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WITMER-ANTORBITAL
extensive medial lamina of the maxillary ascending ramus and
a similar medial lamina from the lacrimal bone such that there
is an extensive antorbital fossa (Figs. 7, 8). These thin medial
laminae are well preserved in Lesothosaurus diagnosticus
(BMNH RUB23, R11956, RUB17, R8501), Heterodontosaurus
tucki (BMNH R8179 [cast of SAM 337]), and Hypsilophodon
foxii (BMNH R197, R2477) and constrict the size of the internal
antorbital fenestra to a foramen. The maxillary medial larnina
of Hypsilophodon foxii (BMNH R2477) and also Heterodontosaurus
tucki (but not that of the basal heterodontosaurid
Lanasaurus scalpridens; Gow, 1975) has an additional opening
between the antorbital and nasal cavities.
The antorbital cavity is enclosed laterally to a variable degree.
In all the aforementioned taxa, there is a well-developed
supralveolar lamina projecting dorsally, giving the ventral margin
of the external antorbital fenestra a sharp edge. In Lesothosaurus
diagnosticus there is no significant development of a
lateral lamina of the maxillary ascending ramus; the rim around
the extensive antorbital fossa is sometimes distinct (BMNH
R11956) and sometimes not (BMNH RUB17). In Heterodontosaurus
tucki and probably Abrictosaurus consors, but not in
Lanasaurus scalpridens, the rostral rim of the antorbital fossa
more strongly overhangs the cavity. In Hypsilophodon foxii, the
lateral lamina of the ascending ramus is extensive and greatly
restricts the size of the external antorbital fenestra (Fig. 8B). In
all taxa, the lacrimal also develops a lateral lamina that overhangs
and further encloses the cavity (Figs. 7, 8).
The course of the nasolacrimal duct is known in Lesothosaurus
diagnosticus (BMNH R8501) and Hypsilophodon foxii
(BMNH R2477). In the latter (Fig. 8A), the canal passes
through the lacrimal from the orbit, through the rostral ramus,
emerging at the rostral end medial to the medial lamina of the
maxilla as observed in Plateosaurus engelhardti (Fig. 12D). In
Lesothosaurus diagnosticus, only the orbital opening of the canal
is visible but the canal clearly does not open rostrolaterally
and therefore probably had a course similar to that in Hypsilophodon
.foxii. Thus, in both cases, the nasolacrimal duct
passed dorsomedially around the antorbital cavity.
Finally, Lesothosaurus diagnosticus has an additional fossa
on the palatine medial to the choana (mentioned above in the
discussion of the muscular hypothesis; Fig. 7C, D). This fossa
perhaps supported the nasal capsule, but it is more likely that
it is an accessory recess associated with the antorbital cavity
(see section on accessory cavities below).
In higher ornithopods, many of the aforementioned trends are
carried further. In Tenontosaurus tilletti (MOR 682), Dryosaurus
altus (CM 3392; see also Galton, 1983), and Carnptosaurus
dispar (UUVP 5946) the antorbital cavity is further reduced in
size, and lateral laminae from the maxilla and lacrimal (and
jugal in D. altus) constrict the external antorbital fenestra to a
relatively small aperture. The same relationships of antorbital
cavity to choana that were observed in more basal ornithopods
and Lesothosaurus diagnosticus are present in the iguanodontians
Iguanodon ather-eldensis (BMNH R5764, 1 152 1 ; Norman,
1986; Fig. ll), probably I. lakotaensis (SDSM 8656
[cast]; Weishampel and Bjork, 1989), and Ouranosaurus nigeriensis
(MNHN GDF 300 [cast]; Taquet, 1976), but here the
antorbital cavity is very small, does- not deeply excavate the
maxilla or lacrimal, and is enclosed laterally by only the lacrimal
such that a faint fossa extends a short distance rostrally
on the maxilla. Where known, the nasolacrimal canal passes
dorsomedially around the antorbital cavity (Fig. 11B). Hadrosaurids
present an interesting case in that they have a completely
closed external antorbital fenestra, which is walled in not
only by the massive maxilla, but also by the lacrimal, premaxilla,
and a large rostral process of the jugal (Weishampel and
Homer, 1990). The antorbital cavity, however, is retained medially
as a small cavity bounded mostly by maxilla, lacrimal,
CAVITY OF ARCHOSAURS
lac
fen antorb -
rnax
FIGURE 18. Ernausaurus emsti, facial skeleton in left lateral view.
(Modified after Haubold, 1990.)
and palatine; it is open caudally and floored ventrally with foramina
for passage of neurovasculature. The cavity is directly
opposite the choana. It should be noted that Heaton (1972) positioned
the choana much too far caudally, behind the vomer;
instead, the choana of hadrosaurids is lateral to the vomer,
bounded caudally by palatine and laterally by maxilla as in
other ornithopods.
Dinosauria: Thyreophora-Thyreophoran ornithischians
also tend to reduce the antorbital cavity and close the external
antorbital fenestra. Unfortunately, little relevant data are available
in the published material of the basal thyreophoran Scutellosaurus
lawleri (Colbert, 1981). The slightly more derived
thyreophoran, Ernausaurus ernsti (Haubold, 1990), provides
critical data. In this taxon, a well-developed antorbital cavity is
formed within the maxilla and lacrimal (Fig. 18). As in Lesothosaurus
diagnosticus and other forms, the maxilla and lacrimal
have broad medial laminae constricting the internal antorbital
fenestra to a relatively small opening. Although the palatal
elements are not known, the medial surface of the maxilla
shows the palatine articular surface, thus fixing the position of
the choana as directly opposite the internal antorbital fenestra.
There is a low supralveolar lamina (Haubold, 1990), and the
maxilla and lacrimal both have lateral laminae constricting the
size of the external antorbital fenestra. Significantly, Haubold
(1990) described a deep cavity entering the maxilla from the
antorbital cavity. The nasolacrimal canal passes through the lacrimal
bone, dorsomedial to the antorbital cavity.
Scelidosaurus harrisonii, the next higher thyreophoran, is
generally similar but shows greater closure of the cavity. In S.
harrisonii (BMNH R111 l), the internal antorbital fenestra is
not only directly opposite the choana, but the postchoanal strut
of the palatine contributes to the caudal margin of the fenestra.
The massive maxilla is so transversely thick that there is actually
a short canal passing laterally from the internal fenestra
to the main cavity. The antorbital cavity is enclosed by the
maxilla, lacrimal, and probably jugal. There is a distinct antorbital
fossa well excavated into the external surface of the max-