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For walking animals (which we define as encompassing any form of legged movement across the ground, such as walking, running, galloping, etc.), such substrate \u2018roughness\u2019 influences locomotion in a multitude of ways across scales, from roughness that influences how each toe or foot contacts the ground, to larger obstacles that animals must move over or navigate around. Historically, the unpredictability and variability of natural environments has limited the ability to collect data on animal walking biomechanics. However, recent technical advances, such as more sensitive and portable cameras, biologgers, laboratory tools to fabricate rough terrain, as well as the ability to efficiently store and analyze large variable datasets, have expanded the opportunity to study how animals move under naturalistic conditions. As more researchers endeavor to assess walking over rough terrain, we lack a consistent approach to quantifying roughness and contextualizing these findings. This Review summarizes existing literature that examines non-human animals walking on rough terrain and presents a metric for characterizing the relative substrate roughness compared with animal size. This framework can be applied across terrain and body scales, facilitating direct comparisons of walking over rough surfaces in animals ranging in size from ants to elephants.<\/jats:p>","DOI":"10.1242\/jeb.245261","type":"journal-article","created":{"date-parts":[[2023,4,21]],"date-time":"2023-04-21T09:51:43Z","timestamp":1682070703000},"source":"Crossref","is-referenced-by-count":20,"title":["The bumpy road ahead: the role of substrate roughness on animal walking and a proposed comparative metric"],"prefix":"10.1242","volume":"226","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5806-7254","authenticated-orcid":false,"given":"Glenna","family":"Clifton","sequence":"first","affiliation":[{"name":"Department of Biology, University of Portland 1 , OR , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4217-2850","authenticated-orcid":false,"given":"Alyssa Y.","family":"Stark","sequence":"additional","affiliation":[{"name":"Department of Biology, Villanova University 2 , PA , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7516-3646","authenticated-orcid":false,"given":"Chen","family":"Li","sequence":"additional","affiliation":[{"name":"Department of Mechanical Engineering, Johns Hopkins University 3 , MD , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9391-2476","authenticated-orcid":false,"given":"Nicholas","family":"Gravish","sequence":"additional","affiliation":[{"name":"Department of Mechanical and Aerospace Engineering, University of California San Diego, 4 San Diego, CA , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2023,4,21]]},"reference":[{"key":"2023042109494461100_JEB245261C1","doi-asserted-by":"crossref","DOI":"10.1007\/s00265-021-03047-8","article-title":"The characteristics and consequences of African wild dog (Lycaon pictus) den site selection","volume":"75","author":"Alting","year":"2021","journal-title":"Behav. 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