Mosaic assembly of regulatory programs for vascular cambial growth: a view from the Early Devonian

Evidence for secondary growth extends into the Early Devonian, 407 million years ago, raising questions about tempo and mode of origination of this key developmental feature. To address such questions, we analyze anatomy in the four oldest fossil plants with well-characterized woody tissues; one of these represents a new genus, described here formally. The new fossil is documented using the cellulose acetate peel technique and associated methods. We use the paradigm of structural fingerprints to identify developmental components of cambial growth based on fossil anatomy. We integrate developmental inferences within a theoretical framework of modular regulation of secondary growth. The fossils possess structural fingerprints consistent with four different combinations of regulatory mechanisms (modules) acting in cambial growth, representing four distinct modes of secondary growth. The different modes of secondary growth demonstrate that cambial growth is an assemblage of regulatory modules whose deployment followed a mosaic pattern across woody plants, which may represent ancestors of younger lineages that exhibit woody growth. The diverse modes of wood development occupy a wide morphospace in the anatomy of wood in the Early Devonian, suggesting that the origins of secondary growth and of its modular components pre-date this interval.