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Number 3171, 10 pp., 3 figures

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MUSEUM OF NATURAL HISTORY

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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.

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

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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-

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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

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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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