There’s still the question of Effigia’s mandible hanging out there.
The question is: “Is that a predentary or a dentary at the tip?” Fig. 1). Nesbitt (2007) says dentary. I say predentaries. Let’s look at the evidence.
To answer that,
I took a comparative survey of poposaur mandibles (Fig. 1), looking for evolutionary patterns and thereby strive to provide an update to the predentary/dentary question. Surprisingly, in the case of Effigia, when you add in the splenials, which neither Nesbitt nor I did before, the mandibular fenestra becomes substantially reduced. That may be similar to what one sees in Lotosaurus, in which the elements are not jumbled. And that provides more substance to the “predentary” argument. Other than Lotosaurus, the closest sister is Shuvosaurus, which is known from an incomplete mandible (Fig.1) showing similar patterns over the remaining portions. Shuvosaurus has something similar to what I saw in Daemonosaurus, that others consider something else. In any case, at some point, something interesting developed in front of the dentaries in certain phytodinosaurs.
The other question is,
when something similar to a predentary appears in front of the dentary, as in Sacisaurus (Figure 1), should it be considered a “beak” rather than a premaxilla? This bone may be paired, as it is in Sacisaurus, rather than a single median bone, as in the predentary of Heterodontosaurus (Fig. 1).
Figure 1. Poposaur (and kin) mandibles. Here are Daemonosaurus, Poposaurus, Pisanosaurus, Heterodontosaurus, Sacisaurus, Lotosaurus, Effigia and Shuvosaurus. The mandibles of Lotosaurus, Shuvosaurus and Effigia appear to share a common heritage of design. In Effigia the splenial reduces the mandibular fenestra helping to clarify the identify of the dentary and premaxilla (or beak). The extension of the angular to the predentary is unique to this clade.
If all these other mandibles had a premaxilla or beak (or the possibility of one), is there any reason to suspect that Effigia did not?
The original reconstructions of the Effigia mandible
introduced us to the largest mandibular fenestra I have ever seen relative to the size of the jaw. The new reconstruction reduces the fenestra length and, no doubt, produces a stronger jaw with the splenial (lavendar to iris blue bone) laminated to the medial side and edges.
Typically the mandibular fenestra splits the surangular from the angular,
as it does in Heterodontosaurus. However, in Lotosaurus the mandibular fenestra develops largely below the dentary with very little surangular and angular exposure. In Shuvosaurus the same pattern could play out, but unfortunately the key parts are missing (perhaps due to a very large mandibular fenestra?). This is a different pattern than in ornithischians, saurischians and theropods. And this pattern is also different from rauisuchians. Among euarchosauriforms, only in aetosaurs does the very large mandibular fenestra develop largely below the dentary. In others, the fenestra develops midway or beneath the surangular and it doesn’t get to the size seen in Effigia and Lotosaurus.
One final point
The suture between the two premaxillae in Effigia is convoluted like a puzzle piece. In this way they are locking themselves together, convergent with the central or fused premaxilla of ornithischians, but homologous with the premaxilla in Lotosaurus and Shuvosaurus.
If I’m wrong, show me some data. At this point, at least it’s worth talking about.
As always, I encourage readers to see specimens, make observations and come to your own conclusions. Test. Test. And test again.
Evidence and support in the form of nexus, pdf and jpeg files will be sent to all who request additional data.
Ferigolo J and Langer MC 2006. “A Late Triassic dinosauriform from south Brazil and the origin of the ornithischian predentary bone”. Historical Biology 19 (1): 1–11. online pdf.
Nesbitt SJ and Norell MA 2006. Extreme convergence in the body plans of an early suchian (Archosauria) and ornithomimid dinosaurs (Theropoda). Proceedings of the Royal Society B 273:1045–1048. online
Nesbitt S 2007. The anatomy of Effigia okeeffeae (Archosauria, Suchia), theropod-like convergence, and the distribution of related taxa. Bulletin of the American Museum of Natural History, 302: 84 pp. online pdf
AMNH Effigia webpage