New Issue: Orbital Catastrophe Ahead? Read Now

Legs, Feet and Toes

The essential parts for walking on land evolved in water

Join Our Community of Science Lovers!


On supporting science journalism

If you're enjoying this article, consider supporting our award-winning journalism by subscribing. By purchasing a subscription you are helping to ensure the future of impactful stories about the discoveries and ideas shaping our world today.


The evolution of terrestrial creatures from aquatic fish with fins may have begun with the need for a breath of fresh air. Animals with limbs, feet and toes—a group known as the tetrapods (literally, “four-footed”)—arose between 380 million and 375 million years ago. Scientists long believed that limbs evolved as an adaptation to life on terra firma. But recent discoveries have revealed that some of the key changes involved in the fin-to-limb transition occurred while the ancestors of tetrapods were still living in the water.

Tetrapod evolution experts such as Jennifer Clack of the University of Cambridge hypothesize that these early modifications to the bones and joint surfaces of the pectoral fins might have benefited tetrapod ancestors in two key ways. First, they could have allowed the creatures, which lived in the plant-choked shallows, to perform a push-up that raised their heads out of the oxygen-poor water for a breather. (Changes in other parts of the skeleton, such as the skull and neck, also facilitated air breathing.) The protolimbs could have also helped these animals to propel themselves along the bottom and to steady themselves against the current while waiting to ambush prey.

Researchers once thought that the bones making up feet and toes were an evolutionary innovation unique to the tetrapods. But over the past few years analyses of tetrapod forerunners, such as the Tiktaalik fossil unveiled in 2006, have revealed that these bones derive directly from bones in the fish fin. Curiously, the earliest tetrapods and tetrapodlike fish had feet with between six and eight digits, rather than the five of most modern tetrapods. Why tetrapods ultimately evolved a five-digit foot is uncertain, but this arrangement may have provided the ankle joint with the stability and flexibility needed for walking.

Kate Wong is an award-winning science writer and was formerly senior editor for features at Scientific American, where she focused on evolution, ecology, anthropology, archaeology, paleontology and animal behavior. She is fascinated by human origins, which she has covered for nearly 30 years. Recently she has become obsessed with birds. Her reporting has taken her to caves in France and Croatia that Neandertals once called home to the shores of Kenya’s Lake Turkana in search of the oldest stone tools in the world, as well as to Madagascar on an expedition to unearth ancient mammals and dinosaurs, the icy waters of Antarctica, where humpback whales feast on krill, and a “Big Day” race around the state of Connecticut to find as many bird species as possible in 24 hours. Wong is co-author, with Donald Johanson, of Lucy’s Legacy: The Quest for Human Origins. She holds a bachelor of science degree in biological anthropology and zoology from the University of Michigan. Follow her on Bluesky @katewong.bsky.social

More by Kate Wong
Scientific American Magazine Vol 301 Issue 3This article was published with the title “Legs, Feet and Toes” in Scientific American Magazine Vol. 301 No. 3 (), p. 80
doi:10.1038/scientificamerican0909-80b

Subscribe to Support Independent Journalism

Great science journalism requires human expertise, time, effort and creativity. And it costs money. That’s why I and the journalists here at Scientific American hope you’ll join our community.

When you subscribe, you are supporting staff and freelance journalists who are passionate about telling science stories that are true, important and compelling. Our editors and reporters are often experts in their fields, which means they understand the nuances of big discoveries and can untangle the breakthroughs from the hype. With a subscription, you are also supporting rigorous fact-checking to ensure the words we publish are precise and accurate. And you’re supporting original illustrations, graphics and photos that bring you closer to an advanced laboratory, an ice sheet in Antarctica or a space mission in orbit. You’re helping us craft other types of high-quality journalism as well: Our newsletters are carefully written, edited and curated by staffers you have or will come to know and love. Our Science Quickly podcast is based on original reporting, collaboration with editors and scientists and exacting production.

Subscriptions keep this engine running so we can continue to deliver thoughtful, rigorous and independent science journalism to you. In an era of viral misinformation, this work is crucial. If you value what we do, I hope you’ll consider joining us as a subscriber

Thank you,

Jeanna Bryner, Editor in Chief, Scientific American

Subscribe