Can a turtle tuck its head all the way inside its shell?
Turtle shells evolved over the course of 300 million years, but self-defense wasn't the initial driver, researchers think.
It's a long-held idea that turtles can tuck their heads into their shells when threatened. But is it true? And is this protective trick why turtles the world over have shells today?
The answer is that some types of turtles can, and others can't, experts told Live Science. And even though shells can be protective for some of these reptiles, fossil evidence suggests that shells evolved for entirely different reasons.
Tortoises are one type of turtle that can tuck their heads into their shells. This terrestrial subgroup of turtles emerged 50 million years ago, Tyler Lyson, senior curator of vertebrate paleontology at the Denver Museum of Nature and Science, told Live Science. They typically move slowly, so they rely on their shells to protect them from predators. Most tortoises can draw their heads into their shells, which typically also have a domed shape with more space inside to make that possible.
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Several terrestrial turtle species, which split their time between land and water, can do the same.
"Turtles have two ways of tucking the head in," Jason Head, a professor of vertebrate evolution and ecology at the University of Cambridge, told Live Science. "We have what are called the side-neck turtles. They have long necks, and they literally fold the head and neck to the side over one of their arms. And then there are the snake-neck or S-neck turtles, which put a loop into the neck, and can actually pull the neck into the shoulder girdle."
One example is the eastern box turtle (Terrapene carolina carolina), whose bottom shell, known as a plastron, is fitted with a hinge that even allows it to completely close up the shell.
But sea turtles are one group of turtles that cannot pull their heads into their shells. Sea turtles have much sleeker, lighter shells that contain no space for them to tuck their heads inside. "This is to lighten the load," Head said, and it allows sea turtles to swim faster to escape predators.
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How turtle shells evolved
So, how did some turtles develop this lifesaving trick? To find out, we need to explore how turtle shells evolved, which takes us back almost 300 million years in the fossil record.
"The turtle shell is a complicated structure. It's made up of over 50 bones," Lyson said. "Bone" is the key word, because fossils reveal that turtle shells are part of their skeletons. And while the modern turtle's shell looks like a solid unit, it's actually made up of two skeletal features that evolved separately.
"The first thing we see in the evolution of the turtle shell is the broadening of the ribs, and we see that in Eunotosaurus africanus," a creature that lived in southern Africa 260 million years ago, before dinosaurs roamed Earth, Lyson told Live Science. Lyson first described Eunotosaurus' contribution to turtle evolution in a 2013 study. Researchers think that these creatures spent time burrowing underground to escape the heat and that the development of wider ribs supported more muscle mass that enabled them to do that.
Then, in Germany, the 2015 discovery of a 240 million-year-old fossil called Pappochelys showed a shell-less animal with wider upper ribs paired with thicker belly ribs — known as "gastralia" — on its underside. By 220 million years ago, an aquatic animal called Odontochelys found in China had developed a fully unified belly plate — the plastron — partly from the expanding gastralia.
"Myself and others think that the evolution of the plastron was a ballast for basically going deeper into the water column," Lyson explained. It's also possible the plastron developed to protect turtles from predators swimming below, he noted.
The first evidence of a fully formed turtle shell comes from 210 million years ago, in the shape of a fossilized creature called Proganochelys, whose thick upper ribs had fused together with dermal bone, forming a closed carapace, attached to a lower plastron. The opening for the turtle's head was formed from shoulder bones that connected the top and bottom of its shell, Lyson explained.
Most evidence suggests that these reptilian creatures, called Pantestudines, ultimately led to modern-day turtles. However, Head noted that similar features — like widened, overlapping ribs — also developed in other animals millions of years ago, including some thought to be more closely related to mammals.
"It's an active area of research, with new discoveries coming all the time," Head said.
The shells of these turtle ancestors developed as a response to varied evolutionary pressures, but today, the turtle's shell is used primarily for self-defense, Lyson noted. "The modern-day function isn't necessarily related to how that feature arose," he said. "It wasn't until you got the full advent of the shell that it was for protection."
The turtle's resilient shell has seen these creatures through almost 300 million years of history, and Lyson thinks it's one reason they've managed to survive three of Earth's five mass extinctions.
"We see the fossil record, and we can see the line in the sand where dinosaurs and lots of other things go extinct," Lyson said. "And we see turtles marching right across that line."
Evolution quiz: Can you naturally select the correct answers?

Emma Bryce is a London-based freelance journalist who writes primarily about the environment, conservation and climate change. She has written for The Guardian, Wired Magazine, TED Ed, Anthropocene, China Dialogue, and Yale e360 among others, and has masters degree in science, health, and environmental reporting from New York University. Emma has been awarded reporting grants from the European Journalism Centre, and in 2016 received an International Reporting Project fellowship to attend the COP22 climate conference in Morocco.
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