Ancient juniper trees growing on cliffs: toward a long Mediterranean tree-ring chronology

Juniperus phoenicea is a tree that can grow on vertical cliff faces in dry and warm Mediterranean climate conditions. These trees are adapted to extreme growing conditions where the main constraints are verticality, compact hard limestone, and low water supply. They respond to these constraints via various specific features and high longevity. The objective of this study is to confirm whether or not their tree-rings are annual in order to specify growth strategies and accurately date these trees. Trunk morphology, anatomical wood anomalies and radial growth were analyzed on 53 trees in the Ardèche canyon. Crossdating of the ring widths using traditional dendrochronological techniques was unsuccessful, so radiocarbon dating of tree pith was used to assess tree age, and wiggle-match dating was used to test for differences between number of rings counted and radiocarbon dates. Radiocarbon dates span the period 2520–685 BP. Minimal difference between radiocarbon dates and ring counts was apparently small—missing rings occur, but not in large numbers. Tree-ring formation is annual and radial growth is low, which creates stunted old trees. Such old living trees are uncommon in the Mediterranean basin, especially at low elevation. They can provide long tree-ring chronologies back to 792–524 cal BC. Results from the radiocarbon dating indicate that accurate annual dating of these rings may be possible by crossdating. J. phoenicea growing on cliffs offer a valuable model to better understand cliff population ecology and the functional responses of trees that can live in harsh environmental conditions.     

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

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

Mediterranean dwarf shrubs and coexisting trees present different radial-growth synchronies and responses to climate

Due to their diversity and dominance in environmentally harsh sites, Mediterranean dwarf shrubs are a valuable tool to understand the consequences of climatic variability on radial growth in woody plants. We evaluate the dendrochronological potential of three Mediterranean dwarf shrubs versus three coexisting tree species inhabiting cold- (Hormathophylla spinosa vs. Pinus sylvestris), mesic- (Ononis fruticosa vs. Abies alba), and xeric sites (Linum suffruticosum vs. Pinus halepensis). Cross-sectional wood sections of the three shrub species and cores in the case of trees were visually cross-dated and ring-widths were measured and converted into residual growth indices. We used linear mixed-effects models to assess how growth indices respond to local factors and climatic variables. The radial growth of the three dwarf shrub species was more asynchronous, i.e., ring-width series differed among conspecific individuals, than that of coexisting tree species. Growth asynchrony was higher for H. spinosa than for O. fruticosa and L. suffruticosum. Similarly, the ring-width series of O. fruticosa and L. suffruticosum was strongly correlated with that of coexisting tree species, while growth series of H. spinosa and P. sylvestris was not related at all. The growth of the three dwarf shrub species was influenced by the regional climatic conditions, but to a lesser degree than coexisting tree species. The highest responsiveness of growth to climate was observed in Mediterranean dwarf shrubs from xeric sites. However, local conditions are also major drivers of growth in Mediterranean dwarf shrubs as indicated by the stronger asynchrony in ring formation of these species as compared with coexisting trees, particularly in cold sites.     

Does sex matter? Gender-specificity and its influence on site-chronologies in the common dioecious shrub Juniperus communis

In recent years an increasing number of studies have shown shrubs to be reliable proxies of environmental conditions in regions where Trees − due to harsh climate conditions − are absent. Although many shrubs are monoecious, some are dioecious, which poses certain questions related to gender-specific growth as observed for trees in previous studies. Here, we address the questions whether dioecious shrubs, similar to trees, show growth differences between male and female plants, and − if so − whether this difference needs to be considered in terms of sample selection. We chose Juniperus communis. L., the most widely distributed woody plant, and a common and well-studied dioecious shrub species in the northern hemisphere, especially in the Boreal, Subarctic tundra and Alpine regions. Our samples were collected from four sites − three from the Ural Mountains and one site from Kirkenes in Norway. To see if there were differences in radial growth between sexes we performed four different analyses. First, we used multivariate explorative statistics to see if there were gender biased sub-populations and generally found no differences. Secondly, to compare growth over the lifetime of shrubs we computed cumulative annual increments of basal area which revealed no gender-specific growth patterns. Thirdly, to test if differences in radial growth between male and female shrubs affect the resulting site-chronology, we compared individual shrub chronologies with the site-chronology and found a significant differentiation between normalized correlations of gender-specific chronologies to the site-chronology. This significant difference was restricted to an overall comparison, but not evident at individual site-level. Lastly, we compared correlations of gender-specific chronologies and a mean site-chronology with monthly climate records to find only very few meaningful differences in their responses. In summary, we could not detect any clear gender-specific growth pattern in Juniperus communis but observed a trend towards more non-climatic signals in female junipers which may affect the resulting site-chronology.     

Age and Long-term Growth of Trees in an Old-growth Tropical Rain Forest, Based on Analyses of Tree Rings and 14C1

In an old‐growth tropical wet forest at La Selva, Costa Rica, we combined radiocarbon (¹⁴C) dating and tree‐ring analysis to estimate the ages of large trees of canopy and emergent species spanning a broad range of wood densities and growth rates. We collected samples from the trunks of 29 fallen, dead individuals. We found that all eight sampled species formed visible growth rings, which varied considerably in distinctiveness. For five of the six species for which we combined wood anatomical studies with ¹⁴C‐dates (ring ages), the analyses demonstrated that growth rings were of annual formation. The oldest tree we found by direct ring counting was a Hymenolobium mesoamericanum Lima (Papilionaceae) specimen, with an age of ca. 530 years at the time of death. All other sampled individuals, including very large trees of slow‐growing species, had died at ages between 200 and 300 years. These results show that, even in an everwet tropical rain forest, tree growth of many species can be rhythmic, with an annual periodicity. This study thus raises the possibility of extending tree‐ring analyses throughout the tropical forest types lacking a strong dry season or annual flooding. Our findings and similar measurements from other tropical forests indicate that the maximum ages of tropical emergent trees are unlikely to be much greater than 600 years, and that these trees often die earlier from various natural causes.     

Thirty years of in situ tree growth under elevated CO2: a model for future forest responses?

Rising concentrations of atmospheric carbon dioxide have been predicted to stimulate the growth of forest trees. However, long-term effects on trees growing to maturity and to canopy closure while exposed to elevated CO₂ have never been examined. We compared tree ring chronologies of Mediterranean Quercus ilex which have been continuously exposed to elevated CO₂ (around 650 μmol mol^–1) since they were seedlings, near two separate natural CO₂ springs with those from trees at nearby ambient-CO₂‘control’ sites. Trees grown under high CO₂ for 30 years (1964–93) showed a 12% greater final radial stem width than those growing at the ambient-CO₂ control sites. However, this stimulation was largely due to responses when trees were young. By the time trees were 25–30 y old the annual difference in tree ring width between low and high CO₂ grown trees had disappeared. At any given tree age, elevated CO₂ had a relatively greater positive effect on tree ring width in years with a dry spring compared to years with more rainfall between April and May. This indicates a beneficial effect of elevated CO₂ on tree water relations under drought stress. Our data suggest that the early regeneration phase of forest stands can be accelerated in CO₂-enriched atmospheres and that maximum biomass per land area may be reached sooner than under lower CO₂ concentrations. In our study, high CO₂ grown Q. ilex trees reached the same stem basal area at the age of 26 y as control trees at 29 y, i.e. three years earlier (faster turnover of carbon?). Reliable predictions of the future development of forests need to account for the variable responses of trees over their entire lifetime. Such responses to elevated CO₂ can presently only be assessed at such unique field sites.     

Competition and drought limit the response of water-use efficiency to rising atmospheric carbon dioxide in the Mediterranean fir <Emphasis Type="Italic">Abies pinsapo</Emphasis>

The gas-exchange and radial growth responses of conifer forests to climatic warming and increasing atmospheric CO₂ have been widely studied. However, the modulating effects of variables related to stand structure (e.g., tree-to-tree competition) on those responses are poorly explored. The basal-area increment (BAI) and C isotope discrimination (C stable isotope ratio; δ¹³C) in the Mediterranean fir Abies pinsapo were investigated to elucidate the influences of stand competition, atmospheric CO₂ concentrations and climate on intrinsic water-use efficiency (WUEi). We assessed the variation in δ¹³C of tree-rings from dominant or co-dominant trees subjected to different degrees of competition. A high- (H) and a low-elevation (L) population with contrasting climatic constraints were studied in southern Spain. Both populations showed an increase in long-term WUEi. However, this increase occurred more slowly at the L site, where a decline of BAI was also observed. Local warming and severe droughts have occurred in the study area over the past 30 years, which have reduced water availability more at lower elevations. Contrastingly, trees from the H site were able to maintain high BAI values at a lower cost in terms of water consumption. In each population, trees subjected to a higher degree of competition by neighboring trees showed lower BAI and WUEi than those subjected to less competition, although the slopes of the temporal trends in WUEi were independent of the competitive micro-environment experienced by the trees. The results are consistent with an increasing drought-induced limitation of BAI and a decreasing rate of WUEi improvement in low-elevation A. pinsapo forests. This relict species might not be able to mitigate the negative effects of a decrease in water availability through a reduction in stomatal conductance, thus leading to a growth decline in the more xeric sites. An intense and poorly asymmetric competitive environment at the stand level may also act as an important constraint on the adaptive capacity of these drought-sensitive forests to climatic warming. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The age of monumental olive trees (Olea europaea) in northeastern Spain

Trees can reach ages that in some cases amount to thousands of years. In the Mediterranean region, olive trees (Olea europaea) have traditionally been considered a particularly long-lived species. The main objective of this study was to assess the age of large olive trees considered to be millenarian and classified as monumental trees in northeastern Spain. We extracted cores of 14 individuals and obtained 8 sections of trees which had already been cut in the area where the largest olive trees in the northeastern Iberian Peninsula are found. The age of the sampled olive trees was assessed by counting the number of annual growth rings. Tree rings did not cross-date well, neither within nor between individuals, but boundaries between likely annual rings were clearly distinct. We found a linear relationship between DBH and tree age (in years) (Age = 2.11 × diameter(cm) + 88.93, R² = 0.80), which was used to estimate the age of unsampled olive trees. The maximum estimated age (627 ± 110 years) is among the greatest ages reported for olive trees around the world (700 years) and among the oldest trees in Mediterranean ecosystems.     

Ageing culturally significant relic trees in southeast Queensland to support bushfire management strategies

Appropriate fire management strategies are needed to protect forests and large old ecologically and culturally significant trees in natural landscapes. The aim of this study was to determine the age of large old and relic trees of cultural significance that included Cypress Pine (Callitris columellaris F. Muell.), a species that is sensitive to crown scorching fires in a fire‐prone landscape, and to calibrate a tree‐growth‐rate method for estimating tree age. Twelve large trees were dated using radiocarbon (¹⁴C) dating. The trees are located on North Stradbroke Island (Indigenous name: Minjerribah), southeast Queensland (Australia) in a fire‐prone landscape where recent wildfires have destroyed many large trees. The median tree ages ranged from 155 to 369 years. These results suggest an important role of past Indigenous land management practices in protecting Cypress Pine from crown scorching fires. The tree‐growth‐rate‐based method for estimating tree age generally overestimated the age derived from radiocarbon dating. Bias correction factors were developed for correcting various measures of periodic growth rates. This study provides evidence that appropriate low‐intensity fire strategies have the potential to contribute to the survival of forests and conserve large old trees.     

西藏东部主要建群树种径向生长对极端干旱的响应差异

不同坡向、不同海拔树木生长对极端干旱事件的响应可能不同,然而这方面的认识不足。为此,选取西藏东部珠角拉山阴、阳坡的建群树种川西云杉 (Picea likiangensis var. rubescens) 和大果圆柏 (Juniperus tibetica),在不同海拔高度建立了树轮宽度年表,分析了径向生长的气候响应,以及对极端干旱事件的抵抗力和恢复力。结果表明:阳坡大果圆柏和阴坡川西云杉的树木生长对气候的响应存在相似性,均与前一年3-6月、11-12月气温显著正相关,与当年4-5月气温显著负相关,与当年4-5月降水和帕尔默干旱指数 (PDSI, Palmer Drought Severity Index) 显著正相关 (P <0.05)。阳坡大果圆柏的抵抗力显著低于阴坡川西云杉。随着海拔升高,阴坡川西云杉树木个体的抵抗力和恢复力均显著提高,而阳坡大果圆柏树木个体抵抗力、恢复力在不同海拔无显著区别。结合混合效应模型表明树木抵抗力主要受当年4-5月平均最高气温限制,树木恢复力主要受干旱事件后四年4-5月平均最高气温限制 (P <0.01),说明生长季高温引起的极端干旱是树木径向生长下降的主要原因。     

Seasonal precipitation and continentality drive bimodal growth in Mediterranean forests

Tree phenology is sensitive to climate warming and changes in seasonal precipitation. Long xylogenesis records are scarce, thus limiting our ability to analyse how radial growth responds to climate variability. Alternatively, process-based growth models can be used to simulate intra-annual growth dynamics and to better understand why growth bimodality varies along temperature and precipitation gradients. We used the Vaganov-Shashkin (VS) growth model to analyse the main climatic drivers of growth bimodality in eight trees and shrubs conifers (four pines and four junipers) across Spain. We selected eleven sites with different continentality degree and spring/autumn precipitation ratios since we expected to find pronounced bimodal growth in less continental sites with spring and autumn precipitation peaks. The VS model successfully simulated annual growth rates at all sites as a function of daily temperature and soil moisture data. Bimodal growth patterns clustered into less continental sites showing low spring/autumn precipitation ratios. This finding agrees with observed climate-growth associations showing that growth was enhanced by wet-cool winter-to-spring conditions, but also by wet autumn conditions in the most bimodal sites. We observed a stronger growth bimodality in pines compared to junipers. We discuss the spatial variability of climate drivers in bimodality growth pattern and how increasing continentality and shifts in seasonal precipitation could affect growth patterns. Bimodality could be an advantageous response to overcome summer drought in Mediterranean forests. The ability of some species to reactivate growth during autumn might determine their capacity to withstand increasing summer aridity.     

Arboreal and prostrate conifers coexisting in Mediterranean high mountains differ in their climatic responses

In contrast to most high elevation areas, plant growth at Mediterranean mountains is exposed to a summer drought period, which represents an additional climatic constraint to low temperatures. Although arboreal and shrubby conifers coexist at high altitudes, most dendroecological studies have focused on climatic responses of tree species, whereas those of shrubby species remain mostly unexplored. We built tree-ring width chronologies for two conifer species, a shrub (Juniperus sabina) and a tree (Pinus sylvestris), coexisting at three high-altitude localities of the Iberian System mountains, eastern Spain. We analyzed their climate–growth relationships for the period 1950–2009 using correlation analyses and multiple regressions. Coexisting species responded to year-to-year climatic variability in different ways. Radial growth in junipers and pines responded positively to April and May temperatures, respectively. Summer drought constrained growth in both cases, although its impact was stronger on junipers than on pines. Our findings suggest that junipers respond earlier than pines to spring temperatures due to their prostrate morphology which may enhance a fast warming of their cambial meristems after snowmelt. The higher dependence of J. sabina on summer rainfall as compared with co-occurring pines confirms that drought stress negatively impacts secondary growth in Mediterranean mountains. This sensitivity to water availability may be caused by the juniper shallow root systems, which mainly use superficial soil water. The climatic signal registered in J. sabina allows studying the response of other similar shrubby woody species growing in Mediterranean alpine areas to the ongoing climate warming, which could also reduce water availability.     

The multistemmed structure of Juniperus thurifera: adaptive advantage in a severe environment?

A comparative study of radial growth and biomass between multistemmed trees with variable number of stems and single-stemmed trees was carried out to better understand determinism and organisation of multicaulis structure of a juniper species (Juniperus thurifera L.) growing in high Mediterranean mountains (High Atlas, Morocco). It appears that all the stems of the same tree have similar ages and so simultaneous development. Their mean annual radial increments show significant differences because of probable competition for water and nutrient supply and obvious physical competition for space. The multistemmed trees characterized by low number of stems have the same mean annual radial growth as single-stemmed trees and more generally all multistemmed junipers have a higher biomass. The multicaulis structure of Juniperus thurifera thus could be considered as an adaptation to severe environment, characterized not only by hard topographical, edaphic and climatic conditions, but by strong human pressure too.     

Variation in woody plant mortality and dieback from severe drought among soils, plant groups, and species within a northern Arizona ecotone

Vegetation change from drought-induced mortality can alter ecosystem community structure, biodiversity, and services. Although drought-induced mortality of woody plants has increased globally with recent warming, influences of soil type, tree and shrub groups, and species are poorly understood. Following the severe 2002 drought in northern Arizona, we surveyed woody plant mortality and canopy dieback of live trees and shrubs at the forest–woodland ecotone on soils derived from three soil parent materials (cinder, flow basalt, sedimentary) that differed in texture and rockiness. Our first of three major findings was that soil parent material had little effect on mortality of both trees and shrubs, yet canopy dieback of trees was influenced by parent material; dieback was highest on the cinder for pinyon pine (Pinus edulis) and one-seed juniper (Juniperus monosperma). Ponderosa pine (Pinus ponderosa) dieback was not sensitive to parent material. Second, shrubs had similar mortality, but greater canopy dieback, than trees. Third, pinyon and ponderosa pines had greater mortality than juniper, yet juniper had greater dieback, reflecting different hydraulic characteristics among these tree species. Our results show that impacts of severe drought on woody plants differed among tree species and tree and shrub groups, and such impacts were widespread over different soils in the southwestern U.S. Increasing frequency of severe drought with climate warming will likely cause similar mortality to trees and shrubs over major soil types at the forest–woodland ecotone in this region, but due to greater mortality of other tree species, tree cover will shift from a mixture of species to dominance by junipers and shrubs. Surviving junipers and shrubs will also likely have diminished leaf area due to canopy dieback.     

西藏尼木县古树年龄鉴定及生长历史分析

鉴定老龄树的古树等级并了解其生长历史是国家古树名木普查建档工作的重要内容.本研究利用树木年轮学方法,调查估算了西藏拉萨市尼木县20棵胸径较大的核桃和10棵大果圆柏的树龄,分析其生长历史.结果表明: 20棵核桃树中,一级古树、二级古树及三级古树分别为4、7及9棵;10棵柏树中,有2棵一级古树、4棵二级古树、4棵三级古树.历史上的持续低生长现象在某些时间段内2个树种间表现出明显的同步性,主要集中在20世纪40至80年代,但古树生长在个体间表现出较大的生长差异.古树生长与帕默尔干旱指数(PDSI)表现出显著的正相关关系.研究区古树主要分布于村庄和道路两旁,人类活动频繁.古树的径向生长受到该区域气候调控的同时也受到人类活动的干扰,这导致了古树的径向生长变化在不同树种间表现出空间同步性,而在个体间表现出较大的生长差异.因此,相关部门在制定古树保护措施时应同时考虑气候变化和人类活动对古树生长的影响.     

Climatically induced interannual variability in aboveground production in forest-tundra and northern taiga of central Siberia

To investigate the variability of primary production of boreal forest ecosystems under the current climatic changes, we compared the dynamics of annual increments and productivity of the main components of plant community (trees, shrubs, mosses) at three sites in the north of Siberia (Russia). Annual radial growth of trees and shrubs was mostly defined by summer temperature regime (positive correlation), but climatic response of woody plants was species specific and depends on local conditions. Dynamics of annual increments of mosses were opposite to tree growth. The difference in climatic response of the different vegetation components of the forest ecosystems indicates that these components seem to be adapted to use climatic conditions during the short and severe northern summer, and decreasing in annual production of one component is usually combined with the increase of other component productivity. Average productivity in the northern forest ecosystems varies from 0.05 to 0.14 t ha⁻¹ year⁻¹ for trees, from 0.05 to 0.18 t ha⁻¹ year⁻¹ for shrubs and from 0.54 to 0.66 t ha⁻¹ year⁻¹ for mosses. Higher values of tree productivity combined with lower annual moss productivity were found in sites in northern taiga in comparison with forest-tundra. Different tendencies in the productivity of the dominant species from each vegetation level (trees, shrubs, mosses) were indicated for the last 10 years studied (1990–1999): while productivity of mosses is increasing, productivity of trees is decreasing, but there is no obvious trend in the productivity of shrubs. Our results show that in the long term, the main contribution to changes in annual biomass productivity in forest-tundra and northern taiga ecosystems under the predicted climatic changes will be determined by living ground cover.     

Age structure and climate sensitivity of a high Andean relict forest of Polylepis rodolfo-vasquezii in central Peru

For a better understanding of forest ecology, tree-ring studies can provide information on climate sensitivity, tree growth patterns and population age structure that can inform about stand dynamics such as recruitment of new individuals, and other interspecific interactions related to competition and facilitation. Little is known about the ecology of the recently identified high Andean tree species Polylepis rodolfo-vasquezii. Here, we analyzed the relationship between tree size and age of two P. rodolfo-vasquezii forest stands located in the central Peruvian Andes at 11°S in latitude, and compared their growth patterns and climate sensitivity. We measured the height and diameter of each individual tree and collected tree core samples of living trees and cross sections of dead standing trees to generate two centennial tree-ring chronology at Toldopampa (1825–2015 CE) and at Pomamanta (1824–2014 CE) sites. The dendrochronological dates were evaluated by ¹⁴C analysis using the bomb-pulse methods analyzing a total of 9 calendar years that confirm the annual periodicity of this tree species. At the Toldopampa stand most trees ranged from 70 to 80 years old, with a 190-year old individual, being an older and better preserve forest than Pomamanta, with younger trees, probably because more human disturbances due to closer village proximity. No significant relationships were found between tree age and size in the oldest stand alerting that tree diameter should not be used as a metric for estimating tree ages as a general rule. The distinct growth patterns and the size-age relationship observed at the two forests may reflect distinct histories regarding human activities such as fire and logging. Nevertheless, both the Toldopampa and the Pomamanta tree-ring width chronologies exhibited common growth patterns and shared a similar positive response to temperature of the current growing season. Overall, our study confirmed the annual radial growth periodicity in P. rodofolfo-vasquezii trees using an independent method such as ¹⁴C analyses and a strong climate sensitivity of this tree species. These findings encourage the development of an extensive P. rodolfo-vasquezii tree-ring network for ecological and paleoclimate studies in the tropical Andes in South America.     

Tree height and age-related decline in growth in Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.)

Growth and seasonal water use was measured amongst trees growing in an old growth Scots pine forest in the Scottish Highlands. Three sites which differed in their recent management history and contained old and naturally regenerated young trees growing together were monitored in the field. Our results showed a clear decrease in growth efficiency with age, from values of around 0.25 kg m⁻² leaves year⁻¹ in approximately 25-year-old trees to less than 0.1 kg m⁻² leaves year⁻¹ in trees over 200 years old. When the old trees in one of the field sites were released from competition by thinning, their growth efficiency reverted to that of coexisting young trees, indicating that the decline in growth was reversible. This is consistent with the results of a parallel study showing that cambial age had no effect on the physiology or growth of grafted seedlings originating from the same population studied here (Mencuccini et al. 2005). Our detailed study of tree water use in the field showed an overall decrease in whole-tree hydraulic conductance and stomatal canopy conductance with tree height in the unthinned stands, in agreement with the hydraulic limitation hypothesis. However, the effect of this reduction in hydraulic efficiency on growth was comparatively small, and old trees also showed consistently lower nitrogen concentrations in needles, suggesting that hydraulic and nutritional factors combined to produce the decline in growth efficiency with age observed in the studied populations.     

Evaluating the annual nature of juvenile rings in Bolivian tropical rainforest trees

Knowledge on juvenile tree growth is crucial to understand how trees reach the canopy in tropical forests. However, long-term data on juvenile tree growth are usually unavailable. Annual tree rings provide growth information for the entire life of trees and their analysis has become more popular in tropical forest regions over the past decades. Nonetheless, tree ring studies mainly deal with adult rings as the annual character of juvenile rings has been questioned. We evaluated whether juvenile tree rings can be used for three Bolivian rainforest species. First, we characterized the rings of juvenile and adult trees anatomically. We then evaluated the annual nature of tree rings by a combination of three indirect methods: evaluation of synchronous growth patterns in the tree- ring series, ¹⁴C bomb peak dating and correlations with rainfall. Our results indicate that rings of juvenile and adult trees are defined by similar ring-boundary elements. We built juvenile tree-ring chronologies and verified the ring age of several samples using ¹⁴C bomb peak dating. We found that ring width was correlated with rainfall in all species, but in different ways. In all, the chronology, rainfall correlations and ¹⁴C dating suggest that rings in our study species are formed annually.     

Growth response to climate and drought in <Emphasis Type="Italic">Pinus nigra</Emphasis> Arn. trees of different crown classes

Tree-ring chronologies were examined to investigate the influence of climate on radial growth of Pinus nigra in southeastern Spain. We addressed whether drought differentially affected the ring-widths of dominant and suppressed trees and if our results supported the hypothesis that, in a Mediterranean climate, suppressed conifer trees suffer greater growth reductions than dominant trees. Climate–growth relationships were analyzed using response and correlation functions, whereas the effect of drought on trees growth was approached by superposed epoch analysis in 10 dry years. A cool, wet autumn and spring, and/or mild winter enhanced radial growth. Latewood was the most sensitive ring section in both kinds of trees and it was primarily influenced by current year precipitations. Earlywood was mostly influenced by climatic conditions previous to the growing season. In general, May was the most influential month. Pinus nigra was shown to be very drought sensitive tree in the study area. Tree-rings in suppressed trees showed lower growth reductions caused by drought than those of dominant trees. However, dominant trees recovered normal growth faster. Dominant trees showed a more plastic response, and suppression appeared to reduce the effect of climate on tree radial growth. Some possible causes for these effects are discussed. Our results support the essential role of the balance between light and moisture limitations for plant development during droughts and show that it is not appropriate to generalize about the way in which suppression affects climate-growth relationship in conifers.