International Journal of Remote Sensing Vol. 31, No.

16, 20 August 2010, 43194333

Linear spectral mixture analysis of Landsat TM data for monitoring invasive exotic plants in estuarine wetlands
MEIMEI HE, BIN ZHAO*, ZUTAO OUYANG, YANER YAN and BO LI Coastal Ecosystems Research Station of the Yangtze River Estuary, Ministry of Education Key Laboratory for Biodiversity Science and Ecological Engineering, Institute of Biodiversity Science, Fudan University, Shanghai 200433, PR China
(Received 2 February 2008; in final form 21 January 2009) This study assessed the feasibility of spectral mixture analysis (SMA) of Landsat thematic mapper (TM) data for monitoring estuarine vegetation at species level. SMA modelling was evaluated, using w2 test, by comparing SMA fraction images with a precisely classified QuickBird image that has a higher spatial resolution. To clearly understand the strengths and weaknesses of SMA, eight SMA models with different endmember combinations were assessed. When the TM data dimension for SMA and the endmember number required were balanced, a model with three endmembers representing water and two vegetation types was most accurate, whereas a model with five endmembers approximated the actual surface situation and generated a relatively accurate result. Our results indicate that an SMA model with appropriate endmembers had relatively satisfactory accuracy in monitoring vegetation. However, errors might occur in SMA fraction images, especially in models with an inappropriate endmember combination, and the errors were mainly distributed in areas filled with water or near water. Therefore, short vegetation usually submerged during high tide tended to be poorly predicted by SMA models. These results strongly suggest that tide water has a great influence on SMA modelling, especially for short vegetation.

1.

Introduction

Estuarine wetlands, where terrestrial and aquatic species live together, are productive ecosystems that are ecologically important and provide various ecosystem services (Pennings and Callaway 1992, Phinn et al. 1996, Klemas 2001, Zhao et al. 2004). These precious wetlands are subjected to considerable loss that is caused by natural and anthropogenic factors (Yue et al. 2003, Niemi et al. 2004, Levinson 2005). Among these factors, the invasions by exotic plants have become extremely serious because invasive plants can change the vegetation structure and functions of wetland ecosystems through excluding native plants (Pimentel et al. 2000, Underwood et al. 2006). Nowadays, many coastal and estuarine marshes are under serious threat from invasions by exotic plants (Grosholz 2002). A better understanding of spatial dynamics of invasive species in estuarine wetlands will facilitate well-targeted and proactive efforts in controlling spread of invasive species. However, it is hard to investigate vegetation distribution in estuarine wetlands using traditional approaches, such as field sampling and survey, because these approaches are time-consuming, cost inefficient, and
*Corresponding author. Email: zhaobin@fudan.edu.cn
International Journal of Remote Sensing ISSN 0143-1161 print/ISSN 1366-5901 online # 2010 Taylor & Francis http://www.tandf.co.uk/journals DOI: 10.1080/01431160903252343

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sometimes impractical (e.g. the areas are inaccessible) (Ozesmi and Bauer 2002). Fortunately, remote sensing has the potential to update surveys by repeatedly covering large areas (Ozesmi and Bauer 2002, Rosso et al. 2005, Shabanov et al. 2005, Aplin 2006), especially by surveying areas that are difficult to access. Therefore, great progress has been made over the past decades with the application of remote sensing to wetlands (Ramachandran et al. 1998, Klemas 2001, Ozesmi and Bauer 2002). Among the remote sensing techniques, the Landsat platform has provided scientists with medium-resolution satellite imagery for over 30 years, and has undoubtedly become a popular platform for wetland researches. However, a practical difficulty facing us is that the diameters of new patches of invasive plants in coastal wetlands may be less than those covered by the maximum average resolution 30 m of Landsat Thematic Mapper (TM), meaning that the vegetation information acquired by TM is mixed pixel, and often contains several plant species. To overcome this difficulty, many efforts have been made to identify vegetation patches at the sub-pixel level (Levinson 2005, Shanmugam et al. 2006). Some studies have shown that sub-pixel classification techniques, such as spectral mixture analysis (SMA), are likely to give more accurate results than the traditional per-pixel classification techniques (i.e. iterative self-organizing data analysis, ISODATA; and maximum likelihood classification, MLC) (Foody and Cox 1994, Bateson and Curtiss 1996, Pu et al. 2003, Shanmugam et al. 2006). The results of SMA are a set of fraction maps providing not only spatial but also abundance information for each vegetation type under investigation (Settle and Drake 1993, Schmidtlein and Sassin 2004). For this reason, SMA has been used to investigate wetland vegetation for many purposes, such as mapping non-photosynthetic vegetation (NPV) (Roberts et al. 1993) and studying mature stands of chaparral (Roberts et al. 1998). Previous studies have treated all plants as one category (presented as vegetation), without identifying the plant communities at species level, which is obviously inadequate for monitoring invasive plants, especially at the early phase of invasion when the species have only a small invading populations and patches. Some recent studies have started to consider vegetation classification at species level. For example, Li et al. (2005) and Rosso et al. (2005) have investigated the spatial distributions of several marsh plant species using SMA technique on Airborne Visible Infrared Imaging Spectrometer (AVIRIS) data at San Pablo Bay and San Francisco Bay, respectively. Their results have shown that the species abundance maps derived with SMA were in good agreement with their field observations. To classify vegetation at species level, many pioneering efforts have focused on hyperspectral data (Schmidtlein and Sassin 2004, Rosso et al. 2005, Underwood et al. 2006). Because hyperspectral data have large numbers of contiguous bands, more class types could be contained in SMA, which may be limited by using multispectral data. However, these studies have some disadvantages in monitoring vegetation because such images are usually obtained with aerial platforms that depend on occasional and often irregularly timed flights. Therefore, it is practically difficult to monitor the whole invasion process within a particular time frame, because of the lack of historical data (Aplin 2006) and the inadequacy of the repetitive observations. In this respect, our work was an attempt to apply SMA to Landsat TM images, and to study the potential use of TM images to determine the distributions of invasive plants at species level. It should be noted that it is not easy to apply SMA in an estuarine wetland because the complex hydrological conditions make the surface soil highly variable. In estuarine wetlands, various plant communities may occur in heterogeneous

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mosaics, with bare soil and open water. A critical step in SMA is to select the proper components, or endmembers. The complex surface conditions increase the difficulty of endmember selection, especially under the inherent mathematical limitations of SMA, i.e. the number of endmembers cannot exceed the number of spectral bands used. Therefore, another purpose of our work was to determine how endmember schemas affect mapping accuracy, and how the moist background influences SMA modelling. Moreover, our work provided a cross-validation technique for comparing the fractional images derived from SMA of TM data with high-spatial-resolution imagery.

2. 2.1

Materials and methods Study site

Our study area is part of brackish marshes in the Yangtze River Estuary, China (figure 1), which typically consists of marshy land that is growing rapidly (Zhao et al. 2008). It is characterized by continuous expansion because of the large amounts of sediment brought down by the Yangtze River (Ma et al. 2003). The tidal fluctuations are regular and semi-diurnal, and there are two distinct periods of ebb and flood tides during the day. The width of the tideland uncovered by tidewater is about 1.5 km during the neap tide, whereas almost all the tideland is submerged by tidewater during the spring tide (Sun and Cai 2001). The vegetation is dominated by Phragmites australis (hereafter Phragmites), Scirpus mariqueter (hereafter Scirpus) and Spartina alterniflora (smooth cordgrass, hereafter Spartina). Phragmites and Spartina usually

Figure 1.

Location of the study area.

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grow up to 1.52.5 m, whereas Scirpus only reaches a height of 0.30.7 m (Zhao et al. 2009). Spartina was intentionally introduced into Dongtan in 2001 for land reclamation, because of its capacity to increase sediment accretion (Chen et al. 2004). After a burst of growth and expansion for several years, Spartina has become a dominant species, and a rapid succession has occurred. The invasion of Spartina has a tremendous impact on the ecosystems, seriously threatening the native ecosystem and coastal aquaculture. The native species, Phragmites and Scirpus, are being rapidly replaced by Spartina, which modifies the structure of the marsh dramatically (Li et al. 2009). The native species Scirpus usually grows in tidal zones with an elevation between 2.0 and 2.9 m, whereas Phragmites dominates higher zones close to the sea wall, and forms a mosaic with Spartina. The distributions of these species have produced a zonal pattern of plant communities (Wang et al. 2009), providing more information for the classification of the vegetation by remote sensing. The phenologies of Phragmites and Spartina are different. Phragmites begins to germinate in early April, has its rapid growth stage spanning from June to mid August, flowers in mid October, and senesces in late November. In contrast, Spartina emerges in May, has its rapid vegetative growth from June to early September, flowers in late September, and dies away in late December. These differences provide important information for the classification of remotely sensed imageries. 2.2 Data

2.2.1 Remotely sensed data. Two different sets of remotely sensed data, including TM and QuickBird, were used in this study. The Landsat TM data were acquired on 20 April 2006, under clear atmospheric conditions. Six bands with 30 m resolution (TM-1: 0.450.52 mm; TM-2: 0.520.60 mm; TM-3: 0.600.69 mm; TM-4: 0.760.90 mm; TM-5: 1.551.75 mm and TM-7: 2.082.35 mm) were extracted from the TM scene. The original digital number (DN) data were calibrated and transformed into reflectance values with a processing module in the software package ENVI version 4.1. The QuickBird data were obtained on 10 May 2006, and comprised five bands: (1) panchromatic band (450900 nm) with a spatial resolution of 0.61 m; (2) blue band (450520 nm); (3) green band (520600 nm); (4) red band (630690 nm); and (5) infrared band (760900 nm). Bands 25 have a spatial resolution of 2.44 m. An ultimate image composed of the last four bands, with a spatial resolution of 0.61 m, was derived after data fusion. Geometrical calibration and atmospheric calibration were performed first, followed by object-oriented classification using eCognition. Because the spatial resolution of QuickBird (0.61 m) is high enough to identify small plant patches, with some prepared classification by field validation, we found that the QuickBird could achieve a high level of classification accuracy (above 90%). It thus approximated the real surface cover, and could be considered a good proxy to assess the accuracy of SMA modelling. The accuracy of QuickBird images was assessed with a confusion matrix using 150 field points, with ground cover situation and global positioning system (GPS) information used as validation. In order to reduce the disturbances caused by tide water, both the TM imagery and QuickBird imagery were obtained under similar tidal period, which was 147 cm when QuickBird image was taken and 140 cm when TM image was taken, and were further carefully cross-registered and geo-referenced, with root mean square error (RMSE) of less than 0.5 TM pixel. 2.2.2 Field data and spectral characteristics of the main components. To assess the spectral distinction of the dominant plant species, soil and water, and to estimate the

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feasibility of remotely sensed SMA, an Analytical Spectral Device (ASD) FieldSpec Handheld (ASD, Inc., Boulder, CO), with a detection region of 3501050 nm and a spectral resolution of 3 nm, was used for the field measurements. Considering the phenological characteristics of the plants, the spectral data were collected on 10 May 2007, coinciding with the season when the remotely sensed data were obtained. The field spectral data were acquired at 0.51.0 m above the canopy with a field of view of 25 . All data were collected between 10:00 and 14:00 on cloud-free days with neap tide. Samples were designated at separate monoculture patches and nine spectrum measurements were made and averaged for each patch. Meanwhile, plant spectral data used in this paper were all collected in sites without water coverage. The field spectroscopy was used to estimate the spectral distinctions among the major marsh components. According to our field spectral analysis, the spectra of the three dominant species had distinct characteristics (figure 2). The reflectance curve of Scirpus showed differences in the magnitude of reflectance, but the overall shape of the curve was similar to that of Phragmites. The reflectance curve of Spartina was different from those of Phragmites and Scirpus in both magnitude and shape. Because the spectral difference around 600700 nm resulted from the chlorophyll content of the plant leaves (Hunt et al. 2004), the spectral shapes of the three dominant species indicated that Phragmites had the highest chlorophyll content, and Spartina had the lowest. This result was consistent with the field observation that Spartina just started to emerge and its ramets were still under its own standing litter in early May. The spectral differences among the three species provided information on endmember selection for the next step in SMA modelling. Figure 2 also shows that dry mud had higher reflectance than washy mud at all wavelengths, which prompted us to pre-set an endmember for washy mud in the SMA model. Moreover, because spectroscopy was only used to validate the differences between categories, and the spectral properties of the same plant at a certain phonological stage do not basically change from year to year, the spectra collected one year after the satellite images had been taken should be reliable in evaluating spectral separability. In other respects, our fieldwork supported the field observations and validated the point selection as well as located the pure patches for categories.

(a)
60 50 Phragmites Spartina Scirpus

(b )

Reflectance (%)

Dry mud Washy mud

40 30 20 10 0

360 400 440 480 520 560 600 640 680 720 760 800 840 880 920 960

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Figure 2.

Field reflectance spectra of major marsh components.

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2.2.3 Reference preparation for accuracy evaluation. To assess the accuracy of the SMA unmixing results, the synchronous QuickBird image was first classified as a reference for the subsequent evaluation of the accuracy of the TM fraction maps. As our concern was to identify a particular species, especially the invasive Spartina, the final classification was divided into six classes: Phragmites, Spartina, Scirpus, washy mud, water and road. To ensure high classification accuracy of reflecting the real situation, a total of 150 ground data points were collected in the field, and then a Training and Test Area (TTA) was established for the assessment of the confusion matrix. After the initial classification, some minor errors were carefully corrected to ensure that the final results were as accurate as possible. 2.3 Endmember selection and spectral mixture analysis

The flow chart in figure 3 illustrates the framework of the SMA and accuracy assessment in this study, in which one of the critical steps was endmember selection. The criteria for a good set of endmembers include linear independency, spectral representativity and spatial generality (Li et al. 2005). Generally, the appropriate endmembers were acquired with two approaches: (1) using the field- or laboratorymeasured spectra as endmembers; and (2) deriving the representative endmembers

Calibrated TM imagery

Calibrated QuickBird imagery

Minimum noise fraction (MNF) transformation

Normalized difference index transformation Classification

Pure pixel index (PPI) calculation

N-dimensional visualization

NDVI extraction

Endmembers selection

Resolution matching (30 m 30 m)

Spectral mixture analysis (SMA)

Correlation analysis

Endmember fraction derivation

Accuracy assessment

Abundance calculation

Figure 3.

Flow chart illustrating the framework of SMA and accuracy assessment in this study.

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directly from the image spectra (Small 2003). Laboratory- and field-measured spectra are standardized but they do not usually relate well to the remotely sensed images. However, the image-derived spectra as endmembers are more representative. Because some researchers believed that image-derived spectra are the optimal choice for SMA applications (Peddle and Smith 2005, Chen and Vierling 2006), we used image-derived spectra as the endmembers in this study. The endmembers were selected based on the pure pixels index (PPI) calculation and N-dimensional visualization of transformed data as the following procedures. 2.3.1 Endmember selection through calculating the PPI. To reduce the noise in the TM image, a forward minimum noise fraction (MNF) transformation was applied, followed by the PPI calculation for each pixel. The MNF analysis of the TM image showed that the first MNF band had an eigenvalue of 212.33 and the sixth band had an eigenvalue of 2.36, at which many of marsh structural features disappeared. The eigenvalue of the fourth band dropped sharply to 6.08 from a value of 12.11 for the third band. This indicates the first three MNF bands contain the major data variance, while bands 4,6 contain less information. Accordingly, it suggests the intrinsic endmembers for TM data are 3,5. Based on the result of MNF, the PPI introduced determined most of the spectrally pure pixels in the image. Several pure patches of different categories had been identified previously from field observations, and were geographically located on the image with GPS data. The pixels with highest PPI values within the located pure patches were then selected as the endmembers for each category. Five endmembers were finally defined under the data dimension of the TM image. The endmembers and their PPI values are listed in table 1. 2.3.2 Endmember selection through index transformation. Rogers and Kearney (2004) have also suggested that the SMA model can be greatly enhanced if the variability of the inner-endmember category is reduced. To eliminate the influence of water content on the spectra of the soil endmember, as well as some interfering signatures, three normalized difference indices were used here: 1. Normalized difference water index (NDWI): (band 3 band 5) / (band 3 band 5); 2. Normalized difference soil index (NDSI): (band 5 band 4) / (band 5 band 4); 3. Normalized difference vegetation index (NDVI): (band 4 band 3) / (band 4 band 3). Of these, NDWI is used to detect open-water areas because water is more reflective in band 3 than that in band 5 of TM (McFeeters 1996); NDSI is calculated from the band pair based on the representative spectra of soil and NDVI (Deering 1978) is well established as the standard for vegetation indices (Shanmugam et al. 2006). If the three indices are envisaged as axes and oriented vertically, water appears at the top of

Table 1. List of endmembers (EMs). Phragmites Abbreviation PPI value Ph 3056 Spartina Sp 2035 Scirpus Sc 2146 Water W 3722 Washy mud Wm 2027

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Phragmites

Spartina

Washy mud

Figure 4. Distribution of pixels in the NDX transformation data spaces. The three axes in the figure denote NDWI (axis 1), NDVI (axis 2) and NDSI (axis 3), respectively.

the NDWI axis, vegetation at the top of the NDVI axis and most soil at the top of the NDSI axis. Thus, these indices can gather pixels with similar spectra, and form a data space with a triangular data cloud. The points inside the triangle can be seen as mixtures, whereas the points around the vertices are supposed to be pure and could be selected as endmembers. Using this approach, three endmembers, Phragmites, Spartina and washy mud, were selected from the corners of the data cloud for SMA modelling (figure 4). After the endmember selection, the SMA was carried out with a processing module in the software package ENVI version 4. To evaluate the influence of endmember combinations on SMA accuracy, models were grouped into three-, four- and fiveendmember ones. Because the purpose of this study was to distinguish Spartina from Phragmites, the endmembers representing Spartina and Phragmites were always included in each model, whereas other endmembers were combined with these two. Thus, a total of eight SMA models were analysed on the TM image. The endmember combinations are listed in table 2. 2.4 SMA mapping accuracy assessment

To match the spatial resolution of the TM image for later analysis, the classified QuickBird image was regrouped to pixels of 30 m 30 m. The pixel coordinates of TM were located on the QuickBird image, and then a 30 m 30 m grid was generated with the Arc Info Workstation. The grids centre was the coordinates of the TM data. In each grid the areas of the classified categories were counted to represent the real percentage of each endmember. Thus, the calculated percentage can be compared with the abundance ratio derived with the SMA models. Then, w2 tests were pixel-by-pixel performed to compare the model results (model-predicted fraction image from TM) with the real situation (percentages counted from the classified QuickBird image). A significant difference between the pixels in the tests implied the poor accuracy of the SMA model, and vice versa.

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Table 2. SMA models and their prediction accuracy. Number of pixels different from QuickBird image (a 0.05) Endmember Ph Sp W Ph Sp Sc Ph Sp Wm Ph Sp Wm 553 1241 649 718 Percentage of the inaccurate pixels (%) 12.14 27.25 14.25 15.77

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Group 3EMs 3EMs (with NDX transferred data) 4EMs 5EMs

Code 3EMw 3EMsc 3EMwm 3EMnd

Average RMSE 0.007956 0.009485 0.007360 0.000586

4EMwsc 4EMwwm 4EMwmsc 5EM

Ph Sp Sc W Ph Sp W Wm Ph Sp Sc Wm Ph Sp Sc W Wm

1023 714 670 718

22.46 15.68 14.71 15.77

0.001833 0.001896 0.006429 0.001440

3. 3.1

Results and discussion QuickBird classification

The classification accuracy of QuickBird is shown in table 3. The accuracy of the QuickBird images was assessed with a confusion matrix using 150 field points, with the ground cover situation and GPS information used as validation. These results indicate a highly accurate, stable and reliable classification result, with an overall classification accuracy of 93.18% and an overall kappa statistic of 0.9054. As mentioned above, some manual checks and corrections were made to ensure that the classification reached a higher level of accuracy after classification. Thus, the classified image represented the real land-cover situation and could be used as a proxy to assess the accuracy of the SMA unmixing results. 3.2 Unmixing results

3.2.1 c2 tests of pixel-by-pixel comparisons. w2 tests of pixel-by-pixel comparisons between classified QuickBird image and SMA endmember fraction image were made, and the significant results were counted and listed in table 2. Of all eight
Table 3. Accuracy assessment of object-oriented classification of QuickBird images. KIA, Kappa Index of Agreement. Class Phragmites Spartina Scirpus Water Washy mud Road Producer 0.9962 0.9778 0.7914 0.6563 0.9430 0.8539 User 0.9910 0.8939 0.9703 1.0000 0.9362 1.0000 KIA per class 0.9934 0.9692 0.7681 0.6373 0.9360 0.8526 Overall accuracy 0.9319 KIA 0.9055

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SMA models, 3EMw had the highest accuracy and 3EMsc the lowest accuracy, suggesting that PhragmitesSpartinawater was the best endmember combination for SMA modelling, whereas PhragmitesSpartinaScirpus was the worst one. The models 3EMsc and 4EMwsc were less accurate because they included the endmember Scirpus. Model 4EMwmsc also included Scirpus, but was reasonably accurate because it included the endmember washy mud. This phenomenon is comprehensible because Scirpus only grows in the lower tidal areas, with a height of less than 0.7 m, thus a water background greatly influences the distinction of Scirpus. However, Scirpus prefers to grow in a washy-mud habitat, so Scirpus could not be distinguished perfectly, which could not be treated as a good representative endmember in this study. Data transformation is designed to reduce the variation within the endmember types, but it cannot solve the problem of intra-endmember similarity. In our study, a spectral similarity existed among the different plant communities, Spartina with standing litter, and mud, so it could not be eliminated by data transformation. Therefore, the accuracy of model 3EMnd was not obviously improved by transforming the data into normalized difference indices. Rogers and Kearney (2004) have reported that the transformation of normalized difference can reduce the spectral variability of soil, water and some principal vegetation components derived from TM data. However, this transformation did not offer a significant advantage for our classification of vegetation at species level. Because in the classification of model 5EM the ground cover included Phragmites, Spartina, Scirpus, washy mud and water which approximated the real conditions of the area, it should be highly accurate. However, our data dimension, as described above, was less than four, and thus this model was only moderately accurate. It should be noted that, in the three-endmember group, model 3EMsc had a much higher RMSE than others because it included the component Scirpus. Similarly, model 4EMwmsc had the highest RMSE in the four-endmember group because it included Scirpus. This result suggested that an SMA model including a wrong or inappropriate endmember would greatly increase RMSE. Conversely, model accuracy does not always increase as RMSE decreases. For example, 3EMw had the highest accuracy but a fairly high RMSE, whereas 3EMnd had moderate accuracy with the lowest RMSE. This is because extra information from multiple bands helped increase the robustness of the SMA least-squares solution. Thus, RMSE does not seem to be an adequate criterion for evaluating the performance of the SMA model. To analyse the spatial variation of the SMA model that is influenced by environmental factors, the significant results from pixel-by-pixel w2 tests and their values were mapped (figures 5 and 6). As the distance increased from the sea wall to the sea, the cell numbers with high accuracy decreased (figure 5). In particular, in the water-filled areas or areas near water, the endmember fractions were poorly predicted in all SMA models. In comparison, the areas covered by dense vegetation seemed to be predicted well with the SMA models, except model 3EMsc including Scirpus (figure 6). This result indicates that tide water had a great influence on SMA modelling.

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4EMwmsc

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0 80

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Figure 5.

Distance from the sea wall and the number of inaccurate pixels.

4.

Conclusions

The special emphasis of this study was placed on the applicability of SMA with TM data to the detection of invasive plants in estuarine wetlands. w2 tests of comparisons between an SMA-model-predicted fraction and a QuickBird classification image provide a measurement to better understand the reliability, the strengths and the

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Figure 6. Occurrence of inaccurate pixels in the image for each of SMA models. Pixels in red are inaccurate, while blue ones are accurate, assessed by chi-square. A darker colour represents a higher chi-square value.

weaknesses of SMA models applied to such detection. Several SMA models with various endmember combinations were analysed and their results were evaluated. A proper endmember combination generated satisfactory SMA results, and had a relatively good capacity to predict vegetation. However, the assessment of accuracy showed that certain errors occurred in the SMA fraction images. Most of these inaccurate predictions were distributed in areas filled with water or near water, indicating that tidal water had a great effect on SMA modelling. Moreover, transforming the TM data to reduce the endmember spectral variation did not actually improve the accuracy of the SMA modelling, which may not be adequate for the detection of invasive plants. It is noteworthy that the multiple endmember spectral mixture (MESMA) (Roberts et al. 1998) assumes that individual pixels contain limited endmembers, although the whole image may contain many spectrally distinct components, which allow the optimization of each pixel. Therefore, MESMA has a great potential for vegetation monitoring in estuarine wetlands, and further work needs to be done using this technique to monitor invasive plants more efficiently. Acknowledgements This work was supported by the National Basic Research Program of China (No. 2006CB403305), the Science and Technology Commission of Shanghai (No. 07DZ12038-2), the National Natural Science Foundation of China (No. 30870409 and 40471087) and the Program for New Century Excellent Talents in University

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(NCET-06-0364) funded by the Ministry of Education of China. We thank Chongming Dongtan National Natural Reserve and the students in our laboratories for their assistance in field sampling. References
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