Remarkable Fossil Footprints Reveal How Dinosaur Ancestors Walked
Scientists studying ancient fossil footprints have uncovered important evidence about the evolution of dinosaur locomotion. The research suggests that the ancestors of dinosaurs gradually changed how they walked. Over millions of years, these animals evolved from sprawling reptiles into creatures with a more upright stance. The newly published research studying dinosaur ancestors footprints provides a fresh perspective on the evolution of bipedal dinosaurs.
The international research team, led by Dr Lorenzo Marchetti of the Museum für Naturkunde Berlin, examined fossil tracks from the Italian Dolomites. Their findings have been published in the scientific journal “Palaeontology”.
Ancient Footprints from the Italian Dolomites
The researchers studied fossil footprints discovered in GEOPARC Bletterbach, part of the UNESCO World Heritage Site in the Dolomites. These remarkable tracks date from the Late Permian, before the devastating end-Permian mass extinction approximately 252 million years ago. Fossils of early archosauriform reptiles from this period are extremely rare. Consequently, fossil footprints provide valuable evidence about animals that are poorly represented in the skeletal fossil record.
The researchers reassessed the footprints and established a new trace fossil taxon, Pachychirotherium gardenaensis. This scientific name refers to the robust nature of the tracks and their geographical origins in the Gröden Valley. The footprints represent some of the earliest known evidence of archosauriform locomotion.

A trackway of an early archosauriform. The type material of Pachychirotherium gardenaensis. Scale bar equals 5cm. Picture credit: University La Sapienza of Roma.
Picture credit: University La Sapienza of Roma
From Sprawling Reptiles to Upright Walkers
By comparing Permian and Triassic footprints, the scientists reconstructed important changes in the posture of early archosauriforms. During the Late Permian, these animals generally walked with their limbs positioned outwards from their bodies. This sprawling posture resembles the stance seen in many modern lizards. However, their descendants gradually developed a more upright posture during the Triassic. By the Middle Triassic, this evolutionary transition had become well established.
Other important anatomical changes accompanied this transformation. For example, the middle toes became longer, whilst the forefeet became proportionately smaller. Furthermore, these reptiles increasingly walked on their toes rather than placing the entire foot on the ground. This type of locomotion is known as digitigrady. Birds walk on their toes, whereas humans walk on their toes and heels (plantigrade). Together, these adaptations helped early archosauriforms become more energy efficient and better runners.

A close view of the foot of the flightless bird (Rhea). Note the enlarged middle toe and that this bird walked on its toes. Picture credit: Everything Dinosaur.
Picture credit: Everything Dinosaur
Important Steps Towards Dinosaur Bipedalism
The study suggests that the evolution of upright posture, digitigrady and adaptations for running occurred gradually. These changes did not happen simultaneously. Instead, they developed over millions of years as archosauriform reptiles diversified during the Permian and Triassic. Such evolutionary innovations helped establish the anatomical foundations for bipedal locomotion in later dinosaurs and their close relatives.
For example, famous theropod dinosaurs such as Tyrannosaurus rex possessed powerful hind limbs and walked on their toes. However, the new research demonstrates that some of the fundamental adaptations associated with dinosaur locomotion originated long before the first true dinosaurs evolved.

The left metatarsal bones of a T. rex (Titus). Theropod dinosaurs walked on their toes. Note the enlarged middle toe. Picture credit: Everything Dinosaur.
Picture credit: Everything Dinosaur
Fossil Footprints Provide Unique Evolutionary Evidence
Fossilised skeletons help palaeontologists reconstruct the anatomy of extinct animals. In contrast, trace fossils preserve direct evidence of their activities. Footprints can reveal information about posture, walking style and how an animal distributed its weight.
Dr Lorenzo Marchetti emphasised the importance of these discoveries. He explained that fossil footprints preserve remarkable details about archosauriform biology and locomotion. Such evidence is especially valuable when skeletal remains are scarce.
This research highlights the scientific importance of trace fossils. Moreover, it demonstrates how the evolutionary history of dinosaurs can be traced back to their distant Permian relatives.
The Everything Dinosaur team acknowledges the assistance of the Museum für Naturkunde Berlin in the compilation of this article.
The scientific paper: “The new archosauriform track Pachychirotherium and the stepwise acquisition of digitigrady, cursoriality and erect stance in the group between the Permian and Triassic” by Lorenzo Marchetti and Hendrik Klein published in Palaeontology.
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