A phylogenetic epoch is a time interval between successive branching events during which the number of ancestral branches is constant. Read backward from the four sampled leaves: there are four branches until the first merger, then three until the next, then two until their common root. Write the random durations as , with expected values .
The original solution diagram below places time horizontally and labels each branch by its number of descendant leaves. Type A is the pectinate topology; type B is the balanced topology. The coloured time strips identify the three phylogenetic epochs; their drawn widths are schematic, not estimated times.
Figure 1.
Epochs and descendant counts for four sampled lineages
. Time runs from the common root to the sampled leaves. The two panels distinguish the three-plus-one and two-plus-two root splits; numbers on branches count sampled descendants. Details.
Count the number of descendant leaves on each branch during each phylogenetic epoch. From the tips backward, both topologies have branch-size lists during , then during . During , type A has and type B has .
Let be the total length of branches carrying descendant leaves. Adding the relevant phylogenetic epoch durations gives
For example, type A has one external branch and one three-descendant branch throughout , while type B has two two-descendant branches. In either case the total expected branch length is , a useful accounting check. The stem above the common root is excluded because mutations there would be shared by all four samples.
For a haploid population with possible parents, each pair of ancestral lineages merges backward at rate in the Kingman's coalescent approximation. With lineages there are pairs. Hence the mean phylogenetic epoch duration is
Thus , and . For an autosomal sample from diploid individuals, replace the number of gene copies by , giving . A mitochondrial sample naturally uses a haploid maternal effective population size; the common factor cancels in the ratios below.
In the Yule process, each of branches gives birth independently at rate per generation. The next birth therefore has total rate . Its waiting time has an exponential distribution, so the mean phylogenetic epoch duration is
For the ensuing calculation use , and . This is the inter-event waiting-time convention requested in the question, including an observation interval with four branches. Stopping the forward process immediately when the fourth branch is born would instead leave a zero terminal interval and describe a different sampling convention.
Phylogenetic epoch 2026-10-20
A phylogenetic epoch is an interval between successive branching or merging events during which the number of extant ancestral branches is constant. Its expected duration with branches is denoted .