A temnospondyl trackway from the early Mesozoic of western Gondwana and its implications for basal tetrapod locomotion.

BACKGROUND: Temnospondyls are one of the earliest radiations of limbed vertebrates. Skeletal remains of more than 190 genera have been identified from late Paleozoic and early Mesozoic rocks. Paleozoic temnospondyls comprise mainly small to medium sized forms of diverse habits ranging from fully aquatic to fully terrestrial. Accordingly, their ichnological record includes tracks described from many Laurasian localities. Mesozoic temnospondyls, in contrast, include mostly medium to large aquatic or semi-aquatic forms. Exceedingly few fossil tracks or trackways have been attributed to Mesozoic temnospondyls, and as a consequence very little is known of their locomotor capabilities on land. METHODOLOGY/PRINCIPAL FINDINGS: We report a ca. 200 Ma trackway, Episcopopus ventrosus, from Lesotho, southern Africa that was made by a 3.5 m-long animal. This relatively long trackway records the trackmaker dragging its body along a wet substrate using only the tips of its digits, which in the manus left characteristic drag marks. Based on detailed mapping, casting, and laser scanning of the best-preserved part of the trackway, we identified synapomorphies (e.g., tetradactyl manus, pentadactyl pes) and symplesiomorphies (e.g., absence of claws) in the Episcopopus trackway that indicate a temnospondyl trackmaker. CONCLUSIONS/SIGNIFICANCE: Our analysis shows that the Episcopopus trackmaker progressed with a sprawling posture, using a lateral-sequence walk. Its forelimbs were the major propulsive elements and there was little lateral bending of the trunk. We suggest this locomotor style, which differs dramatically from the hindlimb-driven locomotion of salamanders and other extant terrestrial tetrapods can be explained by the forwardly shifted center of mass resulting from the relatively large heads and heavily pectoral girdles of temnospondyls.

Evidence for avian intrathoracic air sacs in a new predatory dinosaur from Argentina.

BACKGROUND: Living birds possess a unique heterogeneous pulmonary system composed of a rigid, dorsally-anchored lung and several compliant air sacs that operate as bellows, driving inspired air through the lung. Evidence from the fossil record for the origin and evolution of this system is extremely limited, because lungs do not fossilize and because the bellow-like air sacs in living birds only rarely penetrate (pneumatize) skeletal bone and thus leave a record of their presence. METHODOLOGY/PRINCIPAL FINDINGS: We describe a new predatory dinosaur from Upper Cretaceous rocks in Argentina, Aerosteon riocoloradensis gen. et sp. nov., that exhibits extreme pneumatization of skeletal bone, including pneumatic hollowing of the furcula and ilium. In living birds, these two bones are pneumatized by diverticulae of air sacs (clavicular, abdominal) that are involved in pulmonary ventilation. We also describe several pneumatized gastralia ("stomach ribs"), which suggest that diverticulae of the air sac system were present in surface tissues of the thorax. CONCLUSIONS/SIGNIFICANCE: We present a four-phase model for the evolution of avian air sacs and costosternal-driven lung ventilation based on the known fossil record of theropod dinosaurs and osteological correlates in extant birds: (1) Phase I-Elaboration of paraxial cervical air sacs in basal theropods no later than the earliest Late Triassic. (2) Phase II-Differentiation of avian ventilatory air sacs, including both cranial (clavicular air sac) and caudal (abdominal air sac) divisions, in basal tetanurans during the Jurassic. A heterogeneous respiratory tract with compliant air sacs, in turn, suggests the presence of rigid, dorsally attached lungs with flow-through ventilation. (3) Phase III-Evolution of a primitive costosternal pump in maniraptoriform theropods before the close of the Jurassic. (4) Phase IV-Evolution of an advanced costosternal pump in maniraptoran theropods before the close of the Jurassic. In addition, we conclude: (5) The advent of avian unidirectional lung ventilation is not possible to pinpoint, as osteological correlates have yet to be identified for uni- or bidirectional lung ventilation. (6) The origin and evolution of avian air sacs may have been driven by one or more of the following three factors: flow-through lung ventilation, locomotory balance, and/or thermal regulation.