Grey wolf genomic history reveals a dual ancestry of dogs.

The grey wolf (Canis lupus) was the first species to give rise to a domestic population, and they remained widespread throughout the last Ice Age when many other large mammal species went extinct. Little is known, however, about the history and possible extinction of past wolf populations or when and where the wolf progenitors of the present-day dog lineage (Canis familiaris) lived1-8. Here we analysed 72 ancient wolf genomes spanning the last 100,000 years from Europe, Siberia and North America. We found that wolf populations were highly connected throughout the Late Pleistocene, with levels of differentiation an order of magnitude lower than they are today. This population connectivity allowed us to detect natural selection across the time series, including rapid fixation of mutations in the gene IFT88 40,000-30,000 years ago. We show that dogs are overall more closely related to ancient wolves from eastern Eurasia than to those from western Eurasia, suggesting a domestication process in the east. However, we also found that dogs in the Near East and Africa derive up to half of their ancestry from a distinct population related to modern southwest Eurasian wolves, reflecting either an independent domestication process or admixture from local wolves. None of the analysed ancient wolf genomes is a direct match for either of these dog ancestries, meaning that the exact progenitor populations remain to be located.

Wall thickness of gas- and marrow-filled avian long bones: measurements on humeri, femora and tibiotarsi in crows (Corvus corone cornix) and magpies (Pica pica).

We studied how the ratio K of the internal to external diameter of gas- and marrow-filled avian long bones follows the biomechanical optima derived for tubular bones with minimum mass designed to fulfil various mechanical requirements. We evaluated radiographs of numerous humeri, femora and tibiotarsi in Corvus corone cornix and Pica pica. The K-values of the gas-filled humerus (K=0.78+/-0.03) and the marrow-filled femur (K=0.79+/-0.02) in Corvus are practically the same, while K of the marrow-filled tibiotarsus (K=0.71+/-0.04) is significantly smaller. The same is true for the gas-filled humerus (K=0.78+/-0.02) and the marrow-filled femur (K=0.77+/-0.02) and tibiotarsus (K=0.67+/-0.05) in Pica. K in Corvus is slightly larger than K in Pica, but the differences are statistically not significant. The standard deviation DeltaK of the tibiotarsi (DeltaK=0.04-0.05) is approximately two times as large as that of the humeri (DeltaK=0.02-0.03) and femora (DeltaK=0.02) in both species. Accepting the assumption of earlier authors that the ratio Q of the marrow to bone density is 0.5, our data show that the marrow-filled tibiotarsi of Corvus and Pica are optimized for stiffness, while the marrow-filled femora are far from any optimum. The relative wall thickness W=1-K of the gas-filled avian humeri studied is much larger than the theoretical optimum W*=1-K*=0.07, and thus these bones are thicker-walled than the optimal gas-filled tubular bone with minimum mass.

Testing the biomechanical optimality of the wall thickness of limb bones in the red fox (Vulpes vulpes).

The optimum for the ratio K of the internal to external diameter of a marrow-filled tubular bone with minimum mass designed to withstand a given type of strength (yield/fatigue, stiffness, fracture or impact) depends on Q = rhom/rhob only, where rhom and rhob are the densities of marrow and bone. With computer-assisted evaluation of radiographs of 62 femurs in the red fox (Vulpes vulpes) we measured the values of K. The mean and standard deviation of K are 0.68 and 0.036, and K changes in the rather wide range from 0.59 to 0.74. Accepting the assumption of earlier authors that Q = 0.50 or 0.44, our data would support the hypothesis that the fox femurs are optimized to withstand yield, fatigue or stiffness strengths. However, since the Q-values are unknown, the possibility cannot be excluded that any studied fox bone with an appropriately selected Q-value is optimized for any strength type. Assuming Q = 0.50 or 0.44, the relative mass increments mu of the investigated fox bones are smaller than 5% under all four mechanical conditions. The evolutionary relevance of such tiny mu-values is questionable.