Stride frequency or length? A phylogenetic approach to understand how animals regulate locomotor speed

Speed regulation in animals involves stride frequency and stride length. While the relationship between these variables has been well documented, it remains unresolved whether animals primarily modify stride frequency or stride length to increase speed. In this study, we explored the interrelationships between these three variables across a sample of 103 tetrapods and assessed whether speed regulation strategy is influenced by mechanical, allometric, phylogenetic or ecological factors. We observed that crouched terrestrial species tend to regulate speed through stride frequency. Such a strategy is energetically costly, but results in greater locomotor maneuverability and greater stability. In contrast, regulating speed through stride length is closely tied to larger arboreal animals with relatively extended limbs. Such movements reduce substrate oscillations on thin arboreal supports and/or helps to reduce swing phase costs. The slope of speed on frequency is lower in small crouched animals than in large-bodied erect species. As a result, substantially more rapid limb movements are matched with only small speed increases in crouched, small-bodied animals. Furthermore, the slope of speed on stride length was inversely proportional to body mass. As such, small changes in stride length can result in relatively rapid speed increases for small-bodied species. These results are somewhat counterintuitive, in that larger species, which have longer limbs and take longer strides, do not appear to gain as much speed increase out of lengthening their stride. Conversely, smaller species that cycle their limbs rapidly do not gain as much speed out of increasing stride frequency as do larger species.