Teeth (Cambridge Manuals in Archaeology)
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Recent works have shown the important contribution of this long-living extinct bovid to our understanding of life history evolution  — . Myotragus balearicus presents an extremely modified lower dentition not shared by any other known ruminant that is interpreted as a functional adaptation to increase feeding efficiency under resource limitation . Adults display a single rodent-like evergrowing incisor, a single premolar p4 and three molars per hemimandible.
The second and third incisor, the canine and the third premolar are lost during anagenetic evolution of the Myotragus lineage. Their eruptive sequence is also unusual among living bovids, as the permanent incisors and the premolars erupt relatively early  , . In all living bovids the permanent first incisor appears after the eruption of the second molar and the fourth premolar is the last tooth to emerge after or subsequently to the third molar . However, the permanent incisor in this extinct bovid emerges just after the first molar eruption and the premolar emerges well before the third molar.
However, Bover and Alcover  and subsequently other authors  proposed that, like in rodents and lagomorphs, the evergrowing incisor of adult specimens of M. They conclude that the evergrowing incisors present in the adult specimens of M. To explain the relatively early eruption of the permanent first incisors, they suggested an acceleration of the incisor development to cope with resource limitation on islands, in agreement with the abovementioned ecological hypotheses .
Here we test whether the pattern relative order and pace absolute age of dental eruption in M. The slow signature of dental eruption relative order in M. Our recent works using hard tissue histology and demography provided evidence that the Myotragus lineage evolved towards a slow life history slow growth, delayed maturity and long life span that enhances fitness under density-dependent resource limitation  — . However, an acceleration of the pace of development of the evergrowing incisors of M.
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The present study sheds light on the selection pressures that shape dental ontogeny in mammals. With the aim of analyzing teeth replacement pattern in the fossil goat-like M. The absolute timing of molar development in M. In the present study, these data were used to calibrate the dental eruption schedule absolute age in M. Moreover, we scanned a juvenile mandible of M. CT-scan images were analysed using the software OsiriX v.
We used the original dental homologies for M. The material of M. The mandible of an early Pleistocene species of M. We obtained permission from these institutions to access the collections. All these specimens were loaned to the authors of this work to carry out radiological study. The dental eruption schedule of M.
In the youngest specimens of the M. The smaller incisor di2 is likely to fall out very early before m1 eruption. The larger deciduous incisor di1 shows an open distal base located at the level of mental foramen, before dp3 alveolus. This is the only incisor present in the jaw when m1 begins alveolus emergence Fig. Note the development and the replacement pattern of the first incisor. The first molar is fully erupted by 9 months, age at which the adult evergrowing incisor i1 is already present but not erupted Fig.
The base of i1 is wide and open and is located after mental foramen, just beneath dp3 alveolus, oriented toward the lingual side of the jaw. Replacement of the deciduous first incisor by the permanent one. A Sagittal plane. B Coronal plane. C Transversal plane. A Buccal side of the lower jaw showing the germ of the permanent first evergrowing incisor and the erupted deciduous first incisor. B Lingual side of the deciduous incisor. Note the lack of enamel.
C Buccal side of the deciduous incisor. When m1 crown formation is complete by around 13 months, di1 is already lost.
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The sole incisors that remain in the jaw i1 are now erupted Fig. This tooth is greatly enlarged and extends distally beneath dp4 in a more lingual position. During ontogeny, this evergrowing tooth can extend even more distally beneath m1. The second molar is fully erupted by about 32 months, and p4 formation starts subsequently Fig. At this stage dp3 is almost depleted and subsequently it is shed. The fourth deciduous premolar falls out and p4 emerges when m2 crown formation is complete.
The third deciduous premolar dp3 is replaced by a very reduced conical shaped permanent premolar p3 that is lost during adulthood.
This vestigial tooth is the result of the evolutionary trend towards the reduction of the premolar series in the Myotragus lineage . There is no evidence of m3 formation at this stage Fig. Finally, the third molar erupts by the age of almost 6 years when the other teeth are already moderately worn Fig.
Note the development and replacement pattern of the fourth premolar.
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As regards the juvenile hemimandible of the early Pleistocene species of M. Therefore, in the early species of the Myotragus evolutionary lineage the first permanent incisor i1 emerged right after the second molar m2. The same applies in living bovids Fig.
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Note that in both species the first permanent incisor erupts just after the second molar. A Mandible of an early Pleistocene M. D CT-scan image of the lower jaw of the living goat. Contrary to the suggestion in Bover and Alcover  , the present study provides further evidence for the successional or secondary nature of the adult evergrowing incisors of M.
Based on the CT-scan study on dental development in M.
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The differentiation of a secondary tooth at the lingual margin of the deciduous enamel organ is a constant trait in eutherian dental development . The secondary incisor, hence, always emerges at a slightly more lingual position than the primary one. However, the strongest evidence for the secondary nature of this evergrowing tooth comes from the fossil record of the Myotragus anagenetic lineage.
The juvenile mandible of M. An important argument for the primary or deciduous nature of the incisors of adult specimens of M. However, incisor enlargement in mammals can occur at different dental positions and generations .
The enlarged lower incisors of rodents and lagomorphs are considered di2 based on the development of small, abnormal di1 on the prenatal jaws, which are probably subsequently resorbed . Nevertheless, this is very different from what happens in M. Moreover, this tooth is noticeably wider and longer relative to earlier species of the genus Myotragus .
Three permanent evergrowing incisors with morphology very similar to that of the adult incisor of M. Based on all these arguments, the adult incisor of M. Therefore, it can be established that replacement teeth incisors and premolars in the fossil bovid appear relatively earlier in sequence than in living bovids.
In addition, the advance in the eruption sequence of the i1 occurs along the anagenetic lineage of the genus Myotragus over a period of about 2. The first permanent incisor in the early Pleistocene M. However, in the late Pleistocene-Holocene species M. This is, to our knowledge, the first evidence from the fossil record of advance of incisor replacement along an evolutionary mammalian lineage.
According to the study of Smith  on patterns of tooth eruption sequence in mammals, M. These data conform to the slow life history described for the extinct species.
The present study shows that m1 of M. Moreover, the potential life span for M. Both, the values of m1 emergence and life span are unusual among extant bovids. They are even rare among artiodactyls in general, except for some larger, slower-growing species such as hippopotamus Hippopotamus amphibius that show the same slow sequence signature of tooth eruption as does M. As regards the pace of dental eruption absolute age , the large evergrowing incisor i1 of M.
This incisor grew relatively fast as it took at most three months to attain its important length before eruption. Therefore, the adult incisor did not only grow fast in relative terms but also in absolute terms. However, the posterior teeth molars and premolar of M. Thus, the molars of M.
This slow pace of development of posterior teeth in M. This decoupling between the pace of anterior and posterior tooth development suggests that the state of dental development at weaning might also be target of selection in agreement with ecological hypotheses . The absolute early eruption of the large rodent-like evergrowing incisor of M. In an ecological context of high intraspecific competition for scarce resources, selection favours increased harvesting efficiency as shows the highly specialized dentition of adults .
Our results indicate that selection also operated at an early age leading to dental precocity at weaning. The early eruption of the adult evergrowing incisors provides the mechanical requirements for processing more abrasive, tougher foodstuff food processing hypothesis , in addition to facilitate weaning and independent juvenile foraging foraging independence hypothesis.
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Each book includes a survey of current archaeological practice alongside essential reference material on contemporary techniques and methodology. Reitz and Elizabeth S. Subject to statutory exception and to the provision of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press. Tooth and periodontium. Mustelidae, skunk permanent dentitions.