Virtual resurrection shows that early four-legged animal couldn’t walk very well

In a small office north of London, Stephanie Pierce from the Royal Veterinary College is watching a movement that hasn’t been seen for 360 million years. On her computer, she has resurrected the long-extinct Ichthyostega – one of the earliest four-legged animals to creep about on land. By recreating this iconic beast as a virtual skeleton, Pierce has shown that while it looked like a giant salamander, it couldn’t possibly have walked like one. It had some of the planet’s earliest bony legs, but they weren’t very good at taking steps.

Ichthyostega hails from the Devonian period, a time in Earth’s history when swimming transformed into walking. Fish invaded the land and evolved into the first tetrapods—four-limbed animals that include amphibians, reptiles, birds and mammals.  Muscular fins used for steering and balance evolved into legs for walking.

Since its discovery over 50 years ago, Ichthyostega has been an icon of this pivotal transition. Some 300 specimens have been found but many are incomplete, flattened or distorted. Pierce’s new model provides the best look yet at the animal’s skeleton. “It makes Ichthyostega a bit more tangible,” she says. “It’s not just a fossil laying in a rock now. It’s an animal that’s coming to life.”

Pierce built her virtual skeleton by putting dozens of Ichthyostega specimens in powerful CT-scanners, choosing only the best preserved ones out of the 300 or so in existence. “The front end of the animal was mainly composed from one beautifully preserved specimen called ‘Mr Magic’,” she says.

It was painstaking work. These fossils are so old that chemically, they are almost identical to the rocks around them. By eye, the bones stand out. To the scanners, they blend in. Pierce spent over two years scanning the specimens and building her model, but the results were worth it. “This has been on the wish-list for years,” says Michael Coates, who studies tetrapod evolution at the University of Chicago.

Those boots weren’t made for walking…

The model showed that Ichthyostega’s shoulders and hips were oddly restricted. They could move back and forth, and up and down, but they couldn’t rotate about their long axis. Hold your arm out and rotate your palm so it faces up then down—Ichthyostega’s shoulder couldn’t do that.

Most modern tetrapods need long-axis rotation in order to walk. Without it, their legs can’t be thrown forward or pulled backward. Ichthyostega’s limitations meant that despite having four limbs, it probably couldn’t have taken a step. It hind feet would never have been planted flat against the ground or supported its weight. It had invaded the land, but it wasn’t striding across it.

“It highlights the fact that the earliest tetrapods are not just ‘gigantic salamanders’, despite a vague similarity in outline,” says Per Ahlberg from Uppsala University. “The limbs and girdles are very different from anything now living.”

Pierce thinks that Ichthyostega moved by paddle with its front limbs, using powerful muscles and flexible elbows to make rowing motions. The closest living analogue is probably the mudskipper – a fish that drags itself along muddy land with its front fins (as in the video below).

Pierce also compared Ichthyostega’s joints and limbs to those of other living animals with sinuous bodies and interesting gaits, including a salamander, crocodile, seal, otter and platypus. Compared to these modern species, Ichthyostega’s hips and shoulders were similarly flexible in most planes of movements, but along their long axis, they could barely rotate.

Some scientists think that the tetrapods evolved limbs before they could walk, and their first members lived in shallow water. Others think that it’s the other way round, and that muscular limbs, hips and shoulders evolved while fish still had fins. The virtual Ichthyostega supports the former idea, since it had limbs but couldn’t walk. But Coates cautions against “fitting a smooth transition from swimmers to walkers.” He says, “Evolutionary transitions needn’t follow linear routes. Ichthyostega probably represents one of multiple experiments among the first tetrapods with limbs, trying-out life in the shallows.”

So… what made those tracks?

Other early tetrapods had similar shoulders and hips, so they probably had the same limitations too. John Hutchinson, who led the new study, plans to find out. His lab is busy reconstructing other early tetrapods including Acanthostega, one of Ichthyostega’s contemporaries, and Pederpes, a later model.

But Ahlberg notes that Ichthyostega had a very unusual and rigid spine, and may not have been representative of other early tetrapods. “Other tetrapods are known to have had more flexible spines” he says, “and this probably allowed them to overcome the limitations of their shoulders and hips”.

This might explain why Ahlberg and others have discovered tracks that pre-date Ichthyostega by around 20 million years, and had become fairly common by the time it evolved. Many of these tracks showed precisely the kind of salamander-like movements that Ichthyostega was apparently incapable of making. They were clearly made by early four-legged tetrapods, and to this date, we don’t know what made them.

Pierce agrees that the final word on Ichthyostega’s movements will have to wait until she can animate its entire skeleton. “The ultimate goal would be to try and create some sort of dynamic movement,” she says. She has applied for a grant to do just that, to model the motions of the entire animal, and compare them to salamanders or crocodiles. “That’s going to take so much time, but it’ll be very interesting,” she says.

PS: I want to point out that in researching this story, I spent a good minute on my living room floor trying to walk without long-axis rotation. It was really hard, and I looked like an idiot. I did a similar thing when I was writing about hummingbird wing movements for Nature. I’m going to christen this Method Science Journalism.

Reference: Pierce, Clack & Hutchinson. 2012. Three-dimensional limb joint mobility in the early tetrapod Ichthyostega. Nature http://dx.doi.org/10.1038/nature11124

Image by Julie Molnar

More on tetrapods:

Fossil tracks push back the invasion of land by 18 million years

Fish fins and mouse feet controlled by the same ancient genetic switch