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NUMBER 89 299<br />

would seem unlikely at the outset because the enantiornithine<br />

birds of China have Hesperornis-like teeth, almost certainly establishing<br />

this structure as primitive for birds. Examination of<br />

the <strong>Lo</strong>ndon specimen of Archaeopteryx shows typical avian<br />

structure, with a waisted crown, expanded root, and oval replacement<br />

pits (Figure 3A-C). Elzanowski and Wellnhofer<br />

(1995:42) deny the presence of expanded roots in Archaeopteryx<br />

in spite of the fact that such roots are clearly shown in<br />

Wellnhofer (1988, pl. 8: figs. 2, 4; 1993, pl. 6: figs. 1, 3, 5). It<br />

also is ironic that the first mention of expanded roots in bird<br />

teeth dates back to the first description of teeth in Archaeopteryx<br />

by Evans (1865), and this was later confirmed by Edmund<br />

(1960). There is an excellent oblique photograph (Wellnhofer,<br />

1993, pl. 5: fig. 9) of the seventh specimen of<br />

Archaeopteryx showing the waisted crown and expanded root<br />

typical of other birds and the so-called interdental plates crossing<br />

as tooth septa. Although the features of bird teeth may be<br />

most easily seen in large isolated specimens from Hesperornis<br />

and Parahesperornis, it is clear that all known bird teeth are<br />

closely similar to each other and that the teeth of Archaeopteryx<br />

are not atypical or especially primitive. The claim for interdental<br />

plates (Elzanowski and Wellnhofer, 1996) in the seventh<br />

skeletal specimen is based on the labial margin of the jaw<br />

being higher than the lingual margin and exposing distinct alveolar<br />

septa (interdental plates?) between the tooth sites. In<br />

fact, this condition is better displayed in the <strong>Lo</strong>ndon maxillary<br />

(Howgate, 1984), where it has been interpreted as a tooth socket<br />

(Martin, 1991). Because the alveolar bone of the sockets and<br />

the attachment bone of the interdental plates are ultimately derived<br />

from the same source, we must carefully describe what is<br />

meant by interdental plates versus sockets. When interdental<br />

plates are present, they generally expose the replacement teeth,<br />

and most of the length of the root is surrounded on the lingual<br />

side by the interdental plates. The jaws of the seventh specimen<br />

of Archaeopteryx are spread and compressed so that we get a<br />

slightly oblique view of the jaws on the slab (see Wellnhofer,<br />

1993, pl. 4: fig. 1). We are indebted to Wellnhofer (1993) for<br />

excellent photographs that clearly show the crown-root juncture<br />

on the teeth of the seventh example and show that the constriction<br />

at the base of the crown is almost at the lingual edge of<br />

the mandible (Figure 3E), so that the replacement teeth are<br />

mostly hidden by the side of the dentary as in other birds.<br />

When we look at typical interdental plates (Figure lG), we<br />

see that not only the socket but also part of lingual side of the<br />

jaw is produced by the interdental plates, and that there is a distinct<br />

groove separating the individual plates, terminating in the<br />

"special foramina" and the replacing teeth. The replacing teeth<br />

lie to the lingual side of the adult tooth as shown by the replacing<br />

tooth in Troodon (Figure IF). These are not the relation­<br />

ships shown in the <strong>Lo</strong>ndon Archaeopteryx (Figure 3A-C) or in<br />

the seventh specimen (Figure 3E). The intersepta of the tooth<br />

sockets of the <strong>Lo</strong>ndon maxilla closely resemble a similar view<br />

of an alligator maxilla (Figure 3F), as well as the sockets of Archaeopteryx<br />

(Figure 3E) so well photographed by Wellnhofer<br />

(1993). A close examination of Wellnhofer's photographs also<br />

shows the intersepta widening again as they come to the labial<br />

edge, as expected in a dorsal view of the socket. If the view<br />

were entirely medial, we would not expect to see this widening,<br />

even if these were interdental plates (see Figure lG).<br />

The condition in coelurosaurs is not as clear as it is in camosaurs.<br />

Compsognathus is reported by Ostrom (1978) to have<br />

small interdental plates. Dromaeosaurs were not thought to<br />

have interdental plates (Colbert and Russell, 1969). According<br />

to Currie (1995), dromaeosaurs have fairly typical interdental<br />

plates forming much of the lingual side of the jaw below the<br />

sockets except that the grooves fuse across, forming a solid<br />

wall. This should indicate a modified tooth replacement, and<br />

indeed it appears that the tooth family may be thrown into diagonal<br />

lines so that replacing teeth are both lingual and posterior,<br />

as, for example, in the overlapping replacement tooth in the ramus<br />

of Deinonychus illustrated by Ostrom (1969). This is not<br />

bird-like, nor is the covering of the interdental plates by special<br />

bones (supradentary) in Allosaurus and Tyrannosaurus (intercoronoid<br />

of Brown and Schlaikjer, 1940). It is clear that coelurosaur<br />

teeth are very similar to the teeth of camosaurs and do<br />

not show the specialized type of tooth replacement found in<br />

birds (Figure 2A-D).<br />

Conclusion<br />

The argument that birds are related to dinosaurs is now most<br />

often restated that birds are dinosaurs. If this is the case, we<br />

would expect their anatomical structures to maintain similarity<br />

under a very rigorous analysis. We see that this is not tme for<br />

almost any aspect of tooth form, implantation, or replacement.<br />

The tooth structures identified as interdental plates in Archaeopteryx<br />

by Wellnhofer (1993) do not agree in detail with those<br />

structures in dinosaurs and can be closely duplicated by crocodilians.<br />

We now know from the abundant Chinese enantiornithine<br />

material that the tooth form of birds is similar in all<br />

known groups of birds and must have been established at least<br />

by the Jurassic. Crocodilians and birds form the inside walls of<br />

their tooth-bearing bones differently from dinosaurs and have a<br />

different mode of tooth replacement. Their common ancestor<br />

with dinosaurs may not have been "thecodont" in the descriptive<br />

sense of that word. Crocodilians have derived features that<br />

prevent them from being ancestral to birds, but a sister-group<br />

relationship is still possible.

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