Ecological Archives E090-233-D1

James R. Kellner, David B. Clark, and Michelle A. Hofton. 2009. Canopy height and ground elevation in a mixed-land-use lowland Neotropical rain forest landscape. Ecology 90:3274.


INTRODUCTION

Light detection and ranging (LiDAR) is powerful remote sensing technology for quantifying the structure and dynamics of forests at large spatial scales (Drake et al. 2003, Roth et al. 2007, Kellner et al. 2009). By recording the return time of emitted laser pulses, aircraft-mounted sensors acquire measurements of canopy and ground elevation at high spatial resolution, providing detailed measurements of vertical and horizontal vegetation structure (Fig. 1). In raw form, LiDAR measurements record the time interval required for an emitted laser pulse to travel from the sensor to reflecting surfaces (e.g., leaves, branches and the underlying ground surface) and return to the sensor. By combining the travel time estimate with three dimensional information on the orientation and position of the LiDAR system at the time of laser firing, the elevation and horizontal position of each reflecting surface can be calculated relative to a reference frame, such as WGS 1984 (e.g., Hofton et al. 2000). Elevation measurements can be processed further to generate images, such as digital elevation or canopy models (Clark et al. 2004), or analyzed directly (Lefsky et al. 1999, Asner et al. 2008, Kellner et al. 2009). The data presented here have been processed, screened, and validated using extensive field studies to generate easting and northing mapped coordinate locations (UTM zone 16 North), and canopy and ground elevation that are precise and accurate.

This data set contains 127,849,839 observations of canopy and ground elevation in 128 km2 of mixed-land-use lowland Neotropical rain forest in the Atlantic lowlands of Costa Rica (Fig. 2). The study area includes all 16 km2 of La Selva Biological Station, a site of long-term research in basic and applied tropical forest ecology, current and abandoned plantations outside the La Selva property, and a portion of Braulio Carrillo National Park along an elevation gradient on Barva Volcano. The study area is defined by a mixed history of historical land use that encompasses terre-firme old-growth forest, secondary forest, swamp forest, and a wide range of current and former human land uses that are representative of contemporary continental rain forests. Previous studies have used these data to quantify size-frequency distributions of canopy gaps, and to determine whether canopy height within old-growth forest at La Selva was close to steady-state equilibrium under the 1997–2006 disturbance regime (Kellner et al. 2009, Kellner and Asner 2009). LiDAR data can be applied to a wide range of questions in basic and applied science, including calculation of above-ground carbon stocks (Drake et al. 2002), photosynthetically-active radiation (Chasmer et al. 2008), rainfall interception and water budgets (Roth et al. 2007), canopy and gap dynamics (Vepakomma et al. 2008, Kellner et al. 2009), individual-based tree demography (Yu et al. 2006), understory light environments and vegetation structure (Asner et al. 2008, Chasmer et al. 2008) and other questions (Turner et al. 2003, Chambers et al. 2007). The data are also a valuable resource from a well-studied tropical rain forest for teaching and education, and provide a quantitative baseline against which future conditions can be assessed.

METADATA

CLASS I. DATA SET DESCRIPTORS

A. Data set identity: Canopy height and ground elevation in a mixed-land-use lowland Neotropical rain forest landscape.

B. Data set identification code: The data are in 511 comma-delimited ASCII text files named tilexxx.txt, where xxx is replaced with the respective tile number from 001–511. Download the entire data set here, or to download selected data tiles, rather than the entire data set, please click here to go to a page listing all 511 files with links. Each file represents a 500 × 500 m contiguous area. Tile locations are shown graphically in Fig. 3, and can be indexed in a GIS using the shapefile (see section V.C.2), or in text format in Table 1.

C. Data set description:

1. Principal investigators:

James R. Kellner, Department of Plant Biology, The University of Georgia, Athens, Georgia, 30602 USA (Department of Global Ecology, Carnegie Institution for Science, Stanford, California, USA). jkellner@stanford.edu

David B. Clark, Department of Biology, University of Missouri, St-Louis, St. Louis, Missouri, 63121 USA and La Selva Biological Station, Puerto Viejo de Sarapiquí, Costa Rica. dbclark@sloth.ots.ac.cr

Michelle A. Hofton, Department of Geography, University of Maryland, College Park, Maryland, 20742 USA. mhofton@umd.edu

2.  Abstract: We obtained spatially extensive canopy height measurements using airborne remote sensing to characterize the structure and dynamics of a tropical rain forest landscape. Light detection and ranging (LiDAR) is a remote-sensing technology that acquires measurements of canopy height and ground elevation. By recording the return time of laser pulses emitted by aircraft-mounted sensors, LiDAR systems quantify the structure and geometry of individual trees and canopy height and enable estimation of the vertical and horizontal distribution of biomass using millions of accurate height measurements. This data set contains 127,849,839 records from 128 square kilometers of tropical wet forest in the Atlantic lowlands of Costa Rica (mean sampling density is 1.99 observations/m2). The study area includes all 16 square kilometers of mixed-land-use forest at the La Selva Biological Station and the lower flanks of Braulio Carrillo National Park. It contains a mosaic of historical and contemporary land use that is representative of contemporary tropical forest landscapes. Field studies demonstrated that LiDAR measurements were precise and accurate throughout the topographic and structural conditions at the site. Each record includes: easting and northing mapped coordinates (UTM Zone 16 North), height above ground (m), interpolated ground elevation (m), and six remote-sensing descriptors (point classification, the return number, the number of returns for the given pulse, intensity, scan angle, and the time of emission of the laser pulse). The data can be applied to a wide range of questions in basic and applied science, and are a valuable resource from a well-studied tropical rain forest for teaching and education. They also provide a quantitative baseline against which future conditions can be assessed.

D. Key words: biomass; carbon; Costa Rica; forest structure; landscape; La Selva Biological Station; LiDAR; light detection and ranging; Neotropics; remote sensing; sustainability.

CLASS II. RESEARCH ORIGIN DESCRIPTORS

A. Overall project description: Not applicable, this is a stand-alone project.

B. Specific subproject description

1. Site description: The study site is 128 km2 of Tropical Wet Forest in the Atlantic lowlands of Costa Rica (Figs. 1, 2, 3). Embedded within the study site is La Selva Biological station, which covers 16 km2. A detailed description of La Selva is in given in McDade et al. (1994).

a. Site type: The original site vegetation is classified as Tropical Wet Forest in the Holdridge life-zone system (Hartshorn and Hammel 1994).

b. Geography: Atlantic lowlands of Costa Rica (Figs. 2, 3). The southwest corner of the study area is at easting 820129 and northing 1141996 and the northeast corner is at easting 830629 and northing 1161996 in UTM zone 16 N (Sanford et al. 1994).

c. Habitat: The site has a mixed-land-use history, including cleared or developed areas, selectively-logged forest, terra-firme old-growth forest, current and abandoned plantations and pastures, second-growth forests of varying ages, and old-growth swamp forest (McDade et al. 1994).

d. Geology, landform: Relief is from 37 to 504 m (WGS 1984) on highly weathered Oxisols, and most of the landscape is on undulating upland plateaus (Kleber et al. 2007). Alluvial terraces near the Puerto Viejo and Sarapiquí rivers support higher nutrient availability through alluvial and colluvial deposition. In the rest of the landscape, nutrient availability is lower, although erosion can rejuvenate nutrient supply on deeply weathered soils (Porder et al. 2006).

e. Watersheds, hydrology: The entire study area is at the foot of Barva Volcano in Braulio Carrillo National Park and adjacent lowlands. All major watersheds originate outside the study area above the upper elevation boundary of the data set.

f. Site history: A detailed site history is given in McDade et al. (1994).

g. Climate: At La Selva mean annual temperature at 45 m elevation is 26 C, and annual rainfall is c. 4 m. There is a moderately drier season from February to May, but all months receive > 100 mm of precipitation (long-term station records at La Selva). Rainfall increases steeply with elevation, reaching c. 8 m at a 600 m elevation site (El Plástico) 2 km from the upper boundary of the data set area.

2. Experimental or sampling design

a. Design characteristics: The study area was selected to represent mixed-land-use lowland rain forest, including all 16 km2 of La Selva Biological Station and additional areas outside the La Selva property including plantations and areas within Braulio Carrillo National Park up to 504 m elevation.

b. Permanent plots: Not applicable.

c. Data collection: Remote sensing data were collected on 13–14 March 2006.

3. Research methods

a. Field/laboratory: Data were collected for 128 km2 of mixed-land-use tropical rain forest on 13–14 March 2006 (Kellner 2008, Kellner et al. 2009). The total number of discrete elevation estimates is 127,849,839 and mean sampling density was 1.99 observations/m2. In the configuration used here, the Leica ALS50 was capable of recording the locations of up to three reflecting surfaces within the returned laser signal for every emitted pulse. These correspond to three discrete elevation estimates for each laser footprint and are termed ‘returns’. The minimum number of returns for a given laser pulse is one. In forested terrain, this may correspond to a laser pulse that passes through the forest canopy and is reflected by the ground surface. The maximum number of returns is three, corresponding to three distinct reflecting surfaces within the canopy. Some pulses do not reach the ground surface, and may be reflected entirely by canopy vegetation. In our data, the variables return and nreturn distinguish the return number, and number of returns for each laser pulse. Each observation was classified as ‘ground’ (class = 2) or ‘vegetation’ (class = 5) using local topographic modeling and visual examination of the three-dimensional distribution of LiDAR observations (Cognocarta GIS LLC, San Jose, Costa Rica). Because LiDAR estimates the elevation of vegetation or ground surfaces relative to a reference ellipsoid (in this case, WGS 1984), quantifying height above ground requires interpolation of a ground surface and subtraction of ground from vegetation elevation (Fig. 1). We processed LiDAR data to generate a raster ground surface (digital terrain model, DTM) using natural neighbor interpolation of LiDAR elevation estimates that were classified as ground. Field studies demonstrate that LiDAR DTM estimates are precise and accurate within old-growth forest and the wider landscape at La Selva (see section III.A.4 and Figs., 4, 5). Canopy height can therefore be calculated by subtracting ground from elevation variables (but see V.G.4.A).

b. Instrumentation: Data were collected by the Leica ALS50 aboard a fixed-wing aircraft on 13–14 March 2006. The Leica ALS50 is a discrete-return scanning laser altimeter.

c. Taxonomy and systematics: Not applicable.

d. Permit history: Permits for study at the site from the Organization for Tropical Studies and the Costa Rican government were maintained over the life of the project.

e. Legal / organizational requirements: See II.B.3.d.

CLASS III. DATA SET STATUS AND ACCESSIBILITY

A. Status

1. Latest update: 12 February 2009

2. Latest archive date: 12 February 2009

3. Metadata status: The metadata are complete and up to date.

4. Data verification: Data quality has been assessed using field-validation studies and exploratory data analysis. We conducted visual and statistical examinations, and quantitatively assessed precision and accuracy in the field. Based on a visual examination, 38 points in the original LiDAR files were omitted from the current version. Each omitted point was an obvious outlier, occurring above an acceptable elevation range. We assessed the accuracy of LiDAR elevation estimates by comparing elevation values from the LiDAR DTM to 4184 ground-surveyed control points that were well distributed within the old-growth forest, and 8798 control points in the entire 16 km2 of La Selva Biological Station. At the location of each control point, elevation was measured in the field using optical leveling techniques. These values were subsequently corrected by adding 11.44 m to account for a known underestimation bias between field measurements and the WGS 1984 reference ellipsoid (Hofton et al. 2002, Clark et al. 2004). Within the 16 km2 landscape at La Selva the relationship between field and LiDAR estimates of ground elevation is: field measured elevation (m) = 0.999 ± (0.051) × LiDAR predicted elevation (m) - 1.289 ± (0.001), Pintercept < 0.001 Pslope < 0.001, r2 = 0.996, RMSE = 1.66 m, n = 8798 (Fig. 4). P values and standard errors for regression parameters (in parentheses) were estimated using a non-parametric bootstrap with 2,000 iterations. The non-parametric bootstrap is appropriate for spatial data, where the assumption of independence among observations is unlikely to hold (Clark 2007). This indicates that LiDAR elevation estimates were precise and accurate throughout the topographic and structural conditions at the site.

Several sources of error can influence LiDAR elevation estimates, including the diameter of the laser pulse relative to the size of openings in the forest canopy, laser power, and how the inbound laser pulse was processed to identify reflecting surfaces. Exogenous to the LiDAR system, topographic slope can introduce bias in LiDAR elevation estimates because steep terrain increases the elevation range over which a given laser pulse is reflected to the sensor (Clark et al. 2004). We reexamined the relationship between field and LiDAR estimates of ground elevation in old-growth forest using the subset of field measurements that were collected within old-growth forest. The relationship between field and LiDAR estimates of ground elevation in old-growth forest is: field measured elevation (m) = 0.999 ± (0.001) × LiDAR predicted elevation (m) - 1.406 ± (0.126) m, Pintercept < 0.001 Pslope < 0.001, r2 = 0.994, RMSE = 1.85 m, n = 4184 (Fig. 4). To determine whether there is slope-induced bias in LiDAR elevation estimates, we regressed the difference between each field and LiDAR estimate of ground elevation on topographic slope calculated using the DTM and standard methods. There is no evidence of slope-induced bias in LiDAR elevation estimates in old-growth forest: field measured elevation - LiDAR predicted elevation (m) = -0.004 ± (0.004) × slope (degrees) -1.381 ± (0.049) m, Pintercept < 0.001 Pslope = 0.146, r2 = 0.000, RMSE = 1.84 m, n = 4184 (Fig. 5). In the 16 km2 La Selva landscape, the relationship is: field measured elevation - LiDAR predicted elevation (m) = -0.004 ± (0.002) × slope (degrees) - 1.351 ± (0.031) m, Pintercept < 0.001 Pslope = 0.058, r2 = 0.000, RMSE = 1.66 m, n = 8798 (Fig. 5). Although the latter relationship is marginally significant, this is a function of the large sample size, as indicated by the small coefficient of determination and effect size, and we conclude that it is not biologically significant for most analyses. Additional field studies comparing forest structure and dynamics quantified using LiDAR and field measurements are in Kellner et al. (2009).

In the context of other studies, results presented here are comparable in precision and accuracy to other LiDAR data for the same site. Hofton et al. (2002) reported a 2.52 m overestimation bias for the Laser Vegetation Imaging Sensor, a large-footprint waveform LiDAR (RMSE = 5.64 m). And Clark et al. (2004) reported 0.97 m overestimation bias using the small-footprint FLI-MAP system (RMSE = 2.29 m). In old-growth forest < 10 °, overestimation bias was slightly greater (1.01 m) and RMSE was 1.95 m (Clark et al. 2004). In our data, overestimation bias was 1.29 m throughout the entire landscape, and 1.41 m in old-growth forest. Precision of the relationships (RMSE) was 1.66 and 1.85 m respectively. The offsets and precision reported here and in Clark et al. (2004) are based on comparison of DTM values to field measurements, rather than direct comparison based on laser pulses. Use of the DTM introduces additional assumptions and uncertainty. Nonetheless, the slope of the relationship between field measurements and DTM elevation is very close to 1.000 (Fig. 4), and the results presented here constrain the contribution of vertical and horizontal uncertainty and interpolation errors to a narrow range.

B. Accessibility

1. The data are available from the Ecological Society of America’s data archives. Duplicate copies of the data are being stored at La Selva Biological Station.

2. Contact persons: (1) James R. Kellner, 260 Panama Street, Stanford, California, 94305. Telephone: +1 808-933-8121. Fax: +1 808-933-8120. E-mail: jkellner@stanford.edu (2) David B. Clark, La Selva Biological Station, Puerto Viejo de Sarapiquí, Costa Rica. Telephone: + 506 2766-6565. Fax: +506 2766-6535  E-mail: dbclark@sloth.ots.ac.cr (3) Michelle A. Hofton, Department of Geography, University of Maryland, College Park, Maryland, 20742, USA. E-mail: mhofton@umd.edu

3. Copyright restrictions: None.

4. Proprietary restrictions: None, although we would like to hear how the data are being used (e.g., for what research questions or teaching exercises).

5. Costs: None

CLASS IV. DATA STRUCTURAL DESCRIPTORS

A. Data set files

1. Identity: AllTiles.zip -- The data are in 511 comma-delimited ASCII text files. Each file is a 500 × 500 m square plot. Plot locations are shown graphically in Fig. 3. To download selected data tiles, rather than the entire data set, please click here to go to a page listing all 511 files with links.

2. Size: File sizes are in Table 1.

3. Format and storage mode: Each file is comma-delimited ASCII text that was compressed using WinZip. The first line of each file contains variable names, subsequent lines contain data. In each file, the variables are: easting (UTM zone 16 North), northing (UTM zone 16 North), elevation (m), intensity, return number, number of returns, class, scan angle rank, GPS time. 

4. Header information: The first line of each file contains variable names.

B. Variable information: Variable names, definitions, units of measurement, data types, and ranges of numeric values are in Table 2. Values of the ground variable = -999 indicate that no interpolated value is available at the location of the LiDAR data pulse. This occurs around the perimeter of the study area where no interpolation is possible and affects 35,593 data records.

CLASS V. SUPPLEMENTAL DESCRIPTORS

A. Data acquisition

1. Data forms or acquisition methods: The data were collected using airborne remote sensing. Acquisition methods are described in section II.B.3.a.

2. Location of completed data forms: Not applicable.

3. Data entry and verification procedures: Quality control and field studies are described in section III.A.4.

B. Quality assurance / quality control procedures: Quality control and field validation studies are described in section III.A.4, Kellner (2008) and Kellner et al. (2009).

C. Related materials

1. GIS data layer: land use history for La Selva Biological Station (lu00.shp). Download LandUseHistoryShapefile.zip

2. GIS data layer: mapped locations of LiDAR data tiles (500_m_tiles.shp). Download 500mTilesShapefile.zip

D.  Computer programs and data processing algorithms

1. Generation of digital terrain model to estimate ground elevation

To estimate ground elevation and canopy height above ground, we generated a digital terrain model (DTM). The DTM was based on a natural neighbor interpolation applied to LiDAR elevation estimates that had been classified as ground returns (i.e., those laser pulses that were likely to have penetrated the canopy and been reflected by underlying topography). We then subtracted the predicted DTM elevation at the location of each laser pulse from its elevation estimate to quantify canopy height above ground. Field validation studies quantifying the precision and accuracy of ground topography and canopy height are in section III.A.4, Kellner (2008) and Kellner et al. (2009). Also see V.G.4.a.

2.  An R function to read the LiDAR data ( RFunction.txt). R is an open-source programming language and environment for statistical computation (R Development Core Team, 2009).

E.  Archiving

1. Archival procedures: The data are stored for long-term access in the GIS laboratory at La Selva Biological Station.

2. Redundant archival sites: Redundant copies of the data are stored by each author at their respective institutions.

F. Publications and results

1.  Kellner, J. R. 2008. Population and community dynamics of tropical rain forest canopy trees. The University of Georgia, Athens, Georgia, USA.

2.  Kellner J. R., D. B. Clark, and S. P. Hubbell. 2009. Pervasive canopy dynamics produce short-term stability in a tropical rain forest landscape. Ecology Letters 12:155–164.

3.  Kellner J. R., and G. P. Asner. 2009. Convergent structural responses of tropical forests to diverse disturbance regimes. Ecology Letters 12:887–897.

G. History of data set usage

         1. Data request history: Not applicable

         2. Data set update history: The data were last updated on February 12, 2009.

         3. Review history: The data were last reviewed on February 12, 2009.

         4. Questions and comments from secondary users:

a. Question number 1. I subtracted ground from elevation to estimate canopy height, but the number is negative. How is this possible?

Answer: Although vegetation height must be ≥ 0 m aboveground, it is possible for vegetation height estimates to be negative. This occurs when interpolation errors in the DTM are propagated through the analysis, or when assumptions of the DTM are violated. For example, the DTM assumes that ground elevation is constant within a given pixel. On steep terrain where this assumption could be seriously violated, some laser pulses may fall beneath the interpolated DTM value. In addition, when the number of laser pulses that reach the ground surface is locally sparse, the DTM can have poor precision or accuracy (for example, see outlying points in Fig. 4 as potential candidates). Elevation estimates for individual laser pulses (i.e., the elevation variable) are probably made with greater precision and accuracy than derived estimates of ground elevation (i.e., the ground variable). This is because ground estimates are based on classification of vegetation and ground returns (class variable = 5 or 2, respectively), and interpolation of a gridded surface for points with class = 2. Both procedures introduce assumptions and error.

b. Question number 2. The number of returns should vary from 1 to 3, but there are > 3 returns associated with the same value of the time variable.

Answer: Although the Leica ALS50 was capable of digitizing very small differences in time associated with laser ranges, the time variable is not sufficiently precise to distinguish between returns. Here is an example. The time variable is recorded with two decimal places of accuracy. This indicates that the smallest recordable difference in time (0.01 s) is associated with the distance traveled by a laser pulse in 0.01 s. Using the speed of light, 299,752,458 m · s-1, this distance is 2,997,524.58 m. Thus, the time variable is useful for determining when a given laser pulse was acquired (i.e., for distinguishing between flight lines, or dates of acquisition), but it is not useful for distinguishing between returns.

c. Question number 3. I have never used LiDAR data. What software do I need to get started?

Answer: In our experience, there are three types of tasks typically performed by ecologists that will dictate software requirements: (1) data processing to generate images, (2) visualization and basic summaries, and (3) statistical computation. Software needs vary widely depending on the objective of a study.

Data processing to interpolate images requires classification of LiDAR data points and interpolation of images from point subsets (e.g., creation of a DTM from points classified as ground). In practice, this could involve elimination of points over buildings, water or outside an acceptable elevation range using a combination of automated and manual editing, followed by interpolation using several algorithms (e.g., Clark et al. 2004). The current standard for LiDAR data processing is TerraScan (TerraSolid Ltd.). It can handle very large numbers of points and allows users to interact with and edit data on the fly. QuickTerrain Modeler (Applied Imagery Inc.) performs many of the same tasks as TerraScan and is excellent for rapid visualization of LiDAR point clouds and interpolated surfaces. Figure 1 was generated using QuickTerrain Modeler. Both pieces of software can perform basic statistical summaries of LiDAR data.

To visualize raster images made using LiDAR data we use ENVI (ITT Visual Information Solutions), which also contains sophisticated image processing routines that are accessible through IDL. For users who want to locate a field site or plots on a LiDAR elevation model, imagery can be visualized using ArcGIS.

For statistical computations, any programming environment that can handle large volumes of data will suffice. We use R, which is an open-source language and environment for statistical computation (R Development Core Team 2009).

ACKNOWLEDGEMENTS

Measurements in the CARBONO plots were partially supported by the Volcan Barva Transect TEAM Project of Conservation International, made possible with a grant from the Gordon and Betty Moore Foundation. Acquisition of LiDAR data was supported by contributions from the NSF (SGER 0533575 and 0223284), NASA, the University of Maryland, TEAM and the University of Alberta (Dr. Arturo Sanchez). This work was partially funded by a Doctoral Dissertation Improvement Grant from the NSF to JRK and Stephen P. Hubbell. We thank the Organization for Tropical Studies and the Center for Remote Sensing and Mapping Science, Department of Geography at The University of Georgia for logistic support and paraforesters Leonel Campos and William Miranda for field assistance. We thank Greg Asner and Ty Kennedy Bowdoin for discussions about LiDAR data that improved this manuscript.

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TABLE 1. Number of records, extent and size (uncompressed) of each data tile.

File number Number of records Minimum X Maximum X Minimum Y Maximum Y Size (kb)
1 5,084 827084.82 827128.99 1161496.01 1161599.16 275
2 129,539 827129.00 827628.98 1161496.00 1161879.26 7,058
3 42,107 827629.00 828051.30 1161496.00 1161696.49 2,330
4 65,880 826877.03 827128.99 1160996.00 1161495.98 3,581
5 193,855 827129.00 827628.99 1160996.00 1161495.99 10,625
6 255,501 827629.00 828128.99 1160996.00 1161495.99 13,951
7 140,520 828129.00 828628.99 1160996.00 1161459.23 7,703
8 43,151 828629.00 829065.57 1160996.00 1161221.53 2,365
9 194,886 826665.87 827128.99 1160496.00 1160995.99 10,616
10 172,094 827129.00 827628.99 1160496.00 1160995.99 9,387
11 233,663 827629.00 828128.99 1160496.00 1160995.99 12,675
12 204,618 828129.00 828628.99 1160496.00 1160995.99 11,313
13 126,609 828629.01 829057.10 1160496.00 1160995.99 6,875
14 57,832 826455.95 826628.99 1159996.00 1160407.95 3,170
15 229,686 826629.00 827128.99 1159996.00 1160495.99 12,489
16 231,380 827129.00 827628.99 1159996.00 1160495.99 12,604
17 233,122 827629.00 828128.99 1159996.00 1160495.99 12,727
18 219,306 828129.00 828628.99 1159996.00 1160495.99 12,037
19 37,088 828629.00 828825.37 1160076.54 1160495.99 2,005
20 158,394 826246.76 826628.99 1159496.00 1159995.99 8,615
21 232,855 826629.00 827128.99 1159496.00 1159995.99 12,855
22 215,291 827129.00 827628.99 1159496.00 1159995.99 11,681
23 215,369 827629.00 828128.99 1159496.00 1159995.99 11,829
24 148,086 828129.00 828591.86 1159496.00 1159995.99 8,026
25 13,104 826079.59 826128.99 1158996.00 1159146.79 741
26 294,246 826129.00 826628.99 1158996.00 1159495.99 16,269
27 243,537 826629.00 827128.99 1158996.00 1159495.99 13,399
28 199,401 827129.00 827628.99 1158996.00 1159495.99 10,799
29 248,732 827629.00 828128.99 1158996.00 1159495.99 13,688
30 117,837 828129.00 828628.99 1158996.00 1159495.99 6,388
31 13,964 828629.00 828779.62 1158996.00 1159130.61 774
32 177,859 825885.11 826128.99 1158496.00 1158995.97 10,062
33 274,607 826129.00 826628.99 1158496.00 1158995.99 15,517
34 308,569 826629.00 827128.99 1158496.00 1158995.99 17,094
35 232,840 827129.00 827628.99 1158496.00 1158995.99 12,712
36 256,335 827629.00 828128.99 1158496.00 1158995.99 13,968
37 203,729 828129.00 828628.99 1158496.00 1158995.99 11,156
38 70,319 828629.00 828814.83 1158496.00 1158995.99 3,902
39 379,055 825637.63 826128.99 1157996.00 1158495.99 12,346
40 314,154 826129.00 826628.99 1157996.00 1158495.99 17,902
41 273,315 826629.00 827128.99 1157996.00 1158495.99 15,013
42 267,684 827129.00 827628.99 1157996.00 1158495.99 14,868
43 300,293 827629.00 828128.99 1157996.00 1158495.99 16,309
44 221,663 828129.00 828628.99 1157996.00 1158495.99 12,252
45 2,143 828629.00 828646.08 1158453.35 1158495.99 120
46 111,375 825408.00 825628.99 1157496.00 1157977.47 6,116
47 370,712 825629.00 826128.99 1157496.00 1157995.99 20,671
48 257,686 826129.00 826628.99 1157496.00 1157995.99 14,304
49 219,543 826629.00 827128.99 1157496.00 1157995.99 12,014
50 282,557 827129.00 827628.99 1157496.00 1157995.99 15,489
51 312,430 827629.00 828128.99 1157496.00 1157995.99 16,937
52 135,589 828129.00 828435.12 1157496.00 1157995.99 7,535
53 12,280 824362.91 824601.09 1156996.00 1157105.71 685
54 2,714 824853.96 824957.20 1156996.01 1157039.84 151
55 187,907 825198.46 825628.99 1156996.00 1157495.99 10,215
56 267,002 825629.00 826128.99 1156996.00 1157495.99 14,736
57 210,808 826129.00 826628.99 1156996.00 1157495.99 11,468
58 198,242 826629.00 827128.99 1156996.00 1157495.99 10,884
59 231,020 827129.00 827628.99 1156996.00 1157495.99 12,513
60 240,026 827629.00 828128.99 1156996.00 1157495.99 13,146
61 128,659 828129.00 828628.99 1156996.00 1157495.99 6,991
62 18,503 828629.00 828922.17 1156996.00 1157137.50 1,005
63 159,748 824154.15 824628.99 1156496.00 1156995.99 8,764
64 203,239 824629.00 825128.99 1156496.00 1156995.99 11,062
65 196,060 825129.01 825628.99 1156496.00 1156995.99 10,667
66 247,072 825629.00 826128.99 1156496.00 1156995.99 13,503
67 218,930 826129.00 826628.99 1156496.00 1156995.99 11,879
68 216,421 826629.00 827128.99 1156496.00 1156995.99 11,854
69 216,165 827129.00 827628.99 1156496.00 1156995.99 11,719
70 258,400 827629.00 828128.99 1156496.00 1156995.99 14,085
71 224,050 828129.01 828628.99 1156496.00 1156995.99 12,356
72 204,528 828629.00 829128.99 1156496.00 1156995.99 11,095
73 125,295 829129.00 829628.98 1156496.00 1156897.63 6,905
74 25,534 829629.02 829967.76 1156496.00 1156658.36 1,395
75 41,081 823945.15 824128.99 1155996.00 1156436.28 2,254
76 234,772 824129.00 824628.99 1155996.00 1156495.99 12,811
77 224,231 824629.00 825128.99 1155996.00 1156495.99 12,190
78 228,873 825129.00 825628.99 1155996.00 1156495.99 12,547
79 305,432 825629.00 826128.99 1155996.00 1156495.99 16,560
80 223,424 826129.00 826628.99 1155996.00 1156495.99 12,196
81 247,934 826629.00 827128.99 1155996.00 1156495.99 13,456
82 211,389 827129.00 827628.99 1155996.00 1156495.99 11,532
83 271,024 827629.00 828128.99 1155996.00 1156495.99 14,867
84 253,411 828129.00 828628.99 1155996.00 1156495.99 13,803
85 267,681 828629.00 829128.99 1155996.00 1156495.99 14,585
86 303,146 829129.00 829628.99 1155996.00 1156495.99 16,466
87 335,279 829629.00 830128.99 1155996.00 1156495.99 18,298
88 44,792 830129.00 830260.32 1156056.95 1156419.07 2,485
89 179,499 823737.27 824128.99 1155496.00 1155995.99 9,830
90 264,252 824129.00 824628.99 1155496.00 1155995.99 14,476
91 234,332 824629.00 825128.99 1155496.00 1155995.99 12,689
92 256,932 825129.00 825628.99 1155496.00 1155995.99 14,107
93 343,415 825629.00 826128.99 1155496.00 1155995.99 18,565
94 251,956 826129.00 826628.99 1155496.00 1155995.99 13,845
95 283,590 826629.00 827128.99 1155496.00 1155995.99 15,293
96 287,815 827129.00 827628.99 1155496.00 1155995.99 15,700
97 312,076 827629.00 828128.99 1155496.00 1155995.99 17,255
98 270,484 828129.00 828628.99 1155496.00 1155995.99 14,607
99 289,739 828629.00 829128.99 1155496.00 1155995.99 15,920
100 332,846 829129.00 829628.99 1155496.00 1155995.99 17,978
101 357,498 829629.00 830102.35 1155496.00 1155995.99 19,641
102 11,365 823528.67 823628.99 1154996.00 1155236.76 628
103 246,199 823629.00 824128.99 1154996.00 1155495.99 13,410
104 206,120 824129.00 824628.99 1154996.00 1155495.99 11,226
105 195,604 824629.00 825128.99 1154996.00 1155495.99 10,648
106 208,344 825129.00 825628.99 1154996.00 1155495.99 11,349
107 279,107 825629.00 826128.99 1154996.00 1155495.99 15,181
108 268,917 826129.00 826628.99 1154996.00 1155495.99 14,589
109 284,807 826629.00 827128.99 1154996.00 1155495.99 15,448
110 267,161 827129.00 827628.99 1154996.00 1155495.99 14,511
111 303,554 827629.01 828128.99 1154996.00 1155495.99 16,510
112 227,348 828129.00 828628.99 1154996.00 1155495.99 12,335
113 229,951 828629.00 829128.99 1154996.00 1155495.99 12,588
114 235,281 829129.00 829628.99 1154996.00 1155495.99 12,789
115 119,369 829629.00 829884.19 1154996.00 1155495.99 6,606
116 93,796 823319.87 823628.99 1154496.00 1154995.99 5,218
117 289,649 823629.00 824128.99 1154496.00 1154995.99 15,893
118 211,573 824129.00 824628.99 1154496.00 1154995.99 11,449
119 251,207 824629.00 825128.99 1154496.00 1154995.99 13,877
120 221,759 825129.00 825628.99 1154496.00 1154995.99 11,951
121 290,625 825629.00 826128.99 1154496.00 1154995.99 15,983
122 245,895 826129.00 826628.99 1154496.00 1154995.99 13,303
123 257,522 826629.00 827128.99 1154496.00 1154995.99 14,019
124 275,317 827129.00 827628.99 1154496.00 1154995.99 15,028
125 310,604 827629.00 828128.99 1154496.00 1154995.99 16,782
126 218,617 828129.00 828628.99 1154496.00 1154995.99 12,018
127 221,636 828629.00 829128.99 1154496.00 1154995.99 12,000
128 249,338 829129.00 829628.97 1154496.00 1154995.99 13,824
129 5,636 829629.00 829665.13 1154914.10 1154995.99 314
130 514 823112.17 823128.99 1153996.13 1154036.88 29
131 174,377 823129.06 823628.99 1153996.00 1154495.99 9,546
132 326,989 823629.00 824128.99 1153996.00 1154495.99 17,990
133 252,836 824129.00 824628.99 1153996.00 1154495.99 13,794
134 277,488 824629.00 825128.99 1153996.00 1154495.99 15,374
135 268,715 825129.00 825628.99 1153996.00 1154495.99 14,822
136 330,164 825629.00 826128.99 1153996.00 1154495.99 18,065
137 285,356 826129.00 826628.99 1153996.00 1154495.99 15,493
138 263,454 826629.00 827128.99 1153996.00 1154495.99 14,361
139 274,657 827129.00 827628.99 1153996.00 1154495.99 15,048
140 306,022 827629.01 828128.99 1153996.00 1154495.99 16,531
141 235,986 828129.00 828628.99 1153996.00 1154495.99 12,909
142 228,847 828629.00 829128.99 1153996.00 1154495.99 12,455
143 139,625 829129.00 829446.29 1153996.00 1154495.99 7,742
144 67,680 822902.88 823128.99 1153496.00 1153995.96 3,784
145 267,842 823129.00 823628.99 1153496.00 1153995.99 14,666
146 353,520 823629.00 824128.99 1153496.00 1153995.99 19,344
147 308,347 824129.00 824628.99 1153496.00 1153995.99 17,261
148 295,574 824629.00 825128.99 1153496.00 1153995.99 16,473
149 291,589 825129.00 825628.99 1153496.00 1153995.99 16,311
150 345,889 825629.00 826128.99 1153496.00 1153995.99 19,048
151 296,424 826129.00 826628.99 1153496.00 1153995.99 16,319
152 284,986 826629.00 827128.99 1153496.00 1153995.99 15,652
153 299,533 827129.00 827628.99 1153496.00 1153995.99 16,246
154 373,983 827629.00 828128.99 1153496.00 1153995.99 20,463
155 298,551 828129.00 828628.99 1153496.00 1153995.99 16,228
156 236,308 828629.00 829128.99 1153496.00 1153995.98 12,976
157 17,152 829129.00 829227.27 1153772.33 1153995.99 954
158 148,706 822694.16 823128.99 1152996.00 1153495.99 8,280
159 274,948 823129.00 823628.99 1152996.00 1153495.99 15,062
160 340,342 823629.00 824128.99 1152996.00 1153495.99 19,034
161 253,796 824129.00 824628.99 1152996.00 1153495.99 14,416
162 241,388 824629.00 825128.99 1152996.00 1153495.99 13,450
163 265,554 825129.00 825628.99 1152996.00 1153495.99 14,905
164 317,647 825629.00 826128.99 1152996.00 1153495.99 17,720
165 258,353 826129.00 826628.99 1152996.00 1153495.99 14,200
166 267,484 826629.00 827128.99 1152996.00 1153495.99 14,667
167 253,391 827129.00 827628.99 1152996.00 1153495.99 13,728
168 327,826 827629.00 828128.99 1152996.00 1153495.99 17,904
169 238,657 828129.00 828628.99 1152996.00 1153495.99 12,919
170 220,141 828629.00 829128.99 1152996.00 1153495.99 12,146
171 134,558 829129.00 829628.98 1152996.00 1153430.63 7,299
172 41,169 829629.00 829930.22 1152996.00 1153188.29 2,282
173 22,742 822486.29 822628.99 1152496.00 1152837.42 1,286
174 228,688 822629.00 823128.99 1152496.00 1152995.99 12,700
175 248,079 823129.00 823628.99 1152496.00 1152995.99 13,655
176 347,426 823629.00 824128.99 1152496.00 1152995.99 19,637
177 225,793 824129.00 824628.99 1152496.00 1152995.99 12,702
178 253,176 824629.00 825128.99 1152496.00 1152995.99 14,279
179 253,009 825129.00 825628.99 1152496.00 1152995.99 14,235
180 298,537 825629.00 826128.99 1152496.00 1152995.99 16,837
181 254,052 826129.00 826628.99 1152496.00 1152995.99 14,304
182 277,433 826629.00 827128.99 1152496.00 1152995.99 15,316
183 243,919 827129.00 827628.99 1152496.00 1152995.99 13,418
184 331,636 827629.00 828128.99 1152496.00 1152995.99 18,045
185 235,289 828129.00 828628.99 1152496.00 1152995.99 12,833
186 225,627 828629.00 829128.99 1152496.00 1152995.99 12,312
187 218,892 829129.00 829628.99 1152496.00 1152995.99 11,900
188 139,122 829629.00 829911.49 1152496.00 1152995.97 7,727
189 116,637 822277.82 822628.99 1151996.00 1152495.99 6,628
190 235,854 822629.00 823128.99 1151996.00 1152495.99 13,228
191 275,523 823129.00 823628.99 1151996.00 1152495.99 15,282
192 346,491 823629.00 824128.99 1151996.00 1152495.99 19,718
193 261,289 824129.00 824628.99 1151996.00 1152495.99 14,604
194 279,160 824629.00 825128.99 1151996.00 1152495.99 15,877
195 275,002 825129.00 825628.99 1151996.00 1152495.99 15,372
196 337,270 825629.00 826128.99 1151996.00 1152495.99 19,046
197 266,626 826129.00 826628.99 1151996.00 1152495.99 15,082
198 251,857 826629.00 827128.99 1151996.00 1152495.99 13,948
199 271,537 827129.00 827628.99 1151996.00 1152495.99 15,216
200 334,099 827629.00 828128.99 1151996.00 1152495.99 18,203
201 249,299 828129.00 828628.99 1151996.00 1152495.99 13,693
202 263,391 828629.00 829128.99 1151996.00 1152495.99 14,277
203 249,085 829129.00 829628.99 1151996.00 1152495.99 13,778
204 16,227 829629.00 829701.01 1152327.10 1152495.99 903
205 3,086 822068.94 822128.99 1151496.00 1151638.91 179
206 185,649 822129.00 822628.99 1151496.00 1151995.99 10,544
207 244,727 822629.00 823128.99 1151496.00 1151995.99 13,783
208 309,959 823129.00 823628.99 1151496.00 1151995.99 17,394
209 334,714 823629.00 824128.99 1151496.00 1151995.99 18,882
210 319,237 824129.00 824628.99 1151496.00 1151995.99 17,941
211 309,301 824629.00 825128.99 1151496.00 1151995.99 17,488
212 301,957 825129.00 825628.99 1151496.00 1151995.99 16,951
213 356,373 825629.00 826128.99 1151496.00 1151995.99 20,050
214 302,295 826129.00 826628.99 1151496.00 1151995.99 17,074
215 287,684 826629.00 827128.99 1151496.00 1151995.99 16,116
216 303,107 827129.00 827628.99 1151496.00 1151995.99 16,929
217 345,409 827629.00 828128.99 1151496.00 1151995.99 18,953
218 253,389 828129.00 828628.99 1151496.00 1151995.99 13,947
219 263,686 828629.00 829128.99 1151496.00 1151995.99 14,311
220 160,750 829129.00 829488.92 1151496.00 1151995.99 8,926
221 72,431 821860.19 822128.99 1150996.00 1151495.98 4,148
222 191,210 822129.00 822628.99 1150996.00 1151495.99 10,788
223 244,963 822629.00 823128.99 1150996.00 1151495.99 13,729
224 269,008 823129.00 823628.99 1150996.00 1151495.99 15,327
225 289,631 823629.00 824128.99 1150996.00 1151495.99 16,209
226 272,262 824129.00 824628.99 1150996.00 1151495.99 15,458
227 257,899 824629.00 825128.99 1150996.00 1151495.99 14,431
228 264,145 825129.00 825628.99 1150996.00 1151495.99 14,866
229 314,817 825629.00 826128.99 1150996.00 1151495.99 17,855
230 249,854 826129.00 826628.99 1150996.00 1151495.99 13,983
231 268,788 826629.00 827128.99 1150996.00 1151495.99 15,243
232 278,033 827129.00 827628.99 1150996.00 1151495.99 15,550
233 282,091 827629.00 828128.99 1150996.00 1151495.99 15,658
234 238,563 828129.00 828628.99 1150996.00 1151495.99 12,983
235 250,603 828629.00 829128.99 1150996.00 1151495.99 13,743
236 47,258 829129.00 829277.47 1151145.28 1151495.99 2,634
237 188,376 821652.56 822128.99 1150496.00 1150995.99 10,680
238 220,774 822129.00 822628.99 1150496.00 1150995.99 12,462
239 248,621 822629.00 823128.99 1150496.00 1150995.99 13,965
240 246,670 823129.00 823628.99 1150496.00 1150995.99 13,944
241 328,402 823629.00 824128.99 1150496.00 1150995.99 18,432
242 277,704 824129.00 824628.99 1150496.00 1150995.99 15,760
243 262,197 824629.00 825128.99 1150496.00 1150995.99 14,674
244 281,305 825129.01 825628.99 1150496.00 1150995.99 15,872
245 320,801 825629.00 826128.99 1150496.00 1150995.99 18,187
246 266,263 826129.00 826628.99 1150496.00 1150995.99 14,883
247 251,730 826629.00 827128.99 1150496.00 1150995.99 14,271
248 244,145 827129.00 827628.99 1150496.00 1150995.99 13,486
249 293,662 827629.00 828128.99 1150496.00 1150995.99 16,332
250 257,300 828129.00 828628.99 1150496.00 1150995.99 14,080
251 207,404 828629.00 829128.99 1150496.00 1150995.99 11,481
252 20,447 829129.00 829427.15 1150496.00 1150631.35 1,113
253 43,645 821444.17 821628.99 1149996.00 1150440.27 2,496
254 280,569 821629.00 822128.99 1149996.00 1150495.99 15,770
255 251,677 822129.00 822628.99 1149996.00 1150495.99 14,141
256 280,964 822629.00 823128.99 1149996.00 1150495.99 15,890
257 274,068 823129.00 823628.99 1149996.00 1150495.99 15,359
258 386,245 823629.00 824128.99 1149996.00 1150495.99 21,895
259 292,153 824129.00 824628.99 1149996.00 1150495.99 16,406
260 281,857 824629.00 825128.99 1149996.00 1150495.99 15,822
261 269,265 825129.00 825628.99 1149996.00 1150495.98 15,227
262 307,658 825629.00 826128.99 1149996.00 1150495.99 17,297
263 273,100 826129.00 826628.99 1149996.00 1150495.99 15,248
264 308,420 826629.00 827128.99 1149996.00 1150495.99 17,102
265 261,140 827129.00 827628.99 1149996.00 1150495.99 14,659
266 358,043 827629.00 828128.99 1149996.00 1150495.99 19,982
267 277,962 828129.00 828628.99 1149996.00 1150495.99 15,414
268 267,499 828629.00 829128.99 1149996.00 1150495.99 14,744
269 229,043 829129.00 829628.99 1149996.00 1150495.99 12,517
270 18,286 829629.01 830128.99 1149996.00 1150404.58 1,007
271 37,783 830129.01 830528.19 1149996.00 1150178.11 2,056
272 41,088 821394.89 821628.99 1149787.74 1149995.99 2,344
273 219,870 821629.00 822128.99 1149593.95 1149995.99 12,416
274 307,951 822129.00 822628.99 1149496.00 1149995.99 17,196
275 304,970 822629.00 823128.99 1149496.00 1149995.99 17,298
276 312,008 823129.00 823628.99 1149496.00 1149995.99 17,463
277 392,677 823629.00 824128.99 1149496.00 1149995.99 22,282
278 330,011 824129.00 824628.99 1149496.00 1149995.99 18,436
279 308,560 824629.00 825128.99 1149496.00 1149995.99 17,377
280 284,733 825129.00 825628.99 1149496.00 1149995.99 16,173
281 333,968 825629.00 826128.99 1149496.00 1149995.99 18,545
282 312,826 826129.00 826628.99 1149496.00 1149995.99 17,721
283 305,983 826629.00 827128.99 1149496.00 1149995.99 17,083
284 301,183 827129.00 827628.99 1149496.00 1149995.99 17,086
285 374,435 827629.00 828128.99 1149496.00 1149995.99 20,831
286 303,958 828129.00 828628.99 1149496.00 1149995.99 17,016
287 297,114 828629.00 829128.99 1149496.00 1149995.99 16,451
288 256,337 829129.00 829628.99 1149496.00 1149995.99 14,149
289 222,672 829629.00 830128.99 1149496.00 1149995.98 12,089
290 133,563 830129.00 830527.84 1149496.00 1149995.99 7,282
291 2,935 822082.84 822128.98 1148996.01 1149096.41 164
292 240,959 822129.02 822628.99 1148996.00 1149495.99 13,588
293 221,801 822629.00 823128.99 1148996.00 1149495.99 12,505
294 291,217 823129.00 823628.99 1148996.00 1149495.99 16,392
295 323,554 823629.00 824128.99 1148996.00 1149495.99 18,236
296 271,446 824129.00 824628.99 1148996.00 1149495.99 15,285
297 257,994 824629.00 825128.99 1148996.00 1149495.99 14,531
298 258,642 825129.00 825628.99 1148996.00 1149495.99 14,521
299 295,841 825629.00 826128.99 1148996.00 1149495.99 16,569
300 246,159 826129.00 826628.99 1148996.00 1149495.99 13,902
301 253,171 826629.00 827128.99 1148996.00 1149495.99 14,257
302 258,117 827129.00 827628.99 1148996.00 1149495.99 14,582
303 314,854 827629.00 828128.99 1148996.00 1149495.99 17,654
304 244,344 828129.00 828628.99 1148996.00 1149495.99 13,773
305 260,453 828629.00 829128.99 1148996.00 1149495.99 14,348
306 245,821 829129.00 829628.99 1148996.00 1149495.99 13,586
307 247,554 829629.00 830128.99 1148996.00 1149495.99 13,447
308 38,639 830129.00 830314.43 1149060.35 1149495.98 2,141
309 115,194 821857.11 822128.99 1148496.00 1148995.98 6,451
310 291,554 822129.00 822628.99 1148496.00 1148995.99 16,603
311 246,337 822629.00 823128.99 1148496.00 1148995.99 13,760
312 273,553 823129.00 823628.99 1148496.00 1148995.99 15,447
313 322,949 823629.00 824128.99 1148496.00 1148995.99 18,071
314 271,654 824129.00 824628.99 1148496.00 1148995.99 15,295
315 258,780 824629.00 825128.99 1148496.00 1148995.99 14,663
316 244,583 825129.00 825628.99 1148496.00 1148995.99 13,667
317 296,254 825629.00 826128.99 1148496.00 1148995.99 16,834
318 236,084 826129.00 826628.99 1148496.00 1148995.99 13,228
319 249,278 826629.00 827128.99 1148496.00 1148995.99 14,151
320 266,700 827129.00 827628.99 1148496.00 1148995.99 14,957
321 324,070 827629.00 828128.99 1148496.00 1148995.99 18,410
322 241,810 828129.00 828628.99 1148496.00 1148995.99 13,455
323 257,406 828629.00 829128.99 1148496.00 1148995.99 14,299
324 265,376 829129.00 829628.99 1148496.00 1148995.99 14,516
325 195,165 829629.00 830101.61 1148496.00 1148995.99 10,658
326 286,451 821631.09 822128.99 1147996.00 1148495.99 16,131
327 270,947 822129.00 822628.99 1147996.00 1148495.99 15,316
328 283,025 822629.00 823128.99 1147996.00 1148495.99 15,868
329 269,036 823129.00 823628.99 1147996.00 1148495.99 15,273
330 365,050 823629.00 824128.99 1147996.00 1148495.99 20,406
331 276,437 824129.00 824628.99 1147996.00 1148495.99 15,583
332 261,792 824629.00 825128.99 1147996.00 1148495.99 14,810
333 262,732 825129.00 825628.99 1147996.00 1148495.99 14,708
334 306,499 825629.00 826128.99 1147996.00 1148495.99 17,305
335 263,106 826129.00 826628.99 1147996.00 1148495.99 14,760
336 264,960 826629.00 827128.99 1147996.00 1148495.99 15,014
337 288,700 827129.00 827628.99 1147996.00 1148495.99 16,196
338 309,741 827629.00 828128.99 1147996.00 1148495.98 17,569
339 264,093 828129.00 828628.99 1147996.00 1148495.99 14,686
340 270,222 828629.00 829128.99 1147996.00 1148495.99 15,018
341 269,879 829129.00 829628.99 1147996.00 1148495.99 14,748
342 74,798 829629.00 829888.62 1147996.01 1148495.99 4,206
343 94,491 821406.00 821628.99 1147496.00 1147991.53 5,315
344 472,956 821629.00 822128.99 1147496.00 1147995.99 26,763
345 302,175 822129.00 822628.99 1147496.00 1147995.99 16,911
346 316,836 822629.00 823128.99 1147496.00 1147995.99 17,962
347 306,540 823129.00 823628.99 1147496.00 1147995.99 17,186
348 399,200 823629.00 824128.99 1147496.00 1147995.99 22,475
349 300,832 824129.00 824628.99 1147496.00 1147995.99 16,984
350 263,865 824629.00 825128.99 1147496.00 1147995.99 14,818
351 298,538 825129.00 825628.99 1147496.00 1147995.99 16,842
352 303,207 825629.00 826128.99 1147496.00 1147995.99 17,018
353 282,869 826129.00 826628.99 1147496.00 1147995.99 15,990
354 268,610 826629.00 827128.99 1147496.00 1147995.99 15,112
355 310,963 827129.00 827628.99 1147496.00 1147995.99 17,567
356 302,978 827629.00 828128.99 1147496.00 1147995.99 16,999
357 296,948 828129.00 828628.99 1147496.00 1147995.99 16,602
358 320,803 828629.00 829128.99 1147496.00 1147995.99 17,891
359 250,622 829129.00 829628.99 1147496.00 1147995.99 13,956
360 5,462 829629.00 829675.26 1147890.27 1147995.99 308
361 224,496 821179.67 821628.99 1146996.00 1147495.99 12,624
362 366,995 821629.00 822128.99 1146996.00 1147495.99 20,778
363 292,447 822129.00 822628.99 1146996.00 1147495.99 16,343
364 292,224 822629.00 823128.99 1146996.00 1147495.99 16,553
365 301,134 823129.00 823628.99 1146996.00 1147495.99 16,845
366 343,500 823629.00 824128.99 1146996.00 1147495.99 19,340
367 255,573 824129.00 824628.99 1146996.00 1147495.99 14,541
368 242,940 824629.00 825128.99 1146996.00 1147495.99 13,548
369 251,362 825129.00 825628.99 1146996.00 1147495.99 14,327
370 300,785 825629.00 826128.99 1146996.00 1147495.99 16,819
371 269,177 826129.00 826628.99 1146996.00 1147495.99 15,319
372 257,028 826629.00 827128.99 1146996.00 1147495.99 14,358
373 238,696 827129.00 827628.99 1146996.00 1147495.99 13,568
374 292,645 827629.00 828128.99 1146996.00 1147495.99 16,342
375 252,483 828129.00 828628.99 1146996.00 1147495.99 14,367
376 235,485 828629.00 829128.99 1146996.00 1147495.99 13,200
377 110,626 829129.00 829462.23 1146996.00 1147495.99 6,199
378 53,457 820953.93 821128.99 1146496.00 1146884.33 3,010
379 322,334 821129.00 821628.99 1146496.00 1146995.99 18,304
380 326,290 821629.00 822128.99 1146496.00 1146995.99 18,288
381 300,856 822129.00 822628.99 1146496.00 1146995.99 16,972
382 264,172 822629.00 823128.99 1146496.00 1146995.99 14,875
383 299,926 823129.00 823628.99 1146496.00 1146995.99 16,907
384 322,504 823629.00 824128.99 1146496.00 1146995.99 18,178
385 252,804 824129.00 824628.99 1146496.00 1146995.99 14,241
386 258,608 824629.00 825128.99 1146496.00 1146995.99 14,552
387 240,123 825129.00 825628.99 1146496.00 1146995.99 13,558
388 309,748 825629.00 826128.99 1146496.00 1146995.99 17,430
389 237,053 826129.00 826628.99 1146496.00 1146995.99 13,392
390 287,755 826629.00 827128.99 1146496.00 1146995.99 16,190
391 253,206 827129.00 827628.99 1146496.00 1146995.99 14,254
392 336,594 827629.00 828128.99 1146496.00 1146995.99 18,953
393 269,078 828129.00 828628.99 1146496.00 1146995.99 15,226
394 241,611 828629.00 829128.99 1146496.00 1146995.99 13,543
395 16,170 829129.00 829249.12 1146713.93 1146995.99 924
396 257,165 820727.85 821128.99 1145996.00 1146495.99 14,460
397 302,972 821129.00 821628.99 1145996.00 1146495.99 17,195
398 410,364 821629.00 822128.99 1145996.00 1146495.99 23,002
399 351,300 822129.00 822628.99 1145996.00 1146495.99 19,873
400 312,880 822629.00 823128.99 1145996.00 1146495.99 17,483
401 301,934 823129.00 823628.99 1145996.00 1146495.99 17,013
402 327,397 823629.00 824128.99 1145996.00 1146495.99 18,513
403 303,940 824129.00 824628.99 1145996.00 1146495.99 17,006
404 279,680 824629.01 825128.99 1145996.00 1146495.99 15,880
405 246,024 825129.00 825628.99 1145996.00 1146495.99 13,750
406 329,286 825629.00 826128.99 1145996.00 1146495.99 18,758
407 269,922 826129.00 826628.99 1145996.00 1146495.99 15,112
408 276,550 826629.00 827128.99 1145996.00 1146495.99 15,656
409 267,970 827129.00 827628.99 1145996.00 1146495.99 15,014
410 346,329 827629.00 828128.99 1145996.00 1146495.99 19,678
411 285,184 828129.00 828628.99 1145996.00 1146495.99 16,015
412 115,274 828629.00 829036.30 1146120.44 1146495.99 6,487
413 63,349 820506.73 820628.99 1145496.00 1145777.09 3,581
414 451,778 820629.00 821128.99 1145496.00 1145995.99 25,442
415 352,211 821129.00 821628.99 1145496.00 1145995.99 19,858
416 480,895 821629.00 822128.99 1145496.00 1145995.99 26,966
417 358,849 822129.00 822628.99 1145496.00 1145995.99 20,181
418 354,254 822629.00 823128.99 1145496.00 1145995.99 19,877
419 323,860 823129.00 823628.99 1145496.00 1145995.99 18,227
420 366,429 823629.00 824128.99 1145496.00 1145995.99 20,751
421 305,125 824129.00 824628.99 1145496.00 1145995.99 17,127
422 301,395 824629.00 825128.99 1145496.00 1145995.99 17,023
423 304,153 825129.00 825628.99 1145496.00 1145995.99 17,035
424 374,214 825629.00 826128.99 1145496.00 1145995.99 21,143
425 328,248 826129.00 826628.99 1145496.00 1145995.99 18,411
426 288,774 826629.00 827128.99 1145496.00 1145995.99 16,462
427 305,846 827129.00 827628.99 1145496.00 1145995.99 17,121
428 380,855 827629.00 828128.99 1145496.00 1145995.99 21,615
429 253,073 828129.00 828545.71 1145496.00 1145995.99 14,262
430 241,378 820308.57 820628.99 1144996.00 1145495.99 13,621
431 329,752 820629.00 821128.99 1144996.00 1145495.99 18,701
432 271,755 821129.00 821628.99 1144996.00 1145495.99 15,215
433 433,121 821629.00 822128.99 1144996.00 1145495.99 24,522
434 318,258 822129.00 822628.99 1144996.00 1145495.99 17,812
435 322,682 822629.00 823128.99 1144996.00 1145495.99 18,146
436 304,292 823129.00 823628.99 1144996.00 1145495.99 17,265
437 380,955 823629.00 824128.99 1144996.00 1145495.99 21,353
438 290,543 824129.00 824628.99 1144996.00 1145495.99 16,390
439 286,683 824629.00 825128.99 1144996.00 1145495.99 16,067
440 275,645 825129.00 825628.99 1144996.00 1145495.99 15,605
441 293,843 825629.00 826128.99 1144996.00 1145495.99 16,496
442 288,433 826129.00 826628.99 1144996.00 1145495.99 16,313
443 230,062 826629.00 827128.99 1144996.00 1145495.99 12,927
444 295,499 827129.00 827628.99 1144996.00 1145495.99 16,734
445 366,106 827629.00 828128.99 1144996.00 1145495.99 20,565
446 142,932 828129.00 828342.87 1144996.00 1145495.99 8,082
447 299,590 820129.90 820628.99 1144496.00 1144995.99 16,961
448 290,074 820629.00 821128.99 1144496.00 1144995.99 16,454
449 319,797 821129.00 821628.99 1144496.00 1144995.99 17,902
450 381,981 821629.00 822128.99 1144496.00 1144995.99 21,650
451 330,772 822129.00 822628.99 1144496.00 1144995.99 18,496
452 314,459 822629.00 823128.99 1144496.00 1144995.99 17,709
453 296,579 823129.00 823628.99 1144496.00 1144995.99 16,866
454 368,691 823629.00 824128.99 1144496.00 1144995.99 20,634
455 275,548 824129.00 824628.99 1144496.00 1144995.99 15,647
456 275,469 824629.00 825128.99 1144496.00 1144995.99 15,398
457 277,572 825129.00 825628.99 1144496.00 1144995.99 15,818
458 364,877 825629.00 826128.99 1144496.00 1144995.99 20,394
459 301,389 826129.00 826628.99 1144496.00 1144995.99 17,104
460 307,510 826629.00 827128.99 1144496.00 1144995.99 17,205
461 241,612 827129.00 827628.99 1144496.00 1144995.99 13,699
462 335,074 827629.00 828128.99 1144496.00 1144995.99 18,920
463 1,548 828129.01 828143.16 1144957.45 1144995.98 88
464 50,999 820228.70 820628.99 1144315.08 1144495.99 2,907
465 177,354 820629.00 821128.99 1144084.92 1144495.99 9,973
466 368,104 821129.00 821628.99 1143996.00 1144495.99 20,721
467 415,128 821629.00 822128.99 1143996.00 1144495.99 23,299
468 399,945 822129.00 822628.99 1143996.00 1144495.99 22,510
469 349,779 822629.00 823128.99 1143996.00 1144495.99 19,729
470 326,276 823129.00 823628.99 1143996.00 1144495.99 18,350
471 361,865 823629.00 824128.99 1143996.00 1144495.99 20,430
472 304,717 824129.00 824628.99 1143996.00 1144495.99 17,146
473 317,674 824629.00 825128.99 1143996.00 1144495.99 17,901
474 368,335 825129.00 825628.99 1143996.00 1144495.99 20,713
475 421,506 825629.00 826128.99 1143996.00 1144495.99 23,851
476 307,818 826129.00 826628.99 1143996.00 1144495.99 17,366
477 152,518 826629.00 826984.83 1143996.00 1144495.99 8,527
478 98 827239.72 827628.73 1144486.48 1144495.98 6
479 37,472 827629.19 827909.48 1144358.43 1144495.99 2,122
480 58,361 821322.47 821628.99 1143856.31 1143995.99 3,313
481 331,390 821629.00 822128.99 1143626.80 1143995.99 18,589
482 392,462 822129.00 822628.99 1143496.00 1143995.99 22,138
483 370,738 822629.00 823128.99 1143496.00 1143995.99 20,971
484 356,131 823129.00 823628.99 1143496.00 1143995.99 19,923
485 454,683 823629.00 824128.99 1143496.00 1143995.99 25,804
486 349,388 824129.00 824628.99 1143496.00 1143995.99 19,594
487 358,256 824629.00 825128.99 1143496.00 1143995.99 20,297
488 378,884 825129.00 825628.99 1143496.00 1143995.99 21,221
489 480,803 825629.00 826128.99 1143496.00 1143995.99 27,313
490 326,830 826129.00 826628.99 1143496.00 1143995.99 18,333
491 19,221 826629.00 826726.46 1143807.68 1143995.98 1,073
492 14,955 822414.75 822628.99 1143398.22 1143495.99 841
493 121,912 822629.01 823128.99 1143170.66 1143495.99 6,864
494 306,166 823129.00 823628.99 1142996.00 1143495.99 17,212
495 378,558 823629.00 824128.99 1142996.00 1143495.99 21,390
496 346,199 824129.00 824628.99 1142996.00 1143495.99 19,468
497 297,543 824629.00 825128.99 1142996.00 1143495.99 16,832
498 375,717 825129.00 825628.99 1142996.00 1143495.99 21,151
499 374,309 825629.00 826128.99 1142996.00 1143495.99 21,241
500 156,209 826129.00 826467.97 1142996.02 1143495.99 8,805
501 5,700 823508.48 823628.99 1142940.71 1142995.98 323
502 154,908 823629.00 824128.99 1142712.49 1142995.99 8,713
503 284,918 824129.00 824628.99 1142496.04 1142995.99 16,175
504 361,152 824629.00 825128.99 1142496.00 1142995.99 20,299
505 354,996 825129.00 825628.99 1142496.00 1142995.99 20,148
506 303,017 825629.00 826128.99 1142496.00 1142995.99 17,074
507 9,012 826129.00 826208.69 1142841.63 1142995.98 512
508 183 824600.79 824628.97 1142483.32 1142495.96 11
509 90,798 824629.05 825128.99 1142254.82 1142495.99 5,093
510 188,517 825129.00 825628.99 1142025.97 1142495.99 10,775
511 104,779 825629.00 825950.96 1141996.96 1142495.99 5,912

 

TABLE 2. Descriptions of variables, including names, definitions, units of measurement, types and ranges of numeric values.

Variable

Variable definition

Units

Type

Range numeric values

EASTING

meters east of the central meridian in UTM zone 16 North

meters

continuous

820129.90 – 830528.19

NORTHING

meters north of the equator

meters

continuous

1141997.00 – 1161879.26

ELEVATION

meters above the WGS 1984 reference ellipsoid

meters

continuous

8.3 – 553.7

GROUND

predicted ground elevation (WGS 1984)

meters

continuous

-999 (missing) – 504.41

CLASS

classification of point as vegetation or ground surface

N/A

integer

5 (veg.) or 2 (ground)

RETURN

pulse return number for a given output pulse

N/A

integer

1 – 3

NRETURN

total number of pulse returns for a given output pulse

N/A

integer

1 – 3

INTENSITY

integer representation of pulse return magnitude

N/A

integer

0 – 255

TIME

laser fire time (since start of GPS week)

seconds

continuous

219239.90 – 226528.91

SCANANGLE

scan angle at time of laser shot (+ is to the right)

degrees

integer

-29 to + 24

 

Fig. 1
 
   FIG. 1. Canopy and ground elevation from LiDAR remote sensing. Each image is an interpolated surface derived from LiDAR data for the same 500 × 500 m section of old-growth forest at La Selva, Costa Rica. (a) The surface model contains estimates of canopy elevation relative to a reference ellipsoid (i.e., sea level). To calculate vegetation height above ground, laser pulses are classified as ‘vegetation’ or ‘ground’. Ground pulses were interpolated to produce a digital terrain model, which estimates ground elevation (b). Subtraction of the digital terrain model from the canopy surface produces a model of canopy height (c). The tallest canopy and emergent trees are red objects in the canopy height model, and canopy gaps are dark blue. This figure was generated using data in tile 152.

 

Fig. 2
 
   FIG. 2. Location of study site in Costa Rica, Central America, and extent of LiDAR data coverage. The red area in northeast Costa Rica is the extent of LiDAR data coverage drawn to scale.

 

Fig. 3
 
   FIG. 3. Extent of LiDAR data coverage and index to data tiles. Each square represents a 500 × 500 m tile. Numbers correspond to the tile data number. The location of La Selva Biological Station (16 km2) is shown for reference and scale. The shapefile index to data tiles used to make this figure and land use history for La Selva are available as related materials (section V.C.1 and V.C.2). Sample sizes and the extent of LiDAR data within each tile are in Table 1. As an example, a visualization of tile 152 is in Fig. 1.

 

Fig. 4
 
   FIG. 4. Validation of LiDAR data in a lowland Neotropical rain forest. Data show the relationship between ground elevation estimates made using field measurements (Y axis) and LiDAR remote sensing (X axis). (a) Within 16 km2 of mixed-land-use lowland forest at La Selva, Costa Rica. The relationship is: field measured elevation (m) = 0.999 ± (0.051) × LiDAR predicted elevation (m) - 1.289 m ± (0.001), Pintercept < 0.001 Pslope < 0.001, r2 = 0.996, RMSE = 1.66 m, n = 8798. (b) The relationship restricted to 778 ha of old-growth. The relationship is: field measured elevation (m) = 0.999 ± (0.001) × LiDAR predicted elevation (m) - 1.406 ± (0.126) m, Pintercept < 0.001 Pslope < 0.001, r2 = 0.994, RMSE = 1.85 m, n = 4184. P values and standard errors for regression parameters (in parentheses) were estimated using a nonparametric bootstrap with 2,000 iterations.

 

Fig. 5
 
   FIG. 5. Validation of LiDAR data in a lowland Neotropical rain forest. Data show the difference in elevation between field and LiDAR estimates (Y axis) and topographic slope (X axis). Topographic slope is a known source of bias in LiDAR data. (a) Within 16 km2 of mixed-land-use lowland forest at La Selva, Costa Rica. The relationship is: field measured elevation - LiDAR predicted elevation (m) = -0.004 ± (0.002) × slope (degrees) - 1.351 ± (0.031) m, Pintercept < 0.001 Pslope = 0.058, r2 = 0.000, RMSE = 1.66 m, n = 8798. (b) The relationship restricted to 778 ha of old-growth. The relationship is: field measured elevation - LiDAR predicted elevation (m) = -0.004 ± (0.004) × slope (degrees) - 1.381 ± (0.049) m, Pintercept < 0.001 Pslope = 0.146, r2 = 0.000, RMSE = 1.84 m, n = 4184. P values and standard errors for regression parameters (in parentheses) were estimated using a nonparametric bootstrap with 2,000 iterations.

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