New middle Pleistocene Galictini (Mustelidae, Carnivora) from the Matsugae cave

deposits, northern Kyushu, West Japan

Author(s): Shintaro Ogino and Hiroyuki Otsuka
Source: Paleontological Research, 12(2):159-166. 2008.
Published By: The Palaeontological Society of Japan
DOI: http://dx.doi.org/10.2517/1342-8144(2008)12[159:NMPGMC]2.0.CO;2
URL: http://www.bioone.org/doi/full/10.2517/1342-8144%282008%2912%5B159%3ANMPGMC
%5D2.0.CO%3B2

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 Pleistocene
Paleontological Research, vol. 12, no. 2, pp. 159–166, June 30,extinct
2008 Galictini from Japan 159
© by the Palaeontological Society of Japan

New middle Pleistocene Galictini (Mustelidae, Carnivora)

from the Matsugae cave deposits, northern Kyushu,
West Japan
SHINTARO OGINO1 and HIROYUKI OTSUKA2
1
Systematics & Phylogeny Section, Primate Research Institute, Kyoto University. Kanrin Inuyama City, Aichi 484-8506, Japan
(e-mail: gotolabo@hotmail.com)
2
Division of Nature System Sciences, Kagoshima University. 1-21-35 Korimoto, Kagoshima City, Kagoshima 890-0065, Japan
(e-mail: hmj4254@hyper.ocn.ne.jp)

Received October 3, 2007; Revised manuscript accepted January 5, 2008

Abstract. In the early 20th century, Tokunaga, followed by Naora, reported a Middle Pleistocene mamma-
lian fauna from the Matsugae limestone cave deposits in northern Kyushu, West Japan. Within this fauna
Naora (1968) described a new extinct species of otter (Lutra nipponica). Recently, we have identified new,
well preserved dentitions of this mustelid in their original collection. We suggest that the definition of L.
nipponica given by Naora should be emended, and that the specimens belong to a new genus of the tribe
Galictini (Mustelidae, Carnivora), namely, Oriensictis gen. nov., according to the following characteristics:
short muzzle; transversely expanded P4; and anteroposteriorly expanded M1. The extinct Chinese otter L.
melina (Pei, 1934) from the Middle Pleistocene cave deposits of Choukoutien locality 1 is also included in
Oriensictis based on the characteristics of M1 and a short mandible. These two occurrences of this genus
are the first records of East Asian galictines, and this suggests that the Galictini might have been extensively
distributed over areas in the Northern Hemisphere until the Middle Pleistocene.

Key words: Carnivora, Galictini, Japan, Mustelidae, Pleistocene.

Introduction

The Matsugae mammal fauna, named by Tokunaga

(1931), was recognized from middle Pleistocene lime-
stone cave deposits in northern Kyushu, West Japan
(Figure 1). The fossil specimens were collected and re-
ported by Tokunaga (1931) and Naora (1944) during the
early 20th century, but they were damaged in Tokyo dur-
ing the Second World War. However, some specimens
from the collection escaped destruction. To date, 12
fossil mammals have been identified in their collections,
which are kept in the National Museum of Japanese
History (Ogino and Otsuka, 2005).
Naora (1968) described a fossil mustelid in this col-
lection as an extinct otter. Based on the morphological
characteristics of the elongated P4 and the smaller M1, Figure 1. Location map of the Matsugae cave on the
northern part of Kyushu Island, West Japan.
he identified a row of upper cheek teeth (NMJH-N0181)
as belonging to “Lutra nipponica,” a species related to
the living common otter (L. lutra). However, we identi- described by Naora (1968) should be emended and
fied new well preserved dentitions of this fossil mustelid the species included in the tribe Galictini (Mustelidae,
during a recent study. The newly identified mustelid Carnivora) based on the tooth morphology. The tribe
specimens suggest that the description of L. nipponica Galictini includes the two living genera Galictis and
160 Shintaro Ogino and Hiroyuki Otsuka
Lyncodon found in Central and South America and
eight extinct Late Tertiary genera: Cernictis, Trigonic-
tis, Sminthosinis, and Lutravus from the New World,
and Trochictis, Pannonictis, Enhydrictis, and ?Melodon
incertum from the Old World (Baskin, 1998; Pilgrim,
1932).
In this report, we assign two extinct species of the
East Asian otter — “Lutra nipponica” (Naora, 1968)
from the middle Pleistocene Matsugae cave and “Lutra
melina” (Pei, 1934) from locality 1 of Choukoutien,
North China — to Oriensictis gen. nov. based on their
tooth characteristics. This is the first report on the distri-
bution of Galictini in the Pleistocene East Asia.
Abbreviations
BMNH = formerly British Museum of Natural His-
tory, now The Natural History Museum, London, United
Kingdom; I.G.F. = Istituto di Geologia dell’Università
di Firenze, Italy; IVPP = Institute of Vertebrate Pa-
leontology and Paleoanthropology, Academia Sinica,
China; MHNL = Muséum d’Histoire Naturelle de Lyon,
France; NMJH-N = Naora collections stored at National
Museum of Japanese History, Japan; PMUU = Paleonto-
logical Museum of Uppsala University, Sweden; RGIH
= Geological Institute of Hungary; UL1 = Universite
Lyon 1, France; UM-Ida = University of Michigan,
Museum of Paleontology.
Systematic paleontology
Order Carnivora Bowdich, 1821
Family Mustelidae Fischer, 1817
Subfamily Galictinae Reig, 1956
Tribe Galictini Baskin, 1998
Genus Oriensictis gen. nov.
Type Species.—Oriensictis nipponica (= Lutra nip-
ponica Naora, 1968).
Etymology.—From the Latin oriens, oriental; and ictis,
weasel-like mammals.
Age and Range of Distribution: Middle Pleistocene
on Kyushu Island, Japan, and middle Pleistocene fissure
deposits in locality 1 of the Sinanthropus (Homo pekin-
ensis) site at Choukoutien, China.
Diagnosis.—Facial region is short and broad; canine
extends straight, and deep groove present on the ante-
rior surface; P1 absent; anterior root of P3 projects to
the buccal side; P4 has strong paracone with a furrow
adjoining the parastyle; lingual shelf extended out-
➡ Figure 2. Oriensictis nipponica (Naora, 1968) Ogino and Otsuka. a and b, NMJH-N0181 (Holotype), a left maxilla fragment with
canine and cheek teeth (P3-M1): a, labial view; b, occlusal view. c and d, NMJH-N0182, a maxilla with left canine, right canine and right
cheek teeth (P3-M1): c, labial view; d, occlusal view. e–g, NMJH-N0183, a mandible fragment with right i1–i3, right canine and left m1: e,
occlusal view; f, labial view; g, buccal view. Scale bar is 2 cm.
ward with a well distinguished conical cusp; protocone
undeveloped; differs from Galictis, Lyncodon, and
Sminthosinis due to its large size; differs from Cernictis
due to loss of p4 posterior accessory cusp; differs from
Lutravus due to loss of P1; differs from Pannonictis and
Trigonictis due to the posteriorly expanded hypocone
shelf; differs from Enhydrictis due to the distinct notch
between the metaconule and the hypocone shelf on M1;
mandible is moderately robust and shorter than the other
galictines; three premolars are rather crowded; the ante-
rior roots of p2 and p3 are located buccally.
Included species.—Oriensictis melina (Pei, 1934);
Oriensictis nipponica (Naora).
Oriensictis nipponica (Naora) n. comb.
(Figure 2a–g)
Lutra nipponica Naora, 1968, p. 3, figs. 1, 2
Holotype.—Specimen Reg. No. NMJH-N0181; partial
left maxilla with canine, P3, P4, and M1 (Figure 2a, b).
Referred materials.—Specimen Reg. No. NMJH-
N0182; partial maxilla with broken left canine, right P4,
and broken right M1, (Figure 2c, d). Specimen Reg. No.
NMJH-N0183; mandible with right i1, i2, i3, canine,
and left m1 (Figure 2e–g).
Horizon and Locality.—The first ossiferous bed of
the Matsugae limestone cave deposits (Naora, 1944),
exposed in the Matsugae Quarry in Kishi, Moji district,
Kitakyushu City, Fukuoka Prefecture, Japan (33°52’N,
130°57’E).
Description.—Specimen Reg. No. NMJH-N0181.
NMJH-N0181 is the holotype of “Lutra” nipponica (O.
nipponica) (Naora, 1968). The specimen is probably
slightly deformed due to compaction.
The upper canine extends straight. A deep vertical
groove is present on the anterior surface of the canine,
and its horizontal section is concave. The groove mea-
sures 0.9 mm at its maximum depth and extends to the
dentin layer.
P1 is absent. P2 possesses two roots, judged based on
observation of its alveoli. The buccal part of P3 is buried
in the alveolar bone by physical pressure during sedi-
mentation. P4 is transversely expanded and larger than
M1. From the occlusal view, it exhibits a right-triangular
shape. The anterobuccal paracone is taller than any other
cusp. It has a furrow adjoining the parastyle. The pos-
terobuccal metacone is undeveloped. The protocone is
the smallest cusp. The posterolingual part of P4 broadly
Pleistocene extinct Galictini from Japan 161
162 Shintaro Ogino and Hiroyuki Otsuka
extends posteriorly and has a well distinguished conical
cusp (hypocone).
M1 comprises four main cusps with reduced peri-
cone. The anterior part curves inward, and the anterior
cingulum (ac) is reduced. The protocone is the highest
and most robust cusp. The paraconule is as high as the
protocone but less robust. The paracone is sharper and
larger than the metacone. The parastyle expands antero-
buccally. A well developed metaconule is positioned
along the posterior part of the metacone. The hypocone
shelf is nearly as wide as the paracone-metacone shelf.
There is a sharp notch between the metaconule and the
hypocone shelf, and the notch is considered to be a de-
rived characteristic in O. nipponica.
Specimen Reg. No. NMJH-N0182. The referred ma-
terial (NMJH-N0182) has an extremely short muzzle.
Upper tooth row (C–M length) is short. The palatal
width between the upper canines on opposite sides (C–C
distance) measures 19.5 mm. The fragment of the basal
part of the zygomatic arch is robust.
The upper incisors have fallen out, and only the al-
veoli are preserved. The alveolus for I3 is the largest in
the incisor row.
The canine of NMJH-N0182 is slightly larger than
that from NMJH-N0181.
The upper premolars are very crowded. P1 is absent,
and P2 has two roots. P3 possesses a sharp edge and
2 pointed accessory cusps, and the anterior root of P3
projects into the buccal side. P4 is larger than M1, and
exhibits a right-triangular shape in occlusal view. The
anterobuccal paracone is taller than any other cusp. The
paracone has a furrow that adjoins the parastyle. The
posterobuccal metacone is undeveloped. The protocone
is the smallest cusp. The posterolingual part of P4 ex-
tends broadly outward with a well distinguished conical
cusp (hypocone), which is slightly larger than the proto-
cone.
The morphological characteristics of M1 are almost
identical to those of specimen NMJH-N0181. The ante-
rior part of M1 is preserved and curves inward, and the
anterior cingulum is reduced without a distinct cone;
the protocone is the highest and most robust cusp; the
paraconule is slightly lower than the protocone and less
robust; the paracone is sharper and larger than the meta-
cone; the parastyle expands anterobuccally.
Specimen Reg. No. NMJH-N0183. NMJH-N0183 is
considered to be derived from the same individual as
NMJH-N0182. The mandible is very short and robust.
The height of the horizontal mandible below p4 mea-
sures 15.3 mm. Double mental foramina on the buccal
surface exist beneath p2. In lateral view, the ventral mar-
gin of the mandible is nearly straight.
All the lower incisors are single-cusped; i3 possesses
a buccal cingulum; i1 and i2 are equal in size, and i3 is
larger than the other incisors.
The lower canine forms a hooklike shape and is
covered with shallow furrows, among which the labial
furrow is the deepest and probably occludes with the
buccal cingulum of i3. The cross section is elongated
anteroposteriorly.
The lower premolars are rather crowded. The crowns
of all the lower premolars are damaged, and only their
roots are preserved. Judging from the remaining roots,
p1 was absent, while p2 possesses two roots and lies im-
mediately behind the lower canine. The anterior root of
p3 projects to the buccal side.
The m1 has an elongated trigonid and a slightly re-
duced talonid with rounded marginal cusps, including
the hypoconid and the entoconid. The tallest cusp is the
paraconid on the trigonid, and the metaconid is placed
slightly behind the paraconid. The transverse width of
m1 is short and a distinct cingulum was observed on the
buccal side. The m2 is recognizable only by the pres-
ence of a single alveolus.
Oriensictis melina (Pei, 1934) n. comb.
(Figure 3l)
Lutra melina Pei, 1934, p. 76, fig. 20, pls. 11–12.
Melodon melina (Pei, 1934) Teilhard de Chardin and Leroy,
1945, p. 27, fig. 15.
Materials.—No holotype was designated by Pei
(1934). The syntypic specimens comprise an isolated
right M1 (IVPP C/C. 667); left mandible with p3, p4,
and m1 (IVPP C/C. 668); left mandible with p2, p3, p4,
and m1 (IVPP C/C. 669); right mandible with p3, m1,
and m2 (IVPP C/C. 670A); and right mandible with m1
(IVPP C/C. 670B).
Horizon and Locality.—Middle Pleistocene fissure
deposits from the Sinanthropus site at Choukoutien,
China (Pei, 1934).
Emended diagnosis.—M1 comprises four main cusps
with reduced pericone and is larger than the M1 in
Oriensictis nipponica; a rounded notch is present be-
tween the metaconule and the hypocone shelf on M1.
The lower premolars are crowded. In occlusal view m2
appears externally circular and projects to the buccal
side.
Description.—M1 is strongly worn, but the cusp
morphology remains observable. In occlusal view, M1
presents its reduced pericone, which differs from that in
Melodon majori and ?Melodon incertum. The parastyle
is buccally elongated. The anterolingual expansion of
the anterior cingulum is weak. The posterior surface of
M1 between the metaconule and the hypocone shelf is
margined by a rounded notch; however, it is less devel-
oped than that of O. nipponica. The horizontal mandible
 Pleistocene extinct Galictini from Japan 163

Figure 3. Occlusal shape of P4 and M1 of twelve species of living and fossil mustelids which discussed in this study. a, Trigonictis
idahoensis; b, Sminthosinis bowleri; c, Pannonictis pliocaenica; d, Enhydrictis galictoides; e, Enhydrictis ardea (MHNL-20.161912); f, Tro-
chictis depereti (UL1-252); g, Melodon majori (PMUU, M3813); h, ?Melodon incertum (PMUU, M3814); i, Galictis vittata (USNM281481);
j, Lyncodon patagonicus (BMNH 3.11.5.13); k, Oriensictis nipponica (NMJH-N0181); l, Oriensictis melina (IVPP C/C. 667). a and b are
modified from Bjork (1970); c and d are modified from Ficcarelli and Torre (1967); g and h are modified from Zdansky (1924). Asterisk
means horizontally turned over from the original figure. Scale bar is 1 cm.

is very short, robust, and nearly straight. Discussion

According to Pei (1934), the thickness of the man-
dible below the p4 is approximately 16 mm on IVPP C/C.
668, 14 mm on IVPP C/C. 669, 12.9 mm on IVPP C/C.
670A, and 13.4 mm on IVPP C/C. 670B.
The p1 is absent. The p2 possesses two roots, and
the alveolus for the posterior root is bifurcated. The p2
is missing on IVPP C/C. 668. The anterior roots of p2
and p3 project to the buccal side. The p4 has a single
main cusp without a posterior accessory cusp but with a
broader posterior shelf behind the main cusp.
The m1 differs from that of Old World Galictini. It
does not have a distinct cingulum on the lingual side,
and it has a wide talonid basin. The protoconid is the
highest cusp and is located in front of the metaconid; the
paraconid is separated from the metaconid by a deep fis-
sure; the talonid margin exhibits a clear basin shape with
a hypoconid and an entoconid; the entoconid exhibits a
slight lingual expansion.
Naora (1968) first described the holotype of “Lutra
nipponica” (= Oriensictis nipponica, NMJH-N0181) as
representing an extinct otter based on the similarity of
its M1 morphology to that of Lutra. However, in the
present study, Oriensictis is defined as a new genus in
the Galictini and differs from the Lutrini in that it has
transversely wide P4, anteroposteriorly long M1 with a
developed hypocone and reduced pericone, and reduced
distinct cingulum on the lingual side of m1. “Lutra”
melina (Pei, 1934) also is placed in Oriensictis based on
the characteristics of M1 and the lower teeth.
Figure 3 shows P4 and M1 from the 10 mustelid
species discussed in this study. The late Miocene to
Pleistocene New World galictine mustelids Trigonictis
idahoensis (Figure 3a) and Sminthosinis bowleri (Figure
3b) are similar to Oriensictis in that they have a deep
furrow on the labial surface of the upper canines, innu-
merable furrows on the lower canines, and the occlusal
164 Shintaro Ogino and Hiroyuki Otsuka
feature of P4. However, the P4 of these two genera pos-
sess a conical protocone, and their lingual shelves are
more slender than in Oriensictis. Pannonictis pliocae-
nica (Figure 3c) is similar to Oriensictis with regard to
the occlusal shape of P4; however, its M1 has a reduced
hypocone and the metaconule has been lost. Enhydrictis
galictoides (Figure 3d) and E. ardea (Figure 3e) differ
from Oriensictis in that they lack a furrow on the ante-
rior ridge of P4. Further, E. galictoides also differs from
Oriensictis because its M1 displays an orbital shape in
the occlusal view. Based on dental features such as a
transversely expanded P4, reduced metacone on M1,
and transversely narrow m1, the Miocene European
Trochictis depereti (Figure 3f) is considered to be a
primitive galictine related to the newly proposed Orien-
sictis. However, T. depereti differs from Oriensictis in
that it lacks a posterolingual cone on P4 and has a lon-
ger row of teeth. Zdansky’s (1924) ?Melodon incertum
(Figure 3g) from the late Miocene to Pliocene in Shanxi,
China, also appears to have a deep labial furrow on the
upper canines, a posterolingual cone on P4, and a uni-
cusp on p4; however, it differs from Oriensictis as it
possesses P1 and an enlarged M1 similar to that in
Melini. Pilgrim (1932) stated that “the character of the
dentition, the wide zygomatic arches, the broad and
low face, and the prominent postorbital processes lead
one to infer that the Chinese genus was not far removed
from Enhydrictis”. However, ?M. incertum has the char-
acteristics of a typical omnivore-like tooth row, such
as Melini, with a long muzzle, small P4, and enlarged
M1. Melodon majori (Figure 3h) also possesses typical
omnivore-like teeth. Modern galictine mustelids Galictis
vittata (Figure 3i) and Lyncodon patagonicus (Figure 3j)
differ from Oriensictis in that they have a concave pos-
terior margin and have lost the metaconule on m1.
Figure 4 shows the ratios of P4 width (P4W)/P4
length (P4L) and M1 width (M1W)/M1 length (M1L).
The analysis confirmed that there is an obvious differ-
ence in the form of P4 and M1 between Oriensictis,
other galictines, and Melodon. Trochictis depereti and
?Melodon incertum exhibit high P4W/P4L ratios, while
Trigonictis idahoensis, Sminthosinis bowleri, Pannon-
ictis pliocaenica, Enhydrictis galictoides, and E. ardea
exhibit low ratios. The ratio of M1W/M1L is high for T.
idahoensis, S. bowleri, P. pliocaenica, and E. galictoi-
des. On the other hand, E. ardea, M. majori, ?M. incer-
tum, O. nipponica, and O. melina exhibit low ratios. T.
depereti exhibits a moderate value for the M1W/M1L
ratio. This analysis indicates that the P4 and M1 ratios
for Oriensictis resemble those for E. ardea, M. majori,
and ?M. incertum.
Pei (1934) first described “Lutra melina” (Oriensictis
melina in the present paper) as a new species belonging
Figure 4. Comparison of the P4 and M1 measurement ratios
of ten species of fossil mustelids. P4 width to P4 length is marked
by squares and M1 width to M1 length by circles. Specimen
numbers and measurements are listed in Table.
to the tribe Lutrini, based on an isolated M1 and four
lower cheek teeth. Later, Teilhard de Chardin and Leroy
(1945) placed it in the genus Melodon, an extinct Mio-
cene Melini, and named it M. melina.
However, “Melodon” melina should be transferred to
Oriensictis based on its dental characteristics. The genus
Oriensictis differs from Melodon (M. majori and ?M.
incertum) in that it lacks P1, the anteroposterior length
of M1 is reduced, the lower premolars are crowded, m1
without the buccal cingulum is narrower, and there is
a completely rounded talonid on m1. Furthermore, O.
melina differs from O. nipponica in that it possesses a
larger M1 that does not have a distinct posterobuccal
notch (Figure 3i and j).
The abovementioned derived characteristics tend to
indicate that the teeth of Oriensictis have acquired a cut-
ting function with a concomitant loss of robustness. For
effective occlusive function, the convex I3 corresponds
to the posteriorly concave lower canine, forming a sharp
cutting surface similar to the carnassials, and the upper
and lower canines provide well developed blades. Thus,
the teeth characteristics of Oriensictis indicate that this
genus was specialized in cutting soft tissue.
Mammals with a remarkably well developed furrow
on the canines are not rare. For example, several pri-
mates, including Old World monkeys (Cercopithecidae)
and gibbons (Hylobatidae) have well developed furrows
on their upper canines, while the South American coati
(Nasua; Procyonidae) has well developed furrows on
the lower canines. All these mammals are mainly fru-
givores. Oriensictis has furrows on both the upper and
lower canines; however, it has relatively well developed
P4 trigon and m1 trigonid, which are the characteristics
typically found in carnivorous mustelids.
The sharp tooth blades imply that the teeth of Orien-
 Pleistocene extinct Galictini from Japan 165

Table 1. Measurements of teeth of Oriensictis. Data of O. melina are taken from Pei (1934). All measurements are in mm. †: width of alveolar.

Oriensictis nipponica I3W CL CW P2L P2W P3L P3W P4L P4W M1L M1W C-M1 L
NMJH-ND181 Left – 7.74 4.00 – – 5.45 – 11.20 8.21 8.02 10.07 –
NMJH-ND182 Right – – – – – 6.72 3.93 11.12 8.26 – 10.59 36.53
Left 2.70† 8.09 5.81 – – – – – – – – –
cL cW m1L m1W tridL taldL c-m2L
NMJH-ND183 Right 10.43 6.42 – – – – –
Left – – 13.23 5.83 4.72 5.64 47.24
Oriensictis melina I3W CL CW P2L P2W P3L P3W P4L P4W M1L M1W C-M1 L
IVPP C/C. 667 Right – – – – – – – – – 9.3 11.7 –
p2L p2W p3L p3W p4L p4W m1L m1W m2L m2W
IVPP C/C. 668 Left – – 5.9 4.0 6.9 4.2 14.0 6.2 – –
IVPP C/C. 669 Right 4.5 3.0 6.0 3.6 7.2 3.7 13.7 6.4 – –
IVPP C/C. 670A Right – – 5.5 3.8 – – 12.5 5.8 3.8 4.0
IVPP C/C. 670B Left – – – – – – 12.3 5.5 – –

Table 2. Ratio of P4 width / P4 length and M1 width / M1 length of extinct Galictini. Data of Trigonictis idahoensis
and Sminthosinis bowleri are taken from Bjork (1970), Pannonictis pliocaenica are taken from Kormos (1931), Enhydrictis
galictoides are taken from Ficcarelli and Torre (1967), Melodon majori and ?Melodon incertum are taken from Zdansky
(1924), Oriensictis melina are taken from Pei (1934). Measurements are in mm.

P4L P4W P4W/P4L M1L M1W M1W/M1L Specimen number

Trochictis depereti 10.34 7.53 0.73 7.09 10.57 1.49 MHNL, 252
Trigonictis idahoensis 11.35 7.03 0.62 6.9 11.52 1.67 UM-Ida, V49728, V53273
Sminthosinis bowleri 8.43 5.39 0.64 5.0 8.37 1.67 BMNH, M. 29242
Pannonictis pliocaenica 13.2 8.3 0.63 6.8 12.8 1.88 RGIH, No. Ob/3594
Enhydrictis galictoides 11.5 7.0 0.61 6.0 10.0 1.67 I.G.F. 6089
Enhydrictis ardea 10.42 6.17 0.59 6.82 8.52 1.25 UL1, 20.161912
Melodon majori 8.6 6.3 0.73 9.2 10.6 1.15 PMUU, M3813
?Melodon incertum 9.4 6.8 0.72 8.3 10.8 1.30 PMUU, M3814
Oriensictis nipponica 11.20 8.21 0.73 8.02 10.07 1.26 NMJHN-0181
Oriensictis nipponica 11.12 8.29 0.75 – 10.59 – NMJHN-0182
Oriensictis melina – – – 9.3 11.7 1.26 IVPP C/C. 667

sictis nipponica might not be adapted for dealing with
hard tissues such as bone.
Kawamura (1991 and 1998) and Konishi and
Yoshikawa (1999) discussed the immigration of land
mammals from the Asian continent to the Japanese Is-
lands during the Pleistocene. They proposed two periods
of land connection between the Japanese Islands and the
Asian continent during the middle Pleistocene. Accord-
ing to them, one of the middle Pleistocene land connec-
tions was formed at 0.63 Ma (southern Chinese Stego-
don stage) and the other at 0.43 Ma (northern Chinese
Palaeoloxodon stage). The northern Kyushu Matsugae
mammal fauna is comparable to the northern Chinese
Palaeoloxodon fauna in elements such as rodents,
rhino, deer, dhole, badger, and large macaque (Naora,
1958 and Ogino and Otsuka, 2005). The two East Asian
galictine species (Oriensictis nipponica and O. melina)
described in the present paper also have a close morpho-
logical relationship with each other, and it is conceivable
that O. nipponica is a representative of the mammalian
fauna that migrated from northern China to the Japanese
islands during the age of the second land connection at
approximately 0.43 Ma.
Conclusion
The records of Galictini from East Asia suggest a new
aspect to our knowledge of the tribe Galictini, which
might have become extinct during the Pleistocene in the
Palaearctic region. To date, ?Melodon incertum (Zdan-
sky, 1924) from Shanxi (middle Miocene) was the only
known galictine from East Asia (Pilgrim, 1932). Orien-
sictis nipponica and O. melina described in this study
suggest that during the Middle Pleistocene, the Galictini
were distributed not only in the New World but also ex-
tensively in areas of the Northern Hemisphere.
166 Shintaro Ogino and Hiroyuki Otsuka
Acknowledgements
We thank the following parsons for loans of speci-
mens and assistance while visiting collections in their
care; Hideji Harunari (National Museum of Japanese
History), Robert Fisher (National Museum of Natural
History, U.S.A.), Didier Berthet (Muséum d’Histoire
Naturelle de Lyon), Anne-Marie Bodergat, Abel Prieur
(Universite Lyon 1), Jerry Hooker (The Natural His-
tory Museum, London) and Jin Chang-Zhu (Institute
of Vertebrate Paleontology and Paleoanthropology,
Chinese Academy of Science). We are grateful to Naoki
Kohno (National Museum of Nature and Science,
Tokyo), Jong-Deock Lim (Seoul National University),
and Mieczysław Wolsan (Polish Academy of Science)
for constructive comments on an earlier draft of this
paper. We also thank Ross MacPhee (American Museum
of Natural History, U.S.A.) and the reviewers for mak-
ing constructive suggestions to improve the manuscript.
This research is supported by the Sanshu Club Overseas
Research Grant (2004) and the Sasakawa Scientific
Research Grant from the Japan Science Society (2005).
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