SYNOPSIS WELL preserved material of the crossopterygian fish Eusthenopteron enables fresh reconstructions and interpretations of its postcranial skeleton to be given. Comparisons throughout with other bony fishes show that it may •« primitive in many features. Similarities with early amphibians such as the screw-shaped glenoid, the form of the humerus (on which an attempt to restore the pectoral musculature is based), the dorsal bicipital ribs and the possibility of a sacral attachment, throw much light on the origin of the tetrapod postcranial skeleton, particularly of the cheiropterygium. A functional analysis of the skeleton of Ensthenopteron is attempted, suggesting that it resembled the pike (Esox) in its mode of life and that it may have been capable of short journeys "walking" overland. The possible selective factors stimulating the evolution of such a fish, and further evolution to the tetrapod stage are discussed.
SynopsisIn a previous paper (Andrews and Westoll 1970) the postcranial skeleton of the best known rhipidistian, Eusthenopteron, was described, and its bearing on the origin of the tetrapod postcranial skeleton discussed. The postcranial remains of other Rhipidistia are now described as far as they are known, and comparisons are made with Eusthenopteron and other forms where relevant. Possible modes of function are considered in relation to the habitats in which these fishes may have lived. These studies have made it necessary to revise rhipidistian classification; the Family Rhizodontidae is re-defined and placed alone in a third Order of Rhipidistia (the Rhizodontida, alongside the better known Osteolepidida and Holoptychiida). Fresh insight has been gained into the following morphological problems: the composition of the osteolepid ring-like centrum, the origin of the tetrapod scapular blade and the diphyletic origin of the tetrapods.
Material of crossopterygian fishes from Foulden, Berwickshire, is important in including the first known entire specimen giving a clear picture of the body form of the Family Rhizodontidae (sensu Andrews 1973). This group is specialised in having numerous subsidiary lateral lines on the body, a ventrally expanded shoulder girdle, fins with peculiar stiffened structure, the pectorals forming greatly enlarged paddles and other fins being reduced in size. The Foulden remains fall into large and small size-ranges occurring largely at different levels in the sequence, but although these may represent growth in one taxon the available parts of the body are mostly complementary. Separate preliminary descriptions of the two size-ranges are therefore given, the small complete form being named ?Strepsodus anculonamensis sp. nov., and the large form remaining unnamed pending further study. An abridged historical account of previously named rhizodont genera (Rhizodus, Strepsodus, Sauripterus and Pycnoctenion) is given in order to explain why the new Foulden form(s) cannot at present be generically assigned more closely. A revised diagnosis of the Family Rhizodontidae leads to a discussion of the functional morphology and mode of life of fish of this type. Analogies with modern forms indicate that, like some sharks and crocodiles, they may have fed by tearing flesh off large prey, rotating or shaking against its inertia. Ways in which variously shaped rhizodont teeth (recurved, sigmoid etc.) may have functioned are suggested. Addenda deal with (1) the possible occurrence at Foulden of a large lungfish and (2) (after considering primitive and advanced rhizodont features) the renaming of Rhizodus ornatus Traquair 1878 from higher in the Scottish lower Carboniferous, as the type of a new genus, Screbinodus.
Westlothiana lizziae is known from the Brigantian of East Kirkton, Scotland. The skull resembles that of later amniotes in the large size of the parietal, the apparent loss of the intertemporal, and the absence of a squamosal notch, palatal fangs and labyrinthine infolding of the marginal teeth, but is primitive in the absence of a transverse flange of the pterygoid. The individual trunk vertebrae resemble those of amniotes; large intercentra are retained, but the neural arch is fused to the centrum. A surprising feature is the presence of 36 presacral vertebrae, as is the relative size of the very small but highly ossified limbs. The humerus is much shorter than the femur, but similar in configuration to that of early amniotes. There are three proximal tarsals as in primitive tetrapods, but an amniote phalangeal count. The presence of massive dorsal as well as ventral scales is a more primitive feature than that of most anthracosaurs.Westlothiana is ‘reptiliomorph’, and is judged to be a stem-group amniote on features of the skull roof, the absence of an otic notch, the gastrocentrous vertebrae and the pedal phalangeal formula. It has not, however, reached the amniote condition in the structure of the tarsus, and the palate is more primitive than that of both early amniotes and the ‘diadectomorphs’.
The Order Adelospondyli, an isolated group of amphibians from the Carboniferous of the Edinburgh area, is redescribed and distinguished from other early tetrapods. Although systematics of genera and species is unsatisfactory, the group seems monophyletic, since as far as they are known, all conform to a single overall structural plan. There is an entire skull roof with a reduced complement of bones in an unique pattern, a highly characteristic marginal dentition, a massive hyoid apparatus, at least 70 holospondylous trunk vertebrae, and (like some other Lower Carboniferous tetrapods) a shoulder region with a heavy dermal girdle but no trace of ossified limbs or endochondral girdle. Adelogyrinids show no special affinity with other Palaeozoic amphibians: the vertebral characters in which they resemble microsaurs, lysorophids, aïstopods, and nectrideans may have evolved more than once among early tetrapods, and the shoulder girdle and neural arch characters shared with labyrinthodonts are probably primitive. The adelogyrinids thus illustrate the more general problems of establishing interrelationships among early tetrapods. Their way of life is discussed.
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