Elevational Variation of Leaf Traits in Montane Rain Forest Tree Species at La Chinantla, Southern México1

Variation in leaf traits of dominant tree species in six montane rain forest communities was analyzed along an elevational gradient ranging from 1220 to 2560 m within a single basin at La Chinantla, Oaxaca, México. Three groups of characters were used: morphological (leaf shape, margin, blade configuration, and phyllotaxy), morphometric (leaf area, leaf mass per area, stomatal density, and blade length/width ratio), and anatomical (thicknesses of blade, palisade [PP], and spongy [SP] parenchymae, PP/SP ratio, and epidermis and cuticle thicknesses). The variation of morphological characteristics was only evident at the highest elevations; in contrast, thickness of leaf blade, PP, SP, as well as leaf mass per area clearly increased along the gradient, whereas leaf area was the only variable that significantly decreased with elevation. Thicknesses of epidermis and of the two cuticles were not significantly correlated with elevation. A classification analysis based on a leaf trait matrix led to the distinction between low and high elevation communities, with an approximate limit between them at ca 2300 to 2400 m. The results are discussed in light of environmental changes occurring along elevational gradients. Leaf characteristics of montane rain forest plants offer important insights about the complex roles of abiotic factors operating in these environments and supplement the traditional physiognomic classification schemes for these communities.     

Direct and indirect effects of environmental factors,spatial constraints,and functional traits on shaping the plant diversity of montane forests

Understanding the relative importance of the factors driving the patterns of biodiversity is a key research topic in community ecology and biogeography. However, the main drivers of plant species diversity in montane forests are still not clear. In addition, most existing studies make no distinction between direct and indirect effects of environmental factors and spatial constraints on plant biodiversity. Using data from 107 montane forest plots in Sichuan Giant Panda habitat, China, we quantified the direct and indirect effects of abiotic environmental factors, spatial constraints, and plant functional traits on plant community diversity. Our results showed significant correlations between abiotic environmental factors and trees (r = .10, p value = .001), shrubs (r = .19, p value = .001), or overall plant diversity (r = .18, p value = .001) in montane forests. Spatial constraints also showed significant correlations with trees and shrubs. However, no significant correlations were found between functional traits and plant community diversity. Moreover, the diversity (richness and abundance) of shrubs, trees, and plant communities was directly affected by precipitation, latitude, and altitude. Mean annual temperature (MAT) had no direct effect on the richness of tree and plant communities. Further, MAT and precipitation indirectly affected plant communities via the tree canopy. The results revealed a stronger direct effect on montane plant diversity than indirect effect, suggesting that single‐species models may be adequate for forecasting the impacts of climate factors in these communities. The shifting of tree canopy coverage might be a potential indicator for trends of plant diversity under climate change.     

Tree Diversity,Forest Structure and Productivity along Altitudinal and Topographical Gradients in a Species-Rich Ecuadorian Montane Rain Forest

We studied the spatial heterogeneity of tree diversity, and of forest structure and productivity in a highly diverse tropical mountain area in southern Ecuador with the aim of understanding the causes of the large variation in these parameters. Two major environmental gradients, elevation and topography, representing a broad range of climatic and edaphic site conditions, were analyzed. We found the highest species richness of trees in valleys <2100 m. Valleys showed highest values of basal area, leaf area index and tree basal area increment as well. Tree diversity also increased from ridges to valleys, while canopy openness decreased. Significant relationships existed between tree diversity and soil parameters (pH, total contents of Mg, K, Ca, N and P), and between diversity and the spatial variability of pH and Ca and Mg contents suggesting a dependence of tree diversity on both absolute levels and on the small-scale heterogeneity of soil nutrient availability. Tree diversity and basal area increment were positively correlated, partly because both are similarly affected by soil conditions. We conclude that the extraordinarily high tree species richness in the area is primarily caused by three factors: (1) the existence of steep altitudinal and topographic gradients in a rather limited area creating a small-scale mosaic of edaphically different habitats; (2) the intermingling of Amazonian lowland plant species, that reach their upper distribution limits, and of montane forest species; and (3) the geographical position of the study area between the humid eastern Andean slope and the dry interandean forests of South Ecuador.     

温带针阔混交林叶片性状随树冠垂直高度的变化规律

自然界中,森林植物叶片的生长随树冠高度呈现明显的垂直分布现象;然而,有关叶片性状随着树冠垂直高度增加的变化规律仍不清楚。为了更好地揭示植物叶片对光环境变化的适应策略以及对资源的利用能力,有必要深入探讨叶片性状与冠层高度的定量关系及其内在调控机制。以中国广泛分布的温带针阔混交林为对象,选取8种主要树种为研究对象(白桦、蒙古栎、水曲柳、大青杨、色木槭、千金榆、核桃楸和红松),通过测定这些物种9个冠层高度的叶片比叶面积(SLA)、叶片干物质含量(LDMC)、叶片氮含量(N)、叶片磷含量(P)、氮磷比(N∶P)和叶绿素含量(Chl)等属性,探讨了针阔混交林叶片性状的差异以及各性状之间的相关关系,进而揭示叶片性状随树冠垂直高度的变化规律。实验结果表明:1)温带针阔混交林内优势树种的部分叶片性状在不同冠层高度之间差异显著。2)随着树冠垂直高度的增加,SLA、LDMC、N、P、N∶P和Chl呈现不同的变化趋势。其中,阔叶树种SLA随着树冠垂直高度的增加而减小;所有树种的LDMC随着树冠垂直高度的增加而增加;不同树种的N、P、N∶P和Chl随着树冠垂直高度的变化规律存在差异。3)对于温带针阔混交林冠层中,SLA与N、P、N∶P均存在显著的正相关关系,高SLA伴随着高的N、P、N∶P,表明植物通过SLA与N、P等性状的协同来提高叶片的光合作用(或对光热资源的利用效率)。本研究通过定量分析探讨温带针阔混交林叶片性状随冠层高度的变化规律,一定程度地揭示了树木对光、热和水资源竞争的适应机制,以及植物叶片的资源利用和分配策略,不仅拓展了传统性状研究的范畴,其相关研究结论也有助于树木生长模型的构建和优化。     

Altitudinal change in soil and foliar nutrient concentrations and in microclimate across the tree line on the subtropical island mountain Mt. Teide (Canary Islands)

Mt. Teide (Tenerife, Spain) is a high volcanic island mountain with an unusually low tree line elevation (2000–2100 m). While searching for the causes of this tree line depression, we analysed the concentrations of total N, available P, and salt-exchangeable Ca, K and Mg in three soil horizons, and the foliar N, P, Ca, K and Mg concentrations in six abundant plant species (trees, shrubs, forbs) along a transect from 1400 (1600) to 3100 m a.s.l. The objective of the study was to detect altitudinal trends in soil and plant nutrient status below and above the tree line. For characterising elevational changes in microclimate and hydrology, we also conducted measurements of air and soil temperatures (T_a and T_s), atmospheric water vapour saturation deficit (D), potential evaporation (E) and soil moisture (θ) on 3–4 occasions along the transect.T_a and T_s did not linearly decrease with elevation but were highest at or immediately below the tree line and were relatively low in the closed Pinus canariensis forest at 1600 m. T_s reached maxima at about 70 °C near the tree line. The absence of a linear temperature decrease with elevation was caused by a stable temperature inversion at about 2000 m and by canopy shading in the forest canopy below the tree line. In June, the topsoil (10 cm) dried out almost completely between 1800 and 3100 m, but remained moist in the pine forest at 1600 m. This gradient reflects the transition from the montane cloud belt to the dry alpine belt higher upslope. The subsoil (30 cm) contained >30 vol% of soil water at all elevations even in June except for the uppermost site (3100 m). Potential evaporation increased with elevation despite a decrease in D. We assume that this is mainly due to the air pressure-dependent increase with elevation in the diffusion coefficient for water vapour in air.The concentrations of N and ‘available’ P (after Olsen) in the mineral topsoil were by far smaller than in alpine soils of other humid mountains which is thought to be a consequence of a very dry and biologically inactive topsoil in the semi-arid alpine belt of Mt. Teide. In contrast to many other mountains, foliar N, P and cation concentrations in the plants did not increase with elevation but either remained unchanged, or decreased as in the tree line species P. canariensis. Nevertheless, P. canariensis probably is not limited by nutrient deficiency at the tree line despite rather low N and P needle concentrations. Rather, drought and heat stress effects on seedling establishment are thought to be the causes of the tree line depression.     

Individual Canopy‐tree Species Effects on Their Immediate Understory Microsite and Sapling Community Dynamics

Canopy trees are largely responsible for the environmental heterogeneity in the understory of tropical and subtropical species‐rich forests, which in turn may influence sapling community dynamics. We tested the effect of the specific identity of four cloud forest canopy trees on total solar radiation, canopy openness, soil moisture, litter depth, and soil temperature, as well as on the structure and dynamics of the sapling community growing beneath their canopies. We observed significant effects of the specific identity of canopy trees on most understory microenvironmental variables. Soil moisture was higher and canopy openness lower beneath Cornus disciflora. In turn, canopy openness and total solar radiation were higher beneath Oreopanax xalapensis, while the lowest soil moisture occurred beneath Quercus laurina. Moreover, Chiranthodendron pentadactylon was the only species having a positive effect on litter depth under its canopy. In spite of these between‐species environmental differences, only C. pentadactylon had significant, negative effects on sapling density and species richness, which may be associated to low seed germination and seedling establishment due to an increased litter depth in its vicinity. The relevance of the specific identity of canopy trees for natural regeneration processes and species richness maintenance depends on its potential to differentially affect sapling dynamics through species‐specific modifications of microenvironmental conditions.     

树冠位置对香樟叶形态性状和异速生长关系的影响

为深入了解树冠位置对植物叶形态性状的影响,在常绿乔木香樟树冠上下2层和东南西北4个方位开展调查取样,系统分析了不同树冠位置间叶形态性状(叶长、叶宽、叶厚、叶柄长、叶柄直径和叶形指数)及其异速生长关系的差异性。结果表明,叶形态性状在不同树冠方位间均差异显著,但上下2层变化趋势不完全一致。在树冠上层,除叶形指数和叶炳长外,其余4个性状均表现为东侧最大。在树冠下层,除叶形指数外,其余5个性状指标均表现为东侧最小。在同一方位上,叶形态性状在上下2层间也存在一定差异,其中叶形指数多为下层高于上层,而其他形态性状多呈相反趋势。此外,树冠层次和方位的交互作用对叶片长、叶片厚、叶柄长和叶柄直径有显著影响。各层次和各方位叶形态性状间多为异速生长关系(即异速生长指数不等于1),且多无显著差异。在所有树冠层次和树冠方位,叶宽与叶厚、叶宽与叶炳长、叶长与叶厚及叶长与叶柄长之间均呈异速生长关系。可见,树冠位置对香樟叶形态性状的影响较大,但形态性状间的异速生长关系相对稳定,这是香樟叶形态性状表型可塑性和内在关系稳定性的重要体现。     

Does irrigation affect leaf phenology in deciduous and evergreen trees of the savannas of northern Australia?

Abstract Soil moisture was augmented experimentally during two successive dry seasons and the intervening wet season in a humid tropical savanna in Darwin, northern Australia. Leaf phenology was monitored in four common tree species Termmalia ferdinandiana and Planchonia careya (both deciduous species), and Eucalyptus miniata and Eucalyptus tetrodonta (both evergreen species). Irrigation produced consistently significant effects in only T. ferdinandiana. In this species leaf-flush was significantly earlier, canopy decline and leaf-fall were significantly later and the attainment of full canopy was earlier in irrigated compared with non-irrigated trees. Litterfall, and the seasonal patterns of contraction and expansion of stems (a measure of stem water status or storage) were not significantly affected by irrigation in any species. Leaf longevity in the deciduous species was 6–8 months; some eucalypt leaves lived for approximately 1 year, but none lived longer than 18 months. Irrigation had relatively little effect on leaf longevity. While variation in soil moisture is a potentially important cue to both leaf-fall and leaf-flush, stem water status and climatic factors such as vapour pressure deficit may also be important climatic cues to phenological behaviour.     

Climate warming delays and decreases seedling emergence in a Mediterranean ecosystem

Temperature and moisture impact strongly on the early stages of a plant's life cycle. Global climate change is altering the environmental cues that seeds receive resulting in compromised seedling emergence and changes to seedling performance. Here, we investigate how temperature and moisture affect these early stages of plant development in four Banksia species collected from a longitudinal climate gradient in southwest Western Australia. A common garden was used to examine the between‐species and among‐population variation in seedling emergence, growth and leaf traits under two soil temperature regimes and three levels of precipitation. We predicted that reduced moisture and increased temperature would delay and reduce total seedling emergence and negatively affect seedling performance. Furthermore, we expected that within species there would be geographically structured variation in response to the treatments. Species differed significantly in all measured traits. Soil warming resulted in strong impacts on regenerative traits, significantly slowing seedling emergence in two species and reducing total seedling emergence in three species. In addition, warming altered seedling performance with significant reductions to the above‐ground leaf biomass ratio of three species. In contrast, response to soil moisture manipulation was minimal across all species but possibly due to issues regarding implementation of an effective moisture treatment. The species that showed the greatest decline in emergence under warmed conditions (B. quercifolia) also showed the smallest vegetative shift; the species with the smallest decline in emergence (B. coccinea) showed a relatively large vegetative shift. Among‐population differences were significant for many traits, however, trait differentiation was inconsistent across species and, contrary to our hypothesis, the variation we observed was not clearly associated with the climate gradient. As these among‐population differences in traits are not easy to predict, we caution the use of simple rules for choosing seed populations for conservation and restoration.     

Correlation between water relations and within-canopy distribution of epiphytic ferns in a Mexican cloud forest

The physiological traits associated with water relations of eight common epiphytic ferns in a Mexican cloud forest were investigated in relation to the distribution of these species within the canopy. Fern distribution was significantly correlated with the relative water content at which stomata close, leaf thickness, stomatal density and size. Trichomanes bucinatum desiccated completely within hours in moderately dry air and was confined to the stem bases, and Asplenium cuspidatum, with no evident adaptations to cope with drought, grew in the second most shaded zone within the tree crowns. Despite growing in a humid cloud forest, all other species had xeric adaptations including coriaceous leaves (Pleopeltis mexicana, Elaphoglossum glaucum), succulent rhizomes (Polypodium puberulum, Phlebodium areolatum), low rates of uncontrolled water loss (all species except P. puberulum), leaf scales (Elaphoglossum petiolatum, Polypodium plebeium), and high cell wall elasticity (all species). P. plebeium and Pl. mexicanum, which grow in the most exposed locations, tolerated water loss beyond the turgor loss point before the stomata closed and appear to be poikilohydric or at least to tolerate high water deficits. Received: 8 June 1997 / Accepted: 30 November 1997     

陕北水蚀风蚀交错带柠条适应不同生境的形态和生理可塑性

以陕北水蚀风蚀交错带4个不同坡位与土壤质地生境[沟底+坝淤绵沙土(A)、梁坡+红黄土(B)、坡顶+绵沙土(C)、坡顶+风沙土盖绵沙土(D)]中生长的柠条为研究对象,研究了其生长、光合和水力性状的可塑性变化,以揭示柠条对不同生境的适应机制。结果表明:(1)生境D和生境A样地1~3m和3m土层的平均含水量明显高于生境B和C;(2)与生境B和C相比,生境A和D中生长的柠条冠幅、株高和新枝长显著增加,叶厚度减小,正午叶水势、净光合速率、气孔导度和蒸腾速率亦显著增加,但不同生境中柠条枝比导水率、Huber值和栓塞程度无显著差异;(3)柠条各测定指标中,新枝长、净光合速率、气孔导度和蒸腾速率表现出较大的可塑性。研究认为,柠条可能主要通过新枝长或光合生理特征的改变来适应不同的水分生境。     

Ecophysiology in Relation to Exposure of Pendant Epiphytic Bryophytes in the Canopy of a Tropical Montane Oak Forest1

We investigated ecophysiological and morphological traits affecting light and water use of four commercially important pendant epiphytic bryophyte species in a montane oak‐bamboo forest in Costa Rica. Predictions based mostly on ecophysiological studies of temperate bryophytes and vascular plants were experimentally tested on tropical montane bryophyte species ranked on the basis of their distributions in the canopy from the most protected (1) to the most exposed sites (4): (1) Phyllogonium viscosum; (2) Pilotrichella flexilis; (3) Dendropogonella rufescens; and (4) Frullania convoluta. As predicted, highly exposed species tended to have higher light saturation and compensation points, higher dark respiration rates, more chlorophyll, higher chlorophyll a:b ratios, and higher nitrogen concentrations. Contrary to predicted trends, exposed‐site species had lower water contents at full saturation; there were no detectable differences among species in the rate of water loss. Rates of carbon gain in all species reached asymptotes with increasing water content, but the moisture compensation point for carbon uptake of the moss D. rufescens was substantially higher than in the other species. Observed patterns result from interactions among processes related to water storage and transport operating at different scales and are determined by various morphological traits including the density, size, and disposition of phylloids, as well as by whole‐clump architecture.     

Trait-based community assembly of understory palms along a soil nutrient gradient in a lower montane tropical forest

Two opposing niche processes have been shown to shape the relationship between ecological traits and species distribution patterns: habitat filtering and competitive exclusion. Habitat filtering is expected to select for similar traits among coexisting species that share similar habitat conditions, whereas competitive exclusion is expected to limit the ecological similarity of coexisting species leading to trait differentiation. Here, we explore how functional traits vary among 19 understory palm species that differ in their distribution across a gradient of soil resource availability in lower montane forest in western Panama. We found evidence that habitat filtering influences species distribution patterns and shifts community-wide and intraspecific trait values. Differences in trait values among sites were more strongly related to soil nutrient availability than to variation in light or rainfall. Soil nutrient availability explained a significant amount of variation in site mean trait values for 4 of 15 functional traits. Site mean values of leaf nitrogen and phosphorus increased 37 and 64%, respectively, leaf carbon:nitrogen decreased 38%, and specific leaf area increased 29% with increasing soil nutrient availability. For Geonoma cuneata, the only species occurring at all sites, leaf phosphorus increased 34% and nitrogen:phosphorus decreased 42% with increasing soil nutrients. In addition to among-site variation, most morphological and leaf nutrient traits differed among coexisting species within sites, suggesting these traits may be important for niche differentiation. Hence, a combination of habitat filtering due to turnover in species composition and intraspecific variation along a soil nutrient gradient and site-specific niche differentiation among co-occurring species influences understory palm community structure in this lower montane forest.     

Foliar C:N Ratio of Ferns along an Andean Elevational Gradient1

We studied the concentration of leaf N and C among 183 fern species along an elevational gradient at 1700 to 3400 m in humid montane forest in the Bolivian Andes at different levels of taxonomic resolution. For two species of Elaphoglossum sampled 8 and 14 times, respectively, there were no elevational trends. Similarly, a contrast of 22 species with wide elevational amplitudes sampled at their highest and lowest locations did not show any change in C or N contents, or in C:N ratios with elevation. At the community level, however, the mean values of C:N ratios for (a) all species found at a given elevation showed a significant decline with increasing elevation and (b) among epiphytic species, higher ratios (i.e., lower relative N content) than among terrestrial species at the same elevation. These trends were opposite to those of the upper soil layer, in which C:N ratios increased with elevation.     

Forest aboveground biomass along an elevational transect in Sulawesi,Indonesia, and the role of Fagaceae in tropical montane rain forests

Aim This study investigates how estimated tree aboveground biomass (AGB) of tropical montane rain forests varies with elevation, and how this variation is related to elevational change in floristic composition, phylogenetic community structure and the biogeography of the dominant tree taxa. Location Lore Lindu National Park, Sulawesi, Indonesia. Methods Floristic inventories and stand structural analyses were conducted on 13 plots (each 0.24 ha) in four old‐growth forest stands at 1050, 1400, 1800 and 2400 m a.s.l. (submontane to upper montane elevations). Tree AGB estimates were based on d.b.h., height and wood specific gravity. Phylogenetic diversity and biogeographical patterns were analysed based on tree family composition weighted by AGB. Elevational trends in AGB were compared with other Southeast Asian and Neotropical transect studies (n = 7). Results AGB was invariant from sub‐ to mid‐montane elevation (309–301 Mg ha^?1) and increased slightly to 323 Mg ha^?1 at upper montane elevation. While tree and canopy height decreased, wood specific gravity increased. Magnoliids accounted for most of the AGB at submontane elevations, while eurosids I (including Fagaceae) contributed substantially to AGB at all elevations. Phylogenetic diversity was highest at upper montane elevations, with co‐dominance of tree ferns, Podocarpaceae, Trimeniaceae and asterids/euasterids II, and was lowest at lower/mid‐montane elevations, where Fagaceae contributed > 50% of AGB. Biogeographical patterns showed a progression from dominant tropical families at submontane to tropical Fagaceae (Castanopsis, Lithocarpus) at lower/mid‐montane, and to conifers and Australasian endemics at upper montane elevations. Cross‐continental comparisons revealed an elevational AGB decrease in transects with low/no presence of Fagaceae, but relatively high AGB in montane forests with moderate to high abundance of this family. Main conclusions AGB is determined by both changes in forest structure and shifts in species composition. In our study, these two factors traded off so that there was no net change in AGB, even though there were large changes in forest structure and composition along the elevational gradient. Southeast Asian montane rain forests dominated by Fagaceae constitute important carbon stocks. The importance of biogeography and species traits for biomass estimation should be considered by initiatives to reduce emissions from deforestation and forest degradation (REDD) and in taxon choice in reforestation for carbon offsetting.     

Leaf litter and irrigation can increase seed germination and early seedling survival of the recalcitrant‐seeded tree Beilschmiedia miersii

Although regeneration of recalcitrant‐seeded tree species can be affected by prolonged drought, especially in Mediterranean regions, little is known about the response of such species to varying site conditions. A field experiment was performed to determine the effect of irrigation and leaf litter cover on seed germination and early seedling survival of the vulnerable recalcitrant‐seeded tree Beilschmiedia miersii (Lauraceae). Two levels of irrigation (non‐irrigated and irrigated units) and three levels of leaf litter depth (0, 5 and 12 cm) were applied to 72 groups of 30 seeds along a wet ravine of the Coastal Range of Central Chile, equally distributed across sites with different levels of canopy cover. Seed germination was significantly increased by irrigation only under closed‐canopy cover, and by leaf litter cover (>5 cm) under all canopy conditions. The effect of leaf litter on germination increased with canopy openness, while the effect of irrigation did not show any tendency. Meanwhile, early seedling survival was significantly increased by irrigation under intermediate canopy cover, and by leaf litter (>5 cm) under closed‐canopy cover. As a result of its overall positive effect on germination, leaf litter should be maintained within B. miersii communities, particularly under intermediate to closed‐canopy conditions, where it can also increase early seedling survival, and both seed germination and early seedling survival might be increased through additional water inputs. The presence of leaf litter might help retaining such inputs, prolonging their effect on regeneration of B. miersii communities. We see this as a baseline assessment of regeneration and persistence that needs further testing on species with similar traits, given the expected increase in the frequency and length of dry periods into the future.     

Leaf litter inputs reinforce islands of nitrogen fertility in a lowland tropical forest

The role of lowland tropical forest tree communities in shaping soil nutrient cycling has been challenging to elucidate in the face of high species diversity. Previously, we showed that differences in tree species composition and canopy foliar nitrogen (N) concentrations correlated with differences in soil N availability in a mature Costa Rican rainforest. Here, we investigate potential mechanisms explaining this correlation. We used imaging spectroscopy to identify study plots containing 10–20 canopy trees with either high or low mean canopy N relative to the landscape mean. Plots were restricted to an uplifted terrace with relatively uniform parent material and climate. In order to assess whether canopy and soil N could be linked by litterfall inputs, we tracked litter production in the plots and measured rates of litter decay and the carbon and N content of leaf litter and leaf litter leachate. We also compared the abundance of putative N fixing trees and rates of free-living N fixation as well as soil pH, texture, cation exchange capacity, and topographic curvature to assess whether biological N fixation and/or soil properties could account for differences in soil N that were, in turn, imprinted on the canopy. We found no evidence of differences in legume communities, free-living N fixation, or abiotic properties. However, soils beneath high canopy N assemblages received ~ 60% more N via leaf litterfall due to variability in litter N content between plot types. The correlation of N in canopy leaves, leaf litter, and soil suggests that, under similar abiotic conditions, litterfall-mediated feedbacks can help maintain soil N differences among tropical tree assemblages in this diverse tropical forest.

     

Altitudinal Change in LAI and Stand Leaf Biomass in Tropical Montane Forests: a Transect Study in Ecuador and a Pan-Tropical Meta-Analysis

Abstract Leaf area index (LAI) is a key parameter controlling plant productivity and biogeochemical fluxes between vegetation and the atmosphere. Tropical forests are thought to have comparably high LAIs; however, precise data are scarce and environmental controls of leaf area in tropical forests are not understood. We studied LAI and stand leaf biomass by optical and leaf mass-related approaches in five tropical montane forests along an elevational transect (1,050–3,060 m a.s.l.) in South Ecuador, and conducted a meta-analysis of LAI and leaf biomass data from tropical montane forests around the globe. Study aims were (1) to assess the applicability of indirect and direct approaches of LAI determination in tropical montane forests, (2) to analyze elevation effects on leaf area, leaf mass, SLA, and leaf lifespan, and (3) to assess the possible consequences of leaf area change with elevation for montane forest productivity. Indirect optical methods of LAI determination appeared to be less reliable in the complex canopies than direct leaf mass-related approaches based on litter trapping and a thorough analysis of leaf lifespan. LAI decreased by 40–60% between 1,000 and 3,000 m in the Ecuador transect and also in the pan-tropical data set. This decrease indicates that canopy carbon gain, that is, carbon source strength, decreases with elevation in tropical montane forests. Average SLA decreased from 88 to 61 cm² g⁻¹ whereas leaf lifespan increased from 16 to 25 mo between 1,050 and 3,060 m in the Ecuador transect. In contrast, stand leaf biomass was much less influenced by elevation. We conclude that elevation has a large influence not only on the leaf traits of trees but also on the LAI of tropical montane forests with soil N (nitrogen) supply presumably being the main controlling factor.     

Global patterns of intraspecific leaf trait responses to elevation

Elevational gradients are often used to quantify how traits of plant species respond to abiotic and biotic environmental variations. Yet, such analyses are frequently restricted spatially and applied along single slopes or mountain ranges. Since we know little on the response of intraspecific leaf traits to elevation across the globe, we here perform a global meta‐analysis of leaf traits in 109 plant species located in 4 continents and reported in 71 studies published between 1983 and 2018. We quantified the intraspecific change in seven morpho‐ecophysiological leaf traits along global elevational gradients: specific leaf area (SLA), leaf mass per area (LMA), leaf area (LA), nitrogen concentration per unit of area (Narea), nitrogen concentration per unit mass (Nmass), phosphorous concentration per unit mass (Pmass) and carbon isotope composition (δ¹³C). We found LMA, Narea, Nmass and δ¹³C to significantly increase and SLA to decrease with increasing elevation. Conversely, LA and Pmass showed no significant pattern with elevation worldwide. We found significantly larger increase in Narea, Nmass, Pmass and δ¹³C with elevation in warmer regions. Larger responses to increasing elevation were apparent for SLA of herbaceous compared to woody species, but not for the other traits. Finally, we also detected evidences of covariation across morphological and physiological traits within the same elevational gradient. In sum, we demonstrate that there are common cross‐species patterns of intraspecific leaf trait variation across elevational gradients worldwide. Irrespective of whether such variation is genetically determined via local adaptation or attributed to phenotypic plasticity, the leaf trait patterns quantified here suggest that plant species are adapted to live on a range of temperature conditions. Since the distribution of mountain biota is predominantly shifting upslope in response to changes in environmental conditions, our results are important to further our understanding of how plants species of mountain ecosystems adapt to global environmental change.     

浙江天童太白山不同群落植物构型比较

植物构型是植株构件在空间上的分配方式,反映了植物对环境的响应策略。通过对浙江天童太白山海拔差异很小的栲树(Castanopsis fargesii)、小叶青冈(Cyclobalanopsis gracilis)和云山青冈(Cyclobalanopsis sessilifolia)群落类型中所有植株的树高、树冠厚度、树冠面积、叶片聚集度、枝下高和距地45 cm基径等植物构型性状,以及树冠曝光指数、土壤含水率、空气温湿度、土壤p H值和风速6个环境因子的测定,分别分析乔灌木层植物构型性状及性状关系在3个群落间的变化规律。结果表明:(1)从栲树到小叶青冈至云山青冈群落,灌木层的树高、树冠厚度、枝下高和距地45 cm基径增加,叶片聚集度减小;乔木层的树高、树冠厚度、树冠面积、枝下高和距地45 cm基径均减小,叶片聚集度增大;(2)3个群落灌木层构型性状间显著相关(P0.001),而乔木层只在中低海拔群落存在相关性;(3)从栲树到小叶青冈至云山青冈群落,乔灌木层的冠层曝光指数显著增加(P0.05);(4)多元逐步回归表明,树冠曝光指数对灌木层构型性状变异的贡献最大,而风速、土壤含水率和p H值对乔木构型性状的变异起主导作用。综上得知,天童太白山乔灌木植物在不同群落间存在构型分异,植物对光资源的竞争是引起灌木构型在不同群落间变化的主要驱动因子,而对乔木植物,其构型变化更多受到风速和土壤含水量的影响。