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This study explores the quantitative relationship between canopy structure and the canopy escape ratio (fesc), measured as the ratio of near-infrared reflectance of vegetation (NIRv) to the fraction of absorbed photosynthetically active radiation (fAPAR). We analyzed the correlation between fesc and key indicators of canopy structure—specifically, leaf area index (LAI) and clumping index (CI)—utilizing both Sentinel-2 satellite data and in situ observations. Our analysis revealed a moderate correlation between fesc and LAI, evidenced by an R2 value of 0.37 for satellite-derived LAI, which contrasts with the lower correlation (R2 of 0.15) observed with field-measured LAI. Conversely, the relationship between fesc and CI proved to be significantly weaker (R2 < 0.1), indicating minimal interaction between foliage distribution and light escape at the canopy level. This disparity in correlation strength was further evidenced in time series analysis, which showed little phenological variation in fesc compared to LAI. Our findings elucidate the complexities of estimating fesc based on the NIRv to fAPAR ratio and underscore the need for advanced methodologies in future research to enhance the accuracy of canopy escape models. [ABSTRACT FROM AUTHOR]

Recently, light use efficiency (LUE) models driven by remote sensing data have been widely employed to estimate the gross primary productivity (GPP) of different terrestrial ecosystems at global or regional scales. Furthermore, the two-leaf light use efficiency (TL-LUE) model has been reported to improve the accuracy of GPP estimation, relative to the big-leaf MOD17 model, by separating the entire canopy into sunlit and shaded leaves through the use of constant clumping index estimation (Ω). However, ignoring obvious seasonal changes in the vegetation clumping index (CI) most likely results in GPP estimation errors since the CI tends to present seasonal changes, especially with respect to the obvious presence or absence of leaves within the canopy of deciduous vegetation. Here, we propose a TL-CLUE model that considers the seasonal difference in the CI based on the TL-LUE model to characterize general changes in canopy seasonality. This method composites monthly CI values into two or three Ω values to capture the general seasonal changes in CI while attempting to reduce the potential uncertainty caused during CI inversion. In theory, CI seasonality plays an essential role in the distribution of photosynthetically active radiation absorbed by the canopy (APAR). Specifically, the seasonal difference in CI values mainly considers the state of leaf growth, which is determined by the MODIS land surface phenology (LSP) product (MCD12Q2). Therefore, the one-year cycle (OYC) of leaf life is divided into two (leaf-off and leaf-on) or three seasons (leaf-off, leaf-scattering, and leaf-gathering) according to this MODIS LSP product, and the mean CI of each corresponding season for each vegetation class is computed to smoothen the uncertainties within each seasonal section. With these two or three seasonal Ω values as inputs, the TL-CLUE model by which the seasonal differences in CI are incorporated into the TL-LUE model is run and evaluated based on observations from 84 eddy covariance (EC) tower sites across North America. The results of the analysis reveal that the TL-LUE model widely overestimates GPP for most vegetation types during the leaf-on season, particularly during the growth peak. Although the TL-LUE model shows that the temporal characteristics of GPP agree with the EC observations in terms of general trends, the TL-CLUE model further improves the accuracy of GPP estimation by considering the seasonal changes in the CI. The result of GPP estimation from the TL-CLUE model shows a lower error (RMSE = 2.46 g C m−2 d−1) than the TL-LUE model (RMSE = 2.75 g C m−2 d−1) and somewhat decreases the eight-day GPP overestimation in the TL-LUE model with a constant Ω by approximately 9.76 and 8.970% when adapting three and two Ωs from different seasons, respectively. The study demonstrates that the uncertainty of seasonal disturbance in the CI, quantified by a standard deviation of approximately 0.071 relative to the mean CI of 0.746, is diminished through simple averaging. The seasonal difference in CI should be considered in GPP estimation of terrestrial ecosystems, particularly for vegetation with obvious canopy changes, where leaves go through the complete physiological processes of germination, stretching, maturity, and falling within a year. This study demonstrates the potential of the MODIS CI application in developing ecosystem and hydrological models. [ABSTRACT FROM AUTHOR]

Monitoring gross primary production (GPP) is necessary for quantifying the terrestrial carbon balance. The near-infrared reflectance of vegetation (NIRv) has been proven to be a good predictor of GPP. Given that radiation powers photosynthesis, we hypothesized that (i) the addition of photosynthetic photon flux density (PPFD) information to NIRv would improve estimates of GPP and that (ii) a further improvement would be obtained by incorporating the estimates of radiation distribution in the canopy provided by the foliar clumping index (CI). Thus, we used GPP data from FLUXNET sites to test these possible improvements by comparing the performance of a model based solely on NIRv with two other models, one combining NIRv and PPFD and the other combining NIRv, PPFD and the CI of each vegetation cover type. We tested the performance of these models for different types of vegetation cover, at various latitudes and over the different seasons. Our results demonstrate that the addition of daily radiation information and the clumping index for each vegetation cover type to the NIRv improves its ability to estimate GPP. The improvement was related to foliage organization, given that the foliar distribution in the canopy (CI) affects radiation distribution and use and that radiation drives productivity. Evergreen needleleaf forests are the vegetation cover type with the greatest improvement in GPP estimation after the addition of CI information, likely as a result of their greater radiation constraints. Vegetation type was more determinant of the sensitivity to PPFD changes than latitude or seasonality. We advocate for the incorporation of PPFD and CI into NIRv algorithms and GPP models to improve GPP estimates. [ABSTRACT FROM AUTHOR]

The clumping index (CI) is a commonly used vegetation dispersion parameter used to characterize the spatial distribution of the clumping or random distribution of leaves in canopy environments, as well as to determine the radiation transfer of the canopy, the photosynthesis of the foliage, and hydrological processes. However, the method of CI estimation using the measurement instrument produces uncertain values in various forest types. Therefore, it is necessary to clarify the differences in CI estimation methods using field measurements with various segment lengths in different forest types. In this study, three 100 m × 100 m plots were set, and the CI and leaf area index (LAI) values were measured. The CI estimation results were compared. The results show that the accuracy of CI estimation was affected by different forest types, different stand densities, and various segment lengths. The segment length had a significant effect on CI estimation with various methods. The CI estimation accuracy of the LX and CLX methods increased alongside a decrease in the segment length. The CI evidently offered spatial heterogeneity among the different plots. Compared with the true CI, there were significant differences in the CI estimation values with the use of various methods. Moreover, the spatial distribution of the CI estimation values using the ΩCMN method could more effectively describe the spatial heterogeneity of the CI. These results can provide a reference for CI estimation in field measurements with various segment lengths in different forest types. [ABSTRACT FROM AUTHOR]

The clumping index (CI) quantifies the foliage grouping within distinct canopies relative to randomly distributed canopies, which plays an important role in the vegetation radiative regime. Among the vegetation structure parameters, the global CI map can be retrieved by using multiangle remote sensing observations. The bidirectional reflectance distribution function (BRDF)/Albedo product (MCD43) of the Moderate-Resolution Imaging Spectroradiometer (MODIS) is the crucial input data of the global CI product, which provides validated spatiotemporal continuous directional reflectance data. To determine the impacts of updating the MCD43 product on the consistency and performance of global CI products, CIs retrieved from different MCD43 versions (Collection V005/V006, C5/6) were compared on a global scale and validated with field-measured CI data. The results showed that the global and seasonal comparisons of C5 and C6 CI data are generally consistent. Compared to C5 CI data, C6 CI data have improved quality with more main algorithm retrievals and fewer case of missing data. The comparisons over the field measurements indicate that both versions of CI data agree with field-measured CI data in terms of values and seasonal variations, while C6 CI data (R2 = 0.89, RMSE = 0.05, bias = 0.02) are closer to field CIs than C5 CI data (R2 = 0.80, RMSE = 0.07, bias = 0.03), indicating a higher accuracy for C6 CI data. The monthly CI is recommended for characterizing the overall seasonal patterns of surface CIs. [ABSTRACT FROM AUTHOR]

Canopy clumping index (CI) is a key structural parameter related to vegetation phenology and the absorption of radiation, and it is usually retrieved from remote sensing data based on an empirical relationship with the Normalized Difference between Hotspot and Darkspot (NDHD) index. A rapid production software was developed to implement the CI algorithm based on the Google Earth Engine (GEE) to update current CI products and promote the application of CI in different fields. Daily, monthly, and yearly global CI products are continuously generated and updated in real-time by the software. Users can directly download the product or work with CI without paying attention to data generation. For the application case study, a change detection algorithm, LandTrendr, was implemented on the GEE to examine the global CI trend from 2000 to 2020. The results indicate that the area of increase trend (28.7%, ΔCI > 0.02) is greater than that of the decrease trend (17.1%, ΔCI < −0.02). Our work contributes toward the retrieval, application, and validation of CI. [ABSTRACT FROM AUTHOR]

Estimating clumping indices is important for determining the leaf area index (LAI) of forest canopies. The spatial distribution of the clumping index is vital for LAI estimation. However, the neglect of woody tissue can result in biased clumping index estimates when indirectly deriving them from the gap probability and LAI observations. It is difficult to effectively and automatically extract woody tissue from digital hemispherical photos. In this study, a method for the automatic detection of trunks from Terrestrial Laser Scanning (TLS) data was used. Between-crown and within-crown gaps from TLS data were separated to calculate the clumping index. Subsequently, we analyzed the gap probability, clumping index, and LAI estimates based on TLS and HemiView data in consideration of woody tissue (trunks). Although the clumping index estimated from TLS had better agreement (R² = 0.761) than that from HemiView, the change of angular distribution of the clumping index affected by the trunks from TLS data was more obvious than with the HemiView data. Finally, the exclusion of the trunks led to a reduction in the average LAI by ~19.6% and 8.9%, respectively, for the two methods. These results also showed that the detection of woody tissue was more helpful for the estimation of clumping index distribution. Moreover, the angular distribution of the clumping index is more important for the LAI estimate than the average clumping index value. We concluded that woody tissue should be detected for the clumping index estimate from TLS data, and 3D information could be used for estimating the angular distribution of the clumping index, which is essential for highly accurate LAI field measurements. [ABSTRACT FROM AUTHOR]

In contrast to herbaceous canopies and forests, savannas are grassland ecosystems with sparsely distributed individual trees, so the canopy is spatially heterogeneous and open, whereas the woody cover in savannas, e.g., tree cover, adversely affects ecosystem structures and functions. Studies have shown that the dynamics of canopy structure are related to available water, climate, and human activities in the form of porosity, leaf area index (LAI), and clumping index (CI). Therefore, it is important to identify the biophysical parameters of savanna ecosystems, and undertake practical actions for savanna conservation and management. The canopy openness presents a challenge for evaluating canopy LAI and other biophysical parameters, as most remotely sensed methods were developed for homogeneous and closed canopies. Clumping index is a key variable that can represent the clumping effect from spatial distribution patterns of components within a canopy. However, it is a difficult task to measure the clumping index of the moderate resolution savanna pixels directly using optical instruments, such as the Tracing Radiation and Architecture of Canopies, LAI-2000 Canopy Analyzer, or digital hemispherical photography. This paper proposed a new method using hemispherical photographs combined with high resolution remote sensing images to estimate the clumping index of savanna canopies. The effects of single tree LAI, crown density, and herbaceous layer on the clumping index of savanna pixels were also evaluated. The proposed method effectively calculated the clumping index of moderate resolution pixels. The clumping indices of two study regions located in Ejina Banner and Weichang were compared with the clumping index product over China's landmass. [ABSTRACT FROM AUTHOR]

The development of near-surface remote sensing requires the accurate extraction of leaf area index (LAI) from networked digital cameras under all illumination conditions. The widely used directional gap fraction model is more suitable for overcast conditions due to the difficulty to discriminate the shaded foliage from the shadowed parts of images acquired on sunny days. In this study, a new LAI extraction method by the sunlit foliage component from downward-looking digital photography under clear-sky conditions is proposed. In this method, the sunlit foliage component was extracted by an automated image classification algorithm named LAB2, the clumping index was estimated by a path length distribution-based method, the LAD and G function were quantified by leveled digital images and, eventually, the LAI was obtained by introducing a geometric-optical (GO) model which can quantify the sunlit foliage proportion. The proposed method was evaluated at the YJP site, Canada, by the 3D realistic structural scene constructed based on the field measurements. Results suggest that the LAB2 algorithm makes it possible for the automated image processing and the accurate sunlit foliage extraction with the minimum overall accuracy of 91.4%. The widely-used finite-length method tends to underestimate the clumping index, while the path length distribution-based method can reduce the relative error (RE) from 7.8% to 6.6%. Using the directional gap fraction model under sunny conditions can lead to an underestimation of LAI by (1.61; 55.9%), which was significantly outside the accuracy requirement (0.5; 20%) by the Global Climate Observation System (GCOS). The proposed LAI extraction method has an RMSE of 0.35 and an RE of 11.4% under sunny conditions, which can meet the accuracy requirement of the GCOS. This method relaxes the required diffuse illumination conditions for the digital photography, and can be applied to extract LAI from downward-looking webcam images, which is expected for the regional to continental scale monitoring of vegetation dynamics and validation of satellite remote sensing products. [ABSTRACT FROM AUTHOR]

Leaf area index (LAI) is one of the key biophysical variables required for crop modeling. Direct LAI measurements are time consuming and difficult to obtain for experimental and commercial fruit orchards. Devices used to estimate LAI have shown considerable errors when compared to ground-truth or destructive measurements, requiring tedious site-specific calibrations. The objective of this study was to test the performance of a modified digital cover photography method to estimate LAI in apple trees using conventional digital photography and instantaneous measurements of incident radiation (Io) and transmitted radiation (I) through the canopy. Leaf area of 40 single apple trees were measured destructively to obtain real leaf area index (LAID), which was compared with LAI estimated by the proposed digital photography method (LAIM). Results showed that the LAIM was able to estimate LAID with an error of 25% using a constant light extinction coefficient (k = 0.68). However, when k was estimated using an exponential function based on the fraction of foliage cover (ff) derived from images, the error was reduced to 18%. Furthermore, when measurements of light intercepted by the canopy (Ic) were used as a proxy value for k, the method presented an error of only 9%. These results have shown that by using a proxy k value, estimated by Ic, helped to increase accuracy of LAI estimates using digital cover images for apple trees with different canopy sizes and under field conditions. [ABSTRACT FROM AUTHOR]

The canopy foliage clumping effect is primarily caused by the non-random distribution of canopy foliage. Currently, measurements of clumping index (CI) by handheld instruments is typically time- and labor-intensive. We propose a low-cost and low-power automatic measurement system called Multi-point Linear Array of Optical Sensors (MLAOS), which consists of three above-canopy and nine below-canopy optical sensors that capture plant transmittance at different times of the day. Data communication between the MLAOS node is facilitated by using a ZigBee network, and the data are transmitted from the field MLAOS to a remote data server using the Internet. The choice of the electronic element and design of the MLAOS software is aimed at reducing costs and power consumption. A power consumption test showed that, when a 4000 mAH Li-ion battery is used, a maximum of 8-10 months of work can be achieved. A field experiment on a coniferous forest revealed that the CI of MLAOS may reveal a clumping effect that occurs within the canopy. In further work, measurement of the multi-scale clumping effect can be achieved by utilizing a greater number of MLAOS devices to capture the heterogeneity of the plant canopy. [ABSTRACT FROM AUTHOR]

In terrestrial ecosystems, leaves are aggregated into different spatial structures and their spatial distribution is non-random. Clumping index (CI) is a key canopy structural parameter, characterizing the extent to which leaf deviates from the random distribution. To assess leaf clumping effects on global terrestrial ET, we used a global leaf area index (LAI) map and the latest version of global CI product derived from MODIS BRDF data as well as the Boreal Ecosystem Productivity Simulator (BEPS) to estimate global terrestrial ET. The results show that global terrestrial ET in 2015 was 511.9 ± 70.1 mm yr−1 for Case I, where the true LAI and CI are used. Compared to this baseline case, (1) global terrestrial ET is overestimated by 4.7% for Case II where true LAI is used ignoring clumping; (2) global terrestrial ET is underestimated by 13.0% for Case III where effective LAI is used ignoring clumping. Among all plant functional types (PFTs), evergreen needleleaf forests were most affected by foliage clumping for ET estimation in Case II, because they are most clumped with the lowest CI. Deciduous broadleaf forests are affected by leaf clumping most in Case III because they have both high LAI and low CI compared to other PFTs. The leaf clumping effects on ET estimation in both Case II and Case III is robust to the errors in major input parameters. Thus, it is necessary to consider clumping effects in the simulation of global terrestrial ET, which has considerable implications for global water cycle research. [ABSTRACT FROM AUTHOR]

The in situ leaf area index (LAI) measurement plays a vital role in calibrating and validating satellite LAI products. Digital hemispherical photography (DHP) is a widely used in situ forest LAI measurement method. There have been many software programs encompassing a variety of algorithms to estimate LAI from DHP. However, there is no conclusive study for an accuracy comparison among them, due to the difficulty in acquiring forest LAI reference values. In this study, we aim to use virtual (i.e., computer-simulated) broadleaf forests for the accuracy assessment of LAI algorithms in commonly used LAI software programs. Three commonly used DHP programs, including Can_Eye, CIMES, and Hemisfer, were selected since they provide estimates of both effective LAI and true LAI. Individual tree models with and without leaves were first reconstructed based on terrestrial LiDAR point clouds. Various stands were then created from these models. A ray-tracing technique was combined with the virtual forests to model synthetic DHP, for both leaf-on and leaf-off conditions. Afterward, three programs were applied to estimate PAI from leaf-on DHP and the woody area index (WAI) from leaf-off DHP. Finally, by subtracting WAI from PAI, true LAI estimates from 37 different algorithms were achieved for evaluation. The performance of these algorithms was compared with pre-defined LAI and PAI values in the virtual forests. The results demonstrated that without correcting for the vegetation clumping effect, Can_Eye, CIMES, and Hemisfer could estimate effective PAI and effective LAI consistent with each other (R2 > 0.8, RMSD < 0.2). After correcting for the vegetation clumping effect, there was a large inconsistency. In general, Can_Eye more accurately estimated true LAI than CIMES and Hemisfer (with R2 = 0.88 > 0.72, 0.49; RMSE = 0.45 < 0.7, 0.94; nRMSE = 15.7% < 24.21%, 32.81%). There was a systematic underestimation of PAI and LAI using Hemisfer. The most accurate algorithm for estimating LAI was identified as the P57 algorithm in Can_Eye which used the 57.5° gap fraction inversion combined with the finite-length averaging clumping correction. These results demonstrated the inconsistency of LAI estimates from DHP using different algorithms. It highlights the importance and provides a reference for standardizing the algorithm protocol for in situ forest LAI measurement using DHP. [ABSTRACT FROM AUTHOR]

The Sanjiang Plain is the largest agricultural reclamation area and the biggest marsh area in China. The regional vegetation coverage in this area is vital to local ecological systems, and vegetation growth is affected by natural and anthropogenic factors. The clumping index (CI) is of great significance for land surface models and obtaining information on other vegetation structures. However, most existing ecological models and the retrieval of other vegetation structures do not consider the spatial and temporal variations of CI, and few studies have focused on detecting factors that influence the spatial differentiation of CI. To address these issues, this study investigated the spatial and temporal characteristics of foliage CI in the Sanjiang Plain, analysing the correlation between CI and leaf area index (LAI) through multiple methods (such as Theil−Sen trend analysis, the Mann−Kendall test, and the correlation coefficient) based on the 2001−2015 Chinese Academy of Sciences Clumping Index (CAS CI) and Global LAnd Surface Satellite Leaf Area Index (GLASS LAI). The driving factors of the spatial differentiation of CI were also investigated based on the geographical detector model (GDM) with natural data (including the average annual temperature, annual precipitation, elevation, slope, aspect, vegetation type, soil type, and geomorphic type) and anthropogenic data (the land use type). The results showed that (1) the interannual variation of foliage CI was not obvious, but the seasonal variation was obvious in the Sanjiang Plain from 2001 to 2015; (2) the spatial distribution of the multiyear mean CI of each season in the Sanjiang Plain was similar to the spatial distribution of the land use type, and the CI decreased slightly with increases in elevation; (3) the correlation between the growing season mean CI (CIGS) and the growing season mean LAI (LAIGS) time series was not significant, but their spatial distributions were negatively correlated; (4) topographic factors (elevation and slope) and geomorphic type dominated the spatial differentiation of foliage CI in the Sanjiang Plain, and the interactions between driving factors enhanced their explanatory power in terms of the spatial distribution of foliage CI. This study can help improve the accuracy of the retrieval of other vegetation structures and the simulation of land surface models in the Sanjiang Plain, providing invaluable insight for the analysis of the spatial and temporal variations of vegetation based on CI. Moreover, the results of this study support a theoretical basis for understanding the explanatory power of natural and anthropogenic factors in the spatial distribution of CI, along with its driving mechanism. [ABSTRACT FROM AUTHOR]

The fire-tolerant eucalypt forests of south eastern Australia are assumed to fully recover from even the most intense fires; however, surprisingly, very few studies have quantitatively assessed that recovery. The accurate assessment of horizontal and vertical attributes of tree crowns after fire is essential to understand the fire's legacy effects on tree growth and on forest structure. In this study, we quantitatively assessed individual tree crowns 8.5 years after a 2009 wildfire that burnt extensive areas of eucalypt forest in temperate Australia. We used airborne LiDAR data validated with field measurements to estimate multiple metrics that quantified the cover, density, and vertical distribution of individual-tree crowns in 51 plots of 0.05 ha in fire-tolerant eucalypt forest across four wildfire severity types (unburnt, low, moderate, high). Significant differences in the field-assessed mean height of fire scarring as a proportion of tree height and in the proportions of trees with epicormic (stem) resprouts were consistent with the gradation in fire severity. Linear mixed-effects models indicated persistent effects of both moderate and high-severity wildfire on tree crown architecture. Trees at high-severity sites had significantly less crown projection area and live crown width as a proportion of total crown width than those at unburnt and low-severity sites. Significant differences in LiDAR -based metrics (crown cover, evenness, leaf area density profiles) indicated that tree crowns at moderate and high-severity sites were comparatively narrow and more evenly distributed down the tree stem. These conical-shaped crowns contrasted sharply with the rounded crowns of trees at unburnt and low-severity sites and likely influenced both tree productivity and the accuracy of biomass allometric equations for nearly a decade after the fire. Our data provide a clear example of the utility of airborne LiDAR data for quantifying the impacts of disturbances at the scale of individual trees. Quantified effects of contrasting fire severities on the structure of resprouter tree crowns provide a strong basis for interpreting post-fire patterns in forest canopies and vegetation profiles in Light Detection and Ranging (LiDAR) and other remotely-sensed data at larger scales. [ABSTRACT FROM AUTHOR]

The foliage Clumping Index (CI) is an important vegetation structure parameter that allows for the accurate separation of sunlit and shaded leaves in a canopy. The CI and its seasonality are critical for global Leaf Area Index (LAI) estimating and ecological modelling. However, the cover of snow tends to reduce the reflectance anisotropy of the vegetation canopy and thus probably influences CI estimates. In this paper, we investigate the influence of snow on the magnitude and seasonal variation of the CI retrieved from Moderate-resolution Imaging Spectroradiometer (MODIS) Bidirectional Reflectance Distribution Function (BRDF) products based on field-measured CI and statistics from the global MODIS CI product. We find that the backup algorithm can effectively correct abnormally large CI values and obtain more reasonable CI retrievals than the main algorithm without any constraints in snow-covered areas. Validation indicates that the time-series CI product shows the potential in investigating the trajectories of the clumping effect in snow seasons. For evergreen forests, the clumping effect is relatively stable throughout the year; however, for deciduous vegetation types, CI values tend to display significant seasonal variations. This study suggests that the latest version of the global MODIS CI product, in which the backup algorithm is used to process the snow-covered pixels, has improved accuracy for CI retrievals in snow-covered areas and thus is probably more suitable as the input parameter for ecological and meteorological models. [ABSTRACT FROM AUTHOR]

Directional gap probability or gap fraction is a basic parameter in the optical remote sensing modeling. Although some approaches have been proposed to estimate this gap probability from remotely sensed measurements, few efforts have been made to investigate the scaling effects of this parameter. This paper analyzes the scaling effect through aggregating the high-resolution directional gap probability (pixel size of 20 meters) estimated from leaf area index (LAI) images of VALERI database by means of Beer's law and introduces an extension of clumping index, Ĉ, to compensate the scaling bias. The results show that the scaling effect depends on both the surface heterogeneity and the nonlinearity degree of the retrieved function. Analytical expressions for the scaling bias of gap probability and Ĉ are established in function of the variance of LAI and the mean value of LAI in a coarse pixel. With the VALERI dataset, the study in this paper shows that relative scaling bias of gap probability increases with decreasing spatial resolution for most of land cover types. Large relative biases are found for most of crops sites and a mixed forest site due to their relative large variance of LAI, while very small biases occur over grassland and shrubs sites. As for Ĉ, it varies slowly in the pure forest, grassland and shrubs sites, while more significantly in crops and mixed forest.