Search

Intraspecific variation in plant defences is expected to be the result of adaptive and plastic responses to environmental conditions, where trade-offs between growth and defences are thought to play a key role shaping phenotypic patterns in defensive investment. Axial resin ducts are costly defensive structures that remain imprinted in the tree rings of conifers, therefore being a valuable proxy of defensive investment along the trees’ lifespan. We aimed to disentangle climate-driven adaptive clines and plastic responses to both spatial and temporal environmental variation in resin duct production, and to explore growth-defence trade-offs. To that aim, we applied dendrochronological procedures to quantify annual growth and resin duct production during a 31-year-period in a Mediterranean pine species, including trees from nine populations planted in two common gardens. Both genetic factors and plastic responses modulated annual resin duct production. However, we found no evidence of adaptive clines with climate gradients driving population differentiation. Our results revealed a marked physiological trade-off between growth and defences, where the slope of the trade-off was genetically variable and associated with climatic gradients. Our results help to enlighten the evolutionary patterns and genetic basis of defensive allocation within species, particularly revealing a key role of growth-defence trade-offs.

Producción Científica, Macaronesian laurel forests are the only remnants of a subtropical palaeoecosystem dominant during the Tertiary in Europe and northern Africa. These biodiverse ecosystems are restricted to cloudy and temperate insular environments in the North Atlantic Ocean. Due to their reduced distribution area, these forests are particularly vulnerable to anthropogenic disturbances and changes in climatic conditions. The assessment of laurel forest trees’ response to climate variation by dendrochronological methods is limited because it was assumed that the lack of marked seasonality would prevent the formation of distinct annual tree rings. The aims of this study were to identify the presence of annual growth rings and to assess the dendrochronological potential of the most representative tree species from laurel forests in Tenerife, Canary Islands. We sampled increment cores from 498 trees of 12 species in two well-preserved forests in Tenerife Island. We evaluated tree-ring boundary distinctness, dating potential, and sensitivity of tree-ring growth to climate and, particularly, to drought occurrence. Eight species showed clear tree-ring boundaries, but synchronic annual tree rings and robust tree-ring chronologies were only obtained for Laurus novocanariensis, Ilex perado subsp. platyphylla, Persea indica and Picconia excelsa, a third of the studied species. Tree-ring width depended on water balance and drought occurrence, showing sharp reductions in growth in the face of decreased water availability, a response that was consistent among species and sites. Inter-annual tree-ring width variation was directly dependent on rainfall input in the humid period, from previous October to current April. The four negative pointer years 1995, 1999, 2008 and 2012 corresponded to severe drought events in the study area. This study gives the first assessment of dendrochronological potential and tree-ring climate sensitivity of tree species from the Tenerife laurel forest, which opens new research avenues for dendroecological studies in Macaronesian laurel forests., Ministerio de Ciencia, Innovación y Universidades (projects PID2019-109906RA-I00, PID2020-118444GA-100 and PID2019-106908RA-I00/AEI/10.13039/501100011033), Ministerio de Ciencia, Innovación y Universidades (predoctoral contract PRE2018-084106), Ministerio de Economía, Industria y Competitividad (project CGL2017-87309-P and postdoctoral grant IJC2019-040571-I), Junta de Castilla y León (projects VA113G19 and IR2020-1-UVA08), Universidad de Valladolid (predoctoral contract 113-2019PREUVA22), Comunidad de Madrid (project S2018/EMT-4338)

Producción Científica, Tree pollarding was a dominant management strategy of European forests for centuries creating open agroforestry landscapes with important cultural and environmental values. This traditional practice has been widely abandoned in last decades with a subsequent impact in terms of biodiversity and cultural loss. Central Spain hosts the largest and best-preserved area of pollarded narrow-leaved ash (Fraxinus angustifolia Vahl.) woodlands in Europe. The main aim of this research is to obtain rigorous historical records of pollarding frequency to get adequate information for traditional ash management. We used dendrochronological techniques to evaluate temporal changes of pollarding frequency and rotation length. We analysed the stand level synchrony and the effect of land property on pollarding activity from 322 trees growing in eight pollard stands in Central Spain. Pollarding events were unequivocally identified at tree level by a characteristic change in growth pattern. We identified 2426 tree-level pruning events with the first event dated in 1777. Historical pruning recurrence ranged between 5 and 10 years with higher pollarding frequency on private lands. Pruning events within each site were synchronous, suggesting the existence of a rotational schema within each stand. Pruning frequency decreased drastically in the 1970s matching with the depopulation of rural areas and the general abandonment of traditional practices. Pollarding practices have recovered in recent decades although with lower intensity and lacking the synchronic historical patterns. Providing technical and economic support to make this traditional activity profitably would have strong environmental revenue due the multiple ecosystem services provided by pollarded ashes., iuFOR Institute Unit of Excellence" of the University of Valladolid, funded by the Junta de Castilla y León and co-financed by the European Union (ERDF "Europe drives our growth") project (CLU-2019-01), Ministerio de Ciencia e Innovación y Agencia Estatal de Investigación - MCIN/AEI/10.13039/501100011033 project PROWARM (PID2020-118444GA-100),(IJC2019-040571-I),(grant PRE2018-084106), Junta de Castilla y León, projects, (VA171P20) and (IR2020-1-UVA08), Publicación en abierto financiada por el Consorcio de Bibliotecas Universitarias de Castilla y León (BUCLE), con cargo al Programa Operativo 2014ES16RFOP009 FEDER 2014-2020 DE CASTILLA Y LEÓN, Actuación:20007-CL - Apoyo Consorcio BUCLE

21 Pags.- 5 Figs.- 1 Appendix. - Suppl. Data., This study links tree-ring growth and gross primary production for a variety of forest types under different environmental conditions across Spain. NOAA-AVHRR satellite imagery data were combined with dendrochronological records and climate data at a fine spatial resolution (1.21 km2) to analyze the interannual variability of tree-ring growth and vegetation activity for different forest biomes from 1981 to 2015. Specifically, we assessed the links between tree-ring width indices (TRWi), the Normalized Difference Vegetation Index (NDVI) and a variety of environmental conditions, represented by climatic variables (air temperature, precipitation, evapotranspiration and water balance) and elevation. The impact of these variables on tree growth was assessed by means of the Predictive Discriminant Analysis (PDA). Results reveal a general positive and significant relationship between inter-annual variability of the NDVI at a high spatial resolution (1.21 km2) and tree-ring growth. Maximum correlations between NDVI and tree-ring growth were recorded when cumulative NDVI values were considered, in some cases covering long time periods (6–10 months), suggesting that tree growth is mainly related to Gross Primary Production (GPP) at annual scale. The relationship between tree-ring growth and inter-annual variability of the NDVI, however, strongly varies between forest types and environmental conditions., This work was supported by the research projects CGL2014-52135-C03-01, CGL2015-69186-C2-1-R, CGL2015-69985-R, CGL2013-48843-C2-1-R, AGL2014-53822-C2-1-R, (BFU2010-21451), PCIN-2015-220, CGL2016-81706-REDT and CGL2017-82216-R financed by the Spanish Commission of Science and Technology and FEDER, 1560/2015: Herramientas de monitorización de la vegetación mediante modelización ecohidrológica en parques continentales financed by the Red de Parques Nacionales, IMDROFLOOD financed by the Water Works 2014 co-funded call of the European Commission, CROSSDRO financed by the AXIS (Assessment of Cross(X) – sectorial climate Impacts and pathways for Sustainable transformation) JPI-Climate co-funded call of the European Commission and INDECIS, which is part of ERA4CS, an ERA-NET initiated by JPI Climate, and funded by FORMAS (SE), DLR (DE), BMWFW (AT), IFD (DK), MINECO (ES), ANR (FR) with co-funding by the European Union (Grant 690462). Financed by the Spanish Commission of Science and Technology and FEDER, and INDECIS, which is part of ERA4CS, an ERA-NET initiated by JPI Climate, and funded by FORMAS (SE), DLR (DE), BMWFW (AT), IFD (DK), MINECO (ES), ANR (FR) with co-funding by the European Union (Grant 690462). Natalia Martin-Hernandez was supported by a doctoral grant by the Aragon Regional Government, Marina Peña-Gallardo was granted by the Spanish Ministry of Economy and Competitiveness and Miquel Tomas-Burguera was supported by a doctoral grant by the Spanish Ministry of Education, Culture and Sport. Raúl Sánchez-Salguero, Antonio Gazol and Gabriel Sangüesa-Barreda were supported by Postdoctoral grants (IJCI-2015-25845, MINECO-FPDI 2013-16600 and FJCI 2016-30121, respectively; FEDER funds). This work also benefited from funding from Xunta de Galicia (PGIDIT06PXIB502262PR, GRC GI-1809, ROCLIGAL-10MDS291009PR), INIA (RTA2006-00117), and Interreg V-A POCTEFA (CANOPEE, 2014-2020-FEDER funds) projects.

Producción Científica, Drought is one of the key natural hazards impacting net primary production and tree growth in forest ecosystems. Nonetheless, tree species show different responses to drought events, which make it difficult to adopt fixed tools for monitoring drought impacts under contrasting environmental and climatic conditions. In this study, we assess the response of forest growth and a satellite proxy of the net primary production (NPP) to drought in peninsular Spain and the Balearic Islands, a region characterized by complex climatological, topographical, and environmental characteristics. Herein, we employed three different indicators based on in situ measurements and satellite image-derived vegetation information (i.e., tree-ring width, maximum annual greenness, and an indicator of NPP). We used seven different climate drought indices to assess drought impacts on the tree variables analyzed. The selected drought indices include four versions of the Palmer Drought Severity Index (PDSI, Palmer Hydrological Drought Index (PHDI), Z-index, and Palmer Modified Drought Index (PMDI)) and three multi-scalar indices (Standardized Precipitation Evapotranspiration Index (SPEI), Standardized Precipitation Index (SPI), and Standardized Precipitation Drought Index (SPDI)). Our results suggest that—irrespective of drought index and tree species—tree-ring width shows a stronger response to interannual variability of drought, compared to the greenness and the NPP. In comparison to other drought indices (e.g., PDSI), and our results demonstrate that multi-scalar drought indices (e.g., SPI, SPEI) are more advantageous in monitoring drought impacts on tree-ring growth, maximum greenness, and NPP. This finding suggests that multi-scalar indices are more appropriate for monitoring and modelling forest drought in peninsular Spain and the Balearic Islands., Ministerio de Economía projects: (CGL2015-69186-C2-1-R (Fundiver)), (CGL2015-69985-R (CLIMED)),( CGL2013-48843-C2-1-R (CoMo-ReAdapt)),( AGL2014-53822-C2-1-R (SATIVA)),( XIRONO (BFU2010-21451)),( CGL2014-52135-C03-01), (PCIN-2015-220), and (CGL2016-81706-REDT (Ecometas Network)), (1560/2015 (ECOHIPRO)), Xunta de Galicia, projects:(PGIDIT06PXIB502262PR, GRC GI-1809, ROCLIGAL-10MDS291009PR), INIA (RTA200600117) e Interreg V-A POCTEFA (CANOPEE, 2014-2020-fondos FEDER), Ministerio de Economía y Empresa, postdoctoral grants (MINECO-FPDI 2013-16600; fondos FEDER), Postdoctoral grants (IJCI-2015-25845)

International audience; Deciphering climatic factors limiting cambial activity is critical to forecast the potential of trees to respond to ongoing climatic change. We explored multiple xylem traits, including tree-ring width, inter-annual micro-density variation and intra-annual density fluctuations (IADF), to unveil the climatic factors constraining cambial activity of a Mediterranean conifer (Pinus pinaster) along a continental-aridity gradient. Secondary growth responded mainly to water availability, explaining as much as 64.7% of earlywood growth variance for earlywood growth. The continuous and non-overlapping timing of the climatic signals of earlywood and late wood growth reflected a continuous water limitation of secondary growth along the growing season. Drought also had an extraordinary impact on minimum (D-min) and maximum (D-max) density, with maximal explained variances reaching 47.4% and 39.1%, respectively. D-min was negatively associated to water availability during the initiation of earlywood formation, whereas D-max responded positively to water availability during two distinct periods: previous winter and the initiation of latewood formation. IADFs in the latewood were quite common along the gradient, occurred in 21-51% of the rings, and responded to episodes of high rainfall and elevated temperature at different phases of latewood formation. Xylem traits identity outperformed site as a driver of climatic signal, revealing the potential of a multi-proxy approach to unveil multiple facets of the xylogenetic cycle. Cambial plasticity, i.e. the ability to adjust the xylogenesis rate and to arrest and resume cambial activity to exploit favorable climatic windows, was critical for Pinus pinaster to thrive within wide climatic envelopes. Nevertheless, the pervasive effect of water availability on all analyzed traits indicates that forthcoming reduced precipitation and increased evapotranspiration, as predicted by climate change models, will negatively impact P. pinaster secondary growth along its whole environmental range.

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.

Non-structural carbohydrates (NSC) play a central role in the construction and maintenance of a tree's vascular system, but feedbacks between the NSC status of trees and wood formation are not fully understood. We aimed to evaluate multiple dependencies among wood anatomy, winter NSC, and phenology for coexisting temperate (Quercus robur) and sub-Mediterranean (Q. pyrenaica) oaks along a water-availability gradient in the NW Iberian Peninsula. Sapwood NSC concentrations were quantified at three sites in December 2012 (N = 240). Leaf phenology and wood anatomy were surveyed in 2013. Structural equation modelling was used to analyse the interplay among hydraulic diameter (Dh), winter NSC, budburst date, and earlywood vessel production (EVP), while the effect of Dh and EVP on latewood width was assessed by using a mixed-effects model. NSC and wood production increased under drier conditions for both species. Q. robur showed a narrower Dh and lower soluble sugar (SS) concentration (3.88–5.08 % dry matter) than Q. pyrenaica (4.06–5.57 % dry matter), but Q. robur exhibited larger EVP and wider latewood (1403 µm) than Q. pyrenaica (667 µm). Stem diameter and Dh had a positive effect on SS concentrations, which were related to an earlier leaf flushing in both species. Sapwood sugar content appeared to limit EVP exclusively in Q. pyrenaica. In turn, Dh and EVP were found to be key predictors of latewood growth. Our results confirm that sapwood SS concentrations are involved in modulating growth resumption and xylem production in spring. Q. pyrenaica exhibited a tighter control of carbohydrate allocation to wood formation than Q. robur, which would play a role in protecting against environmental stress in the sub-Mediterranean area. This work was supported by the Spanish Ministry of Economy and Competitiveness (research projects BFU-21451 and CGL2012-34209) with European Regional Development Fund, and by Xunta de Galicia (research projects 10MDS291009PR and GRC2015/008). Gonzalo Pérez-de-Lis received a PhD FPU-ME grant(grantnumberAP2010-4911)fundedbytheSpanishMinistry of Education. The work was partially inspired within the FPS COST Action FP1106 – STReESS and Ecometas net CGL201453840REDT SI

International audience; Tree rings provide information about the climatic conditions during the growing season by recording them in different anatomical features, such as intra-annual density fluctuations (IADFs). IADFs are intra-annual changes of wood density appearing as latewood-like cells within earlywood, or earlywood-like cells within latewood. The occurrence of IADFs is dependent on the age and size of the tree, and it is triggered by climatic drivers. The variations of IADF frequency of different species and their dependence on climate across a wide geographical range have still to be explored. The objective of this study is to investigate the effect of age, tree-ring width and climate on IADF formation and frequency at a regional scale across the Mediterranean Basin in Pinus halepensis Mill., Pinus pinaster Ait., and Pinus pinea L. The analyzed tree-ring network was composed of P. pinea trees growing at 10 sites (2 in Italy, 4 in Spain, and 4 in Portugal), P. pinaster from 19 sites (2 in Italy, 13 in Spain, and 4 in Portugal), and P. halepensis from 38 sites in Spain. The correlations between IADF frequency and monthly minimum, mean and maximum temperatures, as well as between IADF frequency and total precipitation, were analyzed. A significant negative relationship between IADF frequency and tree-ring age was found for the three Mediterranean pines. Moreover, IADFs were more frequent in wider rings than in narrower ones, although the widest rings showed a reduced IADF frequency. Wet conditions during late summer/early autumn triggered the formation of IADFs in the three species. Our results suggest the existence of a common climatic driver for the formation of IADFs in Mediterranean pines, highlighting the potential use of IADF frequency as a proxy for climate reconstructions with geographical resolution.

Little is known concerning the effects of wildfires on tree radial growth and their climatic response under contrasting regimes of fog water inputs on oceanic islands. On Tenerife, Canary Islands, windward slopes are humid with high-fog frequency due to influence of wet trade winds, while climate on leeward slopes is more arid. We used tree-ring records of Pinus canariensis Sweet ex Spreng. to quantify the effects of a fire of known date on radial growth and determine the main limiting climatic factors for growth. Radial growth patterns and their responsiveness to fire severity and climatic variation differed between windward and leeward slopes. Surface fire did not significantly impact growth, while crown fire caused short-term growth reduction, and even cessation, more pronounced on the windward slope. Growth rates, tree-ring common signal, and climate sensitivity were smaller on the windward slope, with cold winters, and summer water stress limiting growth. On the leeward slope, climate explained a greater amount of growth variation mainly due to negative effects of high October-December sea-level pressures causing dry winter conditions. Contrasting growth dynamics on both slopes may result from diverging physiological effects of water inputs and reduced radiation caused by fog drip. Our findings suggest that dating growth suppressions and absent rings are useful to date past high-severity crown fires in P. canariensis forests, in addition to ordinary fire scars dating indicative of low-severity surface fires. © 2011 Springer-Verlag., This research was partially funded by Consellería de Innovación e Industria, Xunta de Galicia (PGIDIT06PXIB502262PR) and INIA, Spanish Ministry of Science and Innovation (RTA2006-00117).

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., This research was partially supported by the Consejería de Medio Ambiente del Principado de Asturias (SV-PA-00-01).

Long-term patterns of tree establishment and past management practices, in an old-growth pollarded woodland in northern Spain, were studied using dendrochronological techniques. Age distribution revealed three main cohorts of pedunculate oak (Quercus robur L.), Dendrochronologie du chêne pédonculé (Quercus robur L.) dans un vieux bois traité en têtard dans le nord de l'Espagne : modèles d'installation et historique de la gestion. Des modèles à long terme d'installation des arbres et des pratiques de gestion passées ont été étudiées en utilisant les techniques dendrochronologiques, dans un vieux bois traité en têtard du nord de l'Espagne. La distribution des âges révèle trois principales cohortes de chênes pédonculés (Quercus robur L.) concernant l'âge des arbres : < 50 ans, 150-200 ans et 250-500 ans. Les arbres jeunes et les arbres âgés de 200-299 ans étaient en bouquets et spatialement séparés des plus vieux arbres. Les chênes plus âgés que 300 ans montraient une distribution spatiale au hasard. La moyenne des intervalles d'étêtage s'est accrue au cours du temps et la moyenne des suppressions annuelles a décru significativement. La fréquence de l'étêtage et son intensité ont décru depuis 1917 pour les arbres mûrs et depuis 1905 pour les vieux arbres, suggérant un déclin des pratiques d'étêtage au début du 20e siècle. Un retard de 6 ans dans la reprise de croissance a été noté en relation avec la suppression des pousses. Ceci résulte ensemble du rétablissement d'une forte croissance après l'étêtage et de la reprise de croissance dans l'espace de la couronne. Les chênes matures montrent surtout 1-3 signaux d'étêtage, tandis que 4-7 suppressions par arbres étaient plus fréquentes chez les très vieux arbres. En 1774,1792, 1806,1839 et 1859, de petits groupes d'arbres et des arbres dispersés étaient étêtés, tandis qu'en 1905 une réduction générale de la croissance était notée. Les résultats de cette étude ont des connotations significatives conservant la préservation et la gestion des anciens bois traités en têtard.

International audience; The influence of climate and disturbances on tree-ring widths of European beech and pedunculate oak were evaluated in a lowland forest of Northern Spain. From 1925 to 1980, 36% of the variance of beech ring-width and 29% of the oak one was explained by climate. The climatic models showed that low precipitation in July of the previous year limited the radial growth of beech, while oak one was instead restricted by water deficits in July of the current year. Ten main disturbance periods were identified from 1780 to 1997, among which the 1922-1935 one was the most important. Since beech trees showed suppressed growth from 1800 to 1920, probably the forest canopy became denser during this time. The disturbance periods identified in 1922-1935 and 1948-1953 contributed to both increase the growth of beech above the expected, and intensify its climatic response. On the other hand, deviations of oak growth from the expected without-disturbance indices agreed with the disturbance history up to 1850. From 1850 to 1997, oak growth became independent from disturbances sequence, yielding a constant climatic response in 1925-1980. The opposite effects of disturbances on both the radial growth and the climatic response of European beech and pedunculate oak are related to their different tolerance to shade. These results have relevant methodological implications on the analysis of climate-growth relationships, and on the reconstruction of past disturbance regimes by means of dendroecological techniques.; Effet du climat et des perturbations locales sur la croissance radiale de Fagus sylvatica L. et Quercus robur L. dans une forêt naturelle de Cantabria, Nord de l'Espagne. L'influence relative du climat et des perturbations locales sur la croissance radiale du hêtre et du chêne pédonculé a été analysée dans une vieille forêt naturelle du Nord de l'Espagne. Entre 1925 et 1980, 36 % de la variance des largeurs de cernes du hêtre et 29 % de celle du chêne s'expliquent par le climat. Les modèles climatiques élaborés montrent que la croissance radiale du hêtre est limitée par les précipitations du mois de juillet de l'année précédente, alors que celle du chêne l'est par le déficit hydrique du mois de juillet de l'année en cours. Dix périodes de perturbation de la croissance, d'origine non climatique, ont été identifiées entre 1780 et 1997, parmi lesquelles celle de 1922-1935 a été la plus importante. La croissance radiale des hêtres apparaît faible de 1800 à 1920 en raison de la fermeture du couvert forestier au cours de cette période. Puis des perturbations survenues en 1922-1935 et 1948-1953 entraînent une augmentation de la croissance, qui devient alors supérieure au signal commun. Conjointement, la réponse aux contraintes climatiques se renforce au cours des mêmes périodes. Chez le chêne, les déviations de la croissance par rapport au signal commun sont en accord avec l'historique des perturbations locales jusqu'en 1850. Puis la croissance devient indépendante de ces perturbations et converge avec le signal commun. Sa réponse au climat demeure constante de 1925 à 1980.