Growth Rings,Phenology, Hurricane Disturbance and Climate in Cyrilla racemiflora L., a Rain Forest Tree of the Luquillo Mountains,Puerto Rico1

The growth phenology of Cyrilla racemiflora L., the dominant tree species of the montane rain forest, (subtropical lower montane rain forest, sensu Holdridge) of the Luquillo Mountains of Puerto Rico was studied intensively during 1989, and then semiannually through mid-1993 to determine the periodicity of changes in xylem structure. Four trees at 770 m were monitored for flowering, branch elongation, leaf litterfall, and xylem cell growth and differentiation in the lower stem, and these events were related to local seasonal patterns of rainfall and temperature. Hurricane Hugo defoliated study trees in September, 1989. Bud-break and branch elongation in March, 1989 were followed by earlywood xylem cell production in the lower stem in April and the onset of flowering in May. Leaf litterfall was greatest between April and June, coinciding with peak branch growth and new leaf formation. Latewood xylem was produced in December. The general phenological pattern was synchronized between trees and over study years. Vessel diameter and density were monitored along with thickness of earlywood and latewood and the former converted to vessel lumen area, a measure of xylem conductance capacity. Annual growth rings were formed with periods of earlywood and latewood production coinciding with traditional summer (rainy) and winter (dry) seasons, respectively, in the Luquillo Mountains. Hurricane defoliation was followed by heavy flowering in 1990, a year of reduced branch elongation and annual xylem ring width, and was associated with maximum vessel lumen area, as was flowering in 1989, prior to the hurricane. Hurricane Hugo provided a perturbation that, through its elicited stress response, allowed for the demonstration of the interplay between flowering, branching, structural growth of xylem, and xylem function.     

Relating tree growth to rainfall in Bolivian rain forests: a test for six species using tree ring analysis

Many tropical regions show one distinct dry season. Often, this seasonality induces cambial dormancy of trees, particularly if these belong to deciduous species. This will often lead to the formation of annual rings. The aim of this study was to determine whether tree species in the Bolivian Amazon region form annual rings and to study the influence of the total amount and seasonal distribution of rainfall on diameter growth. Ring widths were measured on stem discs of a total of 154 trees belonging to six rain forest species. By correlating ring width and monthly rainfall data we proved the annual character of the tree rings for four of our study species. For two other species the annual character was proved by counting rings on trees of known age and by radiocarbon dating. The results of the climate–growth analysis show a positive relationship between tree growth and rainfall in certain periods of the year, indicating that rainfall plays a major role in tree growth. Three species showed a strong relationship with rainfall at the beginning of the rainy season, while one species is most sensitive to the rainfall at the end of the previous growing season. These results clearly demonstrate that tree ring analysis can be successfully applied in the tropics and that it is a promising method for various research disciplines.     

不同径级油松径向生长对气候的响应

建立了黑里河自然保护区油松年轮宽度年表,通过不同径级油松径向生长对逐月气候因子的响应关系,研究了干旱对不同径级油松径向生长的影响。结果表明:两个径级油松的年轮宽度指数达到极显著相关(R=0.943,P<0.01),其中小径级(平均胸径20 cm)油松年表的平均敏感度显著高于大径级(平均胸径43 cm)油松年表(P<0.01)。不同径级油松均与上年9月、当年2月及当年5—6月的降水显著正相关(P<0.05),与当年6月的平均温度显著负相关(P<0.05),此外,小径级油松还与当年7月的降水显著正相关(P<0.05);降水是影响油松生长的主要气候因子。不同径级油松的径向生长量在干旱年份均显著降低(P<0.01)且小径级油松的生长降低量显著高于大径级油松(P<0.01);不同径级油松生长量在干旱发生后1年左右的时间内均恢复正常且小径级油松恢复速度更快。     

Variation in stem radial growth of the Australian conifer, Callitris columellaris, across the world’s driest and least fertile vegetated continent

Climate change could alter the biogeography of many tree species. However, there have been few studies of tree growth across climatic gradients at a continental scale. Callitris columellaris is a widespread conifer that spans many climates and landscape positions across Australia. Our aim was to determine how stem radial growth of C. columellaris varies with tree size and with the biogeographic factors of rainfall, temperature, soil fertility and inter-tree competition. We sampled cores from trees at 85 sites in biomes ranging from tropical savanna to arid desert and temperate forest, and measured widths of the 100 outermost growth rings. We analysed ring width in relation to changes in tree age and diameter, and also evaluated the influence of the biogeographic factors on the width of the ten most recently formed rings. The average width of outermost rings varied only slightly with stem diameter, because the decrease in ring width with age and diameter within trees is offset by an increase with diameter among trees. Our analyses thus explain the weak, inconsistent relationships often observed between stem diameter and growth rate amongst trees. The most important biogeographic factors were the climatic ones: across Australia, ring width increased with both mean annual rainfall and mean annual temperature. These relationships were largely driven by continental scale differences between the tropical and the southern (arid plus temperate) sites, while relationships within climate zones were comparatively weak. Ring width decreased with intense inter-tree competition but showed little correlation with available soil nitrogen or phosphorus.     

南亚热带气候下三种树木径向生长季节节律研究

用带状测树器对南亚热带气候条件下的落叶树柚木（Ｔｅｃｔｏｎａ ｇｒａｎｄｉｓ Ｌ．ｆ．）、喜树（Ｃａｍｐｔｏｔｈｅｃａ ａｃｕｍｉｎａｔａ Ｄｅｃｎｅ）和常绿树火烧花（Ｍａｙｏｄｅｎｄｒｏｎ ｉｇｅｕｍ Ｋｕｒｚ）１６年生（测定开始时年龄）的立木的径向生长进行了６ａ的测定。研究结果表明,所测立木的年均胸围生长量在１．５～２．５ｃｍ（胸径生长量为０．４８～０．８０ｃｍ）之间,年间有一定的差异。年生长     

民勤绿洲荒漠过渡带梭梭树干液流的时滞特征

应用Granier热扩散探针测定民勤绿洲荒漠过渡带梭梭人工林的树干液流,将液流与饱和水汽压差(VPD)和光合有效辐射(PAR)数据分别进行逐行错位分析,探讨树干液流与环境因子之间的时滞效应.结果表明: 梭梭日液流速率呈现明显的季节变化,6月平均液流速率最大,8月平均液流速率最小.生长季(5-9月)梭梭树干液流与光合有效辐射、饱和水汽压差存在明显的时滞,树干液流滞后于PAR 80 min,提前于VPD 114 min,且不同月份的时滞长短存在差异.尽管梭梭树干液流在日尺度上更加依赖于PAR的变化,但在白天,树干液流与饱和水汽压差间有更紧密的关系.梭梭生长季树干液流与VPD或PAR之间的时滞与树形因子(株高、地径、50 cm处直径、枝下高、冠幅)及夜间液流量的相关性均不显著.     

华北落叶松夜间树干液流特征及生长季补水格局

在宁夏六盘山北侧半干旱区的叠叠沟小流域,采用热扩散探针法在2011年生长季监测了华北落叶松(Larix principisrupprechtii)人工林的树干液流速率,分析了夜间树干液流和补水量的变化特征及与气象、土壤水分等环境因子的关系.结果表明:树干液流速率日变化表现为典型的单峰宽峰曲线,且整个生长季均存在微弱的夜间液流,一般表现为逐渐减小,特别是在晴天,且晴天的变幅显著大于雨天.除生长季中期雨天夜间液流平均速率显著高于晴天,生长季初期及末期雨天与晴天的差异并不显著.生长季内,夜间树干补水总量为11.03 mm,占总蒸腾量的7.22％;5月份的树干补水量最大(4.19mm),其他月份的树干补水量明显减小,在0.9-1.7mm的范围波动.但不同月份间的补水贡献率存在明显差异,表现为生长季末期(9、10月)＞初期(5月)＞中期(6-8月).相关分析表明,日补水量与各气象因子关系不大,仅与降水量显著正相关(P＜0.05),与土壤含水率、日间蒸腾量、日蒸腾总量极显著正相关(P＜0.01).夜间补水的月蒸腾贡献率与月均土壤含水率、月均气温、月均日间蒸腾量、月总蒸腾量等显著相关(P＜0.05);而夜间补水的日蒸腾贡献率与日最高气温、日均气温、日间蒸腾量、日均饱和水汽压差、日总蒸腾量、日均太阳辐射强度、日最低气温、日均空气相对湿度、日降水量、土壤含水率等极显著相关(P＜0.01),经逐步回归分析建立了日补水量蒸腾贡献率与环境因子的多元线性模型.     

The effects of repeated cutting on coppice response of <Emphasis Type="Italic">Terminalia sericea</Emphasis>

Key message

Although exposure to multiple cutting cycles reduces stored reserves, shoot diameter and shoot length, it, however, increases shoot production per resprouting stump and foliar nitrogen.

Abstract

In disturbance-prone environments with fluctuating seasonal rainfall such as savannas, the repeated cutting of the same trees eventually results in a possible decline in tree abundance. The effects of subjecting Terminalia sericea trees to one, two and multiple (eight) cutting events over a period of 2 years on coppice response were investigated in a savanna woodland in South Africa. Resprout shoot diameter, shoot length and the total cumulative diameter were lower in trees exposed to a high number of cutting events compared to trees exposed to one cutting event. Increasing the number of cutting events significantly reduced stem total non-structural carbohydrate levels in trees indicating a depletion of stored reserves. Foliar carbon content remained largely the same, while leaf nitrogen and phosphorus concentrations significantly increased with each cutting event, relative to uncut trees. Results indicate that trees cut once had not replenished depleted carbohydrate reserves even after a period of 18 months during which no cutting took place. Sustainable utilization of this tree species must allow for at least more than 18 months of undisturbed growth between harvests to allow for the replenishing of reserves.

     

Radial growth response of trees to seasonal soil humidity in a subtropical forest

Tree growth is the most important factor in determining the carbon sequestration processes of forest ecosystems. However, the growth phenology (seasonal growth pattern) and responses of tree growth to climatic variables vary considerably among different species, especially between deciduous and evergreen species. Thus, it is crucial to explore the seasonal growth patterns of different tree species in relation to climate to better understand the responses of tree physiology to climate changes, especially in mixed-species forest stands. In this study, we monitored the daily basal area increments of 220 individuals belonging to 15 common broadleaved tree species, nine deciduous and six evergreen species, in mixed-species experimental stands in subtropical China and analysed the relationships between radial stem growth and seasonal climate at a high-temporal resolution. We fitted daily increments of stem diameters with four frequently used nonlinear models and chose the best model for each species. The results showed that the evergreen trees grew faster than the deciduous trees, both annually and within the growing season. The tested nonlinear models (Korf, Weibull, logistic and Gompertz) produced good fits for the growth patterns of all species. Overall, the evergreen species began stem growth earlier and finished later during the growing season than that of the deciduous species. Within the growing season, the radial growth of trees in mixed stands containing both types of species was strongly positively correlated with humidity. In spring, increases in both temperature and moisture increased the daily relative basal area increment of all species. Maximum growth rates occurred when the soil water content reached its highest level and gradually decreased when the soil water content decreased. In summer, high temperatures combined with low amounts of precipitation led to heat-induced summer drought, to which the evergreen trees appeared to be more tolerant than the deciduous trees, which was reflected in the reduced stem growth of the latter. These results indicate the different climate-dependent seasonal growth strategies of evergreen and deciduous trees related to the trade-off described by the leaf economics spectrum, i.e., short-lived leaves with higher assimilation rates in deciduous and longer-lived leaves with a greater drought tolerance in evergreen species.     

Climate controls act at different scales on the seasonal pattern of Quercus ilex L. stem radial increments in NE Spain

Long-term data on radial increment dynamics in Mediterranean species may identify which climatic variables are the main constraints for radial growth and at which temporal scales they act. To this end, we examined stem radial fluctuations in Quercus ilex L., the dominant evergreen oak species in the Western Mediterranean Basin, over a period of 11 years (1994–2004) at a coastal site in north-eastern Spain. We used manual band dendrometers to record girth changes in trees on north- and south-facing slopes. Annual increments measured by dendrometers showed good agreement with annual tree-ring width. North-facing trees showed a lower long-term cumulative radial increment than south-facing trees. The seasonal radial increment pattern of Q. ilex was bimodal, being characterized by a greater increase in May and a lesser, more variable increase peak in September. Both phases corresponded to warm and moist climatic conditions, whereas radial increase of stems stopped in winter and occasionally in summer. Considering the whole year, mean maximum air temperature was the main factor positively affecting radial increment of Q. ilex from short- (5 days) to- long (30 days) time scales, whereas the accumulated precipitation exerted a similar effect at longer (30 days) scales, but only on south-facing trees. In summer, all trees were positively correlated with precipitation at long-time scales (30 days); however, only stem increment of south-facing trees showed a significant relation to the temperature at short-time scales (10 days). We confirmed the dominant role of temperature as the major constraint on radial increment at short time scales, despite most previous studies were mostly biased towards precipitation effects at monthly scales.     

Growth allocation between height and stem diameter in nonsuppressed reproducing Abies mariesii trees

To understand the growth patterns with respect to competition and leaf‐mass increase in reproducing trees, growth allocation between height and stem diameter was examined for nonsuppressed reproducing Abies mariesii trees in a subalpine forest in northern Honshu, Japan. The growth allocation was analyzed by dividing the relative growth rate of the stem volume into the relative contributions of height and stem‐diameter growth. During a 9‐yr period, height growth and seed‐cone production showed obvious annual variation, while stem‐diameter growth recorded moderate variation. For two of three years of seed‐cone production during the 9‐yr period, trees with larger seed‐cone production were associated with less height growth in the following year of seed‐cone production; however, there was no trend of height growth in the year of seed‐cone production. In the following year of mast seeding, trees with larger stem‐volume growth were associated with less height growth. This trend was also shown for the relationship between the cumulative stem‐volume growth during the 9‐yr period and growth allocation to height, suggesting that trees with a larger biomass increase depress the allocation of photosynthate to competition with a large expenditure for reproduction. In contrast to this, trees with a smaller biomass increase might allocate photosynthate to competition with surrounding trees. The results of this study suggest that an increase in reproductive organs during life history and annual variation in reproduction are closely associated with the growth patterns of the stem in A. mariesii trees.     

Climatic impact on tree-ring widths in Abies borisii-regis forests from South-East Albania

The long-term radial growth responses to drought and climatic variability of less-studied species such as Abies borisii-regis (Mattf.) remain poorly understood.We tested the hypothesis that severe short-term drought conditions during summer months will impact the radial growth of A. borisii-regis (Mattf.) trees and such impact will have a more pronounced effect on latewood (LW) than earlywood (EW) width.Correlation analysis was employed to investigate the impact of climatic drivers (temperature, precipitation) and drought, using the Standardized Precipitation Evapotranspiration Index (SPEI) calculated at cumulative time scales (1–12 months), on EW, LW and tree-ring width (TRW) in A. borisii-regis (Mattf.) trees from South-Eastern Albania.We found that EW width was positively correlated with precipitation in July and previous September, while the LW width and TRW was enhanced by the current June–July precipitation. Previous autumn and current summer high temperatures constrained the radial growth in A. borisii-regis (Mattf.) trees, particularly the LW and TRW. All the tree-ring widths components showed the highest significant response to drought at short cumulative time scales (<4 months) mainly during July, August and September. The highest impact of drought was observed for the LW width.Under a future reduction of summer precipitation and temperature increase, the A. borisii-regis (Mattf.) may show a decrease in EW formation, causing a decline of radial growth, leading to a reduction in hydraulic conductivity and carbon uptake in these forests.     

Study on Dynamics of Calorific Value,Biomass and Energy in Calamagrostis epigejos Population in Songnen Grassland (in English)

There were two peaks of seasonal changes of the calorific value in shoot and leaves of Calamagrostis epigejos in middle June and in the early August respectively. The calorific value in stem presented a single peak curve which appeared in the early August. The calorific values in inflorescence and dead standing showed a fluctuation and the peak value of inflorescence was in the early September and that of dead standing was in middle June. The seasonal changes of energy standing crop on the above-ground part synchronized with that of the biomass, which presented a single peak curve. The energy allocated to each organ in different seasons was in the order as leaves >stem >inflorescence in middle June, leaves >stem >dead standing >inflorescence in early July, leaves >stem >inflorescence >dead standing in middle July, and leaves >stem >dead standing >inflorescence from August to September. The vertical allocation of energy in the parts of above-ground was that the energy value gradually increased from the surface to the 20 cm high level and the maximum value at the 10-20 cm high level which made up 26.91% of energy on the above-ground partion, and then it was decreased. In the under-ground portion, the energy value progressively decreased with depth and the maximum value was at 0-10 cm depth layer which made up 69.01% of energy of the under-ground portion.     

Demographic and growth patterns of Pentaclethra macroloba (Willd.) Kuntze,a hyperdominant tree in the Amazon River estuary

Little is known about the life history and environmental factors that regulate the growth rate of hyperdominant trees in flooded Amazonian forests. Pentaclethra macroloba is a hyperdominant tree, and it is widely explored in the Amazon, because its seed oil is a powerful herbal medicine. We evaluated the demographic structure and growth patterns of P. macroloba and tested the effect of the Amazon River flood pulse on its growth. We modeled the growth and determined the age of P. macroloba by analyzing the growth rings of 30 monitored trees in relation to hydroclimatic variables. We also inventoried 240 juvenile and 2072 adult trees arranged in a clustered pattern. The diametric distribution pattern of the juvenile and adult trees was exponential and log-normal, respectively. The trees were found to be up to 102 years old, and 47% of them grew freely toward the canopy. Peak growth in height and diameter occurred at 24 (61.7 cm year^-1) and 46 (9.38 mm year^-1) years, respectively. Pentaclethra macroloba showed cambial dormancy during the seasonal peak of rainfall (R² = 0.41; t = −2.62; p < 0.01) and flooding of the Amazon River (R² = 0.47; t = −3.01; p < 0.01). Increases in rainfall and flood level of the river in the rainy season control the growth rate of P. macroloba, making it a seasonal process. The demographic and growth patterns of P. macroloba respond to the environmental heterogeneity of the estuarine floodplain forest and also reflect its life history over time.     

松嫩草原拂子茅种群热值、生物量和能量动态的研究

拂子茅（Calamagrostis epigejos)全株的叶的热值和季节变化规律相似,呈双峰曲线,2个峰值分别出现在6月中旬和8月初,茎热值的变化呈单峰曲线,峰值出现在8月初,穗笔立枯体的热值变化呈波动型,穗的最大值出现在9月初,立枯估出现在6月中旬,地上部能量现状量的季节变化与生物量变化同步,呈单峰曲线,不同季节能量在各器官中分配比率为：6月中旬为叶＞茎＞穗,7月初为叶＞茎＞立枯体＞穗,7月中旬为叶＞茎＞穗＞立枯体,8月至9月为叶＞茎＞立枯体＞穗,能量的垂直分配,地上部为从地表至20cm高度逐渐增加,最大值在１０－２０cm层占地上部总能量的２６．９１％,然后逐渐下降,地下部能量垂直分布规律是随着深度增加而逐渐减少,能量集中分布在地下0－10cm层在占地下部总能量69．01％。     

Climate variability,tree increment patterns and ENSO-related carbon sequestration reduction of the tropical dry forest species <Emphasis Type="Italic">Loxopterygium huasango</Emphasis> of Southern Ecuador

Key message

Striking hydro-climatic differences of 2 years (wet; dry) dramatically control the increment pattern of L. huasango in varying extent, even causing a “growth collapse” during the La Niña drought 2010/2011.

Abstract

We present the first multi-year long time series of local climate data in the seasonally dry tropical forest in Southern Ecuador and related growth dynamics of Loxopterygium huasango, a deciduous tree species. Local climate was investigated by installing an automatically weather station in 2007 and the daily tree growth variability was measured with high-resolution point dendrometers. The climatic impact on growth behaviour was evaluated. Hydro-climatic variables, like precipitation and relative humidity, were the most important factors for controlling tree growth. Changes in rainwater input affected radial increment rates and daily amplitudes of stem diameter variations within the study period from 2009 to 2013. El Niño Southern Oscillation (ENSO) related variations of tropical Pacific Ocean sea surface temperatures influenced the trees’ increment rates. Average radial increments showed high inter-annual (up to 7.89 mm) and inter-individual (up to 3.88 mm) variations. Daily amplitudes of stem diameter variations differed strongly between the two extreme years 2009 (wet) and 2011 (dry). Contrary to 2009, the La Niña drought in 2011 caused a rapid reduction of the daily amplitudes, indicating a total cessation (‘growth collapse’) of stem increment under ENSO-related drought conditions and demonstrating the high impact of climatic extreme events on carbon sequestration of the dry tropical forest ecosystem.

     

Modelling Growth-Competition Relationships in Trembling Aspen and White Spruce Mixed Boreal Forests of Western Canada

We examined the effect of competition on stem growth of Picea glauca and Populus tremuloides in boreal mixedwood stands during the stem exclusion stage. We combined traditional approaches of collecting competition data with dendrochronology to provide retrospective measurements of stem diameter growth. Several competition indices including stand basal area (BA), the sum of stem diameter at breast height (SDBH), and density (N) for the broadleaf and coniferous species, as well as similar indices considering only trees with diameters greater than each subject (BAGR, SDBHGR, and NGR), were evaluated. We used a nonlinear mixed model to characterize the basal area increment over the past 5, 10, 15, 20, 25, 30, and 35 years as a function of growth of nearby dominant trees, the size of the subject trees, deciduous and coniferous competition indices, and ecoregions. SDBHGR and BAGR were better predictors for spruce, and SDBHGR and NGR were better for aspen, respectively, than other indices. Results showed strongest correlations with long-term stem growth, as the best models integrated growth for 10–25 years for aspen and ≥25 for spruce. Our model demonstrated a remarkable capability (adjusted R²>0.67) to represent this complex variation in growth as a function of site, size and competition.     

Canopy Interception for a Tallgrass Prairie under Juniper Encroachment

Rainfall partitioning and redistribution by canopies are important ecohydrological processes underlying ecosystem dynamics. We quantified and contrasted spatial and temporal variations of rainfall redistribution for a juniper (Juniperus virginiana, redcedar) woodland and a tallgrass prairie in the south-central Great Plains, USA. Our results showed that redcedar trees had high canopy storage capacity (S) ranging from 2.14 mm for open stands to 3.44 mm for closed stands. The canopy funneling ratios (F) of redcedar trees varied substantially among stand type and tree size. The open stands and smaller trees usually had higher F values and were more efficient in partitioning rainfall into stemflow. Larger trees were more effective in partitioning rainfall into throughfall and no significant changes in the total interception ratios among canopy types and tree size were found. The S values were highly variable for tallgrass prairie, ranging from 0.27 mm at early growing season to 3.86 mm at senescence. As a result, the rainfall interception by tallgrass prairie was characterized by high temporal instability. On an annual basis, our results showed no significant difference in total rainfall loss to canopy interception between redcedar trees and tallgrass prairie. Increasing structural complexity associated with redcedar encroachment into tallgrass prairie changes the rainfall redistribution and partitioning pattern at both the temporal and spatial scales, but does not change the overall canopy interception ratios compared with unburned and ungrazed tallgrass prairie. Our findings support the idea of convergence in interception ratio for different canopy structures under the same precipitation regime. The temporal change in rainfall interception loss from redcedar encroachment is important to understand how juniper encroachment will interact with changing rainfall regime and potentially alter regional streamflow under climate change.     

寒温带兴安落叶松林土壤温室气体通量的时间变异

采用静态箱/气相色谱(GC)法,对寒温带兴安落叶松林区6-9月生长季土壤CO2、CH4和N2O通量进行原位测定,研究了土壤温室气体通量的季节和昼夜变化及其与环境因子的关系.结果表明:在生长季,兴安落叶松林土壤为大气CH4的汇,吸收通量为22.3～107.8 μg CH4-C·m-2·h-1,6-9月月均甲烷吸收通量为(34.0±7.1)、(71.4±9.4)、(86.3±7.9)和(40.7-±6.2) μg·m-2·h-1;不同季节土壤CH4昼夜通量的变化规律相同,一天中均在10:00达到最大吸收高峰.土壤CO2日通量呈明显的双峰曲线,月均CO2通量大小顺序为7月＞8月＞6月＞9月.土壤N2O通量变异较大,在-9.1 ～31.7μg·m-2·h-1之间.土壤温度和湿度是影响CO2和CH4通量的重要因子,N2O通量主要受温度的影响.在兴安落叶松林区,10:00左右观测获得的温室气体地-气交换通量,经矫正后可以代表当日气体通量.