In this work, an individual-based approach was used to assess the relative importance of tree age, size, and competition in modulating the individual dendroclimatic response of Quercus robur L. This ...was performed in a multi-aged forest in northwestern Spain under a wet Atlantic climate. All trees in five replicated forest stands with homogeneous soil conditions were mapped and inter-tree competition was quantified with a distance-dependent competition index. Tree rings of cored trees were crossdated and total age was estimated on individuals where the pith was missed. The climatic response was evaluated by bootstrapped correlations of individual tree-ring chronologies with climatic records. Inter-annual growth variation, i.e., mean sensitivity, was independent of tree age and bole diameter, but modulated by competition. Water excess in previous summer-autumn and spring negatively affected growth, while warmer September conditions favored growth. Individual response to climate was independent of tree age, but related to the joint effect of tree bole diameter and competition. Larger oaks in less competitive environments responded more plastically to climatic stress, while smaller trees under high competition levels were less responsive to climate. Strong inter-tree competition reduced growth plasticity but amplified the vulnerability of smaller oaks to the particularly rainy conditions of the study area. These findings suggest that inter-tree competition is a relevant size-mediated extrinsic factor that can potentially modulate individual radial growth variation and its response to limiting climatic conditions in temperate deciduous forests. This study highlights the value of individual-based approach as a useful tool that informs about the relative contribution of factors modulating the climatic response of tree-ring growth.
Tree ring records are among the most valuable resources to create high-resolution climate reconstructions. Most climate reconstructions are based on old trees growing in inaccessible mountainous ...areas with low human activity. Therefore, reconstruction of climate conditions in lowlands is usually based on data from distant mountains. Albeit old trees can be common in humanized areas, they are not used for climate reconstructions. Pollarding was a common traditional management in Europe that enabled trees to maintain great vitality for periods exceeding the longevity of unmanaged trees. We evaluate the potential of pollarded deciduous oaks to record past climate signal. We sampled four pollarded woodlands in Central Spain under continental Mediterranean climate. We hypothesized that pollarded trees have a strong response to water availability during current period without pollarding management, but also in the period under traditional management if pruning was asynchronous among trees. Moreover, we hypothesized that if climate is a regional driver of oak secondary growth, chronologies from different woodlands will be correlated. Pollard oaks age exceeded 500 years with a strong response to Standardized Precipitation-Evapotranspiration Index (SPEI) from 9 to 11 months. Climate signal was exceptionally high in three of the sites (r2 = 0.443–0.655) during low management period (1962–2022). The largest fraction of this climate signal (≈70 %) could be retrieved during the traditional management period (1902–1961) in the three sites where pollarding was asynchronous. Chronologies were significantly correlated since the 19th century for all the studied period, highlighting a shared climate forcing. We identified critical points to optimize pollard tree sampling schema. Our results show the enormous potential of pollarded woodlands to reconstruct hydroclimate conditions in the Mediterranean with a fine spatial grain. Studying pollarded trees is an urgent task, since the temporal window to retrieve the valuable information in pollarded trees is closing as these giants collapse and their wood rots.
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•Pollarded trees are not used for climate reconstructions due to recurrent management.•Pollarded oaks can live >500 years in intensively managed territories.•Pollarded oaks secondary growth encapsulates valuable climate information.•We have few decades left to retrieve pollarded trees environmental information.
Abstract
Rear‐edge tree populations are experiencing a combination of higher temperatures and more intense droughts that might push individuals beyond their tolerance limits. This trend towards ...rising atmospheric CO
2
is concurrent with an increase in intrinsic water use efficiency (iWUE), which theoretically enhances photosynthesis and decrease evapotranspiration rates, consequently improving tree resistance to drought. However, it remains unclear whether iWUE is favouring tree growth under current climate conditions, particularly when climate and iWUE legacy effects are simultaneously considered.
We evaluated this question with an extensive sampling along Iberian rear‐edge (dry) populations comprising four mountain ranges and two distinct altitudes. We simultaneously examined the effects of climate and iWUE on secondary growth using annually resolved basal area increments (BAIs) for the period 1901–2017. We used linear mixed models including second‐order autocorrelation and 1‐year legacy effects of iWUE and summer drought.
BAI and iWUE increased across the studied period. iWUE increase was driven by changes in atmospheric CO
2
concentration and water availability during the growing season. Climate and iWUE exerted direct and lagged effects on beech growth. Water availability during growing season was the main driver of tree growth, combining direct and indirect effects through its impact on iWUE. Legacy effects of water availability and iWUE were more important than growing season conditions. The net effect of iWUE shifted when lagged effects were considered, resulting in a net negative impact on tree growth.
Synthesis
: Our results reveal that climate and iWUE legacy effects must be considered to assess the net iWUE effect on secondary growth. Considering lagged effects, the current increase in iWUE is constraining tree growth. Modelling efforts of tree growth response to climate warming should include climate and iWUE legacy effects to adequately assess terrestrial ecosystem carbon balance.
Key message
Sex and site conditions modulate intra- and inter-annual secondary growth and its climatic sensitivity in a dioecious Mediterranean conifer.
Divergent evolutionary pressures associated ...with differential reproductive costs in dioecious trees may lead to sex-related variation in non-reproductive functions. Sex-related differences may be site-dependent, with different outcomes depending on environmental conditions. We explored the effects of sex and environmental conditions on the climatic control of annual secondary growth and intra-annual wood density fluctuations (IADF) of a dioecious conifer (
Juniperus thurifera
L.) growing in two sites with contrasting hydrological conditions under a continental Mediterranean climate. Different sex-related strategies had variable effects on relative secondary growth, with females outperforming males under more favorable hydrological conditions, and males outperforming females under water-limited conditions. Ring width and IADF formation were driven by climatic factors occurring at different temporal scales. Tree-ring growth depended on factors acting prior to the initiation of the xylogenesis and to conditions directly affecting the duration and pace of cambial activity, and ring width, therefore, integrated a complex signal of factors occurring over a relatively long period, and on an annual cycle. In contrast, IADFs responded to singular short-term events that alleviated drought and promoted cambial reactivation during the summer arrest. Female trees showed a more opportunistic water use, displayed in the stronger ring-width response to June–July conditions. Enhanced cambial sensitivity in females set a lower threshold for IADF occurrence, leading to a higher frequency of IADFs irrespective of site. Intra-annual and inter-annual female growth patterns reflect an opportunistic strategy to benefit from favorable climatic windows.
In order to understand the effects of changes in traditional forest management on Spanish juniper (
Juniperus thurifera L.) forest dynamics, height and radial growth patterns were studied in 107 ...juniper trees in Cabrejas del Pinar (Soria, Central Spain).
Suppressions in height and radial growth were common until the second half of XIXth century. These suppressions occurred at relatively low tree heights, and affected 30% of the sampled trees. The individual suppressions ended abruptly with a sharp increase in height and radial growth. These results fit well into the browsing cessation model, suggesting that an intense browsing pressure shaped this forest. The timing in the reduction of the browsing pressure matches the regional decrease in transhumant livestock that occurred in Spain during the second half of XIXth century. No difference in height-growth rate between suppressed and non-suppressed trees was observed for post-suppression period, indicating a high recovery capacity for
J. thurifera.
We observed a high variability in height-growth rate during post-suppression periods. This variability was poorly related to site factors (competition, landform), being related to tree life (age and presence of past suppression events). Traditional logging based solely on trees morphological traits may be partially responsible of this pattern.
The accuracy of direct (based on increment cores) and indirect (based on age-size relationships) methods of tree age estimation in Fagus sylvatica and Quercus robur was tested. This was done through ...increment cores and stem discs taken in an old-growth forest of Northern Spain. It was found that cross-dating was more precise than ring counting by up to 7 years per tree. Furthermore, cross-dating permitted the estimation of the age of trees with floating ring-width series, which were 7% of cored F. sylvatica and 40% of Q. robur ones. In partial cores with the arcs of the inner rings, the length of the missing radius was estimated with both a geometric method, based on the curvature of the arcs, and a new graphical method, based on the convergence of xylem rays at the pith. The graphical method was more accurate when the radial growth was eccentric, as happens in Q. robur, while both methods showed a similar accuracy for F. sylvatica, whose growth is relatively concentric. Empirical models of initial radial growth (IRG), built to estimate the number of missing rings, reduced the errors associated with other methods that assume constant growth rates. Age estimates obtained from the graphical method combined with the IRG models were within 4% of the actual age. This combination ensured age estimates with a mean accuracy of 8 years for 98% of the F. sylvatica trees, and 4 years for 89% of the Q. robur. In partial cores without the arcs of the inner rings, the length of the missing radius was estimated as the distance to the geometric centre of the tree. In that case, age estimates obtained by extrapolating the mean growth rate of the 20 innermost rings in the cores were from 10 to 20% of actual age, which coincided with results obtained in other tree species with this method. Finally, the age-diameter equations of the different cohorts produced better age estimates (from 8 to 14% of actual age) than equations of the population as a whole (from 20 to 40% of actual age). These results proved that the errors derived from doubtful assumptions, such as concentric radial growth, constant growth and recruitment rates, or the absence of anomalous rings, could be reduced by applying more realistic methods of tree age estimation.
Five contrasting deciduous forest stands were studied to characterize the spatial structural variability in human-influenced forests. These stands are representative of cultural forest types widely ...represented in western Europe: one plantation, two coppices, one wood-pasture forest and one high forest stand. All stems with DBH > 5 cm were measured and mapped, and stem DBH distributions, spatial structure of DBH, spatial point patterns and spatial associations were analysed. Spatial autocorrelation for DBH was calculated with Moran's I correlograms and semivariograms. Complete spatial randomness hypothesis for spatial point patterns, and both independence and random labelling hypotheses for spatial associations were analysed using Ripley's K function. The results showed that tree sizes were conditioned by particular former management systems, which determined unimodal symmetric, positively skewed or compound DBH distributions. Spatial structure was more complex when human influence became reduced. Coppice stands showed clumped spatial patterns and independence among size classes, as a consequence of sexual and vegetative establishment of new stems in open areas. The largest clumping intensity was observed in the wood-pasture with an intermediate disturbance frequency and low inter-tree competition. The high forest stand displayed spatial traits consistent with the gap-dynamics paradigm, such as clumping of smaller trees, random arrangement of larger trees, negative association between juveniles and adults, and high structural heterogeneity. It can be expected that after cessation of human interference, coppices and wood-pastures would evolve to a more heterogeneous structure, probably with a higher habitat and species diversity.
•We studied the climate response and drought́s resilience of N. obliqua tree growth across a climate gradient.•Growth was positively related to rainfall and negatively related to summer temperature ...across the area.•There was a trade-off between resistance and recovery in the response to droughts across the gradient.•Dry range edgés stands were highly resilient to drought and did not show a negative growth trend.•Tolerance to droughts depended more on site conditions than on regional climate.
The climate response and resilience of tree growth to drought events have been widely reported for forests from the Northern Hemisphere. However, studies are much scarcer in the extra-tropical forests of southern South America. Mediterranean and Temperate forests of Chile are suffering from a moderate warming and a sustained precipitation decrease, occurring on top of an unprecedented megadrought since 2010. This study evaluated tree-growth patterns, the climate response and drought resilience of nine secondary Nothofagus obliqua forests across a latitudinal gradient from Mediterranean to Temperate climate in the Andes of Chile (35.7° to 40.3° S). Moreover, to improve the understanding of the spatial variation in productivity patterns, this research assessed trends in the maximum Normalized Difference Vegetation Index (peak in the NDVI) across the gradient for 2001–2018. Tree-growth patterns were highly influenced by stand dynamics, with steep decreasing trends in most of the stands related to a gradual canopy closure. Productivity trends had a flat pattern north of 38oS, but positive trends south of this latitude, which were mostly attributed to stand development. Tree growth was positively related to precipitation in all the sites, with annual and summer rainfall being more important in the north (Mediterranean climate) and south (Temperate climate), respectively. Conversely, maximum temperature had a negative effect on growth in most of the studied forests. This implies that projected warmer and drier conditions may have a detrimental effect on N. obliqua growth during coming decades. The two northern stands, located at the species dry range edge, were among the most resilient to drought and have not been strongly affected by the current megadrought in the area. Overall climate conditions, however, do not define the tolerance of stands to droughts, likely because local environmental and forests conditions play a key role. Although droughts have not strongly impacted the growth of N. obliqua across its distribution so far, future studies should assess the effects of the current long-term megadrought on growth resilience, and physiological studies should address the impacts of droughts and heat waves on forest function beyond what growth can unveil.
Past regeneration patterns of
Quercus robur L. and
Fagus sylvatica L. and their relationship to canopy structure, disturbances and forest-use history were investigated in an old-growth, lowland ...forest in Cantabria, Northern Spain. Dendroecological techniques were used to estimate tree ages and reconstruct disturbance histories in four representative stands which differed in composition and structure. Age estimates and tree locations were used to reconstruct patterns of tree establishment. Documentary sources were also used to report on changes in land-use during the past 250 years. Age structures of both species were discontinuous in time and space, and revealed two main cohorts distributed in even-aged, spatially segregated patches. Juveniles showed a clumped spatial pattern at varying distances, indicating an establishment in more or less extensive patches. Establishment site analyses revealed that
Q. robur regenerated in canopy gaps, while
F. sylvatica recruited throughout the forest floor, but it displayed inhibition near mature conspecifics. A lack of tree establishment from the 1840s to the 1920s coincided with an intense grazing pressure from domestic animals.
F. sylvatica exhibited continuous recruitment in periods of forest protection against grazing, such as from the 1740s to the 1830s and from the 1920s. By contrast,
Q. robur recruitment was dependent on both low grazing intensities and disturbances that resulted in the expansion of existing canopy openings, as occurred in 1943, 1956 and 1967. Past selective cuttings, canopy closure and forest protection could be responsible for the increasing decline of oak dominance. Planting
Q. robur and controlling
F. sylvatica regeneration are therefore advisable to prevent the complete domination of beech in the future forest canopy.
•Beech growth and vessel area respond to precipitation at disparate temporal domains.•Previous summer precipitation favors secondary growth.•Precipitation during vessel expansion phase controls ...vessel area variability.•Water-shortage constraints on growth and anatomy are shared along beech dry edge.
The response of European beech (Fagus sylvatica L.) to climate warming will depend on the ability of their populations to adjust tree performance to water shortage. By exploring inter- and intra-annual variations in secondary growth and mean vessel area (MVA), we assessed the effects of precipitation on cambial activity and hydraulic control during the vessel expansion phase along tree lifes. We sampled beech populations at low and high altitude from four mountain ranges across its southwestern distribution edge. We measured a total of 45,897 rings from 126 trees and 5.5 million vessels from 76 trees. We built chronologies for ring width and MVA between 1950 and 2017, calculated their climate responses and evaluated the effects of region, altitude and chronology type (ring-width vs. MVA) by means of ordinations (PCA) and constrained ordinations (pRDA). Precipitation controlled ring width and MVA along beech's southwestern distribution range, but at different time domains. Ring width responded primarily to summer precipitation during the previous growing season, whereas MVA responded to water availability during the vessel expansion phase, with timing shifting along the ring, according to the moment of vessel expansion. Regional differences were significant, but low, compared with the effect of chronology type. A large part of the variance explained by region was due to the strong difference between Western Pyrenees forests –growing under hyperhumid conditions– and the rest of forests under drier and warmer climate. Only minor differences between altitudes were found for the climate control of ring width and vessel size at annual scale, and no intra-annual effect on climate control of MVA. The stronger effect of chronology type on climatic response compared to the role of geographical location or altitude suggests common climate constraints on secondary growth and xylem anatomy along beech dry edge.
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