Farewell to the requirement for character independence: phylogenetic methods to incorporate different types of dependence between characters

This paper discusses methods to take into account interactions between characters, in the context of parsimony analysis. These interactions can be in the form of some characters becoming inapplicable given certain states of other, primary characters; in the form of only certain states being allowed in some characters when a given state or set of states occurs for other characters; or in the form of transformation costs in some character being higher or lower when other characters have certain states or transformations between states. Character–state reconstructions and evaluation of trees under the assumption of independence may easily lead to ancestral assignments that violate elementary rules of biomechanics, well-established theories relating form and function or ideas about character co-variation. An obvious example is reconstructing an ancestral bird as wingless and flying at the same time; another is reconstructing a protein-coding gene as having a stop codon in some ancestors. If the characters are optimized independently, such chimeric ancestral reconstructions can occur even when no terminal displays the impossible combination of states. A set of conventions (implemented via new TNT commands and options) allows the definition of complex rules of interaction. By recoding groups of characters with proper step–matrix costs (and excluding impossible combinations from the set of permissible states), it is possible to find the ancestral reconstructions that maximize homology (and thus the degree to which similarities can be explained by common ancestry), within the constraints imposed by the rules specified by the user. We expect that considerations of biomechanics, functional morphology and natural history will be a source of many theories on possible character dependences, and that the present implementation will encourage users to take the possibility of character dependences into account in their phylogenetic analyses.