Locomotor energetics in the Indonesian blue-tongued skink (Tiliqua gigas) with implications for the cost of belly-dragging in early tetrapods.

Abstract

During the last decade, biomechanical and kinematic studies have suggested that a belly-dragging gait may have represented a critical locomotor stage during tetrapod evolution. This form of locomotion is hypothesized to facilitate animals to move on land with relatively weaker pectoral muscles. The Indonesian blue-tongued skink (Tiliqua gigas) is known for its belly-dragging locomotion and is thought to employ many of the same spatiotemporal gait characteristics of stem tetrapods. Conversely, the savannah monitor (Varanus exanthematicus) employs a raised quadrupedal gait. Thus, differences in the energetic efficiency of locomotion between these taxa may elucidate the role of energetic optimization in driving gait shifts in early tetrapods. Five Tiliqua and four Varanus were custom-fitted for 3D printed helmets that, combined with a Field Metabolic System, were used to collect open-flow respirometry data including O2 consumption, CO2 production, water vapor pressure, barometric pressure, room temperature, and airflow rates. Energetic data were collected for each species at rest, and when walking at three different speeds. Energetic consumption in each taxon increased at greater speeds. On a per-stride basis, energetic costs appear similar between taxa. However, significant differences were observed interspecifically in terms of net cost of transport. Overall, energy expenditure was ~20% higher in Tiliqua at equivalent speeds, suggesting that belly-dragging does impart a tangible energetic cost during quadrupedal locomotion. This cost, coupled with the other practical constraints of belly-dragging (e.g., restricting top-end speed and reducing maneuverability in complex terrains) may have contributed to the adoption of upright quadrupedal walking throughout tetrapod locomotor evolution.