Age-dependent responses of tree-ring growth and intra-annual density fluctuations of Pinus pinaster to Mediterranean climate

Dendrochronology generally assumes that climate–growth relationships are age independent once the biological growth trend has been removed. However, tree physiology, namely, photosynthetic capacity and hydraulic conductivity changes with age. We tested whether the radial-growth response to climate and the intra-annual density fluctuations (IADFs) of Pinus pinaster Ait. varied with age. Trees were sampled in Pinhal de Leiria (Portugal), and were divided in two age classes: young (<65 years old) and old (>115 years old). Earlywood and tree-ring width of young P. pinaster trees were more sensitive to climate influence while the response of latewood width to climate was stronger in old trees. Young trees start the growing season earlier, thus a time window delay occurs between young and old trees during which wood cells of young trees integrate environmental signals. Young trees usually have a longer growing season and respond faster to climate conditions, thus young P. pinaster trees presented a higher frequency of IADFs compared with old trees. Most of the IADFs were located in latewood and were positively correlated to autumn precipitation. The radial-growth response of P. pinaster to climate and the IADFs frequency were age dependent. The use of trees with different age to create a tree-ring chronology for climate studies can increase the resolution of climatic signals. Age-dependent responses to climate can also give important clues to predict how young and old trees react to climate change.     

Linking tree growth and intra-annual density fluctuations to climate in suppressed and dominant Pinus sylvestris L. trees in the forest-steppe of Southern Siberia

Climate change affects forest dynamics with potential consequences for essential ecosystem services. The retrospective analysis of secondary growth unveils the effect of climate on forests. However, most tree-ring studies focus on dominant trees, and less is known about the climatic response of their neighbor suppressed trees. We evaluated the influence of tree social status (dominant/suppressed) on climate response in Pinus sylvestris L. trees from two sites with contrasting water availability conditions in the forest-steppe ecotone in southern Siberia. Tree-ring width and intra-annual density fluctuations (IADFs) were used as proxies. Late spring to early summer conditions were the main climate drivers in both tree social status, but the climate response of suppressed trees was stronger and had a longer time window (May-June). IADFs’ occurrence was controlled by temperature and its frequency was modulated by local conditions, being more common at the dry site, with tree status just marginally significant. Our results suggest that under the projected warmer and drier climate, suppressed trees in southern Siberia will be prone to increased water shortage, leading to possible higher mortality of more sensitive suppressed trees, with potential consequences for carbon sequestration in the forest-steppe ecosystems in southern Siberia.     

Tree-ring growth and intra-annual density fluctuations of Pinus pinaster responses to climate: does size matter?

Dendroclimatology generally assumes that climate–growth relationships are age and size independent. However, there is evidence that climate response can be unstable across different age/size classes. In addition, the occurrence of some anatomical features, such as intra-annual density fluctuations (IADFs), is age dependent. The present study investigates whether the climate–growth responses and the occurrence of IADFs in an even-aged stand of Pinus pinaster Ait., growing under Mediterranean climate, are also size-dependent. We randomly selected 60 P. pinaster trees falling within two stem diameter classes: small (<27 cm) and large (>35 cm). Tree rings were crossdated, measured and IADFs identified according to their position within the ring. The residual chronologies of both size classes were strongly correlated, suggesting a common signal. In fact, similar growth–climate relationships were observed in large and small trees. The frequency of IADFs was higher in large than in small trees, suggesting that IADFs were more likely to occur in wider rings of fast-growing trees. In both size classes, most of the IADFs were found in latewood. Latewood IADFs were triggered by the combination of dry June, wet September, and warm December, whereas IADFs located at the end of earlywood were triggered by previous winter precipitation and favorable conditions before summer (high precipitation for large trees and lower temperature for small trees). Our results suggest that IADFs can be a mechanism used at the individual level for adaptation to drought in P. pinaster. The climatic signal of IADFs between earlywood and latewood was mediated by stem size suggesting that future tree-ring studies should include trees stratified by size to better estimate the sensitivity of IADFs to climate.     

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.     

Climate impact on growth dynamic and intra-annual density fluctuations in Aleppo pine (Pinus halepensis) trees of different crown classes

Mediterranean environments are of special interest for the study of the relationships between climate, growth and anatomic features. Dendrochronological techniques were applied at eight sampling sites that were selected throughout the natural distribution area of Pinus halepensis in the Iberian Peninsula. The objectives of this paper were: (i) to identify relationships between radial growth and climate for different crown classes of Aleppo pine (P. halepensis Mill.); (ii) to quantify the presence of intra-annual density fluctuations (IADFs) according to crown class and cambial age; (iii) to establish the relationships between IADFs and climate. In the more mesic sites, dominant trees showed higher climatic sensitivity than suppressed trees, while in the more xeric sites suppressed trees showed higher sensitivity than dominant trees. Tree-ring growth of both crown classes correlated positively with precipitation during and prior to the growing season. IADFs were more frequent in young than in old stands without differences between crown classes. Precipitation in April and December was positively correlated to the occurrence of IADFs, while precipitation in July correlated negatively. A higher frequency in IADFs occurred in the last 50 years, which coincides with the increase in drought events in the Iberian Peninsula.     

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.     

The facultative bimodal growth pattern in Quercus ilex – A simple model to predict sub-seasonal and inter-annual growth

In Mediterranean climates, bimodal growth patterns, corresponding to two peaks in radial increment during favorable seasons, have been described in several tree species. However, we lack a better mechanistic understanding of bimodality and its potential responses to the predicted warming and aridification trends. Filling this research gap is important since growth duration affects the capacity of trees to form wood and uptake carbon. Here we used an 11-year (1994–2004) long record of dendrometer data of the Mediterranean Holm oak (Quercus ilex) and compared how climate related to radial increment in trees from the south- and the north-facing slopes. We also related climate variables to tree-ring width and the production of intra-annual density fluctuations (IADFs), which reflects bimodality. In this paper, we introduce a model called VS-Lite2 to simulate tree-growth dynamics, which is a modified version of the process-based Vaganov-Shashkin Lite model. The VS-Lite2 model adequately reproduced the bimodal intra‐annual pattern of radial growth, IADFs, and annual tree growth. Trees from the south-oriented slope grew more, produced more IADFs and showed a more marked bimodal pattern than trees from the north-facing slope. These differences agree with the observation that late-summer drought constrained growth. Therefore, radial-growth models should consider plastic bimodality and micro-environmental conditions in areas subjected to seasonal droughts.     

Does insect induced defoliation affect anatomical structure of oak wood?

We counted the intra-annual density fluctuations (IADFs) of oak (Quercus robur) tree rings sampled in a deciduous broadleaf forest near Kozelsk (Kaluga region) in the Ugra National Park, Central Russia. Three sites in the same forest massif were sampled. We also sampled ash (Fraxinus excelsior) trees in one of these sites as a non-host species for the identification of insect outbreaks. In total, 49 oak and 21 ash trees were sampled and analyses were performed for the period 1878–2015. Within the selected time framework, we observed numerous cases with: 1) high percentage of IADFs, 2) decreased oak tree ring widths compared to that of ashes, 3) considerable quantity of oak trees with abrupt growth changes (AGC), and 4) statistically significant differences in counts of IADFs and AGC between the three sampling sites. These traits were characteristic for several consecutive years, and we interpret these as evidence of the insect herbivory causing growth interruptions during the period of the oak leaf expansion. Other external stress factors including summer droughts and late spring frosts can intensify IADFs manifestations during insect outbreaks, but did not affect oak growth strongly enough to cause growth interruptions without insect interference.     

Structural attributes,tree-ring growth and climate sensitivity of Pinus nigra Arn. at high altitude: common patterns of a possible treeline shift in the central Apennines (Italy)

European black pine (Pinus nigra ssp. nigra Arnold) encroachment at increasing elevation has been analyzed at four treeline ecotones of the central Apennines (Italy). The study sites are located along a North-South gradient of 170 km across Marche and Abruzzo regions in Central Italy. The aims of this study were: (i) to detect possible common patterns of structural attributes of black pine regeneration at the treeline ecotone; (ii) to date the seedlings germination and (iii) to assess the climate influence on the pine upward encroachment process also using intra-annual density fluctuations (IADFs) in tree-rings. We sampled 658 encroached black pine trees above the current treeline to the mountain top. All individuals were mapped and their basal stem diameter, total height, annual height increments and other structural attributes measured. One increment core was extracted from stem base of most samples for cambial age determination and detection of intra-annual density fluctuations (IADF). At two sites we also extracted cores at DBH from forest trees to assess climate–growth relationships of black pine. We used multivariate analysis (PCA) to explore the correlation structure of the main tree attributes, regression analysis to relate radial and height increment and dendroclimatic analysis to assess the influence of climate on tree growth and IADF formation.Most black pine trees were located at high altitude and their structural attributes were similar at the four sites where the pine encroachment process started between 30 and 40 years ago featuring similar germination peaks and growth patterns. Black pine is particularly sensitive to maximum temperatures and IADF occurred in mid-late summer with highest frequency peaks between 2003 and 2004. The pine encroachment process, besides the differences of environmental features and land use histories of the four study sites, appears synchronic and spatially diffused. Consistent tree-growth dynamics and the species adaptation to a warming climate are signals envisaging a possible treeline upward shift.     

Climate control on ring width and intra-annual density fluctuations in <Emphasis Type="Italic">Pinus kesiya</Emphasis> growing in a sub-tropical forest of Manipur,Northeast India

Key message

Growth ring study of Pinus kesiya (khasi pine) growing in sub-tropical forest in Manipur, northeast India was performed to understand climate signatures in ring widths and intra-annual density fluctuations.

Abstract

The growth rings in khasi pine (Pinus kesiya Royle ex Gordon) growing in sub-tropical Reserve Forest in Imphal, Manipur, northeast India were analysed to understand environmental signals present in ring-width series and intra-annual density fluctuations (IADFs). For this the growth ring sequences in increment core samples collected from 28 trees were precisely dated and a ring-width chronology spanning AD 1958–2014 developed. The correlation analyses between ring-width chronology and weather data of Imphal revealed that a cool April–May–June favour tree growth. The wood anatomical features of growth rings revealed the occurrence of IADFs in early- and latewoods. The IADFs in earlywood were found to be associated with reduced precipitation in months from April to July. However, the wetter conditions in late growing season, especially August/September triggered the formation of IADFs in latewood. Our findings endorse that the IADF chronologies of khasi pine could emerge as an important proxy of summer monsoon rainfall in long-term perspective in data scarce region of northeast India.

     

Tree ring anatomy indices of Pinus tabuliformis revealed the shifted dominant climate factor influencing potential hydraulic function in western Qinling Mountains

Trees can adjust xylem anatomical structure related with potential hydraulic functions to cope with climate variability. We therefore need a better understanding of how climate variability constrains wood anatomy and tree radial growth. Pinus tabuliformis dominates natural forests and plantations over the western Qinling Mountains, which is one of the ecologically vulnerable areas in China. Here, we investigated the response of P. tabuliformis tree-ring anatomical structure to climate variability by applying wood anatomy analysis, and evaluated the influences of anatomical traits on potential hydraulic functions and the climate significance of intra-annual density fluctuations (IADFs). We found that with the increasing temperature from spring to summer, the negative effect of temperature on the formation and enlargement of earlywood and transition-wood tracheids was gradually enhanced. However, spring precipitation not only had a direct and positive influence on the formation of earlywood, but also had a delaying impact on the transition-wood cell enlargement. Besides, the smaller earlywood tracheid size of P. tabuliformis could be a substantially characteristic reflecting spring drought. The contribution of lumen diameter on conduit wall reinforcement was dominated in earlywood, while the contribution of cell wall thickness was greater than that of lumen diameter in latewood. The different contributions of anatomical traits on conduit wall reinforcement would further affect the response of potential hydraulic function to climate. IADFs of P. tabuliformis could be a potential indicator to reflect the abnormal summer precipitation events in the western Qinling Mountains. IADFs with strong and weak intensity indicated years with high and low rates of change in mid-summer precipitation, respectively. Future warmer and drier climate in the western Qinling Mountains will likely result in the production of smaller tracheids to ensure hydraulic safety, which means the stronger drought resistant of P. tabuliformis in the future. In this study, we linked the xylem anatomy and potential hydraulics functions with intra-seasonal climate variability in the context of climate warming and drying, and proposed some xylem anatomical indices reflecting potential drought events.     

黄土高原南北样带尺度人工刺槐林对气候响应的敏感性分析

根据黄土高原南北样带尺度的人工刺槐林(Robinia pseudoacacia)的年轮宽度资料,分析了该地区刺槐树木生长趋势,以及刺槐年表对气候响应随降雨梯度变化规律。研究结果表明延安以北的刺槐样点(绥德、神木)年轮指数近期趋于下降,树木有生长衰退现象;而延安以南刺槐样点(延安、富县、宜君、永寿)年轮指数近期趋于上升,树木无生长衰退现象。气候响应结果表明,刺槐年表对气候响应均以延安样点最为敏感,表现年表与温度的负相关关系,以及年表与降雨和干旱指数的正相关关系,而延安以北和以南刺槐样点对气候响应敏感性均较低。黄土高原中部延安地区地处森林草原过渡带,刺槐生长对外界环境变化最为敏感,年表中气候信号也较强;延安以南地区地处森林植被带,气候条件较为适宜刺槐林生长,因而年表中气候信号较弱;延安以北地区地处草原植被带,气候条件比较恶劣,刺槐生长对干旱气候已有一定适应性特征,因而年表中气候信号也较弱。     

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.     

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.     

Intra-annual radial growth and water relations of trees: implications towards a growth mechanism

There is a missing link between tree physiological and wood-anatomical knowledge which makes it impossible mechanistically to explain and predict the radial growth of individual trees from climate data. Empirical data of microclimatic factors, intra-annual growth rates, and tree-specific ratios between actual and potential transpiration (T PET(-1)) of trees of three species (Quercus pubescens, Pinus sylvestris, and Picea abies) at two dry sites in the central Wallis, Switzerland, were recorded from 2002 to 2004 at a 10 min resolution. This included the exceptionally hot and dry summer of 2003. These data were analysed in terms of direct (current conditions) and indirect impacts (predispositions of the past year) on growth. Rain was found to be the only factor which, to a large extent, consistently explained the radial increment for all three tree species at both sites and in the short term as well. Other factors had some explanatory power on the seasonal time-scale only. Quercus pubescens built up much of its tree ring before bud break. Pinus sylvestris and Picea abies started radial growth 1-2 weeks after Quercus pubescens and this was despite the fact that they had a high T PET(-1) before budburst and radial growth started. A high T PET(-1) was assumed to be related to open stomata, a very high net CO2 assimilation rate, and thus a potential carbon (C)-income for the tree. The main period of radial growth covered about 30-70% of the productive days of a year. In terms of C-allocation, these results mean that Quercus pubescens depended entirely on internal C-stores in the early phase of radial growth and that for all three species there was a long time period of C-assimilation which was not used for radial growth in above-ground wood. The results further suggest a strong dependence of radial growth on the current tree water relations and only secondarily on the C-balance. A concept is discussed which links radial growth over a feedback loop to actual tree water-relations and long-term affected C-storage to microclimate.     

Quercus robur survival at the rear edge in steppe: Dendrochronological evidence

Climate change is altering forest ecosystems worldwide, particularly in steppe landscapes, where the rare tree communities are challenged with steadily increasing droughts. In the steppe of Eastern Europe, amid dry conditions, Quercus robur occupies mostly riverine habitats and ravines. Here we study the climate sensitivity and drought vulnerability of a Q. robur population located at the rear edge of the species range, in the steppe of Ukraine. The population occupies two adjacent but clearly contrasting in their microclimatic conditions sites: a river floodplain and a steep-sloping river bank. We develop tree earlywood, latewood, and total ring width site-level chronologies and evaluate their relationship with regional climate variables and the local river's water level using response function analysis. We find that trees growing in the floodplain and at the steep river bank have exhibited slightly different growth patterns. The trees at the flooded site have benefited from water proximity, which facilitated their earlywood growth. These trees have responded positively to the current May and previous September precipitation and previous and current May temperatures. At the non-flooded site, the trees have experienced warm temperatures and the lack of precipitation in June. The extreme drought episodes have triggered a decrease in the latewood and total ring width in trees from both sites. We established that oak growth in the floodplain had been depressed by an unknown stressor around 1900, therefore limiting our ability to identify the more beneficial steppe habitat for Q. robur conclusively. Nevertheless, our results provide a dendrochronological evidence of Q. robur survival in a dry steppe environment and lend new insights into local microclimatic factors contributing to it.     

Long-term changes in tree-ring–climate relationships at Mt. Patscherkofel (Tyrol,Austria) since the mid-1980s

Although growth limitation of trees at Alpine and high-latitude timberlines by prevailing summer temperature is well established, the loss of thermal response of radial tree growth during last decades has repeatedly been addressed. We examined long-term variability of climate–growth relationships in ring width chronologies of Stone pine (Pinus cembra L.) by means of moving response functions (MRF). The study area is situated in the timberline ecotone (ca. 2,000–2,200 m a.s.l.) on Mt. Patscherkofel (Tyrol, Austria). Five site chronologies were developed within the ecotone with constant sample depth (≥19 trees) throughout most of the time period analysed. MRF calculated for the period 1866–1999 and 1901–1999 for ca. 200- and ca. 100-year-old stands, respectively, revealed that mean July temperature is the major and long-term stable driving force of Pinus cembra radial growth within the timberline ecotone. However, since the mid-1980s, radial growth in timberline and tree line chronologies strikingly diverges from the July temperature trend. This is probably a result of extreme climate events (e.g. low winter precipitation, late frost) and/or increasing drought stress on cambial activity. The latter assumption is supported by a <10% increase in annual increments of ca. 50-year-old trees at the timberline and at the tree line in 2003 compared with 2002, when extraordinary hot and dry conditions prevailed during summer. Furthermore, especially during the second half of the twentieth century, influence of climate variables on radial growth show abrupt fluctuations, which might also be a consequence of climate warming on tree physiology.     

Effects of age and site quality on the distribution of biomass in Scots pine (Pinus sylvestris L.)

The distribution of the above-ground and below-ground biomass of Scots pine in southern Finland were investigated in trees of different ages (18–212 years) from two types of growth site. Secondly, some structural regularities were tested for their independence of age and growth site. Trees were sampled from dominant trees which could be expected to have a comparable position in stands of all ages. All stands were on sorted sediments. The biomass of the sample trees (18 trees) was divided into needles, branch sapwood and heartwood, stem sapwood and heartwood, stem bark, stump, large roots (diameter >20 cm), coarse roots (five classes) and fine roots. The amount of sapwood and heartwood was also estimated from the below-ground compartments. Trees on both types of growth site followed the same pattern of development of the relative shares of biomass compartments, although the growth rates were faster on the more fertile site. The relative amount of sapwood peaked after canopy closure, coinciding with the start of considerable heartwood accumulation. The relative amount of needles and fine roots decreased with age. The same was true of branches but to a lesser degree. The relative share of the below-ground section was independent of tree age. Foliage biomass and sapwood cross-sectional area were linearly correlated, but there were differences between the growth sites. Needle biomass was linearly correlated with crown surface area. The fine root to foliage biomass ratio showed an increasing trend with tree age.     

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.     

Influence of climate on multiple tree-ring parameters of Pinus kesiya from Manipur,Northeast India

Tree-rings of Pinus kesiya from southern region of Manipur, Northeast India were used to develop chronologies of multiple tree-ring parameters that are: total-ring width (TRW), earlywood width (EW), latewood width (LW) and adjusted latewood (LWadj). The time span of these chronologies is 39 years (1980–2018 C.E.) and we compared their growth responses with monthly and daily climatic records. The comparison revealed a broadly consistent pattern of climate sensitivity with daily climate exhibiting higher correlation. The climate signals during pre-monsoon (March–May) were recorded in TRW and EW, whereas late-monsoon to post-monsoon climate signals were recorded in LW and LWadj. The spatial correlation analysis of tree-ring parameters and global sea surface temperature (SST) showed a positive relationship between tree growth with tropical Pacific Ocean and Indian Ocean during winter (December–February) and pre-monsoon (March–May) seasons. The LW and LWadj were also correlated with peak summer monsoon (July–August) SST over the tropical Pacific Ocean. IADFs observed in EW (E-IADF) were caused by dry and warm conditions during March–April. IADFs in LW (L-IADF) occurred due to a combination of enhanced rainfall and temperature during post-monsoon (October–November) season. Evidence of stand-specific micro-climatic conditions in the formation IADFs in this species was also found. This study showed that multiple parameters of P. kesiya provides a lucid understanding of climate response on its growth and can be considered as a proxy for studying sub-seasonal changes in past environmental conditions in longer records.