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[dinosaur] Jugal bone in dinosaurs + Psittacosaurus cranial function + jugal and quadratojugal in early birds

Ben Creisler

Dinosaurs in a special issue of The Anatomical Record:

Corwin Sullivan and Xing Xu (2016)
Morphological Diversity and Evolution of the Jugal in Dinosaurs.
The Anatomical Record  300(1): 30–48
DOI: 10.1002/ar.23488

In dinosaurs, as in other reptiles, the homologue of the mammalian zygomatic bone is the jugal. The dinosaurian jugal was primitively triradiate, with posterior, dorsal and anterior processes that respectively contacted the quadratojugal, the postorbital, and the maxilla and lacrimal. However, the jugal evolved along different lines in the three major dinosaurian clades. In theropods this cranial element remained relatively conservative in morphology, apart from being reduced to a rod-like structure in most birds and a few non-avians. In sauropodomorphs the jugal eventually became small, plate-like and nearly restricted to the area below the orbit, even being excluded from the ventral margin of the skull in many derived taxa. Among ornithischians the jugal was highly variable, but in many cases became large and/or adorned with ornamental features such as horns, flanges, and rugosities. The jugal does not appear to have been a site of muscle attachment in most non-avian dinosaurs, but represented an important structural element in the akinetic dinosaurian skull. The conspicuous jugal ornaments seen in many ornithischian dinosaurs, like the less striking ones documented in some saurischians, may have played an important role in the social behavior of the species that possessed them. In many cases they have a weapon-like aspect suggesting use in aggressive displays, if not actual combat, adding to the evidence that agonistic behavior was likely widespread among ornithischians in particular. 



Adam C. Taylor, Stephan Lautenschlager, Zhao Qi and Emily J. Rayfield (2016)
Biomechanical Evaluation of Different Musculoskeletal Arrangements in Psittacosaurus and Implications for Cranial Function. 
The Anatomical Record  300(1): 49–61 
DOI: 10.1002/ar.23489

The masseter muscle complex is a unique feature of extant mammals and their advanced cynodont precursors, originating from the zygomatic arch and inserting onto the lateral surface of the dentary. This muscle complex is absent in sauropsids, with the exception of the neomorphic m. pseudomasseter complex that is unique to psittaciform birds (parrots and cockatiels). The anterior position and anterodorsally inclined line of action of both muscle groups increases leverage of the jaw and is thought to contribute to increased bite force, particularly in psittaciforms. A corollary is that in mammals at least, the masseter places increased load on the zygomatic arch, which may be withstood by soft tissue temporal fascia. Recently the existence of a m. pseudomasster (mPSM) and m. adductor mandibulae externus ventralis (mAMEV) has been proposed in the ornithischian dinosaur Psittacosaurus. Here we use computed tomography, digital restoration of skull anatomy and adductor musculature and computational biomechanics to test how the presence of anterodorsally inclined muscle loads influences stress, strain, deformation and estimated bite forces in the skull of Psittacosaurus. We find that the m. pseudomasseter and m. amev increases bite force with an associated increase in cranial stress and deformation. There is, however, limited osteological evidence for the existence of these two additional muscles in the psittacosaur skull and geometric morphometric informed sensitivity analysis of our finite element models shows that bite position has a greater effect on loading-induced deformation than muscle loading or material property variation.


Min Wang and Han Hu (2016)
A Comparative Morphological Study of the Jugal and Quadratojugal in Early Birds and Their Dinosaurian Relatives.
The Anatomical Record  300(1): 62–75
DOI: 10.1002/ar.23446

The zygoma (called jugal bar) in modern birds accounts for a large portion of the ventral margin of the cranium. As a delicate and rod-like element, the jugal bar is functionally integrated into the avian cranial kinesis, a unique property that allows the beak to be elevated or depressed with respect to the braincase and thus distinguishes birds from all other modern vertebrates. Developmental studies show that the jugal bar of modern birds is formed by the fusion of the jugal and quadratojugal that are ossified from separated centers. However, those two bones are unfused and exhibit complicated morphological variations in basal birds and their dinosaurian relatives. Moreover, the jugal and quadratojugal form rigid articulations with the postorbital and squamosal, respectively, consequently hindering the movement of the upper jaw in most non-avian dinosaurs and some basal birds. A comparative study of the jugal and quadratojugal morphology of basal birds and their close relatives such as dromaeosaurids and oviraptorids elucidates how modern birds has achieved its derived jugal bar morphology, and sheds light on the evolution of the postorbital configuration of birds. We propose that numerous morphological modifications of those two elements (morphology changes and reduced articulation with other elements) took place in early bird history, and some of them pertinent to the refinement of the cranial kinesis.