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1 School of Environmental Sciences, University of the North (Qwaqwa Branch), Private Bag X13, Phuthaditjhaba 9866, Republic of South Africa
2 National Museum, Bloemfontein 9300, Republic of South Africa
3 Department of Earth Sciences, Simon Fraser University, Burnaby, British Columbia, V5A 1S6 Canada
A scratch-marked burrow complex with multiple branching tunnels and terminal chambers was excavated from the Lower Triassic Driekoppen Formation, northeastern Free State, South Africa. The burrow complex is attributed to the therapsid Trirachodon, based on disarticulated but fairly complete skulls and skeletons of at least 20 individuals recovered from a nearby, less well-preserved system.
The entrance shaft slopes gently downward and is characterized by a bilobate floor and vaulted roof. The bilobate floor has a central flat-topped scratch-marked ridge flanked by two smooth grooves, each approximately the width of the occupant. At deeper levels, tunnels display tighter lateral curvatures, progressively decreasing burrow diameters, variable orientations, and some right angle branches. Burrow floors at these levels are vaguely bilobate. Distally, burrows flatten dorso-ventrally, becoming wedge-shaped before they terminate.
These complexes are interpreted as colonial dwelling structures. Numerous branching tunnels and terminal chambers, as well as an enlarged entrance, constitute an unrealistically high expenditure of energy for a single occupant. Furthermore, the bilobate floor is atypical of a single occupant system. Absence of scratch marks in the depressions reflects the regular locomotory activity of Trirachodon. In single-occupant burrows, the center of the structure is worn. The bilobate floor with preferential wear along the flanks suggests routine travel on one side of the tunnel or the other, as multiple occupants moved past one another during daily activity.
The presence of numerous fossilized Trirachodon individuals in terminal chambers at the second locality supports multiple cohabitation. There, the uniform sedimentary fill and vertebrate taphonomy suggest that the occupants were drowned in a flash flood. Vertebrate remains are absent at the first locality, where sediment influx was incremental, permitting the escape of burrow occupants.
This constitutes the earliest record of multiple cohabitation of a burrow complex by tetrapods. The Trirachodon (Cynodontia) of the Early Triassic Cynognathus Zone probably displayed complex social behaviors previously regarded to be restricted to the mammals of the Cenozoic. This fossorial behavior may have been for thermoregulation, protection from predators, sites of reproduction, and the rearing of young.
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