Contrasting leaf chemical traits in tropical lianas and trees: implications for future forest composition

Lianas are an important growthform in tropical forests, and liana abundance and biomass may be increasing in some regions. Explanations for liana proliferation hinge upon physiological responses to changing resource conditions that would favour them over trees. Testing a chemical basis for such responses, we assessed 22 foliar traits in 778 lianas and 6496 trees at 48 tropical forest sites. Growthform differences in chemical allocation occurred on a leaf mass and area basis. Light capture-growth and maintenance-metabolism chemicals averaged 14.5 and 16.7% higher mass-based concentration in lianas than in trees globally, whereas structure and defence chemicals averaged 9.0% lower in lianas. Relative differences in chemical allocation by lianas and trees were mediated by climate with peak differences at about 2500 mm year(-1) and 25 °C. Differences in chemical traits suggest that liana expansion could be greatest in forests undergoing increased canopy-level irradiance via disturbance and climate change.     

海南岛热带山地雨林幼苗幼树光合与叶氮、叶磷及比叶面积的关系

以海南岛吊罗山热带山地雨林101个物种的幼苗幼树为试验材料,测定其光合、叶片氮、磷含量及比叶面积;检验其相关关系,并按乔木,乔灌木(小乔木至大灌木)和灌木3个生活型组进行分组检验。研究结果表明,单位叶面积(Aarea)和单位叶重量的光合速率(Amass)均表现出灌木>乔木>乔灌木,方差分析表明,灌木和乔灌木之间Aarea差异显著;灌木和乔木以及灌木和乔灌木之间Amass差异显著(p<0.05)。Aarea与叶氮含量之间的相关性在不同生态型组和所有物种之间均达到极显著水平(p<0.0001);与叶磷之间的相关关系在灌木(p=0.0038),乔灌木(p=0.0002)以及所有物种(p<0.0001)之间达到极显著水平,但是在乔木中未达到显著水平(p>0.05);与SLA之间在灌木(p=0.0006)、乔木(p<0.0001)和所有物种(p<0.0001)之间达到极显著水平,但是在乔灌木中未达到显著水平(p>0.05)。Amass与叶片氮含量、SLA的相关关系在不同生活型组和所有物种中都达到极显著水平(p<0.0001);与叶磷含量之间的相关性在灌木(p=0.0004),乔灌木(p=0.0018)及所有物种(p<0.0001)中极显著,在乔木生活型组中也达显著水平(p=0.0377)。逐步回归表明,与Aarea相比,Amass估计结果更接近于实际测值。由此可见,海南岛热带山地雨林林下幼苗幼树的光合和叶氮、磷含量及SLA之间相关关系与基于成树的研究非常相似,并且A比A更能稳定体现这种相关性。     

Contrasting leaf phenology of woody species of dry tropical forest

We studied selected leaf traits [leaf area (LA), leaf water content (LWC), leaf fresh weight (LFW), leaf dry weight (LDW), specific leaf area (SLA) and chlorophyll content] of eight woody species (Shorea robusta, Buchanania lanzan, Diospyros melanoxylon, Lagerstroemia parviflora, Lannea coromandelica, Terminalia tomentosa, Holarrhena antidysenterica and Lantana camara) dominant at four sites in a dry tropical deciduous forest over complete two annual cycles (2008–2010). Our results showed that leaf traits varied across species (1.7–11.5 fold), months (1.2–1.5 fold) and sites (1.1–1.3 fold). However, leaf traits showed smaller variation between sites than between species. Leaf lifespan varied from 7 months (L. coromandelica) to 12 months (S. robusta). On the same sites, species differed in the length of deciduous period. The maximum LA, LDW, LFW and LWC were recorded for the semi-evergreen species, SLA for long-deciduous species and chlorophyll content for short-deciduous species, respectively. The coefficient of variation was maximum for LDW and minimum for chlorophyll content. Among the eight woody species, T. tomentosa exhibited the greatest LA, LDW, LFW and LWC. LA, LWC, LFW, LDW, SLA, LD, SD, MD & SE confirm.     

Above-ground biomass investments and light interception of tropical forest trees and lianas early in succession

BACKGROUND AND AIMS: Crown structure and above-ground biomass investment was studied in relation to light interception of trees and lianas growing in a 6-month-old regenerating forest. METHODS: The vertical distribution of total above-ground biomass, height, diameter, stem density, leaf angles and crown depth were measured for individual plants of three short-lived pioneers (SLPs), four long-lived pioneers (LLPs) and three lianas. Daily light interception per individual Phi(d) was calculated with a canopy model. The model was then used to estimate light interception per unit of leaf mass (Phi(leaf mass)), total above-ground mass (Phi(mass)) and crown structure efficiency (E(a), the ratio of absorbed vs. available light). KEY RESULTS: The SLPs Trema and Ochroma intercepted higher amounts of light per unit leaf mass (Phi(leaf mass)) because they had shallower crowns, resulting in higher crown use efficiency (E(a)) than the other species. These SLPs (but not Cecropia) were also taller and intercepted more light per unit leaf area (Phi(area)). LLPs and lianas had considerably higher amounts of leaf mass and area per unit above-ground mass (LMR and LAR, respectively) and thus attained Phi(mass) values similar to the SLPs (Phi(mass)=Phi(area)xLAR). Lianas, which were mostly self-supporting, had light interception efficiencies similar to those of the trees. CONCLUSIONS: These results show how, due to the trade-off between crown structure and biomass allocation, SLPs, and LLPs and lianas intercept similar amount of light per unit mass which may contribute to the ability of the latter two groups to persist.     

Nitrogen and phosphorus resorption in trees of a Mexican tropical dry forest

Resorption efficiency (RE) and proficiency, foliar nutrient concentrations, and relative soil nutrient availability were determined during 3 consecutive years in tree species growing under contrasting topographic positions (i.e., top vs. bottom and north vs. south aspect) in a tropical dry forest in Mexico. The sites differed in soil nutrient levels, soil water content, and potential radiation interception. Leaf mass per area (g m^–2) increased during the growing season in all species. Soil P availability and mean foliar P concentrations were generally higher at the bottom than at the top site during the 3 years of the study. Leaf N concentrations ranged from 45.4 to 31.4 mg g^–1. Leaf P varied from 2.3 to 1.8 mg g^–1. Mean N and P RE varied among species, occasionally between top and bottom sites, and were higher in the dry than in the wet years of study. Senesced-leaf nutrient concentrations (i.e., a measure of resorption proficiency) varied from 13.7 to 31.2 mg g^–1 (N) and 0.4 to 3.3 mg g^–1 (P) among the different species and were generally indicative of incomplete nutrient resorption. Phosphorus concentrations in senesced leaves were higher at the bottom than at the top site and decreased from the wettest to the the driest year. Soil N and P availability were significantly different in the north- and south-facing slopes, but neither nutrient concentrations of mature and senesced leaves nor RE differed between aspects. Our results suggest that water more than soil nutrient availability controls RE in the Chamela dry forest, while resorption proficiency may be interactively controlled by both nutrient and water availability.     

Thermal acclimation of leaf respiration of tropical trees and lianas: response to experimental canopy warming,and consequences for tropical forest carbon balance

Climate warming is expected to increase respiration rates of tropical forest trees and lianas, which may negatively affect the carbon balance of tropical forests. Thermal acclimation could mitigate the expected respiration increase, but the thermal acclimation potential of tropical forests remains largely unknown. In a tropical forest in Panama, we experimentally increased nighttime temperatures of upper canopy leaves of three tree and two liana species by on average 3  ° C for 1 week, and quantified temperature responses of leaf dark respiration. Respiration at 25  ° C (R₂₅) decreased with increasing leaf temperature, but acclimation did not result in perfect homeostasis of respiration across temperatures. In contrast, Q₁₀ of treatment and control leaves exhibited similarly high values (range 2.5–3.0) without evidence of acclimation. The decrease in R₂₅ was not caused by respiratory substrate depletion, as warming did not reduce leaf carbohydrate concentration. To evaluate the wider implications of our experimental results, we simulated the carbon cycle of tropical latitudes (24 ° S–24 ° N) from 2000 to 2100 using a dynamic global vegetation model (LM3VN) modified to account for acclimation. Acclimation reduced the degree to which respiration increases with climate warming in the model relative to a no‐acclimation scenario, leading to 21% greater increase in net primary productivity and 18% greater increase in biomass carbon storage over the 21st century. We conclude that leaf respiration of tropical forest plants can acclimate to nighttime warming, thereby reducing the magnitude of the positive feedback between climate change and the carbon cycle.     

The relationship between tree biodiversity and biomass dynamics changes with tropical forest succession

Theory predicts shifts in the magnitude and direction of biodiversity effects on ecosystem function (BEF) over succession, but this theory remains largely untested. We studied the relationship between aboveground tree biomass dynamics (Δbiomass) and multiple dimensions of biodiversity over 8–16 years in eight successional rainforests. We tested whether successional changes in diversity–Δbiomass correlations reflect predictions of niche theories. Diversity–Δbiomass correlations were positive early but weak later in succession, suggesting saturation of niche space with increasing diversity. Early in succession, phylogenetic diversity and functional diversity in two leaf traits exhibited the strongest positive correlations with Δbiomass, indicating complementarity or positive selection effects. In mid‐successional stands, high biodiversity was associated with greater mortality‐driven biomass loss, i.e. negative selection effects, suggesting successional niche trade‐offs and loss of fast‐growing pioneer species. Our results demonstrate that BEF relationships are dynamic across succession, thus successional context is essential to understanding BEF in a given system.     

Spatial and temporal dynamics of forest canopy gaps following selective logging in the eastern Amazon

Selective logging is a dominant form of land use in the Amazon basin and throughout the humid tropics, yet little is known about the spatial variability of forest canopy gap formation and closure following timber harvests. We established chronosequences of large‐area (14–158 ha) selective logging sites spanning a 3.5‐year period of forest regeneration and two distinct harvest methods: conventional logging (CL) and reduced‐impact logging (RIL). Our goals were to: (1) determine the spatial characteristics of canopy gap fraction immediately following selective logging in the eastern Amazon; (2) determine the degree and rate of canopy closure in early years following harvest among the major landscape features associated with logging – tree falls, roads, skid trails and log decks; and (3) quantify spatial and temporal differences in canopy opening and closure in high‐ and low‐damage harvests (CL vs. RIL). Across a wide range of harvest intensities (2.6–6.4 felled trees ha^?1), the majority of ground damage occurred as skid trails (4–12%), whereas log decks and roads were only a small contributor to the total ground damage (<2%). Despite similar timber harvest intensities, CL resulted in more ground damage than RIL. Neither the number of log decks nor their individual or total area was correlated with the number of trees removed or intensity of tree harvesting (trees ha^?1). The area of skids was well correlated with the ground area damaged (m²) per tree felled. In recently logged forest (0.5 years postharvest), gap fractions were highest in log decks (mean RIL=0.83, CL=0.99) and lowest in tree‐fall areas (RIL: 0.26, CL: 0.41). However, the small surface area of log decks made their contribution to the total area‐integrated forest gap fraction minor. In contrast, tree falls accounted for more than two‐thirds of the area disturbed, but the canopy gaps associated with felled trees were much smaller than for log decks, roads and skids. Canopy openings decreased in size with distance from each felled tree crown. At 0.5 years postharvest, the area initially affected by the felling of each tree was approximately 100 m in radius for CL and 50 m for RIL. Initial decreases in gap fraction during the first 1.5 years of regrowth diminished in subsequent years. Throughout the 3.5‐year period of forest recovery, tree‐fall gap fractions remained higher in CL than in RIL treatments, but canopy gap closure rates were higher in CL than in RIL areas. During the observed recovery period, the canopy gap area affected by harvesting decreased in radius around each felled tree from 100 to 40 m in CL, and from 50 to 10 m in RIL. The results suggest that the full spatial and temporal dynamics of canopy gap fraction must be understood and monitored to predict the effects of selective logging on regional energy balance and climate regimes, biogeochemical processes including carbon cycling, and plant and faunal population dynamics. This paper also shows that remote sensing of log decks alone will not provide an accurate assessment of total forest area impacted by selective logging, nor will it be closely correlated to damage levels and canopy gap closure rates.     

Relative effects of different leaf attributes on sapling growth in tropical dry forest

Aims Soil moisture content (SMC) influences establishment, survival and development of plant species and is considered as the most important limiting factor in tropical dry forest (TDF). In this study we attempt to establish the relationship between leaf attributes and of tree saplings in TDF and address the following questions: (i) how are the functional attributes of dominant tree saplings of TDF affected by seasonal changes in SMC at different habitats?, (ii) what is the relationship of functional attributes with each other?, (iii) how are the functional attributes and their plasticity affected by habitat conditions? and (iv) can the functional attributes in single or in combination predict the growth rate of tree saplings of TDF? The study was conducted on four sites (Hathinala, Gaighat, Harnakachar and Ranitali, listed in order of decreasing SMC) within the tropical dry deciduous forest in northern India.Methods We analysed eight leaf attributes, specific leaf area (SLA); leaf dry matter content (LDMC); leaf nitrogen concentration (leaf N); leaf phosphorus concentration (leaf P); chlorophyll concentration (Chl); mass-based photosynthetic rate (A mass); mass-based stomatal conductance (Gs mass); intrinsic water use efficiency (WUEi) and three growth attributes, relative diameter increment (RDI); relative height increment (RHI); relative growth rate (RGR) of the 10 dominant tree saplings (viz., Acacia catechu, Anogeissus latifolia, Boswellia serrata, Buchanania lanzan, Diospyros melanoxylon, Hardwickia binata, Lagerstroemia parviflora, Lannea coromandelica, Shorea robusta and Terminalia tomentosa) of a TDF and observed the effects of site, season and species for a period of 2 years. Saplings were selected in gradients of deciduousness. Step-wise multiple regression was performed to predict RDI, RHI and RGR from mean values of SMC and leaf attributes.Important findings All the 11 attributes were interrelated and differed significantly among the 10 saplings. Species response varied across sites and seasons. Across the SMC gradient, the attributes showed variable plasticity that differed across species. Among the 10 saplings, the highly deciduous Boswellia serrata showed the maximum plasticity in seven functional attributes. According to the step-wise multiple regressions, 65% variability in RDI and 67% variability in RGR were due to Gs mass, and for RHI, 61% variability was due to A mass. SMC and the other attributes, viz ., SLA, Chl, WUEi and LDMC in combination could contribute only for ~2–6% of the variability in RDI, RHI and RGR, which indicates that other traits/factors, not accounted in this study are also important in modulating the growth of tree saplings in TDFs. In conclusion, growth of the tree saplings in the tropical dry environment is determined by soil moisture, whereas the response of saplings of different tree species is modulated by alterations in key functional attributes such as SLA, Chl, WUEi and LDMC.     

尖峰岭热带山地雨林林冠层乔木某些功能性状的系统发育信号、关联性及其演化模式

热带雨林林冠层具有丰富的物种、功能和系统发育(谱系)多样性,能够显著影响生态系统生物地球化学循环,调节大气水热平衡,缓解气候变化与人为干扰的负面作用。因此热带雨林林冠层功能和谱系生态学研究是目前群落生态学研究的热点之一。选取海南尖峰岭热带山地雨林3个1 hm2样地的林冠层为研究对象,利用样地林冠树种清查数据基于APG III系统发育结构重建样地林冠层系统发育树。利用Blomber's K对和Pagel'sλ指标结合系统发育独立性比较法对组成林冠层乔木树种的结构、化学计量和水力学功能性状(11个性状)的系统发育信号、关联性和演化模式进行研究,以探讨物种亲缘关系对尖峰岭热带山地雨林林冠层多维功能性状关联性及其动态演化的影响。结果表明,有8个功能性状(叶面积、叶厚度和潜在最大高度除外)具有显著的系统发育信号(通过P0.05的显著性检验),Blomber's K值和Pagel'sλ值分别介于0.202—0.392和0.277—0.847之间,表明尖峰岭热带山地雨林林冠层功能性状普遍存在系统发育保守性。在系统发育背景下,林冠层乔木树种结构、化学和水力学功能性状在物种水平普遍存在显著关联性(P0.05),表现出趋同或趋异进化;而且林冠层乔木随物种分化其功能性状分化模式大致呈水平"漏斗"状,姐妹类群功能性状间差异性在物种分化早期(大约1亿2千万年前至6000万年前)明显小于其在中后期(大约6000万年前至今)的差异性,导致了林冠层性状空间在中后期迅速膨大。然而林冠层主要功能性状在系统发育树内部每一节点上姊妹类群分化产生的系统发育独立性比较值绝大部分与零模型随机模拟值并无显著性差异。了解系统发育背景下林冠层功能性状的权衡关系及其随物种演化的分化模式与时间动态为进一步探究热带雨林林冠生态系统功能发挥奠定基础。     

Stem water storage and diurnal patterns of water use in tropical forest canopy trees

Stem water storage capacity and diurnal patterns of water use were studied in five canopy trees of a seasonal tropical forest in Panama. Sap flow was measured simultaneously at the top and at the base of each tree using constant energy input thermal probes inserted in the sapwood. The daily stem storage capacity was calculated by comparing the diurnal patterns of basal and crown sap flow. The amount of water withdrawn from storage and subsequently replaced daily ranged from 4 kg d^–1 in a 0·20-m-diameter individual of Cecropia longipes to 54 kg d^–1 in a 1·02-m-diameter individual of Anacardium excelsum, representing 9–15% of the total daily water loss, respectively. Ficus insipida, Luehea seemannii and Spondias mombin had intermediate diurnal water storage capacities. Trees with greater storage capacity maintained maximum rates of transpiration for a substantially longer fraction of the day than trees with smaller water storage capacity. All five trees conformed to a common linear relationship between diurnal storage capacity and basal sapwood area, suggesting that this relationship was species-independent and size-specific for trees at the study site. According to this relationship there was an increment of 10 kg of diurnal water storage capacity for every 0·1 m² increase in basal sapwood area. The diurnal withdrawal of water from, and refill of, internal stores was a dynamic process, tightly coupled to fluctuations in environmental conditions. The variations in basal and crown sap flow were more synchronized after 1100 h when internal reserves were mostly depleted. Stem water storage may partially compensate for increases in axial hydraulic resistance with tree size and thus play an important role in regulating the water status of leaves exposed to the large diurnal variations in evaporative demand that occur in the upper canopy of seasonal lowland tropical forests.     

Phosphorus limitation of forest leaf area and net primary production on a highly weathered soil

We tested the hypothesis that P was the nutrient limiting net primary production of a nativeMetrosideros polymorpha forest on a highly weathered montane tropical soil in Hawaii. A factorial experiment used all combinations of three fertilizer treatments: nitrogen (N), phosphorus (P) and a mix of other essential nutrients (OE), consisting primarily of mineral derived cations and excluding N and P. P addition, but not N or OE, increased leaf area index within 12 months, foliar P concentration measured at 18 months, and stem diameter increment within 18 months. Stem growth at 18 months was even greater when trees fertilized with P also received the OE treatment. N and P additions increased leaf litterfall and N and P in combination further increased litterfall. The sequence of responses suggests that increased available P promoted an increase in photosynthetic area which led to increased wood production. P was the essential element most limiting to primary production on old volcanic soil in contrast to the N limitation found on young volcanic soils.     

林冠结构和地形对亚热带常绿落叶阔叶林林下幼苗物种多样性和功能多样性的影响

森林结构和地形是森林生态系统最明显的特点,也是影响林下幼苗存活和物种多样性的关键因子。该研究采用半球面摄影方法提取八大公山生长监测样地（共1.2 hm²）林冠结构参数,通过调查地表层木本植物幼苗的组成和多度,获取常见植物幼苗叶片功能性状,结合详细的地形信息,利用空间同步自回归模型探究林冠结构变量及地形因子对幼苗物种多样性及功能多样性的影响。结果表明：（1）八大公山亚热带山地常绿落叶阔叶林林冠结构复杂度较高,最大林冠高的平均值达到19.94 m,叶面积指数、平均叶倾角和林冠覆盖度分别为2.94、30.88°和0.87;（2）林冠结构变量和地形因子能够解释32.6%~48.4%的林下幼苗物种多样性指数变异和28.5%~70.2%的功能多样性变异,但地形因子对幼苗物种多样性的影响很小;（3）预测在亚热带常绿落叶阔叶林高海拔的山坡上,有较低的叶面积指数和平均叶倾角群落有较高的幼苗物种多样性;而在低海拔山脊上,较低的叶面积指数和平均叶倾角群落林下幼苗层有较高的功能多样性。此结果对科研人员和林业工作者开展野外森林更新情况评估和样方调查将有所帮助。     

The impact of tropical forest logging and oil palm agriculture on the soil microbiome

Selective logging and forest conversion to oil palm agriculture are rapidly altering tropical forests. However, functional responses of the soil microbiome to these land‐use changes are poorly understood. Using 16S rRNA gene and shotgun metagenomic sequencing, we compared composition and functional attributes of soil biota between unlogged, once‐logged and twice‐logged rainforest, and areas converted to oil palm plantations in Sabah, Borneo. Although there was no significant effect of logging history, we found a significant difference between the taxonomic and functional composition of both primary and logged forests and oil palm. Oil palm had greater abundances of genes associated with DNA, RNA, protein metabolism and other core metabolic functions, but conversely, lower abundance of genes associated with secondary metabolism and cell–cell interactions, indicating less importance of antagonism or mutualism in the more oligotrophic oil palm environment. Overall, these results show a striking difference in taxonomic composition and functional gene diversity of soil microorganisms between oil palm and forest, but no significant difference between primary forest and forest areas with differing logging history. This reinforces the view that logged forest retains most features and functions of the original soil community. However, networks based on strong correlations between taxonomy and functions showed that network complexity is unexpectedly increased due to both logging and oil palm agriculture, which suggests a pervasive effect of both land‐use changes on the interaction of soil microbes.