Vicente Rozas

Quantitative Tracheid Anatomy Reveals a Complex Environmental Control of Wood Structure in Continental Mediterranean Climate

A detailed understanding of how the climate models the hydraulic system of trees is still lacking, in spite of the need to understand tree response to the ongoing process of climatic change. A systematic correlation of tracheid anatomical features with climate has not been implemented in Mediterranean ecosystems, where climatic change is expected to be particularly intense. We measured ring width and cell number—in addition to intraring position, lumen size, and wall thickness of tracheids—in 10 Juniperus thurifera individuals from north-central Spain. We used this information to perform an exploratory analysis of how these parameters correlated with climatic variables in 1965–2004. Cell number and ring width shared a relatively similar climatic signal, whereas the anatomical variables provided differentiated and diverse signals about climatic conditions during their formation. Earlywood and latewood tracheids differed in controlling factors, with earlywood tracheid size and wall thickness being positively related to rainfall during early summer and latewood tracheid size being positively related to August temperature. Tracheid anatomical variables improved our understanding of climate effects on tree growth and wood formation under harsh environmental conditions, as those experienced in continental Mediterranean climates, where limiting factors show multiple shifts across the year.

New star on the stage: amount of ray parenchyma in tree rings shows a link to climate

Tree-ring anatomy reflects the year-by-year impact of environmental factors on tree growth. Up to now, research in this field has mainly focused on the hydraulic architecture, with ray parenchyma neglected despite the growing recognition of its relevance for xylem function. Our aim was to address this gap by exploring the potential of the annual patterns of xylem parenchyma as a climate proxy. We constructed ring-width and ray-parenchyma chronologies from 1965 to 2004 for 20 Juniperus thurifera trees growing in a Mediterranean continental climate. Chronologies were related to climate records by means of correlation, multiple regression and partial correlation analyses. Ray parenchyma responded to climatic conditions at critical stages during the xylogenetic process; namely, at the end of the previous years xylogenesis (October) and at the onset of earlywood (May) and latewood formation (August). Ray parenchyma-based chronologies have potential to complement ring-width chronologies as a tool for climate reconstructions. Furthermore, medium- and low-frequency signals in the variation of ray parenchyma may improve our understanding of how trees respond to environmental fluctuations and to global change.

Climatic control of intra-annual wood density fluctuations of Pinus pinaster in NW Spain

Tree rings of Pinus pinaster often contain intra-annual density fluctuations (IADFs), which have been attributed to the succession of dry and rainy periods typical of Mediterranean climate, but their formation has not been studied yet under Atlantic climate. We analyzed the occurrence and climatic significance of replicated IADFs in ten monospecific stands in NW Spain. The frequency of IADFs was higher than previously reported for this species under Mediterranean conditions and consistently decreased with increasing elevation. The formation of bands of latewood-like tracheids within the earlywood was favored by dry previous August, cold previous winter and dry April. Bands of earlywood-like tracheids within the early latewood were also favored by low winter temperatures. However, their occurrence was geographically heterogeneous, with two groups of stands being defined by their distances to the shoreline. In coastal stands, cold May–August triggered IADFs formation, while in inland stands their formation was favored by dry May–July. Regional winter temperatures and April water balance were strongly related to the East Atlantic (EA) pattern, which greatly conditioned the occurrence of IADFs in the earlywood and the early latewood. By contrast, the presence of bands of earlywood-like tracheids in the late latewood was independent of the EA pattern, being strongly related to warm conditions in spring and especially to a wet October. The link between regional climate and the EA pattern strongly controlled the physiological processes that determine intra-annual growth dynamics and short-term cell enlargement of P. pinaster in NW Spain.

Earlywood vessels of the sub-Mediterranean oak Quercus pyrenaica have greater plasticity and sensitivity than those of the temperate Q. petraea at the Atlantic–Mediterranean boundary

Key messageEarlywood vessel features indicate different adaptations ofQuercus petraeaandQ. pyrenaica, which are probably related with their corresponding Atlantic and sub-Mediterranean ecological requirements.AbstractWe studied the climatic signal of the earlywood anatomy of a temperate [Quercus petraea (Mattuschka) Liebl.] and a sub-Mediterranean (Quercus pyrenaica Willd.) oak species growing under similar climatic conditions in a transitional area between the Atlantic and Mediterranean regions of the Iberian Peninsula. We hypothesized that both species react differently in their wood anatomy due to their contrasting ecological requirements, and we test the usefulness of earlywood anatomical features to study the behaviour of these ring-porous oaks upon climate. For this, we measured the earlywood vessels, and obtained annual series of several anatomical variables for the period 1937–2006 using dendrochronological techniques, considering whether the vessels belonged to the first row or not. After optimizing the data set by principal component analysis and progressive filtering of large vessels, we selected maximum vessel area and total number of vessels as they resulted to be the optimal variables to describe vessel size and number, respectively. Vessel size of Q. pyrenaica was dependent on precipitation along the previous growing season, whereas it did not show any clear climatic response for Q. petraea. On the contrary, vessel number was related to winter temperature for both species. These relationships observed between climate and anatomy appeared to be stable through time. The results obtained reinforce the utility of earlywood vessel features as potential climate proxies.

Drought-induced increase in water-use efficiency reduces secondary tree growth and tracheid wall thickness in a Mediterranean conifer

In order to understand the impact of drought and intrinsic water-use efficiency (iWUE) on tree growth, we evaluated the relative importance of direct and indirect effects of water availability on secondary growth and xylem anatomy of Juniperus thurifera, a Mediterranean anisohydric conifer. Dendrochronological techniques, quantitative xylem anatomy, and 13C/12C isotopic ratio were combined to develop standardized chronologies for iWUE, BAI (basal area increment), and anatomical variables on a 40-year-long annually resolved series for 20 trees. We tested the relationship between iWUE and secondary growth at short-term (annual) and long-term (decadal) temporal scales to evaluate whether gains in iWUE may lead to increases in secondary growth. We obtained a positive long-term correlation between iWUE and BAI, simultaneously with a negative short-term correlation between them. Furthermore, BAI and iWUE were correlated with anatomical traits related to carbon sink or storage (tracheid wall thickness and ray parenchyma amount), but no significant correlation with conductive traits (tracheid lumen) was found. Water availability during the growing season significantly modulated tree growth at the xylem level, where growth rates and wood anatomical traits were affected by June precipitation. Our results are consistent with a drought-induced limitation of tree growth response to rising CO2, despite the trend of rising iWUE being maintained. We also remark the usefulness of exploring this relationship at different temporal scales to fully understand the actual links between iWUE and secondary growth dynamics.

Large-scale structural variation and long-term growth dynamics of Juniperus thurifera trees in a managed woodland in Soria, central Spain

Abstract• Juniper woodlands have a great conservational and productive importance in Spain. However, basic traits of their structural variation and growth dynamics are unknown.• To characterize the structural variation and assess both height and radial growth patterns, we sampled and measured stem cross-sections every 0.5 m in height from 107 Spanish juniper (Juniperus thurifera L.) trees, which were selected based on a 350 m regular grid over a whole woodland of 3 300 ha in area. We used dendroecological techniques and spatial analysis to study tree structure and growth.• Structural traits such as bole diameter and height were mainly determined by tree life-related parameters such as tree age and growth rate variation, while abiotic factors such as altitude played a minor role in determining structural variation. Over the last 300 years, juniper establishment has been continuous in time but discontinuous in space. Large-scale spatial heterogeneity of tree establishment, and the presence of an early growth suppression and abrupt growth changes in junipers > 100 years old are consequences of an intense management that almost ceased in the late 19th century.• Abrupt growth releases and suppressions were synchronic among different age classes, and coincided with documentary records on livestock decrease and key historical changes in land use. Growth patterns suggest that juniper is a brow sing-tolerant species that is able to survive large periods of intense browsing pressure.Résumé• Les forêts de genévrier ont en Espagne une grande importance à la fois du point de vue de la conservation et de la production. Cependant, des caractéristiques de base de leur variation structurelle et leur dynamique de croissance sont inconnues.• Pour caractériser la variation structurelle et évaluer à la fois la hauteur et les modèles de croissance radiale, nous avons échantillonné et mesuré les sections de troncs tous les 0,5 m en hauteur chez 107 genévriers (Juniperus thurifera L.), qui ont été choisis sur la base d’un réseau régulier de 350 m de côté dans la totalité d’un bois d’une surface de 3 300 ha. Nous avons utilisé les techniques dendroécologiques et l’analyse spatiale pour étudier la structure des arbres et la croissance.• Des caractéristiques structurales comme le diamètre de tronc et la hauteur ont été principalement déterminées par des paramètres concernant la vie de l’arbre comme l’âge et la variation du taux de croissance, tandis que des facteurs abiotiques comme l’altitude ont joué un rôle secondaire dans la détermination de la variation structurelle. Pendant les 300 dernières années, l’établissement du genévrier était continu dans le temps, mais discontinu dans l’espace. L’hétérogénéité spatiale à grande échelle de l’établissement des arbres, la présence d’une première suppression de croissance et des changements brusques de croissance chez des genévriers âgés de plus de 100 ans sont les conséquences d’une gestion intensive qui a presque cessé à la fin du 19e siècle.• Des reprises brusques de croissance et des suppressions étaient synchrones parmi différentes classes d’âge et ont coïncidé avec des rapports documentaires sur la diminution du bétail, clefs des changements historiques dans l’utilisation de la terre. Les modèles de croissance suggèrent que le genévrier est une espèce tolérante à l’abroutissement et qui est capable de survivre à de grandes périodes d’abroutissement intense.

Climatic Signals from Intra-annual Density Fluctuation Frequency in Mediterranean Pines at a Regional Scale

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.

Effects of thinning intensity on radial growth patterns and temperature sensitivity in Pinus canariensis afforestations on Tenerife Island, Spain

Abstract• ContextThe suitability of thinning to prevent forest growth decline as a result of global warming has not been tested extensively in Macaronesian Canary pine (Pinus canariensis Sweet ex Spreng.). • AimsThis study aimed to answer the following questions: (1) are tree growth patterns modified by the aspect and thinning intensity? (2) Is sensitivity to climate modulated by thinning? (3) Do trees of different crown classes respond differentially to thinning intensity and climate? • MethodsWe used tree-ring series from dominant, codominant, and overtopped trees to study the effects of thinning intensity on basal area increments (BAI) and climate sensitivity on windward (wet) and leeward (dry) slopes on Tenerife, Canary Islands. Three replicated blocks of control, light thinning, and heavy thinning stands were set on each slope in 1988, and cores were extracted in 2007. • ResultsHeavy thinning induced growth release and increased BAI, mainly on dominant and codominant trees, whereas light thinning effects were negligible; their impacts were more intense on the windward side. Temperature sensitivity was hardly affected by thinning on leeward, where climate control was stronger. On windward, thinning enhanced the influence of summer temperatures. Upper crown classes were more sensitive overall, but overtopped trees responded better in summer. Thinning intensity and aspect greatly influence growth on Canary pine afforestations, but individual responses are highly dependent on crown classes. In addition, thinning may be less effective to modify growth conditions on leeward slopes, at least if not intense.

Contrasting effects of wildfire and climate on radial growth of Pinus canariensis on windward and leeward slopes on Tenerife, Canary Islands

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.

Non-stationary influence of El Niño-Southern Oscillation and winter temperature on oak latewood growth in NW Iberian Peninsula.

The properties of El Niño-Southern Oscillation (ENSO), such as period, amplitude, and teleconnection strength to extratropical regions, have changed since the mid-1970s. ENSO affects the regional climatic regime in SW Europe, thus tree performance in the Iberian Peninsula could be affected by recent ENSO dynamics. We established four Quercus robur chronologies of earlywood and latewood widths in the NW Iberian Peninsula. The relationship between tree growth and the Southern Oscillation Index (SOI), the atmospheric expression of ENSO, showed that only latewood growth was correlated negatively with the SOI of the previous summer–autumn–winter. This relationship was non-stationary, with significant correlations only during the period 1952–1980; and also non-linear, with enhanced latewood growth only in La Niña years, i.e. years with a negative SOI index for the previous autumn. Non-linear relationship between latewood and SOI indicates an asymmetric influence of ENSO on tree performance, biassed towards negative SOI phases. During La Niña years, climate in the study area was warmer and wetter than during positive years, but only for 1952–1980. Winter temperatures became the most limiting factor for latewood growth since 1980, when mean regional temperatures increased by 1°C in comparison to previous periods. As a result, higher winter respiration rates, and the extension of the growing season, would probably cause an additional consumption of stored carbohydrates. The influence of ENSO and winter temperatures proved to be of great importance for tree growth, even at lower altitudes and under mild Atlantic climate in the NW Iberian Peninsula.