The Island Arc (1994) 3, 522-537

Thematic Article
Summary of taxa and morphological adaptations of the Desmostylia
NORIHISA AND CLAYTON E. RAY3
INUZUKA,' DARK P. DOMNING~
'Department cf Anatomy, University of Tokyo, Faculty of Medicine, Hongo, Bunkyo-ku, Tokyo 11 3, Japan,
2Department of Anatomy, Howard University, 520 W Street NW, Washington, DC 20059, USA and 3Department
of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA

Abstract Desmostylians from the Pacific coasts of Japan and North America are compared and
taxonomy and adaptations in the mammalian order Desmostylia discussed. The taxonomy is
based mainly on teeth and skulls. Feeding adaptations and habitat are discussed using teeth and
restored skeletons, respectively. In the extinct order two families, six genera and 10 species can
now be recognized. Each genus is distinguished by a set of tooth characters. The three species
of the genus Paleoparadoxia have different body sizes. Two species of the genus Desmostylus
can be distinguished on morphological differences in the cranium. As a result it is clear that both
species, D. hesperus, which had long been recognized only in North America, and D. japonicus,
which has hitherto been recognized only in Japan, inhabited both coasts of the Pacific. All genera
of the Order Desmostylia share a common and distinctive body form, interpreted by Inuzuka as
having a laterally placed limb posture. Based on this body form, the optimum habitat is thought
to have been an intertidal sandy beach. The dental morphology is variable, and derived species
show an adaptation for wear by an abrasive diet. In conclusion, we theorize that the Order
Desmostylia was adapted for a coastal habitat from its inception, with each genus adapting to a
different sort of herbivorous diet.

Key words: adaptations, Behemotops, Cornwallius, Desmostylia, Desmostylus, evolution,

Kronokotheriurn, Paleoparadoxia, taxonomy, Vanderhoojius.

INTRODUCTION
The higher classification of mammals has recently
received much reconsideration in the context of
mammalian paleontology. The Order Desmostylia
was once included in Paenungulata (Simpson 1945)
or Subungulata (Romer 1966). McKenna (1975)
erected the Mirorder Tethytheria including only
three Orders, Proboscidea, Sirenia and Desmostylia.
Domning et al. (1986), Tassy and Shoshani (1988)
and others have also included the Desmostylia in the
Tethytheria. Thewissen and Domning (1992)
grouped these three orders, plus Perissodactyla and
Hyracoidea, to make the Mirorder Pantomesaxonia;
however, they were unable to resolve the relation-
ships within the Pantomesaxonia.
Despite prolonged discussion, the taxonomic
position of the Order Desmostylia is still unsettled.
The type genus of the order, Desmostylus (Marsh
1888), was most often placed by many authors in
the Order Sirenia, though other authors placed it
among the proboscideans, multituberculates and
elsewhere, as shown in Table 1. However, because
the genus has many characters different from those
of other sirenians, it has been given increasing
taxonomic isolation: the Family Desmostylidae
(Osborn 1905), Suborder Desmostyliformes (Hay
1915) within the Order Sirenia, a Suborder Desmo-
dontia within the Order Multituberculata (Kishida
1933), or an Order Desmostyloidea (Abel 1933) in
the Subclass Multituberculata. I t finally became the
basis of the independent Order Desmostylia (Rein-
hart 1953). However, the relation between the
Order Desmostylia and other orders has never been
clarified (see Domning et al. 1986; Ray et al.
1994).
 Taxa and adaptations of Desmostylia 523
Table 1 Opinions on the taxonomic affinities of the desmostylians.
Sirenia Proboscidea Multituberculata Monotremata Ungulates Desmostylia
1888 Marsh
1891 Flower and Lydekker
1902 Osborn
1902 Yoshiwara and
Iwasaki
1905 Osborn
1906 Merriam
1910 Osborn
1914 Yatsu Abel
1915 Hay Tokunaga
1918 Matsumoto
1919 Abel
1922 Hannibal Abel Abel
1923 Hay
1925 Schlosser
1927 Sone Honda
1928 Weber
1933 Kishida Romer
1934 Tokunaga
1937 Vanderhoof Ijiri
1938 Sickenberg
1938 Ijiri
1941 Nagao
1944 Naora
1945 Simpson
1945 Romer
1951 Gregory
1953 Reinhart
1955 Colbert
1958 Dechaseaux
1961 Ijiri and Kamei
1964 Mitchell
1966 Shikama
1966 Romer
1968 Thenius
1971 Olson Kurt&
1975 McKenna
1978 Starck
1984 Inuzuka
1986 Domning et al.

In order to solve this problem, it is necessary to TAXONOMY

know the characteristics of the Order Desmostylia.
For this purpose, characters of each genus and
species belonging to the order must be set forth. In
the present paper we reconsider all known genera
and species of the order and attempt to redefine
them and identify distinguishing characters.
The abbreviations used in this paper are as
follows: GSJ, Geological Survey of Japan, Tsukuba;
LACM, Natural History Museum of Los Angeles
County, Los Angeles, California; PV, National Sci-
ence Museum, Tokyo; UCMP, University of Califor-
nia Museum of Paleontology, Berkeley; UHR, De-
partment of Geology and Mineralogy, Hokkaido
University, Sapporo; USNM, US National Museum
of Natural History, Washington, DC.
The name of the extinct mammalian order Desmo-
stylia is derived from the bundled columnar shape
of the molar cusps in the advanced taxa. Cheek
teeth of Behernotops Domning, Ray and McKenna,
1986, however, which is the most primitive known
genus in the order and has only recently been
discovered, are bunodont and do not have columnar
cusps. Therefore, this order cannot be defined by
means of molar shape. In the original description
of the genus Behemotops, the basis on which this
taxon is referred to the order is its possession of
fused roots on P, and, possibly, a postzygomatic
foramen connecting the temporal fossa with the
external auditory meatus.
524 N. Inuxuka et al.
However, Behemotops also shares some charac-
ters of its postcranial morphology in common with
advanced forms like Paleoparadoxia (Reinhart
1959) and Desmostylus. For example, they share a
flat and broad sternum, a medially twisted tibia,
and an inwardly inclined calcaneal tuber. There-
fore, these features of the postcranial skeleton
should be selected as characters defining the order.
The order is further characterized by the follow-
ing morphological features: dorsally protruding
orbits; short cranial base; long paroccipital process;
broadened muzzle with transversely aligned inci-
sors; enlargement of lower canine to form a proc-
umbent tusk; upper molars with more than five
cusps; funnel-shaped and ventrocaudally directed
epitympanic sinus; epitympanic sinus, postzygo-
matic foramen and anterior squamosal foramen
opening into temporal fossa; shortened cervical
vertebrae; compressed humeral shaft; large olecra-
non of the ulna; fused and parallel radius and ulna
in adult; concave radial and convex ulnar distal
articular surfaces; anteroposteriorly flattened fem-
oral shaft; low greater trochanter; well-developed
lesser trochanter; nearly flat patellar surface; thick
patella; marked and laterally inclined tibia1 tuber-
osity; anterolaterally inclined distal articular sur-
face of tibia; fibula much shorter than tibia.
The Order Desmostylia includes a t least two
families, containing possibly six genera and 10
species, and these can be classified as follows:
Order Desmostylia Reinhart, 1953
Family Behemotopsidae Inuzuka, 1987
Genus Behernotops Domning, Ray and
McKenna, 1986
B. proteus Domning, Ray and
McKenna, 1986
B. sp. nov. (the Ashoro I1 specimen)
Family Desmostylidae Osborn, 1905
Genus Paleoparadoxia Reinhart, 1959
P. weltoni Clark, 1991
P. tabatai (Tokunaga, 1939)
P. sp. nov. (the Stanford specimen)
Genus Cornwallius Hay, 1923
C. sookensis Cornwall, 1922
Genus Desmostylus Marsh, 1888
D. japonicus Tokunaga and
Iwasaki, 1914
D. hesperus Marsh, 1888
Genus Kronokotherium Pronina, 1957
K. brevimaxillare Pronina, 1957
Genus Vanderhoojius Reinhart, 1959
V. coalingensis Reinhart, 1959
The Order Desmostylia thus includes six nominal
genera, but Vanderhoojius and Kronokotherium
are represented by only a few specimens, and it is
possible that they are synonyms of Desmostylus.
Fossil desmostylians are only found near the
coasts of the North Pacific and from the Late
Oligocene to the late Middle Miocene. Behemotops
and Cornwallius are Oligocene and the other gen-
era are Miocene. Desmostylus, Paleoparadoxia
and Behemotops occur on both coasts of the Pacific
Ocean, Cornwallius and Vanderhoojius are known
only from the west coast of North America and
Kronokotherium is known only from the Kam-
chatka Peninsula (Fig. 1).
Each genus in the Order Desmostylia can be
distinguished by molar shape and the number of
lower incisors (Table 2). Only the most primitive,
Behernotops, has bunodont teeth; other, more ad-
vanced forms have stylodont teeth. In Paleopara-
doxia the cusp (column) retains a small diameter
and thin enamel. In Cornwallius the cusp is
thicker and has thick enamel but retains a low
crown height. The molars of Desmostylus,
Vanderhoojius and Kronokotherium have higher
crowns and more cusps than those of the other
genera.
Vanderhoojius cannot be distinguished from
Desmostylus on molar shape, but acccording to
the original description of Reinhart (1959),
Vanderhoojius has no lower incisors, in contrast to
Desmostylus, which has a pair of lower incisors.
However, the only known mandibles of Vander-
hoojius are from adult individuals, so it is possible
that this could be an age character and their
incisors had already been shed. The peculiar bone
expansion on the medial surface of the holotype
and other mandibles can also be interpreted as an
age character rather than a generic character,
since such an expansion is also seen in adult
Desmostylus but not in juveniles.
According to Domning’s opinion, a referred topo-
typic skull (USNM 214945, unpubl. data) retains a
sagittal crest, like Desmostylus japonicus (see
below), but differs from all Desmostylus in retain-
ing an anteriorly tapering zygomatic process of the
squamosal. Therefore, Vanderhoofius coalingensis
is doubtless a valid species, but whether it is
generically distinct from Desmostylus is a matter
of arbitrary definition.
Kronokotherium is based on only one specimen,
a left mandible with molars (Pronina 1957). The
lower molar is a high-crowned stylodont tooth with
six thick columnar cusps like that of Desmostylus,
but Kronokotherium differs from most Desmosty-
lus in cusp arrangement. In lower molars of Des-
mostylus, the rows of cusps are usually arranged
 Taxa and adaptations of Desmostylia 525

4 Paleoparadoxia
V Cornwallius
0 Desmostylus
0 Kronokotherium
v Vanderhoojius
.**
..10

Fig. 1 Distribution of genera of the Order Desrnostylia

in parallel and the buccal protoconid and hypoconid
tend to be situated slightly distal to the lingual
metaconid and entoconid, respectively. In Kro-
nokotherium this trend is reversed and the buccal
cusps are situated fairly mesial to the correspond-
ing lingual ones. Also, the horizontal ramus is
extremely short for its height. Therefore, Kro-
nokotherium may be generically distinct from
Desmostylus.
The upper molars of Desmostylus have eight
cusps. The problem of cusp identification in the
Order Desmostylia is which cusps correspond to the
primary trigon. Matsumoto (19 18) first attempted
to identify them, VanderHoof (1937) proposed an
alternative interpretation and Reinhart (1959)
agreed with VanderHoof. Due to lack of ancestral
forms, the homology of the primary cones was
largely speculative, as VanderHoof (1937) noted.
But now, the discovery of a primitive member of
the order, Behemotops, makes it possible to homol-
ogize the cusps, and this Inuzuka (1987) attempted
to do with the cusps not only of the upper molar of
Desmostylus, but also both upper and lower molars
and premolars of all desmostylian genera.
The upper molar of Behemotops has five cusps,
three cusps in the first row and two in the second.
The upper molar of Desmostylus has eight cusps
arranged in four rows (3.2.2.1 from mesial to
distal). So there appeared to be two possibilities:
the five cusps of Behemotops correspond to those of
the first two rows of Desmostylus, or to those of
the second and third rows and an intermediate cusp
of the first row of Desrnostylus. The latter identi-
fication has been selected (Fig. 2) based on the

Table 2 Dental characters that distinguish desmostylian genera,

Crown type Diameter of cusp Crown height Dental formula of lower teeth
~

Behemotops Bunodont Thin LOW 3.1.4.3

Paleoparadoxia Stylodont Thin LOW 3.1.3.3
Cornwallius Stylodont Thick LOW 2.1.3.3
Desmostylus Stylodont Thick High 1.1.3.3
Vanderhoojius Stylodont Thick High 0.1.3.3
526 N. Inuxuka et al.
Behemotops Paleoparadoxia
Fig. 2 Cusp identification of cheek teeth of Desmostylia. See text
following three points: (i) the intermediate cusp of
the first row of Behemotops is smaller than the
other four cusps; (ii) in Desmostylus the four cusps
of the second and third rows are larger than the
three cusps of the first row; (iii) accessory cusps or
tubercles developed on the cingulum in front of the
first row of Behemotops can be regarded as devel-
oping into the lateral cusps of the first row of
Desmostylus. In the same way, cusps of premolars
and of other genera are identified based on size and
arrangement of cusps and accessory cusps or tu-
bercles. In Fig. 2, the hypocone or hypoconid (H) is
shown as well as the trigon or trigonid. Cones of
the mesial talon are designated A, B and C from
their sizes and frequency of appearance, as are the
distal talon X and the distal talonid Y and Z.
Accessory cusps that always appear are shown by
open circles and those cuspules or tubercles that
variably appear are shown by black.
GENUS BEHEMOTOPS
The genus was proposed by Domning et al. in
1986, containing two species based on specimens
Upper
premolar
Upper
molar and
milk molar
@
Lower
premolar
Cornwallius Desmostylus Kronokotherium
from North America, with B. proteus as the type
species. Type specimens of both Behemotops pro-
teus and B. emlongi are mandibles with lower
teeth. The upper teeth of the genus are known
from a whole skeleton from Ashoro-cho, Hokkaido,
Japan found in 1981 (Inuzuka 1989b), and also
from a new North American specimen (Ray et al.
1994).
The dental formula of Behemotops is 3.1.4.3.
The upper molar has five cusps which include three
main cusps, protocone (P), paracone (Pa), and
metacone (M), plus hypocone (H) and a mesial
talon (A) (Fig. 2). Some accessory cusps or tuber-
cles may appear at the positions shown by black
circles in the figure. The lower molar has four
cusps, protoconid (P), metaconid (M), hypoconid
(H) and entoconid (E), and an accessory cusp may
appear distally. The lower premolar has two cusps,
protoconid and metaconid, and low and small distal
tubercles, corresponding to the hypoconid and en-
toconid. Besides these, some tubercles may appear
a t the three points shown by black circles in Fig. 2.
A specimen from Baja California, Mexico, UCMP
36078 (VanderHoof 1942) which has been as-
signed to Cornwallius sookensis, may belong in-

stead to the genus Behernotops, because its unworn
cusps are relatively bunodont.
In the genus Behernotops two species, B. proteus
and B. emlongi, have been described. The type
specimen of B. proteus is a small mandible of a
young individual retaining milk molars, and that of
B. ernlongi is a large mandible of an adult with
only the last molar. The last molar of B. proteus is
not fully formed or erupted, complicating the com-
parison of the two specimens. Later, a third speci-
men of the genus was discovered that had charac-
ters in common with both ‘species’, so B. emlongi
has become a synonym of B. proteus (Ray et al.
1994).
Two fossil skeletons of the genus have been
collected from the early Late Oligocene Morawan
Formation in Ashoro-cho, Hokkaido, Japan. The
first Ashoro specimen (Ashoro I) was obtained
from the lower hard shale member (about 29 Ma)
in 1976 and the second one (Ashoro 11) from the
upper tuffaceous silt member (about 25 Ma) in
1981 (Matsui & Ganzawa 1987; Saito et al. 1988).
The second specimen includes upper and lower
teeth. Compared with the three specimens from
North America, the second Ashoro specimen differs
from B. proteus in the following characters of the
teeth and mandible, and therefore seems to be a
new species.
(1) In the third upper molar of Ashoro 11, the
hypocone and metacone stand side by side and the
intermediate transverse valley between mesial and
distal rows is linear, while in that of B. proteus the
hypocone is situated distal to the metacone and
inclines distally.
(2) In the lower molars the crista obliqua is less
developed and the crown is narrow for the length
of the tooth.
(3) In the fourth lower premolar of Ashoro 11, the
protoconid has only a mesial ridge, while in that of
B. proteus it has both mesial and distal ridges. The
metaconid inclines distally and is small. The hypo-
conid and entoconid are comparatively larger.
(4) In the third lower premolar the protoconid
inclines distally and the metaconid is better devel-
oped than in B. proteus.
(5) Compared with B. proteus, the mandible of
Ashoro I1 is relatively larger and the molars are
relatively smaller.
(6) The mandibular ramus is high and its lateral
surface is deeply concave.
(7) The mandibular notch is deep.
(8) The mandibular condyle is larger, inclines
more medially and is convex sagittally and trans-
versely.
Taxa and adaptations of Desmostylia 527
The first Ashoro specimen is an adult skeleton
including the skull and 14 teeth (Inuzuka 1987). It
has primitive characteristics and a smaller skull
and body than those of Ashoro I1 despite having
molars of the same size. Therefore, Ashoro I
represents the oldest and most primitive taxon in
the order, and is probably a different genus from
Behernotops.
GENUS PALEOPARADOXIA
This genus was established by Reinhart (1959).
The desmostylian genera having low-crowned
stylodont teeth are Cornwallius (Hay 1923) and
Paleoparadoxia, but Paleoparadoxia has never
been clearly distinguished from Cornwallius
before. Reinhart (1959) noted ‘The cingulum of
Paleoparadoxia differs from that of Cornwallius
in extending more than halfway above the crown
of the tooth, completely encircling the crown,
having greater lateral thickness, and forming a far
more prominent part of the tooth.’ However, the
cingiilum of Paleoparadoxia does not always com-
pletely encircle the crown. Mitchell and Repenning
(1963) tried to distinguish three genera, Desmosty-
lus, Cornwallius and Paleoparadoxia, by shape,
height and whether or not the columns were closely
appressed, and by presence or absence and degree
of development of the cingulum. However, whether
the columns are closely appressed varies individu-
ally and cannot be used as a generic character.
Desmostylus has no cingulum but Paleoparadoxia
cannot be distinguished from Cornwallius by the
degree of cingular development. Mitchell and Re-
penning (1963) noted that columnar height was
about one to two diameters of a column in Corn-
wallius and only about half of crown height in
Paleoparadoxia. However, this difference is not
constant and hence also not a usable criterion.
Later, two skulls of Cornwallius were discov-
ered (USNM, undescribed), showing that Cornwal-
lius (Reinhart 1982: fig. 1) can easily be distin-
guished from Paleoparadoxia. Cheek teeth of
Cornwallius can be distinguished from those of the
two described species of Paleoparadoxia, P. taba-
tai and P. weltoni, using crown size, columnar
thickness and enamel thickness. However, a new
species of Paleoparadoxia, represented by the
Stanford specimen (UCMP 81302), is larger than
the other paleoparadoxian species, and its teeth are
comparable to those of Cornwallius in size. Hence,
teeth of these two genera must be distinguished
from each other by shape. Differences between
528 N. Inuxuka et al.
Cornwallius and Paleoparadoxia, therefore, will
be enumerated after this new species has been
described.
The dental formula of Paleoparadoxia was de-
scribed as 3.1.4.3 by Ijiri and Kamei (1961) but
was changed to 3.1.3.3, decreasing by one pre-
molar, by Inuzuka (1986). Determination of the
number of premolars by Ijiri and Kamei (1961) was
based on an X-ray of the diastema between two of
the lower premolars in the Izumi specimen (PV-
05601). Inuzuka (1986) re-examined the same
radiograph to try to confirm their identification, but
found no trace of an alveolus or a root in this
diastema. Based on his observations, it became
clear that there were originally only three pre-
molars (Pz-,) with a diastema between P, and P, in
Paleoparadoxia.
Three species of Paleoparadoxia, P. weltoni, P.
tabatai (Fig. 3), and the new Stanford species, are
smaller, intermediate, and larger in body size,
respectively. Because erupted teeth remain the
same size despite growth in body size, each species
can be recognized by the size of its molars. The
Stanford specimen does not preserve any cheek
tooth, but its tooth size can be estimated by the
sizes of the alveoli remaining in the left mandible.
Crown lengths of the third lower molars of these
three species are 23, 35, and 42 mm (e), respec-
tively.
According to Clark (1991), P. weltoni is very
similar to P. tabatai but differs in having smaller
Paleoparadoxia weltoni
Paleoparadoxia tabatai
Fig. 3 Crania of two species of Paleoparadoxia. Dorsal views, drawn to the
same scale: (a) P. weltoni after Clark (1991); (b) P. tabatai after ljiri and
Kamei (1961).
teeth and distally expanded incisors, and can be
distinguished from all other desmostylians by its
lack of a hypoconulid on M, and a well-developed
paraconid on P,.
GENUS DESMOSTYLUS
This genus was erected by Marsh (1888). Desmo-
stylus has characteristic high-crowned stylodont
teeth with more cusps than those of Paleopara-
doxia and Cornwallius, and no cingulum. The
additional columnar cusps of Desmostylus can be
homologized with accessory cusps or hypertrophied
cingular cusps of Behemotops or Paleoparadoxia
(Fig. 2).
In the genus Desmostylus, several species have
come and gone in the literature (Table 3). In the
United States, after the first description of D.
hesperus Marsh, 1888, D. cymatias Hannibal,
1922, and D. californicus Hay, 1923, were named
but soon became synonyms of D. hesperus
(VanderHoof 1937). American authors have since
regarded Desmostylus hesperus as the only species
of the genus. In Japan, meanwhile, after the des-
cription of D. japonicus (Yoshiwara & Iwasaki
1902; Tokunaga & Iwasaki 1914), a larger species,
D. mirabilis Nagao, 1935, and a smaller one, D.
minor (Nagao, 1937a), were named.
A ten-cusped lower molar from Japan was
named Desmostylella typica (Nagao, 1937b).
However, Reinhart, (1959), re-identified it as a
milk molar of Desmostylus. His identification was
proven to be correct by a fossil mandible with a
similar multi-cusped tooth of a young Desmostylus
(USNM 181742) from Oregon. This and other
specimens show that the fourth upper and lower
milk molars of Desmostylus have many cusps,
accounting for the pattern seen in the holotype of
‘Desmostylella’.
v The holotype of D. mirabilis (the Keton speci-
men, UHR 18466) has a second molar nearly equal
in size with a second molar of the holotype of D.
japonicus (the Togari specimen, PV-05600). That
the Keton second molar looks larger than the Togari
is attributed to greater wear (Inuzuka 1989a). The
holotype of D. minor (the Honto specimen from
Sakhalien), which was distinguished by its small
size, has the same size as the first molar of the
Utanobori specimen (GSJ-F7743) (Inuzuka 1988).
Therefore, these three ‘species’ from Japan are not
distinguishable on molar morphology.
In general, although American authors have
regarded the Japanese specimens as representing
 Taxa and adaptations of Desrnostylia 529
Table 3 History of the specific names in the genus Desmostylus from North America and Japan.

Specimens from North America Specimens from Japan

Author Tooth Oregon skull Skull Togari skull Keton Honto Yuda Utanobori
YPM11900 USNM8191 USNM187310 PV-05600 UHR 18466 tooth tooth GSJ-F7743
1888 Marsh D. hesperus
1914 Tokunaga D. japonicus
& Iwasaki
1915 Hay D. hesperus D. watasei
1922 Hannibal D. cymatias
1924 Hay D. hesperus
1935 Nagao D. japonicus D. mirabilis
1937 Nagao D. minor Desmostylella
1937 Vanderhoof D. hesperus D. hesperus D. hesperus
1937 Ijiri D. hesperus D. japonicus D. japonicus D. japonicus
1954 Ijiri Desmostylella
1959 Reinhart D. hesperus D. hesperms D. hesperus D. hesperms D. hesperus
1959 Yabe D. hesperus D. japonicus D. japonicus D. japonicus
1961 Ijiri & Kamei D. japonicus D. mirabilis
1966 Shikama D. hesperus D.h. japonicus D.h. japonicus D.h. japonicus D.h. japonicus
1982 Inuzuka D. mirabilis
1990 Inuzuka D. hesperus D. japonicus D. japonicus D. hesperus D. hesperus

D. hesperus (e.g. VanderHoof, 1937; Reinhart
1959), Japanese authors have been apt to regard
them as a distinct species, D. japonicus (Ijiri 1937;
Yabe 1959), or subspecies, D. hesperus japonicus
(Shikama 1966).
The reason why Desmostylus from Japan were
regarded as the same species as D. hesperus from
North America is that characters that clearly dis-
tinguish them from each other cannot be found in
the molar shape. In order to solve the problem, it is
necessary to study the range of variation of many
molars of Desmostylus from California and to
compare fossil crania from Japan with those from
North America. So, Inuzuka measured 320 cheek
teeth in the UCMP and LACM and compared seven
fossil skulls from North America with three skulls
from Japan.
Tooth class identification of isolated teeth of
Desmostylus is difficult because the individual vari-
ability in size and shape is great. As Desmostylus
has three molars, if the genus includes only one
species, three peaks should appear in a histogram
of crown sizes, even if the size variation of each
tooth class is large. In Inuzuka’s data, however, at
least four peaks appeared. Also, a second molar in
one large mandible is comparable in size with a
third molar in another individual. These facts sug-
gest the presence of two morphotypes, one large
and one smaller, in D. hesperus.
In the smaller type the second molar is nearly
equal in size to the third molar. In the fourth milk
molars, however, two size categories are not dis-
tinguishable (Figs 4, 5). These facts show that
both types have young of the same size, and the
smaller type completes its growth sooner. There-
fore, the two types in D. hesperus are considered
to represent sexual morphs rather than distinct
species.
Three fossil skulls of Desmostylus have been
described from Japan, the Togari specimen of D.
japonicus, the Keton specimen of ‘D. mirabilis’
(Ijiri & Kamei 1961), and the new Utanobori
specimen. The former two are adult and incomplete
specimens lacking the anterior or posterior part.
The Utanobori specimen is complete, but of a
young individual with the first molars worn. Hence,
morphological differences among these skulls can-
not with confidence be attributed to specific differ-
ences or age variations.
Skulls and mandibles of Desmostylus from Ore-
gon in the USNM include each tooth class from
DP3 to M3 (Table 4). Among them an old individ-
ual with M3 (USNM 26134) has a narrow and high
rostrum very similar to the Utanobori specimen
with M1 from Japan, and another type (USNM
187310) has a wide and low rostrum similar to
that of the Togari specimen and has a sagittal
crest. These skulls and cheek teeth are small and
the canines are also small or lacking. Moreover,
there are two types of skulls with DP4, one type
with a sagittal crest (USNM 181744) and another
without it (USNM 186887, 214743). Therefore,
these seem to be different species rather than
sexual morphs.
Consequently, the Utanobori specimen is identi-
fied as D. hesperus, because it belongs to the same
type as the Oregon skull described by Hay (1915).
The other type belongs to the same group as the
530 N. Inuzuka et al.

60-

50-

40-

5 -
2 :
30-

20- Fig. 4 Upper cheek teeth of Desmosty-

i
D4 /us from North America and Japan. 04:
DP4 of both types; M1, M2, M3: larger
type; m l , m2, m3: smaller type; Kt: Keton
l o [ , , ~ , ( , , , , ( , , , , , , , , , ( , , , I ( , , , , ( , , , , , specimen; LM: Los Angeles County Mu-
20 30 40 50 60 70 80 90 seum specimen; Ut Utanobori specimen.
Length (0)America; ( X ) Japan.

50-

40-

5 -
? .
5
30-

Fig. 5 Lower cheek teeth of Desmostylus

from North America and Japan. 04: DP4 of
20-
both types: M1, M2, M3: larger type; m l ,
m2, m3: smaller type; Kt: Keton specimen;
LM: Los Angeles County Museum speci-
men; To: Togari specimen; UC: University
101, , , , ( , , , I ( , , , . , , , , , ( , , , , , , , , , ( , , , , , of California Museum of Paleontology
20 30 40 50 60 70 80 90 specimen; Ut: Utanobori specimen: (0)
Length America; ( X ) Japan.

Table 4 Character distribution in crania of Desmostylus specimens.

Specimen Dental stage Canine Rostrum Narial opening Nasal rise Frontal Sagittal
width crest
USNM186887 D3-4 - - - Strong Wide Absent
USNM214743 D4 - - - - Wide Absent
USNM181744 D4 - - - - Narrow Present
USNM8 1 91 M1 - - - Strong Wide Absent
Utanobori GSJ-F7743 M1 Absent High Anteriorly Strong Wide Absent
Togari PV-05600 M1-2 Present LOW Dorsally Weak - I

Keton UHR18466 M2 - - - - Wide Absent

USNM187310 M2-3 Present LOW Dorsally Weak Narrow Present
USNM26 134 M3 Absent High Anteriorly Strong Wide Absent
USNM214944 M3 Absent LOW Dorsally Weak Narrow Present
USNM18 1737 M3 - - - - Wide Absent

Desmostylus hesperus
Desmostylus japonicus
Fig. 6 Crania of two species of Desmostylus. Left lateral views, not drawn to
the same scale. (a) D. hesperus after Inuzuka (1988); (b) D. japonicus after
Yoshiwara and lwasaki (1902).
Togari specimen, so it should be called D. japoni-
cus (Fig. 6 ) . That is, Desmostylus includes a t least
two species, D. hesperus and D. japonicus, which
can be distinguished from each other not by size
and shape of the molars but by characters of the
skull and mandible shown in Table 5 . The Keton
specimen, which lacks a sagittal crest, probably
represents D. hesperus.
In conclusion, both species of Desmostylus in-
habited the east and west Pacific coasts in the
Miocene. Fossils of Desmostylus from Japan are
Middle Miocene and ones from America are Early
and Middle Miocene, but most specimens from both
areas are represented only by teeth and they
cannot be assigned to species.
ADAPTATI0NS
SKELETAL RESTORATION
Among the six genera of the Order Desmostylia,
reasonably complete fossil skeletons of three gen-
era, Behemotops, Paleoparadoxia and Desmosty-
lus, are known. These postcranial remains make it
possible to restore the skeleton, by which the
habitat and mode of locomotion of these animals
can be hypothesized. However, the body form and
locomotion of desmostylians have always been con-
Taxa and adaptations of Desmostylia 531
Table 5 Difference of characters between Desmostylus hespe-
"us and D. japonicus.
D. hesperus D. japonicus
Maxillary rostrum High and narrow Low and wide
Direction of narial Rather anterior Rather dorsal
opening
Rise in nasal part Strong Weak
Frontal width Wide Narrow
Sagittal crest Absent Present
Mandibular ramus High and short Low and long
troversial, as vividly demonstrated in the illustra-
tions compiled by Inuzuka (198213).
Inuzuka (1984) restored a skeleton of Desmosty-
lus. His methodology included comparative anat-
omy, functional morphology and fossil posture as
preserved taphonomically, and is designed for sci-
entifically restoring extinct animals with no living
relatives.
His method of comparative anatomy has two
purposes. One is to compare individual bones of
Desmostylus with those of other mammals to find
characters peculiar to the animal in question. The
other is to compare skeletons of living mammals
with each other, especially ungulates, among which
Desmostylus belongs, to find rules of skeletal com-
position. As bones of Desmostylus display some
morphological characteristics peculiar to the lin-
eage, they do not follow all these skeletal rules. So,
some rules to adopt are selected by means of
functional morphology. The method is to restore
the position and degree of development of muscles
based on tuberosities of the bones most character-
istic of Desrnostylus and to determine the positions
of bones in which the muscles can work most
efficiently. The skeletal posture restored by these
two methods was corroborated by examining the
posture displayed by a fossil skeleton lying on the
bedding plane. This study led to the following
conclusions.
Many bones of the postcranial skeleton of des-
mostylians have peculiar shapes, different from
those of all other mammals. Their many character-
istics can be explained on a reasonable anatomical
basis only if one single restored posture is postu-
lated, that which is characterized by laterally ex-
tended stylopodia.
In typical terrestrial mammals with an inferiorly
extended humerus, the fore part of the body is
mainly supported by serratus muscles running like
a hammock from each side of the thorax to the
serratus surface of the scapula (Fig. 7). However,
changing the position of the humerus from beneath
the trunk to laterally, the main weight supporter
532 N. Inuxuka et al.
pectdrales
Desmostylians
radius
n running from the gluteal surface to the greater
Typical land mammals
Fig. 7 Postulated mode of muscle attachment and forelimb posture, after
lnuzuka (1984): (a) cranial views; (b) left lateral views.
naturally changes from the serratus to the pectoral
muscles. That is because this change in direction of
the humerus shifts the position of the scapula from
parasagittal to horizontal. In a posture with later-
ally extended humerus, the role in body support of
the serratus muscles decreases, because the muscle
fibers run more horizontally. On the contrary, pec-
toral muscles running from the sternum to the
humerus would play the main role in body support
instead of the serratus, because the position of the
humerus relative to the trunk is higher in the
lateral limb posture than in the inferior, and the
muscle fibers of pectoral muscles would run more
nearly vertically. The main muscle working as a
body supporter would increase in mass and expand
its bony attachments, resulting in the development
of larger tuberosities. The flat and broad sternum
of Desmostylus can thus be accounted for as
an attachment area for hypertrophied pectoral
muscles, and the ill-developed serratus surface of
the scapula is attributable to reduced serratus
muscles.
In typical terrestrial mammals with inferiorly
extending forelimbs, the antebrachial skeleton is in
pronation so that the toes point forward. As a
result, the radius and ulna run across or fuse in
crossing each other. If, however, the humerus
inclines laterally during bending of the elbow, the
arrangement tends to supinate the antebrachial
skeleton so that the radius and ulna run in parallel.
As the two bones of Desmostylus fuse in parallel,
in the lateral limb posture with laterally extending
elbow the toes point forward. Otherwise, in the
inferior limb posture with the humerus extending
beneath the trunk, the toes point outward. Thus, if
a posture with a laterally extending humerus is
postulated, peculiar morphological features of the
sternum, scapula, radius and ulna in Desmostylus
are reasonably explainable.
As well as the forelimb, many peculiar features
in the hindlimb of Desmostylus are also explain-
able, if a posture with a laterally extending femur
is postulated. In typical mammals with an inferi-
orly extending femur, the hind part of the body is
mainly supported by the gluteus medius muscle
trochanter (Fig. 8). Changing the direction of the
femur from beneath the trunk to laterally, the main
weight supporter changes to the iliopsoas or adduc-
tor muscles, because in this posture the body
weight causes the femur to be abducted. The
adductor muscles originate from the portion of the
hip bone ventral to the acetabulum. The relatively
great breadth of this portion of the bone in Desmo-
stylus can be interpreted as indicating well-
developed adductor muscles. The iliopsoas muscle
inserts on the lesser trochanter of the femur. The
large and distally extended lesser trochanter in
Desmostylus is also explainable since in mammals
with a lateral-type limb, more power is needed for
adduction or flexion of the femur than in mammals
with an inferior-type limb.
The patella in desmostylians is large and thick
for their body size. In a lateral limb posture with
the femur extended laterally, an animal must al-
ways flex the knee joint even in standing, because
the knee faces anterolaterally. This demands devel-
opment of the quadriceps femoris muscle to extend
the knee joint. In this way, the morphological
characteristics of the patella are explainable from
the body posture and muscle function.
In typical land mammals with inferior-type
limbs, the knee and ankle joints are uniaxial hinge
joints. In general, the mode of movement in a joint
is more conservative than the size or proportions of
a bone. Hence, a bone is more easily identified by
 Taxa and adaptations of Desrnostylia 533

Fig. 8 Postulated mode of muscle attach auuuctor

calcis
mint and hindlimb posture in (a) desmo- triceps surae
stylians and (b) typical land mammals, after
lnuzuka (1984), posterolateral views.

an epiphysis than by a diaphysis. In mammals with

lateral-type limbs like Desmostylus, the knee faces
anterolaterally, so the toes also point anterolater-
ally, unless the shaft of the tibia twists inward. In
fact, the toes do point forward due to the medially
twisted tibia1 shaft. This reflects the fact that
desmostylians, with lateral-type limbs, were de-
rived from animals with inferior-type limbs, and by
twisting the tibia managed to retain anteriorly
directed toes in the course of evolution.
In typical mammals with inferior-type limbs,
when the calcaneus is articulated with the talus,
the calcaneal tuber protrudes posteriorly perpen-
dicular to the axis of the ankle joint. However,
desmostylians characteristically have a medially
inclined calcaneal tuber. This character was re-
garded only as an oddity (Shikama 1966), but can
be interpreted more reasonably, considering the
muscle function, as follows. The gastrocnemius Desmostylians
muscle, which originates from the posterior surface
of the distal femur, inserts on the point of the Fig. 9 Comparison of limb postures. Reptiles and mammals after Lesser-
tisseur and Saban (1967); desmostylians after lnuzuka (1984).
calcaneal tuber. In an animal with lateral-type
limbs in which the knee joint is flexed and pro-
trudes anterolaterally, the gastrocnemius muscle Another view of desmostylian posture and loco-
must run obliquely a longer distance than in motion has been expressed by Domning, but has
inferior-type limbs, if the calcaneal tuber keeps the not yet been published. It combines features of the
orientation of the inferior type. In order for the reconstructions made by Shikama in 1974 and
muscles to pass straight between the femur and Kamei in 1975 (see Inuzuka 198213). Domning
the tip of the calcaneus as they do in the inferior proposes that the terrestrial posture of desmo-
type, the calcaneal tuber must incline inward in the stylians deviated from that of ‘normal’ ungulates
lateral limb-type animals. much less than has been supposed, and that des-
Thus if a posture with laterally extending femur mostylian osteology is compatible with a modified
is postulated, peculiar morphological features of inferior-type limb posture. They probably had a
the pelvis, femur, patella, tibia, talus and calcaneus quadrupedal stance with the body well off the
are explainable on a reasonable anatomical basis. ground and the limbs more or less under the body;
As a result, Desmostylus was restored by Inuzuka a strongly arched spine and steeply inclined pelvis;
in a posture with laterally extending stylopodia like slightly abducted elbows and more strongly ab-
a reptile despite its being a mammal (Fig. 9). ducted knees; and a digitigrade foot posture with
534 N. Inuzuka et al.
an extended but not hyperextended wrist, the front
toes pointing anterolaterad and the hind toes point-
ing forward. Domning will defend this alternate
interpretation in a future publication.
Most peculiarities of the desmostylian skeleton
are in fact paralleled in one or another of various
large, slow-moving terrestrial mammals such as
ground sloths and chalicotheres. Like these, des-
mostylians apparently had skeletons well suited to
supporting their weight on the hindquarters. In
Domning’s view, this was most likely necessary
while the animals clambered slowly over very un-
even intertidal areas, and crossing such areas en
route to and from the water. Locomotion in the
water probably resembled that of polar bears, with
the forelimbs providing the principal means of
propulsion and the hindlimbs used for steering.
EVOLUTIONARY PATTERNS
The peculiar osteological characters of Desmosty-
lus on which Inuzuka’s restoration is based are as
follows: paired flat sternebrae; parallel radius and
ulna fused in supinated position; pelvis with ace-
tabulum in the middle; femur with well-developed
lesser trochanter; tibia medially twisted; fibula far
shorter than tibia; calcaneal tuber medially inclined
relative to the axis of the ankle joint.
This set of unique characters is common to
Desmostylus, Paleoparadoxia and Behemotops, so
it can be regarded as characteristic of the Order
Desmostylia. If Inuzuka’s hypothesis that Desmo-
stylus had a crocodiloid posture is correct, the
same interpretation will hold for all desmostylians.
This body form peculiar to the Order Desmo-
stylia is characterized by having the contact points
of the feet well away from a very low center of
gravity (Fig. 10). Such a posture sacrifices body
support and walking efficiency to stability. Because
Fig. 10 Restored skeleton of Desmostyhs
by lnuzuka (1 984), based on the Keton speci-
men from south Sakhalien housed at Hokkaido
University
this direction of morphological change goes against
the trend of terrestrial vertebrate evolution, an
unusual habitat can be inferred: a habitat where
stability of the body would be most advantageous,
that is, where moving water would tend to over-
turn the body. The most likely habitat is an inter-
tidal sandy beach.
The animals of this order seem to have inhabited
such places from the behemotopsian stage onward,
because even Behemotops had already taken on
this special body form. The fact that Paleopara-
doxia and Desmostylus have a similar body form
suggests that they also inhabited a similar habitat
and walked with the same mode of locomotion.
On the other hand, desmostylian teeth diversified
greatly from the ancestral morphotype seen in
Behemotops to some divergently derived types seen
in Paleoparadoxia, Desmostylus and so on, as
mentioned above (Fig. 11). This fact suggests the
evolution of diverse feeding habits, which cannot
yet be fully elucidated. However, the hypsodont
teeth of Desmostylus, a t least, seem to indicate
adaptation to a more abrasive diet, probably one
that contained grit mixed with plant material
scooped from the sea floor or sea shore (see discus-
sion in Domning et al. 1986). In conclusion, we
speculate that the animals of the Order Desmostylia
colonized a unique nearshore-marine habitat, and
subsequently adapted to diverse herbivorous diets.
ACKNOWLEDGEMENTS
Inuzuka would like to thank Drs W. A. Clemens,
J. H. Hutchison, L. G. Barnes, M. Kato and Y.
Tomida for giving him access to specimens of
desmostylians for comparison. He is also pleased to
acknowledge the considerable assistance of Dr
G. V. Shkurkin during his stay in Berkeley.

Fig. 11 Characterization of molar teeth in
various desmostylian genera. Upper row: oc-
clusal view of right upper molar. Middle row:
lateral view of molar. Lower row: schematic
cross section of tooth germ and enamel. Behernotops Paleoparadoxia
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