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The authors have previously studied the trackways of modern horses in order to elucidate the gaits exhibited in fossil horse trackways. Here the authors further refine their methodology and apply it to the study of various fossil horse trackways described in the literature. Miocene horse footprints include three trackways found near Jumilla, Spain of Cremohipparion matthewi of horses trotting at an estimated speed of 2.9-3.4 m/s; three trackways from the same site of Cremohipparion matthewi galloping at around 5.2-5.6 m/s; three trackways of Hippotherium malpassii found near Osoppo, Italy of horses galloping at around 6.2-6.5 m/s; and a trackway of Scaphohippus sumani from Barstow, California, in the United States, of a horse likely engaged in a rack at 2.1 m/s. Pliocene horse footprints include a trackway of Hipparion fissurae located near Elche, Spain, showing a horse trotting at around 3.5 m/s, and three trackways of Eurygnathohippus hasumense from Laetoli, Tanzania, displaying two horses racking at 2.1-3.0 m/s as well as perhaps one performing a running walk at 3.1 m/s. Finally, there is one Pleistocene trackway of Equus lambae located near Cardston, Alberta, Canada showing a horse in a gallop at around 6.6 m/s. Hence fossil horses showed the ability to engage in both laterally coordinated and diagonally coordinated symmetrical gaits, as well as asymmetrical gallops. In short they exhibit the same variety of gaits found in modern horse breeds. Several of the trackways showed evidence of herding behavior with the galloping trackways in Cremohipparion all orientated in the same direction but crossed at a perpendicular angle by three trotting trackways perhaps indicating the traveling behavior of adults with (supervised) juveniles at play. The galloping trackways of Hippotherium were also orientated in a similar direction suggesting herding behavior. The Eurygnathohippus prints show three trackways orientated in the same direction in lateral gaits, with a juvenile trackway close to and crossing over a likely mare trackway, showing a mare closely following her foal as is seen in horse herds today.

New Studies on the Gaits Displayed by Miocene, Pliocene, and Pleistocene Fossil Horse Trackways