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By "running" I mean a land gait that at one point has all propulsion limbs off the ground. Jumping, like a flea or a frog, is not running.

The earliest land vertebrates could not run, I assume. The ability to run would be an advantage when pursuing prey or evading predators.

Our data from the period where running would first evolve is limited to bones and footprints. Is that enough to tell what an extinct animal's gait would be?

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    $\begingroup$ Things like sea scorpions walked on the ocean bed so shouldn't be discounted, especially when buoyancy makes it a lot easier to satisfy for your running criteria. $\endgroup$
    – DKNguyen
    Dec 12, 2022 at 14:34
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    $\begingroup$ Welcome to the site. Interesting question - but you could improve it by telling us what you have found already, and what criteria you measure running by. Note that the help center has a page on "homework" questions and what you can do to avoid getting closed for this reason. $\endgroup$
    – bob1
    Dec 12, 2022 at 23:06

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