Formation Of Intra-annual Density Fluctuations Research Articles

Latewood intra-annual density fluctuations indicate wet summer conditions and enhanced canopy activity in a Mediterranean ring-porous oak

Summary The climatic significance and ecosystem implications of latewood intra-annual density fluctuations (IADFs) are still not fully understood in Mediterranean ring-porous oak species. To solve this issue, we investigated: (i) the climate drivers of radial growth and latewood IADFs in the Mediterranean oak Quercus faginea, and (ii) whether they were correlated to increased canopy cover and greenness as recorded by the Normalized Difference Vegetation Index (NDVI). IADFs were formed in the mid to late latewood and they were characterized by rows of parenchyma cells. Such IADFs were uncommon and only present in 30–50% of sampled trees during 1999 and 2014, both years characterized by high precipitation in late July and early August. This relationship between IADFs formation and summer wet conditions differed from the conditions that enhance Q. faginea annual growth, wet winter-spring conditions, and low summer temperatures. Furthermore, IADFs formation showed a positive relationship with August NDVI values, indicating a correspondence of canopy greenness with secondary growth reactivation. We conclude that latewood IADFs in Q. faginea are a robust proxy of rare late-summer wet conditions and enhanced canopy activity as reflected by increased August NDVI values. Further monitoring of xylogenesis in other Mediterranean hardwoods along climatic gradients is suggested to mechanistically explain IADF production in seasonally dry biomes.

Intra-annual density fluctuations in tree rings are proxies of air temperature across Europe

Intra-Annual Density Fluctuations (IADFs) are an important wood functional trait that determine trees’ ability to adapt to climatic changes. Here, we use a large tree-ring database of 11 species from 89 sites across eight European countries, covering a climatic gradient from the Mediterranean to northern Europe, to analyze how climate variations drive IADF formation. We found that IADF occurrence increases nonlinearly with ring width in both gymnosperms and angiosperms and decreases with altitude and age. Recently recorded higher mean annual temperatures facilitate the formation of IADFs in almost all the studied species. Precipitation plays a significant role in inducing IADFs in species that exhibit drought tolerance capability, and a growth pattern known as bimodal growth. Our findings suggest that species with bimodal growth patterns growing in western and southern Europe will form IADFs more frequently, as an adaptation to increasing temperatures and droughts.

Intra-annual density fluctuations in silver fir are triggered by drought conditions

Key messageIn Abies alba Mill. (silver fir), the frequency of intra-annual density fluctuations (IADFs) increases along the latitudinal transect, from North to South, is higher in pure than in mixed stands, and their formation is linked to spring and/or summer drought conditions.Trees respond to climate, recording information in tree rings and their anatomical features, such as IADFs. The IADFs are regions within the tree ring where density changes in response to tree physiological processes and environmental conditions. The objective of this study was to assess the effect of different forest-stand structures and of climate on IADF formation and frequency in populations of silver fir along a latitudinal gradient in Italy (Trentino, Molise and Calabria regions). In doing so, we aimed to compare the frequency of IADFs in pure or mixed stands and to understand the ability of silver fir to cope with environmental fluctuations. Results showed higher frequency of IADFs in pure than mixed stands (in Trentino and Calabria) and in younger trees (namely in the pure stand in Calabria), because of the higher sensitivity to environmental variability. The formation of IADFs in silver fir emerged as a response to cope with drought. Summer precipitation (both pure and mixed stands in Trentino) and early spring/summer (both pure and mixed stands in Calabria) played a key role in the formation of IADF-type E+. The formation of IADF-type L+, on the other hand, was related to temperature/precipitation in late summer and early autumn (both pure and mixed stands in Molise) and to precipitation in summer (pure stand in Calabria). Our findings support the theory that IADFs are an important structural/functional mechanism for responding to climate fluctuations.

Effects of Intra-Seasonal Drought on Kinetics of Tracheid Differentiation and Seasonal Growth Dynamics of Norway Spruce along an Elevational Gradient

Research Highlights: Our results provide novel perspectives on the effectiveness and collapse of compensatory mechanisms of tracheid development of Norway spruce during intra-seasonal drought and the environmental control of intra-annual density fluctuations. Background and Objectives: This study aimed to compare and integrate complementary methods for investigating intra-annual wood formation dynamics to gain a better understanding of the endogenous and environmental control of tree-ring development and the impact of anticipated climatic changes on forest growth and productivity. Materials and Methods: We performed an integrated analysis of xylogenesis observations, quantitative wood anatomy, and point-dendrometer measurements of Norway spruce (Picea abies (L.) Karst.) trees growing along an elevational gradient in South-western Germany during a growing season with an anomalous dry June followed by an extraordinary humid July. Results: Strong endogenous control of tree-ring formation was suggested at the highest elevation where the decreasing rates of tracheid enlargement and wall thickening during drought were effectively compensated by increased cell differentiation duration. A shift to environmental control of tree-ring formation during drought was indicated at the lowest elevation, where we detected absence of compensatory mechanisms, eventually stimulating the formation of an intra-annual density fluctuation. Transient drought stress in June also led to bimodal patterns and decreasing daily rates of stem radial displacement, radial xylem growth, and woody biomass production. Comparing xylogenesis data with dendrometer measurements showed ambivalent results and it appears that, with decreasing daily rates of radial xylem growth, the signal-to-noise ratio in dendrometer time series between growth and fluctuations of tree water status becomes increasingly detrimental. Conclusions: Our study provides new perspectives into the complex interplay between rates and durations of tracheid development during dry-wet cycles, and, thereby, contributes to an improved and mechanistic understanding of the environmental control of wood formation processes, leading to the formation of intra-annual density fluctuations in tree-rings of Norway spruce.

Xylem and Phloem Formation Dynamics in Quercus ilex L. at a Dry Site in Southern Italy

Quercus ilex L. dieback has been recently reported at numerous Mediterranean sites. Wood and phloem formation dynamics and tree-ring series of anatomical traits can be used to evaluate growth conditions of trees. We monitored cambial activity in Q. ilex trees growing at a site in southern Italy in order to assess how xylem and phloem production are affected by harsh seasonal climatic variation during a dry year. We followed xylogenesis by counting the number of cambial cells and detecting the occurrence of post-cambial cells throughout the year. As phloem did not show clear growth rings and boundaries between them, we followed the development of phloem fibres—their morphological traits during development and the distance from the cambium served as a reference point to evaluate the phloem production during the year. We detected a multimodal pattern in cambial activity, with wood production in three periods of the year and consequent formation of intra-annual density fluctuations (IADFs). The lowest production of xylem cells was observed in the dry late spring and summer period (likely due to the low water availability), while the highest occurred in autumn (the wettest period). Although we could not differentiate between early and late phloem, the analysis of phloem traits was useful to follow the dynamics of phloem production, which is generally difficult in Mediterranean tree species. We found cambial production of phloem throughout the year, even in the periods without xylem production. The results showed that if tree growth was constrained by environmental limitations, the ratio between xylem to phloem cells decreased and, in the most severely affected trees, more cells were formed preferentially in the phloem compared to xylem. We also briefly report the way in which to solve technical problems with tissue preparation due to extreme hardness and to the peculiar structure of Q. ilex wood and outer bark.

A synoptic view on intra-annual density fluctuations in Abies alba

Tree growth, as an archive of environmental conditions, has proven to provide valuable proxies for climate reconstructions. Density measurements were successfully used in the field of dendroclimatology in the past, with recent efforts to obtain sub-annual predictors from density profiles. Particular attention is laid on anomalies in the density profile named intra-annual density fluctuations (IADFs). Density fluctuations in earlywood with higher than ordinary density are assumed to be triggered by drought events in spring or early summer. How they are formed, linked to chemical and anatomical characteristics and their functionality is not well understood, although necessary to interpret growth-climate relationships of this trait. An explorative assessment of radial profiles from silver fir, showing an IADF in earlywood of 2005, included cell wall and lumen area, cellulose to lignin ratio, microfibril angle, tracheid length as well as size and shape of pits, beside density. The resulting synoptic view of these traits depicted deviations at the IADF position, indicating that multiple phases of wood formation are affected by a triggering event. These anatomical and chemical adaptions result in a hydraulically safer tissue with improved mechanical strength. The presented approach provides first insights on multiple involved wood traits and leads towards a more comprehensive understanding of IADF formation.

Tropical cyclone frequency inferred from intra-annual density fluctuations in longleaf pine in Florida, USA

CR Climate Research Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsSpecials CR 78:249-259 (2019) - DOI: https://doi.org/10.3354/cr01573 Tropical cyclone frequency inferred from intra-annual density fluctuations in longleaf pine in Florida, USA Tyler J. Mitchell1,*, Paul A. Knapp1, Jason T. Ortegren2 1Carolina Tree-Ring Science Laboratory, Department of Geography, Environment, and Sustainability, University of North Carolina at Greensboro, PO Box 26170, Greensboro, NC 27402, USA 2Department of Earth and Environmental Sciences University of West Florida, 11000 University Pkwy, Pensacola, FL 32514, USA *Corresponding author: tjmitche@uncg.edu ABSTRACT: We present a new method for identifying historic tropical cyclone activity utilizing frequencies of intra-annual density fluctuations in longleaf pine in western Florida. In addition, in this work we provide information about the causal factors that determine the formation of intra-annual density fluctuations (IADFs) in longleaf pine latewood. Specifically, we test the viability of using late latewood (L+) IADFs in longleaf pine as a proxy for historic tropical cyclone frequency and precipitation for the period 1950-2017. The stabilized frequency of L+ IADF occurrence is significantly (p < 0.01) associated with the Palmer drought severity index (PDSI) for the months June through October, indicating that high amounts of late growing-season moisture promote the formation of IADFs in latewood. We find the strongest relationships between PDSI and IADF occurrence during September and October, indicating the influence of tropical cyclone (TC)-sourced precipitation on IADF formation. High IADF stabilized frequencies (i.e. >0.50) nearly always (88%) coincide with a TC tracking into the study area, and we find a significant (p < 0.01) relationship between TC-sourced precipitation and the stabilized frequency of L+ IADFs. Via this relationship, reconstruction of historic tropical cyclone frequency and precipitation is probable, which would allow for increased understanding of historic tropical cyclone activity prior to the historic climate record. KEY WORDS: Tropical cyclone · Intra-annual density fluctuation · Dendroclimatology · Florida · Longleaf pine Full text in pdf format PreviousNextCite this article as: Mitchell TJ, Knapp PA, Ortegren JT (2019) Tropical cyclone frequency inferred from intra-annual density fluctuations in longleaf pine in Florida, USA. Clim Res 78:249-259. https://doi.org/10.3354/cr01573 Export citation RSS - Facebook - Tweet - linkedIn Cited by Published in CR Vol. 78, No. 3. Online publication date: October 17, 2019 Print ISSN: 0936-577X; Online ISSN: 1616-1572 Copyright © 2019 Inter-Research.

High responsiveness of wood anatomy to water availability and drought near the equatorial rear edge of Douglas-fir

In trees, wood anatomy is a more precise and informative measure of hydraulic responses to drought than radial growth. Tree populations located near the equatorial-range edge of their distribution (rear edge) are used to monitor responses to changes in hydroclimate; however, few studies have assessed the wood anatomy of these populations. We measured tracheid lumen diameter (LD) and cell wall thickness (CWT) in a rear-edge Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) population. We also evaluated the formation of intra-annual density fluctuations (IADFs), characterized by earlywood (EW) tracheids with narrow lumens. We expected that EW LD would be particularly sensitive to hydroclimate variability. We found that EW LD was directly related to precipitation during the previous year’s late winter and the current year’s spring. Wet conditions were associated with wider EW tracheids and improved growth. These correlations peaked when considering cumulative precipitation from January to May. EW IADFs were formed in response to dry conditions during March and April, suggesting that dry spring conditions limit tracheid expansion. The described associations in a rear-edge Douglas-fir population confirm that wood anatomy is highly responsive to drought. We conclude that EW LD is a valuable proxy for hydroclimate reconstruction in Douglas-fir.

Growth, wood anatomy and stable isotopes show species-specific couplings in three Mexican conifers inhabiting drought-prone areas

An improved understanding of how tree species will respond to warmer conditions and longer droughts requires comparing their responses across different environmental settings and considering a multi-proxy approach. We used several traits (tree-ring width, formation of intra-annual density fluctuations – IADFs, wood anatomy, Δ13C and δ18O records) to retrospectively quantify these responses in three conifers inhabiting drought-prone areas in northwestern Mexico. A fir species (Abies durangensis) was studied in a higher altitude and slightly rainier site and two pine species were sampled in a nearby, lower drier site (Pinus engelmannii, Pinus cembroides). Tree-ring-width indices (TRWi) of the studied species showed a very similar year-to-year variability likely indicating a common climatic signal. Wood anatomy analyses done over 3.5 million measured cells, showed that P. cembroides lumen area was much smaller than in the other two species and it remained constant along all the studied period (over 64 years). Instead, cell wall thickness was widest in P. engelmannii and this species presented the highest amount of intra-annual density fluctuations. Climate and wood anatomy correlations pointed out that lumen area was positively affected by winter precipitation for all studied species, while cell-wall thickness was negatively affected by this season's precipitation in all species but P. cembroides. Stable isotope analysis showed significantly lower values of Δ13C for P. cembroides and no significant δ18O differences between the three species, although they shared a common decreasing trend. With very distinct wood anatomical traits (smaller cells, compact morphology), P. cembroides stood out as the better adapted species in its current environment and could be less affected by future drier climate. P. engelmannii and A. durangensis showed high plasticity at wood anatomical level, allowing them to promptly respond to seasonal water availability but likely gives few advantages on future climate scenarios with longer and frequent drought spells.

How needle phenology indicates the changes of xylem cell formation during drought stress in Pinus sylvestris L.

Temperature, water availability and photoperiod are the primary drivers of tree phenological processes. However, there is lack of information how the spring temperature and summer drought affect phenology of both cambium and needles. We evaluated the interplay between xylem and needle development of mature Scots pine trees in three consecutive years differing in the spring temperatures and onset and duration of summer drought. Cambial activity began on the day of a year (DOY) 83–87, while the bud break occurred on DOY 113–119, when also the secondary cell wall of tracheids started its formation. While the timing of bud break correlated with the sum of effective temperatures, the beginning of cambial zone activity did not. The needles were fully unfolded around DOY 170, which corresponded to the transition between earlywood and latewood tracheids. Summer drought did not affect needle development, but it changed the rate of production and morphology of latewood tracheids. Latewood tracheids from the year 2015 when the drought was longest (i.e. until the second half of August in 2015 compared to the duration from June until the middle of July in the two other years) were 32% narrower and they had 34% thinner cell walls than in 2014 and 2016. The improvement of tree water status in July resulted in the formation of intra-annual density fluctuations (IADF). The interplay between needle and cambium phenology provided an insight into how the tree allocates the resources with varying temperature and soil water availability.

Effect of provenance and climate on intra-annual density fluctuations of Norway spruce Picea abies (L.) Karst. in Poland

Understanding the growth pattern of trees of various provenances in response to climatic conditions provides information on the adaptive potential of trees. Intra-annual density fluctuations (IADFs) are anatomical markers of variations in wood density within the annual wood increment. Latewood-like cells within earlywood, resulting from severe drought during spring and early summer, express the structural adaptation of the wood tissue to unfavorable climatic conditions. In this study, we used two 40-year old IUFRO provenance trials of Norway spruce located in the western and north-eastern Poland to i) determine the effect of provenance on the IADF frequency and ii) to define the climatic factors determining the formation of IADF. Three highly polymorphic nuclear microsatellites are used to determine the differentiation among provenances in particular between the northern and southern Polish distribution area.Our results demonstrate that the frequency of all types of IADFs is determined by the growing conditions, and only the IADFs in the transition zone are determined by provenance origin. The highest variability between provenances in response to climatic differences was observed for IADF in transition zone, where temperatures and rainfall of spring and early summer differentiated northern and southern provenances. Moreover, the IADF frequency was influenced by drought, thereby differentiating provenances in terms of their sensitivity to severe conditions. Furthermore, environmental differentiation of climate-growth reaction was expressed by a stronger impact of climatic conditions observed in the western than in the northeastern trial site.The results of this study confirm the dependency of climate related changes in wood characteristics on provenance origin. Therefore, density fluctuations seem to be an appropriate adaptive trait for selecting Norway spruce provenances with varying degree of drought sensitivity.

Wood-trait analysis to understand climatic factors triggering intra-annual density-fluctuations in co-occurring Mediterranean trees

ABSTRACTMediterranean trees and shrubs form intra-annual density fluctuations (IADFs) in tree rings as a sign of their plasticity in wood formation in response to intraseasonal variations of environmental conditions. Different species show a different aptitude to form IADFs, due to their diverse ability to cope with climate stressors, since the occurrence of IADFs may affect plant hydraulics. Dendroecology and quantitative wood anatomy were used to characterise IADFs in Pinus pinea and Arbutus unedo co-occurring at a Mediterranean site in Italy. The relations between climate parameters (i.e. temperature and precipitation) and intra-annual tree-ring traits (i. e. IADF frequency and conduit size) were analysed to highlight the main triggers for IADF formation and their functional role.Data showed that both species are characterised by a high plastic response to climate and formed a high frequency of L-IADFs (occurrence of earlywoodlike conduits in latewood). The two species, although forming the same type of IADFs, showed different sensitivity to environmental factors. Pinus pinea showed a high dependence of tracheid size on temperature, while Arbutus unedo was more sensitive to precipitation in spring and autumn. Arbutus unedo promptly developed more than one IADF per year in response to rainfall events following drought periods.The overall results were useful to compare the aptitude of the two species in forming IADFs and to highlight the factors priming their formation. This is useful to understand wood growth reactions to environmental drivers and to evaluate the adaptive capabilities in these two species, and thus to predict forest reactions after climate changes.

Modeled Tracheidograms Disclose Drought Influence on Pinus sylvestris Tree-Rings Structure From Siberian Forest-Steppe.

Wood formation allows trees to adjust in a changing climate. Understanding what determine its adjustment is crucial to evaluate impacts of climatic changes on trees and forests growth. Despite efforts to characterize wood formation, little is known on its impact on the xylem cellular structure. In this study we apply the Vaganov-Shashkin model to generate synthetic tracheidograms and verify its use to investigate the formation of intra-annual density fluctuations (IADF), one of the most frequent climate tree-ring markers in drought-exposed sites. Results indicate that the model can produce realistic tracheidograms, except for narrow rings (<1 mm), when cambial activity stops due to an excess of drought or a lack of growth vigor. These observations suggest that IADFs are caused by a release of drought limitation to cells formation in the first half of the growing season, but that narrow rings are indicators of an even more extreme and persistent water stress. Taking the example of IADFs formation, this study demonstrated that the Vaganov-Shashkin model is a useful tool to study the climatic impact on tree-ring structures. The ability to produce synthetic tracheidogram represents an unavoidable step to link climate to tree growth and xylem functioning under future scenarios.

INTRA-ANNUAL DENSITY FLUCTUATIONS IN TREE RINGS: HOW, WHEN, WHERE, AND WHY?

Intra-annual density fluctuations (IADFs) in tree rings are generally considered structural anomalies caused by deviations from the “normal course” of xylogenesis during the growing season. This definition is based on the bias that, under “normal conditions”, cambial activity stops once a year. Each tree ring can thus be dated to one calendar year, which is one of the principles of dendrochronology. The formation of IADFs can be triggered directly by environmental changes, especially in precipitation and temperature, that affect cambial activity and cell differentiation. It can also be the result of limited photosynthesis, due to defoliation induced by biotic or abiotic constraints. Often indicated with alternative terms, IADFs were first described in the 1930s, and recently reported for many trees and shrubs from different ecosystems throughout the world, particularly for Mediterranean species. Different types of IADFs have been detected; their formation and structural properties depend on many factors including tree genotype, age, size, rooting depth, habitat, soil, climate, photosynthetic activity, and allocation strategies. Whether IADFs affect the adaptive capability of plants remains, however, unclear. We provide an overview of the main anatomical features of IADFs and their occurrence in tree rings from various environments and climatic regimes. We propose a simplified way of classifying them and discuss the hypotheses about their functional role and the factors triggering their formation. To understand the ecological role of IADFs better, we recommend a multidisciplinary approach, involving wood anatomy, dendroecology, and stable isotopes, which has already been applied for Mediterranean species. We conclude by considering that IADFs appear to be the “rule” rather than “anomalies” in some ecosystems where they help plants cope with fluctuating environmental conditions. Moreover, their anatomical structure represents a valuable proxy of past climatic conditions at a sub-seasonal resolution and may be relevant to adapt hydraulic functioning of living trees to changing climatic conditions.

Timing of False Ring Formation in Pinus halepensis and Arbutus unedo in Southern Italy: Outlook from an Analysis of Xylogenesis and Tree-Ring Chronologies.

Mediterranean tree rings are characterized by intra-annual density fluctuations (IADFs) due to partly climate-driven cambial activity. IADFs are used as structural signals to gain information on relations between environmental conditions and eco-physiological processes during xylogenesis, with intra-annual resolution. To reach an unbiased synchronization of the IADF position within tree rings and seasonal fluctuations in environmental conditions, it is necessary to know the timing of cambial activity and wood formation, which are species- and site-specific processes. We applied the microcoring technique to analyze xylogenesis in Pinus halepensis and Arbutus unedo. To the best of our knowledge, this is the first attempt to study xylogenesis in a hardwood species forming frequent IADFs. Both species co-occur at a site in southern Italy characterized by a Mediterranean climate. To facilitate tree-ring dating and identification of IADFs, we performed traditional dendroecological analysis. We analyzed xylogenesis during summer, which is considered a constraint for xylogenesis and a trigger for IADF formation. We followed the different phases of cell development in the current wood increment with the aim of evaluating whether and which type of IADFs were formed. We additionally analyzed the same phases again in September and in winter to verify the possible formation of IADFs in fall and whether cell production and differentiation was completed by the end of the calendar year. Both species formed the same type of IADFs (earlywood-like cells within latewood), due to temporary growth restoration triggered by rain events during the period of summer drought. At the end of the calendar year, no cells in the phases of enlargement and secondary cell wall deposition occurred. A. unedo was more sensitive than P. halepensis because IADFs were formed earlier in the season and were more frequent in the tree-ring series. The dendro-anatomical approach, combining analysis of tree-ring series and of xylogenesis, helped to detect the period of IADF formation in the two species. Results are discussed in functional terms, highlighting the environmental conditions triggering IADFs, and also in methodological terms, evaluating the applicability of xylogenesis analysis in Mediterranean woods, especially when the formation of IADFs is not uniform around the stem.

Does the Genotype Have a Significant Effect on the Formation of Intra-Annual Density Fluctuations? A Case Study Using Larix decidua from Northern Poland.

Intra-annual density fluctuations (IADFs) can imprint environmental conditions within the growing season and most of the research on IADFs has been focused on their climatic signal. However, to our knowledge, the genetic influence on the frequency and type of IADFs has not been evaluated. To understand if the genotype can affect the formation of IADFs we have used a common garden experiment using eight families of Larix decidua established in two neighboring forest stands in northern Poland. Four types of IADFs were identified using X-ray density profiles: latewood-like cells within earlywood (IADF-type E), latewood-like cells in the transition from early- to latewood (IADF type E+), earlywood-like cells within latewood (IADF-type L), and earlywood-like cells in the border zone between the previous and present annual ring (IADF-type L+). The influence of explanatory variables i.e., families, sites, and years on identified density fluctuations was analyzed using generalized estimating equations (GEE). We hypothesized that trees from different families will differ in terms of frequency and type of IADFs because each family will react to precipitation and temperature in a different way, depending on the origin of those trees. The most frequent fluctuation was E+ and L types on both sites. The most important factors in the formation of IADFs were the site and year, the last one reflecting the variable climatic conditions, with no significant effect of the family. However, the relation between the formation of IADFs and selected climate parameters was different between families. Although, our results did not give a significant effect of the genotype on the formation of IADFs, the different sensitivity to climatic parameters among different families indicate that there is a genetic influence.

Structure and Function of Intra-Annual Density Fluctuations: Mind the Gaps.

Tree rings are natural archives of climate and environmental information with a yearly resolution. Indeed, wood anatomical, chemical, and other properties of tree rings are a synthesis of several intrinsic and external factors, and their interaction during tree growth. In particular, Intra-Annual Density Fluctuations (IADFs) can be considered as tree-ring anomalies that can be used to better understand tree growth and to reconstruct past climate conditions with intra-annual resolution. However, the ecophysiological processes behind IADF formation, as well as their functional impact, remain unclear. Are IADFs resulting from a prompt adjustment to fluctuations in environmental conditions to avoid stressful conditions and/or to take advantage from favorable conditions? In this paper we discuss: (1) the influence of climatic factors on the formation of IADFs; (2) the occurrence of IADFs in different species and environments; (3) the potential of new approaches to study IADFs and identify their triggering factors. Our final aim is to underscore the advantages offered by network analyses of data and the importance of high-resolution measurements to gain insight into IADFs formation processes and their relations with climatic conditions, including extreme weather events.

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

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

Changes in climate-growth relationships and IADF formation over time of pine species (Pinus halepensis, P. pinaster and P. sylvestris) in Mediterranean environments

Background: The Mediterranean basin has experienced an increase in the mean annual temperature, a decrease in the mean annual precipitation, and an increase in the frequency of severe drought periods during the second half of the 20th century. However, winter and spring precipitation has increased and summer precipitation has decreased in the western Mediterranean region. Aim of the study:The objectives of the present study were: i) to compare changes in climate-growth relationships over time for Pinus halepensis, P. pinaster and P. sylvestris in Spain ii) to quantify the presence of intra-annual density fluctuations (IADFs) on the three species, and iii) to define the associated climatic variables. Area of study: 26 sampling sites (8 P. halepensis sites, 8 P. pinaster sites and 10 P. sylvestris sites) were selected in their distribution area in Spain. Main results:Precipitation is the main factor influencing growth and IADF occurrence in the three species. Wet periods during previous winter and spring induced higher growth rates on P. halepensis and P. pinaster, while P. sylvestris was mostly influenced by summer precipitation. However, the influence of these climatic variables on the growth of these species changed over the studied period. The increase of winter and spring precipitation combined with increasingly harsh summer climatic conditions in the second half of the 20th century may have enhanced the importance of precipitation at the beginning of the growing season on the growth of species subject to higher summer drought stress (P. halepensis and P. pinaster) and increased IADF occurrence. Research highlights: Besides reflecting changes in the environmental conditions during the growing season, the inclusion of IADF detection in chronologies adds new information to ring-width chronologies, thereby improving its quality.Key words: Aleppo pine; maritime pine; scots pine; dendroclimatology, IADFs.Abbreviations used: IADF: Intra-annual density fluctuation; AIC: Akaike information criterion; ROC: Receiver operating characteristic.

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.