Araeoscelis gracilis was a superficially lizard-like animal that lived during the mid-Permian, around 275 million years ago, in what is now Texas, USA. About 60cm long (~2′), it had a slender body, proportionally long legs, and a solidly-built skull with strong teeth, suggesting that it was a fast runner that specialized in cracking open the carapaces of thick-shelled prey.

It was one of the last known members of a lineage known as araeoscelidians, which are usually considered to be very early members of the diapsid reptiles – but a recent study has proposed they might have even more ancient roots than that, possibly being a branch of stemamniotes instead.


With its armored head and blade-like jaws, the placoderm fish Dunkleosteus terrelli is an iconic Paleozoic animal.

Living during the Late Devonian, about 375-359 million years ago, in subtropical waters covering parts of what are now North America and Europe, this species is known mostly just from the bony plates that covered its head and thorax. The rest of its skeleton was cartilaginous and rarely ever fossilized (only a few vertebrae and the pectoral fin are currently known), so its full body shape and size is poorly understood, and previous length estimates have ranged all the way up to 10m (33′).

…Except it turns out it wasn’t nearly that big.

Based on its head proportions, along with comparisons to more complete remains of other arthrodire placoderms, recent studies instead come up with a maximum length of about 4m (~13ft) – giving Dunkleosteus a much shorter-but-heavier chunky body shape, more like a tuna than a shark.

But even after this size revision Dunkleosteus would have still been one of the largest animals around at the time, with the ability to snap its jaws open at high speed and an incredibly strong bite force. It was probably specialized to mainly prey on other heavily-armored animals such as other placoderms and shelled cephalopods, and was likely a strong swimmer with a shark-like tail fin.

Preserved stomach contents in one fossil show remains of the fast-swimming cartilaginous fish Orodus – suggesting that much like the modern tuna it resembled, Dunkleosteus was also capable of bursts of high speed.


Noripterus complicidens was a pterosaur that lived in what is now Mongolia and northwest China during the early Cretaceous, about 140 million years ago.

It had a wingspan of up to 4m (~13′), with a head-and-body length of around 1.2m (~4′), and like other dsungaripterids it had a distinctively reinforced skull, with a toothless beak at the front of its jaws and strong widely-spaced teeth further back – an arrangement that was probably used to catch and then powerfully crack open hard-shelled prey.

Unlike other pterosaurs, however, Noripterus also had some very unusual feet.

All other known pterosaurs seem to have had plantigrade hindlimbs, standing and walking with the whole foot on the ground. But Noripterus had toe joints that looked more like those of theropod dinosaurs than other pterosaurs, with a higher level of upward flexibility and potentially a more digitigrade posture standing on just its toes.

With dsungaripterids already having fairly stout body proportions that suggest they spent a lot of time walking around on the ground, Noripterus may have been even more agile and adept at terrestrial locomotion. Digitigrady is generally more efficient for moving at higher speeds, so this pterosaur might have been a runner behaving similarly to modern ground birds, preferring to sprint away from threats on foot rather than launch itself into the air.


Cabarzia trostheidei here lived during the early Permian in what is now Germany, about 295 million years ago.

Despite its very lizard-like appearance it was actually part of the varanopid lineage, a group of scaly amniotes traditionally classified as early synapsids (distant relatives of modern mammals), but which more recently have been proposed to instead be sauropsid reptiles closer related to early diapsids.

It was around 50cm long (1’8″), and its short arms, long legs, slender body, and long tail suggest it was capable of shifting into a bipedal posture when running at high speeds, similarly to some modern lizards – probably mainly to escape from larger predators, but possibly also used to pursue fast-moving prey like flying insects.

And whether varanopids were actually synapsids or sauropsids, this makes Cabarzia the earliest known example of an animal running on two legs.


For around 50 years some very unusual dinosaur tracks have been found in ancient desert sediments in South America: strange footprints showing the impression of only a single toe, a walking style never before seen in any reptiles.

And recently a fossil of what might be the track maker has actually been found.

Named Vespersaurus paranaensis, this new species lived during the Late Cretaceous of Brazil (~90 mya) and was a member of the noasaurid family of theropods, closely related to the weird-jawed Masiakasaurus from Madagascar.

Measuring about 1.5m long (~5′), Vespersaurus was fairly lightly built with legs proportioned for running – and its feet were absolutely unique. Although it had the standard three main toes of a theropod, it bore its weight entirely on the middle toe and held the other digits off the ground. The two raised toes on each foot also had large knife-like claws which may have been used during hunting, vaguely similar to the sickle claws on the feet of dromaeosaurs. But unlike dromaeosaurs these claws weren’t highly curved or pointed, suggesting Vespersaurus used more of a scratching and slashing technique rather than the raptors’ puncture-and-restraint strategy.

Much like ancient horses, it may have developed its single-toed stance as an adaptation for more efficient fast running, possibly to avoid larger predators or to chase down small fast-moving prey like hopping desert mammals.

The known one-toed fossil footprints are actually slightly older than the Vespersaurus fossil, and similar tracks in Argentina have been found dating back to the Late Jurassic (~150mya), so there may have been a long lineage of “one-toed” desert-dwelling noasaurids in South America that haven’t been found yet.


Eudibamus cursoris, a bolosaurid from the Early Permian of Germany (~284-279 mya).

Although very lizard-like in appearance, this animal was actually part of a completely extinct group known as parareptiles – a diverse group of early sauropsids who were once thought to be the ancestors of turtles, but are now considered to instead be the evolutionary cousins to the true reptiles.

With a total length of about 25cm long (8-10″), the structure and proportions of its limbs suggest it could run fast on its hind legs, making it one of the earliest known examples of bipedal locomotion. Since its teeth were adapted for a herbivorous diet, it wasn’t using its speed to chase down prey but was instead probably sprinting away from predators.

But unlike the sprawling running of some modern lizards, Eudibamus may have been capable of holding its legs in a more upright position directly under its body, convergently evolving a more energy-efficient posture similar to that of later bipedal animals like dinosaurs.


Ergilornis rapidus, a 1.2-1.5m tall bird (4′-5′) from the Early Oligocene of Mongolia (~33-28 mya). Closely related to modern cranes, trumpeters, and limpkins, it was part of an extinct group called eogruids – flightless birds which existed across Eurasia for a large portion of the Cenozoic from roughly 40-3 million years ago.

Although the earliest known eogruids were smaller and less specialized, and may even have still been somewhat capable of flying, later forms like Ergilornis had highly reduced wings, long legs adapted for running, and convergently ostrich-like feet with only two toes each.


Boverisuchus magnifrons*, a crocodilian from the early Eocene of Germany (~50-40 mya). Reaching about 3m long (9′10″) it was much more heavily armored than its modern cousins, with an interlocking “exoskeleton” of bony osteoderms covering its body and limbs – leading to it being given the nickname “panzer croc”.

It was adapted for walking and running on land, with relatively long legs and surprisingly hoof-like claws. It may even have carried its weight directly on these hooves similar to mammalian ungulates.

And if that’s not unusual enough, its hind leg musculature suggests it also might have been capable of short bursts of bipedal sprinting.

[ * Originally known as Pristichampsus rollinatii before being reassigned in 2013.]