Lance D. McBrayer

Prey processing in amniotes: biomechanical and behavioral patterns of food reduction.

In this paper we examine the biomechanics of prey processing behavior in the amniotes. Whether amniotes swallow prey items whole or swallow highly processed slurries or boluses of food, they share a common biomechanical system where hard surfaces (teeth or beaks) are brought together on articulated jaws by the actions of adductor muscles to grasp and process food. How have amniotes modified this basic system to increase the chewing efficiency of the system? To address this question we first examine the primitive condition for prey processing representative of many of the past and present predatory amniotes. Because herbivory is expected to be related to improved prey processing in the jaws we review patterns of food processing mechanics in past and present herbivores. Herbivory has appeared numerous times in amniotes and several solutions to the task of chewing plant matter have appeared. Birds have abandoned jaw chewing in favor of a new way to chew--with the gut--so we will detour from the jaws to examine the appearance of gut chewing in the archosaurs. We will then fill in the gaps among amniote taxa with a look at some new data on patterns of prey processing behavior and jaw mechanics in lizards. Finally, we examine evolutionary patterns of amniote feeding mechanism and how correlates of chewing relate to the need to increase the efficiency of prey processing in order to facilitate increased metabolic rate and activity.

How muscles define maximum running performance in lizards: an analysis using swing- and stance-phase muscles

SUMMARY Maximum locomotor performance is crucial for capturing prey, escaping predators and many other behaviors. However, we know little about what defines maximum performance in vertebrates. Muscles drive the movement of the limbs during locomotion, and thus likely play a major role in defining locomotor capacity. For lizards, the iliofibularis, a swing-phase muscle, is often linked to ecology and/or performance. However, stance-phase muscles likely limit performance given that they propel the animal. Using a small semi-arboreal lizard (Sceloporus woodi), we compared how swing- and stance-phase muscles relate to maximum running speed and acceleration. We employed both a level and vertical trackway to elicit ecologically relevant locomotor performance. Six individuals were filmed at 250 frames s–1 in lateral view. Following performance trials, upper and lower hindlimbs were sectioned and assessed using histochemistry. Fast glycolytic, fast oxidative and slow oxidative fibers were detected and counted in the gastrocnemius (GA; stance phase) and iliofibularis (IF; swing phase) muscles. In addition, the mean fiber diameter for each fiber type in each muscle was determined, as was the fiber cross-sectional area. We found that properties of the GA, but not the IF, were positively correlated with performance. Interestingly, certain attributes of the GA were correlated with maximum vertical locomotion whereas others were correlated with maximum level locomotion. We conclude that stance phase, not swing phase, limits maximum performance in this species of lizard. In addition, we highlight the need to include properties of stance-phase muscles and a spectrum of ecologically relevant behaviors when attempting to correlate locomotor physiology with ecology and/or performance.

Thermoregulation in Nocturnal Ecthotherms: Seasonal and Intraspecific Variation in the Mediterranean Gecko (Hemidactylus turcicus)

Abstract Comprehensive investigations of thermoregulation have been primarily performed on diurnal lizards. The nocturnal gecko, Hemidactylus turcicus, was used to test a protocol proposed by P. E. Hertz, R. B. Huey, and R. D. Stevenson in 1993. Measures of body temperatures of field active geckos (Tb) and operative temperatures (Te), the equilibrium body temperatures that animals would attain in given microclimates, were compared to measures of preferred temperatures (Tp) determined in a thermal gradient. Measurements were made on adult males, adult nongravid females, adult gravid females, and juvenile geckos in four seasons (June, August, October, March). Both Tb and Te varied between seasons; however, Tb closely tracked Te. No seasonal patterns existed in Tp; however, juveniles had the lowest Tp, whereas gravid females had the highest. Regardless, in all seasons geckos were ineffective thermoregulators. The low variability in Te is the likely cause for this pattern. For such “thermoconforming” species, we suggest that the magnitude of the variation in Tb, Te, and Tp be included in assessments of how well organisms regulate to their set-point (preferred), temperatures. We conclude that geckos, and possibly many nocturnal ectotherms, thermoregulate during the day when a more variable thermal environment exists.

Getting Up to Speed: Acceleration Strategies in the Florida Scrub Lizard, Sceloporus woodi

Small animals typically rely on quick bursts and intermittent pauses when moving in the wild. Hence, the study of acceleration capacity is important for understanding the ecology and evolution of locomotor performance. In this study, we investigate intraspecific variation in the acceleration capacity of a small lizard (Sceloporus woodi). To quantify animal acceleration performance, the momentum‐impulse theorem is applied to data collected from high‐speed video recordings of individuals accelerating from a standstill and over a subsequent distance of 0.4 m. Unlike earlier studies, the momentum‐impulse approach allows one to directly and precisely quantify the per step contribution to acceleration capacity. Like other small vertebrates, we show that S. woodi is capable of accelerating to near maximum speeds (∼2 m s−1) within ∼0.4 m and needs only a few steps (at least five) to achieve maximum speed. However, considerable intraspecific variation in acceleration capacity exists; individuals take different numbers of steps (two to five steps) over the first 0.4 m, and only some individuals (10 of 19) reach their maximum speed over the first 0.4 m. Only acceleration performance in steps 1 and 2 is predictive of running speed at 0.4 m; accelerations in steps 3, 4, and 5 are not related to individual differences in speed. Individual variation in acceleration strategy is considerable, with individuals using one of three strategies to reach maximum speed. Muscle mass‐specific power during acceleration approaches the maximum power output measured for lizard hindlimb musculature (∼900 W kg−1), suggesting that S. woodi accelerations approach the limit of their musculoskeletal system. This study highlights the utility of the momentum‐impulse approach to study acceleration performance and the importance of elucidating the per step contribution to acceleration capacity.

Effects of Training and Testosterone on Muscle Fiber Types and Locomotor Performance in Male Six-Lined Racerunners (Aspidoscelis sexlineata)

Testosterone (T) is thought to affect a variety of traits important for fitness, including coloration, the size of sexual ornaments, aggression, and locomotor performance. Here, we investigated the effects of experimentally elevated T and locomotor training on muscle physiology and running performance in a nonterritorial male lizard species (Aspidoscelis sexlineata). Additionally, several morphological attributes were quantified to examine other characters that are likely affected by T and/or a training regimen. Neither training alone nor training with T supplementation resulted in increased locomotor performance. Instead, we found that T and training resulted in a decrease in each of three locomotor performance variables as well as in hematocrit, ventral coloration, and testis size. Strikingly, neither the size nor the fiber composition of the iliofibularis or gastrocnemius muscles was different among the two treatments or a group of untrained control animals. Hence, the relationships among T, training, and associated characters are not clear. Our results offer important insights for those hoping to conduct laboratory manipulations on nonmodel organisms and highlight the challenges of studying both training effects and the effects of steroid hormones on locomotor performance.

Sexual Size Dimorphisms and Bite Force in the Northern Alligator Lizard, Elgaria coerulea

Abstract Bite force capacity is a directly related performance correlate of head shape and is an integrative measure of performance in capturing and handling prey, fighting, and mating, especially for males. We investigated head shape and bite force dimorphisms in a small semifossorial lizard (Elgaria coerulea) that exhibits females-larger sexual size dimorphism (FL-SSD) in some populations but not in others. Specifically, we explored how body size, head shape, and bite force compare between the sexes relative to any dimorphisms in body or head size in a coastal population of E. coerulea. Female larger-SSD was confirmed for this population, but it contrasted with males-larger sexual dimorphism in head size (ML-SDHS). Males also had greater bite force than females of similar size. The secretive habit of E. coerulea hampers observations of behavioral interactions among conspecifics. However, it is expected that lizards with FL-SSD will have ML-SDHS if males with greater bite force win fights with other males over access to mates and/or if greater bite force increases copulation success.

ALLOMETRY, SEXUAL SIZE DIMORPHISM, AND NICHE PARTITIONING IN THE MEDITERRANEAN GECKO (HEMIDACTYLUS TURCICUS)

Abstract Hemidactylus turcicus is a small gekkonid lizard native to the Middle East and Asia that is known to exhibit sexual dimorphism in head size. Several potential explanations exist for the evolution and maintenance of sexual dimorphism in lizards. We tested 2 of these competing hypotheses concerning diet partitioning and differential growth. We found no differences in average meal size (volume) or in any single dimension of prey size for similarly sized males and females. Allometric patterns of increases in head size also were measured in males and females. We found that males exhibited a mixture of isometric and positively allometric patterns of head size increase, whereas females exhibited isometric and negatively allometric patterns. Thus, we concluded that sexual dimorphism in head size is not the result of diet partitioning but instead of differential growth patterns following sexual maturity in males and females.

Rock-dwelling Lizards Exhibit Less Sensitivity of Sprint Speed to Increases in Substrate Rugosity

Effectively moving across variable substrates is important to all terrestrial animals. The effects of substrates on lizard performance have ecological ramifications including the partitioning of habitat according to sprinting ability on different surfaces. This phenomenon is known as sprint sensitivity, or the decrease in sprint speed due to change in substrate. However, sprint sensitivity has been characterized only in arboreal Anolis lizards. Our study measured sensitivity to substrate rugosity among six lizard species that occupy rocky, sandy, and/or arboreal habitats. Lizards that use rocky habitats are less sensitive to changes in substrate rugosity, followed by arboreal lizards, and then by lizards that use sandy habitats. We infer from comparative phylogenetic analysis that forelimb, chest, and tail dimensions are important external morphological features related to sensitivity to changes in substrate rugosity.

Performance and three-dimensional kinematics of bipedal lizards during obstacle negotiation

SUMMARY Bipedal running is common among lizard species, but although the kinematics and performance of this gait have been well characterized, the advantages in biologically relevant situations are still unclear. Obstacle negotiation is a task that is ecologically relevant to many animals while moving at high speeds, such as during bipedal running, yet little is known about how obstacles impact locomotion and performance. We examined the effects of obstacle negotiation on the kinematics and performance of lizards during bipedal locomotion. We quantified three-dimensional kinematics from high-speed video (500 Hz) of six-lined racerunners (Aspidoscelis sexlineata) running on a 3 m racetrack both with and without an obstacle spanning the width of the track. The lizards did not alter their kinematics prior to contacting the obstacle. Although contact with the obstacle caused changes to the hindlimb kinematics, mean forward speed did not differ between treatments. The deviation of the vertical position of the body center of mass did not differ between treatments, suggesting that in the absence of a cost to overall performance, lizards forgo maintaining normal kinematics while negotiating obstacles in favor of a steady body center of mass height to avoid destabilizing locomotion.

Testing amniote models of prey transport kinematics: a quantitative analysis of mouth opening patterns in lizards

Two models have been proposed to describe the prey transport kinematics of terrestrial vertebrates (Bramble and Wake, 1985; Reilly and Lauder, 1990). The critical difference between the models is the presence or absence of a slow open-II phase (SO-II) in the gape profile during mouth opening. Each of these models has been applied to lizards, however to date, lizard feeding kinematics have not been adequately quantified to assess the utility of these models for this clade. Neither model has been sufficiently tested due to the lack of a methodology to assess the specific differences between the models. We describe a method that uses explicit mathematical criteria to define the kinematic phases in tetrapod feeding. This slope analysis& is used to precisely quantify and compare the transport kinematics of seven lizard species. Lizard transport kinematics were highly variable both within and across taxa. However, several common gape cycle patterns were identified. The predominant patterns were slow-fast opening (37.3%), fast opening only (22.9%) and slow opening only (21.2%). The most common pattern explicitly fits the prediction of the Reilly and Lauder model while the other two are similar to patterns observed in salamanders. Thus, lizards possess both the slow opening-fast opening pattern predicted for amniotes and the more primitive, simple opening pattern characteristic of more basal tetrapods. Plateau phases were found in only 12.8% of the profiles and only a fourth of these (3.4% of the total) explicitly fit the Bramble and Wake model (slow opening, plateau, fast opening) and two species never exhibited plateaus in their gape cycles. Thus, it is clear that the Bramble and Wake model is not supported as a generalized model for lizards or generalized tetrapods.