Quantifying climate–growth relationships at the stand level in a mature mixed‐species conifer forest

A range of environmental factors regulate tree growth; however, climate is generally thought to most strongly influence year‐to‐year variability in growth. Numerous dendrochronological (tree‐ring) studies have identified climate factors that influence year‐to‐year variability in growth for given tree species and location. However, traditional dendrochronology methods have limitations that prevent them from adequately assessing stand‐level (as opposed to species‐level) growth. We argue that stand‐level growth analyses provide a more meaningful assessment of forest response to climate fluctuations, as well as the management options that may be employed to sustain forest productivity. Working in a mature, mixed‐species stand at the Howland Research Forest of central Maine, USA, we used two alternatives to traditional dendrochronological analyses by (1) selecting trees for coring using a stratified (by size and species), random sampling method that ensures a representative sample of the stand, and (2) converting ring widths to biomass increments, which once summed, produced a representation of stand‐level growth, while maintaining species identities or canopy position if needed. We then tested the relative influence of seasonal climate variables on year‐to‐year variability in the biomass increment using generalized least squares regression, while accounting for temporal autocorrelation. Our results indicate that stand‐level growth responded most strongly to previous summer and current spring climate variables, resulting from a combination of individualistic climate responses occurring at the species‐ and canopy‐position level. Our climate models were better fit to stand‐level biomass increment than to species‐level or canopy‐position summaries. The relative growth responses (i.e., percent change) predicted from the most influential climate variables indicate stand‐level growth varies less from to year‐to‐year than species‐level or canopy‐position growth responses. By assessing stand‐level growth response to climate, we provide an alternative perspective on climate–growth relationships of forests, improving our understanding of forest growth dynamics under a fluctuating climate.     

Environmental controls and patterns of cumulative radial increment of evergreen tree species in montane,temperate rainforests of Chiloé Island,southern Chile

We investigated the local environmental controls on daily fluctuations of cumulative radial increment and cambial hydration of three dominant, evergreen tree species from montane, Coastal rainforests of Chiloé Island, Chile (42° 22′ S). During 2 years (1997–1998 and 1998–1999) we recorded hourly cumulative radial increments using electronic band dendrometers in the long‐lived conifer Fitzroya cupressoides (Cupressaceae), the evergreen broad‐leaved Nothofagus nitida (Nothofagaceae), and the narrow‐leaved conifer Podocarpus nubigena (Podocarpaceae). We also measured soil and cambial tissue hydration using capacitance sensors, together with air and soil temperature and rainfall during the period of the study. In addition, we collected cores of these tree species to evaluate how dendrometer measurements reflect annual tree ring width. One‐year long daily time series of cumulative radial increments suggests that radial growth of Fitzroya cupressoides was initiated slowly in early spring, with a maximum in early summer. Multiple regressions showed positive relations between daily precipitation and radial index (i.e. the difference in cumulative radial increment of two consecutive days) in the three species. According to path analysis there was a significant direct effect of changes in tree hydration on radial index of the three focal species. In emergent, pioneer species such as Nothofagus and Fitzroya, radial index was negatively affected by changes in maximum air temperature and photosynthetically active radiation, probably because of high evapotranspiration demand on warm sunny days. The shade‐tolerant species Podocarpus nubigena was positively affected by photosynthetically active radiation. Our diel scale findings support the use of tree ring widths for reconstructing past climate in these southern temperate forests and provide evidence that rainforest trees may be highly sensitive to future declines in rainfall and temperature increases during summer.     

Three-dimensional distribution of mass movement signals in disturbed Picea abies (L.) Karst.

Dendrogeomorphologic approaches enable annual absolute dating of various potentially hazardous geomorphic processes. This study tested the dating precision of three commonly used dendrogeomorphic approaches used for the extraction of geomorphic signals from the tree ring series of disturbed trees. The first approach is based on the onset of reaction wood occurrence, and the other two approaches are based on the detection of abrupt changes in tree ring eccentricity. Nine 165 cm long stems of maximally 27 year old Norway spruces (Picea abies (L.) Karst.) growing on unstable slope affected by mass movement events of known age were analysed. Several sampling directions on stems were analysed to reveal the best sampling strategy for each tested approach. The relationships between stem curvature and the dating precision of the approaches were also analysed. The results of dating precision showed crucial differences between the tested approaches. Tree stem curvature did not influence dating precision. Overall, we concluded that the approaches based on eccentric increments are less precise than those based on reaction wood analysis when juvenile P. abies trees that grow on landslide slope are sampled. The importance of this study is in providing new insight into dating precision by analysing the whole tree stems of trees influenced by mass movements. This is unique since the approaches based on eccentricity increments have not been previously tested in such a detail.     

Spatial analysis of structural and tree‐ring related parameters in a timberline forest in the Italian Alps

Abstract. We investigated the variability in spatial pattern of some structural, dendrochronological and dendroclimatological features of a mixed Larix decidua‐Pinus cembra forest at the timberline in the eastern Italian Alps at fine geographical and temporal scales. Forest structure variables such as stem diameter, tree height, age and tree‐ring related parameters (yearly growth index, mean sensitivity, first order autocorrelation and some dendroclimatic variables) have been compared at various scale levels. We observed that most of the variables show positive autocorrelated structures due to both forest dynamics and fine‐scale driving forces, probably related to microrelief. Spatial structure of yearly indexed radial growth appears sensitive to extreme climatic events. Secondary succession after past disturbances drives the forest towards a structure governed by a gap regeneration dynamics that seems to ensure the different requirements of the two main tree species present. Small spatial scale studies of forest structures, especially if integrated to dendro‐ecological data, seem an efficient tool to assess the disturbance regime and species sensitivity to environmental change.     

Sea ice predicts long‐term trends in Adélie penguin population growth,but not annual fluctuations: Results from a range‐wide multiscale analysis

Understanding the scales at which environmental variability affects populations is critical for projecting population dynamics and species distributions in rapidly changing environments. Here we used a multilevel Bayesian analysis of range‐wide survey data for Adélie penguins to characterize multidecadal and annual effects of sea ice on population growth. We found that mean sea ice concentration at breeding colonies (i.e., “prevailing” environmental conditions) had robust nonlinear effects on multidecadal population trends and explained over 85% of the variance in mean population growth rates among sites. In contrast, despite considerable year‐to‐year fluctuations in abundance at most breeding colonies, annual sea ice fluctuations often explained less than 10% of the temporal variance in population growth rates. Our study provides an understanding of the spatially and temporally dynamic environmental factors that define the range limits of Adélie penguins, further establishing this iconic marine predator as a true sea ice obligate and providing a firm basis for projection under scenarios of future climate change. Yet, given the weak effects of annual sea ice relative to the large unexplained variance in year‐to‐year growth rates, the ability to generate useful short‐term forecasts of Adélie penguin breeding abundance will be extremely limited. Our approach provides a powerful framework for linking short‐ and longer term population processes to environmental conditions that can be applied to any species, facilitating a richer understanding of ecological predictability and sensitivity to global change.     

Climate-growth relationships for Rocky Mountain juniper (Juniperus scopulorum Sarg.) on the volcanic badlands of western New Mexico,USA

We sampled Rocky Mountain junipers (RMJ) to produce a multi-century tree-ring chronology from a relict lava flow, the Paxton Springs Malpais (PAX), in the Zuni Mountains of western New Mexico. Our objective was to assess crossdating potential for RMJ growing on the volcanic badlands of the region, investigate potential relationships between climate and RMJ growth, and investigate temporal variability in relationships identified between climate and RMJ growing at our site. We hypothesized that, similar to other drought stressed-conifers growing on the lava flows, RMJ responds to climate factors that influence and indicate moisture availability. We found a high average mean sensitivity value (0.53), which indicated the PAX chronology exhibited enough annual variability to capture fluctuations in environmental conditions. The average interseries correlation (0.74) indicated confident crossdating and a significant association of annual growth among trees within the stand. The positive correlation between the PAX chronology and total precipitation for the local water year was significant (r = 0.53; P < 0.001). Significant positive correlations also were identified between monthly PDSI, monthly total precipitation, and RMJ radial growth. Analyses of temporal stability indicated that the positive relationship between RMJ growth at the PAX site and monthly PDSI was the most stable relationship during the period of analysis (1895–2007). More importantly, we identified a unique inverse relationship between radial growth and monthly mean temperature during periods of the preceding year and current growing year, the first such finding of a strong temperature response for a low-mid elevation tree species in the American Southwest. Our results confirm that RMJ samples collected on the Paxton Springs Malpais are sensitive to climate factors that affect moisture availability, further suggesting that RMJ may be suitable for use in dendroclimatic research at additional locations across the broad distribution of the species.     

Changing environmental spectra influence age-structured populations: increasing ENSO frequency could diminish variance and extinction risk in long-lived seabirds

As global climate changes, there is increasing need to understand how changes in the frequencies of environmental variability affect populations. Age-structured populations have recently been shown to filter specific frequencies of environmental variability, favoring generational frequencies, and very low frequencies, a phenomenon known as cohort resonance. However, there has been little exploration of how changes in the spectra of environmental signals will affect the stability and persistence of age-structured populations. To examine this issue, we analyzed a likely example to show how changes in the frequency of an influential climate phenomenon, the El Niño-Southern Oscillation (ENSO), could affect a marine bird population. We used a density-dependent, age-structured population model to calculate the transfer function (i.e., the frequency-dependent sensitivity) of Brandt’s cormorant (Phalacrocorax penicillatus), a representative marine bird species known to be influenced by ENSO. We then assessed how the population would be affected by ENSO forcing that was doubled and halved in frequency. The transfer function indicated this population is most sensitive to variance at low frequencies, but does not exhibit the sensitivity to generational frequencies (cohort resonance) observed in shorter-lived species. Doubling the frequency of ENSO unexpectedly resulted in higher mean adult population abundance, lower variance, and lower probability of extinction, compared to forcing with the historical or reduced ENSO frequency. Our results illustrate how long-lived species with environmentally driven variability in recruitment, including many species of marine birds and fish, may respond in counterintuitive ways to anticipated changes in environmental variability.     

Does a heavy fruit crop reduce the tree ring increment? Results from a 12-year study in a subalpine zone

Life history theory posits that an increased investment in reproduction should result in decreased vegetative investment. Switching resources from growth to reproduction are also expected in species experiencing selection pressure for high variation of seed crops. In this study, we tested whether the reproductive effort of trees, measured as the number of fruits produced, is related to their radial growth increment. We examined a population of Sorbus aucuparia, a fleshy-fruited tree species with highly variable interannual individual fruit production growing in the subalpine zone and under strong selection pressure from pre-dispersal seed predators. We used 12-year data to test the relationship between fruit crop and radial growth increments in current, previous and subsequent years, and found no trade-off between growth and reproduction. For almost all trees, there was no correlation between fruit crop and radial growth increment in the same year or next year. Only a few trees showed a positive correlation between fruit crop and previous-year growth. In the statistics, we took advantage of the high variability of individual fruit crops in high production years. In four heavy crop years, we compared the radial growth increments of trees with heavy crops with those of trees with low fruit crops. Current and next-year radial growth did not differ between trees with low and high fruit crops. In all those years, however, trees having heavy fruit crops had higher previous-year growth increments. We suggest that the harsh subalpine weather conditions account for the lack of a trade-off between growth and reproduction in the studied population.     

High resolution radiocarbon spike confirms tree ring dating with low sample depth

Radiocarbon (¹⁴C) has been used to date carbon-rich objects in Earth science, archeology, and history since the 1940s. New methods, using spikes in ¹⁴C caused by solar proton events, can be used to annually date wood when crossdating is not possible, such as when sample size is low, samples are floating in time, or external disturbances lead to insecure dates. Here, we use a spike in radiocarbon during a solar energetic particle (SEP) event in 774/775 CE to confirm crossdating of a poorly-replicated King Billy pine (Athrotaxis selaginoides) chronology. Low sample depth between 1498 and 1523 CE (two trees) prevented confident dating of the early period of the chronology. Three core samples with strong correlation with the master chronology that likely included the 774/775 CE Miyake SEP event were identified for radiocarbon isotope analysis. We sectioned segments centered on the estimated 774/775 CE date and then isolated the holocellulose in each sample. Samples were sent to an accelerator mass spectrometry (AMS) for radiocarbon measurements. The AMS data confirmed the crossdating accuracy of the tree ring series and reinforces the applicability of this technique to anchor poorly dated tree ring series in time. In addition, we found sample processing with a microtome proved superior for holocellulose extractions and yielded more accurate ¹⁴C measurements. We recommend sampling with a microtome, processing at least three samples per year, and including sample masses greater than 100 ug C to confirm dating using radiocarbon spikes.     

A growth study of Prioria copaifera (Caesalpinaceae) using dendrochronological techniques

The Cativo (Prioria copaifera) forms very homogeneous forests called cativales in the flooded plains of some rivers from Costa Rica to Colombia. For over 70 years Cativo has been the main base of the timber industry in the Colombian Darien area. Because of high productivity and high-dominance of Cativo trees, they represent one of the most prone tropical forests for sustainable forest management. The objective of this research is to model diameter and timber volume growth and growth rates (absolute, mean and relative) of Cativo as a function of age, using tree ring data derived from dendrochronologycal techniques. We evaluated the annual nature of the tree rings by radiocarbon analysis and crossdating techniques. Besides, the diameter and volume growth was modeled using von Bertalanffy's model. As of our results, we estimated the life span of Cativo in 614 years as the time required to reach 99% of the asymptotic diameter. By the mean value we have found that the mean rate of diameter growth is 0.31cm/y. The species requires 90 years to reach 40cm in diameter, the regulated cut diameter in Colombia. We find that Cativo reaches maximum current annual increment (ICA) in diameter at 40 years and in volume at 90 years with rates of 0.5cm/y and 0.032m3/y per tree, respectively. The maximum diameter mean annual increments (MAI) are achieved at 80 years and for the volume at 140 year, with growth rates of 0.45cm/y and 0.018m3/y per tree, respectively. The generated information is useful for the sustainable management of Cativò forests.     

Frequency of partial and missing rings in Acer saccharum in relation to canopy position and growth rate

Locally absent growth rings are known to occur in trees during periods of environmental stress, but little evidence has been available on their frequency in moist temperate forests or on their potential for causing problems in dendroecological studies. In this study, 95 stem disks of Acer saccharum cut from trees representing a wide range of size and age were examined for ring anomalies using three techniques – inspection of disks for partial rings, crossdating of each tree against regional master chronologies, and use of stand history evidence. The number of ring anomalies in Acer saccharum was inversely related to growth rate and vigor. Mean percentage of ring anomalies was 1.3% in dominant canopy trees, 4.6% in trees of the intermediate crown class, and 16.2% in overtopped trees. All of the overtopped trees had partial or missing rings, with a mean of 10 per tree and a range of 2 to 20. Use of crossdating to correct for ring anomalies appeared to be successful in 88% of the canopy trees. However, crossdating results were inconclusive in 32% of the overtopped trees because of low ring width variability, multiple missing rings, and short duration of trouble-free segments. Because overtopped trees are of limited value in reconstructing disturbance history and often cannot be crossdated reliably, we suggest that they normally be excluded from disturbance chronologies.     

Temporal resolutions in species distribution models of highly mobile marine animals: Recommendations for ecologists and managers

While ecologists have long recognized the influence of spatial resolution on species distribution models (SDMs), they have given relatively little attention to the influence of temporal resolution. Considering temporal resolutions is critical in distribution modelling of highly mobile marine animals, as they interact with dynamic oceanographic processes that vary at time‐scales from seconds to decades. We guide ecologists in selecting temporal resolutions that best match ecological questions and ecosystems, and managers in applying these models. We group the temporal resolutions of environmental variables used in SDMs into three classes: instantaneous, contemporaneous and climatological. We posit that animal associations with fine‐scale and ephemeral features are best modelled with instantaneous covariates. Associations with large scale and persistent oceanographic features are best modelled with climatological covariates. Associations with mesoscale features are best modelled with instantaneous or contemporaneous covariates if ephemeral processes are present or interannual variability occurs, and climatological covariates if seasonal processes dominate and interannual variability is weak.     

Climate increases regional tree-growth variability in Iberian pine forests

Tree populations located at the geographical distribution limit of the species may provide valuable information about tree‐growth response to changes on climatic conditions. We established nine Pinus nigra, 12 P. sylvestris and 17 P. uncinata tree‐ring width chronologies along the eastern and northern Iberian Peninsula, where these species are found at the edge of their natural range. Tree‐growth variability was analyzed using principal component analysis (PCA) for the period 1885–1992. Despite the diversity of species, habitats and climatic regimes, a common macroclimatic signal expressed by the first principal component (PC1) was found. Moreover, considering the PC1 scores as a regional chronology, significant relations were established with Spanish meteorological data. The shared variance held by the tree chronologies, the frequency of narrow rings and the interannual growth variability (sensitivity) increased markedly during the studied period. This shows an enhancement of growth synchrony among forests indicating that climate might have become more limiting to growth. Noticeably, an upward abrupt shift in common variability at the end of the first half of the 20th century was detected. On the other hand, moving‐interval response functions showed a change in the growth–climate relationships during the same period. The relationship between growth and late summer/autumn temperatures of the year before growth (August–September, negative correlation, and November, positive correlation) became stronger. Hence, water stress increase during late summer previous to tree growth could be linked to the larger growth synchrony among sites, suggesting that climate was driving the growth pattern changes. This agrees with the upward trend in temperature observed in these months. Moreover, the higher occurrence of extreme years and the sensitivity increase in the second half of the 20th century were in agreement with an increment in precipitation variability during the growing period. Precipitation variability was positively related to tree‐growth variability, but negatively to radial growth. In conclusion, a change in tree‐growth pattern and in the climatic response of the studied forests was detected since the mid‐20th century and linked to an increase in water stress. These temporal trends were in agreement with the observed increase in warmer conditions and in precipitation variability.     

Area, shape and isolation of tropical forest fragments: effects on tree species diversity and implications for conservation

Aim To quantify the influences of forest area, shape and isolation on tree species diversity in Ghana and to compare their significance with the influences of climate (average annual rainfall) and disturbance (fire burn, logging, agriculture). Location The forest zone of southern Ghana, West Africa (between 5 and 8° N). Methods For twenty‐two forest fragments (1) bivariate regression analyses of tree species diversity (number and composition) were employed with forest spatial geometry, climate and disturbance variables. (2) Multivariate regression analyses of tree species number and all seven environmental variables were used to determine the variability in tree species number that could be accounted for by these environmental variables. Results Forest area, shape and isolation accounted for sharply decreasing proportions of variability in tree species diversity. Large forest fragments contained the greatest numbers of tree species and the highest proportions of rare tree species; irregular fragments had high proportions of regenerating, light‐demanding pioneers and mature, animal‐dispersed species and isolated fragments were floristically similar to less isolated fragments. Fire burn and average annual rainfall accounted for small, but nevertheless significant, proportions of variability in tree species diversity. Logging and agriculture were non‐significant variables. Main conclusions (1) Forest area is the most important consideration when planning tropical forest reserves. (2) Management of disturbance should take priority over management of forest shape if higher levels of tree diversity and species quality are to be maintained. (3) If new reserves are to be designated, they should be located within different climatic zones in order to capture a large fraction of the regional biota. (4) Biogeographers have an important role to play in formulating and testing hypotheses at a broad spatial scale and ultimately, informing conservation management within the tropical biome.     

Crossdating Juniperus procera from North Gondar, Ethiopia

The application of dendrochronology in (sub)tropical regions has been limited by the difficulty in finding trees with distinct annual rings that can be crossdated. Here, we report successful crossdating of Juniperus procera trees from North Gondar, Ethiopia. The trees form annual rings in response to a unimodal rainfall regime. The selection of mesic locations ensured that the trees did not respond to intra-seasonal weather anomalies. Crossdating was achieved by comparison of the wood anatomy directly on the surface of the core samples and purpose-adapted skeleton plotting. Wood-anatomical anomalies, such as false and indistinct rings, were regarded as potentially replicated features and used in crossdating. COFECHA yielded site-specific mean series inter-correlations between 0.52 and 0.59. AMS radiocarbon dating during the bomb era indicated that dating uncertainty is ±1 year.     

Preboreal oscillations inferred from Arctica islandica sclerochronology

An increasingly important source of annually resolved palaeoenvironmental proxy data originates from cross-dated incremental chronologies summarizing the shell growth of several individuals. Here, we have analysed annual increment variations in a collection of radiocarbon-dated shells of ocean quahog (Arctica islandica) from early Holocene prodelta deposits in north Norway. Radiocarbon dating of the shell material showed that the increments were formed contemporaneously during Preboreal times. The biologically youngest shell contains 35 annual increments, whereas the oldest shell shows 169 increments. Time-series of annual increments demonstrated clear age trends with the widest increments during the very early years of bivalve life, followed by a notable decline in increment widths as the bivalves aged. Subsequent to removing these biological trends from the series, a sclerochronological cross-dating was carried out and resolved the temporal alignments of the shell growth increment records relative to each other. The resulting shell growth increment chronology evidences vigorous growth variations. Spectral analysis of the chronology revealed 3.7- and 4.3-year periodicities, indicative of Preboreal environmental oscillations. Periodicities of longer period were not detected. Our results prove the value of radiocarbon-dated shell assemblages to build “floating” geochronologies for periods and regions where dead shells from museum collections or seabed are not obtainable. Increasing constructions of such chronologies enhance the potential of sclerochronological cross-dating of annual shell growth increment chronologies to depict and detail annually-resolved climate variability not only for late Holocene, as previously illustrated, but also for early Holocene times, when large-scale oscillations punctuated the global climate dynamics. Development of longer chronologies with higher sample replication remains an attainable interdisciplinary target.     

Interspecific synchrony of seabird population growth rate and breeding success

Environmental variability can destabilize communities by causing correlated interspecific fluctuations that weaken the portfolio effect, yet evidence of such a mechanism is rare in natural systems. Here, we ask whether the population dynamics of similar sympatric species of a seabird breeding community are synchronized, and if these species have similar exceptional responses to environmental variation. We used a 24‐year time series of the breeding success and population growth rate of a marine top predator species group to assess the degree of synchrony between species demography. We then developed a novel method to examine the species group – all species combined – response to environmental variability, in particular, whether multiple species experience similar, pronounced fluctuations in their demography. Multiple species were positively correlated in breeding success and growth rate. Evidence of “exceptional” years was found, where the species group experienced pronounced fluctuations in their demography. The synchronous response of the species group was negatively correlated with winter sea surface temperature of the preceding year for both growth rate and breeding success. We present evidence for synchronous, exceptional responses of a species group that are driven by environmental variation. Such species covariation destabilizes communities by reducing the portfolio effect, and such exceptional responses may increase the risk of a state change in this community. Our understanding of the future responses to environmental change requires an increased focus on the short‐term fluctuations in demography that are driven by extreme environmental variability.     

Linking drought legacy effects across scales: From leaves to tree rings to ecosystems

Severe drought can cause lagged effects on tree physiology that negatively impact forest functioning for years. These “drought legacy effects” have been widely documented in tree‐ring records and could have important implications for our understanding of broader scale forest carbon cycling. However, legacy effects in tree‐ring increments may be decoupled from ecosystem fluxes due to (a) postdrought alterations in carbon allocation patterns; (b) temporal asynchrony between radial growth and carbon uptake; and (c) dendrochronological sampling biases. In order to link legacy effects from tree rings to whole forests, we leveraged a rich dataset from a Midwestern US forest that was severely impacted by a drought in 2012. At this site, we compiled tree‐ring records, leaf‐level gas exchange, eddy flux measurements, dendrometer band data, and satellite remote sensing estimates of greenness and leaf area before, during, and after the 2012 drought. After accounting for the relative abundance of tree species in the stand, we estimate that legacy effects led to ~10% reductions in tree‐ring width increments in the year following the severe drought. Despite this stand‐scale reduction in radial growth, we found that leaf‐level photosynthesis, gross primary productivity (GPP), and vegetation greenness were not suppressed in the year following the 2012 drought. Neither temporal asynchrony between radial growth and carbon uptake nor sampling biases could explain our observations of legacy effects in tree rings but not in GPP. Instead, elevated leaf‐level photosynthesis co‐occurred with reduced leaf area in early 2013, indicating that resources may have been allocated away from radial growth in conjunction with postdrought upregulation of photosynthesis and repair of canopy damage. Collectively, our results indicate that tree‐ring legacy effects were not observed in other canopy processes, and that postdrought canopy allocation could be an important mechanism that decouples tree‐ring signals from GPP.     

Overcoming crossdating challenges to generate ring-width chronologies for Sequoia sempervirens across its native range

We utilize a large dendrochronological dataset for Sequoia sempervirens of 1.29 million rings from 5952 series for 278 trees to generate location and regional ring-width chronologies and to investigate constraints on crossdating. Increment cores were collected at regular height intervals along the trunk via free-hanging ropes with some terrestrial deadwood sampled. Inter-annual chronologies span 86–1687 yr for 47 locations in primary and secondary forests across the native Sequoia range. Shared signals based on declining strength of paired correlations with distance, strong correlations within regions, and groupings of principal components guide chronology creation for northern (1032 yr, 108 trees) and southern (865 yr, 52 trees) regions and four sub-regions. The suite of chronologies provides location-specific references as well as more robust regional indices that reflect distinct signals along a climatically-integrative latitudinal gradient. Secondary forests date to near completion, and we use a subset of data to assess crossdating for trees in primary forests. Among tree and landscape attributes, old age is the most important predictor of undated and missing rings, and undated rings align with periods of reduced biomass production distributed across centuries-long lifespans. Crossdating metrics vary by height above the ground with breast height (BH) and 10 m samples having more undated rings than upper samples. At 10 m, samples have the most missing rings. Buttressing affects the lower trunk as BH series have higher ring widths and lower interseries correlations compared to those up to 70 m. Whole-trunk sampling maximizes success for crossdating old, structurally-complex Sequoia and enables dating of crown damage and assessment of biomass dynamics related to environmental change. Reference chronologies, regional assessments, and crossdating insights collectively guide future study of Sequoia for applications including climatology, fire ecology, and seismology.     

Identification,measurement and interpretation of tree rings in woody species from mediterranean climates

We review the literature dealing with mediterranean climate, vegetation, phenology and ecophysiology relevant to the understanding of tree-ring formation in mediterranean regions. Tree rings have been used extensively in temperate regions to reconstruct responses of forests to past environmental changes. In mediterranean regions, studies of tree rings are scarce, despite their potential for understanding and predicting the effects of global change on important ecological processes such as desertification. In mediterranean regions, due to the great spatio-temporal variability of mediterranean environmental conditions, tree rings are sometimes not formed. Often, clear seasonality is lacking, and vegetation activity is not always associated with regular dormancy periods. We present examples of tree-ring morphology of five species (Arbutus unedo, Fraxinus ornus, Quercus cerris, Q. ilex, Q. pubescens) sampled in Tuscany, Italy, focusing on the difficulties we encountered during the dating. We present an interpretation of anomalies found in the wood structure and, more generally, of cambial activity in such environments. Furthermore, we propose a classification of tree-ring formation in mediterranean environments. Mediterranean tree rings can be dated and used for dendrochronological purposes, but great care should be taken in selecting sampling sites, species and sample trees.