Differences in rhythmicity (relative variance in cycle period) among mammal, fish, and lizard feeding systems have been hypothesized to be associated with differences in their sensorimotor control systems. We tested this hypothesis by examining whether the locomotion of tachymetabolic tetrapods (birds and mammals) is more rhythmic than that of bradymetabolic tetrapods (lizards, alligators, turtles, salamanders). Species averages of intraindividual coefficients of variation in cycle period were compared while controlling for gait and substrate. Variance in locomotor cycle periods is significantly lower in tachymetabolic than in bradymetabolic animals for datasets that include treadmill locomotion, non-treadmill locomotion, or both. When phylogenetic relationships are taken into account the pooled analyses remain significant, whereas the non-treadmill and the treadmill analyses become
Summary1. The life stage suffering the highest predation rate is expected to have the highest escape performance unless developmental or functional constraints interfere. Peak aquatic escape performance in ephemeral pond-breeding amphibians is expected to develop early in the larval period, and metamorphosis is expected to reduce or completely disrupt aquatic escape performance. In anurans, exceptionally low escape performance during metamorphosis creates selection favouring rapid metamorphosis -which minimizes the time individuals spend in the vulnerable transition between tadpole and frog. 2. We investigated the development of aquatic escape performance in the spotted salamander, Ambystoma maculatum (Shaw, 1802), from embryonic development through metamorphosis. We expected performance to peak early in the larval period as hatchlings face high rates of predation but embryos must first develop escape behaviours. We also tested whether escape performance during metamorphosis was intermediate, as predicted by tail fin resorption, or lower than larvae and adults indicating a major physiological disruption. 3. Escape performance shows a complex ontogeny that is first positively influenced by embryonic and early larval development and then negatively correlated with tail resorption and body size. Escape distance was the only performance metric not affected by life stage. In contrast, both escape velocity and duration showed ontogenetic peaks early in the larval period with the lowest performance found in early embryos and adults and intermediate performance during metamorphosis. 4. This pattern suggests that metamorphosis does not impose a major physiological disruption on escape performance. Because spotted salamanders do not pass through a frog-like 'ontogenetic performance valley' during metamorphosis, they may be less subject than anurans to selection favouring rapid metamorphosis. 5. Functional implications of phenotypic variation should be considered in an ontogenetic framework because the relationship between body size and escape performance can be reversed on either side of an ontogenetic performance peak. The assumption that metamorphosis radically disrupts basic functions such as predator evasion does not seem universally warranted and suggests examination of ontogenetic performance trajectories in a diversity of animals with complex life cycles.
To effectively balance investment in predator defenses versus other traits, organisms must accurately assess predation risk. Chemical cues caused by predation events are indicators of risk for prey in a wide variety of systems, but the relationship between how prey perceive risk in relation to the amount of prey consumed by predators is poorly understood. While per capita predation rate is often used as the metric of relative risk, studies aimed at quantifying predator-induced defenses commonly control biomass of prey consumed as the metric of risk. However, biomass consumed can change by altering either the number or size of prey consumed. In this study we determine whether phenotypic plasticity to predator chemical cues depends upon prey biomass consumed, prey number consumed, or both. We examine the growth response of red-eyed treefrog tadpoles (Agalychnis callidryas) to cues from a larval dragonfly (Anax amazili). Biomass consumed was manipulated by either increasing the number of prey while holding individual prey size constant, or by holding the number of prey constant and varying individual prey size. We address two questions. (i) Do prey reduce growth rate in response to chemical cues in a dose dependent manner? (ii) Does the magnitude of the response depend on whether prey consumption increases via number or size of prey? We find that the phenotypic response of prey is an asymptotic function of prey biomass consumed. However, the asymptotic response is higher when more prey are consumed. Our findings have important implications for evaluating past studies and how future experiments should be designed. A stronger response to predation cues generated by more individual prey deaths is consistent with models that predict prey sensitivity to per capita risk, providing a more direct link between empirical and theoretical studies which are often focused on changes in population sizes not individual biomass.
SUMMARYThe limb girdles and lungs of turtles are both located within the bony shell, and therefore limb movements during locomotion could affect breathing performance. A mechanical conflict between locomotion and lung ventilation has been reported in adult green sea turtles, Chelonia mydas, in which breathing stops during terrestrial locomotion and resumes during pauses between bouts of locomotion. We measured lung ventilation during treadmill locomotion using pneumotach masks in three individual Terrapene carolina (mass 304-416 g) and found no consistent mechanical effects of locomotion on breathing performance. Relatively small tidal volumes(2.2±1.4 ml breath-1; mean ± s.d., N=3 individuals) coupled with high breath frequencies(36.6±26.4 breaths min-1; mean ± s.d., N=3 individuals) during locomotion yield mass-specific minute volumes that are higher than any previously reported for turtles (264±64 ml min kg-1; mean ± s.d., N=3 individuals). Minute volume was higher during locomotion than during recovery from exercise(P<0.01; paired t-test), and tidal volumes measured during locomotion were not significantly different from values measured during brief pauses between locomotor bouts or during recovery from exercise(P>0.05; two-way ANOVA). Since locomotion does not appear to conflict with breathing performance, the mechanism of lung ventilation must be either independent of, or coupled to, the stride cycle. The timing of peak airflow from breaths occurring during locomotion does not show any fixed phase relationship with the stride cycle. Additionally, the peak values of inhalatory and exhalatory airflow rates do not differ consistently with respect to the stride cycle. Together, these data indicate that T. carolina is not using respiratory-locomotor coupling and limb and girdle movements do not contribute to lung ventilation during locomotion. X-ray video recordings indicate that lung ventilation is achieved via bilateral activity of the transverse (exhalatory) and oblique (inhalatory) abdominal muscles. This specialized abdominal ventilation mechanism may have originally circumvented a mechanical conflict between breathing and locomotion in the ancestor of turtles and subsequently allowed the ribs to abandon their role in lung ventilation and to fuse to form the shell.
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