Abstract

Mammals as a rule have seven cervical vertebrae, a number that remains remarkably constant. Changes of this number are associated with major congenital abnormalities (pleiotropic effects) that are, at least in humans, strongly selected against. Recently, it was found that Late Pleistocene mammoths (Mammuthus primigenius) from the North Sea have an unusually high incidence of abnormal cervical vertebral numbers, approximately ten times higher than that of extant elephants. Abnormal numbers were due to the presence of large cervical ribs on the seventh vertebra, indicating a homeotic change from a cervical rib-less vertebra into a thoracic rib-bearing vertebra. The high incidence of cervical ribs indicates a vulnerable condition and is thought to be due to inbreeding and adverse conditions that may have impacted early pregnancies in declining populations. In this study we investigated the incidence of cervical ribs in another extinct Late Pleistocene megaherbivore from the North Sea and the Netherlands, the woolly rhinoceros (Coelodonta antiquitatis). We show that the incidence of abnormal cervical vertebral numbers in the woolly rhinoceros is unusually high for mammals (15,6%, n = 32) and much higher than in extant Rhinoceratidae (0%, n = 56). This indicates that woolly rhinoceros lived under vulnerable conditions, just like woolly mammoths. The vulnerable condition may well have contributed to their eventual extinction.

Highlights

  • The number of cervical vertebrae in mammals is remarkably constant at seven, in contrast to other tetrapods, where this number varies considerably (Leboucq, 1898; Schultz, 1961; Starck, 1979; Narita & Kuratani, 2005)

  • We investigated developmental abnormalities in the neck vertebrae, the presence of cervical ribs

  • We compared the resulting incidence of cervical ribs with those that we found in skeletons of all five extant African and Asian species of rhinoceros (Dicerorhinus sumatrensis, Ceratotherium simum, Rhinoceros unicornis, R. sondaicus, Diceros bicornis)

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Summary

Introduction

The number of cervical vertebrae in mammals is remarkably constant at seven, in contrast to other tetrapods, where this number varies considerably (Leboucq, 1898; Schultz, 1961; Starck, 1979; Narita & Kuratani, 2005). The only exceptions are manatees (Trichechus, Sirenia) and extant sloths (Bradypus and Choloepus, Xenarthra). These latter taxa have an exceptional number of cervical vertebrae (Bateson, 1894; Starck, 1979; Varela-Lasheras et al, 2011). The extreme evolutionary conservation of the number of cervical vertebrae in mammals implies that there must be selection against intraspecific variation of this number. The most common variation is a partial or

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