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

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

Age-dependent tree-ring growth responses to climate in Qilian juniper (Sabina przewalskii Kom.)

Qilian juniper (Sabina przewalskii kom.) is one of the dominant tree species on Qinghai-Tibetan Plateau and has been used in dendroclimatological studies. Here we designed a test to examine whether or not the climate–growth responses in tree rings of Qilian Juniper vary with the change in tree's age. A total number of 135 increment cores were sampled from Qilian Juniper trees at five sites, in which 112 cores were selected and grouped into five 100-year age classes for analysis of age-dependent climate–growth relationships. Chronology statistics, response functions and ANOVA F-test were used to test the consistency of five age-class mean chronologies (AGCs). The results showed that mean sensitivity (MS) and standard deviation (SD) did not change significantly with age. Response function analysis indicated that (a) climate accounts for a high amount of variance in tree-ring widths; (b) tree-ring growth has significant positive correlation with mean monthly air temperature of previous October and November, and with total monthly precipitation of current January and June, while has significant negative correlation with mean monthly air temperature of current May; and (c) AGC-2, AGC-3 and AGC-4 have stronger response to climate change than AGC-1 and AGC-5. The ANOVA F-test showed that generally there are significant differences between the first age class and other four age classes, but among the four classes in which trees are older than 200 years, the differences are usually insignificant. Overall the long-lived Qilian Juniper is still an ideal tree species for dendroclimatic reconstruction.     

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

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

Does Soil Organic Layer Thickness Affect Climate–Growth Relationships in the Black Spruce Boreal Ecosystem?

The observed long-term decrease in the regional fire activity of Eastern Canada results in excessive accumulation of organic layer on the forest floor of coniferous forests, which may affect climate–growth relationships in canopy trees. To test this hypothesis, we related tree-ring chronologies of black spruce (Picea mariana (Mill.) B.S.P.) to soil organic layer (SOL) depth at the stand scale in the lowland forests of Quebec’s Clay Belt. Late-winter and early-spring temperatures and temperature at the end of the previous year’s growing season were the major monthly level environmental controls of spruce growth. The effect of SOL on climate–growth relationships was moderate and reversed the association between tree growth and summer aridity from a negative to a positive relationship: trees growing on thin organic layers were thus negatively affected by drought, whereas it was the opposite for sites with deep (>20–30 cm) organic layers. This indicates the development of wetter conditions on sites with thicker SOL. Deep SOL were also associated with an increased frequency of negative growth anomalies (pointer years) in tree-ring chronologies. Our results emphasize the presence of nonlinear growth responses to SOL accumulation, suggesting 20–30 cm as a provisional threshold with respect to the effects of SOL on the climate–growth relationship. Given the current climatic conditions characterized by generally low-fire activity and a trend toward accumulation of SOL, the importance of SOL effects in the black spruce ecosystem is expected to increase in the future.     

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.     

Age-specific responses to climate identified in the growth of Quercus alba

The objective of this research was to determine whether the dendroclimatic responses of young Quercus alba (aged 29–126 years) differ from those of old Q. alba (149–312 years). We collected Q. alba increment cores across a range of size classes from Buffalo Mountain Natural Area Preserve, an oak-hickory forest in southcentral Virginia, USA. Tree cores were crossdated and raw ring widths were detrended to remove the influence of increasing circumference with age, microsite, and local stand dynamics. Standardized ring widths were averaged to develop two master chronologies from the 20 oldest and youngest trees. Ring-width indices were correlated with temperature, precipitation, and Palmer Drought Severity Index (PDSI). Annual tree-ring growth in old and young Q. alba was significantly correlated with precipitation from the previous growing season, but was not significantly correlated with temperature. Only the old trees showed a significant correlation between annual ring width and PDSI. These results may indicate that growth in old trees is more sensitive to drought than in young trees. If future climate change includes the predicted increase in mid-growing season droughts, tree-level responses are likely to be age-dependent with older trees experiencing relatively greater reductions in growth.     

Growth responses of an African savanna tree, <Emphasis Type="Italic">Bauhinia thonningii</Emphasis> Schumacher,to defoliation,fire and climate

This study investigated the growth responses to defoliation, fire-exposure and climate factors of a widespread Africana savanna tree, Bauhinia thonningii Schumacher, at a site in central Zambia. Experimental trees (n = 47) were either exposed to fire (n = 12) in the first half of the dry season or protected from fire (n = 35). Some of the fire-protected trees (n = 12) were subjected to artificial defoliation in two consecutive years. Phenological responses (bud break, leaf flush and leaf production) to fire-exposure and defoliation were monitored on permanently marked sample shoots over a 2-year period. Radial tree growth (diameter at 1.3 m above ground) was measured annually over a 7-year period from 1998 to 2005. During the first two years, defoliation and fire-exposure advanced the onset of bud break and leaf flush but fire-exposed trees produced significantly less leaves than did trees protected from fire, probably because scorching caused more severe shoot die-back than is normal. Leaf production was also significantly affected by experimental treatments and their interaction with year. Although treatments had significant short-term effects on radial growth, previous-year growth significantly influenced current-year growth, thereby confirming the existence of autocorrelation in the time-series growth data of B. thonningii. The interaction between previous-year growth and climate factors explained a significant proportion (25–40%, P < 0.001) of the variance in annual tree growth. Variogram models predicted that a 2-year manual defoliation treatment would shorten the longer-term growth cycle while continuous fire-exposure extended the cycle by one year. The results are useful for the management of savanna trees.     

On the detection of dynamic responses in a drought-perturbed tropical rainforest in Borneo

The dynamics of aseasonal lowland dipterocarp forest in Borneo is influenced by perturbation from droughts. These events might be increasing in frequency and intensity in the future. This paper describes drought-affected dynamics between 1986 and 2001 in Sabah, Malaysia, and considers how it is possible, reliably and accurately, to measure both coarse- and fine-scale responses of the forest. Some fundamental concerns about methodology and data analysis emerge. In two plots forming 8 ha, mortality, recruitment, and stem growth rates of trees ≥10 cm gbh (girth at breast height) were measured in a ‘pre-drought’ period (1986–1996), and in a period (1996–2001) including the 1997–1998 ENSO-drought. For 2.56 ha of subplots, mortality and growth rates of small trees (10–<50 cm gbh) were found also for two sub-periods (1996–1999, 1999–2001). A total of c. 19 K trees were recorded. Mortality rate increased by 25% while both recruitment and relative growth rates increased by 12% for all trees at the coarse scale. For small trees, at the fine scale, mortality increased by 6% and 9% from pre-drought to drought and on to ‘post-drought’ sub-periods. Relative growth rates correspondingly decreased by 38% and increased by 98%. Tree size and topography interacted in a complex manner with between-plot differences. The forest appears to have been sustained by off-setting elevated tree mortality by highly resilient stem growth. This last is seen as the key integrating tree variable which links the external driver (drought causing water stress) and population dynamics recorded as mortality and recruitment. Suitably sound measurements of stem girth, leading to valid growth rates, are needed to understand and model tree dynamic responses to perturbations. The proportion of sound data, however, is in part determined by the drought itself. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Age-related drought sensitivity of Atlas cedar (Cedrus atlantica) in the Moroccan Middle Atlas forests

Age-related tree responses to climate change are still poorly understood at the individual tree level. In this paper, we seek to disentangle the relative contribution of tree age to growth decline and growth–climate relationships in Atlas cedar (Cedrus atlantica Manetti) trees at the Middle Atlas Mountains, northern Morocco. Dendrochronological methods were applied to quantify growth–climate relationships using tree-ring width indices (TRWi) calculated for cedars of two contrasting age groups (old trees, age ≥150 years; young trees, age <150 years). TRWi–climate relationships were assessed at the site and tree levels by using response functions and linear mixed-effects models, respectively. Growth of the studied Atlas cedars was negatively affected by recurrent droughts and by the steep temperature rise since the 1970s. Response functions and mixed-effects models indicated that the decline in tree growth was mainly explained by diminishing precipitation. The negative association between cedar growth and temperature was stronger in old than in young trees. Vulnerability to temperature-induced drought stress in old cedar trees may lead to an impending growth decline. We argue that the age dependence of growth sensitivity to drought must be quantified and considered at the individual tree level when predicting the future dynamics and persistence of cedar forests in the Moroccan Middle Atlas.     

川西亚高山不同年龄紫果云杉径向生长对气候因子的响应

运用树木年轮气候学的基本方法,建立王朗自然保护区紫果云杉在集中分布上限区域的年轮宽度年表,选取差值年表分析不同年龄云杉的径向生长同逐月气候因子的相关及响应关系,结果显示:幼龄组云杉年表的敏感度高于中龄组和老龄组云杉,幼龄组云杉对生长季前及生长季的气温状况显著正相关;中龄组云杉年表仅与当年4月份和7月份的月平均最低气温显著正相关;老龄组云杉的年轮宽度指数同上年生长季(上年8月份)的月平均气温和月平均最低温显著负相关,上年生长季高温的"滞后效应"在老龄组云杉体现的更为突出;幼龄组与中龄组云杉对当年6月份降水持续增加显示出明显的负相关关系,上年12月份的降水会对幼龄组和老龄组云杉径向生长不利。研究表明幼龄组云杉包含的气候信息要优于中龄组和老龄组云杉,在该区域进行相关研究时应根据研究需要选取不同年龄跨度的云杉年表。     

Climate sensitivity of radial growth in European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) at different aspects in southwestern Germany

The climate sensitivity of radial growth in European beech (Fagus sylvatica L.) was analyzed within a narrow valley in the Swabian Alb (southwestern Germany). We collected stem disks from three aspects (NE, NW and SW) of trees belonging to different social classes. Common climatic factors limiting growth across the valley were identified using a principal component analysis (PCA). Further, we performed hierarchical cluster analysis (HCA), redundancy analysis (RDA) and bootstrapped correlation analysis to reveal differences in chronologies and climate-growth relationships between aspect and social class. Climatic variables considered in our analyses were monthly and seasonal data on temperature and precipitation, as well as a self-calibrating Palmer drought severity index (sc-PDSI). We identified drought in the period June–August as the most prominent factor limiting growth across the valley. Dominant and co-dominant trees at the NW and SW aspects were found to be particularly drought sensitive, whereas intermediate trees were less susceptible to drought. Underlying causes of established climate–growth relationships are discussed in the context of drought susceptibility, tree-size modulation and tree physiological processes.     

Interannual growth response of Norway spruce to climate along an altitudinal gradient in the Tatra Mountains, Poland

Ring-width chronologies of Picea abies (L.) Karst. from ten sites in the Tatra Mountains, Poland, were developed to explore growth/climate responses in stands along an altitudinal gradient ranging from 839 to 1468 m a.s.l. There were positive relationships between current-year radial growth and mean monthly temperatures in March, April, June and July, but with increasing elevation, the strength of this correlation declined for March–April and increased for June–July temperatures. The mean monthly temperature in October of the previous year positively influenced radial growth of trees at all sites. Lower mean temperatures in January negatively affected growth of trees at the high-elevation sites. Trees at the low-elevation sites responded positively to a warm early spring, whereas trees at the high-elevation sites showed positive growth responses to higher summer temperatures. Growth of trees at the high-elevation sites was better explained by the temperature regime than was growth at the lower-elevation sites. Therefore, it is likely that these sites may be particularly sensitive to potential effects of temperature change.     

Assessing the climatic sensitivity of Douglas-fir at its northern range margins in British Columbia, Canada

Northern hemisphere tree species growing at their northern range margins may be particularly responsive to climate change and could provide important information regarding future broad-scale responses. We analyzed and compared tree-ring data between five Douglas-fir [Pseudotsuga menziesii var. glauca (Mirb.) Franco] populations growing at the species’ northernmost distribution in British Columbia, Canada, and five populations located 150 km to the southeast. We quantified climate–growth relationships using uni- and multivariate techniques at different temporal scales. Our data suggest that (1) even at its northernmost distributions, precipitation limits long-term mature Douglas-fir radial growth more than temperatures, (2) northernmost Douglas-fir populations are distinct from populations located further within the species’ range in terms of certain key short-term growth responses, and (3) northernmost Douglas-fir growth sensitivities to climate may be increasing over time. In the future, mature Douglas-fir productivity in the northern portion of its range may be primarily limited by precipitation, and responses may be strongest at the species’ range margins.     

Vegetation control effects on untreated wood,crude cellulose and holocellulose δ<Superscript>13</Superscript>C of early and latewood in 3- to 5-year-old rings of Douglas-fir

The stable carbon (C) composition of tree rings expressed as δ¹³C, is a measure of intrinsic water-use efficiency and can indicate the occurrence of past water shortages for tree growth. We examined δ¹³C in 3- to 5-year-old rings of Douglas-fir (Pseudotsuga menziesii (Mirb) Franco) trees to elucidate if decreased water supply or uptake was a critical factor in the observed growth reduction of trees competing with understory herb and shrub vegetation compared to those growing without competition. We hypothesized that there would be no differences in δ¹³C of earlywood in trees growing in plots with competing vegetation and those in plots receiving complete vegetation control during 5 years because earlywood formed early in the growing season when soil water was ample. We also hypothesized that δ¹³C in latewood which was formed during the later half of the growing season when precipitation was low, would be greater (less negative) in trees in plots without vegetation control. We then separated early and latewood from rings for three consecutive years and analyzed their δ¹³C composition. No significant differences in earlywood δ¹³C in years 3–5 were observed for trees in the two vegetation control treatments. δ¹³C of untreated latewood separated from wood cores was greater in 4- and 5-year-old rings of trees growing with competing vegetation compared to trees growing without vegetation competition (i.e., −25.5 vs. −26.3‰ for year 4, and −26.1 vs. −26.8‰ for year 5). Results suggest that water shortages occurred in Douglas-fir trees on this coastal Washington site in the latewood-forming portion of the growing season of years 4 and 5 in the no-vegetation control treatment. We also compared δ¹³C from untreated wood, crude cellulose extracted with the Diglyme–HCl method, and holocellulose extracted with toluene–ethanol to see if the extraction method would increase the sensitivity of the analysis. δ¹³C values from the two extraction methods were highly correlated with those from untreated samples (r ² = 0.97, 0.98, respectively). Therefore, using untreated wood would be as effective as using crude cellulose or holocellulose to investigate δ¹³C patterns in young Douglas-fir.     

秦岭不同龄组巴山冷杉径向生长对气候因子的响应差异

秦岭地区树轮气候学研究已经引起众多学者的重视,但年龄因素对调节树木径向生长-气候响应关系的研究尚未在当地得到广泛关注。运用树轮气候学方法,建立秦岭牛背梁国家级自然保护区低、中、高3个龄组巴山冷杉(Abies fargesii)的树轮宽度差值年表,探究不同龄组巴山冷杉径向生长与覆盖研究区的0.5°×0.5°分辨率格点气候要素之间的响应关系,以期解译年龄差异对巴山冷杉树木径向生长-气候响应特征的潜在影响。结果表明：不同龄组巴山冷杉差值年表统计特征值存在明显差异,平均敏感度和样本总体代表性随年龄增大而递减,但标准差、样本间平均相关系数和第一特征根变异解释量均以中龄组最低,信噪比却以中龄组最高;春季(3-5月)降水的增加以及秋季(9-11月)气温的升高对低龄组巴山冷杉径向生长的促进作用最强,对中龄组树木生长的促进作用明显减弱,而对高龄组树木生长几乎没有影响;气温对巴山冷杉径向生长的促进作用随树龄增高而增强,差值年表与当年2月、8月、上年5月平均温、平均最高温的相关系数均呈现随龄级增大而逐渐递增的趋势。年龄因素对牛背梁国家级自然保护区巴山冷杉径向生长-气候响应关系存在明确影响,随着年龄的增加,巴山冷杉径向生长受降水的制约减弱,受气温的限制增强。因此,在进行树木年轮学研究以及制定森林经营管理策略时应该考虑不同树龄巴山冷杉对气候的响应差异。     

Vertical distribution of epiphytic bryophytes and lichens emphasizes the importance of old beeches in conservation

During storms in 2005, a number of beech trees fell over at Biskopstorp, SW Sweden, offering the opportunity to study epiphytes along entire stems. In total 16 beech trees in four beech stands representing three different age classes were included. For each tree, 2 m segments from the base to the top were surveyed. In total 115 species were found (76 lichens, 39 bryophytes), of which 30 were considered to be of conservation concern (22 lichens, 8 bryophytes). For lichens significantly more species were recorded above 2 m in height, whereas more bryophytes were recorded below 2 m in height. Certain red-listed lichens were recorded only above 2 m in height on old trees. In a second data set from the same area 140 age-determined beech trees were surveyed for species of conservation concern at the heights 0–2 and 2–5 m, respectively. These species were found almost exclusively on old beech trees, and presence at 2–5 m was recorded, with one exception, only on those trees which also had species of conservation concern at 0–2 m. Records of these species correlated significantly to microhabitat variables, i.e. the presence of rough bark and moss cover higher up the stems on the old trees. This study indicates that surveying only the base in really old beech forests can underestimate both the number of species of conservation concern and their population sizes. However, surveys restricted to the base in rather even-aged beech stands catch a large proportion of the trees with species of conservation concern.     

Buffered climate change effects in a Mediterranean pine species: range limit implications from a tree-ring study

Within-range effects of climatic change on tree growth at the sub-regional scale remain poorly understood. The aim of this research was to use climate and radial-growth data to explain how long-term climatic trends affect tree growth patterns along the southern limit of the range of Pinus nigra ssp. salzmannii (Eastern Baetic Range, southern Spain). We used regional temperature and precipitation data and measured sub-regional radial growth variation in P. nigra forests over the past two centuries. A dynamic factor analysis was applied to test the hypothesis that trees subjected to different climates have experienced contrasting long-term growth variability. We defined four representative stand types based on average temperature and precipitation to evaluate climate–growth relationships using linear mixed-effect models and multi-model selection criteria. All four stand types experienced warming and declining precipitation throughout the twentieth century. From the onset of the twentieth century, synchronised basal-area increment decline was accounted for by dynamic factor analysis and was related to drought by climate–growth models; declining basal-area increment trends proved stronger at lower elevations, whereas temperature was positively related to growth in areas with high rainfall inputs. Given the contrasting sub-regional tree-growth responses to climate change, the role of drought becomes even more complex in shaping communities and affecting selection pressure in the Mediterranean mountain forests. Potential vegetation shifts will likely occur over the dry edge of species distributions, with major impacts on ecosystem structure and function.     

Long-Term Growth and Succession in Restored and Natural Mangrove Forests in Southwestern Florida

We compared colonization, growth and succession from 1989 to 2000 in a restored mangrove site and in gap and closed canopy sites in a natural mangrove forest. The restored site was created in 1982 and planted with Rhizophora mangle (≈2 m⁻²) propagules. By 1989, Laguncularia racemosa, with densities up to 12.9 tree m⁻², was a dominant in all plots, although densities were greater at edge plots relative to inner plots, and near open water (west plots) relative to further inland (east plots), and in tall mangrove plots relative to scrub plots. Rhizophora mangle (1989 tree densities about 2 m⁻²) was a codominant in inner and scrub plots, while Avicennia germinans had the lowest densities (<1 tree m⁻²) in all plots. From 1989 to 2000 L. racemosa experienced reduced recruitment and apparent density-dependent mortality of canopy individuals in plots with high initial densities. Scrub plots experienced high rates of colonization by R. mangle and L. racemosa, rapid growth in height of all species (1989–1996), followed by a dieoff of L. racemosa in later years (1997–2000) as the canopy came to resemble that of tall mangrove plots. Colonization and growth rates were lower in gap and closed canopy regions of the natural forest relative to rates in the restored site. After 11 years, densities of L. racemosa were 10–20× lower and R. mangle slightly less in the gap relative to densities in tall mangrove plots in the restored site at the same age. Although the restored stand had converged with the natural forest by 2000 in terms of some factors such as species richness, vegetation cover, litterfall, and light penetration, trees were still much smaller and stem densities much higher. Full development of mature structure and ecological function will likely require decades more development.     

Rainforest fragmentation and the demography of the economically important palm <Emphasis Type="Italic">Oenocarpus bacaba</Emphasis> in central Amazonia

We summarize a long-term study of the effects of edge creation on establishment of the economically important arboreal palm Oenocarpus bacaba in an experimentally fragmented landscape in central Amazonia. Recruitment and mortality of large individuals (≥10 cm diameter-at-breast-height) were recorded within 21 1-ha plots in fragmented and intact forests for periods of up to 22 years. In addition, 12 small (0.7 × 14 m) sub-plots within each 1-ha plot were used to enumerate the abundance of seedlings and saplings (5–400 cm tall). On average, the recruitment of large trees was over two times faster near forest edges, leading to a sharp (90%) increase in the mean population density of large individuals near forest edges, whereas the density of larger trees remained constant in the forest interior. Overall seedling and sapling density was significantly lower in edge than interior plots, but edge plots had a much higher proportion of larger (>100 cm tall) saplings. Our findings demonstrate that forest edges can have complex effects on tree demography and that one must consider all tree life stages in order to effectively assess their effects on plant populations.     

Climatic Stress during Stand Development Alters the Sign and Magnitude of Age-Related Growth Responses in a Subtropical Mountain Pine

The modification of typical age-related growth by environmental changes is poorly understood, In part because there is a lack of consensus at individual tree level regarding age-dependent growth responses to climate warming as stands develop. To increase our current understanding about how multiple drivers of environmental change can modify growth responses as trees age we used tree ring data of a mountain subtropical pine species along an altitudinal gradient covering more than 2,200 m of altitude. We applied mixed-linear models to determine how absolute and relative age-dependent growth varies depending on stand development; and to quantify the relative importance of tree age and climate on individual tree growth responses. Tree age was the most important factor for tree growth in models parameterised using data from all forest developmental stages. Contrastingly, the relationship found between tree age and growth became non-significant in models parameterised using data corresponding to mature stages. These results suggest that although absolute tree growth can continuously increase along tree size when trees reach maturity age had no effect on growth. Tree growth was strongly reduced under increased annual temperature, leading to more constant age-related growth responses. Furthermore, young trees were the most sensitive to reductions in relative growth rates, but absolute growth was strongly reduced under increased temperature in old trees. Our results help to reconcile previous contrasting findings of age-related growth responses at the individual tree level, suggesting that the sign and magnitude of age-related growth responses vary with stand development. The different responses found to climate for absolute and relative growth rates suggest that young trees are particularly vulnerable under warming climate, but reduced absolute growth in old trees could alter the species’ potential as a carbon sink in the future.