The root epidermis of tracheophytes consists of hair-forming cells (HCs) and non-hair cells (NCs). The HC distribution pattern is classified into three types: random (Type I), vertically alternating (Type II), and radial (Type III). Type III is found only in core eudicots and is known to be position-dependent in superrosids with HCs positioned between two underlying cortical cells. However, the evolution of Type III and the universality of its position dependency in eudicots remain unclear. We surveyed the HC distribution in basal and Type III-exhibiting core eudicots and conducted genomic analyses to get insight into whether eudicots share the same genetic network to establish Type III. Our survey revealed no canonical Type III in basal eudicots but a reverse Type III, with NCs between two cortical cells and HCs on a single cortical cell, in Papaveraceae of basal eudicots. Type III-exhibiting species from both superrosids and superasterids showed the canonical position dependency of HCs. However, some key components for Type III determination were absent in the genomes of Papaveraceae and Type III-exhibiting superasterids. Our findings identify a novel position-dependent type of HC patterning, reverse Type III, and suggest that Type III emerged independently or diversified during eudicot evolution.

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