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A 2026 analysis of Permian and Triassic trackways shows that dinosaur ancestors did not switch suddenly from lizard-like sprawling to upright walking. Their broader group, archosauriforms, changed in stages: sprawling locomotion dominated in the Late Permian, a major shift occurred near the end-Permian mass extinction, and erect, toe-walking locomotion was established by the Middle Triassic. The same tracks record longer, more running-adapted toes and shrinking hands before true dinosaurs evolved.
What the new footprint study found
Lorenzo Marchetti and colleagues describe the footprint type Pachychirotherium gardenaensis in Palaeontology (2026; DOI 10.1111/pala.70089). The tracks were first identified at GEOPARC Bletterbach in the Italian Dolomites, a UNESCO World Heritage site. Comparisons with archosauriform hand and foot anatomy indicate that the trackmaker was probably an erythrosuchid or an animal with very similar limbs, although a precise species assignment is not possible.
The study’s broader conclusion is evolutionary rather than taxonomic. Archosauriforms acquired an upright stance step by step. Their limbs moved progressively closer to the body, their weight shifted toward the toes (digitigrady), the central toes became relatively longer (a sign of increasing cursoriality), and the hands became smaller. These changes came before the bipedal walking seen in early dinosaurs and some close relatives.
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How a footprint can reveal posture
A skeleton shows what an animal could do; a trackway records what an animal actually did during one episode of movement. As Lorenzo Marchetti of the Museum für Naturkunde Berlin put it: “It is amazing to see how many details on the biology and locomotion of archosauriforms are preserved in fossil footprints, and this becomes even more important when the skeletons are scarce. In fact, trace fossils are the only fossils to directly register the locomotion of living animals.”
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Pace angulation is the key measurement
Pace angulation is the angle between successive footprints made by the same side of the body in a trackway. In the geometric model used for archosauriform locomotion, a sprawling animal projects its legs outward. Its tracks are spread farther apart, and the resulting pace angle is relatively low. An animal carrying its legs beneath the body places its feet closer to the midline, narrows the trackway and lengthens the stride, producing a higher pace angle.
The angle is not a literal photograph of a limb at one instant. Gait, pelvic rotation, limb proportions and body size can all affect it, so researchers use pace angulation as a comparative indicator alongside footprint shape and anatomy.
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The numerical signal across the Triassic
Kubo and Benton’s 2009 analysis measured 461 published Permian and Triassic trackways. Their median pace-angulation values rise through time:
| Interval | Median pace angulation | What it indicates |
|---|---|---|
| Late Permian | 87.5° | Predominantly sprawling locomotion |
| Early Triassic | 139.5° | Large early increase in limb adduction and stance height |
| Middle Triassic | 149° | Continued refinement and wider distribution of erect walking |
| Late Triassic | 156.5° | Highly erect trackway geometry in the measured sample |
The largest numerical jump occurs between the Late Permian and Early Triassic. Kubo and Benton concluded that an erect archosaur posture appeared within the first six million years of recovery after the end-Permian mass extinction, about 252 million years ago. Earlier interpretations based mainly on skeletons had proposed a transition lasting 15–20 million years. The 2026 study presents the whole evolutionary pattern as stepwise: rapid initial change followed by continued refinement through the Triassic, with erect walking complete by the Middle Triassic.
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Sprawling and erect locomotion compared
| Feature | Sprawling locomotion | Erect locomotion |
|---|---|---|
| Leg position | Limbs project outward from the body | Limbs are held closer beneath the body |
| Trackway width | Broad, with feet far from the midline | Narrower, as the feet approach the midline |
| Pace angulation | Lower values | Higher values and generally longer strides |
| Foot contact | Flatter-footed contact is more likely | Digitigrade, with more weight carried on the toes |
| Foot proportions | Less specialized for sustained speed | Relatively longer middle digits and greater cursorial potential |
| Hands | Relatively larger in the earlier condition | Reduced hands as forelimbs became less important in support |
| Later evolutionary consequence | Constrains efficient high-speed movement | Provides conditions that can precede bipedal walking |
Why the posture change mattered for dinosaurs
Walking with the legs under the body improves mechanical support and reduces the side-to-side swinging required by a sprawled gait. Toe-walking and longer central digits can increase stride efficiency, while smaller hands suggest that the forelimbs were becoming less dominant in weight support. Together, those trends created a body plan more compatible with fast terrestrial movement and, later, raising the front of the body off the ground for bipedal travel.
That sequence matters because it places the foundational change before true dinosaurs. The first dinosaurs inherited an already transformed archosauriform locomotor system rather than inventing upright walking from a fully sprawling ancestor in a single evolutionary step.
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Where the tracks come from
Besides the Italian material, the 2026 report discusses comparable Early Triassic Protochirotherium footprints from Germany, Poland and Morocco. This geographic spread helps researchers test whether the pattern was local or part of a wider archosauriform radiation after the end-Permian crisis. The tracks preserve repeated steps, not just isolated impressions, allowing measurements of stride, trackway width and angular geometry.
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What researchers can and cannot identify
Reliable inferences
- Trackways document the locomotion of a living animal at the time the sediment was exposed.
- Repeated geometry can distinguish broad sprawling from more erect movement when many tracks are available.
- Changes in digits, hand size and foot posture can be compared with archosauriform skeletons.
Limits on trackmaker names
- Most trackways cannot be assigned confidently to an exact species.
- Researchers usually identify a broad group by matching digit number, hand and foot shape, proportions and known skeletons.
- Pachychirotherium gardenaensis is therefore best discussed as an archosauriform track, probably erythrosuchid-like, rather than proof that one named species made every similar print.
- Pace angulation should support, not replace, anatomical and sedimentological evidence because gait and body form can alter the measurement.
The evolutionary timeline in one view
- Late Permian: Archosauriform trackways show low median pace angulation (87.5°), consistent with sprawling limbs.
- End-Permian recovery: Within roughly six million years after the mass extinction at about 252 million years ago, the median rises sharply to 139.5° in the Early Triassic.
- Early to Middle Triassic: Erect walking spreads and is refined, alongside digitigrade contact, longer central digits and reduced hands.
- Middle Triassic: The 2026 study considers the transition to erect walking complete across the archosauriform pattern it documents.
- Later Triassic: Median pace angulation reaches 156.5° in the Kubo–Benton dataset, while the locomotor groundwork for later bipedal dinosaurs is already in place.
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