Understanding how climate affects xylem formation is critical for predicting the impact of future conditions on tree growth and functioning in the Mediterranean region, which is expected to face warmer and drier conditions. However, mechanisms of growth response to climate at different temporal scales are still largely unknown, being complicated by separation between spring and autumn xylogenesis (bimodal temporal pattern) in most species such as Mediterranean pines. We investigated wood anatomical characteristics and carbon stable isotope composition in Mediterranean Pinus pinea L. along tree-ring series at intra-ring resolution to assess xylem formation processes and responses to intra-annual climate variability. Xylem anatomy was strongly related to environmental conditions occurring a few months before and during the growing season, but was not affected by summer drought. In particular, the lumen diameter of the first earlywood tracheids was related to winter precipitation, whereas the size of tracheids produced later was influenced by mid-spring precipitation. Diameter of latewood tracheids was associated with precipitation in mid-autumn. In contrast, tree-ring carbon isotope composition was mostly related to climate of the previous seasons. Earlywood was likely formed using both recently and formerly assimilated carbon, while latewood relied mostly on carbon accumulated many months prior to its formation. Our integrated approach provided new evidence on the short-term and carry-over effects of climate on the bimodal temporal xylem formation in P. pinea. Investigations on different variables and time scales are necessary to disentangle the complex climate influence on tree growth processes under Mediterranean conditions.

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Forecasted warmer and drier conditions will probably lead to reduced growth rates and decreased carbon fixation in long-term woody pools in drought-prone areas. We therefore need a better understanding of how climate stressors such as drought constrain wood formation and drive changes in wood anatomy. Drying trends could lead to reduced growth if they are more intense in spring, when radial growth rates of conifers in continental Mediterranean climates peak. Since tree species from the aforementioned areas have to endure dry summers and also cold winters, we chose two coexisting species: Aleppo pine (Pinus halepensisMill., Pinaceae) and Spanish juniper (Juniperus thuriferaL., Cupressaceae) (10 randomly selected trees per species), to analyze how growth (tree-ring width) and wood-anatomical traits (lumen transversal area, cell-wall thickness, presence of intra-annual density fluctuations-IADFs-in the latewood) responded to climatic variables (minimum and maximum temperatures, precipitation, soil moisture deficit) calculated for different time intervals. Tree-ring width and mean lumen area showed similar year-to-year variability, which indicates that they encoded similar climatic signals. Wet and cool late-winter to early-spring conditions increased lumen area expansion, particularly in pine. In juniper, cell-wall thickness increased when early summer conditions became drier and the frequency of latewood IADFs increased in parallel with late-summer to early-autumn wet conditions. Thus, latewood IADFs of the juniper capture increased water availability during the late growing season, which is reflected in larger tracheid lumens. Soil water availability was one of the main drivers of wood formation and radial growth for the two species. These analyses allow long-term (several decades) growth and wood-anatomical responses to climate to be inferred at intra-annual scales, which agree with the growing patterns already described by xylogenesis approaches for the same species. A plastic bimodal growth behavior, driven by dry summer conditions, is coherent with the presented wood-anatomical data. The different wood-anatomical responses to drought stress are observed as IADFs with contrasting characteristics and responses to climate. These different responses suggest distinct capacities to access soil water between the two conifer species.

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An improved understanding of how tree species will respond to warmer conditions and longer droughts requires comparing their responses across different environmental settings and considering a multi-proxy approach. We used several traits (tree-ring width, formation of intra-annual density fluctuations -IADFs, wood anatomy, ∆ 13 C and δ 18 O records) to retrospectively quantify these responses in three conifers inhabiting drought-prone areas in northwestern Mexico. A fir species (Abies durangensis) was studied in a higher altitude and slightly rainier site and two pine species were sampled in a nearby, lower drier site (Pinus engelmannii, Pinus cembroides). Tree-ring-width indices (TRWi) of the studied species showed a very similar year-to-year variability likely indicating a common climatic signal.Wood anatomy analyses done over 3.5 million measured cells and showed that P. cembroides lumen area was much smaller than in the other two species and it remained constant along all the studied period (over 64 years). Instead, cell wall thickness was widest in P. engelmannii and this species presented the highest amount of intra-annual density fluctuations. Climate and wood anatomy correlations pointed out that lumen area was positively affected by winter precipitation for all studied species, while cell-wall thickness was negatively affected by this season's precipitation in all species but P. cembroides. Stable isotope analysis showed significantly lower values of Δ 13 C for P. cembroides and no significant δ 18 O differences between the three species, although they shared a common decreasing trend. With very distinct wood anatomical traits (smaller cells, compact morphology), P. cembroides stood out as the better adapted species in its current environment and could be less affected by future drier climate. P. engelmannii and A. durangensis showed high plasticity at wood anatomical level, allowing them to promptly respond to seasonal water availability but likely gives few advantages on future climate scenarios with longer and frequent drought spells.