A new species of Chungchienia (Tillodontia, Mammalia) from the Eocene of Lushi, China

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A New Species of Chungchienia
(Tillodontia, Mammalia) from the
Eocene of Lushi, China

MINCHEN CHOW,! JINGWEN WANG,' AND JIN MENG!”

ABSTRACT

A new species of Chungchienia from the Eocene
of Lushi, Henan, is described. Based on dental
features such as a distinctive hypoconulid lobe on
m3, the new material decisively reallocates
Chungchienia, previously believed to be either an
edentate, a tillodont, or a taeniodont, to the order
Tillodontia. The new species shows that Chung-
chienia is a highly specialized tillodont, having
rootless hypsodont cheek teeth with only the labial

surface covered with enamel. Shared similarities
such as a long diastema before p3 suggest that
Chungchienia is more closely related to the tro-
gosine genus Jillodon from North America than
to other tillodonts. Phylogenetic relationships of
tillodonts indicate possible multiple dispersal
events in a two-way exchange of tillodonts be-
tween North America and Asia during the Eocene.

INTRODUCTION

Tillodontia is an extinct group of early Ter-
tiary mammals known in North America,
Europe and Asia. Five tillodont genera have
been reported from North America: Estho-
nyx Cope, 1874; Trogosus Leidy, 1871; Til-
lodon Gazin, 1953; Megalesthonyx Rose,

1972; and Azygonyx Gingerich, 1989. Two
genera were reported from Europe: Plesies-
thonyx Lemoine, 1891, and Franchaius
Baudry, 1992. Several taxa from Asia have
also been included in Tillodontia, including
Adapidium Young, 1937; Basalina Dehm and

' Institute of Vertebrate Paleontology and Paleoanthropology, Beijing.
2 Department of Vertebrate Paleontology, American Museum of Natural History.

Copyright © American Museum of Natural History 1996

ISSN 0003-0082 / Price $1.60

2 AMERICAN MUSEUM NOVITATES

Oettingen-Spielberg, 1958; Kuanchuanius
Chow, 1963a; Lofochaius Chow et al., 1973;
Meiostylodon Wang, 1975; and Dysnoetodon
Zhang, 1980. More recently, Ting and Zheng
(1989) restudied Interogale and Anchilestes,
which were previously assigned to Anagalida
(Huang and Zheng, 1983; Qiu and Li, 1977),
and found them to be primitive tillodonts.

Conventionally, tillodonts have been re-
lated one way or another to arctocyonid con-
dylarths (Gregory, 1910; Gazin, 1953; Van
Valen, 1963; Rose, 1972; Szalay, 1977), al-
though McKenna (1975) placed the Tillo-
dontia in the Mirorder Eparctocyona that ac-
commodates Arctocyonia, Tubulidentata,
Dinocerata, Embrithopoda, and Artiodac-
tyla. With discoveries of early pantodont and
tillodont material from China, the hypothesis
that tillodonts are akin to pantodonts was
proposed and became more popular (Chow
and Wang, 1979; Gingerich and Gunnell,
1979; Lucas and Schoch, 1981; Stucky and
Krishtalka, 1983; Lucas, 1993). Although
Lucas believed that the similarities between
tillodonts and pantodonts are mostly among
derived forms of these two groups, a tillo-
dont-pantodont relationship may still be sug-
gested by the possible sister taxon of tillo-
donts, i.e., Deltatherium (Chow and Wang,
1979; Lucas, 1993), because McKenna (1975)
considered Deltatherium to be a pantodont
(but see Lucas, 1993). However, while Lucas
considered Deltatherium the sister group to
Tillodontia instead of being included in the
order, McKenna (personal commun.) now
prefers to put the genus in the order on the
basis of a premolar loss, although Lucas
thought that the lack of Pl/pl was a con-
verged feature in Deltatherium and Estho-
nychidae.

Based on their new analysis of Interogale
and Anchilestes, Ting and Zheng (1989) con-
cluded that tillodonts probably have affinities
with anagalids, an alternative hypothesis that
calls for future testing to determine whether
or not these two genera are actually tillodonts
(Lucas, 1993). Regardless of their relation-
ships with pantodonts or anagalids, the fossil
record points to an Asian origin of tillodonts
and their dispersal from Asia to North Amer-
ica across Beringia during the late Paleocene
and early Eocene (Krause and Maas, 1990).

While previous studies focused on Asian

NO. 3171

taxa that may represent roots and stems of
tillodonts that are still obscured in phyloge-
netic bushes of early mammals, we report
here a new species of Chungchienia that cer-
tainly qualifies as a new twig in the tillodont
clade, based on material collected by J. Wang
and Y. Tong from the Lushi Basin, Henan.
Chungchienia was proposed by Chow (1963b)
and has long been considered either an eden-
tate or a taeniodont, but was recently placed
in Tillodontia (Schoch, 1986). Because of the
new light shed on these relationships by the
Lushi\material described here, the tillodont
afinity of Chungchienia is confirmed. This
genus is believed to be the most specialized
tillodont known. Chungchienia further di-
versifies Asian tillodonts and indicates the
possibility of two-way interchange of this or-
der between Asia and North America during
the Eocene, given the derived similarities
shared by this genus and the North American
Tillodon.

ORDER TILLODONTIA MARSH, 1875
FAMILY ESTHONYCHIDAE COPE, 1883
SUBFAMILY TROGOSINAE GAZIN, 1953
Chungchienia Chow, 1963

INCLUDED SPECIES: C. sichuanica and C.
lushia, new species.

REVISED DIAGNOSIS: Large tillodont; all
known teeth (2, p3 and/or p4, m1-3) root-
less; 12 extremely large, gliriform, and with
enamel covering the entire anterior surface;
a long diastema before p3; all cheek teeth
hypsodont (both root and crown hypsodon-
ty), with enamel restricted to the entire labial
surface of the tooth body that extends deeply
into the mandible; flat grinding surfaces of
teeth, owing to early wear of cusps; p3 and/
or p4 columnar, showing no division of tri-
gonid and talonid; trigonid and talonid of
m1l-—2 forming two lobes (tooth double-co-
lumnar) with the trigonid lobe slightly nar-
rower than the talonid lobe; crown of m3
trilobed (tooth tri-columnar) owing to a large
hypoconulid.

Chungchienia lushia, new species

HOoLotyPe: A left 12, a right p3 (or p4), a
left m1, a right m2 and m3 (V10805; Institute

1996 CHOW ET AL.: A NEW SPECIES OF TILLODONT 3

Fig. 1.

of Vertebrate Paleontology and Paleoanthro-
pology, Beijing, P.R.C.) (fig. 1, 2).
LOCALITY AND AGE: Specimens were col-
lected from a pit in a small valley about 300
m southwest of Xiaowan village, Lushi
County, Henan province. The locality is be-
tween the Xiejiagou and Mengjiapo sites re-
ported by Chow et al. (1973). Age determi-

Society of Vertebrate Paleontology
401 N. Michigan Ave.

Chicago, IL 60611-4267
Phone: (312) 321-3708 Fax: (312) 245-1085
E-mail: SVP@SBA.COM

Lingual (upper) and labial (lower) views of the second left lower incisor (V10805).

nation of the Xiaowan site is difficult based
on scattered fossils. However, given its oc-
currence between Mengjiapo and Xiejiagou
in which middle Eocene taxa such as Hy-
rachyus and Uintatherium were found (Tong
and Wang, 1980, 1981), beds yielding C. /u-
shia are probably within sediments of age
equivalent to the Irdin Manha Fm. (middle

4 AMERICAN MUSEUM NOVITATES NO. 3171

2 3 4 5 6 7

Fig. 2. Top to bottom: crown, lingual, and labial views of a right p3 (or p4), a left m1, a right m2
and m3 (V10805).

1996

Eocene) of Inner Mongolia or slightly youn-
ger.

ETYMOLOGy: Species name adopted from
the county Lushi.

D1AGNOsIs: Differs from C. sichuanica in
being larger and p3 (p4) less transversely
compressed; enamel band on p3 (p4) about
70% broader than that of C. sichuanica and
its posterior border with the dentine not
marked by groove; dentine consisting of a
homogeneous labial and rugose lingual por-
tions, contrasting with the two-layered den-
tine in C. sichuanica (see Chow, 1963b: fig.
1).

DESCRIPTION: The left 12 is complete. It is
a long, gliriform, and robust tooth, with the
enamel wrapping around the anterior and
most of the labial surfaces throughout the
entire tooth body. The enamel layer becomes
thinner from ventral to labial side. The tip
of the tooth is chisel-shaped, convex anteri-
orly, and flat posteriorly. The labial surface
of the tooth is uneven in that the lateral side
is a longitudinal ridge about 10 mm higher
than the lingual side. From the tip to the
proximal end, the incisor is 260 mm long.

The p3 (p4) is columnar and rootless, arch-
ing anterolaterally. The anterolabial surface
is covered with a longitudinal enamel band,
measuring 15 mm wide near the crown and
13 mm near the base, throughout the entire
tooth body. Most of the tooth surface is ex-
posed dentine. The enamel layer is decorated
with transverse as well as longitudinal fine
lines. The enamel is brown in color, in con-
trast with the white dentine, and its anterior
and posterior boundaries with the dentine are
sharply defined. The dentine is dense and is
filled with minute cracks, which may be post-
mortem fractures. There is a groove on the
lingual side of the tooth, indicating an orig-
inally double-rooted tooth. The crown out-
line is roughly square, bearing a wear surface
that slopes posteriorly.

The m1 is both root and crown hypsodont,
therefore rootless, and bowed laterally and
leaning forward. It tapers gradually from the
crown to the root end. The trigonid and tal-
onid form the anterior and posterior lobes,
with the posterior one being slightly wider
and longer than the anterior one. The enamel
is present only on the labial surface, whereas
the lingual side of the tooth is bare dentine

CHOW ET AL.: A NEW SPECIES OF TILLODONT 5

with a rough surface. The trigonid and tal-
onid portions of the tooth are separated by a
deep longitudinal groove on both the lingual
and labial sides; therefore, the tooth is dou-
ble-columned. Along the bottom of the labial
groove, a narrow band of white dentine is
exposed, so the enamel is discontinuous
therein. The trigonid and talonid bear no re-
maining cusps but form grinding surfaces,
with the trigonid being slightly higher than
the talonid. The dentine consists of two areas:
labially it is dense and uniform, whereas lin-
gually it is rugose and filled with minute
cracks. Loss of enamel may be responsible
for this morphology, but it seems unlikely
that this is a preservational artifact. There is
no distinct boundary between these two por-
tions of dentine. The wear surface of the la-
bial side of the crown is smooth and lower,
in contrast to the rough and higher wear sur-
face of the lingual portion. This suggests that
the labial dentine, although it appears to be
denser, is not any harder than the lingual den-
tine.

The m2 is similar to m1 in general mor-
phology but is larger in all dimensions. The
dentine of the two molars is stained with light
purple color. The m3 differs considerably
from m1-2 in having a large hypoconulid
lobe; therefore, the m3 is triple-columnar with
the crown outline being much longer but nar-
rower than that of m1 and m2. The trigonid
is higher than the talonid, although the
boundary between them is not distinct. The
dentine is totally white, indicating that the
condition of the m3 is slightly different from
that of the other color-stained cheek teeth; it
is probably from a different individual as well.
The enamel surrounding the hypoconulid
column extends to the lingual side of the tooth,
and the lingual surface is bare dentine oth-
erwise. Unlike the other cheek teeth that ta-
per from the crown toward the root, m3 flares
slightly in the opposite direction. The dentine
on the lingual portion of the tooth is rougher
in texture, but not so much as that in m1 and
m2. The labial enamel is continuous near the
crown from the trigonid column to hypocon-
ulid column, and is increasingly separated by
broader dentine bands toward the base. In all
cheek teeth, the enamel layer from crown end
to root end shows no significant change in
thickness.

6 AMERICAN MUSEUM NOVITATES

Measurements of V10805 (mm):

Length Width
12 43 28
p3 (p4) 19 17
ml 24 24
m2 28 26
m3 43 22
DISCUSSION

The link between C. sichuanica and C. lu-
shia is p3 (or p4), which is the only tooth that
permits comparison. They are considered to
be the same tooth, either a p3 or a p4. They
are similar in general morphology, including
the distribution of enamel, curvature of the
tooth, and wear pattern of the crown. On the
other hand, they have several differences, as
listed in the diagnosis. Judged from its po-
sition relative to the diastema, the only pre-
served tooth in the Sichuan specimen should
be a premolar rather than m1, as previously
identified by Chow (1963b). Chow also ob-
served a longitudinal trough along the ventral
side of the mandible and suspected that it
was for housing a large chisellike incisor. Such
an incisor is confirmed by the Lushi speci-
mens.

Considering the single tooth to be a root-
less, cylindrical ml, Chow (1963b) placed
Chungchienia into Edentata when he first
proposed this taxon based on the specimen
collected from Sichuan, Henan (=Honan).
Romer (1968: 219) was surprised by the
abrupt appearance ofan Asian edentate, sup-
posedly a ground sloth, and commented:
“Both in time and space we are far, far re-
moved from any situation in which we would
expect a ground sloth.’’ Then he suggested
that Chungchienia is a taeniodont. Although
Chow et al. (1973) still regarded Chungchi-
enia an edentate, an alternative and more
recent concept endorses Romer’s view that
Chungchienia is a taeniodont (Ding, 1979;
Hoffstetter, 1982; Russell and Zhai, 1987;
Rose and Emry, 1993). A substantial phy-
logenetic analysis of Chungchienia since its
publication was given by Ding (1987), who
discussed the phylogenetic position of Er-
nanodon, a possible edentate from South
China. Because the enamel band, narrow but
distinctive, does not fit the overall edentate

NO. 3171

dental evolution characterized by enamel de-
generation, and because a long diastema and
other features in the lower dentition do not
compare with those of any taeniodont, Ding
concluded that it is equally difficult to place
Chungchienia into either Edentata or Tae-
niodontia on the basis of available material.
Schoch (1986), however, explicitly assigned
Chungchienia to Tillodontia, which is sup-
ported by the Lushi specimens.

The tillodont identification of the Lushi
specimens was suggested by M. C. McKenna
(personal commun.) in a brief examination
of the specimens and has proven to be cor-
rect. The decisive reallocation of Chungchi-
enia into Tillodontia on the basis of the new
material resolves the uncertainties that re-
mained from its affiliation with either eden-
tates or taeniodonts. Quite convincingly, as
shown in the Lushi specimens, the cheek teeth
of Chungchienia are not those of an edentate.
The enlarged second lower incisor, the dou-
ble-columned and triple-columned lower
molars consequent to prominent trigonid and
talonid lobes, and the unilateral distribution
of enamel on the labial surface of cheek teeth,
are typical of neither edentates nor taenio-
donts. In this regard, edentates and taenio-
donts are more similar to each other in hav-
ing columnar cheek teeth in which the tri-
gonid and talonid are no longer recognizable.

The lower molars of tillodonts have a
U-shaped trigonid with a metastylid and a
somewhat rounded talonid basin; the cheek
teeth are unilaterally high crowned even in
primitive forms. The talonid basin of m3 is
elongate because of the enlarged hypoconu-
lid, and a third lobe is universally present in
tillodonts. As a highly derived form,
Chungchienia possesses derived features that
are not present in other tillodonts. However,
most of these can be interpreted as advanced
in a morphological spectrum ranging from
primitive to derived features. For instance,
the hypsodont teeth and the unilateral dis-
tribution of the enamel on cheek teeth are
already initiated in more primitive forms such
as Tillodon and Trogosus. As stated by Gazin
(1953: 33): “The external hypsodonty of the
lower teeth as compared with the more
brachydont appearance of the lingual wall is
characteristic of all the tillodonts, but be-

1996

comes increasingly distinctive in the later
forms.” It must be pointed out that in all
tillodonts except Chungchienia, the hypso-
donty applies to tooth crown only. In those
forms, cheek teeth are primitively rooted and
the enamel does not extend into the jaw bones.
Nonetheless, the outline of the tooth crowns
of Chungchienia, particularly that of m3, re-
mains little changed, which reveals its rela-
tionships. The trilobed crown shape of m3
in Chungchienia is most comparable to that
of large tillodonts: Ti/lodon and Trogosus. The
wear pattern of the cheek teeth is such that
the labial side of dentine is more worn than
the lingual side, which also resembles that of
other tillodonts.

In conventional classification, Tillodontia
consists of a single family, Esthonychidae
Cope, 1883, which was divided into two sub-
families (Gazin, 1953): Esthonychinae and
Trogosinae. This classification was supported
by Rose (1972) who pointed out that Es-
thonychinae is characterized by large but
rooted second incisors and that Trogosinae
possesses large gliriform second incisors that
grow from persistent pulp. However, a large
and rooted second incisor appears to be prim-
itive for tillodonts; therefore it is inadequate
for diagnosing a taxon within the group. Re-
cent phylogenetic analyses confirmed the
monophyly of Trogosinae and showed the
paraphyly of Esthonychinae (Baudry, 1992;
Lucas, 1993; fig. 3). Lucas redefined Trogo-
sinae by including Megalesthonyx and Adap-
idium in the subfamily, based not on gliri-
form second incisors as advocated by Rose
but on a set of characters that allows a broad-
er conception of Trogosinae including Me-
galesthonyx and Adapidium. Inclusion of
primitive Asian forms, such as Lofochaius
and Meiostylodon in Esthonychidae and Jn-
terogale and Anchilestes in the Tillodontia,
was considered to be questionable (Ginger-
ich, 1989; Lucas, 1993). Although phyloge-
netic relationships of tillodonts have been ex-
plored (Baudry, 1992; Lucas, 1993), there is
no comprehensive taxonomy encompassing
all these taxa. In addition, problems exist
concerning the taxonomic status of some til-
lodont taxa such as Kuanchuanius and Ple-
siesthonyx (e.g., Baudry, 1992; Lucas, 1993).

Chungchienia is readily assignable to the

CHOW ET AL.: A NEW SPECIES OF TILLODONT 7

subfamily Trogosinae in any case, because it
possesses large, gliriform second incisors and
reduced talonids on p3-p4 (Gazin, 1953;
Rose, 1972; Lucas, 1993). Chungchienia is
similar to Tillodon and Trogosus in having
greatly enlarged body size, deepened man-
dibular symphysis, and in the position of the
mental foramen. Chungchienia is further-
more similar to Tillodon in several aspects:
a greatly elongate rostrum suggested by the
large second incisor, a narrow M3 reflected
by the narrow m3 (Lucas, 1993), and a dis-
tinct diastema before p3. As Gazin (1953: 49)
noted: “‘Lower canine and p2 are spaced and
well separated from p3” in Tillodon. The lon-
ger diastema in Chungchienia, as shown in
the Sichuan specimen, may have resulted
from loss of p2 and/or p3. In Tillodon, p3 is
small relative to p4. This tooth may be lost
in Chungchienia, which is why we consider
the tooth of C. sichuanica to be either a p3
or a p4. The canine and p2 are small and the
canine is somewhat vertical in Jillodon, dif-
fering from the condition in C. sichuanica,
in which a large alveolus in the mandible
suggests a sizable and procumbent canine
(Chow, 1963b).

Among Asian forms, Kuanchuanius is be-
lieved to be a member of the subfamily Tro-
gosinae (Rose, 1972; Baudry, 1992), although
Lucas (1993) further regarded it as a junior
synonym of Jrogosus. In spite of its taxo-
nomic position, Kuanchuanius, like Trogo-
Sus, appears more primitive than 7i//odon in
being smaller and having a narrow gap be-
tween p2 and p3. Another possible tillodont
of China is represented by an enlarged incisor
(i2) from the middle Eocene Guangzhuang
Formation of Shandong. This specimen was
called ““Unbestimmabare Ungulatenzahne”’
by Zdansky (1930) and is believed to be
Chungchienia (Lucas, personal commun.),
which, if confirmed, broadens the distribu-
tion of the genus.

Given the derived similarities of Chungchi-
enia and North American Tillodon, it is pos-
sible that Chungchienia is a derivative of
North American forms rather than of Asian
tillodonts, although it can be argued that those
similarities are a result of parallel evolution.
The relationship shown in figure 3 suggests
that migration of tillodonts between Asia and

8 AMERICAN MUSEUM NOVITATES

NO. 3171

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Fig. 3. Two recent phylogenetic hypotheses of tillodonts: A, modified from Lucas, 1993; and B, from
Baudry, 1992, with Chungchienia inserted in both. Lucas’ hypothesis employs Deltatherium as the sister
taxon of Tillodontia, whereas Baudry’s includes all tillodont genera known, without providing an out-
group. Features diagnosing various nodes were provided by Lucas (1993: table 14.2) and Baudry (1992:

fig. 12).

North America may have occurred more than
once and that a dispersal from North Amer-
ica to Asia during the middle or late Eocene
is possible.

ACKNOWLEDGMENTS

We thank Y. Tong of the Institute of Ver-
tebrate Paleontology and Paleoanthropology,
Beijing, for generously providing the Lushi

specimens. We acknowledge S. Ding, S. G.
Lucas, M. C. McKenna, K. D. Rose, R. H.
Tedford and Y. Wang for comments that im-
proved the manuscript. Chow’s research was
partially supported by the Frick Laboratory
Endowment Fund from the Department of
Vertebrate Paleontology, AMNH. Meng’s re-
search is supported by the Frick Postdoctoral
Fellowship and an NSF grant (DEB-
9508685).

1996

CHOW ET AL.: A NEW SPECIES OF TILLODONT 9

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