Three-dimensional hindlimb kinematics of water running in the plumed basilisk lizard (<i>Basiliscus plumifrons</i>)

Hsieh, S. Tonia

doi:10.1242/jeb.00679

Cited by 66 publications

(75 citation statements)

References 47 publications

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“…This difference could be related to the different methodologies used: treadmill (Irschick and Jayne, 1999) vs. track (Hsieh, 2003; present study).…”

Section: Discussionmentioning

confidence: 68%

Bipedal locomotion in Tropidurus torquatus (Wied, 1820) and Liolaemus lutzae Mertens, 1938

et al. 2008

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Bipedalism has evolved on numerous occasions in phylogenetically diverse lizard families. In this paper we describe, for the first time, bipedal locomotion on South American lizards, the sand-dweller Liolaemus lutzae and the generalist Tropidurus torquatus. The lizards were videotaped running on a racetrack and the sequences were analyzed frame by frame. The body posture, as a whole, diverged a lot during bipedal locomotion between the two species, even though there was no difference regarding their sprint performance. The locomotor behavior of L. lutzae is, in general, more similar to the one observed on other sand-dweller lizards. Certain particularities are common, such as the digitigrade posture at footfall and throughout stance, trunk angles; and tail posture. In contrast, T. torquatus exhibited high trunk angles and dragged its tail, in a posture compared to basilisks. This body posture could be related to certain characteristics and obstacles of a microhabitat such as the one around lakes and streams (basilisks) and the one with compact shrubby vegetation (T. torquatus).Keywords: bipedality, kinematics, locomotion, Tropidurus, Liolaemus. Bipedalismo em Tropidurus torquatus (Wied, 1820) e Liolaemus lutzae Mertens, 1938Resumo O bipedalismo apareceu em inúmeras ocasiões em famílias de lagartos filogeneticamente diversas. Neste trabalho, descrevemos, pela primeira vez, locomoção bipedal em lagartos sul-americanos, o lagarto de hábito terrestre Liolaemus lutzae e o generalista Tropidurus torquatus. Os lagartos foram filmados correndo em uma pista de corrida e as seqü-ências foram analisadas quadro a quadro. A postura corporal, como um todo, divergiu bastante durante a locomoção bipedal das duas espécies, mesmo quando não houve diferença significativa em relação ao desempenho locomotor. O comportamento locomotor de L. lutzae é, em geral, similar ao observado em outros lagartos especialistas em solos arenosos. Certas particularidades são comuns, como a postura digitígrada no momento de apoio e durante a fase de apoio; ângulos do corpo e postura da cauda. Em contraste, T. torquatus exibiu altos ângulos corporais e arrastava sua cauda, numa postura comparada à dos lagartos basiliscos. Este tipo de postura corporal pode estar relacionado com certas características e obstáculos de um microhabitat como ao redor de rios e riachos (basiliscos) e uma vegetação de arbustos compacta (T. torquatus).

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“…This difference could be related to the different methodologies used: treadmill (Irschick and Jayne, 1999) vs. track (Hsieh, 2003; present study).…”

Section: Discussionmentioning

confidence: 68%

Bipedal locomotion in Tropidurus torquatus (Wied, 1820) and Liolaemus lutzae Mertens, 1938

et al. 2008

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“…Basilisks within the examined size range tend to slap the water with their foot in a slight toe down position (7). By contacting the water at an acute angle, the total generated force has both a vertical and fore-aft component.…”

Section: Resultsmentioning

confidence: 96%

“…Beyond the moment the model foot contacts the water, however, the relevance of forces produced during passive model deceleration to basilisk lizard water running is limited. Foot movements when submerged are highly complex during actual water runs as basilisks actively sweep their foot through the water in threedimensions (7). As a result, forces produced by a model passively decelerating only vertically through water because of hydrodynamic drag would not reflect additional thrust and side forces generated during an actual step.…”

Section: Resultsmentioning

confidence: 99%

“…Lizards ran at sprint velocities within the range reported in another study (1.3-1.6 m͞s; see ref. 7). Each running sequence was filmed with two synchronized high-speed Redlake Imaging (Morgan Hill, CA) PCI500 cameras operating at 250 frames per s with a shutter speed of 1͞1,000 s. One camera filmed a dorsal view of the lizard through a mirror so we could determine the position of the lizards' foot relative to a laser light sheet illuminating the region of analysis in the water (see DPIV).…”

Section: Methodsmentioning

confidence: 99%

“…Basilisk lizard water-running kinematics are highly complex and are described in detail elsewhere (7). To facilitate analyses, each stride was divided into three phases based on foot kinematics: the slap, stroke, and recovery phases (7) (Fig.…”

Section: Methodsmentioning

confidence: 99%

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Running on water: Three-dimensional force generation by basilisk lizards

Hsieh

Lauder

2004

Proc. Natl. Acad. Sci. U.S.A.

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113

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Water provides a unique challenge for legged locomotion because it readily yields to any applied force. Previous studies have shown that static stability during locomotion is possible only when the center of mass remains within a theoretical region of stability. Running across a highly yielding surface could move the center of mass beyond the edges of the region of stability, potentially leading to tripping or falling. Yet basilisk lizards are proficient water runners, regularly dashing across bodies of water to evade predators. We present here direct measurements of time-averaged force produced by juvenile plumed basilisk lizards (Basiliscus plumifrons) while running across water. By using digital particle image velocimetry to visualize fluid flow induced by foot movement, we show that sufficient support force is generated for a lizard to run across water and that novel strategies are also required to run across a highly yielding surface. Juvenile basilisk lizards produce greatest support and propulsive forces during the first half of the step, when the foot moves primarily vertically downwards into the water; they also produce large transverse reaction forces that change from medial (79% body weight) to lateral (37% body weight) throughout the step. These forces may act to dynamically stabilize the lizards during water running. Our results give insight into the mechanics of how basilisk lizards run across water and, on a broader scale, provide a conceptual basis for how locomotor surface properties can challenge established rules for the mechanics of legged locomotion.Basiliscus plumifrons ͉ hydrodynamics ͉ particle image velocimetry ͉ locomotion R unning across water is a dramatic example of a locomotor function once thought to be limited to small-bodied invertebrates (e.g., see refs. 1-4). Although considerable literature is available that examines the biomechanics of movement across solid surfaces, very little is known about how softer, more yielding surfaces, such as water, affect legged locomotion. Water provides an unusual challenge, because it readily yields to any applied force. As a result, it would be expected that any animal attempting to walk across water would sink toward the supporting limb.There are few vertebrates capable of running across water. Waterfowl are known to slap the water with their feet during flapping take-off. However, they obtain a great deal of lift from their wings. Anolis lizards and baby green iguanas have been reported to run short distances across water. This capability, however, seems to be limited primarily to smaller-bodied individuals. Basilisk lizards (Basiliscus sp.) are unique in that they regularly run across water, using only their feet as a source of both lift and thrust. This behavior is prevalent among hatchlings through adults and is made more spectacular by their large size range: Hatchlings weigh Ϸ2 g, whereas adults can weigh Ͼ200 g. Basilisk lizards, therefore, serve as interesting model organisms for examining the mechanics of this remarkable locomotor fe...

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Biomechanical modeling and sensitivity analysis of bipedal running ability. I. Extant taxa

Hutchinson

2004

Journal of Morphology

128

212

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I used a simple mathematical model of the inverse dynamics of locomotion to estimate the minimum muscle masses required to maintain quasi-static equilibrium about the four main limb joints at mid-stance of fast running. Models of 10 extant taxa (a human, a kangaroo, two lizards, an alligator, and five birds) were analyzed in various bipedal poses to examine how anatomy, size, limb orientation, and other model parameters influence running ability. I examined how the muscle masses required for fast running compare to the muscle masses that are actually able to exert moments about the hip, knee, ankle, and toe joints, to see how support ability varies across the limb. I discuss the assumptions and limitations of the models, using sensitivity analysis to see how widely the results differed with feasible parameter input values. Even with a wide range of input values, the models validated the analysis procedure. Animals that are known to run bipedally were calculated as able to preserve quasi-static equilibrium about their hindlimb joints at mid-stance, whereas non-bipedal runners (iguanas and alligators) were recognized as having too little muscle mass to run quickly in bipedal poses. Thus, this modeling approach should be reliable for reconstructing running ability in extinct bipeds such as nonavian dinosaurs. The models also elucidated how key features are important for bipedal running capacity, such as limb orientation, muscle moment arms, muscle fascicle lengths, and body size. None of the animals modeled had extensor muscle masses acting about any one joint that were 7% or more of their body mass, which provides a reasonable limit for how much muscle mass is normally apportioned within a limb to act about a particular joint. The models consistently showed that a key biomechanical limit on running ability is the capacity of ankle extensors to generate sufficiently large joint moments. Additionally, the analysis reveals how large ratite birds remain excellent runners despite their larger size; they have apomorphically large extensor muscles with relatively high effective mechanical advantage. Finally, I reconstructed the evolution of running ability in the clade Reptilia, showing that the ancestors of extant birds likely were quite capable runners, even though they had already reduced key hip extensors such as M. caudofemoralis longus.

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Three-dimensional hindlimb kinematics of water running in the plumed basilisk lizard (Basiliscus plumifrons)

Cited by 66 publications

References 47 publications

Bipedal locomotion in Tropidurus torquatus (Wied, 1820) and Liolaemus lutzae Mertens, 1938

Bipedal locomotion in Tropidurus torquatus (Wied, 1820) and Liolaemus lutzae Mertens, 1938

Running on water: Three-dimensional force generation by basilisk lizards

Biomechanical modeling and sensitivity analysis of bipedal running ability. I. Extant taxa

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