The two descendant branches of an equal-rate Jukes-Cantor model compose to a single branch of duration . More explicitly, the model is symmetric, so summing over the ancestral base gives and a total mismatch probability . Thus use the pairwise difference fraction directly with , rather than halving the differences and correcting the halves separately:This is the parameter in the printed formula of (c). For a mutation rate measured as actual changes per site per year, the conversion derived in (b) givesThe latter is directly comparable to the naive estimate in (a): allowing repeated changes raises that estimate by about . The exponent parameter and actual event rate are two conventions for the same fitted process, not two different biological estimates.
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