Scale effects and morphological diversification in hindlimb segment mass proportions in neognath birds

Introduction: In spite of considerable work on the linear proportions of limbs in amniotes, it remains unknown whether differences in scale effects between proximal and distal limb segments has the potential to influence locomotor costs in amniote lineages and how changes in the mass proportions of limbs have factored into amniote diversification. To broaden our understanding of how the mass proportions of limbs vary within amniote lineages, I collected data on hindlimb segment masses - thigh, shank, pes, tarsometatarsal segment, and digits - from 38 species of neognath birds, one of the most speciose amniote clades. I scaled each of these traits against measures of body size (body mass) and hindlimb size (hindlimb length) to test for departures from isometry. Additionally, I applied two parameters of trait evolution (Pagel's lambda and delta) to understand patterns of diversification in hindlimb segment mass in neognaths. Results: All segment masses are positively allometric with body mass. Segment masses are isometric with hindlimb length. When examining scale effects in the neognath subclade Land Birds, segment masses were again positively allometric with body mass; however, shank, pedal, and tarsometatarsal segment masses were also positively allometric with hindlimb length. Methods of branch length scaling to detect phylogenetic signal (i.e., Pagel's lambda) and increasing or decreasing rates of trait change over time (i.e., Pagel's delta) suffer from wide confidence intervals, likely due to small sample size and deep divergence times. Conclusions: The scaling of segment masses appears to be more strongly related to the scaling of limb bone mass as opposed to length, and the scaling of hindlimb mass distribution is more a function of scale effects in limb posture than proximo- distal differences in the scaling of limb segment mass. Though negative allometry of segment masses appears to be precluded by the need for mechanically sound limbs, the positive allometry of segment masses relative to body mass may underlie scale effects in stride frequency and length between smaller and larger neognaths. While variation in linear proportions of limbs appear to be governed by developmental mechanisms, variation in mass proportions does not appear to be constrained so.