Comparative Wood Anatomy in Pinaceae with Reference to Its Systematic Position

The wood anatomy of 132 species of the genera Abies, Cathaya, Cedrus, Keteleeria, Larix, Nothotsuga, Picea, Pinus, Pseudolarix, Pseudotsuga and Tsuga was studied to determine the elements that characterise the xylem of each genus and discuss possible groupings by wood anatomy for comparison with clades established by molecular phylogeny. The presence of resin canals and ray tracheids supports the family Pinaceae, although the absence of ray tracheids in Keteleeria and their occasional presence in Abies and Pseudolarix weakens it. Based on wood structure, Pinaceae clearly supports division into two groups, coinciding with molecular phylogeny: Pinoideae (Cathaya-Larix-Picea-Pinus-Pseudotsuga) and Abietoideae (Abies-Cedrus-Keteleeria-Nothotsuga-Pseudolarix-Tsuga). Although differences between genera are slight in Pinoideae, the Abietoideae group presents problems such as the presence of only axial resin canals in Keteleeria and Nothotsuga, absence of ray tracheids in Keteleeria and presence of traumatic radial resin canals in Cedrus. However, other features such as pitted horizontal walls and nodular end walls of ray parenchyma cells, indentures, scarce marginal axial parenchyma and presence of crystals in ray parenchyma strengthen the Abietoideae group.