Locomotion, Energetics, Performance, and Behavior: A Mammalian Perspective on Lizards, and Vice Versa.

Animals are constrained by their abilities and by interactions with environmental factors, such as low ambient temperatures. These constraints range from physical impossibilities to energetic inefficiencies, and may entail trade-offs. Some of the constraints related to locomotion and activity metabolism can be illustrated through allometric comparisons of mammals and lizards, as representative terrestrial vertebrate endotherms and ectotherms, respectively, because these lineages differ greatly in aerobic metabolic capacities, resting energetic costs, and thermoregulatory patterns. Allometric comparisons are both useful and unavoidable, but "outlier" species (unusual for their clade) can also inform evolutionary scenarios, as they help indicate extremes of possible adaptation within mammalian and saurian levels of organization. We compared mammals and lizards for standard metabolic rate (SMR), maximal oxygen consumption during forced exercise (VO2max), net (incremental) cost of transport (NCT), maximal aerobic speed (MAS), daily movement distance (DMD), daily energy expenditure (DEE) during the active season, and the ecological cost of transport (ECT = percentage of DEE attributable to locomotion). (Snakes were excluded because their limbless locomotion has no counterpart in terrestrial mammals.) We only considered lizard SMR, VO2max, NCT, MAS, and sprint speed data if measured at 35-40 °C. On average, MAS is ∼7.4-fold higher in mammals, whereas SMR and VO2max are ∼6-fold greater, but values for all three of these traits overlap (or almost overlap) between mammals and lizards, a fact that has not previously been appreciated. Previous studies show that sprint speeds are similar for smaller mammals and lizards, but at larger sizes lizards are not as fast as some mammals. Mammals move ∼6-fold further each day than lizards, and DMD is by far the most variable trait considered here, but their NCT is similar. Mammals exceed lizards by ∼11.4-fold for DEE. On average for both lineages, the ECT is surprisingly low, somewhat higher for lizards, and positively allometric. If a lizard and mammal of 100 g body mass were both to move their entire DMD at their MAS, they could do so in ∼21 and 17 min, respectively, thus de-emphasizing the possible importance of time constraints. We conclude that ecological-energetic constraints related to locomotion are relatively more likely to occur in large, carnivorous lizards. Overall, our comparisons support the idea that the (gradual) evolution of mammalian endothermy did not necessarily require major changes in locomotor energetics, performance, or associated behaviors. Instead, we speculate that the evolution of thermoregulatory responses to low temperatures (e.g., shivering) may have been a key and "difficult" step in this transition.