Causes of reduced leaf‐level photosynthesis during strong El Niño drought in a Central Amazon forest

Sustained drought and concomitant high temperature may reduce photosynthesis and cause tree mortality. Possible causes of reduced photosynthesis include stomatal closure and biochemical inhibition, but their relative roles are unknown in Amazon trees during strong drought events. We assessed the effects of the recent (2015) strong El Niño drought on leaf‐level photosynthesis of Central Amazon trees via these two mechanisms. Through four seasons of 2015, we measured leaf gas exchange, chlorophyll a fluorescence parameters, chlorophyll concentration, and nutrient content in leaves of 57 upper canopy and understory trees of a lowland terra firme forest on well‐drained infertile oxisol. Photosynthesis decreased 28% in the upper canopy and 17% in understory trees during the extreme dry season of 2015, relative to other 2015 seasons and was also lower than the climatically normal dry season of the following non‐El Niño year. Photosynthesis reduction under extreme drought and high temperature in the 2015 dry season was related only to stomatal closure in both upper canopy and understory trees, and not to chlorophyll a fluorescence parameters, chlorophyll, or leaf nutrient concentration. The distinction is important because stomatal closure is a transient regulatory response that can reverse when water becomes available, whereas the other responses reflect more permanent changes or damage to the photosynthetic apparatus. Photosynthesis decrease due to stomatal closure during the 2015 extreme dry season was followed 2 months later by an increase in photosynthesis as rains returned, indicating a margin of resilience to one‐off extreme climatic events in Amazonian forests.     

胡杨枝芽生长特征及其展叶物候特征

以5个不同发育阶段的胡杨(Populus euphratica Oliv.)个体为研究对象,观测记录了枝芽展叶物候、枝芽生长特征和叶形变化的空间分布规律。结果表明:不同发育阶段的胡杨个体以及同一个体树冠的不同层次,其枝芽生长及其展叶物候期表现出不同的时空特征。随着树龄的增加和树冠层次的增高(由基向顶),当年新生枝条长度、枝条叶片数和叶形指数逐渐减小,但叶面积和叶片干重逐渐增大。5个不同发育阶段胡杨个体均表现出展叶物候始于树冠顶层,依次向下结束于树冠基部;展叶物候期共性表现在枝芽萌动期均在4月上旬,起始展叶期集中在4月中旬,展叶终期则在5月上旬到下旬;树龄较大的个体其枝芽萌动期、起始展叶期、展叶终期较树龄较小的个体早;其枝芽萌动期到展叶终期的时间进程较树龄较小的个体短;不同发育阶段的个体枝芽萌动期出现的时间较为离散,起始展叶期和展叶终期出现的时间较为集中。相关分析表明,出叶周期与枝条长度、枝条叶片数量和叶形指数呈极显著正相关,与叶面积和叶片干重呈显著负相关。     

Trade-offs between light interception and leaf water shedding: a comparison of shade- and sun-adapted species in a subtropical rainforest

Species in high-rainfall regions have two major alternative approaches to quickly drain off water, i.e., increasing leaf inclination angles relative to the horizontal plane, or developing long leaf drip tips. We hypothesized that shade-adapted species will have more pronounced leaf drip tips but not greater inclination angles (which can reduce the ability to intercept light) compared to sun-adapted species and that length of leaf drip tips will be negatively correlated with photosynthetic capacity [characterized by light-saturated net photosynthetic rates (A _max), associated light compensation points (LCP), and light saturation points (LSP)]. We tested this hypothesis by measuring morphological and physiological traits that are associated with light-interception and water shedding for seven shade-adapted shrub species, ten sun-adapted understory shrub species, and 15 sun-adapted tree species in a subtropical Chinese rainforest, where mean annual precipitation is around 1,600 mm. Shade-adapted understory species had lower LMA, A _max, LSP, and LCP compared to understory or canopy sun-adapted species; their leaf and twig inclination angles were significantly smaller and leaf drip tips were significantly longer than those in sun-adapted species. This suggests that shade-adapted understory species tend to develop pronounced leaf drip tips but not large leaf inclination angles to shed water. The length of leaf drip tips was negatively correlated with leaf inclination angles and photosynthetic capacity. These relationships were consistent between ordinary regression and phylogenetic generalized least squares analyses. Our study illustrates the trade-offs between light interception and leaf water shedding and indicates that length of leaf drip tips can be used as an indicator of adaptation to shady conditions and overall photosynthetic performance of shrub species in subtropical rainforests.     

Growth Response of a Chrysanthemum Crop to the Environment. III, Effects of Radiation and Temperature on Dry Matter Partitioning and Photosynthesis

Vegetative crops of chrysanthemum were grown for 5 or 6 weekperiods in daylit assimilation chambers. Crop responses to differentradiation levels and temperatures were analysed into effectson dry matter partitioning, specific leaf area, leaf photosynthesisand canopy light interception. The percentage of newly formed dry matter partitioned to theleaves was almost constant, although with increasing radiationor decreasing temperature, a greater percentage of dry matterwas partitioned to stem tissue at the expense of root tissue.There was a positive correlation between the percentage of drymatter in shoot material and the overall carbon: dry matterratio. Canopy photosynthesis was analysed assuming identical behaviourfor all leaves in the crop. Leaf photochemical efficiency wasonly slightly affected by crop environment. The rate of grossphotosynthesis per unit leaf area at light saturation, P_A (max),increased with increasing radiation integral, but the same parameterexpressed per unit leaf dry matter, P_w (max) was almost unaffectedby growth radiation. In contrast, P_A (max) was hardly affectedby temperature but P_w (max) increased with increasing growthtemperature. This was because specific leaf area decreased withdecreasing temperature and increased with decreasing radiation.There was a positive correlation between canopy respirationintegral and photosynthesis integral, and despite a four-foldchange in crop mass during the experiments, the maintenancecomponent of canopy respiration remained small and constant. Canopy extinction coefficient showed no consistent variationwith radiation integral but was negatively correlated with temperature.This decrease in the efficiency of the canopy at interceptingradiation exactly cancelled the increase in specific carbonassimilation rate that occurred with increasing growth temperature,giving a growth rate depending solely on the incident lightlevel. Chrysanthemum, dry matter partitioning, photosynthesis, specific leaf area     

华北落叶松人工林蒸散及产流对叶面积指数变化的响应

定量评价林地蒸散和产流等水文过程对冠层叶面积指数(LAI)的响应,对于深入认识森林植被的生态水文过程及其发生机制,实现半干旱区林水综合管理和区域可持续发展是非常必要的。应用集总式生态水文模型BROOK90,模拟分析了不同降水年型(丰水年、平水年、枯水年)下,位于半干旱区的宁夏六盘山叠叠沟小流域内华北落叶松(Larix principis-rupprechtii)人工林的水文过程对冠层LAI变化的响应关系。结果发现:林分总蒸散量、冠层截留量、蒸腾量与LAI都呈显著的正相关关系(R~20.99,P0.01),而土壤蒸发量、产流量则与LAI均呈显著的负相关关系(R~20.99,P0.01);在不同的降水年型下,各水文过程变量与LAI的关系都可以很好地用指数函数来表达,且都存在着一个LAI阈值。当LAI低于阈值时,各水文过程变量随LAI的变化幅度较大;但高于阈值时,各变量的变化十分缓慢并趋于稳定。在不同降水年型下,各变量LAI阈值之间存在着一定的差异。一般地,丰水年各变量的LAI阈值要大于枯水年,尤其是冠层截留和土壤蒸发。在丰水年,各水文过程变量随LAI增加而变化的速率要比在平水年、枯水年更快,说明在水分充足年份中各变量的波动更多取决于LAI变化,而在水分亏缺的年份中则可能更多地受到水分条件的限制。模拟结果表明,通过减少冠层LAI(如间伐)导致的林分的降低蒸散耗水和增加产流的作用是有限的,这是由于林分蒸散降低的幅度要比LAI降低的幅度小。例如,在平水年,当LAI从4.2变为2.0(减少幅度52.4%)时,林分年蒸散仅从357.2 mm减少至333.9 mm(减少幅度6.5%)。     

不同光照梯度的遮荫处理对绒毛番龙眼幼苗生长的影响

在不同光照梯度,即100%自然全光照(natural sunlight,NS)、37.3%NS、15.5%NS、4.2%NS、1.6%NS和0.6%NS的人工遮荫条件下,研究了西双版纳季节雨林冠层树种绒毛番龙眼(Pometia tomentosa)幼苗的早期生长和定居后的生长特点。结果表明,光照是影响幼苗生长的重要环境因子。生长早期的幼苗基径和复叶数随遮荫程度的增加而降低;主根长、根冠比、总干重和单株叶面积均以37.3%NS处理最大;比叶面积随遮荫程度的增加而增大,而相对生长率则降低;幼苗株高在0.6%NS下增长最快,表明种子中贮藏的营养物质对幼苗的早期生长可能具有重要作用。37.3%NS处理对定居后绒毛番龙眼幼苗的生长最有利,幼苗的株高、基径、复叶数、叶轴长、复叶最多小叶数、单株叶面积、相对生长率和净同化率均在37.3%NS处理下获得最大增长;幼苗总干重随光照强度的减弱而降低;比叶面积在15.5%NS处理时最大。幼苗比叶面积和根冠比在生长过程中的波动可能是光照和土壤水分共同作用的结果。     

Physiological comparisons of two soybean cultivars differing in canopy photosynthesis. I. Variation in vertical 14CO2 labelling and dry weight partitioning

Differences in canopy apparent photosynthesis (CAP) among soybean [Glycine max (L.) Merr.] genotypes have been shown to be correlated to seed yields. Since the physiological basis for such differences in CAP is unknown, two cultivars known to differ in CAP, Tracy and Davis, were studied during the 1978–1980 growing seasons. The CAP and dry weights of component plant parts were determined. In 1978 and 1979, ¹⁴CO₂ uptake by vertical leaf strata was determined and specific leaf weight (SLW) and leaf area index (LAI) were determined for corresponding strata in 1979 and 1980. Measurements were taken on several dates during reproductive growth. With the exception of CAP, all measurements (¹⁴C uptake, dry weights) were made in layers within the canopy. CAP on some dates were significantly higher in Tracy than in Davis and integrated CAP values from a certain growth period, labeled as R5 to R7, averaged 16 percent higher in Tracy for the three years studied. No differences in the relative recovery of ¹⁴C from different layers of leaves in the canopy were found. This indicates that variations in canopy structure or leaf orientation did not play a major role in the CAP differences between cultivars. The differences seem related to variations in leaf dry weights. Overall, Tracy exhibited 13.5, 19.2, and 13.2 percent greater leaf dry weights than Davis during 1978, 1979, and 1980, respectively. These differences in leaf dry weight seem largely due to a differences in the SLW. Data from these experiments indicate that differences in soybean CAP values were associated with differences in SLW.Abbreviations CAP Canopy Apparent Photosynthesis - CER Carbondioxide Exchange Rates - EST Eastern Standard Time - LAI Leaf Area Index - LSD Least Significant Difference - POPOP 1,4-bis-[2(5-phenyloxazdyl)]-benzene - PPO 2,5-diphenyloxazole - SLW Specific Leaf Weight     

Ecological differentiation in xylem cavitation resistance is associated with stem and leaf structural traits

Cavitation resistance is a critical determinant of drought tolerance in tropical tree species, but little is known of its association with life history strategies, particularly for seasonal dry forests, a system critically driven by variation in water availability. We analysed vulnerability curves for saplings of 13 tropical dry forest tree species differing in life history and leaf phenology. We examined how vulnerability to cavitation (P₅₀) related to dry season leaf water potentials and stem and leaf traits. P₅₀‐values ranged from ?0.8 to ?6.2 MPa, with pioneers on average 38% more vulnerable to cavitation than shade‐tolerants. Vulnerability to cavitation was related to structural traits conferring tissue stress vulnerability, being negatively correlated with wood density, and surprisingly maximum vessel length. Vulnerability to cavitation was negatively related to the Huber‐value and leaf dry matter content, and positively with leaf size. It was not related to SLA. We found a strong trade‐off between cavitation resistance and hydraulic efficiency. Most species in the field were operating at leaf water potentials well above their P₅₀, but pioneers and deciduous species had smaller hydraulic safety margins than shade‐tolerants and evergreens. A trade‐off between hydraulic safety and efficiency underlies ecological differentiation across these tropical dry forest tree species.     

Vegetative Phenology and Growth of a Facultatively Deciduous Bamboo in a Monsoonal Climate1

Canopy closure, leaf flush, and ramet recruitment in Bambusa arnhemica, a semelparous, clumping bamboo from the Australian monsoonal tropics, were monitored monthly for 2.5 years at three sites along a flood gradient. Bambusa arnhemica was facultatively deciduous, remaining evergreen at a downslope riparian site but suffering total loss of canopy on a hillside for up to 4 mo during the dry season. Leaf flush was flexible, occurring after as little as 25 mm of rain at the onset of wet season, in response to unusual dry season storms, and apparently also in response to fire independent of rainfall. New culms emerged soon after leaf flush early in the wet season. Culm growth took place during the middle and late wet season, with peak elongation rates of 15–30 cm/day. Some growth continued into the dry season, mostly on branches and leaves of new culms at riparian sites. Not all culms completed elongation before the onset of the dry season, and those that did not were permanently stunted. The demands of culm elongation may limit the occurrence of bamboo in wet‐dry climates to areas with predictable and sustained wet season rainfall, but the flexibility of branching and leaf processes facilitates coping with, and permits exploitation of less predictable pre‐ and postmonsoonal rains. The bamboo growth form and phenological patterns differ markedly from those of dicotyledonous trees and shrubs.     

Mean residence time of leaf number, area, mass, and nitrogen in canopy photosynthesis

Mean residence time (MRT) of plant nitrogen (N), which is an indicator of the expected length of time N newly taken up is retained before being lost, is an important component in plant nitrogen use. Here we extend the concept MRT to cover such variables as leaf number, leaf area, leaf dry mass, and nitrogen in the canopy. MRT was calculated from leaf duration (i.e., time integral of standing amount) divided by the total production of leaf variables. We determined MRT in a Xanthium canadense stand established with high or low N availability. The MRT of leaf number may imply longevity of leaves in the canopy. We found that the MRT of leaf area and dry mass were shorter than that of leaf number, while the MRT of leaf N was longer. The relatively longer MRT of leaf N was due to N resorption before leaf shedding. The MRT of all variables was longer at low N availability. Leaf productivity is the rate of canopy photosynthesis per unit amount of leaf variables, and multiplication of leaf productivity by MRT gives the leaf photosynthetic efficiency (canopy photosynthesis per unit production of leaf variables). The photosynthetic efficiency of leaf number implies the lifetime carbon gain of a leaf in the canopy. The analysis of plant-level N use efficiency by evaluating the N productivity and MRT is a well-established approach. Extension of these concepts to leaf number, area, mass, and N in the canopy will clarify the underlying logic in the study of leaf life span, leaf area development, and dry mass and N use in canopy photosynthesis.     

Differences in leaf phenology between juvenile and adult individuals of two tree species in a seasonally dry tropical woodland

In temperate forests, juvenile trees anticipate leaf phenology compared to adults, thus avoiding shading and herbivory. This is also expected to occur in seasonal tropical forests due to intense herbivory and shading during the rainy season; however, the anticipation of leaf phenology by juveniles in seasonal tropical forests has yet to be demonstrated. Stem‐succulent species are expected to be prone to juvenile phenological anticipation because these species are able to use water stored in their stems for leaf flushing in the dry season. We investigated this hypothesis by comparing leaf phenology (bud break, leaf expansion) of juveniles and adults of two species with contrasting wood densities in the transition between dry and rainy seasons in a tropical dry woodland. We also investigated the level of light limitation that juveniles experience in the rainy season. Both species exhibited bud break during the dry season, but only expanded their leaves with the occurrence of the first rains. In general, the stem‐succulent species had a more precocious bud break; however, anticipation by juveniles occurred only in the species with more dense wood. Canopy openness was lower than in temperate deciduous forests, but the fact that the full expansion of leaves occurred only with rainfall indicates that bud break in anticipation of canopy closure contributes only to keeping leaf photosynthetic balance from going negative, and not to higher carbon gain. The importance of anticipated budding for escaping herbivory remains an alternative explanation in need of investigation.     

Growth dynamics of root and shoot hydraulic conductance in seedlings of five neotropical tree species: scaling to show possible adaptation to differing light regimes

The dynamics of growth (shoot and root dry weights, surface areas, hydraulic conductances, and root length) were measured in seedlings of five neotropical tree species aged 4–16 months. The species studied included two light-demanding pioneers (Miconia argentea and Apeiba membranacea) and three shade-tolerant young- or old-forest species (Pouteria reticulata, Gustavia superba, and Trichilia tuberculata). Growth analysis revealed that shoot and root dry weights and hydraulic conductances and leaf area all increased exponentially with time. Alternative methods of scaling measured parameters to reveal differences that might explain adaptations to microsites are discussed. Scaling root conductance to root surface area or root length revealed a few species differences but nothing that correlated with adaptation to light regimes. Scaling of root surface area or root length to root dry weight revealed that pioneers produced significantly more root area and length per gram dry weight investment than shade-tolerant species. Scaling of root and shoot hydraulic conductances to leaf area and scaling of root conductance to root dry weight and shoot conductance to shoot dry weight also revealed that pioneers were significantly more conductive to water than shade-tolerant species. The advantages of scaling hydraulic parameters to leaf surface area are discussed in terms of the Ohm's law analogue of water flow in plants. Received: 24 March 1997 / Accepted: 17 November 1997     

Specific leaf area as a superior predictor of changes in field layer abundance during forest succession

Question : How accurately can a suite of suggested functional traits predict plant species response to succession from semi‐open woodland to closed deciduous canopy forest? Location : Southeastern Sweden. Methods : Abundance of 46 field‐layer plant species in a temperate deciduous forest, measured as frequency of occupied plots, was estimated in 1961, 1970 and 2003. Abundance change over time across species was tested for correlations with functional traits and literature information on habitat preference. Results : Increase in abundance was positively correlated with specific leaf area (SLA), weakly negatively correlated with seed mass and not significantly correlated with plant height or start, peak and length of the flowering period. Change in abundance was correlated with the Ellenberg light indicator value, whereas no correlations were found with Ellenberg values for nitrogen, calcium and moisture, or forest preference according to the literature. Conclusions : SLA was a better predictor of how field layer plants responded to succession from semi‐open woodland to closed canopy forest than empirically‐derived measures of habitat preference. The same holds for SLA in relation to seed size, indicating that interactions in the established life‐cycle phase are more important than the recruitment phase for species response to succession.     

Impact of tree leaf phenology on growth rates and reproduction in the spring flowering species Trillium erectum (Liliaceae)

We investigated the impact of overstory tree leaf phenology on growth rates, carbon allocation pattern, and fruit characteristics in the spring flowering species, Trillium erectum (Liliaceae). Air temperature, overstory canopy closure, and T. erectum phenology were monitored at three locations following a latitudinal gradient in Québec, Canada. Northern sugar maple trees leaf out at cooler temperatures than more southern populations, while Trillium development was initiated at the same soil temperature irrespective of the latitude. Therefore, in northern areas, the time between initiation of T. erectum leaf expansion and canopy closure was shorter than in southern areas, which left less time for northern plants to accumulate reserves before canopy closure. Differences in growth patterns were noted between T. erectum populations. From a south-north gradient, investment to reproduction, total plant biomass, and annual growth rate decreased, while specific leaf area and stem height increased, indicating shade acclimation. The length of the high light period in early spring seems to be a determinant for spring flowering plants' growth and reproduction and may explain the northern distribution limit of some of these species.     

Morphological and phenological shoot plasticity in a Mediterranean evergreen oak facing long-term increased drought

Mediterranean trees must adjust their canopy leaf area to the unpredictable timing and severity of summer drought. The impact of increased drought on the canopy dynamics of the evergreen Quercus ilex was studied by measuring shoot growth, leaf production, litterfall, leafing phenology and leaf demography in a mature forest stand submitted to partial throughfall exclusion for 7 years. The leaf area index rapidly declined in the throughfall-exclusion plot and was 19% lower than in the control plot after 7 years of treatment. Consequently, leaf litterfall was significantly lower in the dry treatment. Such a decline in leaf area occurred through a change in branch allometry with a decreased number of ramifications produced and a reduction of the leaf area supported per unit sapwood area of the shoot (LA/SA). The leafing phenology was slightly delayed and the median leaf life span was slightly longer in the dry treatment. The canopy dynamics in both treatments were driven by water availability with a 1-year lag: leaf shedding and production were reduced following dry years; in contrast, leaf turnover was increased following wet years. The drought-induced decrease in leaf area, resulting from both plasticity in shoot development and slower leaf turnover, appeared to be a hydraulic adjustment to limit canopy transpiration and maintain leaf-specific hydraulic conductivity under drier conditions.     

典型阔叶红松林主要树种叶性状的垂直变异及经济策略

分析不同树种叶片性状的变化有助于了解植物群落结构。该文通过对典型阔叶红松(Pinus koraiensis)林15种阔叶树种的比叶质量、叶片厚度、叶干物质含量、叶绿素含量指数、叶片碳、氮、磷含量的测定, 分析了冠层高度对叶性状及叶性状间相关关系的影响。结果表明, 水曲柳(Fraxinus mandshurica)和大青杨(Populus ussuriensis)上层的比叶质量显著大于下层, 而其他树种冠层间的比叶质量无显著变化; 叶绿素含量指数在白桦(Betula platyphylla)和春榆(Ulmus japonica)冠层间的分布分别为上层显著大于下层和上层显著大于中层; 单位质量氮含量在水曲柳的中层显著大于上层。叶片性状间存在着广泛的相关性, 比叶质量与叶片厚度、干物质含量在三层间均呈显著正相关关系, 而有些性状, 只在一或二个冠层中存在一定的相关性。山杨(Populus davidiana)和大青杨的叶片倾向于选择光合能力较低、营养浓度较低、呼吸速率较慢的一端, 而黄檗(Phellodendron amurense)和山槐(Maackia amurensis)叶片更倾向于光合能力强、营养物质浓度高的一端。不同树种对光照响应的差异可能会改变不同冠层中叶片的形态和化学性状, 从而有助于群落构建和物种共存。     

黑果枸杞功能性状对氮磷添加的响应及其可塑性

分析养分添加对荒漠植物功能性状的影响,对揭示其响应和适应环境变化的规律至关重要.本研究以黑果枸杞为材料,设置3个氮磷(NP)添加量(低、中、高)和N/P(5∶1、15∶1、45∶1),量化分析了整株、根、茎、叶和果实性状对NP添加的响应.结果 表明:黑果枸杞功能性状差异化响应了NP添加量和比例,随NP添加量的增加,生物...     

Vertical variation and economic strategy of leaf trait of major tree species in a typical mixed broadleaved-Korean pine forest

分析不同树种叶片性状的变化有助于了解植物群落结构。该文通过对典型阔叶红松(Pinus koraiensis)林15种阔叶树种的比叶质量、叶片厚度、叶干物质含量、叶绿素含量指数、叶片碳、氮、磷含量的测定, 分析了冠层高度对叶性状及叶性状间相关关系的影响。结果表明, 水曲柳(Fraxinus mandshurica)和大青杨(Populus ussuriensis)上层的比叶质量显著大于下层, 而其他树种冠层间的比叶质量无显著变化; 叶绿素含量指数在白桦(Betula platyphylla)和春榆(Ulmus japonica)冠层间的分布分别为上层显著大于下层和上层显著大于中层; 单位质量氮含量在水曲柳的中层显著大于上层。叶片性状间存在着广泛的相关性, 比叶质量与叶片厚度、干物质含量在三层间均呈显著正相关关系, 而有些性状, 只在一或二个冠层中存在一定的相关性。山杨(Populus davidiana)和大青杨的叶片倾向于选择光合能力较低、营养浓度较低、呼吸速率较慢的一端, 而黄檗(Phellodendron amurense)和山槐(Maackia amurensis)叶片更倾向于光合能力强、营养物质浓度高的一端。不同树种对光照响应的差异可能会改变不同冠层中叶片的形态和化学性状, 从而有助于群落构建和物种共存。     

Carbon limitation leads to suppression of first year oak seedlings beneath evergreen understory shrubs in Southern Appalachian hardwood forests

Inhibition of canopy tree recruitment beneath thickets of the evergreen shrubs Rhododendron maximum L. and Kalmia latifolia L. has long been observed in Southern Appalachian forests, yet the mechanisms of this process remain unresolved. We present a first-year account of suppression of oak seedlings in relation to Rhododendron and Kalmia basal area, light and resource availability, seedling performance and the rates of seedling damage (i.e., herbivory). We found no evidence of first-year seedling suppression or significant resource deficiencies beneath thickets of K. latifolia in mature mixed hardwood stands. Suppression beneath R. maximum was apparent during the first growing season. We found that seedling biomass, light availability prior to canopy closure, and seedling tissue C:N ratios were negatively correlated with R. maximum basal area. Basal area of R. maximum was positively correlated with seedling mortality rates, soil [Al], and early-growing season leaf herbivory rates. Seedling growth was positively correlated with light and tissue C:N, while negatively correlated with soil [Al]. Overall, our results support the inhibition model of shade-mediated carbon limitation beneath dense understory shrubs and indicate the potential importance of herbivory and aluminum toxicity as components of a suppression mechanism beneath R. maximum thickets. We present a causal model of first year inhibition beneath R. maximum in the context of our findings and the results of prior studies.     

Litter Decomposition Within the Canopy and Forest Floor of Three Tree Species in a Tropical Lowland Rain forest,Costa Rica

The rain forest canopy hosts a large percentage of the world's plant biodiversity, which is maintained, in large part, by internal nutrient cycling. This is the first study to examine the effects of site (canopy, forest floor) and tree species (Dipteryx panamensis, Lecythis ampla, Hyeronima alchorneoides) on decay rates of a common substrate and in situ leaf litter in a tropical forest in Costa Rica. Decay rates were slower for both substrates within the canopy than on the forest floor. The slower rate of mass loss of the common substrate in the canopy was due to differences in microclimate between sites. Canopy litter decay rates were negatively correlated with litter lignin:P ratios, while forest floor decay rates were negatively correlated with lignin concentrations, indicating that the control of litter decay rates in the canopy is P availability while that of the forest floor is carbon quality. The slower cycling rates within the canopy are consistent with lower foliar nutrient concentrations of epiphytes compared with forest floor-rooted plants. Litter decay rates, but not common substrate decay rates, varied among tree species. The lack of variation in common substrate decay among tree species eliminated microclimatic variation as a possible cause for differences in litter decay and points to variation in litter quality, nutrient availability and decomposer community of tree species as the causal factors. The host tree contribution to canopy nutrient cycling via litter quality and inputs may influence the quality and quantity of canopy soil resources.