Scaling effect on the mid-diaphysis properties of long bones—the case of the Cervidae (deer)

Abstract

How skeletal elements scale to size is a fundamental question in biology. While the external shape of long bones was intensively studied, an important component of their organization is also found in their less accessible inner structure. Here, we studied mid-diaphyseal properties of limb long bones, characterizing notably the thickness of their cortices (bone walls), in order to test whether body size directly influences bone inner organization. Previous examinations of scaling in long bones used broad samplings to encompass a wide range of body sizes. To account for the effect of confounding factors related to different lifestyles, we focused our comprehensive sampling on a mammalian clade that comprises various body sizes but a relatively uniform lifestyle, the Cervidae. Positive allometry was found in femoral cross-sectional shape, indicating greater directional bending rigidity in large-sized taxa. None of the compactness parameters scaled allometrically in any of their bones. The cortices of sampled zeugopodial bones (tibia and radius) were found as significantly thicker than those of stylopodial bones (femur and humerus). Furthermore, while the mean relative cortical thickness values for both stylopodial and zeugopodial bones are close to mass-saving optima, the variance for the stylopodial bones is significantly lower. This suggests that mass saving is less intensively selected in zeugopodial bones. Finally, the long-legged Elk (Alces) and the short-legged dwarf Cretan deer (Candiacervus) featured rather thin and thick cortices, respectively, suggesting that the acquisition of a different limb proportion is accompanied by a modification of the relative mid-diaphyseal cortical thickness.