Maximum Likelihood and Minimum-Steps Methods for Estimating Evolutionary Trees from Data on Discrete Characters

Felsenstein, J. (Department of Genetics SK-50, University of Washington, Seattle, Washington 98195). 1973. Maximum likelihood and minimum-steps methods for estimating evolutionary trees from data on discrete characters. Syst. Zool. 22:240-249.The general maximum likelihood approach to the statistical estimation of phylogenies is outlined, for data in which there are a number of discrete states for each character. The details of the maximum likelihood method will depend on the details of the probabilistic model of evolution assumed. There are a very large number of possible models of evolution. For a few of the simpler models, the calculation of the likelihood of an evolutionary tree is outlined. For these models, the maximum likelihood tree will be the same as the "most parsimonious" (or minimum-steps) tree if the probability of change during the evolution of the group is assumed a priori to be very small. However, most sets of data require too many assumed state changes per character to be compatible with this assumption. Farris (1973) has argued that maximum likelihood and parsimony methods are identical under a much less restrictive set of assumptions. It is argued that the present methods are preferable to his, and a counterexample to his argument is presented. An algorithm which enables rapid calculation of the likelihood of a phylogeny is described. [Evolutionary trees: maximum likelihood. ] The first systematic attempt to apply standard statistical inference procedures to the estimation of evolutionary trees was the work of Edwards and Cavalli-Sforza (1964; see also Cavalli-Sforza and Edwards, 1967). At about the same time, the "parsimony' or minimum evolutionary steps method of Camin and Sokal (1965) gave a great impetus to the development of welldefined procedures for obtaining evolutionary trees. Edwards and Cavalli-Sforza concerned themselves with data from continuous variables such as gene frequencies and quantitative characters. The CaminSokal approach, on the other hand, was developed for characters which are recorded as a series of discrete states. Although some taxonomists have declared that the problem of guessing phylogenies should be viewed as a problem of statistical inference (Farris, 1967, 1968; Throckmorton, 1968), until recently there have been no attempts to explore the relationship between the statistical inference and minimum-steps approaches. Recently, Farris (1973) has presented a detailed argument that, under certain reasonable assumptions, the maximum-likelihood method of statistical inference appropriate to discrete-character data is precisely the parsimony method of Camin and Sokal. In this paper, I will examine the application of maximum likelihood methods to discrete characters, and will show that parsimony methods are not maximum likelihood methods under the assumptions made by Farris. They are maximum likelihood methods under considerably more restrictive assumptions about evolution. METHODS OF MAXIMUM LIKELIHOOD Suppose that we want to estimate the evolutionary tree, T, which is to be specified by the topological form of the tree and the times of branching. We are given a set of data, D, and a model of evolution, M, which incorporates not only the evolutionary processes, but also the processes of sampling by which we obtained the data. This model will usually be probabilistic, involving random events such as changes of the environment, occurrence of favorable