Figuring out the upside-down skull of Yanoconodon

Figure 1. Yanoconodon fossil in situ. See the skull in closeup in figure 2.

Figure 1. Yanoconodon fossil in situ. See the skull in closeup in figure 2. The published tracing is distorted here to match the underlying photo.

Wikipedia reports, “Yanoconodon was a small mammal, barely 5 inches (13 centimetres) long. It had a sprawling posture, Yanoconodon was a Eutriconodont, a group composing most taxa once classified as “triconodonts” which lived during the time of the dinosaurs. These were a highly ecologically diverse group, including large sized taxa such as Repenomamus that were able to eat small dinosaurs, the arboreal Jeholodens, the aerial volaticotherines and the spined Spinolestes. Yanoconodon is inferred to be a generalized terrestrial mammal, capable of multiple forms of locomotion.

Figure 1. Yanoconodon is exposed in ventral view. Even so, if you employ DGS, even on a fuzzy photo, you can put together a reconstruction that shares several traits with Repenomamus.

Figure 2. Yanoconodon is exposed in ventral view. Even so, if you employ DGS, even on a fuzzy photo, you can put together a reconstruction that shares several traits with Repenomamus.

Mammal-like reptiles?
Wikipedia also reports, “The Yanoconodon holotype is so well preserved that scientists were able to examine tiny bones of the middle ear. These are of particular interest because of their “transitional” state: Yanoconodon has fundamentally modern middle ear bones, but these are still attached to the jaw by an ossified Meckel’s cartilage. This is a feature retained from earlier stem mammals, and illustrates the transition from a basal tetrapod jaw and ear, to a mammalian one in which the middle ear bones are fully separate from the jaw. Despite this feature Yanoconodon is a true mammal. It is thought that the feature was retained during early embryo development,[4] whereas it is lost in most other mammal groups. The intermediate anatomy of the middle ear of Yanocodon is said to be a “Rosetta Stone”[5] of mammalian middle ear evolution.”

In the large reptile tree (LRT, 1037 taxa) Yanoconodon, Repenomamus, Jeholodens and Spinolestes are not mammals, but very close to the base of the Mammalia. Both clades share Pachygenelus as last common ancestor. So that means the ‘transitional state’ mentioned above is indeed outside the Mammalia. Other paleontologists consider this list of taxa to be mammals, but here the mammal-like traits they had were developed in parallel and not quite to mammal standards.

Figure 4. Repenomamus reconstructed using DGS methods. The manus and feet are loose figments at present. Despite its predatory nature, note the reduction in canines, a clade trait.

Figure 4. Repenomamus reconstructed using DGS methods. The manus and feet are loose figments at present. Despite its predatory nature, note the reduction in canines, a clade trait.

The skull of Yanoconodon
(Fig. 2) can be largely, but not completely, reconstructed based on the visible bones. The skull is low and wide and without the typical constriction anterior to the jugals. The anterior teeth are large and spike-like while the posterior teeth are molariform. Large teeth typically require deep roots and deep bones to house those roots. The mandibles are as long as the skull. The small orbits are far forward on the skull and the temporal fenestra are correspondingly large.

Figure 2. The origin and radiation of stem mammals and crown mammals. Compare the LRT tree (above) to a recent cladogram by Close et al. 2015.

Figure 2. The origin and radiation of stem mammals and crown mammals. Compare the LRT tree (above) to a recent cladogram by Close et al. 2015.

With the new data on Yanocondon
several taxa within the LRT shifted places, but not far and still within the derived Cynodontia. Something about the Mammalia helped them survive several extinction events that the derived Tritylodontia (= Pseudomammalia) succumbed to. Pseudomammalia LOOK like mammals, but are not mammals. They continued to exist into the Early Cretaceous and some, like Repenomamus, were quite large.

References
Close RA, Friedman M, Lloyd GT and Benson RBJ 2015. Evidence for a mid-Jurassic adaptive radiation in mammals. Current Biology. 25(16): 2137–2142. 
Luo Z, Chen P, Li G, and Chen M 2007.
 A new eutriconodont mammal and evolutionary development in early mammals. Nature 446:15. online Nature

wiki/Yanoconodon

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