Cells and tissues in the vegetative sporophytes of early land plants

Remarkable preservation in coalified and pennineralized fossils from Upper Silurian and Lower Devonian sediments deposited some 420 to 390 million years ago provides insight into the major anatomical innovations associated with the early stages in the colonization of the land by higher plants. Using uniformitarian principles, such information, combined with gross morphology, can then be used to reconstruct the pioneers as growing, metabolizing and reproducing organisms, as well as allowing assessment of affinity, although apart from the lycophytes, they have no close relationship with extant groups. In considering vascular tissues, diversity is exemplified by descriptions of the metaxylem in protosteles of Zosterophyllopsida and Drepanophycopsida (putative lycophytes), of Psilophyton (Trimerophytopsida), of the Rhyniaceae (including Rhynia gwynnevaughanii, certain Taeniocrada spp., Sennicaulis) and of Cooksonia pertoni, a Lower Devonian representative of the organization found in the oldest pteridophyte-like land plants. Aglaophyton major is included as a plant with bryophyte-like vascular tissues in a branching sporophyte with cuticle, stomata and intercellular space system typical of the homoiohydric tracheophyte. In all water-conducting cells, interpretation of the primary and secondary wall results from comparisons involving the anatomy and chemistry of extant examples and an understanding of taphonomic processes. Phloem is only rarely preserved and usually identified from its position around the xylem. In contrast, dermal features are better known, because of penetration of the resilient cuticle between epidermal cells. They appear conservative. Thus stomata with two guard cells look remarkably similar to extant forms in surface view, and by consideration of their relationship with surrounding epidermal cells and of extent of cuticularization, seem to have operated in a similar way to those in certain mosses and ferns. Cuticular ornamentation (papillae, striations) and epidermal outgrowths (unicellular and multicellular) are described and their adaptive significance conjectured. Ground tissue systems are best preserved in Rhynie Chert permineralizations where zonation of the cortex is tentatively related to photosynthetic and structural roles. The latter function is also attributed to thick-walled, outer cortical cells recorded in many zosterophylls and in Psilophyton, although the chemical nature of the walls themselves remains equivocal. Problems of identification of roots in plants possessing axial organization with exarch xylem, and in leafy plants with smooth lateral branching systems are briefly addressed. Finally axis apices in Rhynia givynne-vaughanii and sections showing stages in the maturation of Asteroxylon stems are described from the Rhynie Chert