Turtling the salamander: tail movements mitigate need for kinematic limb changes during walking in tiger salamanders (Ambystoma tigrinum) with restricted lateral movement

Lateral undulation and trunk flexibility offer performance benefits to maneuverability, stability, and stride length (via speed and distance traveled). These benefits make them key characteristics of the locomotion of tetrapods with sprawling posture, with the exception of turtles. Despite their bony carapace preventing lateral undulations, turtles are able to improve their locomotor performance by increasing stride length via greater limb protraction. The goal of this study was to quantify the effect of reduced lateral flexibility in a generalized sprawling tetrapod, the tiger salamander (Ambystoma tigrinum). We had two potential predictions: (1) either salamanders completely compensate by changing their limb kinematics, or (2) their performance (i.e., speed) will suffer due to the reduced lateral flexibility. This reduction was performed by artificially limiting trunk flexibility by attaching a 2-piece shell around the body between the pectoral and pelvic girdles. Adult tiger salamanders (n = 3, SVL = 9 cm-14.5 cm) walked on a 1 m trackway under three different conditions: unrestricted, flexible shell (Tygon tubing), and rigid shell (PVC tubing). Trials were filmed in a single, dorsal view, and kinematics of entire midline and specific body regions (head, trunk, tail), as well as the fore and hindlimbs, were calculated. Tygon individuals had significantly higher curvature than both PVC and unrestricted individuals for the body, but this trend was primarily driven by changes in tail movements. PVC individuals had significantly lower curvature in the trunk region compared to unrestricted individuals or Tygon; however, there was no difference between unrestricted and Tygon individuals suggesting the shells performed as expected. PVC and Tygon individuals had significantly higher curvature in the tails compared to unrestricted individuals. There were no significant differences for any limb kinematic variables among treatments including average, minimum, maximum angles. Thus, salamanders respond to decreased lateral movement in their trunk by increasing movements in their tail, without changes in limb kinematics. These results suggest that tail undulations may be a more critical component to sprawling-postured tetrapod locomotion than previously recognized.

How moles walk; it's all thumbs

A recurring theme in the evolution of tetrapods is the shift from sprawling posture with laterally orientated limbs to erect posture with the limbs extending below the body. However, in order to invade particular locomotor niches, some tetrapods secondarily evolved a sprawled posture. This includes moles, some of the most specialized digging tetrapods. Although their forelimb anatomy and posture facilitates burrowing, moles also walk long distances to forage for and transport food. Here, we use X-ray Reconstruction Of Moving Morphology (XROMM) to determine if the mole humerus rotates around its long axis during walking, as it does when moles burrow and echidnas walk, or alternatively protracts and retracts at the shoulder in the horizontal plane as seen in sprawling reptiles. Our results reject both hypotheses and demonstrate that forelimb kinematics during mole walking are unusual among those described for tetrapods. The humerus is retracted and protracted in the parasagittal plane above, rather than below the shoulder joint and the ‘false thumb’, a sesamoid bone (os falciforme), supports body weight during the stance phase, which is relatively short. Our findings broaden our understanding of the diversity of tetrapod limb posture and locomotor evolution, demonstrate the importance of X-ray-based techniques for revealing hidden kinematics and highlight the importance of examining locomotor function at the level of individual joint mobility.

Forelimb posture in neoceratopsian dinosaurs: implications for gait and locomotion

Ceratopsid dinosaurs traditionally have been restored with sprawling forelimbs and were considered unable to run at high speeds. An alternative view restores the ceratopsids as rhinoceros-like with parasagittal forelimb kinematics and the ability to run faster than extant elephants. Several anatomical difficulties concerning the mounting of ceratopsid skeletons with nearly parasagittal forelimbs stem not from the forelimb itself, but from errors in rib and vertebral articulation. Matching a skeletal restoration to a probable ceratopsid trackway shows that the hands were placed directly beneath the glenoids, and that manual impressions were directed laterally, not medially as in sprawling reptiles. Pedal impressions in trackways are medial to the manual impressions, owing to the slightly averted elbow and to the asymmetrical distal femoral condyles, which directed the crus slightly medially. The limbs of ceratopsians of all sizes display substantial joint flexure, strongly indicating that the elephantine forelimb posture that has sometimes been suggested as the alternative to a sprawling posture is erroneous. The articular surfaces of uncrushed ceratopsian scapulocoracoids and forelimb joints confirm that the forelimb operated in a near-parasagittal plane with the elbows only slightly averted. The maximal running speed of even the largest ceratopsids is inferred to have significantly exceeded that of elephants and was probably broadly similar to that of rhinos.

Locomotion in alligator mississippiensis: kinematic effects of speed and posture and their relevance to the sprawling-to-erect paradigm

In terms of locomotory posture, amphibians and lizards are considered to be sprawlers, mammals and dinosaurs are considered to be erect, and extant crocodilians are considered to be intermediate because they use the 'high walk', a semi-erect posture where the body is held half-way between the sprawling and erect grades during locomotion. In addition, crocodilians occasionally use a sprawling posture. Extant crocodilians, therefore, provide an interesting model in which to investigate the sprawling-to-erect transition in vertebrate evolution. This study quantifies the sprawl and high walk kinematics of the alligator Alligator mississippiensis moving at different speeds on a treadmill and compares them with kinematic data available for other vertebrates. These data allow us to examine the effects of speed on crocodilian postures and to examine how crocodilian locomotion relates to the sprawling-to-erect paradigm in vertebrate locomotion. Our results show that the crocodilian sprawl is not functionally equivalent to the primitive sprawling behaviors exhibited by salamanders and lizards. In fact, although the high walks and sprawls of alligators exhibit some kinematic differences, they are actually much more similar than expected and, essentially, the crocodilian sprawl is a lower version of a high walk and could be termed a 'low walk'. In terms of the sprawling-to-erect transition, the high walk has knee kinematics intermediate between those of birds and non-archosaurian tetrapods, but alligators increase speed in a way completely different from other terrestrial vertebrates (distal rather than proximal limb elements are used to increase speed). These kinematic data viewed in the light of the fossil and phylogenetic evidence that modern crocodilians evolved from erect ancestors suggest that modern crocodilians have secondarily evolved a variable semi-erect posture and that they are problematic as an intermediate model for the evolutionary transition from sprawling to erect postures in archosaurs.

Sprawling locomotion in the lizard Sceloporus clarkii: quantitative kinematics of a walking trot

Although the hindlimb is widely considered to provide the propulsive force in lizard locomotion, no study to date has investigated the kinematic patterns of the lizard hindlimb during running for more than one stride for a single individual. The quantitative kinematics of the hindlimb, pelvis and backbone are described here for two individuals of the lizard Sceloporus clarkii using a fast walking trot on a treadmill moving at a constant speed of 0.833 m s-1. Pelvic rotation, femoral retraction, knee flexion and posterior movement of the foot all begin before the foot hits the substratum and, thus, there is a terminal portion of the swing phase during which the limb is retracting. Pelvic rotation (to the opposite side), femoral protraction and knee flexion all begin before the foot leaves the substratum. The foot, however, continues to move posteriorly into the early swing phase. Thus, limb retraction and protraction movements do not directly correlate with footfall phases. Axial bending involves a rough standing wave with two nodes, one centered on each limb girdle. In Sceloporus clarkii, the foot clearly remains lateral to the knee and, thus, has a more sprawling posture than that of any other vertebrate studied to date. Therefore, the generalization that the 'lacertilian' foot passes under the knee joint is no longer supported. The kinematics of sprawling locomotion in Sceloporus clarkii are compared and contrasted with the general understanding of lizard locomotion based on qualitative work to date. Comparisons with other tetrapods reveal a fundamental functional dichotomy in hindlimb retraction mechanics in salamanders and mammals versus lizards that may be related to a key morphological difference in the saurian caudifemoralis muscle.