Comparative study of the body proportions in Elephantidae and other large herbivorous mammals

In this study, we aimed to achieve three objectives: (1) to precisely characterize the body plans of Elephantidae and other large herbivorous mammals; (2) based on this analysis, to determine whether the body plans of the extinct woolly mammoth (Mammuthus primigenius) and steppe mammoth (M. trogontherii) differ from those of modern-day Elephantidae: the Asian elephant (Elephas maximus), the African bush (Loxodonta africana), and forest (L. cyclotis) elephants; (3) to analyze how the body plans have changed in extant perissodactyls and proboscideans compared with their Paleogene ancestors. To accomplish this, we studied mammoth skeletons from the collections of Russian museums and compared this data with a large number of skeletons of extant elephantids, odd-toed, and even-toed ungulates, as well as their extinct relatives. We showed that three genera of Elephantidae are characterized by a homogeneous body plan, which is markedly different from other large herbivores. Elephantids break the interrelationship, that exists in artiodactyls and perissodactyls, between the total length of the head and neck on one side and the limb's segments on the other. Their limbs are very tall (inferior in this regard among large ungulates only to the giraffe), and, contrary to the other large herbivorous mammals, elongated due to the length of the proximal segments. This allows them to effectively utilize the principle of inverted pendulum (straight-legged walking) in locomotion. The biggest differences in the body plan of mammoths compared with extant elephants are a markedly larger pelvis, elongated fore- and hindlimbs (due to the increased relative length of their proximal segments), and different proportions of the skull. The body plans of plesiomorphic Paleogene proboscideans and perissodactyls differed markedly from their descendants in every body part; these differences are related, on the one hand, to the allometric growth, and on the other hand, to the advancement of the locomotor apparatus in the course of their evolution. The most notable difference in the body plan between Paleogene proboscidean Moeritherium and extant Elephantidae is the ~2-fold increase in relative limb height.