Exploring wood anatomy,density and chemistry profiles to understand the tree-ring formation in Amazonian tree species

Long-term analysis of tree growth using annual tree rings is increasingly in demand for tropical tree species. The basis of these studies has traditionally been the anatomical identification of the annual ring boundary. However, the structure of these annual rings has been sparsely explored for complementary physical and chemical wood traits. Here, we explore the relationships among wood density features and chemical elements (S, K, Ca, Mn) involved in the annual tree ring formation of 12 tropical tree species from non-flooded forest in the southern Amazon basin. Transverse wood sections were used for each species to determine: 1) macroscopic distinction (radial growth and wood density), 2) microscopic analyse of vessels, axial and ray parenchyma (anatomy) and 3) X-ray densitometry (physical) and X-ray fluorescence (chemical). For some species, the profiles of wood density, and Ca and Mn content showed intra- and inter-annual patterns that allowed to define and characterize the growth boundary of tree rings. Ca, K and S were mainly distributed in axial parenchyma cells, and around vessels, whereas, Mn was mainly distributed in fibres. Our results showed significant species-specific correlations between tree-ring width, density and concentrations of Ca, K and Mn. The anatomical characterization and the complementary information provided by the density and chemical profiles in some Amazonian species can represent a valuable proxy to improve the definition of annual ring-boundaries and improve the understanding of long-term growth and physiological patterns.     

Tree-ring distinctness,dating potential and climatic sensitivity of laurel forest tree species in Tenerife Island

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.     

Climate–growth relationships of the dominant tree species from semi-arid savanna woodland in Ethiopia

Long-term climate–growth relationships, were examined in tree rings of four co-occurring tree species from semi-arid Acacia savanna woodlands in Ethiopia. The main purpose of the study was to prove the presence of annual tree rings, evaluate the relationship between radial growth and climate parameters, and evaluate the association of El Niño and drought years in Ethiopia. The results showed that all species studied form distinct growth boundaries, though differences in distinctiveness were revealed among the species. Tree rings of the evergreen Balanites aegyptiaca were separated by vessels surrounding a thin parenchyma band and the growth boundary of the deciduous acacias was characterized by thin parenchyma bands. The mean annual diameter increment ranged from 3.6 to 5.0 mm. Acacia senegal and Acacia seyal showed more enhanced growth than Acacia tortilis and B. aegyptiaca. High positive correlations were found between the tree-ring width chronologies and precipitation data, and all species showed similar response to external climate forcing, which supports the formation of one tree-ring per year. Strong declines in tree-ring width correlated remarkably well with past El Niño Southern Oscillation (ENSO) events and drought/famine periods in Ethiopia. Spectral analysis of the master tree-ring chronology indicated occurrences of periodic drought events, which fall within the spectral peak equivalent to 2–8 years. Our results proved the strong linkage between tree-ring chronologies and climate, which sheds light on the potential of dendrochronological studies developing in Ethiopia. The outcome of this study has important implications for paleoclimatic reconstructions and in restoration of degraded lands.     

Living on the edge: Legacy of water availability on Tetraclinis articulata secondary growth under semiarid conditions in Morocco

Tetraclinis articulata is a xerothermic Mediterranean conifer native from NW Africa that grows under semiarid conditions in the Atlas Mountains. This species is particularly well adapted to water scarcity and forms xeric woodlands subjected to recurring drought, aridification and overexploitation. Wood of T. articulata has been considered of limited dendrochronological value due to abundant anatomical anomalies in their growth rings. We studied tree-ring growth and tree-ring features to check its dendrochronological potential and evaluate their climatic signals and drought legacy effects at two semiarid sites with contrasting elevation in Morocco. Tree-ring boundaries were properly identified on wood cores from 60 to 68% of the sampled trees. Intra-annual wood density fluctuations (IADF), microrings, missing rings and rings with undefined limits were most abundant at the low-elevation coastal site than at the mountain site. Microring and missing ring abundance was inversely related to tree-ring growth, independent of cambial age, and mostly dependent on drought occurrence. IADF frequency was independent of tree-ring growth but inversely related to cambial age. A complex response of IADF formation to cool March to June conditions was found at the low-elevation coastal site, while IADF formation in the mountain site was related to May water availability. Undefined ring limits were observed only at the low-elevation site, showing positive relationships with elevated maximum temperatures in previous winter. Previous winter rainfall favored radial growth, but a significant legacy effect of water availability was evident up to three years prior to growth. Tree-ring growth was mostly related to water availability 33 and 26 months prior to growth at low-elevation and mountain sites, respectively. The obtained results reveal a very plastic cambial activity and a strong ability of T. articulata to withstand drought even for long periods.     

Growth-ring boundaries of tropical tree species: Aiding delimitation by long histological sections and wood density profiles

Recent methodological advances have opened new perspectives for tropical dendrochonological studies by facilitating the visualization, delimitation, and analyses of tree-rings. One of those improvements was brought by X-ray densitometry, which allows building radial wood density profiles at microscopic scale. Furthermore, recent methods allow for cutting long histological sections to study anatomical variations along the entire radius of trees. These techniques have mainly been applied to low wood-density species from temperate and Mediterranean regions, with only limited applications in the tropics. Here we provide an improved protocol that allows for obtaining long histological sections of tropical woods, apply it to six species with varying wood densities 0.45−0.85 g cm⁻³ (Eucalyptus grandis, Tectona grandis, Acacia mangium, Cedrela fissilis, Hymenaea courbaril, and Copaifera duckei), and explore potential applications for tree-ring analyses. We provide instructions on core-microtome knife adjustments, procedures for softening and sectioning long histological samples of high wood-density species. We also present a multi-proxy approach that combines X-ray density profiles with the histological sections that improve the characterization and distinction of the various and complex tropical growth rings anatomical markers (fibre zone, marginal parenchyma, and ring porosity). This multi-proxy approach also opens the door for obtaining quantitative anatomy and physical parameters of tropical species with (intra-annual resolution. Our proposed approach is thus not only an additional tool to improve ring-boundary delimitation of tropical species, but it also paves the way to more innovative, borderline approaches in tropical dendrochronology.     

xRing—An R package to identify and measure tree-ring features using X-ray microdensity profiles

Climate influences tree-ring density and ring-density variables extracted from X-ray images have been widely used for climate reconstructions. The R package xRing was developed to identify and measure tree rings on X-ray microdensity profiles automatically. This package is available for free and it offers functions to visualize and calibrate X-ray images, to detect tree-ring borders and to identify earlywood-latewood transition using wood density variations at the inter- and the intra-ring scale. The most important functions are calibrateFilm, detectRings, correctRings, detectEwLw, and getDensity. Outputs of these functions are S3 objects, for which specific methods are provided, including plot and print. The non-linear relationship between optical density of the film and wood density is defined by the function calibrateFilm. The function detectRings detects tree rings using wood density profiles as input. This function uses the difference between local maximum and minimum values to identify tree-ring borders automatically. The correctRings function is used to call a Graphical User Interface (GUI) to visualize and to correct tree-ring borders manually. After correcting tree-ring borders, the detectEwLw function is used to compute earlywood and latewood widths by dividing rings according to relative intra-ring density changes. The getDensity function computes for each tree ring the minimum (maximum) density and the mean earlywood, latewood and whole-ring density. Finally, a list with dataframes with tree-ring width and density variables can be obtained using the function getRwls. One of the major advantages of xRing package is that requires little knowledge of R language, but at the same time it can be easily changed or adapted by experienced users.     

Potential of Schinopsis lorentzii for dendrochronological studies in subtropical dry Chaco forests of South America

The lack of knowledge about species with well-delimited annual rings has hampered the development of dendrochronological records in the subtropical Chaco region of South America. In this contribution, we present the first tree-ring chronology of Schinopsis lorentzii (Anacardiaceae), a dominant species in the semi-arid Chaco. Cross sections were collected near Las Lajitas, Salta, Argentina, and processed following the methods commonly used in dendrochronology. Annual growth variations between radii from a single individual and between radii from different trees were highly correlated. To determine the climatic parameters that control radial growth, we compared annual tree-ring variations against regional temperature and precipitation records. Correlation functions indicate that tree growth is highly influenced by spring–summer rainfall variations, which represent more than 80% of the total annual precipitation. The chronology, which covers the interval from 1829 to 2004, provides a context for the unprecedented increase in precipitation since the mid-1970s in the region. The climatic-sensitivity of S. lorentzii provides a unique opportunity to reconstruct precipitation variations during past centuries in the extensive semiarid regions of subtropical South America.     

Dendrochronological analysis and dating of wooden artefacts from the prehistoric copper mine Kelchalm/Kitzbühel (Austria)

The potential of dendrochronological analysis of wood found in prehistoric and historic mining areas in Tyrol has remained unattended for a long time. For the first time, systematically analysed wooden artefacts from a prehistoric mining area in Tyrol (Kelchalm near Kitzbühel) can be presented here.The investigated artefacts, related to mining and everyday life, were found in the course of archaeological excavations, which were carried out between 1932 and 1953 by Richard Pittioni and Ernst Preuschen. Taking an adequate number of tree rings and well-preserved wood wane into account, 21 pieces of mining timber were pre-selected for a dendrochronological analysis. The identified wood species are spruce (Picea abies, n=18) and fir (Abies alba, n=3). The length of the established tree-ring series ranges from 13 to 145. We cross-dated the tree-ring series of seven wooden artefacts among each other, which resulted in a spruce-fir tree-ring record of 153 values (Kelchalm mean curve). The last tree ring measured of the Kelchalm spruce-fir mean curve dates back to 1237 BC. This accurate dendro-result dates the Bronze Age mining activities at the Kelchalm to about two centuries earlier than the long-lasting assumption proposed by Richard Pittioni. His assumption was based on the typology of ceramic and metal artefacts.The established dendro-date for the Bronze Age mine at the Kelchalm matches with available ¹⁴C results from other important copper-mining areas in the north-eastern Alps (NE Alps). The activities at these other sites are dated between the 17th and 6th century BC. Furthermore, the radiocarbon dating, as well as the dendro-result from the Kelchalm, suggests a transition from earlier mined copper-ore deposits in the eastern areas of the NE Alps, to the later mined ore deposits in the western section. This has led to both parallel and sequential mining activities in several ore districts during the last two millennia BC in the NE Alps.     

Dendroclimatological assessment of Polylepis rodolfo-vasquezii: A novel Polylepis species in the Peru highlands

In spite of enormous diversity in tree species, dendrochronological records in the tropical Andes are very scarce. Therefore, it is necessary to increase the search for new tree species with high dendrochronological characteristics in the tropical Andes, including the humid Puna of Peru. We present the first tree-ring chronology from Polylepis rodolfo-vasquezii, a recently described tree species in the Central Andes of Peru between 4000 and 4400 m elevation. Fifty trees were sampled in the district of Comas, Peru. After establishing the anatomical characteristics that delimit the annual growth rings, we developed a ring-width chronology by applying conventional dendrochronological techniques. The chronology covers the period 1869–2015 (157 years) and is well replicated from 1920 to present (> 20 samples). The statistics used to evaluate the quality of the chronology indicate that the P. rodolfo-vasquezii has similar values of MS, RBAR and EPS to those shown by other Polylepis spp chronologies. To determine the main climatic factors controlling the growth of P. rodolfo-vasquezii, we compared our chronology with local and regional temperature and precipitation records. Growth season temperature (November to May) seems to be the main climatic factor modulating inter-annual variations in the growth of this species. The sensitivity to inter-annual temperature variations highlights the potential of P. rodolfo-vasquezii to provide climatically sensitive dendrochronological records in the Central Andes. To our knowledge, this is the first tree-ring record in South America displaying significant relationships with temperature over the tropical Atlantic Ocean.     

Automated 3D tree-ring detection and measurement from X-ray computed tomography

Tree ring analysis is essential to reveal the environmental information encoded in the wood structure. It provides quantitative data on the anatomical structure which can be used, for example, to measure the impact of the fluctuating environment on the tree growth, to support global vegetation models and for the dendrochronological analysis of archaeological wooden artefacts. Currently, several imaging-based methods for tree-ring detection and tree-ring feature estimation exist. However, despite advances in computer vision and edge recognition algorithms, detection of tree-rings is mostly limited to two-dimensional (2D) datasets and performed manually in some cases. This paper describes a new approach to estimate the three-dimensional (3D) structure of tree rings and their width automatically from X-ray computed tomography data. This approach relies on a modified Canny edge detection algorithm, which is capable of detecting fully connected tree-ring edges throughout the image stack. Our results show that this approach performs well on six tree species having conifer, ring-porous and diffuse-porous ring boundary structures. In our study, image denoising proved to be a critical step to achieve accurate results. A major advantage of this procedure is that it requires very little to no user interaction rendering it a reproducible procedure for tree-ring width measurements. As it also provides 3D representations of the ring edges, it also may be used in the future for the inspection of anatomical features.     

The utility of bulk wood density for tree-ring research

Bulk wood density measurements are recognized for their utility in ecology, industry, and biomass estimations. In tree-ring research, microdensitometric techniques are widely used, but their ability to determine the correct central tendency has been questioned. Though rarely used, it may be possible to use bulk wood density as a tool to check the accuracy of and even correct microdensitometric measurements. Since measuring bulk wood density in parallel with X-ray densitometry is quickly and easily done, we suspect that its omission is largely due to a lack of awareness of the procedure and/or its importance. In this study, we describe a simple protocol for measuring bulk wood density tailored for tree-ring researchers and demonstrate a few possible applications. To implement real-world examples of the applications, we used a sample of existing X-ray and Blue Intensity (BI) measurements from 127 living and dead Pinus sylvestris trees from northern Sweden to produce new measurements of bulk wood density.We can confirm that the central tendency in this sample material is offset using X-ray densitometry and that the diagnosis and correction of X-ray density is easily done using bulk wood density in linear transfer functions. However, this approach was not suitable for our BI measurements due to heavy discoloration. Nevertheless, we were able to use bulk wood density to diagnose and improve the use of deltaBI (latewood BI – earlywood BI) with regard to its overall trends and multi-centennial variability in a dendroclimatological application. Moreover, we experimented with percent of latewood width, scaled with bulk wood density, as a time- and cost-effective proxy for annual ring density. Although our reconstruction only explains about half of the variation in ring density, it is most likely superior to using fixed literature values of density in allometric equations aimed at biomass estimations.With this study, we hope to raise new awareness regarding the versatility and importance of bulk wood density for dendrochronology by demonstrating its simplicity, relevance, and applicability.     

Intra-annual variability of anatomical structure and δ<Superscript>13</Superscript>C values within tree rings of spruce and pine in alpine,temperate and boreal Europe

Tree-ring width, wood density, anatomical structure and ¹³C/¹²C ratios expressed as δ¹³C-values of whole wood of Picea abies were investigated for trees growing in closed canopy forest stands. Samples were collected from the alpine Renon site in North Italy, the lowland Hainich site in Central Germany and the boreal Flakaliden site in North Sweden. In addition, Pinus cembra was studied at the alpine site and Pinus sylvestris at the boreal site. The density profiles of tree rings were measured using the DENDRO-2003 densitometer, δ¹³C was measured using high-resolution laser-ablation-combustion-gas chromatography-infra-red mass spectrometry and anatomical characteristics of tree rings (tracheid diameter, cell-wall thickness, cell-wall area and cell-lumen area) were measured using an image analyzer. Based on long-term statistics, climatic variables, such as temperature, precipitation, solar radiation and vapor pressure deficit, explained <20% of the variation in tree-ring width and wood density over consecutive years, while 29–58% of the variation in tree-ring width were explained by autocorrelation between tree rings. An intensive study of tree rings between 1999 and 2003 revealed that tree ring width and δ¹³C-values of whole wood were significantly correlated with length of the growing season, net radiation and vapor pressure deficit. The δ¹³C-values were not correlated with precipitation or temperature. A highly significant correlation was also found between δ¹³C of the early wood of one year and the late wood of the previous year, indicating a carry-over effect of the growing conditions of the previous season on current wood production. This latter effect may explain the high autocorrelation of long-term tree-ring statistics. The pattern, however, was complex, showing stepwise decreases as well as stepwise increases in the δ¹³C between late wood and early wood. The results are interpreted in the context of the biochemistry of wood formation and its linkage to storage products. It is clear that the relations between δ¹³C and tree-ring width and climate are multi-factorial in seasonal climates.     

Dendrochronology of maritime pine in the middle of the Atlantic Ocean

The Azores Archipelago, located in the North Atlantic Ridge, experiences heavy rainfall and mild temperatures with weak seasonal differences due to oceanic influence. To our knowledge, there have been no dendrochronological studies in the Azores. The aim of this study is to explore the dendrochronological potential of Pinus pinaster Ait. growing in this archipelago and to determine what limiting factor is regulating tree growth. To do so, we have sampled adult maritime pine trees growing in a plantation, in the Pico island of the Azores.Tree ring boundaries were not always easily distinguished, suggesting that in some years cambial activity did not stop during winter. Despite this, it was possible to successfully crossdate the tree-ring series and to establish a tree-ring width chronology with a strong common signal. Climatic correlations revealed a positive response to spring precipitation but no temperature signal in the tree-ring width chronology. Tree-ring width was also negatively correlated with the North Atlantic Oscillation (NAO) and the sea level pressure (SLP) in May − June.Intra-annual density fluctuations (IADFs), which are anatomical features formed in response to variations in environmental conditions during the growing season, were present in 85% of the tree rings. IADFs were identified based on its position within the ring: type E⁺, characterized as a transition wood from early- to latewood; type L, the most frequent, characterized as earlywood-like cells within latewood; and type L⁺, characterized as earlywood-like cells between latewood and earlywood of the next tree ring. Each IADF type presented a unique climatic signal: type E⁺ was positively correlated with early summer precipitation and early spring temperature; type L was positively correlated with early autumn precipitation and temperature; and type L⁺ was positively correlated with late autumn precipitation.In conclusion, the tree-ring width chronology established for maritime pine growing in the Pico Island of Azores contains a clear climatic signal for spring precipitation, whereas IADFs frequency correlated better with precipitation later in the growing season. For this reason, we suggest that IADFs should be included in future dendrochronological studies in the Macaronesia Biogeographical region since they can improve the climatic signal present in tree-ring width chronologies.     

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.     

Improved tree-ring visualization using autofluorescence

The great diversity of wood anatomical features found in trees worldwide results in a broad variety of growth-ring boundary types that are not always easy to recognize, especially in tropical woods. However, the presence of clearly visible limits between tree rings is essential for any tree-ring studies. Here, we propose the use of autofluorescence of wood in order to enhance tree-ring visualization. The multispectral light emitted from the fluorescence stereomicroscope can be filtered in specific wavelengths to improve the visualization of wood anatomical features. To evaluate the effectiveness of this technique, we compared visualization under natural light, GFP (green fluorescent protein) filter, RFP (red fluorescent protein) filter and UV filter. We tested this technique with a set of 38 tree species with different types of growth-ring boundaries. Although results are species-specific, fluorescence has been shown to improve the visualization of growth-ring boundaries by enhancing the contrast among cell types. It may highlight fibrous zones (e.g. Cavanillesia arborea, Aspidosperma polyneuron), different porosity patterns (e.g. Myracrodruon urundeuva), secretory canals (e.g. Copaifera langsdorffii), and parenchyma bands (e.g. Tipuana tipu). Fluorescence allows the visualization of growth-ring boundaries in species that were previously described as having indistinct growth rings under natural light. For species with clear tree-ring boundaries such as Cedrela fissilis and Hymenaea courbaril, this approach aids the identification of false rings. In addition to the visualization of growth-ring boundaries, autofluorescence may be useful for other qualitative and quantitative analyses of wood anatomy, such as wood identification and automated measurements of anatomical features. Scientists struggling with tree-ring counting and cross-dating due to difficult tree-ring visualization may find fluorescence useful. It may also aid to identify new species suitable for tree-ring studies.     

Dendrochronological evidence of cockchafer (Melolontha sp.) outbreaks in subfossil tree-trunks from Tovačov (CZ Moravia)

In dendrochronological dating, we encounter the trouble that some samples cannot be dated due to the occurrence of anomalies in the tree-ring series. When dating subfossil wood for the extension of existing master chronologies, this may be an especially unwanted circumstance as subfossil wood is scarce and each sample is valuable. In Moravia (Czech Republic) outbreaks of the cockchafer (Melolontha sp.) used to appear with a disastrous effect on agricultural crops, and, for a dendrochronologist, on samples of wood to be dated. Oak ring width reduction caused by cockchafer induced defoliation can superimpose climate induced growth variations and may complicate dating of historical or subfossil samples. For this study, 33 samples of sub-fossil trunks were assessed; 19 of which were attacked by cockchafer. For comparison, we analysed a total of nine living oak trees from the forest district Strá?nice-Vracov, recently attacked by the cockchafer. Occurrence of the cockchafer was reported there in 1999, 2003 and 2007.We found clear growth patterns with cyclic narrow rings every 4 years. This allowed us to create an artificial curve expressing the recurring cycles of cockchafer outbreaks which assisted us in the detection of the same pattern in subfossil tree growth. The tree-ring series of the attacked living oak stand as well as the 19 individual tree-ring series from subfossil oak trunks could not be dated using standard chronologies and showed a high resemblance to the cockchafer life cycle. Additionally, we found out that the living oak stand was attacked in 1983, 1987, 1991 and 1995 as well. The results support the hypothesis that the periodical reduction observed in tree-rings at regular increments is caused by cockchafer infestation. This evidence is further supported by characteristic anatomical features in early wood. A mean tree-ring series was created from the individual tree-ring series of subfossil oaks even though radiocarbon dating showed completely different ages of samples. This means that the affected tree-ring series cannot be used for dendrochronological dating and dendroclimatological analyses. These series cannot be used without preceding mathematical and statistical modification.     

Comparison of whole wood and cellulose carbon and oxygen isotope series from Pinus nigra ssp. laricio (Corsica/France)

Stable isotopes in tree rings have widely been used for palaeoclimate reconstructions since tree rings record climatic information at annual resolution. However, various wood components or different parts of an annual tree-ring may differ in their isotopic compositions. Thus, sample preparation and subsequent laboratory analysis are crucial for the isotopic signal retained in the final tree-ring isotope series used for climate reconstruction and must therefore be considered for the interpretation of isotope–climate relationships. This study focuses on wood of Corsican Pine trees (Pinus nigra ssp. laricio) as this tree species allows to reconstruct the long-term climate evolution in the western Mediterranean. In a pilot study, we concentrated on methodological issues of sample preparation techniques in order to evaluate isotope records measured on pooled whole tree-ring cellulose and whole tree-ring bulk wood samples. We analysed 80-year long carbon and oxygen chronologies of Corsican Pine trees growing near the upper tree line on Corsica. Carbon and oxygen isotope records of whole tree-ring bulk wood and whole tree-ring cellulose from a pooled sample of 5 trees were correlated with the climate parameters monthly precipitation, temperature and the self-calibrating Palmer Drought Severity Index (sc-PDSI). Results show that the offsets in carbon and oxygen isotopes of bulk wood and cellulose are not constant over time. Both isotopes correlate with climate parameters from late winter and summer. The carbon and oxygen isotope ratios of cellulose are more sensitive to climatic variables than those of bulk wood. The results of this study imply that extraction of cellulose is a pre-requisite for the reconstruction of high-resolution climate records from stable isotope series of P. nigra ssp. laricio.     

Inter-annual and seasonal variability of radial growth, wood density and carbon isotope ratios in tree rings of beech (Fagus sylvatica) growing in Germany and Italy

We investigated the variability of tree-ring width, wood density and ¹³C/¹²C in beech tree rings (Fagus sylvatica L.), and analyzed the influence of climatic variables and carbohydrate storage on these parameters. Wood cores were taken from dominant beech trees in three stands in Germany and Italy. We used densitometry to obtain density profiles of tree rings and laser-ablation-combustion-GC-IRMS to estimate carbon isotope composition (δ ¹³C) of wood. The sensitivity of ring width, wood density and δ ¹³C to climatic variables differed; with tree-ring width responding to environmental conditions (temperature or precipitation) during the first half of a growing season and maximum density correlated with temperatures in the second part of a growing season (July–September). δ ¹³C variations indicate re-allocation and storage processes and effects of drought during the main growing season. About 20% of inter-annual variation of tree-ring width was explained by the tree-ring width of the previous year. This was confirmed by δ ¹³C of wood which showed a contribution of stored carbohydrates to growth in spring and a storage effect that competes with growth in autumn. Only mid-season δ ¹³C of wood was related to concurrent assimilation and climate. The comparison of seasonal changes in tree-ring maximum wood density and isotope composition revealed that an increasing seasonal water deficit changes the relationship between density and ¹³C composition from a negative relation in years with optimal moisture to a positive relationship in years with strong water deficit. The climate signal, however, is over-ridden by effects of stand density and crown structure (e.g., by forest management). There was an unexpected high variability in mid season δ ¹³C values of wood between individual trees (−31 to −24‰) which was attributed to competition between dominant trees as indicated by crown area, and microclimatological variations within the canopy. Maximum wood density showed less variation (930–990 g cm⁻³). The relationship between seasonal changes in tree-ring structure and ¹³C composition can be used to study carbon storage and re-allocation, which is important for improving models of tree-ring growth and carbon isotope fractionation. About 20–30% of the tree-ring is affected by storage processes. The effects of storage on tree-ring width and the effects of forest structure put an additional uncertainty on using tree rings of broad leaved trees for climate reconstruction.     

Dendrochronological potential of four neotropical dry-forest tree species: Climate-growth correlations in northeast Brazil

Tropical dry forests (TDF) are highly important tropical forest ecosystems. Yet, these forests are highly threatened, usually neglected and only poorly studied. Understanding the long-term influences of environmental conditions on tree growth in these forests is crucial to understand the functioning, carbon dynamics and potential responses to future climate change of these forests. Dendrochronology can be used as a tool to provide these insights but has only scantly been applied in (dry) tropical forests. Here we evaluate the dendrochronological potential of four Caatinga neotropical dry forest tree species – Aspidosperma pyrifolium, Ziziphus joazeiro, Tabebuia aurea, and Libidibia ferrea – collected in two locations in northeastern Brazil (Sergipe state). We provide an anatomical characterization of the ring boundaries for the four species and investigate correlations of their growth with local and regional climatic variables. All four species form annual rings and show high inter-correlation (up to 0.806) and sensitivity (up to 0.565). Growth of all species correlated with local precipitation as well as with sea-surface temperatures in the tropical Atlantic and/or tropical Pacific oceans. We also show teleconnections between growth and the El Niño South Oscillation. The strong dependence of tree on precipitation is worrisome, considering that climate change scenarios forecast increased drought conditions in the Caatinga dry forest. Including more species and expanding dendrochronological studies to more areas would greatly improve our understanding of tree growth and functioning in TDFs. This type of knowledge is essential to assist the conservation, management and restoration of these critical tropical ecosystems.