Seasonal cambium activity in the subtropical rain forest tree Araucaria angustifolia

Information on the timing and dynamics of tree ring formation is essential to assess the seasonal behavior of secondary wood growth and its associated environmental influences. Araucaria angustifolia is a dominant species in highland pluvial ecosystems of southeastern South America. Previous investigations indicated that their growth rings are formed annually, but no information exists about the timing of growth ring formation and the environmental triggers influencing cambium activity. In this paper we examine inter- and intra-annual cambial activity in A. angustifolia, through anatomical and dendrochronological evidence at two study sites, and model the relationships between regional climate variation and intra-annual tree ring formation. The results confirm the annual growth ring formation in A. angustifolia and indicate that its growth season extends from October to April. Day length and temperature were the main environmental factors influencing the seasonal cambium activity. Our results evidence the dendrochronological potential of A. angustifolia for ecological and climatological studies in southeastern South America.     

Microstructure alignment of wood density profiles: an approach to equalize radial differences in growth rate

We studied intra-annual wood density profiles of Douglas-fir tree rings (Pseudotsuga menziesii [Mirb.] Franco) in southwestern Germany. Growth rates are variable around the tree circumference. This leads to differences in wood formation, which can be observed in the shape of the density profiles of the same tree ring measured in different radial directions. Due to this spatial variation in density profiles, we need a reliable method to determine an average profile, which preserves the common characteristics of the data. To this end, we developed a multiple interval-based curve alignment (MICA) procedure. It identifies characteristic points within the profiles such as minima, maxima and inflection points. These reference points are shifted gradually against each other within a proportionally defined baseline interval. Using our progressive alignment approach, we are able to calculate an average profile that represents very well the characteristics of all measured curves of a specific tree ring. We applied the procedure to get year-specific average profiles using various trees. This results in representative mean density profiles that preserve the density variations common to all aligned profiles. Individual noise is reduced, thereby enabling the analysis of the impact of weather variations on wood density.     

Plastic Response of Tracheids in Pinus pinaster in a Water-Limited Environment: Adjusting Lumen Size instead of Wall Thickness

The formation of wood results from cambial activity and its anatomical properties reflect the variability of environmental conditions during the growing season. Recently, it was found that wood density variations in conifers growing under cold-limited environment result from the adjustment of cell wall thickness (CWT) to temperature. Additionally, it is known that intra-annual density fluctuations (IADFs) are formed in response to precipitation after the summer drought. Although IADFs are frequent in Mediterranean conifers no study has yet been conducted to determine if these structures result from the adjustment of lumen diameter (LD) or CWT to soil water availability. Our main objective is to investigate the intra-ring variation of wood anatomical features (LD and CWT) in Pinus pinaster Ait. growing under a water-limited environment. We compared the tracheidograms of LD and CWT for the years 2010–2013 in P. pinaster growing in the west coast of Portugal. Our results suggest a close association between LD and soil moisture content along the growing season, reinforcing the role of water availability in determining tracheid size. Compared with CWT, LD showed a higher intra- and inter-annual variability suggesting its strong adjustment value to variations in water availability. The formation of a latewood IADF appears to be predisposed by higher rates of cell production in spring and triggered by early autumn precipitation. Our findings reinforce the crucial role of water availability on cambial activity and wood formation in Mediterranean conifers, and emphasize the high plasticity of wood anatomical features under Mediterranean climate.     

Precision of dating insect outbreaks using wood anatomy: the case of Anoplophora in Japanese maple

To control invasive Anoplophora outbreaks, it is crucial to accurately date infestation dynamics. Dating of Anoplophora outbreaks is possible as these xylobiont insects induce wounds in living trees by forming e.g., exit holes. This study investigates to what precision these wounds can be dated with dendrochronological techniques. In an experimental setting, we studied the precision of wound dating on Acer palmatum, an ornamental tree occasionally containing larvae of A. chinensis. We studied the development of wound reactions at the beginning, during and after the growing season, both in relation to leaf phenology and intra-annual tree-ring growth. We found that the precision of dating exit holes is limited due to the highly variable intra-annual tree-ring growth whereby only an accurate distinction can be made between wounds originated during, or after tree-ring formation. The resolution was improved using local growth—the local reactivation of xylem growth around the wound—as a marker for wounds that occurred at the end of the growing season. We conclude that the intra-annual precision of dating Anoplophora outbreaks in Acer palmatum in the temperate North-western European climate is limited to three distinct phases: (i) The period of dormancy and leaf emergence (ca. October until April/beginning of May), when the wounds are located at the tree-ring boundary (ii) The period of tree-ring growth in which wounds are located within the tree ring (ca. end of April/beginning of May until late August/beginning of September), (iii) end of growing season (ca. end of August/September) in which local growth occurs.     

The potential of intra-annual density information for crossdating of short tree-ring series

The crossdating of tree-ring series is typically based on tree-ring width sequences, which is a crude abstraction of the growth signal stored in tree rings. In contrast, intra-annual wood density data allows a much more detailed comparison of wood growth processes and new measurement techniques scale well to measure large amounts of samples. Thus, chronologies of intra-annual densitometric curves can be built. Here, we investigate to what extent intra-annual wood density information can improve crossdating. We evaluate different approaches on a data set of Norway spruce trees (Picea abies) and compare the results to standard methods that are based on ring width or maximum density. Our results show that intra-annual densitometric data indeed increases crossdating success rate notably for short tree ring series that cover less than 25 years.     

Climatic control of intra- and inter-annual wood-formation dynamics of Scots pine in northern Finland

Boreal forests are highly sensitive to climate and human impacts and therefore suitable as biological indicator for environmental changes. In this context, our study was aimed at getting deeper insight into the climate-dependence of the onset, intensity and end of wood formation of Scots pine during the growing season.We monitored the intra-annual growth dynamics of, on average, 42-year-old Scots pine trees over five consecutive years, 2000-2004, at two sites located 80 and 300 km south of the tree line in northern Finland. For that purpose, the cambium of the trees was weekly wounded with a pin and the resulting wound tissue, microscopically detectable in transverse thin-sections through the newly built wood, was taken as a time marker. During this 5-year study period, the intra-annual wood formation at the southern site was mainly positively associated with summer temperature. However, at the northern site such an association was either entirely missing or negative. At both sites, two thirds of the radial growth was produced within only 4 weeks from mid-June to mid-July, independent of whether the growing season started earlier or later.Moreover, we measured the widths of all tree rings from bark to pith (inter-annual growth) of the same study trees and assembled them to 51-year long tree-ring site chronologies. Since 1999, these two site chronologies - after having run fairly parallel over the preceding decades - were running in divergent directions thus corroborating our results derived from the intra-annual climate/growth analysis. Whereas the chronology of the southern site follows the average temperature of May and July very closely from 1961 up to 2004, the chronology of the northern site follows the July temperature, but only up to 1998, and from 1999 to 2004 is running just opposite to the distinctly rising July temperature. During the same period, there was - unlike in the years before - nearly no snow cover in May at the northern site, whereas at the southern site there was no change of the normally existing slight snow cover in May. This deviating weather situation may have led to a temperature-induced, temporary drought stress for the Scots pines at the northern site.     

High-resolution climatic analysis of wood anatomical features in Corsican pine from Corsica (France) using latewood tracheid profiles

Key message

We propose a new methodology to identify intra-annual density fluctuations in latewood using cell features and relative radial position within the latewood of pine trees growing on Corsica, France. Climatic forcing of latewood wood anatomical anomalies was analyzed.

Abstract

We analyzed latewood anatomical features from Corsican pine (Pinus nigra ssp. laricio) of high-elevation sites in Corsica (France) derived from digital images of the wood surface. Latewood of each ring during the period 1950–2008 was partitioned into ten equal parts P1–P10. Mean values of the cell parameters cell lumen area (CLA), radial cell width (RCW), radial cell wall thickness (CWT), and modeled latewood density (MLD) were calculated for P1–P10. The cellular profiles for each cell parameter were subjected to principal component analyses. It was possible to quantify macroscopically visible variations of wood anatomy like intra-annual density fluctuations (IADFs) by latewood profiles of different cell parameters. A combination of cell parameter characteristics including their relative radial position within latewood provides a quantification of the cell anatomical variations in an IADF. Individual cell parameter chronologies and principal components of cell parameter profiles were correlated with climate data to determine the climatic forcing on latewood formation. Average cell parameter profiles and deviations from the long-term means are able to describe “normal” and “anomalous” environmental conditions during latewood formation. Cell feature anomalies throughout the latewood during individual years allow the reconstruction of past weather conditions with a high temporal resolution.     

Cambial activity of Pinus elliottii var. densa reveals influence of seasonal insolation on growth dynamics in the Florida Keys

We determined the temporal and seasonal dynamics of intra-annual cell formation of south Florida slash pine (Pinus elliottii Engelm. var. densa Little & Dor.), the southernmost native pine in the United States and the foundation species of globally endangered pine rockland ecosystems. To assess intra-annual cambial activity and identify possible relationships between cell production and climatic factors, wood micro-cores were extracted monthly from six trees during the period March 2010 to March 2011. The results confirmed annual growth ring formation in P. elliottii var. densa and indicated that its growing season extends from February to December, with a short period of dormancy that varied little between individuals. Within the growing season, earlywood cells were produced from February to July, latewood cells produced from July to December, and intra-annual density fluctuations (IADFs) occurred in the growth rings of four of six trees between July and August. A principal component analysis indicated a homogeneous response of cambial activity among trees to site-specific climatic factors. The first principal component axis explained 71?% of the total variance in cell production during the study period. Our results indicated that the dynamics of seasonal cambial activity of P. elliottii var. densa are controlled by solar radiation (r?=?0.51, p?<?0.10) in the Florida Keys. The nature of our data allow us to only speculate on the ecophysiological processes responsible for IADFs in P. elliottii var. densa, and additional research is needed to better understand the relationship between their formation and the environment in the Lower Florida Keys.     

The number of days on which increment occurs is the primary determinant of annual ring width in Callitris intratropica

Key message

The number of days on which a measureable increment occurred, and the average rate of stem growth, rather than the overall duration of the wet season, were the main determinants of ring width in young Callitris intratropica trees. These effects were amplified by competition.

Abstract

Dendroclimatology of tropical tree species is an important tool for understanding past climatic variability at low latitudes where long-term weather records are often absent. Despite the growing number of published tropical tree-ring chronologies, however, still little is known of the factors that control annual ring formation in tropical tree species. In this paper we used an endemic Australian conifer, Callitris intratropica, to study the intra-annual dynamics of seasonal growth and xylem formation, and the effects of environmental conditions and competition, on growth ring formation. We combined high-resolution growth and climate monitoring (every 15 min for 2 years) with less frequent cambial sampling. Trees exhibited marked reductions in growth during certain periods within the rainy season when rainfall was not as regular and VPD was high. Overall, we found that ring width was most influenced by the number of days when increment occurred; regardless of how early the growing season began or ended, and by the rates of tracheid production. The effect of competition was also important. Trees growing in dense groves had narrower annual rings (4.6 mm) than trees that were growing in the open (6.7 mm), due to less active cambia, slower rates of xylem production and expansion and more increment days, although the overall growing season duration was also shorter in grove trees.     

Discrete versus continuous analysis of anatomical and δ13C variability in tree rings with intra-annual density fluctuations

Intra-annual density fluctuations (IADFs) are anomalies of tree rings where wood density is abruptly altered after sudden changes in environmental conditions. Their characterisation can provide information about the relationship between environmental factors and eco-physiological processes during tree growth. This paper reports about the variability of anatomical traits and stable carbon isotopic composition along tree rings as resulting from the application of two different methodological approaches: (a) the separation of each ring into different regions (earlywood, latewood and IADF) and the comparison of anatomical and isotopic parameters measured in those specific sectors and (b) the analysis of such features in continuum along ring width. Moreover, different parameters of vessels (i.e. ecd—equivalent circle diameter, elongation, sphericity and convexity of vessel lumen) were considered to identify those more appropriate for the representation of intra-annual anatomical variations. The analysis was conducted on Arbutus unedo L. growing on the Elba Island (Italy); tree rings of this species form IADFs with features clearly responsive to the environmental conditions experienced during plant growth. Results showed that the first approach, although more suitable to obtain data for subsequent statistical comparisons and for the calculation of correlations with environmental parameters, suffers from elements of subjectivity due to the size and position of the selected tree-ring regions. The in continuum method allows a clearer identification of the variation of tree-ring properties along ring width. Regarding anatomical parameters, shape indexes were not suitable indicators of intra-annual variability. The overall analysis suggested that using both methodologies in synergy helps to gain complete information and avoid misleading interpretations of IADFs in tree rings.     

Adjustment Capacity of Maritime Pine Cambial Activity in Drought-Prone Environments

Intra-annual density fluctuations (IADFs) are anatomical features formed in response to changes in the environmental conditions within the growing season. These anatomical features are commonly observed in Mediterranean pines, being more frequent in younger and wider tree rings. However, the process behind IADF formation is still unknown. Weekly monitoring of cambial activity and wood formation would fill this void. Although studies describing cambial activity and wood formation have become frequent, this knowledge is still fragmentary in the Mediterranean region. Here we present data from the monitoring of cambial activity and wood formation in two diameter classes of maritime pine (Pinus pinaster Ait.), over two years, in order to test: (i) whether the differences in stem diameter in an even-aged stand were due to timings and/or rates of xylogenesis; (ii) if IADFs were more common in large trees; and (iii) if their formation is triggered by cambial resumption after the summer drought. Larger trees showed higher rates of cell production and longer growing seasons, due to an earlier start and later end of xylogenesis. When a drier winter occurs, larger trees were more affected, probably limiting xylogenesis in the summer months. In both diameter classes a latewood IADF was formed in 2012 in response to late-September precipitation, confirming that the timing of the precipitation event after the summer drought is crucial in determining the resumption of cambial activity and whether or not an IADF is formed. It was the first time that the formation of a latewood IADF was monitored at a weekly time scale in maritime pine. The capacity of maritime pine to adjust cambial activity to the current environmental conditions represents a valuable strategy under the future climate change conditions.     

The impact of the 2003 summer drought on the intra-annual growth pattern of beech (Fagus sylvatica L.) and oak (Quercus robur L.) on a dry site in the Netherlands

Climate change is expected to result in more extreme weather conditions over large parts of Europe, such as the prolonged drought of 2003. As water supply is critical for tree growth on many sites in North-Western Europe, such droughts will affect growth, species competition, and forest dynamics. To be able to assess the susceptibility of tree species to climate change, it is necessary to understand growth responses to climate, at a high temporal resolution. We therefore studied the intra-annual growth dynamics of three beech trees (Fagus sylvatica L.) and five oak trees (Quercus robur L.) growing on a sandy site in the east of the Netherlands for 2 years: 2003 (oak and beech) and 2004 (oak). Microcores were taken at 2-week intervals from the end of April until the end of October. Intra-annual tree-ring formation was compared with prior and contemporary records of precipitation and temperature from a nearby weather station.The results indicate that oak and beech reacted differently to the summer drought in 2003. During the drought, wood formation in both species ceased, but in beech, it recovered after the drought. The causes of species-specific differences in intra-annual wood formation are discussed in the context of susceptibility to drought.     

Growth dynamics and potential for cross-dating and multi-century climate reconstruction of Podocarpus falcatus in Ethiopia

Podocarpus falcatus is an indigenous evergreen conifer species of tropical mountain forests in southeastern Ethiopia, showing potential tree ages of around 500 years. To study the influence of seasonal climate on the growth pattern of P. falcatus, we combined high-resolution electronic dendrometer measurements with wood anatomical investigations of microcores from the outermost stem parts collected in monthly intervals. At any time of the year sufficient rain events are able to cause cambial activity in P. falcatus. This permanent growing readiness leads to irregular wood formation with the formation of intra-annual density fluctuations and missing rings. Wood anatomical studies of microcores collected around the circumference of a mature P. falcatus revealed locally different activity status of the cambium on different lobes of the stem. Tree-ring width measurements of stem disks resulted in tentative tree ages that were confirmed by radiocarbon dating of selected wood samples. Although our efforts to cross-date ring-width series from several stem disks were not successful, further sampling in areas with different rainfall regimes, additional radiocarbon dating and measurements of stable isotopes hopefully would enable the establishment of a multi-century-long tree-ring series for climate reconstruction.     

A phenology-based approach to the analysis of conifers intra-annual xylem anatomy in water-limited environments

Xylem structure and cambial phenology (i.e. onset and cessation of cambial cell division) of conifers growing under severe water-limitations can change dramatically in relation to moisture availability. In hyperarid environments, analytical tools commonly used to investigate intra-annual variability of xylem anatomy (i.e. tracheidograms), may fail to capture the complexity of tree phenological responses to environmental conditions. This greatly limits our ability to accurately date the onset of intra-annual density variations, including the transition between earlywood and latewood. I present a new approach for developing phenological tracheidograms (“pheno-tracheidograms”) calibrated to account for the seasonal variations in cell division rates. Pheno-tracheidograms were developed for a population of Pinus ponderosa in the Mojave Desert (Nevada, USA) during the period 2015–2016 in order 1) to determine the onset date of latewood formation and 2) to investigate relationships between environmental conditions, lumen diameter, and cell wall thickness targeting specific climatic windows for each tracheid. Pheno-tracheidograms were standardized at the tree-level, showing more flexibility compared to tracheidograms standardized according to a pre-determined number of cells. By displaying cellular parameters with respect to the date of formation of the tracheid to which each measurement is associated, pheno-tracheidograms allowed to determine the beginning of latewood formation with daily resolution. Lumen diameter was significantly correlated with the onset date of cellular enlargement, while cell wall thickness showed a weaker relationship with the beginning of secondary wall deposition. Soil moisture positively affected the duration of cell enlargement and tracheid lumen diameter, particularly in the earlywood, while cell wall thickness was not significantly influenced by environmental conditions. Pheno-tracheidograms represent an empirical, yet effective way to date intra-annual xylem structures and to investigate high-resolution climate-anatomy relationships in conifer species from arid environments characterized by high phenological plasticity.     

Comparison of methods for the demarcation between earlywood and latewood in tree rings of Norway spruce

The precise demarcation between earlywood and latewood is important for the detailed analysis of intra-annual tree ring features. Different techniques based on visual assessment, wood anatomy analysis and X-ray densitometry have been developed and are currently used for this purpose. Depending on the chosen method, tree species and environmental conditions, the results can significantly vary. Thus, it is important to determine the technique optimal for a particular research. Here, we investigated Norway spruce (Picea abies) tree rings to examine the agreement among the following demarcation methods: (1) direct visual assessment, (2) Mork’s index (anatomical definition of the transition from earlywood to latewood based on cell wall-lumen ratio) and (3) fixed and floating density thresholds applied to intra-ring density profiles. The aim was to modify both the Mork’s criterion and density thresholds on the basis of reference values given by visual identification of earlywood/latewood transition. A total of 231 tree rings were analysed by all methods. Our results showed that the usage of floating threshold (defined for each ring separately based on density profiles) is more reliable in comparison with fixed threshold (the same threshold value used for all tree rings and samples). Statistical analysis revealed the best correspondence between visual identification of earlywood/latewood transition and demarcation based on the standard Mork’s index and the floating density threshold derived as 80 % of maximum latewood density. In terms of Mork’s index calibration, the results showed that to determine latewood cells in Norway spruce trees growing in temperate conditions, it is sufficient to use an index value equal to 0.83. The results are applicable for the studied spruce population growing in a temperate climate. The methodology itself, however, is universal and can help to calibrate criteria for earlywood-latewood demarcation under specific conditions.     

Cambial activity and intra-annual xylem formation in roots and stems of Abies balsamea and Picea mariana

Background and Aims

Studies on xylogenesis focus essentially on the stem, whereas there is basically no information about the intra-annual growth of other parts of the tree. As roots strongly influence carbon allocation and tree development, knowledge of the dynamics of xylem production and maturation in roots at a short time scale is required for a better understanding of the phenomenon of tree growth. This study compared cambial activity and xylem formation in stem and roots in two conifers of the boreal forest in Canada.

Methods

Wood microcores were collected weekly in stem and roots of ten Abies balsamea and ten Picea mariana during the 2004–2006 growing seasons. Cross-sections were cut using a rotary microtome, stained with cresyl violet acetate and observed under visible and polarized light. The number of cells in the cambial zone and in differentiation, plus the number of mature cells, was counted along the developing xylem.

Key Results

Xylem formation lasted from the end of May to the end of September, with no difference between stem and roots in 2004–2005. On the contrary, in 2006 a 1-week earlier beginning of cell differentiation was observed in the stem, with cell wall thickening and lignification in roots ending up to 22 d later than in the stem. Cell production in the stem was concentrated early in the season, in June, while most cell divisions in roots occurred 1 month later.

Conclusions

The intra-annual dynamics of growth observed in stem and roots could be related to the different amount of cells produced by the cambium and the patterns of air and soil temperature occurring in spring.Key words: Abies balsamea, boreal forest, cambium, cell differentiation, cell wall thickening, lignification, Picea mariana, root, stem, xylem     

樟子松树轮δ13C的年内变化特征及其对气候要素的响应

对大兴安岭北部两株樟子松(Pinus sylvestris var.mongolica)树轮样品的年内稳定碳同位素比率(δ13C)进行测定,结果表明:樟子松树轮年内δ13C值在不同生长阶段总体表现出每年生长季中期最高、早期次之、晚期最低的变化特征.δ13C的年内变化趋势在幼龄期至速生期变化剧烈,成熟期至衰老期相对平缓.从幼龄期至衰老期的整个生长阶段,同时期年内δ13C的变动幅度基本为晚材大于早材.幼龄期年内晚材的δ13C一直明显高于早材,而成熟期年内早晚材δ13C的差别逐渐减小,至衰老期年内晚材δ13C已低于早材且无显著差别.树轮δ13C的年内变化主要体现在生长季中后期,即早晚材之间的过渡段至晚材.年内不同时段的d13C序列与同时段的宽度去除生长趋势序列(去趋势序列)之间的相关性随生长季节的推移而逐渐降低.当年早材宽度与前一年晚材宽度显著正相关,当年早材δ13C序列与前一年晚材宽度和当年早材宽度的去趋势合并序列呈现较显著的负相关性,与前一年晚材δ13C序列或宽度去趋势序列之间均未表现出显著的相关性.分析结果表明:早材的形成很可能来源于前一年光合作用的产物,在利用树轮年内不同材质宽度或δ13C序列进行气候环境重建时需要考虑这一点.年内早材、过渡段和晚材三个时段的δ13C分别对应于4月下旬至6月中旬土壤湿度较大、温度上升较快的时期,6月下旬至7月中旬降水增加、温度达到最高而相对湿度降低的时期,以及7月下旬至9月中旬降水增加、温度下降而相对湿度较大的时期.     

Revealing Polylepis microphylla as a suitable tree species for dendrochronology and quantitative wood anatomy in the Andean montane forests

In the tropical Andes climate change is expected to increase temperatures and change precipitation patterns. To overcome the lack of systematic weather records that limits the performance of climate models in this region, the use of the environmental information contained in tree rings from tropical Andean species have been found useful to reconstruct spatio-temporal climate variability. Because classical dendrochronology based on ring-width patterns is often challenging in the tropics, alternative approaches such as Quantitative Wood Anatomy (QWA) based on the measurement and quantification of anatomical traits within tree rings can be a significant advance in the field. Here we assess the dendrochronological potential of Polylepis microphylla and its climate sensitivity by using i) classic dendrochronological methods to generate the first Tree-ring Width (TRW) chronology for this tree species spanning from 1965 to 2018; ii) radiocarbon (¹⁴C) analyses as an independent validation method to assess the annual periodicity of the tree growth layers; and iii) QWA to generate tree-ring annual records of the number (VN) and size (VS) of vessels to investigate the climate sensitivity of these anatomical traits. The annual periodicity in P. microphylla radial growth was confirmed by both dendrochronological and ¹⁴C analyses. We found that VN and VS are promising new proxies to reconstruct climate variability in this region and that they provide different information than TRW. While TRW provides information at inter-annual resolution (i.e., year-to-year variability), VN and VS generated with sectorial QWA provide intra-annual resolution for each stage of the growing process. The TRW and the anatomical traits (i.e., VN and VS) showed strong positive correlation with maximum temperature for different periods of the growing season: while VS is higher with warmer conditions prior to the growing season onset, tree-rings are wider and present higher number of vessels when warmer conditions occur during the current growing season. Our findings pointed out the suitability of P. microphylla for dendrochronological studies and may suggest a good performance of this species under the significant warming expected according to future projections for the tropical Andes.     

Characteristics of a multi-species conifer network of wood properties chronologies from Southern Australia

Progress in quantitative wood anatomy has resulted in a growing number of increasingly understood proxies from the tree-ring archive. Much of this work has been based on tree species in the Northern Hemisphere. Here, we present and examine a relatively dense network of wood property chronologies (wood density, tracheid radial diameter, cell wall thickness and ring width) from several species in Tasmania, southern Australia. We ask how the relationships amongst the different types of chronologies differ within and amongst species. We also consider how each chronology responds to monthly climate. In general terms, and similar to findings in the Northern Hemisphere, relationships between the various wood properties and climate are stronger than those between climate and ring width chronologies. An important exception to this is the highest elevation Lagarostrobos franklinii site. Additionally, strongest response to climate for the wood properties generally occurs for the concurrent growing season compared to the prior growing season for ring width. Relationships amongst the various chronology types differ for the various species, with L. franklinii also showing some variation in these relationships by site (possibly associated with elevation). Results suggest there is considerable value in further exploring the potential for developing anatomical wood chronologies for climate reconstruction from other species for which ring widths do not exhibit a strong climate signal.     

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.