Utility of information in photographs taken upwards from the floor of closed-canopy deciduous broadleaved and closed-canopy evergreen coniferous forests for continuous observation of canopy phenology

Hemispherical photographs taken on forest floors are used to monitor seasonal changes in canopy openness or leaf area index in ecological studies. Those analyses usually use black and white images converted from the original colour images. Photographs taken by downwards-facing cameras installed on towers are used to provide detailed information on leaf expansion, maturation and senescence of various tree species through the analysis of red, green and blue ‘digital numbers’ (DN_RGB) extracted from those images. To examine the usefulness of colour information encoded in upwards hemispherical photographs in monitoring canopy phenological characteristics, we examined the consistency of DN_RGB values between downwards and upwards images in deciduous broadleaved and evergreen coniferous forests in Japan. In the deciduous broadleaved forest, the DN_RGB values in the upwards images were able to detect canopy phenology almost as well as those in the downwards images. However, we found the effects on DN_RGB of (1) the spatial heterogeneity among observed points, (2) low-vegetation (before the beginning of leaf-expansion and after the end of leaf-fall period) and (3) white balance settings. In the evergreen coniferous forest, in contrast, the DN_RGB values in the upwards images did not capture canopy phenology. These different results may be attributable to the light attenuation characteristics in the canopies due to the geometries of leaves and branches. Thus, the DN_RGB values obtained from upwards images are almost as good as those of downwards images for monitoring detailed canopy phenology in deciduous broadleaved forests with a closed canopy.     

Spectral vegetation indices as the indicator of canopy photosynthetic productivity in a deciduous broadleaf forest

Aims Understanding of the ecophysiological dynamics of forest canopy photosynthesis and its spatial and temporal scaling is crucial for revealing ecological response to climate change. Combined observations and analyses of plant ecophysiology and optical remote sensing would enable us to achieve these studies. In order to examine the utility of spectral vegetation indices (VIs) for assessing ecosystem-level photosynthesis, we investigated the relationships between canopy-scale photosynthetic productivity and canopy spectral reflectance over seasons for 5 years in a cool, temperate deciduous broadleaf forest at 'Takayama' super site in central Japan.Methods Daily photosynthetic capacity was assessed by in situ canopy leaf area index (LAI), (LAI × V cmax [single-leaf photosynthetic capacity]), and the daily maximum rate of gross primary production (GPP max) was estimated by an ecosystem carbon cycle model. We examined five VIs: normalized difference vegetation index (NDVI), enhanced vegetation index (EVI), green–red vegetation index (GRVI), chlorophyll index (CI) and canopy chlorophyll index (CCI), which were obtained by the in situ measurements of canopy spectral reflectance.Important findings Our in situ observation of leaf and canopy characteristics, which were analyzed by an ecosystem carbon cycling model, revealed that their phenological changes are responsible for seasonal and interannual variations in canopy photosynthesis. Significant correlations were found between the five VIs and canopy photosynthetic capacity over the seasons and years; four of the VIs showed hysteresis-type relationships and only CCI showed rather linear relationship. Among the VIs examined, we applied EVI–GPP max relationship to EVI data obtained by Moderate Resolution Imaging Spectroradiometer to estimate the temporal and spatial variation in GPP max over central Japan. Our findings would improve the accuracy of satellite-based estimate of forest photosynthetic productivity in fine spatial and temporal resolutions, which are necessary for detecting any response of terrestrial ecosystem to meteorological fluctuations.     

干旱胁迫下雷竹叶片叶绿素的高光谱响应特征及含量估算

植物叶片的反射光谱特征与叶绿素含量密切相关。以重要的笋用竹种雷竹(Phyllostachys violascens)为研究对象,采用盆栽及控水试验方法研究了2年生雷竹在干旱胁迫条件下冠层叶片反射光谱的响应特征,分析了叶片叶绿素含量与不同波段光谱反射率一阶微分值以及光谱特征参数之间的相关关系,并以雷竹叶绿素含量敏感波段及构建的植被指数与叶绿素含量进行了拟合。结果表明,重度缺水处理后雷竹叶片叶绿素含量显著降低,在可见光区叶片光谱反射率随叶绿素含量的降低而增加,以波长493、639、693、756 nm等处的光谱反射率一阶微分值与叶绿素含量的相关性较高。雷竹叶片叶绿素含量与光谱特征参数如绿峰反射率、红谷反射率、蓝边面积、绿峰面积之间的相关性较高。与已有的植被指数相比基于雷竹叶绿素含量敏感波段修正后的植被指数与叶绿素含量相关性优于原植被指数。基于反射率一阶微分值构建的多元回归方程以及修正的绿色归一化植被指数(m GNDVI)构建的回归方程拟合效果较好,为雷竹叶绿素含量的较优估算方程。研究结果可以为雷竹叶绿素含量的快速无损测定以及季节性干旱条件下雷竹林的科学经营及灾后评估提供依据。     

Vertical gradients in photosynthetic physiology diverge at the latitudinal range extremes of white spruce

Light availability drives vertical canopy gradients in photosynthetic functioning and carbon (C) balance, yet patterns of variability in these gradients remain unclear. We measured light availability, photosynthetic CO₂ and light response curves, foliar C, nitrogen (N) and pigment concentrations, and the photochemical reflectance index (PRI) on upper and lower canopy needles of white spruce trees (Picea glauca) at the species' northern and southern range extremes. We combined our photosynthetic data with previously published respiratory data to compare and contrast canopy C balance between latitudinal extremes. We found steep canopy gradients in irradiance, photosynthesis and leaf traits at the southern range limit, but a lack of variation across canopy positions at the northern range limit. Thus, unlike many tree species from tropical to mid-latitude forests, high latitude trees may not require vertical gradients of metabolic activity to optimize photosynthetic C gain. Consequently, accounting for self-shading is less critical for predicting gross primary productivity at northern relative to southern latitudes. Northern trees also had a significantly smaller net positive leaf C balance than southern trees suggesting that, regardless of canopy position, low photosynthetic rates coupled with high respiratory costs may ultimately constrain the northern range limit of this widely distributed boreal species.     

Threshold values of canopy reflectance indices and chlorophyll meter readings for optimal nitrogen nutrition of tomato

Sustainable and optimal economic N management requires correct and timely on‐farm assessment of crop N status to detect N deficiency or excess. Optical sensors are promising tools to assess crop N status throughout a crop or at critical times. The ability of optical sensor measurements of canopy reflectance (Crop Circle ACS 470) and leaf chlorophyll (SPAD 502 chlorophyll meter) to assess crop N status was evaluated weekly throughout an indeterminate tomato crop. Strong linear relationships with the optical sensor measurements were obtained, throughout most of the crop, for both (a) crop N content for ranges of 1.5–4.5%, and (b) the nitrogen nutrition index (NNI) for ranges of 0.4–1.3. The relationships of the optical sensor measurements to crop NNI were generally equal to or slightly better than with crop N content. Indices based on reflectance in the red, the normalised difference vegetation index (NDVI) and the red vegetation index (RVI), were the best predictors of crop N status in terms of goodness of fit, earliness and maintenance of relationships throughout the crop. SPAD chlorophyll readings and reflectance indices based on reflectance in the green, the normalised difference vegetation index on greenness (GNDVI) and the green vegetation index (GVI), were good indicators of crop N status for most of the crop, but with lower goodness of fit in the latter part of the crop. The linear relationships between sensor indices and readings with NNI or crop N content, each week, demonstrated the potential for using proximal canopy reflectance indices such as NDVI and RVI, and chlorophyll meter for monitoring crop N status of indeterminate tomato crops. Threshold values for optimal crop N nutrition for canopy reflectance indices and for chlorophyll meter readings were derived for each day of measurement from the relationships between optical sensor measurements and NNI by solving for NNI = 1. The threshold values obtained for each index and type of measurement varied during the crop cycle. The approach developed for determining threshold values from NNI can facilitate on‐farm use of optical sensors for monitoring crop N status, by enabling assessment of whether crop N status is excessive, deficient or adequate.     

Diverse Optical and Photosynthetic Properties in a Neotropical Dry Forest during the Dry Season: Implications for Remote Estimation of Photosynthesis1

Using optical and photosynthetic assays from a canopy access crane, we examined the photosynthetic performance of tropical dry forest canopies during the dry season in Parque Metropolitano, Panama City, Panama. Photosynthetic gas exchange, chlorophyll fluorescence, and three indices derived from spectral reflectance (the normalized difference vegetation index, the simple ratio, and the photochemical reflectance index) were used as indicators of structural and physiological components of photosynthetic activity. Considerable interspecific variation was evident in structural and physiological behavior in this forest stand, which included varying degrees of foliage loss, altered leaf orientation, stomatal closure, and photosystem II downregulation. The normalized difference vegetation index and the simple ratio were closely related to canopy structure and absorbed radiation for most species, but failed to capture the widely divergent photosynthetic behavior among evergreen species exhibiting various degrees of downregulation. The photochemical reflectance index and chlorophyll fluorescence were related indicators of photosynthetic downregulation, which was not detectable with the normalized difference vegetation index or simple ratio. These results suggest that remote sensing methods that ignore downregulation cannot capture within‐stand variability in actual carbon flux for this diverse forest type. Instead, these findings support a sampling approach that derives photosynthetic fluxes from a consideration of both canopy light absorption (e.g., normalized difference vegetation index) and photosynthetic light‐use efficiency (e.g., photochemical reflectance index). Such sampling should improve our understanding of controls on photosynthetic carbon uptake in diverse tropical forest stands.     

Is there a temporal niche separation in the leaf phenology of savanna trees and grasses?

Aim It has been proposed that, in tropical savannas, trees deploy their leaves earlier in the growing season and grasses deploy their leaves later. This hypothesis implies a mechanism that facilitates the coexistence of trees and grasses in savannas. If true, this hypothesis would also allow algorithms to use differences in the phenological timing of grass and tree leaves to partition the relative contribution of grasses and trees to net primary production. In this study we examine whether a temporal niche separation between grasses and trees exists in savanna. Location A semi‐arid, subtropical savanna, Kruger National Park, South Africa. Methods We use a multi‐spectral camera to track through an entire growing season the normalized difference vegetation index (NDVI) of individual canopies of grasses and trees at eight sites arranged along a precipitation and temperature gradient. Results Among trees, we identified two distinct phenological syndromes: an early flushing syndrome and a late‐flushing syndrome. Leaf flush in the tree strategies appears to pre‐empt rainfall, whereas grass leaf flush follows the rain. The growing season of trees is 20 (late‐flushing trees) to 27 (early flushing trees) days longer than that of the grasses. Main conclusions We show that grasses and trees have different leaf deployment strategies. Trees deployed leaves at lower temperatures than grasses and retained them for longer at the end of the growing season. The timing of the increase in NDVI is, however, similar between grasses and late‐flushing trees and this complicates the separation of grass and tree signals from multi‐spectral satellite imagery.     

Increased leaf reflectance in tropical trees under elevated CO2

Globally increasing atmospheric CO₂ concentrations are known to affect many aspects of plant physiology and development; however, little attention has been given to leaf and canopy optical properties. Three tropical trees in the Leguminosae, an important canopy tree family in many tropical forests, responded similarly to an experimental doubling of CO₂ partial pressure with a 9–23% increase in spectral leaf reflectance to light in the visible (400–700 nm) waveband. Decreased leaf chlorophyll content under elevated CO₂ may explain part of the observed increase in reflectance. However, analyses that statistically corrected for chlorophyll content effects on reflectance still indicated a significant CO₂ effect. This results, in conjunction with the spectral pattern of the response, suggests that the primary mechanism is increased optical masking of chlorophyll under elevated CO₂. The magnitude of the increase in leaf reflectance is sufficient to suggest that increased canopy reflectance of tropical forests (and possibly other terrestrial ecosystems) may be an important negative feedback in the response of global net radiative climate forcing to increasing atmospheric CO₂.     

Evaluation of reflectance spectroscopy indices for estimation of chlorophyll content in leaves of a tropical tree species

The effectiveness of eight spectral reflectance indices for estimating chlorophyll (Chl) content in leaves of Eugenia uniflora L., a tropical tree species widely distributed throughout the world and a key species for ecosystem restoration projects, was evaluated. Spectral reflectance indices were tested using sun and shade leaves with a broad variation in leaf mass per area (LMA). Shortly after plants were exposed to chilling temperatures, there was a dramatic visible change in some sun leaves from green to red. Prior to testing Chl-related reflectance indices, the green and red leaves were separated according to the anthocyanin reflectance index (ARI). Slightly green to dark green leaves corresponded to an ARI value less than 0.11 (n = 107), whereas slightly red to red leaves corresponded to an ARI value greater than 0.11 (n = 35). To estimate leaf Chl, two simple reflectance indices (SR₆₈₀ and SR₇₀₅), two normalized difference indices (ND₆₈₀ and ND₇₀₅), two modified reflectance indices (mSR₇₀₅ and mND705), a modified Chl absorption ratio index (mCARI₇₀₅) and an index insensitive to the presence of anthocyanins (CI_re) were evaluated. Good estimates of leaf Chl content were obtained using the reflectance indices tested regardless of the presence of anthocyanins and changes in LMA. Based on the coefficients of determination (r ²) and the root mean square errors (RMS?_c) the best results were obtained with reflectance indices measured at wavelengths of 750 and 705 nm. Considering the performance of the models the best reflectance indices to estimate Chl contents in E. uniflora leaves with a broad variation in LMA and anthocyanin contents was SR₇₀₅ and mCARI₇₀₅.     

落叶阔叶林冠层光合有效辐射分量的遥感模拟与分析

冠层绿色叶片(光合组分)的光合有效辐射分量(绿色FPAR)真实地反映了植被与外界进行物质和能量交换的能力,获取冠层光合组分吸收的太阳光合有效辐射,对生态系统生产力的遥感估算精度的提高具有重要的意义。研究以落叶阔叶林为例,基于SAIL模型模拟森林冠层光合组分和非光合组分吸收的光合有效辐射,研究冠层FPAR变化规律以及与植被指数的相关关系。结果表明,冠层结构的改变会影响冠层对PAR的吸收能力,冠层绿色FPAR的大小与植被面积指数及光合组分面积比相关;在高覆盖度植被区,冠层绿色FPAR占冠层总FPAR的80%以上,非光合组分的贡献较小,但在低植被覆盖区,当光合组分和非光合组分面积相同时,绿色FPAR不及冠层总FPAR的50%;相比于NDVI,北方落叶阔叶林冠层EVI与绿色FPAR存在更为显著的线性相关关系(R~20.99)。     

Plant ecophysiological processes in spectral profiles: perspective from a deciduous broadleaf forest

The need for progress in satellite remote sensing of terrestrial ecosystems is intensifying under climate change. Further progress in Earth observations of photosynthetic activity and primary production from local to global scales is fundamental to the analysis of the current status and changes in the photosynthetic productivity of terrestrial ecosystems. In this paper, we review plant ecophysiological processes affecting optical properties of the forest canopy which can be measured with optical remote sensing by Earth-observation satellites. Spectral reflectance measured by optical remote sensing is utilized to estimate the temporal and spatial variations in the canopy structure and primary productivity. Optical information reflects the physical characteristics of the targeted vegetation; to use this information efficiently, mechanistic understanding of the basic consequences of plant ecophysiological and optical properties is essential over broad scales, from single leaf to canopy and landscape. In theory, canopy spectral reflectance is regulated by leaf optical properties (reflectance and transmittance spectra) and canopy structure (geometrical distributions of leaf area and angle). In a deciduous broadleaf forest, our measurements and modeling analysis of leaf-level characteristics showed that seasonal changes in chlorophyll content and mesophyll structure of deciduous tree species lead to a seasonal change in leaf optical properties. The canopy reflectance spectrum of the deciduous forest also changes with season. In particular, canopy reflectance in the green region showed a unique pattern in the early growing season: green reflectance increased rapidly after leaf emergence and decreased rapidly after canopy closure. Our model simulation showed that the seasonal change in the leaf optical properties and leaf area index caused this pattern. Based on this understanding we discuss how we can gain ecophysiological information from satellite images at the landscape level. Finally, we discuss the challenges and opportunities of ecophysiological remote sensing by satellites.

     

Differences in nitrogen use efficiency between leaves from canopy and subcanopy trees

We examined the effects of increasing light availability along a vertical gradient within a forest community on the efficiency of leaf nitrogen (N) use in individual trees. The N contents of green and senescent leaves in canopy and subcanopy trees of an evergreen coniferous species, Podocarpus nagi, and an evergreen hardwood species, Neolitsea aciculata, were analyzed in a mixed forest community at Mt Mikasa, Nara City, Japan. The inverse of N concentration (N_C) in senescent leaves was used as an index of N use efficiency (NUE) at the leaf-level. The leaf-level NUE was higher in canopy trees than in subcanopy trees in both P.nagi and N.aciculata, although soil N mineralization rates around canopy and subcanopy trees did not differ significantly. The N_C in green leaves was lower in canopy trees than in subcanopy trees. The ratio of resorbed N in senescent leaves to the N content in green leaves was higher in canopy trees than in subcanopy trees. The higher leaf-level NUE of canopy trees was partly a result of lower N_C in living tissues and partly because of greater N resorption during senescence. The present study suggested that the leaf-level NUE could be increased in response to an imbalance between soil N and light availability caused by spatial community structure.     

Development of a stable isotope index to assess decadal-scale vegetation change and application to woodlands of the Burdekin catchment, Australia

Forty-four study sites were established in remnant woodland in the Burdekin River catchment in tropical north-east Queensland, Australia, to assess recent (decadal) vegetation change. The aim of this study was further to evaluate whether wide-scale vegetation ‘thickening’ (proliferation of woody plants in formerly more open woodlands) had occurred during the last century, coinciding with significant changes in land management. Soil samples from several depth intervals were size separated into different soil organic carbon (SOC) fractions, which differed from one another by chemical composition and turnover times. Tropical (C₄) grasses dominate in the Burdekin catchment, and thus δ¹³C analyses of SOC fractions with different turnover times can be used to assess whether the relative proportion of trees (C₃) and grasses (C₄) had changed over time. However, a method was required to permit standardized assessment of the δ¹³C data for the individual sites within the 13 Mha catchment, which varied in soil and vegetation characteristics. Thus, an index was developed using data from three detailed study sites and global literature to standardize individual isotopic data from different soil depths and SOC fractions to reflect only the changed proportion of trees (C₃) to grasses (C₄) over decadal timescales. When applied to the 44 individual sites distributed throughout the Burdekin catchment, 64% of the sites were shown to have experienced decadal vegetation thickening, while 29% had remained stable and the remaining 7% had thinned. Thus, the development of this index enabled regional scale assessment and comparison of decadal vegetation patterns without having to rely on prior knowledge of vegetation changes or aerial photography.     

Spatial and seasonal heterogeneity in understory light conditions caused by differential leaf flushing of deciduous overstory trees

In a deciduous broad-leaved forest, we investigated the seasonality and heterogeneity of understory light conditions in relation to the leaf phenology of overstory trees. Fisheye photographs were taken from spring to autumn to estimate direct and indirect light conditions above the understory. Spatial variation in daily direct photosynthetic photon flux density (PPFD) was highest in late May, when the early and the intermediate-flushing trees had finished flushing and the late-flushing trees had just started flushing. After whole canopy closure, spatial variation in direct PPFD became low. Thus, asynchronous overstory leaf flushing in spring resulted in spatial heterogeneity in understory light conditions. These results suggest that the leaf phenological patterns of overstory trees are an important factor in the formation of the understory community.     

交通绿化带植物配置对空气颗粒物的净化效益

交通绿化带有显著的净化街道空气的环境效益。通过对上海浦东某交通干道旁侧绿化带不同季节大气中总悬浮颗粒物(TSP)的测定,定量研究了绿化带对TSP的净化效益;同时对研究区域内植物的配置情况用郁闭度和疏透度进行了表征,并研究了其与TSP的净化效益之间的相关性。研究结果表明:交通绿化带对由机动车引起的TSP污染有明显的净化作用,其中夏、秋季净化百分率较高,春、冬季较低;为达到较高的TSP净化效益,交通绿化带宽度应不小于5m,最佳为10m,采用先灌后乔的配置方式,并更多的选用常绿树种;绿地对TSP的净化百分率同植物群落的郁闭度成正相关,同疏透度成负相关关系;绿地内植物郁闭度的最佳范围为0.70～0.85,疏透度的最佳范围为0.25～0.33。研究结果可为评价现有交通绿化带植物配置情况和日后的城市绿地规划和建设提供技术依据。     

基于小波分析的大豆叶绿素a含量高光谱反演模型

 2003和2004年分别在长春市良种场和中国科学院海伦黑土生态实验站实测了大田耕作与水肥耦合作用下大豆（Glycine max）冠层高光谱反射率 与叶绿素a含量数据,对光谱反射率、微分光谱与叶绿素a含量进行了相关分析;采用归一化植被指数(Normalized diffe rence vegetation index, NDVI)、土壤调和植被指数(Soil-adjusted vegetation index, SAVI)、再归一植被指数(Renormalized difference vegetation index, RDVI)、第二修正比值植被指数(Modified second ratio index, MSRI)等建立了大豆叶绿素a反演模型;应用小波分析对采集的光谱反 射率数据进行了能量系数提取,并以小波能量系数作为自变量进行了单变量与多变量回归分析,对大豆叶绿素a进行了估算。研究结果表明,大 豆叶绿素a 与可见光光谱反射率相关性较好,并在红光波段取得最大值(R²>0.70),但在红边处,微分光谱与大豆叶绿素a的相关性较反射率好 得多,在其它波段则相反;由NDVI、SAVI、RDVI、MSRI等植被指数建立的估算模型可以提高大豆叶绿素a的估算精度（R²>0.75）;小波能量系 数回归模型可以进一步提高大豆叶绿素a含量的估算水平,以一个特定小波能量系数作为自变量的回归模型,大豆叶绿素a回归决定系数R²高达 0.78;多变量回归分析结果表明,大豆叶绿素a实测值与预测值的线性回归决定系数R²均高达0.85。以上结果表明, 小波分析可以对高光谱进 行特征变量提取,并可在一定程度上提高大豆生理参数反演精度。     

水稻冠层结构变化对二向反射率的影响

通过对植被二向反射特性的研究,可反演植被冠层结构信息,如叶面积指数、平均叶角、株高、覆盖率等。在大田晚稻（秀水63）移栽后第26、35、41、49、62、67和86d,对水稻冠层结构及二向反射率进行实测,并分析了二向反射率随冠层结构变化规律,结果表明,植被冠层二向反射率对入射角与观测角的敏感性随着植被冠层结构的变化而变化,蕴涵着丰富的植被结构信息。     

Influence of Seasonal Variations in Soil Water Availability on Gas Exchange of Tropical Canopy Trees

Seasonal variations in environmental conditions influence the functioning of the whole ecosystem of tropical rain forests, but as yet little is known about how such variations directly influence the leaf gas exchange and transpiration of individual canopy tree species. We examined the influence of seasonal variations in relative extractable water in the upper soil layers on predawn leaf water potential, saturated net photosynthesis, leaf dark respiration, stomatal conductance, and tree transpiration of 13 tropical rain forest canopy trees (eight species) over 2 yr in French Guiana. The canopies were accessed by climbing ropes attached to the trees and to a tower. Our results indicate that a small proportion of the studied trees were unaffected by soil water depletion during seasonal dry periods, probably thanks to efficient deep root systems. The trees showing decreased tree water status (i.e., predawn leaf water potential) displayed a wide range of leaf gas exchange responses. Some trees strongly regulated photosynthesis and transpiration when relative extractable water decreased drastically. In contrast, other trees showed little variation, thus indicating good adaptation to soil drought conditions. These results have important applications to modeling approaches: indeed, precise evaluation and grouping of these response patterns are required before any tree‐based functional models can efficiently describe the response of tropical rain forest ecosystems to future changes in environmental conditions.     

Tree growth and death in a tropical gallery forest in Brazil: understanding the relationships among size, growth, and survivorship for understory and canopy dominant species

We studied the relationships between size, growth, and survival for two functional groups, the persistent canopy and understory dominant tree species in a tropical gallery forest in Southeastern Brazil. In 28 plots of 10?×?30?m we tagged, identified, and measured the diameter of all trees with diameter at ground level ??5?cm in 1993/1994, 1998, and 2004. We choose the three dominant canopy species (Protium spruceanum, Copaifera langsdorffii, and Pera glabrata) and two dominant understory species (Ixora brevifolia and Trichilia emarginata) for the comparisons. We assessed the relationship between previous growth rates and mortality, variation in growth and mortality rates among size classes, and temporal correlations in growth rates. Trees (whole community) with null or negative growth had a higher mortality, although this effect was not significant for individual species. Growth patterns were consistent along consecutive periods of evaluation for canopy species, but not for understory species. Canopy species had higher and more variable growth rates than understory species, which we attribute to greater access to light at the canopy level and also to a wider range of light conditions experienced during growth. Canopy species and one understory species, T. emarginata, showed accelerated growth as they became larger. Mortality rates were higher for the smallest trees for the community overall and for P. spruceanum.     

Optical trait indicators for remote sensing of plant species composition: Predictive power and seasonal variability

Most plant species feature similar biochemical compositions and thus similar spectral signals. Still, empirical evidence suggests that the spectral discrimination of species and plant assemblages is possible. Success depends on the presence or absence of faint but detectable differences in biochemical (e.g., pigments, leaf water and dry matter content) and structural properties (e.g., leaf area, angle, and leaf structure), i.e., optical traits. A systematic analysis of the contributions and spatio-temporal variability of optical traits for the remote sensing of organismic vegetation patterns has not yet been conducted. We thus use time series of optical trait values retrieved from the reflectance signal using physical models (optical trait indicators, OTIs) to answer the following questions: How are optical traits related among patterns of floristic composition and reflectance? How variable are these relations in space and time? Are OTIs suitable predictors of plant species composition?We conducted a case study of three temperate open study sites with semi-natural vegetation. The canopy reflectance of permanent vegetation plots was measured on multiple dates over the vegetation period using a field spectrometer. We recorded the cover fractions of all plant species found in the vegetation plots and extracted gradients of species composition from these data. The physical PROSAIL leaf and canopy optical properties model was inverted with random forest regression models to retrieve time series of OTIs for each plot from the reflectance spectra. We analyzed these data sets using correlation analyses. This approach allowed us to assess the distribution of optical traits across gradients of species composition. The predictive performance of OTIs was tested in relation to canopy reflectance using random forest models.OTIs showed pronounced relationships with floristic patterns in all three study sites. These relationships were subject to considerable temporal variability. Such variability was driven by short-term vegetation dynamics introduced by local resource stress. In 72% of all cases OTIs out-performed the original canopy reflectance spectra as indicators of plant species composition. OTIs are also easier to interpret in an ecological sense than spectral bands or features. We thus conclude that optical traits retrieved from reflectance data have a high indicative value for ecological research and applications.