In this study, we used a previously developed osteometry‐based method to calculate available range of motion in presacral intervertebral joints in artiodactyls. We have quantified all three directions of intervertebral mobility: sagittal bending (SB), lateral bending (LB), and axial rotation (AR). This research covers 10 extant families of artiodactyls from 33 genera and 39 species. The cervical region in artiodactyls is the most mobile region of the presacral vertebral column in SB and LB. Mobility is unevenly distributed throughout the joints of the neck. The posterior neck joints (C4–C7) are significantly more mobile (on average by 2.5–3.5°) to anterior joints (C2–C4) and to the neck–thorax joint (C7–T1) in SB and LB. An increase in the relative length of the cervical region in artiodactyls is accompanied by an increase in the bending amplitudes (SB: Pearson r = 0.781; LB: r = 0.884). Animals with the most mobile necks (representative of Giraffidae and Camelidae) are 2–3 times more mobile in SB and LB compared to species with the least mobile necks. The thoracic region in artiodactyls, as in other mammals, is characterized by the greatest amplitudes of AR due to the tangential orientation of the zygapophyseal articular facets. The lowest AR values in the thoracic region are typical for the heaviest artiodactyls—Hippopotamidae. The highest AR values are typical for such agile runners as cervids, musk deer, pronghorn, as well as large and small antelopes. SB mobility in the posterior part of the thoracic region can be used by artiodactyls during galloping. The highest values of SB aROM in the posterior part of the thoracic region are typical for small animals with high SB mobility in the lumbar region. The lumbar region in mammals is adapted for efficient SB. Both the cumulative and average SB values in the lumbar region showed correspondence to the running type employed by an artiodactyl. The greatest SB amplitudes in the lumbar region are typical for small animals, which use saltatorial and saltatorial–cursorial running. An increase in body size also corresponds to a decrease in lumbar SB amplitudes. The lowest SB amplitudes are typical for species using the so‐called mediportal running. Adaptation to endurance galloping in open landscapes is accompanied by a decrease in lumbar SB amplitudes in artiodactyls. The consistency of the approach used and the wide coverage of the studied species make it possible to significantly expand and generalize the knowledge of the biomechanics of the vertebral column in artiodactyls.

In this paper, we develop and validate an osteometry-based mechanistic approach to calculation of available range of motion (aROM) in presacral intervertebral joints in sagittal bending (SB), lateral bending (LB), and axial rotation (AR). Our basic assumption was the existence of a mechanistic interrelation between the geometry of zygapophysial articular facets and aROM. Trigonometric formulae are developed for aROM calculation, of which the general principle is that the angle of rotation is given by the ratio of the arc length of motion to the radius of this arc. We tested a number

This study is the first analysis of mobility in the lumbosacral joint of even-toed ungulates covering the full range of body masses and running forms. In this study, we modified a previously developed osteometry-based method to calculate the available range of motion (aROM) in the lumbosacral joint in artiodactyls. We quantified all three directions of intervertebral mobility: sagittal bending (SB), lateral bending (LB), and axial rotation (AR). This research covers extant artiodactyls from 10 families, 57 genera, and 78 species. The lumbosacral joint in artiodactyls is on average almost twice as mobile in SB as the average intralumbar joint (aROM 15.68° vs 8.22°). In all artiodactyls, the first sacral prezygapophyses are equipped with postfacet fossae determining the available range of lumbosacral hyperextension. SB aROM in the lumbosacral joint in artiodactyls varies almost sevenfold (from 4.53° to 31.19°) and is closely related to the body mass and running form. An allometric equation was developed for the first time, for the joint angular amplitude of motion, exemplified by the artiodactyl lumbosacral SB aROMs, as a power function of body mass, the power coefficient value being close to −0.15. High SB aROM at the lumbosacral joint is characteristic of artiodactyls with at least one of the following characteristics: high cumulative and average SB aROM in the lumbar region (Pearson r = 0.467-0.617), small body mass (r = −0.531), saltatorial or saltatorial-cursorial running form (mean = 16.91-18.63°). The highest SB aROM in the lumbosacral joint is typical for small antelopes and Moschidae (mean = 20.24-20.27°). Among these artiodactyls SB aROMs in the lumbosacral joint are on par with various carnivores. Large and robust artiodactyls, adapted predominantly to mediportal and stilt (running on extremely tall limbs) running forms, have 2-3 times smaller SB aROMs in the lumbosacral joint. Adaptation to endurance galloping in open landscapes (cursorial running form) is accompanied by smaller lumbar and lumbosacral SB aROMs compared to that in saltatorial-cursorial artiodactyls of the same body mass. The wide range of species studied makes it possible to significantly expand the knowledge of relations of the mobility of the lumbosacral joint in artiodactyls to body mass and running form.

The vertebral column is a hallmark of vertebrates; it is the structural basis of their body and the locomotor apparatus in particular. Locomotion of any vertebrate animal in its typical habitat is directly associated with functional adaptations of its vertebrae. This study is the first large‐scale analysis of mobility throughout the presacral region of the vertebral column covering a majority of extant odd‐toed ungulates from 6 genera and 15 species. In this study, we used a previously developed osteometry‐based method to calculate available range of motion. We quantified all three directions of intervertebral mobility: sagittal bending (SB), lateral bending (LB), and axial rotation (AR). The cervical region in perissodactyls was found to be the most mobile region of the presacral vertebral column in LB and SB. Rhinoceroses and tapirs are characterized by the least mobile necks in SB among odd‐toed and even‐toed ungulates. Equidae are characterized by very mobile necks, especially in LB. The first intrathoracic joint (T1–T2) in Equidae and Tapiridae is characterized by significantly increased mobility in the sagittal plane compared to the typical thoracic joints and is only slightly less mobile than typical cervical joints. The thoracolumbar part of the vertebral column in odd‐toed ungulates is very stiff. Perissodactyls are characterized by frequent fusions of vertebrae with each other with complete loss of mobility. The posterior half of the thoracic region in perissodactyls is characterized by especially stiff intervertebral joints in the SB direction. This is probably associated with hindgut fermentation in perissodactyls: the sagittal stiffness of the posterior thoracic region of the vertebral column is able to passively support the hindgut heavily loaded with roughage. Horses are known as a prime example of a dorsostable galloper among mammals. However, based on SB in the lumbosacral part of the backbone, equids appear to be the least dorsostable among extant perissodactyls; the cumulative SB in equids and tapirs is as low as in the largest representatives of artiodactyls, while in Rhinocerotidae it is even lower representing the minimum across all odd‐toed and even‐toed ungulates. Morphological features of small Paleogene ancestors of rhinoceroses and equids indicate that dorsostability is a derived feature of perissodactyls and evolved convergently in the three extant families.

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