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What constitutes a morphological character versus character state has been long discussed in the systematics literature but the consensus on this issue is still missing. Different methods of classifying organismal features into characters and character states can dramatically affect the results of phylogenetic analyses. Here, I show that the modular structure of the gene regulatory network (GRN) underlying trait development, and the hierarchical nature of GRN evolution, essentially remove thedoi:10.1101/420471 fatcat:yv33l5asgffj7jly3qrig7wf4u