) is the site of lateral origin of the much larger
pars anterointernus (Heyning 1989a; synonymies
in Mead & Fordyce 2009). It is more likely that
145
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
the maxillary crest of n. gen., much higher than
the ridges described in M. carlhubbsi, was the site
of origin for the pars anterointernus. In modern
ziphiids, fibers of this muscle insert on the nasal
passages (Heyning 1989a).
PALAEOBIOGEOGRAPHY
AND PALAEOECOLOGY
e 18 diagnostic partial skulls recovered from
the Atlantic Ocean floor off the coasts of Spain
and Portugal are referred to the six genera (two
of them are new) and four species (all of them
are new) listed below.
– Choneziphius leidyi n. sp. (three specimens from
Galicia, 2 from Portugal);
– Tusciziphius atlanticus n. sp. (four from Galicia,
one from Portugal);
– Globicetus hiberus n. gen., n. sp. (two from
Galicia, one from Portugal);
– Imocetus piscatus n. gen., n. sp. (three from
Portugal);
– aff. Caviziphius sp. (one from Galicia);
– aff. Ziphirostrum sp. (one from Galicia).
Several other ziphiid skulls, including two speci-
mens trawled off the Galician coast and identified
by us as belonging to I. piscatus n. gen., n. sp.,
are not described here in detail because they are
kept in private collections.
In addition to the fossil ziphiids, the following
fragmentary remains were also collected: five iso-
lated teeth of stem physeteroids; three periotics,
eight tympanic bullae, and a skull fragment all
belonging to mysticetes (Balaenidae, Balaenopteri-
dae and Cetotheriidae); and several vertebrae and
teeth of sharks (some belonging to Cosmopolitodus
hastalis and Carcharocles megalodon).
On the whole, the fossil ziphiid associations
of Galicia and Portugal show the same compo-
sition, with the exception of two fragmentary
skulls collected off the Galician coast and referred
to aff. Caviziphius sp. and aff. Ziphirostrum sp.
A similar condition is present today as the ziphiid
communities, and more generally the cetacean
assemblages, off Galicia and Portugal are substan-
tially similar (Covelo & Martínez 2001). Even if
we cannot demonstrate that all these taxa lived
at the same time, this past ziphiid diversity is
roughly similar to the present diversity; six extant
species have been recorded by strandings and/
or sightings off the Atlantic coast of the Iberian
Peninsula: Hyperoodon ampullatus, Mesoplodon
bidens (Sowerby, 1804), M. densirostris, M. euro-
paeus (Gervais, 1855), M. mirus True, 1913, and
Ziphius cavirostris (see Reiner 1979; Valverde &
Galan 1996; Valverde 1997; López et al. 2002;
Kiszka et al. 2007; Smith 2010). e geographic
distribution of at least a part of these extant taxa
being strongly related to topographic parameters
(depth and slope) (Smith, 2010), we think that
the concentration of large and heavy, not easily
transported, fossil specimens at important depths
in the proximity of local topographic features (e.g.,
Nazaré Canyon off Portugal or Ortegal Spur off
Galicia) might reflect roughly similar ecological
preferences.
Acknowledging the lack of precise stratigraphic
data (see below), a comparison between the fossil
ziphiid association of the Iberian Atlantic coast
and those of other areas (see the systematic para-
graph below) reveals that: 1) Tusciziphius atlanticus
n. sp. is also recorded in South Carolina (USA),
whereas another species of Tusciziphius (T. crispus)
was found in Tuscany (Italy); 2) Choneziphius is
also reported in South Carolina and in the North
Sea (Belgium and e Netherlands), with a few
large indeterminate specimens (roughly similar
to C. leidyi n. sp.) and smaller specimens (re-
ferred to C. planirostris in the North Sea); and 3)
Caviziphius and Ziphirostum are recorded in the
area of Antwerp (Belgium). e fossil record of
Globicetus n. gen. and Imocetus n. gen. is instead
restricted for now to the Iberian Atlantic coast.
Unfortunately the fossil ziphiid samples from
eastern North America and Mediterranean are
still too fragmentary for a comprehensive com-
parison between these associations. e fossil
ziphiid record outside the North Atlantic Ocean
and Mediterranean Sea is even more fragmentary,
with two major exceptions: Peru and South Africa.
From the middle Miocene-Pliocene Pisco For-
mation of Peru, Messapicetus gregarius Bianucci,
Lambert & Post, 2010, Ninoziphius platyrostris
146 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
Muizon, 1983, and Nazcacetus urbinai Lambert,
Bianucci & Post, 2009 are recorded. None of these
three genera is present in the Atlantic Iberian fossil
association, but Messapicetus is reported in Italy
with a different species (M. longirostris Bianucci,
Landini & Varola, 1992; see Bianucci et al. 1992)
and is tentatively reported from Maryland, east
coast of USA (Fuller & Godfrey 2007), whereas
Ninoziphius Muizon, 1983 is also recorded, based
on a fragmentary specimen, from the east coast of
USA (Muizon & DeVries 1985; Morgan 1994).
e fossil ziphiid association recovered by
trawling from the sea floor off the South African
coast, also characterized by a low stratigraphic
resolution, is very diversified, being represented
by at least 11 species and 9 genera (Bianucci
et al. 2007, 2008). Surprisingly, none of the
taxa recorded in South Africa is listed in the
Atlantic Iberian fossil association or any other
North Atlantic realm assemblages, possibly evi-
dencing an ecological and/or physical barrier
between North and South Atlantic, in addition
to expected temporal gaps between different
communities (and between species from a same
region). An analysis of the extant ziphiid com-
munity reveals substantial differences, even if less
pronounced, between the beaked whales off the
Iberian Peninsula and South African coasts (Ross
1984; Mead 1989b; López et al. 2002; Dalebout
et al. 2003, Kiszka et al. 2007; Smith 2010). At
the genus level the widely ranging Hyperoodon,
Mesoplodon, and Ziphius are recorded in both
areas, whereas Berardius Duvernoy, 1851 and
Indopacetus Moore, 1968 are not recorded in
the North Atlantic. At the species level, among
the eight ziphiid species living off the South
African coasts, only the widely ranging Mesoplo-
don densirostris, M. mirus and Ziphius cavirostris
are also recorded off the Atlantic coast of the
Iberian Peninsula.
e different modern cetacean compositions
of the northern and southern hemispheres are
directly related to the presence of an equato
-
rial warm water mass, representing an efficient
barrier to dispersal, but also generating vicari-
ant speciation (e.g., in the genera Berardius and
Hyperoodon, both containing antitropical spe-
cies) during temporary cooling event(s) of the
oceanic waters (Davies 1963; Hare et al. 2002).
It is possible that most of the fossil ziphiids of
Iberia and South Africa were restricted to cold
and/or temperate waters and consequently were
not able to cross the warm equatorial barrier, as
today for species of Berardius and Hyperoodon.
GEOLOGICAL SETTING
AND AGE OF THE SPECIMENS
On the Ortegal Spur, off the northwestern corner
of the Iberian Peninsula, one of the areas where
fossil ziphiids were found (Fig. 1), the Neogene
sedimentary succession lies above about 1200 m of
Late Jurassic-Eocene deposits and is represented by
Oligo-Miocene silty marl and foraminiferal ooze,
associated with slope breccias or conglomerates,
indicating a relatively deep-water depositional
environment (Wallrabe-Adams et al. 2005; Jané
et al. 2010). It is probable that the fossil ziphiids
originate from one or more phosphorite episodes
within this succession (see below). e Miocene
sediments are irregularly covered by Plio-Quater-
nary deposits consisting of alternated silt and clay
laminas interbedded with coarse sediments (Jané
et al. 2010).
e Nazaré Canyon, in the area where the fossil
ziphiids from Portugal were found, represents one
of the late Variscan faults that cut transversally the
Mesozoic rifted Iberian margin (Pinheiro et al. 1996).
At the end of Mesozoic the rifting ended; during
Eocene and Miocene this area suffered a compres-
sional episode that reactivated the old Variscan
structures (Pinheiro et al. 1996). Published studies
on the sediments outcropping on the sea bottom
of the Nazaré Canyon report that a large portion
is covered with Holocene mud (Koho et al. 2007;
Masson et al. 2011). e age of rocks of the area
where the fossil ziphiids were found include Miocene
and Pliocene (Badagola 2008; LNEG-LGM 2001).
According to the geological map of the continental
platform, the only Miocene rocks that might have
been the source of the skulls here described are in
an area around 39°18’N and 9°47’W and about
160 m of depth, in the Mar da Ericeira, which has
147
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
been dated as Aquitanian to Langhian, early to
middle Miocene (Badagola 2008).
Although we have not found data about the
presence of phosphorite levels in the specific areas
where the fossil ziphiids were collected, North
Atlantic phosphorite deposits dated to Cenozoic
are reported both off Spain and Portugal (Riggs &
Sheldon 1990). ese sediments are related to the
Upper Early Miocene-Middle Miocene (20-14 Ma)
episode of phosphogenesis associated with the
TB2 second-order eustatic sea-level fluctuation
(Haq et al. 1987; Riggs & Sheldon 1990). is
interval of age is compatible with those of the
fossil ziphiids examined, considering that the
oldest unquestionable records of this family are
from late early Miocene-middle Miocene (Bi-
anucci et al. 2005; Lambert & Louwye 2006).
Unfortunately precise ages are not available for
most of the fossil ziphiids from other localities
in the world that are related to those examined
here. Indeed, most of these fossil ziphiids are from
phosphorite deposits and/or their precise strati-
graphical position is unknown. Moreover, since
only one Iberian species (Tusciziphius atlanticus
n. sp.) is also reported from a locality outside
Iberia, faunal correlations remain difficult. As
mentioned above, T. atlanticus n. sp. was found
reworked on the bottom of Morgan River (South
Carolina, USA), and a late Miocene-Pliocene
age can only be proposed (Post et al. 2008).
Tusciziphius is also recorded from Italy, with a
more precise early Pliocene age (Bianucci 1997;
Bianucci et al. 2001), but with a different spe-
cies (T. crispus). A precise dating is not available
for most of the Neogene ziphiids of the North
Sea (Belgium and the Netherlands) (Bianucci &
Post 2005; Lambert 2005; Lambert & Louwye
2006). is is unfortunately especially true for
genera also reported (Choneziphius), or tenta-
tively reported (Caviziphius and Ziphirostrum),
from Iberian Peninsula. Nevertheless Lambert
(2005) mentioned a skull of Ziphirostrum mar-
ginatum and a skull of Choneziphius planirostris,
both collected from the Tortonian (late Miocene)
Deurne Sands Member of the Diest Formation
(Antwerp, Belgium). e genus Choneziphius is
also reported from Phosphate Beds of South Car-
olina, USA, and from reworked sediments of the
Lee Creek Mine, North Carolina (Whitmore &
Kaltenbach 2008), with fragmentary skulls that,
based on their large size, could be conspecific
with C. leidyi n. sp. Interestingly, even if the age
of these Northern American ziphiids is not well
defined (see Post et al. 2008), Riggs & Sheldon
(1990: table 18.1) refer the South and North
Carolina phosphorites to the same episode (TB2)
than the Iberian phosphorites. In conclusion, the
comparison with fossil ziphiids from other local-
ities only provides minor clues for the definition
of the age of the phosphoritized Iberian ziphi-
ids. A late early Miocene-middle Miocene age,
as suggested by TB2 phosphorites, is probable,
even if the few well-dated extra-Iberian ziphiids
(belonging to different species) are younger (late
Miocene or early Pliocene). However, we cannot
exclude a priori that all or part of these fossil
ziphiids were collected from younger phosphorite
episodes (e.g., the TB3, Late Miocene-Pliocene),
for now not documented from the sea floor off
the Atlantic coasts of Spain and Portugal. New
data on the local geology and stratigraphy and
the dating of associated phosphorites through
radiometric methods are needed to better define
the very approximative ages provided here for
these specimens. Such additional data would be
crucial to further support the palaeobiogeographic
and palaeoecological hypotheses discussed above
(see Pyenson et al. 2009 for an example of the
importance of a detailed sedimentological and
stratigraphical analysis for the understanding of
fossil marine mammal localities).
CONCLUSION
e systematic study of 40 partial fossil ziphiid
skulls dredged from the Atlantic Ocean floor off
Portugal and Spain lead to the description of two
new genera, Globicetus n. gen. and Imocetus n. gen.,
and four new species, G. hiberus n. gen., n. sp.,
I. piscatus n. gen., n. sp., Choneziphius leidyi n. sp.,
and Tusciziphius atlanticus n. sp. In addition, mem-
bers of the genera Caviziphius and Ziphirostrum are
tentatively reported.
148 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
e phylogenetic analysis of the new taxa places
them in the subfamily Ziphiinae here redefined,
also including Choneziphius planirostris, Tusciziphius
crispus, and the genera Izikoziphius and Ziphius.
Bizarre elements observed on the skull of Glo-
bicetus n. gen. (large premaxillary spheroid) and
T. atlanticus n. sp. (medial premaxillary bulge, likely
sexually dimorphic), are commented from a func-
tional point of view; these structures are confronted
to various hypotheses proposed for the function of
pachyosteosclerotic parts of the rostrum in several
ziphiid lineages.
Other peculiar features of the skull of Imocetus
(spur-like rostral maxillary crests and long maxillary
crests limiting a large facial basin) and Choneziphius
spp. (excrescences on the maxilla at the rostrum
base) are interpreted as areas of origin for rostral
and facial muscles.
e palaeobiogeography of Neogene ziphiids is
discussed in the light of the new discoveries. Differ-
ences in the composition of cold to temperate north-
ern and southern hemisphere ziphiid communities
might be explained by a warm equatorial barrier.
Finally, by comparison with other fossil ziphiid
assemblages in the world and on the basis of a few
geological and stratigraphic data for the Atlantic
Ocean floor off Portugal and Spain, a late early to
middle Miocene age is very tentatively proposed
for the studied specimens. However, this hypothesis
should be considered cautiously; a younger age can-
not be excluded for part or all of these specimens.
Acknowledgements
We would like to thank the following fishermen
who donated specimens to SGHN: Miguel Ángel
Iglesias and the crew of the ship Nuevo Richard
from Cedeira; José Castro Sambás and his son
Daniel from Muxía; Luis from Camelle; José An-
tonio González and Lino from Cedeira; the crew
of the ship Gonzacove Uno; Manuel Ángel Iglesias
from Cedeira. anks are also due to Carlos Filipe
Alexandre, Estevão Anastácio da Cruz, Francisco
Reiner, Ildo Hermógenes Marques da Silva, José
Augusto, Luciano Mesquita, and Mário Estevens
who donated the Portuguese specimens to the ML
or provided informations about the locations, and
to Remmert Schouten and Pedro Viegas for calling
our attention to Portuguese specimens. We also
thank for access to the collections under their care
the following persons and/or institutions: P. Agnelli
(Museo di Zoologia, Università di Firenze), CEM-
MA (Coordinadora para o Estudio dos Mamíferos
Mariños, Galicia), E. Cioppi and S. Dominici (IGF),
F. J. Cristobo (IEO), J. I. Díaz (SGHN), X. Guerra
and I. Fraga (MHNUSC), G. Lenglet (Institut royal
des Sciences naturelles de Belgique, Brussels), J. R.
García and Lucía (Museo Marítimo de Asturias
in Luanco), J. G. Mead and C. W. Potter (United
States National Museum of Natural History, Smi-
thsonian Institution, Washington DC), S. van de
Mije, H. van Grouw, and R. van Zelst (Nationaal
Natuurhistorisch Museum Naturalis, Leiden), and
H. van der Es (NMR). Finally we wish to thank
the reviewers C. de Muizon (Muséum national
d’Histoire naturelle, Département Histoire de la
Terre, Paris), and N. D. Pyenson (United States
National Museum of Natural History, Smithsonian
Institution, Washington DC) for their constructive
comments that greatly improved the manuscript,
as well as the editors D. Merle and A. Ohler for
nomenclature points.
REFERENCES
ABEL O. 1905. — Les odontocètes du Boldérien (Mi-
ocène supérieur) des environs d’Anvers. Mémoires du
Musée royal d’Histoire naturelle de Belgique 3: 1-155.
AU W. W. L. 1993. — e Sonar of Dolphins. Springer
Verlag, New York, 277 p.
B
ADAGOLA
A. P. L. 2008. — Evolução morfo-tectónica
da plataforma continental do Esporão da Estremadura.
Tese de mestrado, Geologia Dinâmica (Geodinâmica).
Faculdade de Ciências, Universidade de Lisboa, avai-
lable at http://repositorio.ul.pt/handle/10451/1308
B
IANUCCI
G. 1997. — e Odontoceti (Mammalia
Cetacea) from Italian Pliocene. e Ziphiidae. Pal-
aeontographia Italica 84: 163-192.
BIANUCCI G. & POST K. 2005. — Caviziphius altiros-
tris, a new beaked whale from the Miocene southern
North Sea basin. Deinsea 11: 1-6.
B
IANUCCI
G., L
ANDINI
W. & V
AROLA
A. 1992. — Messa-
picetus longirostris, a new genus and species of Ziphiidae
(Cetacea) from the late Miocene of “Pietra Leccese”
(Apulia, Italy). Bolletino della Società Paleontologica
Italiana 31: 261-264.
149
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
B
IANUCCI
G., L
ANDINI
W. & V
AROLA
A. 1994. —
Relationships of Messapicetus longirostris (Cetacea,
Ziphiidae) from the Miocene of South Italy. Bollettino
della Società Paleontologica Italiana 33 (2): 231-241.
B
IANUCCI
G., M
AZZA
P., M
EROLA
D., S
ARTI
G. &
CASCELLA A. 2001 — e Early Pliocene mammal
assemblage of Val di Pugna (Tuscany, Italy) in the
light of calcareous plankton biostratigraphical data
and paleoecological observations. Rivista Italiana
di Paleontologica e Stratigrafia 107 (3): 425-438.
B
IANUCCI
G., L
ANDINI
W., V
ALLERI
G., R
AGAINI
L. &
V
AROLA
A. 2005. — First cetacean fossil records
from Ecuador, collected from the Miocene of Es-
meraldas Province. Rivista Italiana di Paleontologia
e Stratigrafia 111 (2): 345-350.
B
IANUCCI
G., L
AMBERT
O. & P
OST
K. 2007. — A
high diversity in fossil beaked whales (Odontoceti,
Ziphiidae) recovered by trawling from the sea floor
off South Africa. Geodiversitas 29 (4): 561-618.
B
IANUCCI
G., P
OST
K. & L
AMBERT
O. 2008. — Beaked
whale mysteries revealed by sea floor fossils trawled
off South Africa. South African Journal of Science
104 (3-4): 140-142.
BIANUCCI G., LAMBERT O. & POST K. 2010. — High
concentration of long-snouted beaked whales (genus
Messapicetus) from the Miocene of Peru. Palaeon-
tology 53 (5): 1077-1098.
BRISSON M.-J. 1762. — Regnum Animale in classes IX
distributum, sine synopsis methodica. eodorum
Haak, Paris, 296 p.
BUFFRÉNIL V. DE & CASINOS A. 1995. — Observations
histologiques sur le rostre de Mesoplodon densirostris
(Mammalia, Cetacea, Ziphiidae) : le tissu osseux
le plus dense connu. Annales de Sciences Naturelles,
13e série, 16: 21-32.
BUFFRÉNIL V. DE & LAMBERT O. 2011. — Histology
and growth pattern of the pachy-osteosclerotic
premaxillae of the fossil beaked whale Aporotus
recurvirostris (Mammalia, Cetacea, Odontoceti).
Geobios 44: 45-56.
B
UFFRÉNIL
V.
DE
, Z
YLBERBERG
L., T
RAUB
W. & C
ASINOS
A. 2000. — Structural and mechanical characteristics
of the hyperdense bone of the rostrum of Mesoplodon
densirostris (Cetacea, Ziphiidae): summary of recent
observations. Historical Biology 14: 57-65.
COVELO P. & MARTÍNEZ J. 2001 — Varamientos de
mamíferos marinos en las costas de España y Portugal
entre 1996 y 1998: Atlancetus. Galemys 13: 93-106.
C
RANFORD
T. W. 1999. — e sperm whale’s nose:
sexual selection on a grand scale? Marine Mammal
Science 15 (4): 1133-1157.
C
RANFORD
T. W., A
MUNDIN
M. & N
ORRIS
K. S.
1996. — Functional morphology and homology
in the Odontocete nasal complex: implications
for sound generation. Journal of Morphology 228:
223-285.
CRANFORD T. W., MACKENNA M. F., SOLDEVILLA M.
S., WIGGINS S. M., GOLDBOGEN J. A., SHADWICK
R. E., KRYSL P., ST. LEGER J. A. & HILDEBRAND J.
A. 2008. — Anatomic geometry of sound trans-
mission and reception in Cuvier’s beaked whale
(Ziphius cavirostris). e Anatomical Record 291
(4): 353-378.
C
UVIER
G. 1823. — Recherches sur les ossements fos-
siles, 5 (1ère partie). G. Dufour et E. D’Ocagne,
Paris, 405 p.
D
ALEBOUT
M. L., M
EAD
J. G., B
AKER
C. S., B
AKER
A.
N. & VAN HELDEN A. L. 2002. — A new species
of beaked whale Mesoplodon perrini sp. n. (Cetacea:
Ziphiidae) discovered through phylogenetic analyses
of mitochondrial DNA sequences. Marine Mammal
Science 18 (3): 577-608.
DALEBOUT M. L., ROSS G. J. B., BAKER C. S., ANDER-
SON R. C., BEST P. B., COCKCROFT V. G., HINSZ H.
L., PEDDEMORS V. & PITMAN R. L. 2003. — Ap-
pearance, distribution, and genetic distinctiveness
of Longman’s beaked whale, Indopacetus pacificus.
Marine Mammal Science 19 (3): 421-461.
D
AVIES
J. L. 1963. — e antitropical factor in cetacean
speciation. Evolution 17 (1): 107-116.
D
U
B
US
B. A. L. 1868. — Sur différents Ziphiides
nouveaux du Crag d’Anvers. Bulletin de l’Académie
Royale des Sciences de Belgique 25: 621-630.
DU BUS B. A. L. 1872. — Mammifères nouveaux du
Crag d’Anvers. Bulletin de l’Académie royale des Sci-
ences de Belgique 34: 491-509.
D
UVERNOY
G. 1851. — Mémoire sur les caractères
ostéologiques des genres nouveaux ou des espèces
nouvelles de cétacés vivants ou fossiles. Annales de
Sciences naturelles, Zoology 15: 52-68.
E
STEVENS
M. & Á
VILA
S. P. 2007. — Fossil whales
from the Azores. Açoreana. Revista de Estudos Aço-
reanos 5: 140-161.
FLOWER W. H. 1867. — Description of the skeleton
of Inia geoffrensis and the skull of Pontoporia bla-
invillii, with remarks on the systematic position of
these animals in the Order Cetacea. Transactions of
the Zoological Society of London 6: 87-116.
FORDYCE R. E. & CULLEN D. J. 1979. — A Miocene
ziphiid whale (Odontoceti: Cetacea) from Central
Chatham Rise, East of New Zealand. New Zealand
Oceanographic Institute Records 4 (6): 45-53.
F
ULLER
A. J. & G
ODFREY
S. J. 2007. — A late Miocene
ziphiid (Messapicetus sp.: Odontoceti: Cetacea) from
the St. Marys Formation of Calvert Cliffs, Mary-
land. Journal of Vertebrate Paleontology 27: 535-540.
GOLOBOFF P. 1993. — Estimating character weights
during tree search. Cladistics 9: 83-91.
GOWANS S. & RENDELL L. 1999. — Head-butting in
northern bottlenose whales (Hyperoodon ampullatus):
a possible function for big heads? Marine Mammal
Science 15 (4): 1342-1350.
150 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
GRAY J. E. 1821. — On the natural arrangement of
vertebrose animals. e London Medical Repository
15: 296-310.
G
RAY
J. E. 1850. — Catalogue of the Specimens of
Mammalia in the Collections of the British Muse-
um. Part I – Cetacea. Richard and John E. Taylor,
London, 153 p.
HAQ B. U., HARDENBOL J. & VAIL P. R. 1987. — Chrono-
logy of fluctuating sea levels since the Triassic. Science
235: 1156-1166.
HARDY M. T. 2005. — Extent, development and function
of sexual dimorphisms in the skulls of the Bottlenose
whales (Hyperoodon spp.) and Cuvier’s beaked whale
(Ziphius cavirostris). Master thesis, School of Biolog-
ical Sciences, University of Wales, Bangor, UK, 99 p.
HARE M. P., CIPRIANO F. & PALUMBI S. R. 2002. —
Genetic evidence on the demography of speciation in
allopatric dolphin species. Evolution 56 (4): 804-816.
H
EYNING
J. E. 1984. — Functional morphology in-
volved in intraspecific fighting of the beaked whale,
Mesoplodon carlhubbsi. Canadian Journal of Zoology
62: 1645-1654.
HEYNING J. E. 1989a. — Comparative facial anatomy of
beaked whales (Ziphiidae) and a systematic revision
among the families of extant Odontoceti. Contribu-
tions in Science, Natural History Museum of Los Angeles
County 405:1-64.
HEYNING J. E. 1989b. — Cuvier’s beaked whale Ziphi-
us cavirostris G. Cuvier, 1823, in RIDGWAY S. H. &
HARRISON R. (eds), Handbook of Marine Mammals,
vol. 4: River Dolphins and the Larger Toothed Whales.
Academic Press, London: 289-308.
H
ORIKAWA
H., T
AZAKI
K. & K
ANNO
T. 1987. — [Fossil
Ziphiidae from Koshiji-Syo off Sado Island, Central
Japan]. Bulletin of the Sado Museum 9: 225-230 (in
Japanese).
HUGGENBERGER S., RAUSCHMAN M. A., VOGL T. J. &
O
ELSCHLÄGER
H. A., 2009. — Functional morphol-
ogy of the nasal complex in the harbor porpoise
(Phocoena phocoena L.). e Anatomical Record
292: 902-920.
JANÉ G., MAESTRO A., ERCILLA G., LÓPEZ-MARTÍNEZ
J., DE ANDRÉS J. R., CASAS D., GONZÁLEZ- ALLER
D. & C
ATALÁN
-M
OROLLÓN
M. 2010. — Occurrence
of pockmarks on the Ortegal Spur continental mar-
gin, Northwestern Iberian Peninsula. Marine and
Petroleum Geology 27: 1551-1564.
KISZKA J., MACLEOD K., VAN CANNEYT O., WALK-
ER
D. & R
IDOUX
V. 2007. — Distribution, encounter
rates, and habitat characteristics of toothed cetaceans
in the Bay of Biscay and adjacent waters from plat-
from-of-opportunity data. ICES Journal of Marine
Science 64 (5): 1033-1043.
K
OHO
K. A., K
OUWENHOVEN
T. J., D
E
S
TIGTER
H. C. &
VAN DER ZWAAN G. J. 2007. — Benthic foramin-
ifera in the Nazaré canyon, Portuguese continental
margin: influence of sedimentary disturbance on
fauna. Marine Micropaleontology 66: 27-51.
L
AMBERT
O. 2005. — Systematics and phylogeny
of the fossil beaked whales Ziphirostrum du Bus,
1868 and Choneziphius Duvernoy, 1851 (Cetacea,
Odontoceti), from the Neogene of Antwerp (North
of Belgium). Geodiversitas 27 (3): 443-497.
LAMBERT O. & LOUWYE S. 2006. — Archaeoziphius
microglenoideus, a new primitive beaked whale
(Mammalia, Cetacea, Odontoceti) from the Middle
Miocene of Belgium. Journal of Vertebrate Paleon-
tology 26: 182-191.
L
AMBERT
O., B
IANUCCI
G. & P
OST
K. 2009. — A
new beaked whale (Odontoceti, Ziphiidae) from
the middle Miocene of Peru. Journal of Vertebrate
Paleontology 29 (3): 911-922.
LAMBERT O., BIANUCCI G. & POST K. 2010. —
Tusk-bearing beaked whales from the Miocene of
Peru: sexual dimorphism in fossil ziphiids? Journal
of Mammalogy 91 (1): 19-26.
L
AMBERT
O., B
UFFRÉNIL
V.
DE
& M
UIZO N
C.
DE
2011. — Rostral densification in beaked whales:
diverse processes for a similar pattern. Comptes
Rendus Palevol 10: 453-468.
LANKESTER E. R. 1870. — A new ziphioid cetacean
from the Suffolk Bone-bed (Choneziphius packardi).
Quarterly journal of the Geological Society of London
26: 502-507.
LEIDY J. 1876. — Remarks on fossils from the Ash-
ley Phosphate Beds. Proceedings of the Academy of
Natural Sciences of Philadelphia 1876: 80-81; 86-87.
LEIDY J. 1877. — Description of vertebrate remains,
chiefly from the Phosphate Beds of South Caro-
lina. Journal of the Academy of Natural Sciences of
Philadelphia, 2nd series 8: 209-261.
LNEG-LGM 2001. — Carta Geológica de Portugal à
escala 1:1.000.000 [Geological map]. Lisboa. ISBN:
978-989-675-005-3.
L
ÓPEZ
A., S
ANTOS
M. B., P
IERCE
G. J., G
ONZÁLEZ
A. F., VALEIRAS X. & GUERRA A. 2002. — Trends
in strandings and by-catch of marine mammals in
northwest Spain during the 1990s. Journal of the
marine Biological Association of the U.K. 82: 1-9.
MACLEOD C. D. 2002. — Possible functions of the
ultradense bone in the rostrum of Blainville’s beaked
whale (Mesoplodon densirostris). Canadian Journal
of Zoology 80: 178-184.
MACLEOD C. D. & HERMAN J. S. 2004. — Devel-
opment of tusks and associated structures in Me-
soplodon bidens (Cetacea, Mammalia). Mammalia
68: 175-184.
MASSON D. G., HUVENNE V. A. I., DE STIGTER H.
C., WOLFF G. A., KIRIAKOULAKIS K., ARZOLA R.
G. & B
LACKBIRD
S. 2010. — Efficient burial of
carbon in a submarine canyon. Geology 38: 831-834.
MEAD J. G. 1975. — Anatomy of the external nasal
151
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
passages and facial complex in the Delphinidae
(Mammalia : Cetacea). Smithsonian Contribution
to Zoology 207: 1-67.
MEAD J. G. 1989a. — Bottlenose whales Hyperoodon
ampullatus (Forster, 1770) and Hyperoodon planifrons
Flower, 1882, in RIDGWAY S. H. & HARRISON R.
(eds), Handbook of Marine Mammals, vol. 4: River
Dolphins and the Larger Toothed Whales. Academic
Press, London: 321-348.
M
EAD
J. G. 1989b. — Beaked whales of the genus
Mesoplodon, in R
IDGWAY
S. H. & H
ARRISON
R.
(eds), Handbook of Marine Mammals, vol. 4: River
Dolphins and the Larger Toothed Whales. Academic
Press, London: 349-430.
MEAD J. G. & FORDYCE R. E. 2009. — e therian
skull: a lexicon with emphasis on the odontocetes.
Smithsonian Contributions to Zoology 627: 1-248.
MIJÁN I. 2007. — Hallazgos de restos fósiles de Hy-
peroodon sp. (Cetacea, Ziphiidae) en las costas
gallegas (NO España). Revista de Biología Marina
y Oceanografía 42 (3): 253-260.
M
IYAZAKI
N. & H
ASEGAWA
Y. 1992. — A new species
of fossil beaked whale, Mesoplodon tumidirostris sp.
nov. (Cetacea, Ziphiidae) from the Central North
Pacific. Bulletin of the National Science Museum,
Tokyo, Ser. A 18 (4): 167-174.
M
ORGAN
G. S. 1994. — Miocene and Pliocene marine
mammal faunas from the Bone Valley Formation of
Central Florida, in B
ERTA
A. & D
EMERÉ
T. A. (eds),
Contributions in marine mammal paleontology
honoring Frank C. Whitmore, Jr. Proceedings of the
San Diego Society of Natural History 29: 239-268.
MUIZON C. DE. 1984. — Les Vertébrés de la Formation
Pisco (Pérou). Deuxième partie : les Odontocètes
(Cetacea, Mammalia) du Pliocène inférieur du
Sud-Sacaco. Travaux de l’Institut français d’Études
andines 27: 1-188.
MUIZON C. DE & DEVRIES T. J. 1985. — Geology
and paleontology of late Cenozoic marine deposits
in the Sacaco area (Peru). Geologische Rundschau
74: 547-563.
P
INHEIRO
L. H., W
ILSON
R. C. L., P
ENA
D
OS
R
EIS
R.,
WHITMARSH R. B. & RIBEIRO A. 1996. — e
Western Iberia Margin: a Geophysical and Geolog-
ical Overview, in W
HITMARSH
R. B., S
AWYER
D. S.,
KLAUS A. & MASSON D. G. (eds), Proceedings of the
Ocean Drilling Program, Scientific Results 149: 3-23.
POST K., LAMBERT O. & BIANUCCI G. 2008. — First
record of Tusciziphius crispus (Cetacea, Ziphiidae)
from the Neogene of the US east coast. Deinsea
12: 1-10.
PYENSON N. D., IRMIS R. B., LIPPS J. H., BARNES L.
G., M
ITCHELL
E. D. J
R
. & M
AC
L
EOD
S. A. 2009. —
Origin of a widespread marine bonebed deposited
during the middle Miocene Climatic Optimum.
Geology 37 (6): 519-522.
REINER F. 1979. — Nota sobre un raro ziphioid, Me-
soplodon densirostris, Blainville 1817, nas costas de
Portugal. Museo Marino Cascais, Memorios Series
Zoologie 1: 1-12.
RIGGS S. R. & SHELDON R. P. 1990. — Paleoceano-
graphic and paleoclimatic controls of the temporal
and geographic distribution of upper Cenozoic
continental margin phosphorites, in BURNETT W.
C. & R
IGGS
S. R. (eds), Phosphate deposits of the
world. Volume 3: Neogene to modern phosphorites.
Cambridge University Press, Cambridge: 207-222.
ROBINEAU D. 1973. — Sur deux rostres de Mesoplo-
don (Cetacea, Hyperoodontinae). Mammalia 37
(3): 504-513.
ROSS G. J. B. 1984. — e smaller cetaceans of the
south east coast of southern Africa. Annals of the
Cape Provincial Museums of Natural History 15
(2): 173-410.
SCHENKKAN E. J. 1973. — On the comparative anat-
omy and function of the nasal tract in odontocetes
(Mammalia, Cetacea). Bijdragen tot de Dierkunde
43: 127-159.
S
MITH
J. 2010. — e ecology of Cuvier’s Beaked
Whale, Ziphius cavirostris (Cetacea: Ziphiidae), in
the Bay of Biscay. Unpublished PhD thesis, Gradu-
ate School of the National Oceanography Centre,
Southampton, 214 p.
S
WOFFORD
D. L. 2001. — PAUP*. Phylogenetic
Analysis Using Parsimony (*and other methods).
Version 4b10. Sinauer Associates, Sunderland,
Massachusetts (software).
TAZAKI K., HORIKAWA H. & MIYAZAKI S. 1987. —
[Fossil Ziphiidae from Hyotan-Guri off Sado Is-
land, Central Japan]. Bulletin of the Sado Museum
9: 219-223 (in Japanese).
T
YLER
P., A
MARO
T., A
RZOLA
R., C
UNHA
M. R., D
E
S
TIGTER
H., G
OODAY
A., H
UVENNE
V., I
NGELS
J., K
IRIAKOULAKIS
K., L
ASTRAS
G., M
ASSON
D., O
LIVEIRA
A., P
ATTENDEN
A., V
ANREUSEL
A., V
AN
W
EERING
T., V
ITORINO
J.,
W
ITTE
U. & W
OLFF
G. 2009. — Europe’s grand canyon:
Nazaré submarine canyon. Oceanography 22 (1): 46-57.
VALVERDE J. A. 1997. — Notes on a specimen of Bla-
inville’s beaked whale Mesoplodon densirostris (De
Blainville, 1817) stranded on the coast of Donana,
Huelva, southern Spain. European Research on Ce-
taceans 10: 184-189.
V
ALVERDE
J. A. & G
ALAN
J. M. 1996. — Notes on a spec-
imen of Gervais’ beaked whale Mesoplodon europaeus
(Gervais), Ziphiidae stranded in Andalucia, Southern
Spain. European Research on Cetaceans 10: 177-183.
WALLRABE-ADAMS H. J., ALTENBACH A. V., KEMPE A.,
KUHNT W. & SCHÄFER P. 2005. — Facies develop-
ment of ODP Leg 173 sediments and comparison
with tectono-sedimentary sequences of compressional
Iberian plate margins – a general overview. Journal of
Iberian Geology 31 (2): 235-251.
152 GEODIVERSITAS • 2013 • 35 (1)
Bianucci et al.
W
HITMORE
F. C. J
R
. & K
ALTENBACH
J. A. 2008. —
Neogene Cetacea of the Lee Creek Phosphate Mine,
North Carolina. Virginia Museum of Natural History
Special Publication 14: 181-269.
WHITMORE F. C. JR., MOREJOHN G. V. & MULLINS,
H. T. 1986. — Fossil beaked whales – Mesoplodon
longirostris dredged from the ocean bottom. National
Geographic Research 2 (1): 47-56.
ZBYSZEWSKI G. 1954. — Découverte d’une mandibule
de Palaeoziphius dans le Miocène de Melides. Comu-
nicações dos Serviços Geológicos de Portugal 35: 51-55.
ZIOUPOS P., CURREY J. D., CASINOS A. & BUFFRÉ-
NIL V. DE 1997. — Mechanical properties of the
rostrum of the whale Mesoplodon densirostris, a
remarkably dense bony tissue. Journal of Zoology
241 (4): 725-737.
Submitted on 8 June 2011;
accepted on 28 October 2011;
published on 29 March 2013.
153
Fossil beaked whales fished off the Iberian Peninsula
GEODIVERSITAS • 2013 • 35 (1)
APPENDIX 1
Coding of the characters for taxa not considered in the previous analysis (Bianucci etal. 2010). 0, primitive state; 1, 2, 3, derived
states; a, variable between 0 and 1; ?, missing character.
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29
Choneziphius planirostris 2030112a11101?0102????????1? ?
Choneziphius leidyi 1030113011 1 0 1 ?0102????????1? ?
Tusciziphius crispus ?0301032?1102100021???????1? ?
Tusciziphius atlanticus 10301032?1102100021???????1? ?
Globicetus 10301032?311210002????????1? ?
Imocetus 20310032?31011?002????????1? ?
