others, these two measures of branch length are not always in direct proportion. The expected number of substitutions per site per year is often indicated with the Greek letter mu (μ). A model is said to have a strict molecular clock if the expected number of substitutions per year μ is constant regardless of which species' evolution is being examined. An important implication of a strict molecular clock is that the number of expected substitutions between an ancestral species and any of its present-day descendants must be independent of which descendant species is examined. Note that the assumption of a strict molecular clock is often unrealistic, especially across long periods of evolution. For example, even though rodents are genetically very similar to primates, they have undergone a much higher number of substitutions in the estimated time since divergence in some regions of the genome. This could be due to their shorter generation time, higher metabolic rate, increased population structuring, increased rate of speciation, or smaller body size. When studying ancient events like the Cambrian explosion under a molecular clock assumption, poor concurrence between cladistic and phylogenetic data is often observed. There has been some work on models allowing variable rate of evolution. Models that can take into account variability of the rate of the molecular clock between different evolutionary lineages in the phylogeny are called “relaxed” in opposition to “strict”. In such models the rate can be assumed to be correlated or not between ancestors and descendants and rate variation among lineages can be drawn from many distributions but usually exponential and lognormal distributions are applied. There is a special case, called “local molecular clock” when a phylogeny is divided into at least two partitions (sets of lineages) and a strict molecular clock is applied in each, but with different rates. Time-reversible and stationary models Many useful substitution models are time-reversible; in terms of the mathematics, the model does not care which sequence is the ancestor and which is the descendant so long as all other parameters (such as the number of substitutions per site that is expected between the two sequences) are held constant. When an analysis of real biological data is performed, there is generally no access to the sequences of ancestral species, only to the present-day species. However, when a model is time-reversible, which species was the ancestral species is irrelevant. Instead, the phylogenetic tree can be