PROBLEMS OF THE ORIGIN OF REPTILES

The fossil records of the four living reptilian orders can be traced into the Triassic. The earlier ancestry of the turtles has not been established. Squamates and rhyncho-cephalians evolved from the Late Permian eosuchians; crocodiles from the thecodonts. The ancestry of the eosuchians and thecodonts is to be found in the central stock of Permo-Carboniferous reptiles, the captorhinomorphs. The earliest captorhino-morphs, from the Lower Pennsylvanian, are already fully developed reptiles. The limnoscelids and solenodonsaurids are more primitive forms, retaining features typical of anthracosaurian amphibians. Neither reptiles nor any appropriate ancestors are known prior to the Lower Pennsylvanian. Because of the absence of any true ancestors, the nature of the amphibian-reptilian transition must be studied on the basis of amphibians contemporary with the early reptiles. The Permian seymouriamorphs have long been accepted as relicts of the group which gave rise to reptiles, although Seymouria itself is specialized in many features of its anatomy. The Middle Pennsylvanian genus Gephyrostegus appears to resemble much more closely the anatomy expected in the ancestors of reptiles. This genus forms the basis for consideration of the anatomical, physiological and behavioural changes which culminated in the origin of reptiles. Study of the earliest known reptiles and their closest relatives among contemporary amphibians indicates that the initial adaptation leading to the emergence of the class was assumption of a terrestrial habit, with accompanying small body size. The small body size of the immediate ancestors of reptiles would have made it possible for them to produce sufficiently small eggs that they could develop in damp places on land without initially being supported and protected by extraembryonic membranes. The rapid increase in body size in all lineages of Pennsylvanian reptiles indicates the prior development of an amniotic egg. Fundamental to the emergence of reptiles was modification in the jaw mechanism from the kinetic inertial system of amphibians to a static pressure system. The latter was presumably developed in order for the developing reptiles to utilize more active terrestrial prey. This change in the jaw mechanism is reflected in the reorganization of the palate which serves as a morphological basis for denning the establishment of reptilian status. At approximately the same stage as the change in palatal structure, the definitive reptilian vertebral pattern was developed. The apparent closure of the otic notch and the probable reorientation of the stapes in the amphibian-reptilian transition presumably resulted from the decrease in relative skull size and do not appear to be related to any change in hearing ability. The tympanum probably maintained the same relative relationship with the squamosal and supratemporal throughout this transition. On the basis of the present fossil record, all adequately known Palaeozoic reptiles appear to have had a common ancestry among the predecessors of the known gephyro-stegids. The family Diadectidae is the only important group whose specific relationships cannot be established. On the basis of this study, the following taxonomic changes are suggested: the family Limnoscelidae should not be included among the captorhinomorphs. The seymouria-morph concept should be restricted to forms having the specializations of Seymouria, the discosauriscids and kotlassids. Gephyrostegids should be specifically excluded from the Seymouriamorpha and should be included in a separate taxon among the anthra-cosaurs of equal rank with embolomeres and seymouriamorphs.