New insights into Paleozoic charophyte morphology and phylogeny.

Examination of Paleozoic charophyte fructifications using microscopy and high-resolution x-ray synchrotron microtomography has revealed that most of them have a utricle that forms a supplementary calcified cover around the gyrogonite. All Paleozoic families with utricles have been assigned to the Sycidiales. We consider the Moellerinaceae to occupy a central position in the phylogeny of the Charophyta. From these, one group of descendants constitutes the gyrogonites inside the utricles of the Sycidiales; a second descendant goup is thought to be the utricle-free ancestors of the Charales prior to inversion of spiralization and reduction in cell number. The Sycidiales have a multilayered wall and an internal vesicle, but their families are distinguished by diversity in orientation of external cells, complexity of the utricle wall, and in presence or absence of antheridia. The solidly packed structure of the utricle is believed to be an organ protecting the zygote against dessication. We interpret the morphological similarities between Paleozoic Sycidiales and Mesozoic Clavatoraceae, both with a utricle, as homoplasous rather than expressing a true phylogenetic relationship. We confirm that some umbellids might correspond to utricles of charophytes.