COMPLEX TETRAPOD BURROWS FROM MIDDLE TRIASSIC RED BEDS OF THE ARGANA BASIN (WESTERN HIGH ATLAS, MOROCCO)

Abstract

Although burrowing ability has been widespread in tetrapods for more than 300 million years, subsurface dwelling structures that indicate communal behavior are poorly evidenced from pre-Cenozoic strata. Here we present recently discovered tetrapod burrows from Middle Triassic red beds of the Argana Basin in central Morocco, whose complexity suggests an origin by gregarious animals. The well-preserved burrows occur in interbedded mudstones and sandstones interpreted as channel and overbank deposits of ephemeral, braided streams. All burrows originate from the top of thick-bedded sandstones and descend as moderately inclined (10°–30°), partially spiral tunnels to laterally extended, branched chambers in underlying mudstones. Tunnel segments are biconvex to planoconvex in cross section, up to 20 cm wide and 12 cm in maximum height and exhibit transverse scratch marks along the ceilings and sidewalls. Distinctive burrow characteristics include a laterally sinuous geometry (wavelength λ = 38–45 cm; amplitude A = 5–10 cm) of the tubelike passages and the presence of grouped alcoves in terminal chambers. We attribute the burrows to procolophonids or therapsids based on closely associated tetrapod tracks and the limited diameter of the excavations. Our findings represent the second oldest record of communal fossorial behavior by tetrapods and the oldest example from low-latitude areas. Beyond providing refuge from predators, these elaborate underground structures probably functioned as a buffer against diurnal or seasonal variations of air temperature and humidity in a semiarid habitat that was situated just north of the paleoequator.