Quadrate bone

Abstract: The structure and evolution of the mandible, suspensorium, and stapes of mammal-like reptiles and early mammals are examined in an attempt to determine how, why, and when in phylogeny the precursors of the mammalian tympanic bone, malleus, and incus (postdentary jaw elements and quadrate) came to function in the reception of air-borne sound. The following conclusions are reached. It is possible that at no stage in mammalian phylogeny was there a middle ear similar to that of “typical” living reptiles, with a postquadrate tympanic membrane contacted by an extrastapes. The squamosal sulcus of cynodonts and other therapsids, usually thought to have housed a long external acoustic meatus, possibly held a depressor mandibulae muscle. In therapsids an air-filled chamber (recessus mandibularis of Westoll) extended deep to the reflected lamina and into the depression (external fossa) on the outer aspect of the angular element. A similar chamber was present in sphenacodontids but pterygoideus musculature occupied the small external fossa. The thin tissues superficial to the recessus mandibularis served as eardrum. Primitively, vibrations reached the stapes mainly via the anterior hyoid cornu, but in dicynodonts therocephalians, and cynodonts, vibrations passed mainly or exclusively from mandible to quadrate to stapes and the reflected lamina was a component of the eardrum. In the therapsid phase of mammalian phylogeny, auditory adaptation was an important aspect of jaw evolution. Auditory efficiency, and sensitivity to higher sound frequencies, were enhanced by diminution and loosening of the postdentary elements and quadrate, along with transference of musculature from postdentary elements to the dentary. These changes were made possible by associated modifications, including posterior expansion of the dentary. Establishment of a dentray-squamosal articulation permitted continuation of these trends, leading to the definitive mammalian condition, with no major change in auditory mechanism except that in most mammals (not monotremes) the angular, as tympanic, eventually became a non-vibrating structure.

Abstract: A detailed description is given of the osteology of the holotype of Sphenosuchus. The skull, particularly the braincase, is excellently preserved and shows a wealth of anatomical detail. Sphenosuchus was one of the largest of the early crocodylomorphs, with a skull length of 192 mm and an estimated total length of 1.4 m. The primary head of the quadrate meets the prootic and squamosal but not the opisthotic (or laterosphenoid); quadrate and pterygoid are not fused to the braincase and the basipterygoid articulation is free. The braincase and some other skull bones are pneumatized. The otic capsule is crocodilian but the subcapsular buttress (ossified subcapsular process) does not enclose the vagus nerve or the internal carotid artery. The scapula blade is triangular; the coracoid has a long posteroventral extension which is thought to have articulated firmly with a large interclavicle. Clavicles were absent. Metatarsal I is reduced; metatarsals II and IV are symmetrical about III, which is longest. A paired series of dorsal scutes was present. Sphenosuchus is considered to have been cursorial and carnivorous. Comparison is made between the pneumatic spaces in the Sphenosuchus skull and those of modern crocodiles and birds, and homologies are discussed. Representatives of the main cavities found in the crocodilian skull are present in the skull of Sphenosuchus, in some cases in a less clearly defined state. On the other hand, certain pneumatic spaces in the Sphenosuchus skull are not found in the modern crocodile but resemble cavities in the bird skull. The courses of the internal carotid and stapedial arteries are reconstructed; the latter is considered to have passed through the postquadrate foramen, temporal canal and anterior temporal foramen as it does in modern forms. The problem of the position of the stapedial artery in the crocodile is discussed. It is believed that enclosure of the artery took place as a result of the forward migration of the quadrate head, leading to the formation of a temporal canal. Detailed comparisons are made between the otic capsule of Sphenosuchus and those of modern crocodiles and birds, which it closely resembles. Changes in otic capsule structure in archosaurs to give the crocodilian or bird condition, starting from a primitive form like Euparkeria, are outlined. The skull is believed to have been kinetic, and the quadrate streptostylic, in the juvenile Sphenosuchus. The parts of the proximal end of the crocodilian quadrate are differentiated; in particular, the \`true' head is distinguished from the anterodorsal process. Although very reduced in the modern crocodile, the \`true' head is in the same morphological position as in Sphenosuchus; contact with the laterosphenoid has been brought about, not by further forward movement of the head, but by geniculation of the upper portion of the bone. The anterodorsal process is considered to have arisen as a result of the dorsal migration of an anterolateral projection somewhat similar to that of the thecodontian Stagonolepis. This change was also responsible for the elongation of the quadratojugal in crocodylomorphs. The validity of the order Crocodylomorpha is discussed. It is concluded that the most important steps in crocodylomorph evolution, particularly in the skull, had taken place in sphenosuchians, hence they should be included in the same taxon as protosuchians and more advanced crocodilians, rather than with thecodontians.