Unexpected morphological diversity in ancient dogs compared to modern relatives.

Dogs are among the most variable species today, but little is known about the morphological variability in the early phases of their history. The Neolithic transition to farming may have resulted in an early morphological diversification as a result of changes in the anthropic environment or intentional selection on specific morphologies. Here, we describe the variability and modularity in mandible form by comparing 525 dog mandibles from European archaeological sites ranging from 8100 to 3000 cal. BC to a reference sample of modern dogs, wolves, and dingoes. We use three-dimensional geometric morphometrics to quantify the form of complete and fragmented mandibles. We demonstrate that an important morphological variability already existed before the Bronze Age in Europe, yet the largest, smallest, most brachycephalic or dolichocephalic extant dogs have no equivalent in the archaeological sample, resulting in a lower variation compared to modern relatives. The covariation between the anterior and posterior parts of the mandible is lower in archaeological dogs, suggesting a low degree of intentional human selection in early periods. The mandible of modern and ancient dogs differs in functionally important areas, possibly reflecting differences in diet, competition, or the implication of ancient dogs in hunting or defence.

Masticatory system integration in a commensal canid: interrelationships between bones, muscles and bite force in the red fox.

The jaw system in canids is essential for defence and prey acquisition. However, how it varies in wild species in comparison with domestic species remains poorly understood, yet is of interest in terms of understanding the impact of artificial selection. Here, we explored the variability and interrelationships between the upper and lower jaws, muscle architecture and bite force in the red fox (Vulpes vulpes). We performed dissections and used 3D geometric morphometric approaches to quantify jaw shape in 68 foxes. We used a static lever model and bite force estimates were compared with in vivo measurements of 10 silver foxes. Our results show strong relationships exist between cranial and mandible shape, and between cranial or mandible shape on the one hand and muscles or estimated bite force on the other hand, confirming the strong integration of the bony and muscular components of the jaw system. These strong relationships are strongly driven by size. The functional links between shape and estimated bite force are stronger for the mandible, which probably reflects its greater specialisation towards biting. We then compared our results with data previously obtained for dogs (Canis lupus familiaris) to investigate the effect of domestication. Foxes and dogs differ in skull shape and muscle physiological cross-sectional area (PCSA). They show a similar amount of morphological variation in muscle PCSA, but foxes show lower variation in cranial and mandible shape. Interestingly, the patterns of covariation are not stronger in foxes than in dogs, suggesting that domestication did not lead to a disruption of the functional links of the jaw system.

Bite force and its relationship to jaw shape in domestic dogs.

Previous studies based on two-dimensional methods have suggested that the great morphological variability of cranial shape in domestic dogs has impacted bite performance. Here, we used a three-dimensional biomechanical model based on dissection data to estimate the bite force of 47 dogs of various breeds at several bite points and gape angles. In vivo bite force for three Belgian shepherd dogs was used to validate our model. We then used three-dimensional geometric morphometrics to investigate the drivers of bite force variation and to describe the relationships between the overall shape of the jaws and bite force. The model output shows that bite force is rather variable in dogs and that dogs bite harder on the molar teeth and at lower gape angles. Half of the bite force is determined by the temporal muscle. Bite force also increased with size, and brachycephalic dogs showed higher bite forces for their size than mesocephalic dogs. We obtained significant covariation between the shape of the upper or lower jaw and absolute or residual bite force. Our results demonstrate that domestication has not resulted in a disruption of the functional links in the jaw system in dogs and that mandible shape is a good predictor of bite force.