Studies into the function of structures are crucial for making connections between morphology and behaviour of organisms, but are still rare for the terrestrial Testudinidae. We investigated the kinematics of shoulder girdle and forelimb motion in Hermann's tortoise Testudo hermanni using biplanar X-ray fluoroscopy with a twofold aim: firstly, to understand how the derived shapes of shoulder girdle and carapace together influence rotation of the girdle; and, secondly, to understand how girdle rotation affects forelimb excursion. The total degree of shoulder rotation in the horizontal plane is similar to a species with a less domed shell, but because of the long and nearly vertically oriented scapular prong, shoulder girdle rotation contributes more than 30% to the horizontal arc of the humerus and nearly 40% to the rotational component of step length. The antebrachium and manus, which act as a functional unit, contribute roughly 50% to this component of the step length because of their large excursion almost parallel to the mid-sagittal plane. This large excursion is the result of the complex interplay between humerus long-axis rotation, counter-rotation of the antebrachium, and elbow flexion and extension. A significant proportion of forelimb step length results from body translation that is due to the propulsive effect of the other limbs during their stance phases. Traits that are similar to other tortoises and terrestrial or semi-aquatic turtles are the overall slow walk because of a low stride frequency, and the lateral-sequence, diagonally coupled footfall pattern with high duty factors. Intraspecific variation of carapace shape and shoulder girdle dimensions has a corresponding effect on forelimb kinematics.
The authors declare no competing or financial interests.
M.S. and M.M. analyzed the data and interpreted the findings. M.S. wrote and revised the manuscript. M.M. executed the experiments and collected the data. M.S.F. designed the experiments, interpreted the findings, and revised the manuscript.
This research project was funded solely by the budget of the Institute of Systematic Zoology and Evolutionary Biology, Friedrich-Schiller-University Jena.
All raw x-ray movies are available at the Jena Collection of X-ray Movies (http://szeb.thulb.uni-jena.de/szeb/templates/master/template_szeb/index.xml).
Supplementary information available online at http://jeb.biologists.org/lookup/doi/10.1242/jeb.137059.supplemental
- Received January 1, 2016.
- Accepted June 20, 2016.
- © 2016. Published by The Company of Biologists Ltd