Phylogenetic relationships of Echinocactus inferred with the total evidence matrix. Maximum parsimony strict consensus tree of 2 most parsimonious trees of 863 steps with a consistency index of 0.790 and a retention index of 0.770. Values above/below nodes indicate maximum parsimony bootstrap (mpb) and posterior probabilities (pp), respectively. Black circles indicate synapomorphic characters identified by the character optimisation analyses. Numbers above/below circles indicate characters and character states, respectively (see Appendix 2). The HEA clade is labelled showing the three genera: Astrophytum, Echinocactus and Homalocephala. Line drawings illustrate the morphological synapomorphies for respective clades.