Different xylogenesis responses to atmospheric water demand contribute to species coexistence in a mixed pine–oak forest

Abstract

Seasonal patterns of wood formation (xylogenesis) remain understudied in mixed pine–oak forests despite their contribution to tree coexistence through temporal niche complementarity. Xylogenesis was assessed in three pine species (Pinus cembroides, Pinus leiophylla, Pinus engelmannii) and one oak (Quercus grisea) coexisting in a semi-arid Mexican forest. The main xylogenesis phases (production of cambium cells, radial enlargement, cell-wall thickening and maturation) were related to climate data considering 5–15-day temporal windows. In pines, cambium activity maximized from mid-March to April as temperature and evaporation increased, whereas cell radial enlargement peaked from April to May and was constrained by high evaporation and low precipitation. Cell-wall thickening peaked from June to July and in August–September as maximum temperature and vapour pressure deficit (VPD) increased. Maturation of earlywood and latewood tracheids occurred in May–June and June–July, enhanced by high minimum temperatures and VPD in P. engelmannii and P. leiophylla. In oak, cambial onset started in March, constrained by high minimum temperatures, and vessel radial enlargement and radial increment maximized in April as temperatures and evaporation increased, whereas earlywood vessels matured from May to June as VPD increased. Overall, 15-day wet conditions enhanced cell radial enlargement in P. leiophylla and P. engelmannii, whereas early-summer high 15-day temperature and VPD drove cell-wall thickening in P. cembroides. Warm night conditions and high evaporation rates during spring and summer enhanced growth. An earlier growth peak in oak and a higher responsiveness to spring–summer water demand in pines contributed to their coexistence.