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Some widely distributed timber species, such as oaks, are vulnerable to low soil water availability and drought. Therefore, selecting broadleaved minor species with lower sensitivity to drought could be an alternative in seasonally dry areas. However, the growth performance of these minor hardwood species is unknown, particularly under warm and dry conditions at the southern limit of their distribution range. We assessed the radial growth response to drought by correlating tree-ring and climate data in coexisting service trees (Sorbus domestica) and two oak species (Quercus faginea, Quercus pyrenaica). Trees were sampled in three Mediterranean sites located in Spain with different precipitation patterns. We used the Vaganov–Shashkin (VS) growth model to infer the main climate constraints of growth. To ascertain whether climate is changing tree phenology, we also simulated changes in xylem onset and cessation timings and compared them with leaf falling dates in service trees. Service trees showed a trend towards advancing leaf fall, but this was not related to xylem growth. Oaks responded more to a drought index than service trees. The strongest responses corresponded to droughts peaking from spring to summer, but oaks tended to respond to longer droughts (9–13 months) than service trees (2–7 months). These different responses are due to the positive responses of all species to high precipitation in the growing season, but the sensitivity of oak growth to warm summer conditions and increased atmospheric water demand. The VS model indicated a more bimodal growth pattern in services trees than in oaks, which could allow the former to recover better after a dry summer. Service trees could be more widely used as a source of valuable timber wood under Mediterranean continental conditions.

Different leaf (evergreen vs. deciduous habit) and xylem (diffuse-vs. ring-porous wood) traits represent contrasting strategies to face seasonal changes in water availability and temperature. However, how contrasting leaf and xylem habits of coexisting tree species affect stem wood formation and tree-ring development remains poorly understood. Here, we investigated the spatio-temporal patterns of wood formation in two deciduous oaks (Quercus faginea and Quercus petraea) and two evergreen oaks (Quercus ilex and Quercus suber) coexisting in seasonally dry Mediterranean forests along an aridity gradient in Spain. We hypothesized that growth responses to drought and intra- and inter-annual growth patterns would differ between functional groups. We simulated intra- and inter-annual growth using a modified version of the Vaganov-Shashkin (VS) process-based, growth model. The VS model simulations were used to estimate growth changes under a high emission scenario (RCP 8.5) for the current distribution of the study oak species and to forecast their future performance under warm (4.8 ​°C) conditions in the Iberian Peninsula. Our simulations indicate that climate warming would induce a shortening of the ring-growth season and a reduction of radial growth in evergreen and deciduous Mediterranean oaks, particularly in dry sites from southern and eastern Iberia currently occupied by Q. ilex and Q. faginea. Evergreen oaks may better recover after dry periods than deciduous oaks by resuming growth after the summer drought. Low soil water availability in spring would be more detrimental to growth of deciduous oaks. Process-based growth models should be refined and validated to better forecast changes in tree growth as a function of climate.

Despite growing interest in predicting plant phenological shifts, advanced spring phenology by global climate change remains debated. Evidence documenting either small or large advancement of spring phenology to rising temperature over the spatio-temporal scales implies a potential existence of a thermal threshold in the responses of forests to global warming. We collected a unique data set of xylem cell-wall-thickening onset dates in 20 coniferous species covering a broad mean annual temperature (MAT) gradient (−3.05 to 22.9°C) across the Northern Hemisphere (latitudes 23°–66° N). Along the MAT gradient, we identified a threshold temperature (using segmented regression) of 4.9 ± 1.1°C, above which the response of xylem phenology to rising temperatures significantly decline. This threshold separates the Northern Hemisphere conifers into cold and warm thermal niches, with MAT and spring forcing being the primary drivers for the onset dates (estimated by linear and Bayesian mixed-effect models), respectively. The identified thermal threshold should be integrated into the Earth-System-Models for a better understanding of spring phenology in response to global warming and an improved prediction of global climate-carbon feedbacks.

Intra-annual wood density fluctuations are valuable indicators of the role of the local climate and provenance origins on tree growth. These intra-ring wood features were the most informative about local adaptation and phenotypic plasticity ofPinus halepensisMill. in a provenance trial. This highlights the importance of taking provenance into account when selecting forest reproductive material to mitigate the effects of climate change. Context: It is essential to disentangle the influence of climate from the effect of provenance on inter- and intra-annual wood traits to identify populations’ responses to climate changes. This will be even more relevant in the drought-prone Mediterranean region, where species have high genetic differentiation among populations. Aims: We investigated the effects of provenance and climate driving Pinus halepensis Mill. wood traits, as imprinted phenotypic adaptations to the Mediterranean climate. Methods: Inter- and intra-annual wood trait variation (ring-, early-, and latewood width, wood density) was studied by microdensitometry in 8 provenances from Spain, France, Greece, and Italy in a common-garden experiment. We related wood traits and density fluctuations with inter-annual climate variations at the trial site and the provenance origin. Results: Provenances showed distinct responses to climate regarding tree growth, wood density, and intra-annual fluctuations. Density fluctuations in latewood were more frequent than in earlywood. Climate at the provenance origin was a more important force for density fluctuations than climatic variations at the planting site, with different trends between coastal and inland provenances, and higher frequency of latewood fluctuations in provenances from arid and semiarid sites. Conclusion: Differences in P. halepensis wood density fluctuations confirmed the imprinted origin provenance control in response to climate variations and indicated a relation with local adaptation of these xylem anatomical

Tree species inhabiting riparian forests under Mediterranean climate have evolved to face summer water shortage but may fail to cope with future increases in drought severity. Thus, understanding tree growth phenological variations in response to environmental conditions is necessary to assess the impact of seasonal drought in riparian forests. In this study, we investigated the response of stem radial growth to climate in the narrow-leaved ash (Fraxinus angustifolia) over its distribution in southern Europe. We simulated intra- and inter-annual growth patterns using the Vaganov-Shashkin (VS) model considering five sites subjected to summer drought but showing different climate conditions. The growth pattern in this species varied from unimodal in cool-wet sites to facultative bimodal in warm-dry sites. Bimodal patterns were characterized by two growth peaks coinciding with favorable climate conditions in spring and autumn. The spring growth peak occurs earlier (May) in warm-dry sites than in wet-cool sites (June–July). The variation in the season growth length and growth timing suggests different strategies adopted by this species to cope with summer drought. The VS model revealed different growth patterns across which would be relevant in predicting the response of this and other riparian tree species to climate warming and aridification. Differences in the length of the growing season, timings of growth peaks and the shift from unimodal to bimodal growth patterns should be considered when assessing growth adjustments to future climate scenarios.

Climate warming is expected to lengthen the growing season of tree species and enhance radial growth rates. Alternatively, a longer growing season could not lead to improved radial growth if wood production depends more on growth rate than on growing season length. We test these ideas by comparing leaf phenology data and the estimated start and end dates of wood formation predicted by the VS-Lite growth model. We analyzed long-term series of leaf unfolding and fall dates and reconstructed radial growth of two pine species under contrasting climatic conditions: Scots pine (Pinus sylvestris) in a Russian boreal site and Aleppo pine (Pinus halepensis) in a Spanish Mediterranean site. On average, leaf onset occurred in days 99 and 163 in P. halepensis and P. sylvestris, respectively, about 40 days earlier than the estimated start date of wood formation. The onset of leaf unfolding advanced 2.1 days per decade in P. sylvestris in response to warmer May temperatures. Radial growth was enhanced by warm-wet spring-summer conditions in P. sylvestris and by wet soil conditions from prior winter up to current summer in P. halepensis. In this species the growing season length and the radial growth rate were not coupled because the growing season length shortened during cool-wet periods whereas growth rates increased. In P. sylvestris leaf onset was delayed during years with low growth rates suggesting a potential coupling between warmer spring conditions, earlier leaf onset and enhanced growth whenever soil water content is high enough. Overall, we show that longer growing seasons do not necessarily imply higher radial growth rates.

Wood formation during the growing season is shaped by the intra-annual variability of climatic conditions. In the Mediterranean, the cambial activity is seasonally constrained by winter low temperature and summer drought, resulting in bimodal growth patterns. Although bimodal growth is an ecologically important adaptation of woody species to seasonally dry environments, its variability across space and among species remains unclear. We combined direct monitoring of intra-annual radial growth using dendrometers and xylogenesis micro-sampling with indirect wood formation modeling to assess growth bimodality for two coexisting oak species across six sites in North-Eastern Spain. We focused on species with two contrasting life strategies, evergreen with diffuse-porous wood (Quercus ilex) and winter deciduous with ring-porous wood (Q. faginea/Q. pubescens). Study sites spanned from oceanic areas with high autumn precipitation to dry continental inland regions. We hypothesized that growth bimodality reflects the interaction between local precipitation patterns and species-specific traits. Our results revealed cambial activity in autumn after summer quiescence across the entire region and for both oak species. However, growth bimodality and the relative contribution of the autumn growth peak to the total annual growth was geographically structured and species-specific. Growth was unimodal under continental and Atlantic conditions. By contrast, growth bimodality was significant along the Mediterranean coast, where precipitation showed a bimodal pattern. Moreover, evergreen Q. ilex showed more growth bimodality compared with deciduous Q. faginea/Q. pubescens at Mediterranean sites. Different intra-annual distribution of radial growth could facilitate coexistence of both oak species through temporal niche partitioning. The growth bimodality helps to compensate for reduced growth in summer by autumn cambial resumption and, therefore, the differences in its intensity between si