Vertebrate tracks offer important paleobiological and paleoenvironmental insights as indicated in the increasing number of studies of modern and fossil tracks in recent decades (Laporte and Behrensmeyer, 1980; Loope, 1986; McKeever, 1991; Platt and Hasiotis, 2008; Milàn et al., 2007; and references therein). Whereas fine-grained sediments typically preserve the greatest detail, a track record of many organisms is archived in sand-rich strata representing paleo-shorelines, river banks, and dunes (Frey and Pemberton, 1986; Lockley, 1986). Despite the dynamic nature and coarse texture of sandy media ( = substrates), many studies report abundant...

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