Size and shape of terrestrial animals

TL;DR

This study demonstrates that the size-dependent frontal aspect ratio in terrestrial animals is driven by natural terrain roughness, influencing stability needs across diverse species regardless of their anatomy or behavior.

Contribution

It reveals that terrain roughness, rather than anatomy or behavior, primarily influences the evolution of animal body shape for stability during terrestrial locomotion.

Findings

Size-dependent scaling of frontal aspect ratio across species

Fractal terrain roughness explains shape evolution

Shared ancestry does not account for shape scaling

Abstract

Natural selection for terrestrial locomotion has yielded unifying patterns in the body shape of legged animals, often manifesting as scaling laws. One such pattern appears in the frontal aspect ratio. Smaller animals like insects typically adopt a landscape frontal aspect ratio, with a wider side-to-side base of support than center of mass height. Larger animals like elephants, however, are taller than wide with a portrait aspect ratio. Known explanations for postural scaling are restricted to animal groups with similar anatomical and behavioural motifs, but the trend in frontal aspect ratio transcends such commonalities. Here we show that vertebrates and invertebrates with diverse body plans, ranging in mass from 28 mg to 22000 kg, exhibit size-dependent scaling of the frontal aspect ratio driven by the need for lateral stability on uneven natural terrain. Because natural terrain exhibit scale-dependent unevenness, and the frontal aspect ratio is important for lateral stability during locomotion, smaller animals need a wider aspect ratio for stability. This prediction is based on the fractal property of natural terrain unevenness, requires no anatomical or behavioural parameters, and agrees with the measured scaling despite vast anatomical and behavioural differences. Furthermore, a statistical phylogenetic comparative analysis found that shared ancestry and random trait evolution cannot explain the measured scaling. Thus, our findings reveal that terrain roughness, acting through natural selection for stability, likely drove the macroevolution of frontal shape in terrestrial animals.