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To see how, consider if Philip or Joanna carry a deleterious mutation which is recessive in its expression. Consider if you have a dozen first cousins to choose from vs. This is extremely suggestive.
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Correlation is not causation, but correlation is necessary for an inference of causation. Over time this sort of process would result in radically increased inbreeding coefficients. In the case of the Hapsburgs there is no need for de novo mutations, all humans carry within them numerous deleterious alleles which are masked because they only express recessively.
But it is nice to see it laid out in a precise manner. Consider that lower infant morality in modern societies, and the large families the norm in many developed societies. In addition to inbreeding due to unions between close relatives, ancestral inbreeding from multiple remote ancestors makes a substantial contribution to the inbreeding coefficient of most kings.
In some ways inbreeding can be thought of as a way to crank down long term effective population size and increase the power of stochastic processes within a gene pool. His tongue was so large that he was barely able to speak. Finally you have to include in the inbreeding coefficient of that common ancestor. That might make a big difference in terms of the probability of exogamy or not.
As you can see above iterated generations of inbreeding tend to result in some serious problems. His brief life consisted chiefly of a passage from prolonged infancy to premature senility.
This increases the pool of possible mates in many extended families. Since the paper is Open Access, I recommend you just go straight to it. Assuming that the grandparents, of whom there are only two, are unrelated. In plain English, the higher the inbreeding coefficient the higher mortality.
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Though in many cultures cousin-marriage is preferred for various reasons, until recently it was extant at lower frequencies than is the case today. The Spanish Hapsburgs had a higher infant mortality than Spanish commoners.
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The Samaritans of Israel are even engaging in a program of marrying out to reduce the high frequency of birth defects. Not only did the Spanish Hapsburgs have a high infant mortality rate, but it got higher as their inbreeding become more advanced.
The kings of the Spanish Habsburg dynasty frequently married close relatives in such a way that uncle-niece, one piece dresses in bangalore dating first cousins and other consanguineous unions were prevalent in that dynasty. His intellect was similarly disabled.
That being said, smaller families sizes in many developed nations in the near future might mean that there is a natural social break on the practice of cousin marriage. Even then, he was unable to walk properly, because his legs would not support him and he fell several times. The chart to the left plots out the process of repeated brother-sister matings within an isolated population over time. If drift becomes powerful enough it can lead the population toward extinction through mutational meltdown. But this a story of genetics as well as history, because historians have long assumed impressionistically that there was something rotten in the gene pool of the Spanish Hapsburgs.
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