Unresolved problems on the origin and early evolution of land plants

The origin of land plants or embryophytes from the Charophyceae is generally accepted today by the botanists. In fact, numerous morphological, cytological, ultrastructural, biochemical and molecular characters are shared in these organisms. A fundamental problem is still constituted by the evolution of the sporophyte, i.e. the appearance of two different phase cycles (gametophyte/sporophyte alternance), although two theories ("antithetic" and "homologous") try to explain this evolutionary event.However, another phylogenetic dilemma is represented, in my opinion, either by the formation of bryophytes or by the transition from these first land plants to the pteridophytes, considering them at whole organism level. The bryophyte gametophyte is the most elaborate of the land plants. It presents several complex characters, principally the growth developmental form, the appearance of multicellular sex organs, antheridia and archegonia. Also the sporophyte shows a complicated structure that is not found in the other land plants or tracheophytes. The sporangium, in particular, exhibits some intricate morphological traits such as the peristome of true mosses for spore dispersion, the elaters of liverworts and the indeterminate growth in the hornworts. The pteridophytes are represented especially by their dominant sporophyte. This latter has the capacity to produce multiple sporangia and, in many cases, two kinds of spores which develop in male and female gametophyte (heterosporous pteridophytes). Another important characteristic of this sporophyte is its ability to become independent of the gametophyte. However, one of the most innovative character is the formation of true vascular elements (xylem and phloem). All these very large evolutionary jumps are discussed on the basis of the phyletic gradualistic neo-Darwinian theory and the punctuated equilibrium theory of Eldredge and Gould. In this context other genetic evolutionary mechanisms are also considered.Nevertheless, the origin of bryophytes and pteridophytes remain, at the moment, a mystery.