Transitional changes in arborescent lignophytes at the Devonian–Carboniferous boundary

It is usually considered that after the extinction of the Devonian tree Archaeopteris, no new arborescent lignophytes were established before the late Tournaisian. A reassessment of this pattern is presented here based on a three-fold approach: a re-evaluation of the taxic diversity of Tournaisian lignophyte trees based on descriptions of new plants from palaeotropical latitudes, a study of the patterns of phenotypic changes occurring among early lignophytes using a principal coordinate analysis and a phylogenetic analysis of the affinities of the arborescent taxa. The best supported results indicate that a substantial taxonomic and phenotypic diversity of arborescent lignophytes was already established in the first part of the Tournaisian, including some taxa that persisted until the Serpukhovian. Two genera may have originated in the Late Devonian and crossed the Devonian–Carboniferous boundary. Fewer originations and a decrease in phenotypic diversity occurred in the Viséan. The phenotypic distinctiveness of tree stems compared with those of other growth forms in the lignophytes is assessed. We propose a scenario in which the presence of lignophyte trees is continuous across the Devonian–Carboniferous boundary, with arborescent taxa distinct from Archaeopteris already present in the latest Devonian, possibly in upland floras, and diversifying significantly soon after the Devonian–Carboniferous boundary.