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Implantation and Replacement of Bird Teeth<br />

ABSTRACT<br />

Study of the teeth of the Mesozoic birds Hesperornis, Parahesperornis,<br />

Ichthyornis, Cathayornis, and Archaeopteryx provides<br />

new evidence for avian tooth implantation and replacement. Birds<br />

share with crocodilians and, to a lesser extent, mammals, a complex<br />

mode of tooth implantation, with deep sockets walled lingually<br />

by the dentary, maxilla, or premaxilla. These walls crowd<br />

the replacing teeth so mat early in ontogeny the teeth migrate labially<br />

and continue their development under the crown of their predecessor.<br />

They thus form a vertical tooth family, as opposed to the<br />

horizontal tooth family found in dinosaurs and most other tetrapods.<br />

Birds, crocodilians, and mammals have root cementum on<br />

their teeth and presumably attach teeth to the socket with periodontal<br />

ligaments. The sockets in mammals and presumably in<br />

birds are formed by the outside of the periodontal sac, whereas<br />

cementum is deposited by the inside of the sac. Bird teeth are initially<br />

formed in a groove, and ontogenetically the sockets (in<br />

socket-forming species) form first at the front of the jaw. Socket<br />

formation then proceeds posteriorly, as in crocodilians. Young<br />

dinosaurs have the lingual side of the jaw around the teeth open, so<br />

that the roots are exposed. The sockets form around dinosaur teeth<br />

as bone of attachment, which is probably the same periodontal<br />

bone that forms sockets in mammals, crocodilians, and birds. The<br />

sites of new tooth formation extend lingually within the so-called<br />

"special foramina" that separate the interdental plates. The interdental<br />

plates represent the surrounding attachment bone and are<br />

similar to the attachment bone in pleurodont lizards. In fact, dinosaurs<br />

might be characterized as having a superpleurodonty that<br />

results in sockets.<br />

Introduction<br />

In our previous paper on avian teeth (Martin et al., 1980), we<br />

called attention to numerous features shared by crocodilians<br />

and birds but not found in theropod dinosaurs. At that time, we<br />

were unaware of how fundamentally different the whole dental<br />

Larry D. Martin andJ.D. Stewart<br />

Larry D. Martin, Natural History Museum and Department of Ecology<br />

and Evolutionary Biology, University of Kansas, Lawrence, Kansas<br />

66045, United States. J.D. Stewart, <strong>Lo</strong>s Angeles County Museum,<br />

900 Exposition Boulevard, <strong>Lo</strong>s Angeles, California 90007, United<br />

States.<br />

295<br />

system is in crocodilians and dinosaurs and how similar the<br />

dentition is in crocodilians and birds.<br />

The characteristic tooth morphology of crocodilians and<br />

birds includes a flattened, unserrated crown that becomes constricted<br />

as it approaches the crown/root juncture. The tooth is<br />

narrow at this point and then expands into a cement-covered<br />

root at least as broad as the crown and usually broader. <strong>Res</strong>orption<br />

begins as circular to oval pits in the lingual side of the root,<br />

and the replacement tooth has most of its formative history beneath<br />

the tooth that it will replace (below or above depending<br />

on lower or upper dentition). This morphology is found in all<br />

of the Triassic and Early Jurassic crocodilians that we have<br />

been able to examine. For instance, this tooth form is very<br />

clearly shown in acid-prepared specimens from the Liassic<br />

(Early Jurassic) marine crocodilian Pelagosaums in The Natural<br />

History Museum, <strong>Lo</strong>ndon, collections.<br />

ACKNOWLEDGMENTS.—We are grateful to Alan Charig,<br />

Cyril Walker, and Angela Milner of The Natural History Museum,<br />

<strong>Lo</strong>ndon (formerly the British Museum (Natural History);<br />

BMNH) for access to specimens; P. Wellnhofer (Bayerische<br />

Staatssamlung, Munich) generously shared access to specimens<br />

and insights, as did P. Currie (Tyrrell Museum, Drumheller)<br />

and G. Edmund (Royal Ontario Museum, Toronto). C.<br />

Bennett and John Chom read the manuscript, and we especially<br />

thank Zhonghe Zhou for helpful suggestions. The photography<br />

staff at BMNH made the excellent ultraviolet photographs, and<br />

the drawings are by Mary Tanner (Craneview Studio, North<br />

Platte, Nebraska) and A. Aase (University of Kansas Natural<br />

History Museum). Funding was provided by National Science<br />

Foundation grant DEB 7821432 and National Geographic Society<br />

grant 2228-80.<br />

Discussion<br />

Aside from the nature of the teeth themselves, their mode of<br />

implantation in vertebrates also has proven to be useful in<br />

working out relationships. The earliest reptiles had acrodont<br />

teeth, as are found in the labyrinthodont amphibians and the<br />

captorhinomorph reptiles (Figure lA). In the earliest diapsid<br />

reptile known (Petrolacosaums), this condition has been modified<br />

(Reisz, 1981) by the upward (in the lower dentition) exten-

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