UNIVERSITY OF CALIFORNIA Santa Barbara An Assessment of Hyperspectral and Lidar Remote Sensing for the Monitoring of Tropical Rain Forest Trees A Dissertation submitted in partial satisfaction of the requirements for the degree Doctor of Philosophy in Geography by Matthew Loren Clark Committee in charge: Professor Dar A. Roberts, Chair Professor Oliver A. Chadwick Professor David B. Clark Professor Phaedon C. Kyriakidis September 2005 The dissertation of Matthew Clark is approved. ____________________________________________ Oliver A. Chadwick ____________________________________________ Phaedon C. Kyriakidis ____________________________________________ David B. Clark ____________________________________________ Dar A. Roberts, Committee Chair September 2005 An Assessment of Hyperspectral and Lidar Remote Sensing for the Monitoring of Tropical Rain Forest Trees Copyright © 2005 by Matthew Loren Clark iii ACKNOWLEDGEMENTS This research would not have been possible without financial support from NASA Headquarters, through an Earth System Science Fellowship Grant NGT530436, and a Teaching Assistantship from the UCSB Geography Department. I am also deeply thankful to the AVIRIS team at NASA’s Jet Propulsion Laboratory, who let me use their portable spectrometer and who paid the costs to ship the instrument to Costa Rica for my field campaign. In particular, I would like to thank Rob Green and Jessica Faust for training and logistical support during my graduate studies. Another critical part of this research was access to unique tropical rain forest datasets. The Spectral Information Technology Application Center (SITAC) collected the HYDICE imagery and the U.S. Army Corps of Engineers Topographic Engineering Center collect the FLI-MAP lidar data used in this research. Both datasets were graciously donated to the Organization for Tropical Studies (OTS), who manages the La Selva Biological Station in Costa Rica. OTS provided logistical support for these campaigns, as well as for my field work in Costa Rica. Topographic field data in my research was based upon work supported by the National Science Foundation under Grant DEB-0129038. Old-growth tree data were collected by David and Deborah Clark, with support from the National Science Foundation (Grant DEB-0129038) and the Andrew W. Mellon Foundation. Plotscale tree height measurements were provided by Jack Ewel, whose work is supported by the National Science Foundation (Grant DEB-9975235) and by the Andrew W. Mellon Foundation. iv I would like to thank my main academic mentors in my doctoral research, Dar Roberts and David Clark. Dar taught me to appreciate the physics behind remote sensing and he propelled me forward with his keen intellect, enthusiasm and energetic teaching abilities. David instilled in me a profound appreciation for tropical scientific research and the wonders of the La Selva forest. His enthusiasm for bridging the gap between remote sensing and forest ecology inspired me to return to graduate school and to conduct my research at La Selva. In addition, I would like to thank my committee members, Phaedon Kyriakidis and Oliver Chadwick, for their academic guidance in helping me complete my dissertation. Thank you to my parents, Adrianne and Don Clark, as well as my sister Phoebe Kleiger. My mother has been a solid pillar of emotional support and a wealth of wisdom during my entire academic career. I would not have been able to complete my doctoral program without my loving wife, Ana Horta. I am deeply grateful to have Ana in my life, and I thank her for her love, companionship and willingness to leave her family and country and to put her career on hold so that I could complete my doctoral degree. I would like to thank past and present OTS staff, especially Robert Matlock, Jorge Jimenez, Cristián Coronas, Orlando Vargas, and Antonio Trabucco for their assistance in helping bring my research to fruition. I am greatly indebted Leonel Campos and William Miranda at La Selva, who provided me with warm companionship and invaluable expertise while conducting my field work. I also thank my colleagues in the VIPER lab (and honorary members) for their friendship, computer code, morale and lively discussions: Phil Dennision, Kerry v Halligan, Carl Legleiter, Izaya Numata, Seth Peterson, Becky Powell, Dylan Prentiss, and Carlos Souza. I also thank the Geography Department staff, especially Michelle Kueper, Connie Padilla, and Beilei Zhang. Finally, I would like to thank Deborah Clark, Stephanie Bohlman and Jim Kellner for their friendship and enthusiastic support of my research. vi VITA OF MATTHEW LOREN CLARK September 2005 EDUCATION Bachelor of Arts in Integrative Biology and Environmental Science, University of California, Berkeley, December 1993 Master of Science in Ecosystem Analysis and Conservation, University of Washington, Seattle, December 1998 Doctor of Philosophy in Geography, University of California, Santa Barbara, September 2005 (expected) PROFESSIONAL EMPLOYMENT 2003-2005: Graduate Student Researcher, Department of Geography, University of California, Santa Barbara 2001-2002: Teaching Assistant, Department of Geography, University of California, Santa Barbara 1998-2001: GIS Laboratory Manager, La Selva Biological Station, Organization for Tropical Studies, Costa Rica 1996-1998: Graduate Research Assistant, Long Term Ecological Research Network (LTER), University of Washington, Seattle 1994-1996: Senior Biochemical Technician, Genentech, Inc., South San Francisco AWARDS National Aeronautics and Space Administration Earth Systems Science Fellowship, 2002-2005 California Space Grant Fellowship, 2004 PUBLICATIONS Clark, M.L., Roberts, D.A., & Clark, D.B. (2005). Hyperspectral discrimination of tropical rain forest tree species at leaf to crown scales. Remote Sensing of Environment, 96(3-4), 375-398. Clark, M.L., Clark, D.B., & Roberts, D.A. (2004). Small-footprint lidar estimation of sub-canopy elevation and tree height in a tropical rain forest landscape. Remote Sensing of Environment, 91(1), 68-89. Clark, D.B., Read, J.M., Clark, M.L., Murillo Cruz, A., Fallas Dotti, M., & Clark, D.A. (2004). Application of 1-m and 4-m resolution satellite data to studies of tree demography, stand structure and land-use classification in tropical rain forest landscapes. Ecological Applications, 14(1), 61–74. Clark, M.L., Roberts, D.A., Gardner, M. & Weise, D.R. (2004). Estimation of Hawaiian Islands fire fuel parameters from AVIRIS imagery. Proc. 13th Annual JPL Airborne Earth Science Workshop, Jet Propulsion Laboratory, Pasadena, CA. Powell, R.L., Matzke, N., de Souza, Jr., C, Clark, M.L., Numata, I., Hess, L.L., & Roberts, D.A. (2004). Sources of error in accuracy assessment of thematic landvii cover maps in the Brazilian Amazon. Remote Sensing of Environment, 90, 221234. Clark, M.L. (1998). An Analysis of Western Olympic Peninsula Forest Structure Using Combined Synthetic Aperture Radar and Landsat Thematic Mapper Images. Master of Science Thesis, University of Washington, Seattle, WA. 214 pp. PRESENTATIONS Clark, M.L., Clark, D.B, & Roberts, D.A. “Remote sensing of tropical rain forest structure with small-footprint lidar”, Ecological Society of America & International Congress of Ecology, 90th ESA Annual Meeting, August 7-12, 2005, Montreal, Canada. Clark, M.L., Clark, D.B, & Roberts, D.A. “Lidar estimation of sub-canopy elevation and tree height in a tropical rain forest landscape”, Guest lecture presentation, University of California, Geography Dept., May 27, 2005, Santa Barbara, CA Clark, M.L., Roberts, D.A., & Clark, D.B. “Hyperspectral discrimination of tropical rain forest tree species at leaf to crown scales”, American Society of Photogrammetry and Remote Sensing Annual Conference, March 7-11, 2005, Baltimore, MD Clark, M.L., Roberts, D.A., Gardner, M. & Weise, D.R. “Estimation of Hawaiian Islands fire fuel parameters from AVIRIS imagery”, 13th Annual JPL Airborne Earth Science Workshop, March 31-April 2, 2004, Pasadena, CA. Clark, M.L., Clark, D.B., & Roberts, D.A. “Lidar estimation of sub-canopy elevation and tree heights at La Selva”, Visiting researcher colloquium, April, 14, 2004, La Selva Biological Station, Costa Rica. Clark, M.L., Powell, R., Matzke, N., de Souza, Jr., C., Numata, I., Hess, L.L., Roberts, D.A. “Accuracy assessment of remote sensing products using airborne videography: A case study from Rondônia, Brazil”, Invasive Exotic Plants: Approaches for the Florida Landscape, Conference and Workshop, February 1214, 2003, Miami, FL. Clark, M.L. Evaluación de sucesión usando sensores remotos comerciales. Guest lecture presentation, Universidad de Tucumán, April 22, 2003, San Miguel de Tucumán, Argentina. Clark, M.L. (1999). “Teledetección y ArcView GIS”. World Bank Central American Ecosystem mapping project workshop, June, 1999, Panama City, Panama. viii ABSTRACT An Assessment of Hyperspectral and Lidar Remote Sensing for the Monitoring of Tropical Rain Forest Trees by Matthew L. Clark The main objective this research was to assess two types of emerging remote sensing technology, hyperspectral and lidar sensors, for the automated discrimination of tropical rain forest tree (TRF) species. The hyperspectral data contain information on the biochemical and structural properties of crowns, while the lidar data contain structural information. I hypothesized that these two datasets combined would permit greater species classification accuracy than either dataset alone. Working in an old-growth TRF in Costa Rica, canopy-emergent individual tree crowns (ITCs) for seven target species were manually digitized with reference to high spatial resolution hyperspectral and lidar datasets that were acquired from airborne sensors. Multispectral and hyperspectral classification was performed using pixel- and crown-scale spectra and spectral angle mapper (SAM), maximum likelihood (ML), and linear discriminant analysis (LDA) classifiers. Pixel-majority and crown-scale ITC classifications were significantly more accurate with hyperspectral data relative to multispectral data, revealing the importance of the ix spectral detail offered by hyperspectral imagery. Additional techniques were explored to best harness this spectral information. These included incorporating hyperspectral metrics into decision trees (DTs) and multiple endmember spectral mixture analysis (MESMA). The best spectral-based classification accuracy was with crown-scale spectra and a relatively simple LDA procedure. These results suggested that hyperspectral imagery need not be acquired at a very high spatial resolution or analyzed with sophisticated techniques to provide adequate discrimination of species. Leaf phenology was important in mapping TRF tree species. Leaf-off trees had distinct volume-scattering and spectral mixing properties that influenced classifier variable selection as well as final classification accuracy. Crown-scale hyperspectral data were combined with structural data from the lidar sensor in LDA and DT classifiers. There were significant differences in the majority of lidar-derived structural metrics among the study tree species; however, the addition of this information to the classifiers did not improve classification accuracies. Although lidar data was not useful for species discrimination, it did provide an unprecedented view of canopy topography and sub-canopy elevation that is difficult to measure using traditional techniques. x TABLE OF CONTENTS CHAPTER 1: Introduction............................................................................................ 1 CHAPTER 2: Lidar estimation of sub-canopy elevation and tree heights ................. 17 CHAPTER 3: Discrimination of tree species at multiple scales................................. 71 CHAPTER 4: Classification of tree species with absorption features and binary decision trees............................................................................................................. 132 CHAPTER 5: Classification of tree species with multiple endmember spectral mixture analysis ........................................................................................................ 186 CHAPTER 6: Comparison of lidar and hyperspectral data for tree classification ... 221 CHAPTER 7: Conclusions........................................................................................ 254 REFERENCES.......................................................................................................... 273 APPENDIX I: List of Acronyms .............................................................................. 306 APPENDIX II: Summary of spectral metrics (Chapter 4)........................................ 309 xi CHAPTER 1: Introduction 1.1. Background Mapping and monitoring of threats to tropical rain forest biotic diversity Tropical rain forests (TRF) now cover only 6.4 percent of the Earth’s terrestrial surface (9.5 x 108 km2) yet they maintain a large proportion of the world’s biotic diversity (Thomas et al., 2004; Whitmore, 1990). There are many reasons to be concerned about TRF biotic diversity. Monetary benefits include timber and nontimber (e.g., fruits, nuts, medicines) products (Fearnside, 1999). The vast genetic diversity contained in TRF could provide future economic wealth and utility, such as through undiscovered pharmaceuticals or food resources (Fearnside, 1999; Laurance, 1999). Besides these utilitarian benefits of TRF, many authors have written about the ethical reasons to protect the irreplaceable species and human cultures maintained by these forests (Raven, 1988; Laurance, 1999). Often neglected in evaluating TRF value are the many ecosystem processes or “services” that an intact forest provides, such as flood control, soil conservation, carbon storage and regional and global climate regulation (Laurance, 1999). For example, it is estimated that tropical forests store 59% and 27% of global carbon stocks in vegetation biomass and soils, respectively (Dixon et al., 1994). Deforestation and forest fragmentation are well-documented immediate threats to tropical forest biodiversity (Achard et al., 2002; Fearnside, 1999; Phillips, 1997; Skole & Tucker, 1997). Digital images from satellites have been crucial for understanding the spatial extent and temporal dynamics of these threats because the 1 sensors provide continuous spatial and frequent temporal measurements of reflected radiance from TRF canopies. Mapping of tropical forests has primarily relied on medium spatial resolution satellite imagery from multispectral sensors, such as Landsat Thematic Mapper (TM) with 30-m pixels and 6 optical bands. This imagery is relatively inexpensive and has permitted the mapping of general forest cover classes needed to calculate the rate and extent of deforestation and forest fragmentation (Cochrane et al., 1999; Roberts et al., 2002; Skole & Tucker, 1993; Steininger et al., 2001). However, variability in forest types due to high tree diversity and natural and human disturbances results in complex radiance signals that are difficult to discriminate using coarse spectral and spatial resolution sensors, leading to significant errors in estimates of land cover area and temporal change (Achard et al., 2002; Cochrane, 1999; Foody, 2003; Powell et al., 2004; Skole & Tucker, 1993). Tropical forest degradation is also a threat to biodiversity. Factors causing degradation include: 1) the selective extraction of plants, such as the removal of commercially-valuable trees (i.e., selective logging); 2) selective removal of animals (i.e., defaunation); 3) invasive species; 4) climate change; 5) changing atmospheric composition, such as elevated CO2 on plant growth; 6) changing tree mortality and recruitment rates; 7) fire intrusion into forests; and 8) forest fragmentation (Cochrane et al., 1999; Nepstad et al. 1996; Phillips, 1997; Uhl & Kauffman, 1990). Many of these biodiversity threats have direct and indirect effects on tropical trees, which are the major components of forest structure and largely determine the 2 microclimate, substrate and food resource environments that sustain the rich plant and animal biota in the forest. One example of an indirect effect on biodiversity is climate change. Warmer global temperatures and associated changes in precipitation patterns linked to greenhouse gas emissions may alter tree growth, recruitment and mortality rates, thereby creating new assemblages of trees (Clark, Piper et al., 2003; Laurance et al., 2004; Phillips, 1997). If these altered tree communities fail to sustain the complex interactions among trees, pollinators, seed dispersers, herbivores, symbiotic fungi and other species that are common in tropical forests, then overall biodiversity will likely decline (Laurance et al., 2004). One recent global-scale study concluded that climate-change effects on tropical forests over the next 50 years may pose as much risk to species survival as deforestation (Thomas et al., 2004). Our scientific understanding, monitoring, and management of these threats to biodiversity are greatly hindered by a lack of spatially- and temporally-extensive information on tree demography (Clark, Read, et al., 2004), such as species, location, growth, and survivorship. Forest degradation affects one to several trees, and it may go undetected in coarse spatial resolution imagery unless it causes clear spectral change. For example, selective logging may increase fractions of exposed soil or dead vegetation, permitting detection in coarse resolution pixels (Asner et al., 2002; Souza & Barreto, 2000). Individual tree crowns (ITCs) are not resolved in coarse-resolution imagery from a sensor such as Landsat. Most available data on tree demography come from relatively small, sparsely-sample field plots with infrequent re-sampling intervals. It is difficult to generalize such field data to the 3 landscape, regional and global scales needed for understanding the important processes affecting biodiversity (Foody et al., 2003; Tuomisto, Poulsen et al., 2003; Phillips et al., 2003). However, more intensive sampling over broader spatial scales is generally not possible due to inaccessibility or cost constraints. For more detailed maps of ITCs over broad spatial scales, scientists and land managers in the tropics have relied on visual interpretation of aerial photographs from film cameras (Clément & Guellec, 1974; Herwitz et al., 1998; Myers & Benson, 1981; Trichon, 2001). Aerial photography has limited use in the tropics because these environments are mainly located in developing countries, where imagery is prohibitively expensive or otherwise difficult to obtain (Clark, Read, et al., 2004). In addition, because photo-interpretation relies on human intervention, it is prone to inconsistencies among interpreters (Congalton & Mead, 1983; Myers and Benson, 1981). Emerging remote sensing technology for individual tree crown analyses A new generation of high spatial resolution (< 4 m), multispectral airborne and spaceborne digital sensors now exists that can resolve ITCs as groups of digital image pixels (McGraw et al., 1998; Gougeon & Leckie, 2003). In particular, the launches of the commercial IKONOS and Quickbird satellites in 1999 and 2001 (Space Imaging, Thornton, CO; Digital Globe, Longmont, CO), respectively, have greatly increased the availability of high spatial resolution imagery for applications in tropical forests (reviewed in Hurtt et al., 2003). Preliminary TRF studies in Costa 4 Rica and Brazil have shown that IKONOS imagery can resolve ITCs and permit crown size measurements (Asner et al., 2002; Clark, Read, et al., 2004; Read et al., 2003) and the tracking of tree mortality (Clark, Castro, et al., 2004). These details of crown structure, along with other canopy-scale information such as presence/absence of tree species, logging roads and patios, may greatly improve the discrimination of selective-logging impacts (Clark, Read, et al., 2004; Read et al., 2003; Souza & Roberts, 2005). This capability has immense potential for monitoring forest degradation and insuring regulatory and certification compliance over broad scales (Read et al., 2003). In high spatial resolution optical imagery, each ITC encompasses many pixels with an n-dimensional spectrum that can be used for automated species discrimination. Current research shows that species discrimination is best accomplished by aggregating pixels into their respective crowns for object-based (as opposed to per-pixel) classification using each crown’s spectral and spatial properties (Gougeon, 1995; Leckie, Gougeon, Hill et al., 2003; Meyer et al., 1996; Preston et al., 1999). Locating and delineating these image objects can be accomplished with manual digitization (Asner et al., 2002; Clark, Read, et al., 2004; Lamar et al., 2005; Read et al., 2003). For consistency and reduction of production costs, ITC inventory over large spatial extents will require automated algorithms for crown detection and delineation. Automated ITC delineation is a sub-discipline of image segmentation (Warner et al., 1999). Recently developed algorithms are reviewed extensively in Hill and Leckie (1999), McGraw et al. (1998), and Key et 5 al. (2001). Techniques generally involve collapsing spectral data into one illumination/albedo band (Warner et al., 1999) from which radiometric maxima and minima are analyzed to locate tree centers (maxima) and define crown edges (minima), respectively (Gougeon, 1999; Key et al., 2001; Culvenor, 2002). Most segmentation algorithms were developed for forestry applications in coniferdominated stands (e.g., Brandtberg & Walter, 1998; Gougeon, 1999; Lamar et al., 2005). There are still many challenges to crown delineation in complex, old-growth hardwood forests, where trees may intertwine and overlap, inter-tree shadows are narrow, and canopy-gap shadows are prevalent (McGraw et al., 1998; Warner et al., 1999), causing automated segmentation schemes to identify clusters of trees rather than individual crowns (Culvenor, 2002). High spatial resolution optical sensors typically measure a few spectral regions in broad bands (i.e., multispectral). Vegetation spectra are controlled by similar factors, such as chlorophyll and water absorption in leaf tissues, and as a consequence their spectral differences are subtle and substantial spectral detail may be needed to distinguish among species (Price, 1994; Cochrane, 2000). Airborne and spaceborne hyperspectral optical sensors offer such detailed spectral information by measuring the visible to shortwave infrared regions of the electromagnetic spectrum (400-2500 nm) in over 100 channels. The Hyperion sensor on the EO-1 platform is the only spaceborne sensor offering hyperspectral data over the large spatial extents needed for regional ITC mapping; however, the 30-m resolution of the imagery has precluded ITC classification research. High spatial resolution hyperspectral data (typically > 4 m) is now available from commercial (e.g., HyMap 6 [Integrated Spectronics Pty Ltd., Baulkham Hills NSW, Australia]) and experimental (e.g., Airborne Visible/Infrared Imaging Spectrometer [AVIRIS; Jet Propulsion Laboratory, NASA, Pasadena, CA USA]) airborne sensors. It is anticipated that this high spatial and spectral resolution imagery will make the automated classification of tropical species a reality (Cochrane, 2000); however, this hypothesis has remained untested with an airborne or spaceborne hyperspectral sensor. Another promising technology for ITC-level analysis is LIght Detection And Ranging (lidar). Lidar systems are an active form of remote sensing that uses a scanning laser to measure surface heights (Baltsavias, 1999a). These sensors have emerged as the premier instruments for the generation of detailed digital terrain models (DTMs: Kraus & Pfeifer, 1998; Petzold et al., 1999) and the estimation of forest height (Næsset, 1997; Magnussen & Boudewyn, 1998; Persson et al., 2002) and aboveground biomass (Drake, Dubayah, Clark et al., 2002). Small-footprint lidar sensors record the three-dimensional height distribution of crown components, providing information on crown structure that is useful for conifer species discrimination (Brandtberg et al., 2003; Holmgren & Persson, 2004). There have been no attempts to discriminate species with lidar in a tropical forest environment. Automated, computer-based classification of tropical rain forest trees In visual interpretation of aerial photographs, the species of tree crowns are distinguished using crown color and structural properties, such as branch architecture, canopy position, contour shape, size, foliage cover and texture 7 (Fournier et al., 1995; Trichon, 2001). These spectral and structural properties should be equally important for automated, computer-based ITC classification. Hyperspectral imagery provides visible color information, but also includes the near infrared and shortwave infrared regions of the electromagnetic spectrum beyond the range of human vision. Tree spectral response is largely determined by the principle tissues in their crown—leaves and bark (Asner, 1998; Roberts et al., 2004). In general, the optical properties of these tissues are controlled by surface and internal structure and biochemical concentrations, such as water, chlorophyll, lignin, and cellulose. At leaf scales, many species spectra may be similar in shape because there are only a few factors that control leaf reflectance (Price 1994, Poorter et al, 1995; Cochrane 2000). Furthermore, there may be significant spectral variation between individuals of a single species or within one individual’s crown, which could limit our ability to discriminate species through spectral techniques (Cochrane, 2000). This is especially true in the tropics, where epiphylls quickly colonize species with long-lived leaves (Roberts, Nelson et al. 1998) thereby increasing conspecific (within species) leaf spectral variation. Unique species spectra are more likely found at branch or crown scales, where crown structure exerts an influence on overall spectral response in the sensor’s ground instantaneous field of view (GIFOV). At these scales, important structural properties such as branch architecture, leaf arrangement and morphology, and species phenology (e.g., leaf turnover, flowering) combine to determine the relative mixtures and illumination of photosynthetic vegetation, non-photosynthetic materials, and shadows that form the radiance signal measured by the sensor (Asner et al., 1998, Roberts et al., 2004). 8 For imagery with a spatial resolution finer than the scale of a crown, the spatial arrangement of spectral vectors (i.e., pixels) may encode additional information on crown structure. Few studies have used pixel spatial information for automated ITC species discrimination, and there is inconclusive evidence that it greatly improves forest composition classification (Franklin et al., 2000; Leckie, Gougeon, Walsworth, & Paradine, 2003, Wang et al., 2004; although see Franklin et al., 2001; Zhang, et al., 2004). A more straight-forward approach to remote measurement of crown structure is through the use of small-footprint lidar sensors, since their measurements respond directly to the physical arrangement of crown tissues (Brandtberg et al., 2003; Holmgren & Persson, 2004). 1.2. Research overview and objectives The main goal of this research is to investigate high spatial resolution hyperspectral and lidar sensors for automated tropical rain forest species classification. The study site and data sets are described below in Section 1.3. This research involved several analytical decisions and constraints, including: 1) selection of tree species of interest, 2) crown detection and delineation methods, 3) classification schemes, 4) spatial scale of observation, and 5) calculation and selection of pertinent spectral and structural variables. I focused on emergent individuals of tree species which could be visually identified both in the remote sensing data and in the field. Species were chosen based on their importance in ecological research or conservation efforts and my ability to find a sufficiently-large sample size. I opted to manually delineate crowns 9 so that I could focus on questions regarding species discrimination rather than tree detection and delineation (i.e., object segmentation). Several different classification schemes have been used for ITC species classification, including neural networks (Gong et al., 1997), linear discriminant analysis (Brandtberg et al., 2003; Fung et al., 1998; Gong et al., 1997; Holmgren & Persson, 2004), spectral angle mapper (Xiao et al., 1999), spectral mixture analysis (Xiao et al., 2004), decision trees (Preston et al., 1999), nearest-neighbor (Wang et al., 2004); parallelpiped (Meyer et al., 1996), and the popular Gaussian maximum likelihood classifier (Gougeon, 1995; Meyer et al., 1996; Leckie & Gougeon, 1999; Key et al., 2001; Leckie, Gougeon, Hill et al., 2003). I devote three chapters to an exploration of the relative trade-off among five classification techniques: maximum likelihood (ML), spectral angle mapper (SAM), linear discriminant analysis (LDA), decision trees (DT), and multiple endmember spectral mixture analysis (MESMA). These analyses explored several methods for isolating and extracting information from the hyperspectral and lidar data for optimal species discrimination. The issue of spatial scale of spectral measurement, and its effect on species separability is also analyzed and discussed. The general objectives of this research were to: 1. Develop hyperspectral techniques for classifying tropical rain forest tree species 2. Identify the optimal spectral regions and spatial scale for species discrimination 10 3. Evaluate the importance of lidar-derived crown structure information for species discrimination 4. Assess lidar technology for ecological analyses of tropical rain forests 1.3. Study site and remote-sensing datasets Study site overview: The La Selva Biological Station My research was conducted using data acquired at the La Selva Biological Station (LSBS), located in the Atlantic lowlands of north-east Costa Rica in the Sarapiquí canton (84°00'13.0" W, 10°25'52.5" N). LSBS is a 1614-ha reserve that contains a mixture of old-growth terra firme, swamp, secondary and selectivelylogged forests, as well as plantations, developed areas with buildings and mowed grass, and abandoned pastures with large grasses, shrubs and remnant trees (Fig. 1.1). Precipitation averages 4244 mm annually, with a comparatively dry season from January to April and a second smaller dry season from August to October (Frankie et al, 1974; Organization for Tropical Studies [OTS] meteorological data 1957-2003, http://www.ots.ac.cr). The old-growth forest (Fig. 1.1) is classified as a Tropical Wet Forest in the Holdridge Life Zone System and is characterized by a species-rich, multi-layered community of trees, palms, lianas, and other terrestrial and epiphytic plants (Holdridge, 1971; Hartshorn & Hammel, 1994). There are at least 400 species of hardwood trees in the reserve. Although some overstory trees can be completely deciduous for a part of the year, mainly in the dry season, the canopy is considered evergreen (Frankie et al., 1974; Hartshorn & Hammel, 1994). The geomorphology of the landscape is structured by two main features: 1) highly 11 eroded lava flows that contain a system of alternating ridges and stream valleybottoms separated by steep slopes, and 2) flat to gently undulating alluvial terraces (Sanford et al., 1994). The reserve is covered by a 50 x 100-m grid, oriented 32 degrees from North, with permanent monuments at each grid intersection. The local grid coordinate system permits researchers to accurately geo-locate field data in a local Geographic Information System (GIS) Cartesian coordinate system. La Selva 1 Costa Rica Km HYDICE extent FLI- MAP extent Rivers Land Use Developed Areas Selectively- logged Old- growth Forest Secondary Forest Pasture Plantation Swamp Figure 1.1. The La Selva Biological Station study site and extent of HYDICE hyperspectral and FLI-MAP lidar datasets. Hyperspectral data For this research, I had access to a unique, high spatial resolution hyperspectral dataset from the airborne HYperspectral Digital Imagery Collection Experiment 12 (HYDICE) sensor (Basedow et al., 1995). The U.S. Spectral Information Technology Application Center (SITAC) flew HYDICE over LSBS in March 30, 1998, which corresponds to the end of the drier season in the region. HYDICE is a push-broom, indium-antimonide hyperspectral sensor that measures 210 bands covering the 400-2500 nm region of the electromagnetic spectrum (Basedow et al., 1995). The LSBS flights were flown at a 3.17 to 3.20-km altitude between 7:558:27 am local time (13:55 to 14:27 UTC). Six runs of 0.5-km wide, variable length, 1.6-m spatial resolution data (0.5 mrad instantaneous field of view; IFOV) were acquired over old-growth forest, secondary forest, selectively-logged forest, tree plantations, pastures and the nearby town of La Guaria. The section of the dataset used in my research is shown in Figure 1.1. Pre-processing of the HYDICE imagery is described in Chapter 3. Lidar data Another unique dataset used in this research was from a high spatial resolution lidar sensor called FLI-MAP (John E. Chance & Associates, Lafayette, Louisiana). These data were acquired from a helicopter on September 12 and 13, 1997. FLIMAP is a small-footprint, first-return 900-nm laser sensor that has a 8000-Hz pulse rate, 30-degree scan angle, 2-mrad beam divergence, typically 9-point/m2 sampling density, 30-cm footprint spacing, and a rated vertical accuracy of ~10-cm (Blair & Hofton, 1999, Huising & Gomes Pereira, 1998). The 10-cm footprints (Hofton et al., 2002) were converted to a Triangular Irregular Network (TIN) (John E. Chance & Associates), and the final product delivered for analysis by the data distributor 13 (Fig. 2a) was a raster digital surface model (DSM) interpolated from the TIN with 0.33-m cell support (each containing a height). The DSM covered a 754-ha area of the LSBS reserve (Fig. 1.1). 1.4. Description of Chapters Chapter 2 is an assessment of small-footprint lidar technology for the estimation of ground elevation and tree heights in tropical landscapes. This research provided the foundation for Chapter 6, which sought to combine lidar and hyperspectral data for ITC species discrimination. In Chapter 2, I developed a method to retrieve canopy heights, as a canopy height model (DCM), from the original lidar height surface (the DSM). The DCM was then used in Chapter 6 for ITC species discrimination (discussed below). A key step in creating an accurate DCM was to estimate sub-canopy terrain elevation from the DSM. With the DCM as a final goal, I developed a ground-retrieval scheme to find terrain points. I then evaluated geostatistical techniques for interpolating a DTM from the points. The accuracy of the lidar-derived DTM was rigorously tested with a comparison to field-survey points. I discuss how differences in DTM accuracy are related to terrain slope and land-use factors. Next, the DSM and the derived DTM were used to calculate the DCM, and the accuracy of lidar-derived estimates of stem heights at individual tree and plot scales was assessed with reference to comparable field measurements. This research is considered an extreme test of lidar technology because: 1) there are several forest types with dense, structurally-complex canopies that severely restrict 14 ground-level light transmission and lidar returns (Clark et al., 1996; Drake, Dubayah, Clark et al., 2002; Montgomery & Chazdon, 2001), 2) all vegetation classes are in “leaf-on” conditions, exacerbating ground-retrieval difficulties, and 3) the study site covers a range of terrain conditions. A version of this research is published in Remote Sensing of Environment (Clark, Clark, & Roberts, 2004). Chapter 3 begins my analyses of ITC species discrimination. In this chapter, I examine the relative trade-offs between spectral regions, spatial scale of measurement, and traditional classification schemes for species discrimination using hyperspectral reflectance bands. Field spectrometer and airborne hyperspectral reflectance spectra were acquired from seven species of emergent trees at LSBS, permitting analyses at leaf, pixel and crown scales. My main objectives in Chapter 3 were to: 1) use statistical tests to determine if spectral variation among TRF tree species (interspecific) is greater than spectral variation within species (conspecific), thereby permitting spectral-based species discrimination; 2) identify the spatial scale and spectral regions that provide optimal discrimination among TRF emergent tree species; 3) develop an analytical framework for the species-level classification of ITCs based on their pixel- or crown-scale reflectance spectra; 4) assess the relative importance of narrowband hyperspectral versus broadband multispectral imagery for species identification; and, 5) assess the efficacy of three traditional classifiers: LDA, ML, and SAM. This research is published in Remote Sensing of Environment (Clark et al., 2005). In Chapter 4, I developed a purely hyperspectral-based method for ITC species classification. I first calculated a suite of hyperspectral metrics that respond to 15 crown structure and photosynthetic pigments, water and other biochemical absorption features. At tissue, pixel and crown scales, I tested for significant differences in mean response of these spectral metrics among my study tree species. I then assessed the utility of these metrics for ITC species discrimination using a DT classification scheme. A manuscript of this chapter is ready for submission to Remote Sensing of Environment. Chapter 5 explores MESMA for the species-level classification of ITCs. Starting with a large spectral library of image and laboratory spectra, I devised an automated approach to select optimal endmembers for two- and three-endmember MESMA models. In particular, the selection scheme sought to find within-species specialists while excluding among-species generalists. The selected endmembers were then used in MESMA to classify ITC species. This chapter will be submitted simultaneously with Chapter 4 for publication in Remote Sensing of Environment. The objective of Chapter 6 is to extend the spectral-based analyses from Chapters 3 and 4 to include lidar-derived, crown structure metrics calculated from the DCM (Chapter 2). I first assessed how crown structure, as quantified by lidar metrics, varies among species. I then explored the benefits of lidar-derived structure information for species classification. In Chapter 7, I summarize the findings from Chapters 2-6 and outline general conclusions about these new forms of remote sensing technology for tropical tree species discrimination. I also make recommendations for future research. 16 CHAPTER 2: Lidar estimation of sub-canopy elevation and tree heights 2.1. Introduction Digital terrain models (DTMs) and canopy height estimates are two important remote sensing products for studies of TRF ecology and management. DTMs describe the variation of elevation across a landscape and have been used in applications including mapping drainage basin geomorphology (Yin & Wang, 1999), flood modeling (Bates & De Roo, 2000), calculation of biophysical controls on vegetation distribution (e.g., temperature, solar radiation) (Dymond & Johnson, 2002), and spatial analysis of soil properties (Gessler et al., 2000). Canopy height estimates from remote sensing technology have a variety of potential applications in TRF, such as calculating surface roughness for atmosphereland interaction models (Raupach, 1994), spatial analyses of forest dynamics, such as canopy gap formation, distribution and turn-over (Birnbaum, 2001), identifying plant species (Brandtberg et al., 2003; Holmgren & Persson, 2004), mapping of wildlife habitat (Hinsley et al., 2002), modeling canopy rain interception (Herwitz & Slye, 1995), and modeling light penetration (Clark et al., 1996; Montgomery & Chazdon, 2001). Vegetation height is allometrically related to forest structure parameters, such as estimated aboveground biomass (Brown et al., 1995). Because roughly half of biomass is composed of carbon, improvements in our ability to map biomass through remote sensing will translate into better estimates of carbon stocks and flux at broad scales. Such advances are particularly important in tropical forests, which contain a large proportion of terrestrial carbon, and consequently have the 17 greatest potential to increase atmospheric carbon dioxide from deforestation (Dixon et al, 1994). Although passive optical and active synthetic aperture radar (SAR) signals and associated metrics are sensitive to forest aboveground biomass variation (Ranson et al., 1997; Sader et al., 1989), biomass estimates from these sensors tend to saturate at the high biomass levels typically found in tropical forests (Imhoff, 1995; Luckman et al., 1998, Steininger 2000). There have been relatively few published applications that have used smallfootprint lidar sensors in tropical rain forest environments (Blair & Hofton, 1999; Hofton et al., 2002). However, studies using the large-footprint LVIS sensor in Costa Rica have shown that there is immense potential of lidar technology for TRF research and monitoring efforts (Blair & Hofton, 1999; Drake, Dubayah, Clark et al., 2002, Drake, Dubayah, Knox et al., 2002; Hofton et al., 2002; Weishampel et al., 2000). For example, Drake, Dubayah, Clark et al. (2002) showed that lidar metrics applied to large-footprint waveforms can accurately predict aboveground biomass over a wide range of tropical forest conditions without saturation. 2.1.1. Digital terrain models and lidar Photogrammetry has been the traditional source of broad-scale DTMs throughout the world. There has been an increase in the use of IfSAR (interferometric synthetic aperture radar) and lidar active sensors as more economic alternatives for producing DTMs (Hodgson et al., 2003; Petzold et al., 1999). In many developing countries, where most tropical forests occur, the best publicly available topographic information is from the Shuttle Radar Topography Mission (SRTM), which used 18 IfSAR technology to produce a global terrestrial DTM with a 90-m horizontal resolution (Rabus et al., 2003). However, the relative vertical accuracy of this product at the 50-100 km scale is roughly 6 m due to errors from several systematic and random factors that can only be partially reduced by data post-processing. Lidar-derived DTMs estimated in open areas or under areas with low vegetation can have vertical accuracies ranging from 0.06 to 0.61-m root-mean-square error (RMSE) (Cobby et al., 2001; Huising & Gomes Pereira, 1998). Trunks, branches and leaves in dense vegetation tend to cause multiple-scattering reflections or absorption of the emitted laser energy so that fewer backscattered returns are reflected directly from the ground (Harding et al., 2001; Hofton et al., 2002). This effect increases with more canopy closure, canopy depth (or volume) and structural complexity (Harding et al., 2001; Hodgson et al., 2003; Hofton et al., 2002), and it is expected to be more severe for first-return only lidar systems because recorded returns generally come from the canopy (Magnussen & Boudewyn, 1998). The result is that the RMSE between the lidar-derived DTM and reference elevation data tends to increase in areas of dense vegetation because: 1) there are fewer groundreturn samples for DTM surface interpolation, and 2) those samples selected as ground may actually be understory vegetation, logs or rocks (Cobby et al., 2001, Hodgson et al. 2003; Raber et al., 2002). When non-ground samples are included in the interpolation, mean-signed residual error will be positive (lidar-reference), i.e., the lidar DTM overestimates the reference elevation. When actual ground samples are mistakenly filtered out of the data before DTM interpolation, as discussed below, results are less predictable and depend on local topographic curvature. DTM peaks 19 may be clipped or valleys filled due to inadequate retrieval of ground samples, resulting in the underestimation or overestimation of local elevation, respectively. Additional sources of error in lidar-derived DTMs include vertical and horizontal error in positioning the laser platform, laser scan angle, surface reflectivity, and slope of the terrain, all of which combined can add 0.20-2.00 m of error to an elevation estimate (Baltsavias, 1999b; Hofton et al., 2002; Huising & Gomes Pereira, 1998; Kraus & Pfeifer, 1998). The filtering of vegetation from sub-canopy ground returns for the interpolation of DTMs, or “bald Earth”, has been an active area of research. However, most algorithms for small-footprint lidar data are proprietary, and reported instrument and DTM accuracies are often poorly documented and generally assumed to be measured under optimal conditions—flat areas with no vegetation (Baltsavias, 1999a; Huising & Gomes Pereira, 1998;). Some vegetation-filtering techniques include morphological filters and statistical analyses of heights in a neighborhood, and may be fully-automated or involve some human interpretation (Huising & Gomes Pereira, 1998). Kraus and Pfeifer (1998) used an automated, iterative technique that interpolated a mean surface from the lidar cloud of xyz points and then successively removed or down-weighted points with residuals higher than a specified threshold. A relatively simple approach is to find local-minima relative to neighboring samples at a specified scale and/or search configuration (Cobby et al., 2001; Petzold et al., 1999). Resulting ground samples (i.e., local minima) must then be interpolated to form a surface. 20 There have been relatively few studies that provide rigorous accuracy assessments of lidar-derived DTMs under dense forest canopy with leaves, be they simple or composite leaves in hardwood forests or needles in conifer forests. Working with last-return lidar data flown over a leaf-on pine/deciduous forest landscape, Hodgson et al. (2003) identified ground points through a combination of proprietary software and human interpretation. A comparison of DTM elevation against 1470 survey-grade field measurements had an overall RMSE of 0.93 m. DTM error differed significantly by land use. Although RMSE was 0.33 m for low grass, it increased to 1.22 m and 1.53 m for the more structurally-complex scrubs/shrub and deciduous vegetation types, respectively. Furthermore, these researchers found that in the dense, multi-layered shrub/scrub class, there was a highly significant increase in DTM error of roughly 2 m from lowest (0-2 deg.) to steepest (6-8 deg.) slopes, which the authors attributed to vertical inaccuracies over relatively short horizontal distances under complex canopy. Cobby et al. (2001) developed an automated ground-retrieval scheme for a floodplain environment that included deciduous forests with leaf-on conditions. An initial DTM was interpolated from local-minima cells retrieved from non-overlapping, 5 x 5-pixel windows (10-m side) overlaid on a last-return DSM (2-m support). The final DTM was achieved by tailoring the ground-retrieval algorithm to short and tall vegetation classes. While terrain under short vegetation could be predicted with a 0.17-m RMSE (n=5), the RMSE was 3.99 m (n=12) under deciduous forests on steeper slopes (10-15 degrees). Using last-return filtered proprietary methods, Reutebuch et al. (2003) found that a DTM under conifer plantations had a 0.32-m RMSE and a +0.22-m 21 mean-signed error (overestimated, n=347 reference points). There was only a slight 0.15-m increase in mean error between clearcut and uncut forest classes. Hodgson et al. (2003) used a triangulated irregular network (TIN) to interpolate the DTM from ground points, while Cobby et al. (2001) used bilinear interpolation. Additional interpolation techniques include co-variance linear weighting approaches, such as inverse distance weighting described below (Reutebuch et al., 2003), which can help smooth the high-frequency effects of spurious vegetation points, especially when applied in iterative filtering schemes (Lohmann & Koch, 1999). 2.1.2. Geostatistical methods for DTM generation I used geostatistical techniques to interpolate a DTM from 0.33-m support ground cells derived from a vegetation-filtering algorithm. In the interpolation process, cells were treated as xyz-coordinate samples (i.e., points). Two common geostatistical interpolation algorithms, inverse distance weighted (IDW) (Bartier & Keller, 1996; Isaaks & Srivastava, 1989) and ordinary kriging (OK) (Goovaerts, 1997; Isaaks & Srivastava, 1989; Lloyd & Atkinson, 2002a, 2002b), were considered in this research. Both techniques involve a weighted linear combination of neighboring data samples in estimating values at unsampled locations. In the case of IDW, weighting of neighboring samples is based solely upon an estimation location-to-sample inverse-distance function, along with a user-specified power weight factor (Bartier & Keller, 1996). The further away a sample is from the estimation location, and the greater the power weight factor, the less influence the sample will have on the estimate value. OK uses a distance-covariance model to 22 weight samples based on their distance from the estimation location, as well as to down-weight samples that are clustered in space—an effect which tends to smooth the variance in the surface (Isaaks & Srivastava, 1989). The kriging covariance model is generally developed with reference to an empirical semivariogram (Goovaerts, 1997; Isaaks & Srivastava, 1989). Recently, Lloyd and Atkinson (2002a, 2002b) reported that the more sophisticated OK technique was more accurate than IDW when interpolating DTMs from photo-interpreted (Lloyd and Atkinson 2002a) or lidar-derived (Lloyd and Atkinson 2002b) elevation points. 2.1.3. Lidar vegetation height and forest structure estimation With discrete-return small-footprint systems, vegetation height is calculated as the difference between the original footprint heights and the bald-Earth DTM. The result is a set of estimated canopy heights with a footprint-scale support. Alternatively, a DSM is interpolated from footprint heights and subtracted from the DTM, thereby creating a canopy surface with height values recorded in square pixels or cells, i.e., a digital canopy model (DCM). Small-footprint lidar technology now permits the detection and segmentation of individual tree crowns from fine spatial resolution DCMs (Brandtberg et al., 2003; Persson et al., 2002). Metrics applied to either DCM cells or footprint heights from within crown segments have been used to estimate the height and structure of individual crowns (Brandtberg et al., 2003; Gaveau & Hill, 2003; Næsset & Økland, 2002; Persson et al., 2002; Popescu et al., 2003). Such measures also hold promise for tree species classification (Brandtberg et al., 2003; Holmgren & Persson, 2004). 23 Persson et al. (2002) estimated individual conifer tree heights from DCM-cell maxima with a model r2 of ~0.98 and RMS error of 0.63 m. Næsset & Økland (2002) estimated conifer tree height with the maximum of first-return footprints within crowns. The regression model explained 75% of the variance with a prediction RMS error of 0.23 m. Tree height was slightly overestimated by 0.18 m (3.15-m s.d.) in Næsset & Økland (2002), yet underestimated by 1.13 m in Persson et al. (2002). Næsset & Økland (2002) concluded that to reduce underestimation, high footprint density is needed to increase the probability of detecting the tops of conifer crowns. I know of only two studies that estimated heights for hardwood individuals. Working in West Virginia with forests in leaf-off winter conditions, Brandtberg et al. (2003) estimated hardwood tree heights with first-return footprints from within crown segments. A regression model explained 69% of the variance (RMSE not reported). There was an underestimation of height for taller trees, which the authors attributed to the low probability of the laser detecting the maximum crown height with leaf-off conditions, as well as random error in field measurements and inaccuracy in ground retrieval. In the United Kingdom, Gaveau and Hill (2003) estimated leaf-on hardwood tree and shrub heights from a first-return DCM with a model that explained 95% of the variance (1.89-m RMSE). The authors also presented strong evidence that an underestimation of tree heights (-2.12 m meansigned error) resulted from laser pulses penetrating into the crown before reflecting a detectable first-return signal. The depth of signal penetration depended on variation of foliage and branches in the crown at the fine scale of a lidar footprint. 24 Stand-scale height of conifer-dominated forests have been predicted with varying levels of success from small-footprint lidar metrics (Næsset, 1997, 2002, Næsset & Økland, 2002). Several authors have shown that laser underestimation can be minimized by comparing a quantile of ground measurements (i.e., mean, maximum height) to a certain quantile of upper-most canopy returns in the stand (Magnussen & Boudewyn, 1998; Næsset,1997, 2002). For example, Næsset (1997) found that the mean of lidar maxima (footprints) from a grid-overlay of square cells (i.e., 15 x 15 m) estimated average conifer stand height with a mean-signed error of -0.40 m to 1.90 m (8-20 m tree height; 1.5-ha stands; RMSE not reported). For mixed conifer and hardwood forests with 6 to 29-m tall trees, Næsset (2002) used multipleregression analyses to relate several lidar metrics to mean and dominant tree height at plot (0.02 ha) and stand (average 1.6 ha) scales. Lidar data included first and last pulses and metrics included the maximum, mean, density and various percentquantile measurements within the plots. Model cross-validation RMS errors for young to mature forest plots ranged from 0.05 to 0.07 m (R2=0.82 to 0.95) and 0.07 to 0.08 m (R2=0.74 to 0.93) for mean and dominant plot-scale height, respectively. Stand-scale heights were estimated as the mean of plot-scale lidar estimates, and resulting models explained 92% and 87% of the variance for mean and dominant tree height, respectively (RMSE not reported). I am not aware of any studies that have used small-footprint lidar to estimate stand-scale height for areas composed of evergreen, tropical tree species (i.e., hardwood trees, palms). However, one recent study by Lim et al. (2003) focused on leaf-on hardwood forests in Ontario, Canada. Plot-scale (0.04 ha) Lorey’s mean 25 height was estimated using either the mean or maximum of lidar pulses (first and last return). Regression models explained 66% and 86% of the variance, and it was found that tree height was underestimated (RMSE not reported). Fine-scale lidar data has also been used to estimate stand-scale biophysical properties, such as volume, biomass, crown diameter and diameter breast height (Lim et al., 2003; Popescu et al., 2003). Advances in analyzing large-footprint waveforms have produced metrics that can estimate biophysical parameters over dense conifer and broadleaf forests with considerable accuracy [see Lefsky et al. (2002) for a recent review of the literature]. Drake, Dubayah, Clark et al. (2002) used multiple-regression and metrics from LVIS waveforms to predict plot-scale (025 to 0.5 ha) basal area, aboveground biomass, and quadratic-mean stem diameter over a range of tropical forest types at the La Selva Biological Station, Costa Rica; their resulting models explained 93%, 72% and 93% of the variance, respectively, and models did not saturate with increasing forest height and complexity. 2.2. Methods 2.2.1. Topographic reference data The DTM derived in this research was compared to 3859 in situ elevation points (Table 2.1) that were surveyed in 1991 using optical-leveling techniques (Hofton et al., 2002). In the comparison, co-located survey points and DTM cells were determined by linking points to the nearest 1-m DTM cell centroid based on x,y coordinates. Survey data covered the range of land-use and geomorphic conditions found in the reserve. There were 1321 points measured at grid intersections, while 26 the remaining points did not have permanent markers and were located off the grid, mainly in the northwest to southeast direction between grid intersections. All survey points were transformed to the Universal Transverse Mercator (UTM), WGS-84 datum coordinate system from the local La Selva coordinate system using a leastsquares affine transformation based on 5 differential-GPS points, as used by Hofton et al. (2002). DGPS measurements were taken from permanent towers or in open areas to avoid obstruction by vegetation. Differential corrections were performed using data from a base station at La Selva. The transformation had an overall RMSE of 0.57 m (XRMS = 0.48 m, YRMS = 0.31 m). The vertical values of the survey points were shifted from the local reference mean sea level (MSL) to the WGS-84 ellipsoid MSL using the constant 11.44-m offset published by Hofton et al. (2002). This offset was verified by selecting the 4 to 5 nearest FLI-MAP (discussed below) lidar returns around three grid monuments in open areas with very flat, mowed lawn. The average vertical offset between surveyed and lidar-derived elevations was also found to be 11.44 m. Neither the FLI-MAP data nor the reference data were referenced relative to a geoid, and so the elevation offset between the datasets is constant across the study extent DTM error was analyzed relative to slope and land-use factors. A total of 932 survey points at grid intersections in old-growth forest were further classified into one of four slope classes based on field measurements from a separate forest structure study (Clark et al., 1999; field data summarized in Table 2.1). Slope classes included: 0-3, 3-10, 10-20 and > 20 degrees. A total of 2060 survey points were classified into one of seven land-use categories based on an overlay operation 27 with an existing year 2000 land-use map (Fig. 2.1) derived from historic aerial photographs, IKONOS imagery and survey maps (OTS, unpublished data). The land-use categories, from short-simple to tall-structurally-complex vegetation, were: Developed Areas, Pastures, Plantations (abandoned and current), Secondary forests (1 to 34 years old), Selectively-logged Forest (including 50 year-old, abandoned agroforestry areas), Swamp forests and Old-growth forests. To limit the confounding effect of slope on DTM error (discussed in results), only survey points on slopes less than 10 degrees were considered in the land-use analysis. A mask delineating areas with slopes less than 10 degrees was created by sampling the DTM-derived slope (Arc/Info 8.0.1, Environmental Systems Research Institute, Redlands, California) after filtering with an averaging 3x3-cell filter. 2.2.2. Individual tree heights In the field, the maximum height of a crown was estimated with a handheld laser range-finder (Impulse-200LR, Laser Technology Inc., Englewood, Colorado) for individual trees taller than 20-m. Several readings of the crown apex (i.e., highest foliage) were taken per individual from different locations (if possible) and trees were included in analyses if their measurement standard deviation was less than 1 m. The mean of the readings for an individual established the tree’s height. Only the > 20-m height class was considered because field measurements were taken 3-4 years after the FLI-MAP overflight, and rapid tree growth in smaller tree-height classes during this time would confound the analysis. 28 The heights of 21 trees in pasture with isolated crowns were measured in the field during July, 2001 (~ 9 species, Table 2.2). The crown center (centroid) for each pasture tree was estimated using DGPS readings from a Trimble Pro XL GPS and on-site base station (Trimble Navigation Limited, Sunnyvale, California) and subsequent visual spatial adjustment of the DGPS points with reference to the DCM. Between February and October 2000, the heights of 59 old-growth forest emergent trees were measured (11 species, Table 2.2) in the field and their trunks were geo-located relative to the closest grid marker using a tape measure and compass. Trunk locations were then transformed to UTM coordinates (Hofton et al., 2002) and crown centroids were identified through visual adjustment of trunk points with reference to the DCM. I selected canopy-emergent trees in old-growth forest for analysis because 1) they are major components of overall forest biomass (Clark & Clark, 1996), 2) they were easy to locate unequivocally in the DCM, and 3) their height growth in the 3 years between field measurement and the lidar overflight was expected to be minimal. 2.2.3. Plot-scale stem heights For plot-scale reference data, I used stem heights from a 1997 census of agroforestry plantations within La Selva (Menalled et al., 1998). The 32 plantation plots considered in this research were on 1-year (n=9), 4-year (n=9) and 16-year (n=14) cutting cycles and had stems spaced 2-m apart at roughly equal densities at the time of the census. Each plot had a total area of 0.04, 0.08, and 0.13 ha, respectively, and the oldest trees in a plot were 6 years old in 1997. All trees were 29 one of three species: Hyeronima alchorneoides (Euphorbiaceae), Cedrela odorata (Meliaceae), or Cordia alliodora (Borginaceae). Seven of the fourteen 16-yr rotation plots included a mixture of one of the three dominant tree species as well as a sub-canopy palm Euterpe oleracea (mean height 7.13 m) and a smaller understory shrub Heliconia imbricata (mean height 1.53 m). The 2-dimensional area of each field plot (i.e., a rectangle polygon) was digitized in a GIS and located with visual reference to the DCM. In the field, height measurements were recorded with a laser range-finder for every individual stem in the plot. Plot mean height (Table 2.2) was calculated in two ways: 1) as the average of heights for all stems in the plot (termed “all stems”), and 2) the average of heights for trees in the plot (termed “tree stems”). Note that the difference between these two metrics lies in the inclusion of a mix of canopy (tree) and sub-canopy (palm and shrub) stem heights in the plot mean-height calculation for 7 of the 32 plots; the mean height of the other 25 plots, which did not include the palms and shrubs, did not differ between the two methods. 2.2.4. Accuracy of field height measurements I established the vertical accuracy of the laser range-finder in two ways. In one case, five height estimates at 10, 20, 30 and 40-m distances from a 15.00-m pole in an open area were found to have an overall vertical mean error of +0.18 ± 0.14 SD m (slight overestimation). There was no significant effect on the mean height measured with distance (Kruskal-Wallis non-parametric ANOVA, α=0.5). Next I analyzed the stem height of actual trees. Crown apices were more difficult to 30 identify from the ground than the top of the 15-m pole. Four trees in open areas had range-finder height estimates made before being cut down. The actual height of the tree was then measured from its horizontal position on the ground, including the stump. The four trees had an average ground-measured height of 27.3 m and a range-finder estimate mean error of +1.45 ± 1.67 SD m (overestimation). In summary, reference measurements using the laser range-finder have their own associated error, mostly due to the human observer’s inability to identify the apex of the tree crown, rather than distance from the target. The overall bias appears to be an overestimation of tree height by the laser range-finder technique. 2.2.4. Small-footprint lidar data The small-footprint lidar dataset used in this research (FLI-MAP) was introduced in Chapter 1. Previous lidar research that used this dataset include analyses of LVIS pseudo-waveforms synthesized from the DSM cells (Blair & Hofton, 1999) and a comparison of LVIS to FLI-MAP sub-canopy elevation retrieval (Hofton et al., 2002). In the latter study, FLI-MAP elevation readings over areas of structurallycomplex vegetation (e.g., old-growth forest) were found to be mostly from canopy, not the ground, as is to be expected from a first-return sensor. However, in a 25-m diameter circular area, there were at least some FLI-MAP hits that coincided with survey elevations (Hofton et al., 2002), which may be due in part to sensor’s relatively high sampling density (Huising & Gomes Pereira, 1998). 31 2.2.5. Ground retrieval and DTM interpolation To create a DTM, FLI-MAP data processing involved two major steps: 1) identifying ground-return cells in the DSM, and 2) subsequent geostatistical interpolation of ground cells to form a DTM. I developed two simple groundretrieval algorithms that sought a balance between processing speed and the minimization of elevation error over the range of vegetation and terrain conditions found at La Selva. 2.2.6. Local-minima ground-retrieval scheme The local-minima algorithm proceeded as follows: a grid of non-overlapping, square cells was overlaid on top of the original DSM. Within each grid cell, one local-minima DSM cell (0.33-m support) was selected and identified as a ground return. This procedure resulted in a population of ground-return cells for each of the five grid scales considered independently: 5, 10, 15, 20 and 30 m. At a relatively coarse scale (e.g., a 25-m diameter circle), there is expected to be at least one lidar pulse that penetrates to or near the ground surface. This pulse would register as a cell of relatively low height in the DSM; and thus, the above local-minima scheme is analogous to selecting the lowest return in a square footprint of a specified scale (i.e., 5 m, 10 m, etc.). Ground-return cells identified at each scale were then used in separate geostatistical interpolation schemes (described below) that generated DTMs with a 1-m cell size. Samples from each DTM were compared to 3859 co-located reference points. The overall RMS errors of the resulting DTMs were used as the basis for the selection of a final ground-retrieval/interpolation scheme. 32 2.2.7. DTM interpolation schemes The inverse distance weighted (IDW) and ordinary kriging (OK) geostatistical techniques were assessed for the interpolation of the final DTM. In both procedures, DSM ground-return cells (0.33-m support) were treated as representing a cloud of xyz-coordinate points, with the cell’s centroid defining the x and y coordinates and the cell’s DSM height defining the z value. Note that these points have less variance than they would if derived directly from the original lidar footprints, instead of from the DSM. This is because DSM cell values were smoothed from TIN-interpolation and subsequent rasterization. IDW interpolation of ground points was conducted using Arc/Info 8.0.1 (Environmental Systems Research Institute, Redlands, California). The IDW interpolation moved in a circular window of 50-m radius and included at least 4 data points. The IDW power parameter specifies how sample points are weighted with distance from the interpolation node. A lower power relaxes the weighting of near points, causing more smoothing in the interpolated surface (Isaaks & Srivastava, 1989). Initial experiments based on minimizing RMSE identified that a value of 2 was the most appropriate IDW power for this data set. For OK, an isotropic normal-score transform variogram (Goovaerts, 1997) for each set of ground points (i.e., 5 m, 10 m, etc. scales) was manually fitted using Variowin v2.21 (Pannatier, 1996). These models contained a short- and long-range nested structure and a very small nugget effect (Goovaerts, 1997; Isaaks & Srivastava, 1989). Ordinary kriging of the normal-score data points was performed 33 using the GSLIB v2.0 software package (Deutsch & Journel, 1992) with a 250-m search radius, and the inclusion of 4 to 12 data points at each interpolation node. 2.2.8. Iterative-addition ground-retrieval scheme An additional ground-retrieval technique was tested that iteratively added in ground points at successively finer scales (Fig 3). Point selection proceeds through 3 iterations, each with 3 steps (in Fig. 2.3; Iterations 1-3 are rows and steps are columns). In the first step of the first iteration (Fig. 2.3; row 1, col. 1), a coarsescale grid (i.e., 20-m cells) is overlaid on the original DSM and one 0.33-m minima cell is selected from within each 20 x 20-m cell. In Step 2 (Fig.3; row 1, col. 2), DSM minima cells are converted to xyz points. In Step 3 (Fig. 2.3; row 1, col. 3), a first-pass DTM is created from IDW interpolation with a 5-m support. The first iteration is essentially the local-minima scheme as described above, but with a 5-m IDW interpolation. For Step 1 of Iterations 2 and 3 (Fig. 2.3; rows 2 & 3, col. 1), the coarse-scale grid is reduced by 5-m (i.e., from 20 to 15 m, or 15 to 10 m) and overlaid on the DSM; again, minima cells are identified, converted to xyz points, and then added to the population of minima points from the previous iteration. In Step 2 of Iteration 2 & 3 (Fig. 2.3; rows 2 & 3, col. 2), minima-point z values are subtracted from the DTM resulting from the previous iteration (rows 1 or 2, col. 3), and individual points 0.25-m higher than the DTM (marked with “+” in Fig. 2.3) are removed from the population of minima points (Fig. 2.3; rows 2 or 3, col. 2). In Iteration 2, Step 3 (Fig. 2.3; row 2, col. 3) another preliminary DTM is interpolated with IDW for use 34 in the third iteration. At the end of the Iteration 3, a final population of ground points is interpolated with IDW or Ordinary Kriging (OK) with a 1-m support (Fig. 2.3; row 3, col. 3). The effect on the mean-signed error of the final DTM using a 0.25, 0.50 and 1.0m residual threshold was evaluated; and subsequently, 0.25 m was deemed a conservative residual threshold in that it allowed finer-scale variation to be added back to the population of ground-return points, while it minimized the risk of including understory vegetation points (and subsequently inflating the mean-signed error). To minimize processing time, an IDW interpolator with 5-m support was chosen for generating intermediary DTMs (Fig. 2.3; rows 1 or 2 only, col. 3); future research should test the impact of using a finer-scale interpolation (i.e., 1 m) or an ordinary kriging interpolator on the accuracy of the final DTM product. 2.2.9. Sink removal Roughly 0.5% of the DSM-derived, xyz points from the ground-retrieval schemes contained very low, spurious points that created local sinks in the interpolated DTM. Some sinks were prevented from entering the population of ground points by imposing two restrictions on DSM local-minima cells: 1) that the minima cell not lie 60-m lower than its highest neighboring DSM cell (no trees or other objects in the scene are expected to be taller than 60-m high), and 2) the cell not lie lower than the minimum study site elevation (30 m). Sink xyz points that persisted through the ground-retrieval process were automatically removed by calculating their sink depth from a first-pass DTM interpolation. All sink points greater than 5-m in depth were 35 deleted from the population of ground points, and the DTM surface was reinterpolated. 2.2.10. Digital Terrain Model (DTM) accuracy assessment For each DTM interpolated, error was calculated by subtracting DTM elevation from the elevation measured at co-located field-survey points (DTM – reference; sensu Hodgson et al., 2003). Calculated error statistics included mean-signed error, mean absolute error (MAE), and RMSE (Isaaks & Srivastava, 1989). The final DTM with the lowest RMSE was submitted to a more rigorous analysis that sought to understand the variation of error across the TRF landscape. I asked the following questions: 1) Does DTM error differ among slope inclination categories in old-growth forest? 2) Does DTM error differ among land-use categories? As reviewed in Section 2.1.1, relatively dense and structurally-complex vegetation decreases DTM accuracy. Lidar range error also increases non-linearly with greater surface-inclination angle (Baltsavias, 1999a); and, when dense vegetation types occur in areas of steep terrain, these combined vertical errors decrease the accuracy of the interpolated surface over relatively short horizontal distances (Hodgson et al. 2003). The square-root transformation of the mean absolute error (SqrtMAE) was used in an ANOVA to test for statistical significance of the above hypotheses (sensu 36 Hodgson et al. 2003), with α=0.05. The square-root transformation of MAE was chosen to adjust the positively-skewed distribution of MAE to a normal distribution, as required by ANOVA. Since the minimum distance between reference points was 0.28 m, the classical ANOVA assumption of independence of residuals was suspect; therefore, I opted to use the generalized least-squares ANOVA with a residual variance-covariance weighting, formulated by Gotway and Cressie (1990). The weightings take into consideration residual autocorrelation and reduce chances of committing Type I errors. This technique requires a variance-covariance model of the SqrtMAE residuals (SqrtMAE minus a trend component). The trend component of the SqrtMAE points was estimated using ordinary kriging of the local mean (Goovaerts, 1997). An isotropic variogram of the resulting SqrtMAE residuals was modeled, and this model in conjunction with the data-to-data distances of the survey points were used to estimate the SqrtMAE residual variance-covariance sub-matrices for each class (e.g., Old-growth, Developed Areas, etc.), as required by the spatial ANOVA (Gotway and Cressie, 1990). 2.2.11. Stem height accuracy assessment A digital canopy model (DCM) was calculated by subtracting the final 1-m support DTM, bald-Earth surface from the lidar DSM (i.e., DCM = DSM – DTM, Fig. 2.2). The DCM maintained the 0.33-m support of the DSM. Individual tree and plot-scale stem heights were estimated from the DCM using metrics that had comparable spatial supports and calculations as those used in the field (sensu Drake, Dubayah, Knox et al., 2002; Næsset, 1997). 37 Plot-scale DCM metrics considered the same 2-dimensional area as field plots, i.e., same spatial support. In field measurements, plot mean height was calculated as the average height of stems spaced 2-m apart. Following Næsset (1997), I devised a comparable lidar metric that averaged the DCM local-maxima (i.e., maximum height) from each 2 x 2-m cell in a grid overlaying each plot. Alternatively, I also calculated the average of all heights (i.e., all DCM cells) in the plot extent. These two lidar metrics are referred to as Mean2x2 and MeanALL, respectively. Field measures of individual tree height located the highest leaf or twig in a crown (i.e., crown apex). Similarly, I based DCM metrics for estimating tree height on a sample of cells from within the crown. All DCM cells within 5 horizontal meters of the crown centroid were sampled and tree height was estimated as the maximum value of the cells (referred to as “Maximum”). Since crowns had multiple peaks of high foliage, field measures may have missed the highest crown apex. In consideration of this potential field error, I devised another metric that averaged DCM cells whose values were above the 95-percent quantile (sensu Ritchie et al., 1993; referred to as Mean95). At individual tree and plot scales of analysis, height error was calculated as the difference between the DCM and field height metrics (DCM – field, sensu Næsset, 1997). These errors, which are residuals from a 1:1 relationship, were summarized with the statistics mean-signed error, MAE and standard deviation (Isaaks & Srivastava, 1989). I also evaluated the linear relationship between lidar and field height metrics with regression analyses, which provided a coefficient of determination (r2) value 38 and a model RMSE (referred to as RMSEm). RMSEm was calculated from prediction residuals (sensu Drake, Dubayah, Clark et al., 2002; Næsset, 2002, Næsset & Økland 2002) and is the general error expected if the regression model were applied to DCM-derived estimates, i.e., calibration using ground measurements. Because I did not have an independent set of field data for model validation, model prediction residuals needed for calculating RMSEm were acquired through a common cross-validation procedure (Drake, Dubayah, Clark et al., 2002; Næsset, 2002; Næsset & Økland 2002; Popescu et al., 2003). 2.3. Results 2.3.1. DTM generation and accuracy assessment The five local-minima ground-retrieval scales tested in this research (i.e., 5, 10, 15, 20 and 30 m) had overall RMSE errors ranging from 2.29 to 5.09 m, using either IDW or OK for surface interpolation (data not shown). The scale with the lowest RMSE for both interpolation methods was found to be 20 m. For this tropical landscape and lidar sampling density, 20 m appears to be the near-optimum scale to identify ground returns with the local-minima approach, and so only the results from this scale will be discussed (Table 2.3). This optimal scale is likely determined by the average crown dimensions and canopy gap characteristics in old-growth forest, which comprises 69% of the study area. By visual interpretation of colorized 1-m IKONOS imagery, Clark, Read, et al. (2004) measured mean maximum crown diameter to be 19.6 m for all trees within old-growth plots. In this research, mean maximum crown diameter of the very largest, canopy-emergent trees in old-growth 39 forests were measured from the ground to be 27.2 m (n=36). At 15-m scales or finer, there was a higher probability that a DSM local-minima cell could lie completely within one large crown or many connected crowns. If these crowns are vertically dense, or have understory trees underneath, the DSM local minima would likely be from upper- or lower-canopy leaves or branches, not the ground. At 30-m scales or greater, there was a higher chance of selecting a canopy-gap DSM cell with a height from or near the ground; however, a trade-off exists in that local minima at coarser scales fail to sample fine-scale topography. As shown in Figure 2.4 (green points), the 20-m local-minima scheme identified DSM-cell minima (subsequently converted to xyz points) on the periphery of dense vegetation, such as in the shortest plantation plots, on the edges of emergent tree crowns and in more open swamps. The 20-m scale, iterative-addition scheme could identify more of these local-minima cells (xyz points), while still avoiding the areas of densest vegetation (Fig. 2.4, green and red points together). The population of xyz points for each ground-retrieval scheme were used to model variograms needed for OK interpolation. The normal-score variogram from the 20-m scale, iterative-addition dataset (method used for final DTM interpolation, Fig. 2.5a) was found to have a zero nugget effect, exponential short-range structure of 380-m range with a sill of 0.31, and an additional long-range spherical structure spanning to a 2000-m range, comprising an additional 0.20 of the total sill (maximum sill modeled in study extent was 0.51). Considering all 3859 survey points distributed throughout the landscape, the correlation (r) between DTM and reference elevation was +0.99 (p<0.0001) for IDW 40 interpolation and +1.00 (p<0.0001) for OK interpolation, using either groundretrieval scheme. In terms of RMSE, OK also performed better than IDW for DTM interpolation. There was a 0.18-m RMSE difference between the best IDW and OK interpolated surfaces (Table 2.3). The inclusion of finer-scale local-minima points using the iterative-addition scheme (i.e., iteratively adding local-minima from 20,15 to 10-m scales) improved the OK-interpolation RMSE by 0.10 m, resulting in an overall RMSE of 2.29 m (Fig. 2.5b; Table 2.3). All DTMs had a positive meansigned error, and so they tended to overestimate elevation. Although this overestimation was up to 0.87-m higher with OK relative to IDW interpolation, the mean absolute error (MAE) using OK was lower (Table 2.3). Indeed, the OK DTM was significantly more accurate than IDW interpolation, using either groundretrieval technique (n=3859, paired t-test of sqrtMAE, p<0.0001 for both comparisons). As expected, OK was found to reduce the variance of errors (Table 2.3, standard deviation). This smoothing of the variance across space tends to minimize the influence of spurious understory vegetation or downed trunks that are inevitably included in the DTM interpolation. In contrast, IDW does not exhibit this desirable smoothing effect as strongly as OK. The OK iterative-addition method was selected to generate the final DTM because the surface had a relatively low error variance and the lowest overall RMSE and MAE. This DTM was free of obvious underestimation (i.e., sinks) or overestimation (i.e., peaks) errors and revealed remarkable fine-scale detail of geomorphology (Fig. 2.2b). As expected, elevation error in the final DTM was not distributed evenly across the landscape. Mean absolute error (MAE) was significantly different between slope 41 inclination classes in old-growth forest (p<0.0001). Mean-signed error was positive (overestimation of DTM elevation) and increased 0.62 m from the lowest to the steepest slope classes (Fig. 2.6, +1.07-m [0-3°] vs. +1.69-m [>20°] mean-signed error). The difference in MAE between slopes 0-3 and 0-10 degrees was not significantly different (paired t-test; p=0.68); however, the MAE on slopes ≤ 10 degrees was significantly less than on slopes 10-20 and > 20 degrees (Fig. 2.6, lines connecting classes; p < 0.004 both comparisons). Under old-growth forest, RMS error was lowest on slopes ≤ 10 degrees (2.21 m, 0-3° & 3-10° combined) and highest on slopes greater than 20 degrees (3.09 m; Fig. 2.6), indicating that very steep slopes had the largest overestimation bias and error variance (Isaaks & Srivastava, 1989). I next assessed DTM elevation error relative to land-use. The slope error analysis indicated that slopes ≤ 10 degrees had statistically similar DTM errors, so I limited land-use analyses to survey points on slopes ≤ 10 degrees to avoid the confounding effects of slope error. For all land-use classes on slopes less than 10 degrees, the overall RMSE for the DTM was 1.72 m (Table 2.4). There was a highly significant difference in mean absolute error between the land-use classes (p<0.0001). The classes with relatively few trees or shrubs, Developed Areas and Abandoned Pastures, had a slight elevation underestimation (mean-signed error) of -0.58 and -0.28 m, respectively (Fig. 2.7; Table 2.4). As expected, DTM elevation was overestimated (positive mean-signed error) under forest canopies (Fig. 2.7; Table 2.4). DTM error was most severe in old-growth forests, which had extremely dense, multi-layered canopies. 42 Old-growth forests had significantly higher MAE (pair-comparisons with other classes, p<0.05), the largest mean-signed error (+1.01-m overestimation), and highest RMSE (1.95 m) relative to all other classes (Fig 7; Table 2.4). In contrast, the classes with very little to no canopy cover, Developed Areas and Abandoned Pastures, had the lowest RMS errors (1.02 and 1.10 m, respectively). Interestingly, classes with continuous canopies of relatively low tree height, Secondary forests and Agroforestry Plantations, had statistically similar absolute errors (Fig 7., lines). 2.3.2. Landscape view of canopy-surface heights Differences in land-use depicted in Figure 2.1 are clearly observable as mean height and textural variations in the DCM (Fig. 2.8). As expected, mean canopysurface height is greatest for old-growth, swamp and selectively-logged forests, which have relatively low human disturbance (Table 2.5). Several linear strips of low elevation running southwest to northeast indicate areas where no lidar footprints were recorded due to gaps in the flight-line (Fig. 2.8). These areas were avoided in tree-height analyses. 2.3.3. Individual- tree height estimation In estimating old-growth tree height, the Maximum lidar metric had a significantly lower mean absolute error than the Mean95 lidar metric (Table 2.6; p<0.001, t-test). The relative MAE error for Maximum and Mean95, respectively, was 8.1% and 8.7% of the mean height for individual emergent trees measured in the field (Tables 2.2 & 2.6). Mean-signed error was negative, indicating that the DCM 43 metrics underestimate actual emergent tree heights in old-growth forest. Linear regression models for predicting old-growth tree height explained 51% and 50% of the variance (RMSEm of 4.15 and 4.19 m) for Maximum and Mean95, respectively (Fig. 2.9 a & b). Lidar metric underestimation can be seen as points clustering above the 1:1-relationship line in the scatter plot (Figure 2.9 a & b). As with old-growth heights, pasture tree heights were underestimated by the DCM metrics (Table 2.6, negative mean-signed error). However, DCM height estimates of pasture trees had a lower MAE, with 2.33 and 2.84 m for Maximum and Mean95, respectively (Table 2.6). Mean absolute errors using Maximum and Mean95 were 7.4% and 9.0%, respectively, of the mean field height of pasture trees (Tables 2.2 & 2.6). For pasture trees, the Maximum lidar metric had a significantly lower mean absolute error than the Mean95 lidar metric (Table 2.6; p<0.001, t-test). Height errors were also less variable than those of old-growth trees (Table 2.6, standard deviations; Fig. 2.9, 1:1-relationship line). For pasture trees, the linear models relating lidar-derived to field-derived height explained 95% of the variance (using either lidar metric) and relationships were stronger than those observed for oldgrowth tree heights (Fig. 2.9 c & d vs. Fig. 2.9 a & b). Model RMS errors for pasture trees (2.41 and 2.48 m for Maximum and Mean95, respectively) were roughly half of those expected in applying old-growth height models (Fig. 2.9). In summary, direct estimation of individual tree height from the DCM was more accurate (i.e., less absolute error) when using the Maximum lidar metric, which calculated height as the maximum DCM cell within 5-m radius of the crown centroid. Furthermore, pasture height estimates using either DCM metric had less 44 absolute error and stronger linear relationships with field measurements than those from old-growth forest (Table 2.6; Fig. 2.9 a & c). 2.3.4. Plot-scale height estimation In terms of mean-signed error, MAE and standard deviation of errors, the MeanALL lidar metric out-performed the Mean2x2 method in estimating plot mean height, calculated from all or tree-only stems (Table 2.7). Considering tree-only stems, mean-signed error was slightly negative (-0.36 m) when using the mean of all DCM cells in the plot (i.e., MeanALL metric), indicating a slight underestimation of plot mean height. In contrast, I found that the Mean2x2 lidar metric overestimated mean stem height (+1.55 mean-signed error). Mean absolute error was significantly less for MeanALL relative to Mean2x2 when considering the plot mean height of all or tree-only stems (p<0.0001, paired t-tests). All plot-scale regression models were highly significant (p<0.001). Using the MeanALL lidar metric, the mean height of tree stems in plantation plots was predicted with a model r2 of 0.97 and RMSEm of 1.08 m (Fig. 2.10b—trees-only plots), whereas this relationship dropped to an r2 of 0.84 and RMSEm rose to 2.26 m when predicting mean height for all canopy and sub-canopy stems in plots (Fig. 2.10d—all stem plots). A similar pattern was observed in using the Mean2x2 lidar metric (Fig. 2.10 a & c). Height was overestimated for the 7 all-stem plots, and so those points fall below the regression line (Fig. 2.10 c & d—open circles) and decrease the overall strength of the models. In general, the best plot-scale relationship was between the MeanALL lidar metric and the trees-only field metric. Errors between 45 this combination of lidar and field metrics had the lowest 1:1-relationship bias (0.36-m mean-signed error), strongest linear relationship (r2 = 0.97) and lowest model RMSE (1.08 m). 2.4. Discussion 2.4.1. Lidar ground retrieval in a tropical landscape The overall 2.29-m RMSE of the DTM generated using an iterative groundretrieval and ordinary kriging interpolation scheme (Table 2.3) fell short of the decimeter accuracies reported for other laser sensors (Cobby et al., 2001; Huising & Pereira, 1998). However, those accuracies were achieved only under ideal conditions favoring retrieval of ground points, such as in areas with relatively flat terrain, without vegetation or with deciduous vegetation in a leaf-off state. In this tropical wet forest landscape, areas exhibiting these characteristics are rare. The “bald Earth” geomorphology in the study area is not flat due to its volcanic history and leaf-on canopy persists throughout the entire year. In comparing areas of similar slope (< 10 degrees), my DTM error is 0.79 m greater than the error (1.72 vs. 0.93-m RMSE) observed by Hodgson et al. (2003) when analyzing a lidar-derived DTM from a temperate-zone, deciduous (leaf-on) landscape. My research used a fullyautomated technique to retrieve ground xyz points from a DSM, which was interpolated from the original lidar footprints. In contrast, Hodgson and colleagues used proprietary software to identify ground-return footprints and then used a visualinterpretation step to remove spurious xyz points prior to DTM interpolation. Due to my first-return lidar data, DSM-interpolation smoothing, fully-automated ground 46 retrieval, and extremely-dense vegetation, I would expect my DTM to have relatively greater overall error than the Hodgson et al. (2003) study. Despite the many challenges in using lidar over a TRF landscape, my observed DTM error is well below the USGS maximum-permitted RMSE of 7 m for Level-3 DTMs, the highest-quality nationwide products publicly offered in the United States (Hodgson et al. 2003); and furthermore, this lidar-generated DTM is a vast improvement over previously-available topographic data at this important tropical research site. I found that DTM accuracy followed a gradient in human-disturbance intensity. The DTM had the greatest RMS error (1.95 m) in old-growth forests. The structure of these forests is composed of multi-layered leaves and branches maintained by a fine-scale, tree-fall disturbance regime. In effect, this structure is a dense media which acts to absorb or multi-scatter photons of near-infrared laser light, thereby reducing the probability that photons reach the ground and return to the sensor; consequentially, my ground-retrieval algorithm had a greater chance of mistaking sub-canopy returns for ground returns in old-growth forests, and the accuracy of the interpolated DTM decreased. In contrast, DTM accuracy was 0.93-m greater in developed areas of the reserve (1.02-m RMSE), which had scattered overstory trees and shrubs, mowed grass and cobble roads. Even under these more ideal groundretrieval conditions, the algorithm failed to entirely separate ground from overlying vegetation height. When I considered just those survey points in developed areas with mowed grass or roads, overstory vegetation > 3-m away, and on flat terrain, RMSE was 0.58 m and mean-signed-error was -0.49 ± 31sd m (n=20). This error can be considered the elevation error related to sensor artifacts when using first47 return laser pulses, i.e., footprint xyz positioning, scan angle (Baltsavias, 1999b; Huising & Gomes Pereira, 1998). A last-return, small-footprint system would likely provide better ground-retrieval performance in this densely-vegetated landscape. Indeed, last-return systems are generally deployed for DTM-generating campaigns (Hodgson et al., 2003; Lefsky et al., 2002) because there is a higher probability that laser returns will come from ground reflections. Flying a lidar sensor with higher sample density (i.e., more frequent postings) is also expected to increase the chances of detecting the ground (Hodgson et al., 2003). In structurally-complex old-growth forest, I found vertical errors to increase 0.67 m on the steepest slopes relative to flattest areas (Fig. 2.6, > 20 vs. 0-3-deg. class RMSE). In comparison, Hodgson et al. (2003) found a 2-m greater RMSE on flat versus steep slopes under structurally-complex, leaf-on vegetation (0-2 vs. 6-8 deg. classes). Observed DTM vertical error has both horizontal and vertical error components (Hodgson et al., 2003), such as footprint positioning and instrument range errors, respectively. As mentioned above, these factors combined may contribute 0.58-m to DTM elevation error. On steep slopes, vertical errors between DSM-derived ground points translate into large DTM interpolation errors relative to flat areas, which have more points of similar elevation that act to down-weight the impact of a spurious point during interpolation. Also, most DTM error analyses assume that errors in reference data are negligible. In this research, field-survey data suffer from horizontal, planimetric errors (~0.57 m) due to the transformation between the local and UTM WGS-84 coordinate and datum systems. On the steepest slopes (44 degrees [Table 2.1]), planimetric error could introduce up to 48 0.40-m vertical error in reference data. When I considered only reference points on slopes ≤ 10 degrees, RMS error decreased from 2.29 to 1.72 m (all land-use). This 0.57-m difference in DTM error can easily be explained by greater reference error combined with instrument-related vertical error on steep slopes. However, my data do not permit me to rigorously assess these differences in lidar- and referencerelated vertical errors. Compared to previously published accuracies achieved with the large-footprint LVIS sensor at LSBS (Hofton et al., 2002), FLI-MAP elevation estimation had better overall performance across the entire landscape (5.64-m [LVIS] vs. 2.29-m [FLI-MAP] RMSE). Also, the DTM from my research performed better than LVIS on flatter areas under the range of vegetation conditions in the study area (1.72-m [FLI-MAP, slopes ≤ 10º] RMSE vs. 2.42-m [LVIS, slopes ≤ 3º]). In the LVIS study, the estimated elevation at centroids of 25-m diameter footprints were compared directly to the nearest reference point, and so steep slopes compounded the vertical error associated with planimetric discrepancies between centroids and survey points. In contrast, the FLI-MAP DSM had a 0.33-m spatial support that permitted more precise planimetric location of ground-retrieved samples. In addition, fine spatial support provided denser ground sampling relative to the LVIS footprints. Due to these benefits of using a small-footprint sensor, my observed elevation error was relatively low compared to the Hofton et al. (2002) study, even on the steepest slopes and under the densest vegetation in the landscape (2.42 to 3.09-m RMSE from flat to steep slopes in old-growth forest). 49 As was found by Lloyd and Atkinson (2002b), ordinary kriging was a better DTM interpolator of lidar ground-retrieved samples than the more conventional IDW technique. Although the mean-signed error was smaller for IDW, OK interpolations had smaller mean absolute errors, lower RMS errors, and smaller error variance. The variance-dampening effect afforded by OK is desirable because it tends to reduce the impact of spurious sub-canopy vegetation samples that have inadvertently passed through the vegetation filter. There are other variants of kriging that could be useful to DTM interpolation of lidar data. For example, cokriging (Goovaerts, 1997) the ground samples with a co-varying variable, such as land-use type, optical reflectance data or a textural variable calculated from the original lidar data, could provide a robust means of adjusting the data covariance weighting to various landscape units. 2.4.2. Lidar estimation of tropical vegetation height Individual tree heights were underestimated by the lidar metrics (Table 2.6, negative mean-signed error). These results concur with those by Brandtberg et al., (2002), Gaveau and Hill (2003), and Persson et al. (2002). The FLI-MAP sensor used in my research detected first-return signals from densely-sampled small footprints, and so it is unlikely that this underestimation error is entirely from the laser missing the upper-most reaches of the crown (Næsset & Økland, 2002). Based on recent findings by Gaveau and Hill (2003) for leaf-on hardwood trees, I believe underestimation is partly caused by laser pulses penetrating below crown surfaces until inner-crown materials reflect a detectable first-return signal. 50 Some of the discrepancy between lidar and field estimates is also related to DTM error. In abandoned pastures, the DTM tends to slightly underestimate elevation by 0.28 m on average and RMS error was 1.10 m (Table 2.4). In contrast, the DTM error under old-growth forests was significantly greater. The DTM was overestimated by 1.01 m on average and RMS error increased to 1.95 m (Table 2.4). In abandoned pastures, broad areas without tree canopy permitted more laser energy to reach the ground and return to the sensor, resulting in better DTM accuracy. In old-growth forest, dense canopy causes DTM overestimation that effectively “clips” trees at their base when the DCM is calculated. In terms of RMSE, DTM-related error can account for up to 47% and 42% of the model error observed for old-growth and pasture trees, respectively (Fig. 2.10, Maximum). I found that DTM error increases on steep slopes in old-growth forests; however, slope was not linearly related to the absolute difference between lidar- and field-derived old-growth tree heights (Maximum metric [Fig. 2.9a], r2 = 0.003; p=0.68). I therefore conclude that slope-related effects on tree height estimates are not severe. Another source of random error in the predictive models is from field reference data (Brandtberg et al., 2003; Gaveau & Hill, 2003; Persson et al., 2002). In my research, field measurements of individual tree heights were taken 3 years after the lidar mission, and many trees could have grown higher in that time. This growth would result in the field estimates being higher than the lidar measurements, thus adding to the underestimation bias seen here. Although my field data included only trees with precise measurements (i.e., measurement std. dev. ≤1 m), high precision does not ensure absolute accuracy. In this research, I found that directly-measured 51 stem height was overestimated by the laser range-finder. Range-finder estimation precision was between 0.14 to 1.67 m (s.d. of errors). Again, this field measurement error would create an additional source of apparent underestimation in lidar-derived estimates and may account for up to 69% and 40% of model RMSE for pasture and old-growth tree heights, respectively (Fig. 2.9, Maximum). Given the difficulties in estimating elevation and field stem heights in oldgrowth forest, it is not surprising that regression models had a stronger linear relationship for pasture trees than old-growth trees (r2 0.95 vs. 0.51; Fig. 2.9 a & c, Maximum;). The strength of the relationship for pasture trees is greater than that observed by Brandtberg et al. (2003) in estimating deciduous trees in leaf-off conditions (r2 0.69), and the same as that observed by Gaveau and Hill (2003) in estimating leaf-on hardwood trees and shrubs (r2 0.95). All studies used the firstreturns from small-footprint lidar data to estimate vegetation height and observed an underestimation for tall trees. My research and the Brandtberg et al. (2003) study used the local-maxima of samples from within a crown, while Gaveau and Hill (2003) compared DCM cells directly to field points measured over crowns with high xyz-coordinate precision. The two-return lidar data used by the other two studies, as well as the leaf-off conditions in the Brandtberg et al. (2003) study, likely permitted better ground retrieval and higher DTM accuracy relative to that which can be expected with a first-return lidar flown over an evergreen TRF landscape. Relatively high DTM error in old-growth forests likely explains why old-growth TRF tree heights were not as reliably predicted as those of tropical pasture trees (my study) and temperate-zone hardwood trees (Gaveau & Hill, 2003). Despite the limitations 52 of my elevation and field-height estimates, my pasture tree calibration model was surprisingly strong relative to both the Brandtberg et al. (2003) and Gaveau and Hill (2003) models. Hardwood trees in leaf-on conditions were underestimated by -1.58 m and -2.12 (mean-signed error) in my study and the Gaveau and Hill (2003), respectively. Exposed leaves in hardwood trees favor the detection of the crown surface because they readily transmit and multiple-scatter near-infrared laser light (Grant, 1987). In contrast, the Brandtberg et al. (2003) model had weaker accuracy because leaf-off crowns expose only twigs and branches to the sensor and allow greater laser penetration into the canopy before returning a signal. Regression models for estimating individual conifer tree height had RMS errors of 0.23 and 0.63 m (r2 0.75 and ~0.98) in Næsset & Økland (2002) and Perrson et al. (2002), respectively. In my research, I observed a 2.41-m model RMSE for pasture trees (r2 0.95; Fig 9 c, Maximum) and Gaveau and Hill (2003) observed a 1.98-m model RMSE for deciduous hardwoods and shrubs (r2 0.95). The two conifer studies used first and last return, small-footprint lidar data. Assuming DTM error is equal between the studies, it thus appears that lidar calibration models (i.e., linearregression models) have a lower RMS error in estimating conifer tree heights than they do for leaf-on, hardwood trees. However, more comparative studies between tropical and temperate species, tree conditions (i.e., stress, senescence), lidar sensors and associated analytical methods are needed to confirm this accuracy assessment. The linear correlation between lidar and reference measurements of plot mean height of tree stems was very strong (Fig. 2.10 b). The 0.97 r2 exceeded the 0.380.49, 0.91, 0.82-0.95, and 0.91 r2 values reported for conifer-dominated stands by 53 Magnussen and Boudewyn (1998), Næsset (1997, 2002), and Næsset & Økland (2002), respectively, and the 0.68 r2 value for leaf-on hardwood stands (Lim et al., 2003) As was found in this and other lidar studies (Lim et al., 2003; Magnussen & Boudewyn, 1998; Næsset, 1997), mean tree height at the plot scale was slightly underestimated by the MeanALL lidar metric (Table 2.7). In my research, this underestimation (negative mean-signed error) was 0.36 m, while it was 2.1 to 4.1 m and 0.70 m in the Næsset (1997) and Magnussen and Boudewyn (1998) studies, respectively. At the stand scale, conifer underestimation likely results because too few footprints detect the upper-most twigs and branches of conical trees (Magnussen & Boudewyn, 1998). In my research, laser penetration into the hardwood tree canopy is likely an issue (Gaveau & Hall, 2003), and square DSM cells may not always record the height of the highest surface material within the cell—an underestimation that would persist in DCM heights. Furthermore, my field metric considered the mean heights of tree stems (i.e., maximum height of individual crowns), whereas the MeanALL lidar metric averaged all canopy-surface heights (i.e., DCM cells) in the plot, not just DCM crown-maxima, and this lidar measurement is expected to be low relative to the field measurement (Magnussen & Boudewyn, 1998). Although the Mean2x2 metric was designed to isolate DCM crown-maxima (i.e., stem heights of trees spaced 2-m apart), the metric overestimated mean tree height by an average of 1.55 m (Table 2.7). These results are contrary to those of Næsset (1997), who found that a local-maxima grid-overlay metric reduced mean-signed error to insignificant levels. However, the Næsset (1997) and other “grid-overlay” 54 studies have used last-return lidar systems with relatively sparse sampling densities, properties which combined lower the probability of recording a return from the upper-most canopy. This problem may have been compounded by the geometry of conifer crowns in the previous studies (Magnussen & Boudewyn, 1998; Næsset, 1997, 2002). In contrast to TRF tree crowns, which are broad and generally hemispherical in shape, conical crowns expose much less upper-most canopy material to a lidar sensor (Magnussen & Boudewyn, 1998). A grid-overlay lidar metric can compensate for missing the crown by selecting only the highest heights in the dataset. In the case of FLI-MAP, which has both a high sampling density and first-returns, there is less chance of missing the upper-canopy, especially over these agroforestry plots that contained even-aged stands with a relatively flat uppercanopy layer of broad leaves. In this case, the grid-overlay metric needlessly overcompensates the plot-height estimate upward relative to the reference height. Given the dense upper-canopy layer in the older plots, FLI-MAP first-return signals were not expected to be sensitive to sub-canopy stem heights. This subcanopy insensitivity was revealed clearly in linear-regression analyses. When considering all stems in plots, regression models had weaker relationships and higher model RMS errors than those built when considering just tree-stem heights (Fig. 2.10). The averaging of canopy and sub-canopy stem heights from the field lowered the mean height for seven of the reference plots, and since FLI-MAP was sensitive mainly to the upper-canopy trees, the lidar-metric estimation was high relative to the field measurement. 55 As was found by Næsset & Økland (2002) for conifer trees, the estimation of height for individual hardwood trees (Table 2.6) was not as accurate as that found for plot-scale estimation (Table 2.7). This finding is not surprising, because at the plot scale, both lidar and field measurement errors are minimized by averaging many co-located lidar and reference data points. 2.5. Conclusions In this chapter, I found that a small-footprint, first-return lidar sensor can be used to predict sub-canopy elevation with 2.29-m accuracy in a tropical landscape. The accuracy of the elevation surface was significantly affected by vegetation cover, with largest errors detected in areas with structurally-complex old-growth forests, especially on steep slopes. The digital terrain model that resulted from this analysis had a 1-m spatial support, which makes it an ideal input for other ecological or management applications such as modeling of inundation zones, hill-slope processes, and habitat associations. I showed that small-footprint lidar systems have potential for the estimation of individual tree height of tropical, hardwood species in leaf-on conditions. For oldgrowth forest emergent trees and isolated abandoned-pasture trees greater than 20-m tall, individual tree heights could be estimated directly from a lidar canopy-height surface (i.e., DCM) with mean absolute errors that were 8.1% and 7.4% of mean field heights, respectively. Models for individual old-growth and pasture trees explained 59% and 95% of the variance, with model RMS errors of 2.41-m and 4.15-m, respectively; however, as was found in other studies (Brandtberg et al., 56 2003; Persson et al., 2002), it was difficult to separate lidar-related error from reference measurement error. Plot-scale, mean stem height for trees within plantation stands was estimated from the DCM with much greater accuracy than for individual tree heights. The best plot-scale models explained 87% of the variance when estimating the mean height of all canopy and sub-canopy stems in plots, while 97% of the variance when considering just canopy-tree stems. These results are encouraging in that a stand (i.e., a plot in this study) is often an important scale for many broad-scale studies and management decisions. 57 Table 2.1. Topographic reference data. Count Min Max Elevation (m) 3859 38.78 141.37 Slope (deg) a 932 0.00 44.00 a Measured only in old-growth forest. Mean 70.93 12.94 Median 62.38 12.00 Table 2.2. Height reference data for individual trees and plots (units in meters). Count Min Max Mean Median S.D. Old-growth trees 59 31.00 56.39 45.04 44.34 5.76 Pasture trees 21 20.01 51.19 31.64 27.54 10.73 Plots (All stems) 32 0.38 17.53 6.19 4.70 5.27 Plots (Tree stems) 32 0.38 18.53 7.28 4.70 6.17 Table 2.3. Error of Digital terrain model (DTM) elevation estimates (n=3859, units in meters).a Interp.d (Ground retrievale) RMSE b MAE c Mean S.D. IDW (Local-minima) 2.47 1.78 +0.68 2.37 IDW (Iterative-addition) 2.47 1.75 +0.08 2.47 OK (Local-minima) 2.39 1.69 +1.10 2.13 OK (Iterative-addition) 2.29 1.60 |+0.97 2.07 d e Interp. (Ground retrieval ) Median Min Max IDW (Local-minima) 0.69 -14.80 17.64 IDW (Iterative-addition) 0.29 -16.29 16.54 OK (Local-minima) 0.94 -14.77 18.62 OK (Iterative-addition) 0.83 -17.14 16.89 a Error is calculated as DTM – field-survey elevation (deviation from 1:1 relationship). b RMSE is root mean-square error. c MAE is mean absolute error. d Interpolation schemes are: IDW = inverse distance weighted, or OK = ordinary kriging. e Ground-retrieval schemes are: Local-minima = minimum DSM cell in a 20x20-m grid, or Iterative-addition = iteratively adding local-minima from 20,15 to 10-m scales. 58 Table 2.4. DTM error on slopes ≤10 degrees summarized by land use (in meters).a Land-use Class Mean ± SD (RMSE) Developed Areas -0.53 ± 0.88 (1.02) Abandoned Pastures -0.28 ± 1.08 (1.10) Selectively-logged Forest +0.21 ± 1.61 (1.62) Secondary Forest +0.49 ± 1.36 (1.44) Agroforestry Plantations +0.62 ± 1.08 (1.24) Swamp Forest +0.72 ± 1.48 (1.64) Old-growth Forest +1.01 ± 1.66 (1.95) All classes +0.66 ± 1.59 (1.72) a DTM interpolation from OK, iterative-addition scheme. Table 2.5. Canopy-surface height summarized by land use.a Mean Land-use Class No. Cells Area (Ha) (m) S.D. (m) Abandoned Pastures 2,126,838 23.6 3.3 5.0 Developed Areas 826,562 9.2 8.0 8.2 Agroforestry Plantations 3,287,534 36.5 12.8 9.0 Secondary Forest 5,395,687 60.0 12.9 8.2 Old-growth Forest 43,675,481 485.3 20.5 8.8 Swamp Forest 4,210,094 46.8 20.5 11.1 Selectively-logged Forest 3,271,990 36.4 20.8 11.0 a Calculated by from the digital canopy model (DCM), 0.33-m cells. Table 2.6. Estimation error of individual tree heights.a Units are in meters. MAE b Mean S.D. Min Max c Old-growth Trees (n=59) Maximum d 3.67 -2.11 4.11 -5.02 11.59 d Mean95 3.94 -2.72 4.15 -4.40 12.10 Pasture Trees c (n=21) Maximum d 2.33 -1.58 2.40 -5.11 2.68 Mean95 d 2.84 -2.30 2.46 -5.64 2.35 a d c Error is calculated as: lidar – field height metric (deviation from 1:1 relationship) b MAE is mean absolute error. c Field metric is the maximum height in an individual crown (old-growth emergent or isolated pasture trees). d Lidar metrics are: Maximum = maximum value of DCM cells; Mean95 = mean of cells above 95% quantile. 59 Table 2.7. Estimation error of plot-mean stem heights.a Units are in meters. MAE b Mean S.D. Min Max Tree stems c (n=32) Mean2x2d 1.74 +1.55 1.55 -0.52 4.92 MeanALLd 0.90 -0.36 1.09 -3.69 1.60 All stems c (n=32) Mean2x2d 2.82 +2.63 2.78 -0.52 8.08 MeanALLd 1.64 +0.73 2.32 -3.69 5.91 a d c Error is calculated as: lidar – field height metric (deviation from 1:1 relationship) b MAE is mean absolute error c Field metrics are mean of individual stem heights for tree species only (Tree stems) or trees and other species (All stems). d Lidar metrics are: Mean2x2 = mean of maximum DCM values in a 2x2-m grid overlaid on plot; MeanALL = mean of all DCM values in plot 60 ! La Selva 0 0.5 1 2 Km Costa Rica Lidar data extent Rivers Land Use Developed Areas Selectively- logged Forest Old- growth Forest Secondary Forest Abandoned Pasture K Agroforestry Plantation Swamp Forest Figure 2.1. La Selva Biological Station study area land-use and lidar data extent. 61 A Lidar Height (m) 128.8 45.2 DSM B Elevation (m) 92.6 45.2 DTM C Vegetation Height (m) 48.6 0.0 DCM Figure 2.2. A 3-dimensional perspective of a 250 x 250-m subset of the lidar raster products, all covering the same geographic extent in an old-growth forest: a) the unprocessed lidar height surface (i.e., digital surface model, DSM), b) the estimated sub-canopy elevation surface (i.e., digital terrain model, DTM), and c) the estimated vegetation height surface (i.e., digital canopy model, DCM) resulting from the subtraction of the DTM (b) from the DSM (a). In (c), canopy emergent trees are redtone, concave domes. 62 Iteration 1 20 m 5-m IDW Iteration 2 15 m 5-m IDW Iteration 3 10 m or 1-m IDW Step 1. DSM grid overlay find 0.33-m DSM minima cell in each coarse-scale grid cell (e.g., 20 x 20 m) Step 2. Calculate elevation residuals (point-DTM) 1-m OK Step 3. Interpolate DTM from filtered xyz points remove xyz points (+) with residuals > 0.25 m Figure 2.3. Conceptual flow of the iterative-addition ground-retrieval scheme. Abbreviations are: digital surface model (DSM), digital terrain model (DTM), inverse distance weighed interpolation (IDW), and ordinary kriging interpolation (OK). Note: to create the figure, the same 60 x 60-m extent was placed on top of the actual DSM, and ground samples outside the extent were included in the DTM interpolation (Step 3, all iterations). 63 A Height: 0 B 47 m Height: 0 54 m Figure 2.4. An example of ground-retrieved samples (i.e., xyz points) for a) agroforestry plantations, and b) an old-growth forest with an open swamp. Points shown are for the 20-m local-minima retrieval scheme (green dots) and the iterativeaddition scheme spanning 20, 15 and 10-m scales (green and red dots combined). The underlying gray-scale surface is from the final DCM (Fig 2.2c; Fig. 2.8). Extent is 210 x 180 m. 64 A B 0.5 140 0.4 Survey Elevation (m) Semivariance 120 0.3 0.2 OK iterative-addition method n = 3859 r = 1.00 RMSE = 2.29 m 100 80 60 0.1 Model variogram Empirical variogram 40 0.0 0 200 400 600 800 1000 40 Lag Distance (m) 60 80 100 120 140 DTM Elevation (m) Figure 2.5. a) Empirical and modeled variograms from elevation points retrieved from the iterative-addition algorithm (20-m starting scale). b) Relationship between lidar-estimated ground elevation (sampled from the final DTM [Fig. 2.2b]) and fieldsurveyed elevation. 65 5 RMSE Mean-signed Error Observed Error (m) 4 187 3 323 138 284 2 1 0 0-3 3-10 10-20 20 + Slope Class (Degrees) Figure 2.6. Root-mean-square error (RMSE) and mean-signed error distribution by slope class within old-growth forest. Numbers above the bars indicate the number of reference points for each category. Lines connecting classes indicate groups of homogenous mean absolute error (α = 0.05; t-tests account for heteroscedasticity [Gotway & Cressie, 1990]). 66 RMSE Mean-signed Error Observed Error (m) 3 2 1 0 191 53 196 171 130 204 1115 -1 De vel Ab S Se Pla Sw Old c and elec tive onda ntatio amp F -grow on ed r n e l th ore y ys dP Are st ast logge Fores as ure t d op Figure 2.7. Root-mean-square error (RMSE) and mean-signed error distribution by land-use class for all samples with slopes ≤ 10 degrees. Numbers below the bars indicate the number of reference points for each category. Lines connecting classes indicate groups of homogenous mean absolute error (α = 0.05; t-tests account for heteroscedasticity [Gotway & Cressie, 1990]). 67 Vegetation Height (m) Plantations 60 Abandoned Pasture 0 Secondary Forest Old-growth Forest Swamp Forest ± 0 0.5 1 Km Figure 2.8. Landscape-scale overview of the final digital canopy model. 68 A B 60 60 Old-growth Emergent Trees n = 59 2 r = 0.51 RMSEm = 4.15 m 50 Tree Height (m) Tree Height (m) 50 40 30 Old-growth Emergent Trees n = 59 2 r = 0.50 RMSEm = 4.19 m 40 30 y = 8.64 + 0.85 x y = 8.87 + 0.85 x 20 20 20 30 40 50 60 20 40 50 Mean95 Lidar Height (m) C D 60 60 60 Isolated Pasture Trees n = 21 2 r = 0.95 RMSEm = 2.41 m 50 Tree Height (m) 50 Tree Height (m) 30 Maximum Lidar Height (m) 40 30 Isolated Pasture Trees n = 21 2 r = 0.95 RMSEm = 2.48 m 40 30 y = 4.15 + 0.91 x y = 4.79 + 0.91 x 20 20 20 30 40 50 60 20 Maximum Lidar Height (m) 30 40 50 60 Mean95 Lidar Height (m) Figure 2.9. Individual-tree height regression models for old-growth emergent trees (a & b) and isolated pasture trees (c & d). Reference data are the maximum height within a tree’s crown. Lidar metrics include Maximum (a & c) or Mean95 (b & d). Solid lines are the fitted models and dotted lines are the 1:1-relationships. RMSEm is the root-mean-square error of the model (i.e., cross-validation prediction residuals). 69 A B 20 20 Tree stems only n = 32 2 r = 0.96 RMSEm = 1.35 m 15 Plot Mean Height (m) Plot Mean Height (m) 15 Tree stems only n = 32 2 r = 0.97 RMSEm = 1.08 m 10 5 10 5 y = -0.41 + 0.87 x y = -0.07 + 1.06 x 0 0 0 5 10 15 20 0 10 15 Lidar MeanALL Height (m) C D 20 20 20 All stems n = 32 2 r = 0.87 RMSEm = 2.02 m All stems n = 32 2 r = 0.84 RMSEm = 2.26 m 15 Plot Mean Height (m) 15 Plot Mean Height (m) 5 Lidar Mean2x2 Height (m) 10 5 10 5 y = -0.06 + 0.7 x y = 0.38 + 0.84 x 0 0 0 5 10 15 20 0 Lidar Mean2x2 Height (m) 5 10 15 20 Lidar MeanALL Height (m) Figure 2.10. Plot mean height regression models. Reference data are the plot mean of stem heights calculated from only tree stems (a & b; filled circles) or from trees, sub-canopy palms and shrub stems (c & d; open circles). Plot-scale lidar metrics include Mean2x2 (a & c) or MeanALL (b & d). Solid lines are the fitted models and dotted lines are the 1:1-relationships. RMSEm is the root-mean-square error of the model (i.e., cross-validation prediction residuals). 70 CHAPTER 3: Discrimination of tree species at multiple scales 3.1. Introduction 3.1.1. Hyperspectral discrimination of tropical tree species Hand-held, airborne and spaceborne hyperspectral optical sensors measure spectral information in over 100 narrow bands spanning the visible (VIS=437-700 nm), near-infrared (NIR=700-1327 nm), and two shortwave-infrared (SWIR1=14671771 nm; SWIR2=1994-2435 nm) regions of the electromagnetic spectrum (region ranges adapted from Asner, 1998). It is anticipated that the automated classification of tropical species may be possible with hyperspectral imagery that is both fine enough to resolve individual tree crown (ITC) objects and also measures pertinent discriminatory spectral features from 400 to 2500 nm (Cochrane, 2000); however, this hypothesis has remained untested with an airborne or spaceborne hyperspectral sensor. In this chapter, field spectrometer and high spatial resolution hyperspectral data offer an unprecedented opportunity to explore the spatial-scale dependency of spectral reflectance in the remote identification of tree species. Below I briefly discuss important factors that influence plant reflectance at various spatial scales and that may affect the automatic discrimination of tree species. 3.1.2. Reflectance properties of vegetation at leaf, branch and crown scales Leaf-scale reflectance spectra are controlled by 1) leaf biochemical properties (e.g., water, photosynthetic pigments, structural carbohydrates), which create wavelength-specific absorption features, and 2) leaf morphology (e.g., cell-wall 71 thickness, air spaces, cuticle wax), which affects photon scattering (Asner, 1998, Grant, 1987; Roberts et al., 2004; Woolley, 1971). VIS spectral variability among species is low due to strong absorption by chlorophyll (Cochrane 2000; Poorter et al, 1995). High NIR transmittance and reflectance result from photon scattering within leaf air-cell wall interfaces, such as in spongy mesophyll (Gausman, 1985; Grant, 1987; Woolley, 1971). In SWIR1 and SWIR2, water absorption tends to obscure other absorption features produced by biochemical constituents (e.g., lignin and cellulose) (Asner, 1998; Gausman, 1985). Branch-scale spectra, such as from a high resolution pixel (e.g., < 4 m) or measured in situ with a hand-held spectrometer, are a mixture of radiance determined by the proportion, physical arrangement, and reflective and transmittive properties of crown tissues, including leaves, stems, branches, fruits, and flowers. Photon multiple-scattering among these components will tend to increase the expression of leaf biochemical absorption features, especially within crowns with large, densely-distributed and/or horizontally-oriented leaves (Asner, 1998). Finescale shadows cast within the branch may depress overall reflectance. Relative to leaf scales, these factors are known to increase branch-scale spectral variability and enhance separability of northern-latitude conifer and broadleaf trees (Roberts et al., 2004). Fung and colleagues (1998) used laboratory-derived, branch-scale hyperspectral data (400 to 900 nm, 90 evenly-spaced bands) and a linear discriminant classifier to discriminate 12 subtropical tree species. An overall accuracy of 84% was achieved and individual species Producer’s accuracies ranged from 56 to 100%. Species spectral separability was attributed to the effect of leaf72 size variation expressed at the branch scale. Gong et al. (1997) found that a neural network classifier applied to sunlit first-derivative spectra (6-8 cm spatial resolution, in situ) could classify 6 conifer species with an average overall accuracy of 91%. At the crown scale, the three-dimensional architectural arrangement of foliage and non-photosynthetic components determines the amount of photon volumetricscattering and attenuation within the crown (Asner, 1998). van Aardt and Wynne (2001) have shown that the VIS, NIR and SWIR1 regions are useful for discriminating species of temperate forest conifer and hardwood species when using in situ crown-scale hyperspectral data (sunlit sides of crowns). Spectral derivatives provided the best overall classification accuracies, which were 84% for conifer species and 93% for hardwood species. Cochrane (2000) provides the only investigation of TRF crown-scale hyperspectral data for automated species recognition (350-1050 nm data). The study used laboratory spectra from 11 tree species to simulate branch and crown scales. Target species discrimination was possible at crown scales, while it deteriorated at branch and leaf scales. Crown-scale spectra were best separated in the VIS-NIR transition (i.e., the “red edge”) and NIR regions. However, because the analysis used simulated branch and crown scale spectra, it is not known how non-photosynthetic vegetation or volumetric crown scattering will affect tree species spectral separability. 3.1.3. Challenges to tree species discrimination in tropical rain forests Tropical rain forests pose challenging obstacles to ITC classification. TRF tree communities are characterized by high diversity and relative rarity of individuals, so 73 large image extents are needed to find representative training samples. Many trees occur below a dense overstory canopy, preventing their detection by a passive optical sensor. In lowland tropical forests, a relatively constant growing season fosters a diversity of phenological traits, and leaf flush and flowering may follow annual or irregular cycles with no overriding community-scale patterns (Newstrom et al., 1994). Therefore, strategically-timed over-flights to capture spectrallyimportant phenological events (sensu Key et al., 2001) may be done for only a few tree species that have well-characterized phenology. Moreover, leaf-turnover and flower display may be asynchronous among and within individual crowns of the same species, thereby increasing conspecific variability in leaf- to crown-scale spectra. For example, long-lived leaves within a crown may be covered with epiphylls, which combined with leaf necrosis decrease VIS and increase NIR reflectance (Roberts, Nelson et al., 1998). Depending on the density of leaves in a crown, which may vary in time, radiance from understory shrubs, sub-canopy trees, lianas, bark lichens, canopy soil, and epiphytes may mix with a target species radiance and increase conspecific spectral variability. It is not yet clear whether these spectral components will increase branch- and crown-scale within-species variation to a level that inhibits among-species spectral discrimination (Castro-Esau et al., 2004; Cochrane, 2000). 3.1.4. Objectives In this chapter, I examine the relative trade-offs between spectral features, spatial scale of measurement, and classification schemes for the automated classification of 74 individual TRF tree species using their reflectance properties. Field spectrometer and airborne hyperspectral reflectance spectra (161 bands, 437-2434 nm) were acquired from seven species of emergent trees in a lowland tropical rain forest, permitting analyses at leaf, pixel and crown scales. My main objectives were to: • Determine if spectral variation among TRF tree species (interspecific) is greater than spectral variation within species (intraspecific), thereby permitting spectral-based species discrimination. • Identify the spatial scale and spectral regions that provide optimal discrimination among TRF emergent tree species. • Develop an analytical procedure for the species-level (floristic) classification of individual tree crowns using their reflectance spectra. • Assess the relative importance of narrowband hyperspectral versus broadband multispectral information for species identification of TRF trees. 3.2. Methods 3.2.1. Data sets and pre-processing 3.2.1.1. Canopy-emergent trees To select my study species, I conducted field surveys and took advantage of a Geographic Information System (GIS) database of tree locations from a long-term tree demography study at LSBS (the TREES project; Clark et al., 1998). My preliminary analysis involved 544 individual trees belonging to 27 species and led 75 me to focus my efforts on seven species (Table 3.1) of canopy emergents for which there were sufficient individuals in the hyperspectral imagery for a representative sample. Emergent trees with large, exposed crowns provided a large sample of pixels that were less influenced by spectral shadowing or scattering by neighboring trees, and they were easy to locate in the orthorectified hyperspectral imagery. Furthermore, five of the seven study species (BAEL, DIPA, HYME, HYAL and LEAM) are under analysis in the TREES project, providing opportunities to generalize local-scale research (e.g., demographic changes) to broader spatial scales using remote sensing. A total of 214 individuals of the seven study species were identified in the hyperspectral imagery through field surveys conducted between January 2000 and July 2001 (Table 3.2; Fig. 3.1). The trunk coordinates in the LSBS grid system (see Chapter 2) were surveyed by measuring the distance and angle of the trunk from the nearest grid tube. These trunk coordinates were then converted from LSBS grid coordinates to the UTM projection, WGS-84 datum coordinate system using a leastsquares affine transformation with RMSE of 4.8 m (OTS, unpublished data). There is little long-term data on leaf and flowering phenology of the study species. Some overstory tree species are deciduous and completely drop and flush leaves, generally beginning in the first dry season, while others are evergreen and continuously flush small amounts of leaves throughout the year (Table 3.1). Hyperspectral imagery was acquired on March 30, 1998, at the end of the first dry season, and all study trees were expected to have high mature leaf cover (i.e., high leaf area index) except DIPA and LEAM (personal observation, April 12, 2004; 76 literature data [Frankie et al., 1974; O’Brien, 2001]; summarized in Table 3.1). Although I do not have field observations from my study individuals during the image acquisition, O’Brien (2001) estimated leaf cover of BAEL, DIPA, HYME and LEAM individuals at LSBS that were 30-60 cm diameter above buttress and unobstructed or emergent crowns. Data included March through April, 1998 and showed that a relatively large proportion of DIPA and LEAM individuals had low mature leaf cover, while BAEL and HYME individuals had higher mature leaf cover. An example Balizia crown is shown in Figure 3.2a. 3.2.1.2. Leaf-scale spectra A shotgun was used to shoot 152 leaf samples down from crowns of individual study trees in August, 2002 (Table 3.3). Three to five individual trees from each of the seven study species were selected for sampling, and 2-10 leaves per individual were shot down from the upper, sun-exposed part of the crown. Leaf samples included a range of maturity and health. Individual leaflets > 1 cm width were sampled from separate leaves for species with compound leaves (CEPE, DIPA, HYME) and leaflets were analyzed as leaves. For scale considerations discussed below, BAEL compound leaves were analyzed rather than individual leaflets. Bidirectional reflectance properties of the “leaves” (i.e., leaves or leaflets) were measured in a darkroom at LSBS. Leaf samples were put in a plastic bag with a moist paper towel and stored in a cooler with ice until refrigerated in the laboratory. All samples were measured within 12 hours of collection. A single 150-W halogen lamp was placed with a 25-degree incident angle and 53 cm above a matte-black, 77 5%-reflective box. An ASD FieldSpec spectrometer (Analytical Spectral Devices, Boulder, CO, USA) sensor with an 8° fore-optic was positioned 7.1 cm at nadir above the box center, yielding a 1-cm sensor field of view (FOV). The spectrometer was optimized with a white Spectralon® panel (Labsphere, North Sutton, NH, USA) placed in the box center, and the instrument was re-optimized using the panel after measuring every 5 to 7 leaf samples. Leaf samples were placed in the box center with adaxial (upper) surfaces to the sensor and radiance was measured 5 times per leaf. Bidirectional reflectance of a single leaf sample will vary across its surface depending on biochemical variation (e.g., chlorophyll concentration, leaf necrosis, epiphyll cover), structural properties (e.g., cuticle texture, mesophyll depth), and illumination and sensor geometry. To capture this potential spectral variation from an individual leaf, leaf orientation and position relative to the sensor FOV were varied with each of the 5 radiance measurements (i.e., the leaf was moved while the sensor remained stationary). Radiance spectra from the Spectralon® panel were used as a standard to convert leaf radiance to percent reflectance. The final leaf reflectance spectrum was an average of the five reflectance spectra from each leaf. Individual leaflets of BAEL (Balizia) leaves were smaller than the sensor FOV, thus causing the black background to mix with the radiance signal. To counteract this effect, each Balizia leaf was stacked on top of 3 other Balizia leaves to simulate a dense layer of leaves. The leaf-stack position and orientation was haphazardly varied with each radiance measurement. 78 3.2.1.3. Hyperspectral image pre-processing Hyperspectral imagery came from the HYDICE sensor, which is described in Chapter 1. HYDICE runs were delivered as 16-bit calibrated radiance data. The morning data acquisition avoided afternoon cloud cover yet the 56.3° to 48.4° solar zenith angles (92° to 94° azimuth angles) during the flight caused deep tree shadows that are particularly noticeable in old-growth forest canopy gaps (Fig. 3.2b). Individual tree crowns are clearly resolved in this high spatial and spectral resolution imagery (Fig. 3.2b). I orthorectified the LSBS sections of HYDICE runs 6, 9, 12 and 15 using the Erdas IMAGINE OrthoBASE software package (Leica Geosystems GIS & Mapping, LCC, Atlanta, GA, USA). Runs were segmented into 800-m long blocks and each block was orthorectified with 21 to 75 ground control points collected by visually matching emergent tree crown centers in HYDICE imagery to co-located crown centers in a 0.3-m lidar digital canopy model (DCM; Chapter 2). Terrain distortions in the imagery were corrected in the orthorectification processing with a 10-m resolution digital terrain model (DTM: OTS, unpublished data), which was originally derived from Laser Vegetation Imaging Sensor (LVIS) lidar data (Rocchio, 2000). Orthorectification used a nearest-neighbor interpolator and georegistered the HYDICE imagery in the Universal Transverse Mercator (UTM), WGS-84 datum projection of the DCM and DTM reference data (Fig. 3.1 shows spatial extent of runs). The ACORN v4.0 (Analytical Imaging and Geophysics LLC, Boulder, Colorado) atmospheric correction package was used for calibrating radiance values 79 to surface reflectance. Although atmospheric water vapor can be calculated on a per-pixel basis (Gao and Goetz, 1990), low signal-to-noise in principal water absorption bands for HYDICE (Basedow et al., 1995) produced considerable spatial error in water vapor estimates; and therefore, atmospheric corrections were performed with a fixed precipitable water vapor of 32 mm. A tropical atmospheric model was used with atmospheric visibility of 100 km. Water vapor and visibility parameters were established based on visual assessment of old-growth tree spectra and an empirical, minimum root-mean square error (RMSE) comparison with fieldcollected spectra. Field spectra were measured in August, 2002 with an ASD FieldSpec spectrometer and included gravel road, cement, tile, exposed soil, wood planks, green metal roof-tops, and mowed lawn targets that were located within the HYDICE runs. An 8° fore-optic was positioned about 1.5 m above a target and sensor radiance was converted to reflectance using an in situ white Spectralon® calibration panel. Five individual reflectance measurements were averaged to create a target spectrum over a 1 m2 area, and then several of these spectra were collected over a homogenous area of the target and then averaged. Wavelength calibration differed among runs by 0.61 to 2.67 nm per wavelength (HYDICE metadata, SITAC). Atmospheric correction was performed using each run’s respective band centers and full-width half-maximum (FWHM) parameters. A common set of band center wavelengths were calculated by averaging bands centers from the four runs, and reflectance values from each run were then linearlyinterpolated to this common set of center wavelengths. Band centers were spaced an average distance of 6 nm in VIS, 14 nm in NIR, 12 nm in the SWIR1 and 9 nm in 80 SWIR2. Post-calibration reflectance artifacts (e.g., spikes near water absorption features) were minimized with a 3-channel box-car filter. Bands with extreme noise in spectral regions less than 437 nm and greater than 2435 nm, as well as bands in the strong water absorption features 1313-1466 nm and 1771-1994 nm, were removed from analyses. An example comparison between final HYDICE reflectance and field-measured ASD reflectance for a wooden-plank suspension bridge and a nearby tree crown (Pentaclethra macrophylla) is shown in Fig. 3.3. The HYDICE reflectance spectra were generally the same shape, but reflectance was much lower than field spectrometer measurements in the SWIR1 and SWIR2 regions. This pattern was observed in comparing HYDICE with other field spectra. NIR reflectance peaks were high and water absorption features centered at 980 nm and 1200 nm were deep relative to field spectra, especially for the wooden bridge. These artifacts in HYDICE derive from a combination of poor radiometric calibration, sensor noise, atmospheric noise (e.g., water vapor absorption) and the difference in time between HYDICE and field measurements. Field measurements were taken on August 2, 2002 (9:30 am) while HYDICE was acquired on March 30, 1998 (8 am). If atmospheric conditions were constant over the reserve, all HYDICE artifacts should be common to all HYDICE spectra because the same parameters were used to convert each pixel to reflectance. The mismatch between HYDICE reflectance and expected reflectance affects my analysis in three ways: 1) in comparing HYDICE spectra to laboratory leaf spectra, 2) possibly shifting band selection towards bands 81 with higher signal to noise, and 3) limiting the comparison of my results with other sensors. 3.2.1.4. Simulated broadband, multispectral imagery HYDICE reflectance spectra were convolved using sensor-specific spectral response functions to simulate IKONOS, Landsat ETM+, and ASTER (Advanced Spaceborne Thermal Emission and Reflection Radiometer) imagery. Although each of these sensors has a different spatial resolution (i.e., 4 m IKONOS, 30 m ETM+, 15-30 m ASTER), the spatial resolution of simulated imagery was fixed at the 1.6 m of HYDICE imagery. Also, by using simulated imagery, the same artifacts evident in HYDICE were incorporated into simulated spectra. Therefore using simulated multispectral imagery reduced the effects of spatial scale and radiometric artifacts on inter-sensor comparisons. 3.2.1.5. Pixel- and crown-scale spectra from individual tree crowns Field-surveyed trunk locations were overlaid on the orthorectified HYDICE mosaic and the polygons representing the 2-dimensional area of the tree crowns were manually digitized over the imagery. I used the DCM as a visual aid to determine a crown’s shape in areas with shaded pixels. The average crown area for the study species was 444 m2, with each crown comprising from 41-662 pixels (Table 3.2). Hereafter I refer to the digitized crown polygons as individual tree crowns (ITCs: Gougeon & Leckie, 2003). 82 My analyses differentiated between all (PixelALL) and sunlit-only (PixelSUN) pixels within each ITC. Sunlit pixel spectra were designated as all pixels within an ITC that had reflectance greater than or equal to the crown’s mean 800-nm (NIR) reflectance (Gougeon, 1995). Crown-scale spectra were calculated by averaging either all (CrownALL) or sunlit-only (CrownSUN) pixel spectra within each ITC. 3.2.2. Data analysis 3.2.2.1. Testing of within and among species spectral variability Spectral separability of species should be optimal if different species have high statistical distance in feature space and within-species variation is less than amongspecies variation. I tested the null hypothesis that within- and among-species spectral variation are equal with a non-parametric multivariate analysis of variance technique (NPMANOVA) first developed for use with ecological distance matrices (Anderson, 2001; McArdle & Anderson, 2001). In remote sensing applications, the spectral angle is a metric used for comparing the degree of similarity between two spectra (Kruse et al., 1993). Unlike Euclidean distance, the spectral angle is insensitive to linearly-scaled differences among spectra such as those caused by illumination. In my implementation of the NPMANOVA, the distance between each spectrum to every other within- and among-species spectrum was calculated using the spectral angle and Euclidean distance, and distances were stored in N x N distance matrices (N = number of observations). In the calculation of spectral distance, spectra were analyzed using the entire 161-bands or limited to the VIS, NIR, SWIR1, SWIR2 regions. A pseudo-F statistic was calculated as the ratio of 83 among to within species sums of squares (Anderson, 2001; McArdle & Anderson, 2001). The null hypothesis tested was that within and among species spectral variation was equal, which would make the F-ratio close to one. The significance of the F-ratio was tested against a null distribution of F created by 5000 random permutations of the distance matrix (Anderson, 2001). For pixel-scale NPMANOVA tests, 200 pixels for each species were randomly selected from crowns for the respective species. For leaf- and crown-scale NPMANOVA tests, I used all available spectra due to limited sample sizes. I performed NPMANOVA tests using the DISTLM2 v.5 software program (Anderson, 2004). 3.2.2.2. Species classification schemes I explored three popular supervised classification schemes for TRF tree classification: spectral angle mapper (SAM), linear discriminant analysis (LDA), and the maximum likelihood (ML) classifier. SAM is a spectral matching technique (Kruse et al., 1993). The spectral angles between each sample spectrum and several reference spectra are calculated to reduce the hyperspectral data cube from an ndimensional spectral space to a similarity space with dimensions equal to the number of reference spectra (i.e., classes). SAM classification was accomplished by assigning each sample spectrum to the class with the closest similarity (i.e., lowest spectral angle), and no maximum-angle threshold was used to minimize false detections. LDA is a common classifier that has been used in previous ITC classification research (Gong et al., 1997; Fung et al., 1998; van Aardt & Wynne, 2001). For LDA classification, the pooled within-class covariance matrix and 84 predictor variables (e.g., reflectance values) from training samples are used to build classification equations, or discriminant functions for each class (Duda & Hart, 1973; Tabachnick & Fidell, 1989). A class is chosen based on the highest a posteriori probability calculated from the functions. The most important assumption of LDA classification is that all classes share the same covariance matrix (i.e., homogeneity). In the ML classifier, each class mean, standard deviation and covariance matrix are estimated from the training data to evaluate a sample’s class a posteriori membership probability (Duda & Hart, 1973). ML has been widely used in ITC species classification (Gougeon, 1995; Key et al., 2001; Leckie, Gougeon, Hill et al., 2003; Meyer et al., 1996). Supervised classification schemes are often stymied by the large dimensionality of hyperspectral imagery. Fine resolution spectral bands are often correlated and so represent redundant information. Also, sensor noise such as stripes from bad detectors or atmospheric attenuation may be greater in certain bands and this noise may increase class variance and decrease class separability. With the ML classifier in particular, it has been shown that the within-class covariance matrix can be poorly estimated when there are few training samples relative to the data dimensionality, leading to a decrease in classifier performance called the Hughes phenomenon (Duda & Hart, 1973; Jackson & Landgrebe, 2001). Hence, the use of ML is limited for hyperspectral remote sensing of forested areas because image dimensionality is high while training data are expensive or difficult to acquire. A common solution to this dilemma is to reduce data dimensionality through spectral feature (i.e., band) selection. In this chapter, I used a forward stepwise selection method based on 85 discriminant analysis (Tabachnick & Fidell, 1989; van Aardt & Wynne, 2001). This method was implemented using the SAS STEPDISC procedure (SAS Institute Inc., Cary, NC, USA) with the significance criteria set at α = 0.05 for all analyses except the crown scale, which had criteria set to α =0.20. Following feature selection, SAM, LDA and ML classifiers were applied to leaf-, pixel- and crown-scale spectra to assess how the spatial scale of spectral measurements affects species classification accuracy. At each scale, the ndimensional spectral space was varied to include the full-spectra dataset (161 bands), spectral regions (i.e., VIS, NIR), or LDA stepwise-selected bands. Spectral regions were sub-sampled to include only 10 bands per region. These bands were evenlyspaced with an average spacing of 23 nm (VIS), 55 nm (NIR), 25 nm (SWIR1), and 47 nm (SWIR2). The same set of classifiers was also applied to the simulated broadband multispectral data. All LDA classification was accomplished using the “MASS” package in the R statistical environment (R Development Core Team, 2004; MASS 7.2-12, R v2.0) while SAM and ML classification was performed in ENVI v4.1 and IDL v6.1 (RSI, Inc., Boulder, CO, USA). 3.2.2.3. Crown-scale and “pixel-majority” ITC classification A major objective of this chapter was to assess optical remote sensing for operational, ITC species discrimination. Current research shows that tree species discrimination is best accomplished by aggregating pixels into their respective crowns for object-based (as opposed to pixel-based) classification using spectral and spatial properties (Gougeon, 1995; Leckie, Gougeon, Hill et al., 2003; Meyer et al., 86 1996). In this chapter, the species of ITCs in the HYDICE imagery were determined by 1) the class assigned from crown-scale spectra, CrownALL or CrownSUN and, 2) taking the majority value of the classified pixels (PixelALL or PixelSUN) within each ITC, i.e., the “winner-takes-all” rule (Meyer et al., 1996). I refer to this latter approach (2) as “pixel-majority” ITC classification. 3.2.2.4. Classifier training and accuracy assessment For pixel-scale classifications, 300 randomly-selected training pixels were sampled from the crown objects for each species (300 pixels x 7 species = 2100 training pixels). Pixels were sampled from the whole crown (PixelALL) or from sunlit regions of the crown (PixelSUN), and each crown was sampled unless it had fewer than 40 pixels. Each classifier (LDA, ML or SAM) was applied to the remaining non-training pixels within each crown. I sampled 300 training pixels per species to provide a robust estimation of ML class covariance statistics. For pixelscale testing, 300 non-training classified pixels per species were randomly selected (300 pixels x 7 species = 2100 test pixels). A new set of 300 test pixels were randomly selected for each classifier-band combination analyzed. For crown-scale and pixel-majority analyses, classification training and testing were performed with cross-validation due to the limited number of ITCs (Duda & Hart, 1973; Krzanowski, 2001). I sequentially left one ITC out and trained classifiers with pixel- or crown-scale spectra from the remaining 213 ITCs (i.e., “leave-one-out” cross validation). For pixel-majority classification, each withheld crown was classified based on the majority-class rule. Cross validation provides a 87 slightly biased estimate of true classifier accuracy (Krzanowski, 2001). Statistical differences among classifications were tested with the Z statistic calculated from the Kappa statistic and variance (Congalton, 1991). Leaf-scale classification was also performed with a similar cross-validation procedure. 3.3. Results 3.3.1. Reflectance properties at different scales 3.3.1.1. Leaf-scale spectra Leaf-scale spectra for the seven tree species (Fig. 3.4) showed typical patterns of vegetation: low VIS reflectance caused by absorption by chlorophyll and other pigments, high NIR reflectance due to multiple-scattering within the leaf structure, weak NIR water absorption features at 980 and 1200 nm, and moderate reflectance in SWIR1 and SWIR2 with peaks at 1650 and 2200 nm caused by dominant water absorption features at 1400, 1900 and 2700 nm (Gausman, 1985; Roberts et al., 2004). There was considerable variation in reflectance within species, especially in the NIR and SWIR (Fig. 3.4). Several factors can cause leaf spectral variation within a given species, including epiphyll cover, herbivory, necrosis, maturation of the mesophyll, and the concentration of chlorophyll and water. Seven percent of all upper-canopy leaves sampled had epiphyll coverage. As seen for HYME leaves of roughly the same age (Fig. 3.5a), epiphyll coverage tends to lower the green peak and NIR reflectance, possibly due to more absorption of light by epiphylls covering the adaxial leaf surface. Herbivory is another factor that affected 4% of the leaves 88 sampled. For LEAM leaves of the same age, some leaves had light brown-colored leaf mines caused by an insect. As the percentage of these mines increased, there was less VIS absorption (higher reflectance) likely due to lower amounts of photosynthetic pigments from eaten leaf material, and increased NIR and SWIR reflectance (Fig. 3.5b) likely due to lower water content and more exposed nonphotosynthetic leaf material (e.g., residual, dried leaf veins). Finally, leaf age is an important factor because it determines time exposed to epiphylls and herbivory, as well as internal leaf architecture and chemical properties. For TEOB, young thin leaves had high red and low NIR, SWIR1 and SWIR2 reflectance relative to mature, thicker leaves (Fig. 3.5c). Thin leaves are compact and have fewer air-cell wall refractive discontinuities than mature leaves, causing lower NIR-SWIR reflectance (Gausman, 1985). Also, lower chlorophyll content in young leaves likely accounts for higher VIS reflectance (i.e., lower VIS absorption) in the blue (450 nm) and red (680 nm) regions (Woolley, 1971; Gausman, 1985). As the leaf senesces, lower concentrations of chlorophyll greatly reduce the amount of absorption throughout the VIS, thereby increasing reflectance (Fig. 3.5c). 3.3.1.2. Pixel-scale spectra Pixel-scale spectra revealed the same general patterns of NIR scattering and chlorophyll and water absorption as seen in leaf-scale spectra (Fig. 3.6). Relative to leaf-scale spectra, there was an overall reduction (darkening) of percent reflectance in PixelALL and PixelSUN spectra (Table 3.4). Darkening of spectra is partly due to fine-scale shadows within branches of leaves and other crown materials, especially 89 in the PixelALL samples. However, some of the darkening in the SWIR1 and SWIR2 regions of pixel spectra was due to poor HYDICE radiometric calibration. Roberts et al. (2004) found that biochemical absorption properties of leaves were accentuated at the pixel scale by the multiple-scattering of photons among leaves and other crown tissues within the hyperspectral sensor’s FOV. HYDICE spectra show evidence of this phenomenon; photon scattering can partially explain why NIR water absorption features at 980 nm and 1200 nm were deeper in pixels relative to leafscale samples (Fig. 3.6). The arrangement and density of crown tissues will govern the crown scattering environment and the degree to which leaf biochemical properties are accentuated at pixel or crown scales. Considering sunlit pixels (PixelSUN), DIPA and LEAM had 40.2% and 37.9% NIR reflectance, respectively, while the other species had between 43.8% (CEPE) to 51.5% (TEOB) mean reflectance (Fig. 3.6). Relatively low NIR reflectance makes DIPA and LEAM appear purple in the image (Fig. 3.2b, RGB 1651 nm: 835 nm: 661 nm). Individuals of these deciduous species had low crown foliage density (i.e., leaf area index, LAI) during HYDICE image acquisition in the late dry season. With fewer leaves in the crown, there was less photon scattering and subsequently, NIR reflectance was low relative to leaf-on species. The variation in scattering environments among species with different crown LAI, explains why the NIR standard deviation was much higher in pixel scales than in leaf scales (Table 3.4). 90 3.3.1.3. Crown-scale spectra Crown scale spectra were an average of pixel spectra. The averaging of spectra decreased CrownALL and CrownSUN variance in all spectral regions relative to PixelALL and PixelSUN variance, respectively (Table 3.4, Fig. 3.7). As observed with pixel scales, the mixture of pixels from bright and dark, shadowed parts of the crown tended to lower average reflectance for CrownALL spectra relative to CrownSUN spectra. 3.3.2. Among- and within-species spectral distances Using spectral angle as a distance metric, among-species (interspecific) spectral variability was significantly greater (p≤0.001) than within-species (conspecific) variability for all spectral regions at leaf and pixel scales (Table 3.5). However, species differences at the crown scale were mainly focused in the NIR region. Using Euclidean distance, which includes variation due to illumination, among-species variability was significantly greater (P≤0.001) than within-species variability at all scales and all spectral regions. The greater separation of species with Euclidean distance over spectral angle distance at crown scales indicates that crown-level illumination differences among species, possibly due to varying crown LAI, tend to increase species separability. There was no advantage to using just sunlit pixels at crown scales (CrownALL vs. CrownSUN). However, at pixel scales sunlit samples had greater separability (i.e., higher F-ratios) than when considering all samples. The NIR, SWIR1 and SWIR2 regions of sunlit pixels had particularly high F-ratios. 91 3.3.3. Selected spectral features 3.3.3.1. Leaf-scale For discriminating species, 90% of the 10 most important wavelengths selected by the stepwise procedure were concentrated in the NIR and SWIR1 regions (Fig. 3.8a; Fig. 3.9a), where there was relatively large variation in percent reflectance (Table 3.4). Balizia leaves had the highest NIR variability (6.7% s.d. for BAEL, 3.3 to 5.5% s.d. for other species). BAEL variability is likely caused by measuring their reflectance from leaf stacks. Photons have more opportunity for scattering within stacks of leaves, thereby increasing the NIR plateau, broadening water absorption features, and increasing overall NIR variability. SWIR2 had lower variability than NIR or SWIR1 yet SWIR2 comprised 25% of the bands selected when considering 20 bands (Fig. 3.9b). The VIS had the lowest spectral variability, and only 10% of the bands selected were from VIS when considering 10 or 20 bands. The selected VIS bands were in the blue absorption feature at 449 nm, the green peak at 568 nm, and the red-edge at 719 nm (Fig. 3.8a). 3.3.3.2. Pixel-scale Leaf-scale band selection is best compared to the PixelSUN samples because they have a similar range of illumination. Of the 20 most important bands, there were 4 more NIR and 4 fewer SWIR1 bands selected for PixelSUN relative to leaf-scale spectra (Fig. 3.8a-b). In terms of the percentage of bands selected (Fig. 3.9a & b) and NPMANOVA F-ratios (Table 3.5), the NIR region was particularly useful in pixel-scale species discrimination, especially with sunlit samples. Selected NIR 92 bands were concentrated in the red-edge and the plateaus surrounding the water absorption features at 980 and 1200 nm (Fig. 3.8b), while less so at leaf scales (Fig. 3.8a). There was high NIR variability relative to other regions (Table 3.4), which is likely caused by species differences in crown LAI. Low-LAI deciduous species (e.g., DIPA, LEAM) had lower NIR reflectance relative to high-LAI species (e.g., HYAL, TEOB). These differences in crown architecture among species thus create distinctive variation in maximum NIR reflectance that permits clearer NIR species discrimination with pixel spectra relative to leaf spectra, which are not influenced by crown architecture. SWIR1 may be less useful at pixel scales relative to leaf scales due to a combination of greater water absorption and lower sensor signal to noise. When considering the best 20 discriminatory bands, NIR was less important when all sunlit and shaded pixels were analyzed, likely due to less photon scattering in shaded pixels (Fig. 3.8b; Fig. 3.9b). With PixelALL samples, there were a similar number of bands in the VIS, NIR and SWIR1 regions. VIS bands were spread across blue-green edge (491 nm), yellow edge (575 and 619 nm) and red well (670 nm) features, while SWIR1 bands were evenly spaced across the region. 3.3.3.3. Crown-scale With the stepwise band selection procedure, only 42 and 41 bands were significant at the α=0.05 level for CrownALL and CrownSUN spectra, respectively. The significance criteria was substantially relaxed to α=0.2 in order to select 60 bands for comparison with leaf and pixel scales. The twenty most important bands (all at α=0.05) were concentrated in the VIS, NIR and SWIR2 regions for CrownALL 93 spectra (Fig. 3.8c; Fig. 3.9b). For CrownSUN spectra, there were 2 fewer bands in VIS and 2 more bands in SWIR2 relative to CrownALL spectra. As with the pixelscale, NIR bands were clustered in the red edge and on the peaks bordering the water absorption features. 3.3.4. Classification of tree species at leaf, pixel and crown scales 3.3.4.1. Hyperspectral, narrowband classification The overall accuracies of tree species classifications with different narrowband combinations and classification schemes are presented in Table 3.6. Leaf-scale maximum likelihood (ML) analysis was limited to 10 bands due to the low number of training samples per class (Table 3.3). Linear discriminant analysis (LDA) had the highest accuracy (89.5%) for leaves when the analysis was isolated to the 10 best bands, while ML with 10 bands was 8.6% less accurate (Z=6.66, p≤0.05). Balizia was the only species that had no inter-class confusion with the LDA classifier and 10+ bands nor with LDA and NIR or SWIR1 bands, which can be attributed to its distinct spectral leaf-stack properties. LDA accuracy was significantly higher with 20 bands (Z=3.15; p≤0.05) and overall accuracy reached 100% with 40 bands (no significant difference between 20 and 30 to 60 bands). Both band selection and NPMANOVA identified the SWIR1 region as important to species separability at leaf scales, while the VIS region was not as important (Fig. 3.9, Table 3.5). In agreement with these findings, 10 SWIR1 bands provided the highest classification accuracy of the spectral regions while 10 VIS bands produced relatively low accuracy (Table 3.6, LDA or ML). Relative to LDA and ML 94 classifiers, the Spectral Angle Mapper (SAM) classifier had relatively poor performance (< 51% overall accuracy) and differences among band combinations will not be discussed. At the pixel scale, ML generally had higher overall accuracy than LDA for most band combinations, using all or sunlit-only pixels (Table 3.6). SAM had very low performance, with no accuracy exceeding 50%. In the ML and LDA analyses, PixelSUN classifications were significantly more accurate than when using PixelALL samples (Table 3.6, arrows). Best pixel-scale performance was with 40-60 bands from PixelSUN samples with either classifier (differences not significant). ML classification with 161 bands (full-spectra) was significantly lower than when using 20 to 60 bands due to the Hughes phenomenon, while full spectra information did not dramatically change LDA accuracy. Classification accuracies for both LDA and ML were significantly higher with 10 bands from across the entire spectra relative to selecting 10 bands from specific spectral regions. The LDA classifier applied to crown-scale spectra produced some of the highest species classification accuracies using airborne HYDICE imagery. For all band combinations, LDA accuracies were not significantly greater with CrownALL versus CrownSUN samples. LDA classification accuracy using 10 optimally-selected bands was significantly greater than using all 161 bands or 10 evenly-spaced bands in the VIS, NIR, SWIR1 and SWIR2 spectral regions. The best accuracy was 92.1% when the LDA classifier was applied to 30 CrownALL stepwise-selected bands (Tables 3.6 & 3.7). However, 20 bands provided only 2.8% less overall accuracy and were not significantly different than 30 bands. 95 Of the 30 bands used in the CrownALL classification, 30.0% were within the NIR region while 23.3% were in each of the other 3 spectral regions. Forty-one percent of misclassified ITCs involved confusion between DIPA and LEAM. This result was expected because both species had similar reflectance properties due to low-LAI crowns. Although it was not possible to compare LDA and ML classifiers for data dimensionalities > 10 bands due to limited training data, the cross-validation accuracies between LDA and ML with 10-bands were significantly higher for LDA—7.5% and 7.0% higher with CrownALL or CrownSUN, respectively. As with other scales of analysis, SAM crown-scale classification accuracy was relatively low. The highest SAM accuracy achieved was only 53.7%. In all LDA and ML classification analyses, there were no minimum a posteriori probability criteria set for assigning samples to a class; and therefore, there were no unclassified samples. The criteria was omitted to permit a direct comparison of LDA and ML with the SAM classifier. For crown-scale LDA classifications, I also experimented with 50%, 75% and 90% probability thresholds, where a crown-scale spectrum (i.e., ITC) was classified as “unknown” if its maximum class a posteriori probability was lower than the specified threshold. As the probability threshold was increased (i.e., made more conservative), more ITCs were labeled as unclassified and the overall accuracy dropped (Fig. 3.10). The decrease in overall accuracy was less severe as more bands were added to the analysis. With 30 bands, overall classification accuracy was 8.9% significantly higher with no probability threshold relative to a 90% threshold (Tables 3.7 & 3.8). A higher probability threshold acted 96 to increase omission errors by switching correctly-classified ITCs to “unknown”, thereby decreasing the class Producer’s accuracies. In this example, 19 correctlyclassified ITCs did not meet the threshold criteria and so were left unclassified. On the other hand, a more conservative threshold decreased commission errors, thereby increasing class User’s accuracies. For example, the 90% threshold identified 3 ITCs that had been confused between DIPA and LEAM due to low a posteriori probabilities. With the threshold set, these crowns were left unclassified and User’s accuracy increased 2.5% and 11.3% for DIPA and LEAM, respectively (Tables 3.7 & 3.8). In my classification analyses, only 7 of over 400 tree species were classified. In an operational classification, a probability threshold or other technique will be necessary to ensure that ITCs that are not target species remain unclassified. I tested the crown-scale, threshold classifier against non-target ITCs. A total of 30 emergent crowns comprising 14 non-target species were identified in the field and digitized over the imagery, following methods used for the 7 study species. I then classified these non-target ITCs with the 30-band, LDA classifier and a 90% a posteriori probability threshold. The classifier was only trained with the 7 study species (214 ITCs). Five of the non-target ITCs were actually classified as “unknown,” whereas 17 were classified as HYAL, 6 as DIPA, 1 as BAEL and 1 as CEPE. Misclassifications were not random. Crowns that were misclassified as DIPA had very similar spectral properties as true DIPA crowns—appearing as purple ITCs in false-color imagery (e.g., Fig. 3.2b)—suggesting that they had low-LAI crowns. In contrast, ITCs misclassified as HYAL had spectral properties of the high-LAI 97 crowns typical of true HYAL crowns. The classifier thus appears most attuned to differentiating ITC phenology and structure rather than clear species distinctions. 3.3.4.2. Multispectral, broadband classification Classification accuracies from simulated broadband sensors (Table 3.9) followed the same general patterns across scales as discussed for narrowband data: LDA and ML classifiers outperformed SAM; there was an increase in accuracy with more bands; and, LDA and ML accuracies were generally higher at crown scales relative to pixel scales. As with narrowband analyses, the LDA classifier was particularly strong at the crown scale. For CrownALL ASTER spectra (9 bands), overall accuracy was 76.2%, which was 7.9% and 14.0% lower (both significant) than the accuracies achieved with 10 and 30 narrow bands, respectively (Table 3.6). With CrownALL LDA analyses, there was a non-significant 7% increase in overall accuracies with 6 ETM+ bands over 4 IKONOS bands, while there were 9.8% and 16.8% significant increases with 9 ASTER bands over IKONOS and ETM+ bands, respectively. 3.3.5. Pixel-majority classification using within-crown pixels 3.3.5.1. Hyperspectral, narrowband classification ITCs were next assigned a species label based on the majority class of pixelscale, narrowband spectra within each crown object (Table 3.6, pixel-majority). Pixel-majority ITC classification accuracy was not significantly different using PixelALL or PixelSUN within-crown spectra. The best overall pixel-majority accuracies with each classifier-band combination were generally higher than those 98 achieved with comparable pixel-scale classifications, except for LDA with sunlit pixels and 40, 60 or 161 bands (Table 3.6). The highest pixel-majority accuracy, 85.5%, was achieved with a LDA classifier applied to 30 bands selected from PixelSUN spectra (Table 3.10), although accuracy was not significantly less with 10 and 20 band combinations, nor with PixelALL spectra. For the 30-band LDA classification (PixelSUN), I assessed the percentage of within-crown classified pixels that had the same species label as their corresponding ITC. If a within-crown pixel and its corresponding ITC species labels were the same, the pixel was considered “correctly classified”. If pixels were randomly classified, then each crown would likely have only 14% (1 out of 7) of its pixels correctly classified, and ITCs would be assigned to the class that had a pixel majority by chance. I found that the mean percentage of correctly-classified pixels within correctly-labeled ITCs was 89.9% (range 40.0% to 100.0%). Therefore, although there were misclassified pixels within ITCs, I am confident that crowns were not assigned a correct label by chance. For mislabeled ITCs, the mean of correctly-classified pixels within each crown was 13.7% (range 0.0% to 47.8%) This indicates that there was substantial spectral variation among individual crowns of a single species—some crowns had very poor pixel classification accuracy, while others had very high accuracy. For example, TEOB crowns were all correctly labeled and individual crowns had an average 91.6% accuracy. In contrast, only 57% of LEAM crowns were correctly-labeled, and correct crowns only averaged 78.9% pixel accuracy. As expected, mislabeled LEAM crowns were mainly dominated by DIPA pixels, the other low-LAI species. 99 As recommended by Meyer et al. (1996), I next set a 35% pixel-majority threshold for determining the class of an ITC; that is, a crown was labeled unclassified if the majority class comprised less than 35% of within-crown pixels. With the threshold set and a LDA classifier applied to 30 PixelSUN bands, only 2 ITCs were affected. One ITC was a HYME that had been misclassified as a BAEL, while the other ITC was a BAEL that had been misclassified as a DIPA. Since no correctly-labeled ITCs were affected by the threshold, overall accuracy remained at 85.5%. However, the threshold helped decreased the commission error for BAEL and DIPA by 2.3% and 0.9%, respectively, and so the threshold appears useful for filtering ITCs with low accuracies. 3.3.5.2. Multispectral, broadband classification Pixel-majority LDA and ML classification accuracies were significantly greater for 10 to 30 narrowband, HYDICE imagery than for simulated IKONOS and ETM broadband imagery, which had 4 and 6 bands, respectively (Table 3.9). Overall accuracy with ASTER data (9 bands) and an ML classifier was 78.0% (PixelSUN), which was not significantly lower than the pixel-majority accuracies achieved using 10 to 30 HYDICE bands with either the LDA or ML classifiers. 3.4. Discussion 3.4.1. Spatial scale and the spectral classification of TRF tree species There was a decrease in classification accuracy from fine to coarser scales of spectral measurement (i.e., leaf to pixel and crown scales). Some of this trend can 100 be explained by the differences in sensors used. Leaf-scale spectra had a relatively high ratio of signal to noise because they were measured in a controlled laboratory environment with a well-calibrated, high spectral resolution instrument. In contrast, pixel- and crown-scale spectra had considerably more noise due to poor sensor radiometric calibration and atmospheric effects. Leaf spectral variability among individuals of a certain species, or even within a single crown, was attributed to differences in internal leaf structure and biochemistry (e.g., water, chlorophyll content, epiphyll cover and herbivory). Leaves have nonLambertian properties and physical differences in adaxial leaf cuticle (e.g., microtopography, wax, leaf hairs) affect first-surface specular reflectance, especially in the VIS region with large incident and/or view angles (Grant, 1987). Another source of variability among leaves of the same species was thus introduced by measuring laboratory bidirectional (as opposed to hemispherical) reflectance with varying leaf orientations. Despite the multiple factors causing spectral variation, I found that leaf spectral variability among species was significantly greater than that within species. Crossvalidation classifications confirmed that leaf-scale reflectance could discriminate among species with >89% overall accuracy using as few as 10 optimally-positioned bands. Important bands were concentrated in the NIR and SWIR1, where diffusereflectance dominates and variability is largely controlled by internal leaf structure and water content (Gausman, 1985; Grant, 1987). Pixel-scale measurements acquired with the airborne hyperspectral sensor were dominated by leaf-scale spectral properties. Water absorption was enhanced at this 101 coarser scale by multiple-scattering of photons among leaves, stems and branches (Asner, 1998; Roberts et al., 2004). In the NIR region, high levels of multiplescattering caused the 980 and 1200 nm water absorption features to deepen, and overall NIR variability increased. The band-selection scheme for pixel spectra identified important bands bordering the NIR water absorption features, possibly detecting species differences in photon scattering caused by fine-scale crown architecture (e.g., LAI). Bands in the visible part of the spectrum were also important at pixel scales when considering all sunlit and shaded samples. The bluegreen edge, yellow edge and red well bands chosen may be sensitive to species differences in spectral properties caused by their relative spectral mixing of leaf and bark fractions; however, my data do not allow me to test this hypothesis. I originally hypothesized that isolating sunlit regions of crowns for pixel-scale analysis would lower within-species spectral variance and enhance species separability. This hypothesis was confirmed by larger sunlit-sample F-ratios with spectral angle and Euclidean distance metrics (Table 3.5). LDA and ML classifications also showed significant improvements in accuracy with PixelSUN over PixelALL samples (Table 3.6). Gong et al. (1997) found similar results when classifying conifer spectra acquired at 6-8 cm spatial scales. At the crown scale, the architectural arrangement of crown components, such as leaves and branches, controls the relative amounts of shading and complex, anisotropic multiple-scattering of photons relative to the illumination and view geometry—described by each crown’s bidirectional reflectance distribution function (BRDF). Crown-scale spectra in my research were a linear average of within-crown 102 pixel spectra. The averaging of shadowed and bright pixels lowered mean crownscale reflectance, and as is expected by averaging, crown-scale variance was low relative to pixel-scale variance. Band selection indicated that the NIR and SWIR2 were the main regions producing crown-scale separability among species. As with pixel scale spectra, NIR reflectance is largely controlled by structural properties (e.g., density and arrangement of leaves) that influence the photon scattering environment and subsequent NIR reflectance. SWIR2 variability among species may be related to two factors: overall differences in crown water concentration that affects the expression of water absorption features at 1400, 1900 and 2700 nm (Gausman, 1985; Roberts et al., 2004); and, ligno-cellulose absorption features that may be expressed when high fractions of non-photosynthetic woody tissues are exposed to the sensor (Asner, 1998; Curran, 1989), such as in a low-LAI deciduous crown. Other methods for capitalizing on species-level differences caused by crown structure and their influence on high spatial resolution imagery are discussed in Section 3.4.2. The SAM classifier was the least successful of the classifiers, regardless of the spatial scale or spectral region considered. This result was surprising since there were highly significant statistical separations of species with the spectral angle metric at leaf and pixel scales (Table 3.5). SAM does not use second-order statistics (e.g., covariance), and basing a classification on a single distance metric appears ineffective given within-species spectral diversity. LDA was highly accurate at all scales of analysis, indicating that spectral covariance information—pooled for all species—is important for species 103 discrimination. The leaf-scale and crown-scale ML classifiers may not have performed as well as LDA because of two factors. For one, the bands selected by the stepwise procedure were optimized for LDA classification. Other band-selection techniques, such as using Bhattacharyya distance (Haertel & Landgrebe, 1999), may improve ML classification accuracy. Furthermore, ML requires a large number of training samples for adequate estimation of the class covariance matrices from higher-dimensional data that may contain redundant and noisy information. Only with pixel-scale and pixel-majority classifications did I have a large enough sample size to adequately assess ML with high dimensional data (i.e., 20+ bands). Although ML performed slightly better at pixel scales relative to LDA, pixel-majority classifications were generally higher with LDA. The apparent advantage of the ML classifier at pixel scales may be spurious because only subsets of pixels from ITCs were used for training and testing in the analyses, while pixel-majority classifications used all ITC pixels through cross-validation. The crown-scale ML classifications suffered from a lack of samples (individual trees) when estimating class covariance matrices. Collecting large training sets is challenging in TRF because target species have low densities, thus requiring large field surveys of forest that is often difficult to access. The LDA classifier appears more appropriate for TRF species discrimination because it strengthens the covariance estimation by pooling information from all species. Results from my pixel-scale LDA classification analysis can be compared to a study by Fung et al. (1998), who used laboratory measurements of branch spectra to classify 12 subtropical tree species with 84% overall accuracy (Producer’s 104 accuracies from 56 to 100%). LDA and 90 bands from VIS to NIR were used in the analysis. In my analysis of sunlit pixels (i.e., branch scale) and 60 bands (VIS to SWIR2 sampled), overall classification accuracy was 85% and Producer’s accuracies ranged from 74% to 95%. Overall accuracy did not increase when using the fullspectrum of 161 bands. These results are encouraging since my airborne data suffers from multiple factors that could confound species discrimination, such as mixed pixels in training and testing data, variable illumination and viewing geometry, and noise introduced by atmospheric conditions and non-target biological organisms (e.g., lianas). At all scales of observation, I noted an increase in accuracy with increased data dimensionality to a certain level. Results using the LDA and ML classifier revealed a general increase in accuracy up to 30 bands, while including more bands yielded equal or lower accuracy. My results confirm the benefits of hyperspectral over multispectral data for TRF tree identification. At all scales, the best accuracies with hyperspectral data were higher than those achieved with simulated multispectral imagery. Here I have applied fairly conventional analytical techniques that select optimal bands and then apply classifiers to narrowband reflectance values. Band selection was a necessary analytical step to isolate the most important bands for reliable classifier parameter estimation given my training data limitations (Duda & Hart, 1973). However, one major advantage of contiguous hyperspectral bands is their continuous description of spectral space, allowing measurements of the shape and position of key spectral features, such as liquid water absorption features in NIR. Purely hyperspectral 105 analytical techniques exist and include spectral shape filters (Cochrane, 2000) and analyses of spectral first- and second-order derivatives. First-order derivatives have been shown to improve tree species classifications over the use of reflectance spectra (Gong et al., 1997; van Aardt & Wynne, 2001). However, current research has relied upon data from portable spectrometers in anticipation of high spatial and spectral resolution data from future airborne or spaceborne sensors. Abiotic and biotic noise, such as atmospheric water vapor, epiphytes, and lianas, will complicate the radiance from a TRF canopy acquired by an airborne or satellite sensor, and much research is needed to test hyperspectral-based classification techniques under these challenging conditions. 3.4.2. Classification of individual tree crowns Classifications of ITCs using crown-scale spectra had relatively high accuracies. The maximum accuracy achieved was 92% with the LDA classifier and 30 bands. Producer’s accuracies ranged from 70% (CEPE) to 100% (TEOB), and User’s accuracies ranged from 81% (LEAM) to 100% (TEOB). My overall classification accuracy is higher than the 65% accuracy Leckie and Gougeon (1999) observed for temperate hardwood classification with crown-scale spectra, and it is similar to the 93% accuracy achieved by van Aardt and Wynne (2001) using in situ crown-scale spectra to classify 3 hardwood tree species (second derivatives used, sunlit samples). As reported in studies with conifer trees (Gougeon, 1995; Leckie, Gougeon, Hill et al., 2003), there was no evidence that sunlit crown spectra could be more accurately classified than by averaging the spectra from all pixels within the crown. My results 106 are encouraging considering that with visual interpretation of tropical tree species, Clément and Guellec (1974) could only identify their target species with 73% accuracy, and Myers and Benson (1981) visually-interpreted only 22% of their species with >75% accuracy. I therefore conclude that spectral-based classification of TRF tree species is possible, and accuracy is comparable or potentially greater than from visual interpretation of aerial photographs. Furthermore, computer-based classification permits the automation and removal of subjectivity from the process. In the crown-scale LDA classification, there was inter-species confusion between individuals of Lecythis and Dipteryx (LEAM and DIPA, Table 3.7). This confusion is attributed to the deciduous phenology of these species—individuals had very low crown LAI and similar spectral properties. Bark lichen, epiphytes, and understory plants are also more likely to be exposed to the sensor in low-LAI crowns, and spectra from these components could dilute tree species spectral differences. Crown-scale spectra from DIPA and LEAM crowns were thus distinct from other species, but confused between the two species. Despite this confusion, DIPA crowns had 92.6% Producer’s accuracy and 89.3% User’s accuracy (Table 3.7). Furthermore, the User’s accuracy could be increased to 91.8% by applying a LDA probability threshold (Table 3.8). These results are encouraging because large Dipteryx trees have an important ecological function in providing a major seed resource and nesting cavities for the endangered Great green macaw (Ara ambigua) (pers. comm., Powell 2001). Deforestation in the Sarapiquí region has mostly eliminated large Dipteryx trees outside of protected areas, thereby contributing to a dramatic decline in the macaw population. Remote sensing 107 technology that can identify large Dipteryx crowns may contribute to macaw conservation efforts by providing a rapid and cost-effective means to map macaw habitat and migration corridors across the region. In this chapter, another ITC classification technique was to label crown objects using the majority class of classified within-crown pixels. Relative to crown-scale LDA classification with 30 optimal bands, the pixel-majority classification scheme had 8.5% and 2.4% lower accuracy with PixelALL and PixelSUN samples, respectively (Table 3.6). However, one operational advantage of the pixel-majority approach to ITC classification is that robust training statistics (e.g., the covariance matrix) can be estimated from a relatively small number of individual crowns per species because each crown contains many pixels. Furthermore, the pixel-majority approach allows for the inevitable spectral variation within crowns that leads to misclassified samples. I observed that correctly-labeled ITCs had a mean within-crown pixel class accuracy of 90%. ITCs with more mixed-class pixels had more suspect species labels and were often mislabeled. I found that a pixel-majority threshold (e.g., majority class must have ≥35% of within-crown pixels) could be used to improve species User’s accuracy by excluding low-accuracy ITCs and reducing commission errors; and, overall accuracy was left unchanged with the threshold. In contrast, crown-scale spectra blur within-crown variation. Basing an ITC species label on a single, crown-scale spectrum classification is risky due to relatively weak spectral separation among species (Table 3.5, Euclidean distance). My research benefited from distinct phenological differences among species, which undoubtedly helped crown-scale species discrimination. The pixel-majority approach to classification 108 may prove useful if ITCs are more spectrally confused due to similar phenology and crown architecture. IKONOS and ETM+ overall accuracies were both lower than 67% for crownscale and pixel-majority ITC classifications. Since I used 1.6-m simulated multispectral data, none of my tests considered the actual resolution of existing sensors. The IKONOS multispectral sensor provides 4-m resolution images and so could be amenable to either a crown-scale or pixel-majority classification scheme. However, I found that 1.6-m resolution pixels with IKONOS-simulated bands only provided 62% overall accuracy and DIPA User’s accuracy was 64% (LDA, CrownSUN). However, nine ASTER bands could classify ITCs with 77% overall accuracy with CrownSUN spectra, and DIPA User’s accuracy was 76%. ASTER has 2 VIS, 1 NIR, 1 SWIR1 and 5 SWIR2 bands, while IKONOS contains only 3 VIS and 1 NIR bands. These additional SWIR2 bands in ASTER help discriminate species. This conclusion is supported by my hyperspectral, narrowband analyses, which also indicated that SWIR2 was important for species discrimination. At crown scales, ten narrow bands from HYDICE imagery—with wavelength positions in all spectral regions—produced 85% overall accuracy and DIPA User’s accuracy was 93%. I therefore conclude that a high spatial resolution sensor with 10+ channels across the VIS to SWIR2 spectrum is necessary to classify TRF tree species with reasonable accuracy (i.e., ≥85%). For a satellite sensor, finer spectral resolution requires coarser fields of view due to limited surface photon flux. Likewise, airborne sensors can cover a larger swath if flown at a higher altitude with coarser spatial resolution pixels. The crown-scale results indicate that a spatial 109 resolution at the scale of tree crowns is adequate for species discrimination. However, my technique calculated crown-scale spectra by averaging only those spectra from within the ITC footprint, and in an operational situation, coarse-scale square pixels (e.g., 10 to 30 m) may subsume background plant species, soil and shadows, thereby producing mixed spectra and decreased classification accuracy. Future studies should thus examine the sensitivity of classification accuracy to spatial resolutions expected in operational circumstances (i.e., >1.6 m but less than the scale of a crown). I analyzed emergent trees for two reasons: to be certain crowns could be identified in the imagery given georegistration errors, and to sample an adequate number of pixels per crown. Higher spatial resolution sensors will allow the investigation of more TRF tree species, especially co-dominant individuals that do not have broad, emergent crowns. Leaf-scale analyses indicated that spectral measurements with a very fine FOV are more accurately classified than spectrallymixed pixels from coarser spatial scales. For example, leaf spectra were classified with 89% accuracy with just 10 bands and LDA classifier, while pixel and crownscale accuracies were < 85% for the same combination of bands and classifier (Table 3.6). However, these leaf-scale results are based on controlled laboratory conditions with a few species; spectra measured at this fine of scale from an airborne sensor will include atmospheric effects and spectral mixing from photon scattering among various crown tissues and other plant species. It is thus unclear if airborne digital sensors with very high resolutions (i.e., leaf scale) will allow species discrimination with relatively few bands (e.g., Meyer et al., 1996). 110 3.5. Conclusions My results confirm that species of tropical rain forest (TRF) trees can be discriminated based on their spectral reflectance properties. Individual tree crowns (ITCs) were successfully classified with 92% overall accuracy using 30 optimallyselected bands from crown-scale reflectance spectra and a linear discriminant analysis classifier. Pixel-majority ITC classification, which labeled crowns based on within-crown classified pixels, was not as accurate as crown-scale spectra classification. At a fixed 1.6-m spatial scale, crown-scale ITC classification was significantly more accurate with hyperspectral narrowband data (10 band HYDICE) relative to accuracies achieved with multispectral broadband data (simulated IKONOS, Landsat ETM+ and ASTER). This chapter represents the first use of high spectral and spatial resolution imagery acquired over TRF canopy for automated discrimination of individual tree species. Similar to laboratory-based analyses by Cochrane (2000), my results indicate that there are spectral differences among species that permit classification at leaf to crown scales; however, there is also temporal and spatial spectral variation within populations and even single individuals of TRF tree species that will inevitably decrease classification accuracy. A major challenge is to develop classification schemes that can maximize the spectral, spatial and temporal information content of digital imagery while accommodating inherent variation within species. In Chapters 4 and 5, I explore more elaborate classification 111 techniques for tree species discrimination that seek to fully exploit the hyperspectral properties of the HYDICE data. 112 Table 3.1. Study tree species attributes (adapted from Frankie et al., 1974, O’Brien, 2001 and personal observation). Leaf cover is for late-March to early-April, and is what would be expected for the majority of individuals for each species based on available literature data and personal field observations. Tree species Code Leaf Leaf 3/30 [family or sub-family] Phenology Exchange Leaf Functional Timing Cover Group Balizia elegans BAEL Deciduous Annual High (Ducke) Barneby & Grimes [Mimosoideae] Ceiba pentandra CEPE Deciduous Annual High Gaertn. [Bombacaceae] Dipteryx panamensis DIPA Deciduous Annual Low (Pittier) Record & Mell [Papilionoideae] Hyeronima alchorneoides HYAL Evergreen Continuous High Allemão [Euphorbiaceae] Hymenolobium mesoamericanum HYME Deciduous Sub-annual High Lima [Papilionoideae] Lecythis ampla LEAM Deciduous Annual Low Miers [Lecythidaceae] Terminalia oblonga TEOB Evergreen Continuous High (Ruiz & Pav.) Steud. [Combretaceae] Table 3.2. Summary of characteristics of individual tree crowns from HYDICE data. Species names defined in Table 3.1. Tree No. of Crown area, m2 All pixels/crown Sunlit pixels/crown species crowns mean (range) mean (range) mean (range) BAEL 29 358 (108-699) 140 (42-273) 68 (19-131) CEPE 10 766 (361-1695) 299 (141-662) 153 (62-338) DIPA 81 519 (141-1167) 203 (55-456) 98 (28-227) HYAL 34 388 (159-635) 152 (62-248) 78 (34-118) HYME 14 479 (108-1009) 187 (42-394) 99 (23-185) LEAM 21 349 (164-630) 136 (64-246) 67 (31-136) TEOB 25 312 (105-543) 122 (41-212) 64 (21-110) All 214 444 (105-1695) 174 (41-662) 87 (19-338) 113 Table 3.3. Laboratory leaf spectra summary. Species names defined in Table 3.1. Tree species No. of samples No. trees sampled (leaves per tree) BAEL 16 3 (5-6) CEPE 15 3 (5) DIPA 30 5 (3-10) HYAL 23 4 (6-10) HYME 30 3 (6-10) LEAM 14 3 (2-6) TEOB 24 4 (5-7) All 152 25 Table 3.4. Mean and standard deviation (s.d.) of percent reflectance across spectral regions. Means and s.d. were calculated on a band-by-band basis and then averaged for each spectral region. VIS NIR SWIR1 SWIR2 Full Spectra Leaf 5.8 (2.8) 41.3 (6.8) 26.3 (5.5) 12.7 (4.2) 20.9 (4.7) PixelALL 2.7 (1.4) 34.8 (13.5) 15.1 (6.3) 5.5 (2.6) 14.4 (5.9) PixelSUN 3.4 (1.1) 44.4 (10.3) 20.2 (4.8) 7.5 (2.3) 18.4 (4.6) CrownALL 2.7 (0.8) 35.6 (6.9) 15.2 (2.9) 5.8 (1.7) 14.6 (3.0) CrownSUN 3.5 (0.8) 45.0 (7.9) 19.7 (3.1) 6.9 (1.7) 18.5 (3.4) 114 Table 3.5. Non-parametric multivariate analysis of variance (NPMANOVA), comparing among- and within-species spectral variation using spectral angle and Euclidean distance. Values are F ratios. In all analyses, 5000 permutations were used to test significance. Significance is: ns = not significant, * = p≤0.05, ** = p≤0.01, *** = p≤0.001. Crown Scale Spectral Region Bands Leaf Scale a Pixel Scale Spectral Angle Distance Full spectra 161 n/a 119.7 *** 5.4 * Full spectra (sunlit) 161 16.1 *** 162.3 *** 7.7 *** VIS 43 n/a 68.3 *** 1.3 ns VIS (sunlit) 43 13.4 *** 103.3 *** 2.5 ns NIR 46 n/a 113.0 *** 6.0 *** NIR (sunlit) 46 12.6 *** 137.2 *** 8.3 *** SWIR1 25 n/a 94.2 *** 0.8 ns SWIR1 (sunlit) 25 26.6 *** 128.1 *** 1.4 ns SWIR2 48 n/a 32.3 *** 3.0 ns SWIR2 (sunlit) 48 10.2 *** 78.3 *** 3.6 * Euclidean Distance Full spectra 161 n/a 28.6 *** 3.7 *** Full spectra (sunlit) 161 13.4 *** 180.3 *** 3.7 *** VIS 43 n/a 13.1 *** 4.6 *** VIS (sunlit) 43 4.1 *** 51.2 *** 4.5 *** NIR 46 n/a 34.1 *** 2.8 *** NIR (sunlit) 46 12.8 *** 195.7 *** 2.6 *** SWIR1 25 n/a 16.4 *** 4.4 *** SWIR1 (sunlit) 25 18.4 *** 133.7 *** 4.4 *** SWIR2 48 n/a 61.2 *** 3.8 *** SWIR2 (sunlit) 48 18.6 *** 157.4 *** 3.8 *** a Artificial illumination from halogen lamp 115 Table 3.6. Overall accuracy of classifiers using leaf-, pixel- and crown-scale narrowband (HYDICE) spectra. Leaf-scale data were simulated HYDICE spectra from laboratory measurements. “Pixel-majority” refers to ITC classification using a class-majority rule applied to classified within-crown pixels. Arrows represent the direction of significant improvement in overall accuracy between using all and sunlit-only samples (ns = not significant at α=0.05). Pixel d Crown c Pixel-majority c Leaf c Bands a All All Sunlit All Sunlit All Sunlit Linear Discriminant Analysis (LDA) 10 a 89.5 67.5 → 72.1 84.1 ns 84.6 74.8 ns 81.8 20 a 98.0 75.0 → 81.7 89.3 ns 85.5 81.3 ns 83.2 a 30 99.3 79.0 → 83.2 92.1 ns 87.9 83.6 ns 85.5 40 a 100.0 79.2 → 86.5 90.2 ns 89.7 84.1 ns 84.1 a 50 100.0 79.7 → 85.2 89.7 ns 91.1 83.6 ns 85.5 60 a 100.0 80.6 → 85.4 89.7 ns 90.2 84.1 ns 84.1 161 (Full) 88.8 80.9 → 85.5 61.2 ns 67.8 81.8 ns 83.6 b VIS 60.5 45.1 → 53.1 65.9 ns 66.4 62.1 ns 67.3 NIRb 80.9 54.7 → 61.5 67.8 ns 67.3 60.3 ns 66.4 SWIR1 b 88.8 55.1 → 59.8 69.2 ns 68.7 64.0 ns 71.5 b SWIR2 81.6 47.8 → 57.6 75.7 ns 76.6 64.5 ns 69.6 Maximum Likelihood (ML) 10 a 80.9 69.5 → 76.1 76.6 ns 77.6 77.6 ns 79.4 20 a n/a 76.4 → 86.2 n/a 84.6 ns 82.7 30 a n/a 79.5 → 86.7 n/a 82.7 ns 80.4 40 a n/a 81.6 → 86.9 n/a 82.2 ns 80.4 50 a n/a 81.2 → 87.3 n/a 79.0 ns 79.9 a 60 n/a 79.9 → 87.6 n/a 78.0 ns 77.6 161 (Full) n/a 68.3 → 79.1 n/a 71.0 ns 71.5 VIS b 64.5 49.7 → 58.7 30.4 ns 27.1 62.6 ns 67.3 b NIR 71.7 59.6 → 66.3 50.0 ← 37.9 69.2 ns 70.1 SWIR1 b 82.2 53.5 → 66.2 34.1 → 46.7 66.8 ns 69.2 b SWIR2 71.1 49.2 → 58.1 39.7 ns 49.1 65.4 ns 71.0 116 Table 3.6 (continued). Spectral Angle Mapper (SAM) 10 a 46.1 42.4 ns 44.1 45.8 ns 50.9 48.6 ns 42.5 a 20 50.7 37.9 → 48.9 46.3 ns 53.7 48.1 ns 56.5 30 a 48.0 39.8 → 48.6 46.7 ns 47.7 46.3 ns 50.9 40 a 48.7 39.2 → 46.4 45.8 ns 50.5 46.3 ns 52.8 a 50 47.4 38.9 → 47.0 46.7 ns 51.4 47.7 ns 53.3 60 a 46.1 38.0 → 48.4 46.3 ns 50.5 46.3 ns 50.0 161 (Full) 48.7 38.7 → 48.4 44.4 ns 48.6 43.9 ns 47.2 b VIS 35.5 38.8 → 42.1 14.0 ns 12.6 41.6 ns 43.0 NIR b 38.2 42.3 → 47.1 37.4 ns 39.3 48.6 ns 49.5 b SWIR1 37.5 33.5 → 38.0 45.3 ns 51.9 37.4 ns 41.6 SWIR2 b 36.8 31.6 → 36.8 45.8 ns 44.9 43.0 ns 39.3 a. Bands selected using a Linear Discriminant Analysis (LDA) forward, stepwise selection procedure. Significance criteria α =0.05 for leaf, pixel and crown objects, α = 0.2 for CrownALL and CrownSUN spectra). b. VIS, NIR, SWIR1 and SWIR2 regions have 10 evenly-spaced bands. c. Accuracy results are from cross validation of samples. d. Training and testing data were two mutually-exclusive sets of 300 randomly-selected pixels per species. 117 Classification 118 Table 3.7. Error matrix for crown-scale classification using 30 bands and a Linear Discriminant Analysis classifier with no a posteriori probability threshold (Kappa = 0.90). Field Reference Species BAEL CEPE DIPA HYAL HYME LEAM TEOB Total 27 1 28 BAEL 7 1 8 CEPE 1 2 75 1 1 4 84 DIPA 1 33 34 HYAL 1 13 14 HYME 1 3 17 21 LEAM 25 25 TEOB Unknown 29 10 81 34 14 21 25 214 Total 93.1% 70.0% 92.6% 97.1% 92.9% 81.0% 100.0% Prod. % Species names defined in Table 3.1. 118 User % 96.4% 87.5% 89.3% 97.1% 92.9% 81.0% 100.0% 92.1% Classification 119 Table 3.8. Error matrix for crown-scale classification using 30 bands and a Linear Discriminant Analysis classifier with a 90% a posteriori probability threshold (Kappa = 0.79). Field Reference Species BAEL CEPE DIPA HYAL HYME LEAM TEOB Total User % 25 1 26 96.2% BAEL 6 1 7 85.7% CEPE 1 1 67 1 3 73 91.8% DIPA 1 32 33 97.0% HYAL 1 11 12 91.7% HYME 1 12 13 92.3% LEAM 25 25 100.0% TEOB 2 3 10 2 2 6 25 Unknown 29 10 81 34 14 21 25 214 Total 83.2% Prod. % 86.2% 60.0% 82.7% 94.1% 78.6% 57.1% 100.0% Species names defined in Table 3.1. 119 Table 3.9. Classification results for simulated broadband spectra. Bands are for IKONOS, Landsat ETM+ and ASTER sensors. Scales follow Table 3.6. Pixel e Crown d Pixel-majority d Leaf d a Bands All All Sunlit All Sunlit All Sunlit Linear Discriminant Analysis (LDA) IKONOS a 44.7 42.7 → 49.2 59.3 ns 61.7 52.3 ns 59.3 ETM+ b 57.9 48.6 → 55.8 66.4 ns 66.4 60.3 ns 64.5 ASTER c 80.3 56.4 → 64.7 76.2 ns 77.1 66.8 ns 72.0 Maximum Likelihood (ML) IKONOS a 61.2 45.3 → 55.1 50.5 ns 50.9 48.1 ns 51.9 b ETM+ 73.7 54.0 → 62.4 62.6 ns 62.1 62.6 ns 65.9 ASTER c 83.6 59.4 → 70.4 72.9 ns 72.4 77.6 ns 78.0 Spectral Angle Mapper (SAM) IKONOS a 40.1 45.3 ← 33.6 20.1 ns 31.3 20.1 ns 31.8 ETM+ b 34.2 32.5 → 35.9 38.3 ns 39.7 34.1 ns 41.1 ASTER c 40.1 30.6 → 36.1 41.1 ← 31.3 38.3 ns 37.9 a. 4 bands - 483, 551, 663, 794 nm. b. 6 bands - 479, 561, 661, 835, 1651, 2209 nm. c. 9 bands - 555, 658, 805, 1655, 2166, 2207, 2264, 2333, 2394 nm. d. Accuracy results are from cross validation of samples. e. Training and testing data were two mutually-exclusive sets of 300 randomly-selected pixels per species. 120 Classification 121 Table 3.10. Error matrix for pixel-majority classification using 30 bands, sunlit pixels, a Linear Discriminant Analysis classifier, and no pixel-majority threshold (Kappa = 0.82). Field Reference Species BAEL CEPE DIPA HYAL HYME LEAM TEOB Total User % 24 3 3 1 1 32 75.0% BAEL 5 5 100.0% CEPE 2 5 76 1 1 8 93 81.7% DIPA 2 1 30 33 90.9% HYAL 11 11 100.0% HYME 1 12 13 92.3% LEAM 25 25 100.0% TEOB 1 1 2 Unknown 29 10 81 34 14 21 25 214 Total 82.8% 50.0% 93.8% 88.2% 78.6% 57.1% 100.0% 85.5% Prod. % Species names defined in Table 3.1. 121 La Selva 1 Km Costa Rica Tree crowns HYDICE extent Rivers Land Use Developed Areas Selectively- logged Old- growth Forest Secondary Forest Pasture Plantation Swamp Figure 3.1. The La Selva Biological Station study site and extent of HYDICE hyperspectral imagery. The 214 study crowns are labeled with points. 122 A B SPECIES BAEL CEPE DIPA HYAL HYME LEAM TEOB Figure 3.2. A) View of old-growth Tropical Wet Forest at the La Selva Biological Station. The canopy-emergent tree in the foreground is Balizia elegans. B) Example of 1.6-m spatial resolution HYDICE hyperspectral imagery over oldgrowth canopy (Red: 1651 nm [SWIR2], Green: 835 nm [NIR], Blue: 661 nm [Red]) with overlaid individual tree crown polygons. Map scale is 1:3000. 123 0.50 HYDICE Canopy 0.45 HYDICE Bridge ASD Canopy 0.40 ASD Bridge Reflectance 0.35 0.30 0.25 0.20 0.15 0.10 0.05 0.00 350 850 1350 1850 2350 Wavelength (nm) Figure 3.3. Reflectance spectra from airborne HYDICE and field ASD spectrometers for a wooden bridge (over dark water) and a Pentaclethra macrophylla crown. The Pentaclethra ASD spectrum was acquired from the bridge. 124 Reflectance Reflectance Reflectance 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 BAEL (N =16) 850 1350 1850 2350 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 2350 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 2350 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 DIPA (N = 30) 850 1350 1850 HYME (N = 30) 850 1350 1850 CEPE (N = 15) 850 1350 1850 2350 HYAL (N = 23) 850 1350 1850 2350 LEAM (N = 14) 850 1350 1850 2350 Reflectance Wavelength (nm) TEOB (N = 24) 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 850 1350 1850 2350 Wavelength (nm) Figure 3.4. Leaf-scale mean (bold line) and standard deviation (±1 S.D., thin line) of reflectance by species. Species codes are listed in Table 3.1. 125 A 0.60 100% Reflectance 0.50 50% 0.40 0% 0.30 0.20 0.10 0.00 400 650 900 1150 1400 1650 1900 2150 2400 Wavelength (nm) B 0.60 20% Reflectance 0.50 10% 0.40 0% 0.30 0.20 0.10 0.00 400 650 900 1150 1400 1650 1900 2150 2400 Wavelength (nm) C 0.60 Senesced Reflectance 0.50 Mature Young 0.40 0.30 0.20 0.10 0.00 400 650 900 1150 1400 1650 1900 2150 2400 Wavelength (nm) Figure 3.5. Leaf-scale variation in spectral properties. A) Hymenolobium mesoamericanum leaves: percent area covered by a single species of epiphyll. B) Lecythis ampla leaves: percent area of leaf herbivory (i.e., light brown-colored mines) caused by a leaf-mining insect. C) Terminalia oblonga leaves: leaf aging from young to senesced leaves. Note: all spectra in Fig. 3.5 were from upper-canopy leaves and were included in leaf-scale spectral analyses except for the Terminalia senesced leaf, which was collected on the ground. 126 Reflectance Reflectance Reflectance 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 BAEL (N =300) 850 1350 1850 2350 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 2350 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 2350 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 DIPA (N =300) 850 1350 1850 HYME (N = 300) 850 1350 1850 CEPE (N = 300) 850 1350 1850 2350 HYAL(N = 300) 850 1350 1850 2350 LEAM (N = 300) 850 1350 1850 2350 Reflectance Wavelength (nm) TEOB (N = 300) 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 850 1350 1850 2350 Wavelength (nm) Figure 3.6. Mean (bold line) and standard deviation (±1 S.D., thin line) of reflectance by species for sunlit pixels. Species codes are listed in Table 3.1. 127 Reflectance Reflectance Reflectance 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 BAEL (N = 29) 850 1350 1850 2350 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 2350 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 2350 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 DIPA (N = 81) 850 1350 1850 HYME (N = 14) 850 1350 1850 CEPE (N = 10) 850 1350 1850 2350 HYAL (N = 34) 850 1350 1850 2350 LEAM (N = 21) 850 1350 1850 2350 Reflectance Wavelength (nm) TEOB (N = 25) 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 350 850 1350 1850 2350 Wavelength (nm) Figure 3.7. Crown-scale mean (bold line) and standard deviation (±1 S.D., thin line) of reflectance by species for CrownALL spectra. Species codes are listed in Table 3.1. 128 Leaf Scale A 1.20 Reflectance 1.00 All 20 All 10 0.80 0.60 0.40 0.20 0.00 350 850 1350 1850 2350 Wavelength (nm) Pixel Scale B 1.20 Sunlit 20 Sunlit 10 Reflectance 1.00 All 20 All 10 0.80 0.60 0.40 0.20 0.00 350 850 1350 1850 2350 Wavelength (nm) Crown Scale C 1.20 Sunlit 20 Sunlit 10 Reflectance 1.00 0.80 All 20 All 10 0.60 0.40 0.20 0.00 350 850 1350 1850 2350 Wavelength (nm) Figure 3.8. Mean reflectance spectra for the seven study species at A) leaf, B) pixel and C) crown scales from Figures 3.4, 3.6 and 3.7, respectively. Pixel and crown spectra were calculated from sunlit-only (Sun) and all (All) pixel spectra prior to band selection. PixelSUN and CrownALL spectra are displayed in B and C, respectively. Dots above spectra represent the best 10 and 20 bands selected by the stepwise-selection procedure. 129 100 Percent 80 60 SWIR2 SWIR1 40 NIR VIS 20 0 Leaf Pixel (All) Pixel Crown Crown (Sun) (All) (Sun) 100 Percent 80 SWIR2 SWIR1 NIR VIS 60 40 20 0 Leaf Pixel (All) Pixel Crown Crown (Sun) (All) (Sun) Figure 3.9. The percentage of 10 (A) and 20 (B) stepwise-selected bands within each spectral region at leaf, pixel and crown scales. 130 Percent Accuracy 100 90 80 70 60 50 40 30 20 10 0 90% 75% 50% No Probability 0 5 10 15 20 25 30 35 40 45 50 55 60 Number of Bands Figure 3.10. Crown-scale classification overall accuracy with the addition of bands. Bands were added based on their ranking using a stepwise procedure and the classifier was linear discriminant analysis (LDA). The a posteriori LDA probability required for a crown to be classified was adjusted to four different thresholds: 1) no probability threshold and 50%, 75% and 90% thresholds. 131 CHAPTER 4: Classification of tree species with absorption features and binary decision trees 4.1. Introduction Digital data from high spatial resolution satellite or airborne optical sensors provide a new means of mapping and monitoring individual tree crowns (ITCs) over larger areas and with more frequency than traditional field-based techniques (Clark, Read, et al., 2004; Gougeon & Leckie, 2003). Research in tropical forests has shown that the spatial and temporal resolution of this imagery permits the remote measurement of ITC growth and mortality, as well as canopy disturbances such as gap formation from selective logging (Clark, Read et al., 2004; Clark, Castro et al., 2004; Hurtt et al., 2003; Read et al., 2003). In temperate forests, high spatial resolution multispectral imagery has been used to detect, delineate and classify the species of tree crowns (Gougeon & Leckie, 2003; McGraw et al., 1998). In tropical forests, however, within species (conspecific) variation in reflectance properties due to crown architecture, phenology and other biotic factors severely limits the specieslevel discrimination of ITCs with low spectral resolution, multispectral data (Chapter 3). Hyperspectral sensors measure the visible to shortwave infrared region (4002500 nm) of the electromagnetic spectrum in over 100 channels, and this increased spectral resolution may permit species discrimination based on subtle differences in their reflectance spectra. When viewed from above by an airborne sensor, spectral 132 variation among tree species is controlled by two principal factors: biochemistry and structure. Leaves, woody branches and trunks, flowers and fruits each have their own biochemical properties that create distinct absorption features in reflectance spectra. Green leaves have photosynthetic pigments—mainly chlorophyll-a, -b, or c and caroteniods (carotenes, xanophylls) (Gates et al., 1965; Raven et al., 1992). These pigments absorb light in the blue region of the electromagnetic spectrum (near 445 nm), but only chlorophyll pigments absorb light in the red region (645-680 nm). Green leaves also have weak water absorption features in the near-infrared region at 970 nm and 1240 nm, respectively (Gao & Goetz, 1990; Gates et al., 1965). In contrast, bark spectra typically have no photosynthetic pigment signal, but instead have absorption features in the shortwave infrared region due to biochemical compounds, such as lignin, cellulose and proteins (Curran, 1989; Elvridge, 1990). The internal and surface structures of plant tissues also play significant roles in multiple-scattering and transmission of photons within their canopies (Gates et al., 1965; Grant, 1987). At a coarser scale, species-level differences in crown structure, such as leaf and branch area, angular distribution, and clumping, affect the expression of tissue biochemical absorption features in branch- to crown-scale reflectance spectra (Asner, 1998). I hypothesize that if tropical rain forest tree species differ in crown structure or tissue biochemistry, then hyperspectral metrics that target key absorption features should be useful for species-level individual tree crown classification. Several recent studies have correlated hyperspectral reflectance bands or derived metrics 133 with canopy physicochemical properties to classify temperate forest composition (Fuentes et al, 2001; Kokaly et al., 2003; Martin et al., 1998; Ustin & Xiao, 2001). Cochrane (2000) noted differences in red-edge metrics for tropical rain forest tree species when analyzing laboratory spectra, but there have been no studies that have used hyperspectral-derived metrics for tropical tree species classification. In this chapter, I explore hyperspectral metrics and a decision-tree classification scheme for automatic classification of individual tropical rain forest tree species from high spatial resolution imagery. My main objectives were to: 1) compute a suite of hyperspectral metrics that respond to crown structure and absorption features from photosynthetic pigments, water and other biochemicals; 2) at tissue, pixel and crown scales, test for significant differences in mean response of these spectral metrics among tropical tree species; and, 3) assess ITC classification accuracy using binary decision trees applied to spectral metrics. 4.2. Background 4.2.1. Hyperspectral metrics There are many methods for measuring the absorption features expressed in hyperspectral data. One approach is to use a ratio-based index that contrasts an absorption feature band with a band from a relatively stable spectral region (Elvidge & Chen, 1995; Peñuelas et al., 1997b). More sophisticated techniques involve using the multiple, contiguous reflectance bands within a hyper-spectrum to identify the shape and position of key absorption features (Broge & Leblanc, 2000; Clark, 134 Swayze et al., 2003). Common metrics are based on first- and second-order derivatives of reflectance spectra (Demetriades-Shah et al., 1990; Elvridge & Chen, 1995) and absorption feature area, width or depth (Clark, Swayze et al., 2003; Gong et al., 2002; Kokaly et al., 2003; Pu, Ge et al., 2003). A first-order derivative spectrum is the slope of the reflectance spectrum at every point. The first- and second-order derivatives can be used to resolve overlapping absorption features and identify inflection points (Demetriades-Shah et al., 1990). Below I discuss potential spectral metrics related to key vegetation absorption features. In this discussion, I refer to three regions of the electromagnetic spectrum: visible (VIS=437-700 nm), near-infrared (NIR=700-1327 nm), and shortwave-infrared (SWIR=1467-2435 nm). 4.2.2. Absorption features related to photosynthesis Vegetation indices are common in remote sensing and generally measure properties of photosynthetic pigment absorptions. I investigated the following vegetation indices (formulas in Table 4.1): Simple Ratio Vegetation Index (SR; Jordon, 1969; Tucker, 1979), Normalized Difference Vegetation Index (NDVI; Rouse et al., 1973), Soil-Adjusted Vegetation Index (SAVI; Huete, 1988), Photochemical Reflectance Index (PRI; Gamon et al., 1997), Atmospherically Resistant Vegetation Index (ARVI; Kaufman & Tanré, 1992), Enhanced Vegetation Index (EVI; Huete et al., 2002), and the Red-edge Vegetation Stress Index (RVSI; Merton, 1998). The SR, NDVI, SAVI, ARVI, and EVI indices were developed for broadband multispectral sensors (e.g., Landsat, AVHRR), but they can be 135 formulated with narrowband wavelengths from hyperspectral data (Elvidge & Chen, 1995; McGwire et al. 2000). The popular SR and NDVI measure the contrast in red-light chlorophyll absorption relative to high NIR reflectance caused by internal leaf and canopy scattering (Tucker, 1979). At relatively coarse spatial scales (e.g., >900 m2), these indices have been linked to forest biophysical parameters such as leaf area index (LAI), aboveground biomass, and primary productivity (Elvidge & Chen, 1995; Spanner et al. 1994), and have contributed to forest-type classification (Steininger, 1996), mapping of plant species richness (Oindo & Skidmore, 2002), and change detection (Ferreira et al., 2003). The NDVI and SR are sensitive to background soil and atmospheric contamination, and SAVI and ARVI were introduced to minimize these two factors, respectively (Huete et al., 1985; Huete, 1988; Kaufman & Tanré, 1992). At coarse spatial scales, the EVI minimizes both soil and atmospheric noise and is more sensitive to vegetation structure (e.g., LAI, canopy architecture) than NDVI (Huete et al., 2002). The PRI is a narrowband ratio that is sensitive to changes in reflectance at 531 nm caused by interconversions of xanthophyll cycle pigments, which are related to photosynthesis light-use efficiency across species, functional types, and nutrient treatments (Gamon et al., 1997; Peñuelas et al., 1997a). Few studies have investigated structural and chemical variation among plant species as expressed through these ratio-based indices. Using field measurements of 6 tree species in Africa, Franklin et al. (1991) found strong relationships between NDVI and crown biomass for each species. Nagler et al. (2004) found that NDVI 136 calculated from fine spatial resolution digital imagery was significantly different among individual riparian species. They attributed variation among species to leaf angle effects on light attenuation rather than LAI. In exploring narrowband indices for species-independent chlorophyll estimation, Sims and Gamon (2002) noted that relationships were weakened by species differences in leaf surfaces (e.g., waxes, pubescence), causing variation in first-return reflection. First-order derivative bands derived from hyperspectral reflectance spectra have been used to estimate foliar content of chlorophyll, nitrogen, phosphorus, and potassium in conifer and mixed-deciduous forest canopies (Gong et al., 2002; Martin & Aber, 1997; Smith et al., 2002). Derivative analysis is useful for finding the inflection point on the tail of an absorption feature. The wavelength of the red-edge inflection point (RE-λ), between 680 nm and 740 nm, is correlated to chlorophyll concentration, LAI and vegetation stress (Blackburn, 1998; Demetriades-Shah et al., 1990; Horler et al., 1983; Pu, Gong et al., 2003), and it may be useful for discriminating tropical rain forest tree species (Cochrane, 2000). 4.2.3. Water absorption features Green vegetation spectral reflectance in the 900 to 2500 nm region of the spectrum is dominated by liquid water absorption, and narrowband sensors can exploit measurements in this region to estimate vegetation moisture content. Two useful water indices are the Water Band Index (WBI; Peñuelas et al., 1997b) and the Normalized Difference Water Index (NDWI; Gao, 1996), which measure NIR water absorption features at 970 nm and 1240 nm, respectively (Table 4.1). These indices 137 are not influenced by chlorophyll absorption, and so they are complementary to indices based on red absorption (Gao, 1996). Near-infrared water absorption features are also useful for estimating leaf equivalent water thickness (EWT), which is the amount of liquid water in vegetation needed to account for an observed water absorption feature (Roberts et al., 1997). The metric is calculated by fitting a Beer-Lambert model of light extinction through a water-absorbing medium to the vegetation spectrum covering either the 950–1000 or 1150–1260 nm absorption regions (Roberts et al., 1997). Research has shown that EWT and other hyperspectral-derived metrics correlate with moisture content and structure of canopy components (Dennison et al., 2003; Roberts, Brown et al.,1998; Roberts et al., 2003; Roberts et al., 2004; Serrano et al., 2000; Ustin et al., 1998). 4.2.4. Other biochemical absorption features Several other biochemicals in vegetation create absorption features across the 400-2500 nm spectrum measured by hyperspectral sensors (Curran, 1989; Elvidge, 1990). Cellulose is a prevalent structural chemical that has NIR and SWIR combination-band and overtone absorptions at 1.22, 1.48, 1.93, 2.10, 2.28, 2.34 and 2.48 µm (Elvidge, 1990). Lignin is another important chemical in plants, with combination-bands and overtones at 1.45, 1.68, 1.93, 2.05-2.14, 2.27, 2.33, 2.38 and 2.50 µm. Other biochemicals include proteins, xylan, tannins and waxes (Curran, 1989; Elvidge, 1990). Typically, absorptions by these biochemicals are not isolated but overlap due to broadening by multiple scattering and overlap with absorptions at similar wavelengths (Curran, 1989). Green leaves contain 40-80% water by weight, 138 and thus water absorption tends to mask the more subtle SWIR absorptions caused by other chemicals (Curran, 1989; Elvridge, 1990; Kokaly & Clark, 1999). Remote sensing of vegetation has not focused on SWIR features for estimating biochemical concentrations or discriminating species, largely because these regions are sampled with only one to a few bands in multispectral sensors, if at all. Imaging spectroscopy provides an unprecedented opportunity to exploit the information content of SWIR features (Curran, 1989; Elvidge, 1990). In one novel study, bands normalized to the depth or area of SWIR absorption features were found to be highly correlated to biochemicals such as nitrogen, lignin, and cellulose in dry leaves (Kokaly & Clark, 1999). Using similar techniques, Curran et al. (2001) found that VIS, NIR and SWIR features were useful for predicting chlorophyll, water, nitrogen, cellulose, lignin and other plant chemicals. 4.2.3. Spectral mixture analysis Another analytical technique is spectral mixture analysis (SMA), which models reflectance spectra as a linear combination of two or more dominant or “pure” spectral components, or endmembers (reviewed in Keshava & Mustard, 2002). SMA outputs are the fractional abundance of each endmember and a root-meansquare error (RMSE) model fit. Endmembers are selected from a library of field, laboratory, image-extracted or “virtual” spectra. Typically one endmember is designated as shade, which models variation in illumination. Other bright endmember(s) represent major spectral components—typically green photosynthetic vegetation (GV), non-photosynthetic vegetation (NPV: litter, bark, branches) and 139 soil in vegetation mapping applications (Roberts et al., 1993). Although SMA can be performed on multispectral data, the model solution is strengthened with hyperspectral data because there are more high-signal bands, which may help distinguish spectral mixtures based on the unique spectral shape (i.e., absorption features) of their endmembers. 4.3. Methods 4.3.1. Canopy-emergent trees Analyses focused on the classification of canopy emergent individuals of seven tree species (Table 4.2). Chapter 3 provides details about the species and number of study trees (Table 3.1) and how their ITC polygons were digitized on the HYDICE hyperspectral imagery. As explained in Chapter 3, some overstory tree species are completely deciduous, generally beginning in the first dry season, while others are evergreen and continuously flush small amounts of leaves throughout the year (Table 4.2). Hyperspectral imagery was acquired on March 30, 1998 (Chapter 1), at the end of the first dry season, and all study trees were expected to have high mature leaf cover except DIPA and LEAM (Table 4.2). However, BAEL and HYME had relatively fine compound leaves and so their mature leaf cover was expected to have a lower LAI relative to the broadleaf species CEPE, HYAL and TEOB. I thus expected LAI for the study species to be low for DIPA and LEAM, moderate for BAEL and HYME, and high for CEPE, HYAL and TEOB. 140 4.3.2. Bark spectra Laboratory leaf and bark spectra were acquired for tissue-scale assessment of metrics and SMA. I measured leaf spectra in a laboratory at LSBS (Fig. 4.1a; detailed methods in Chapter 3). Bark specimens from the seven study species (Table 4.2), as well as from Cedrela odorata, Cordia alliodora, Laetia procera, Pentaclethra macroloba, and Simarouba amara were sampled in the station vicinity in April, 2004. Most samples were from the branches of young trees that were retrieved with a shotgun, pole clipper, rope, or saw, while some DIPA trunk bark was sampled by climbing emergent trees. Bark specimens had an average width of 6 cm and length of 20 cm. Fresh specimens were stored in sealed plastic bags in a refrigerator at 6° C for 1.5 months and their spectral properties were measured with an ASD full-range spectrometer (Analytical Spectral Devices, Boulder, CO, USA) sensor with an 8° fore-optic. Specimens were blotted with a paper towel to remove surface water and were placed in a 5%-reflective box and illuminated with a 250 W halogen bulb. The fore-optic was positioned 10 cm at nadir above the box center, yielding a 1.4-cm sensor field of view (FOV). To capture spectral variation from each specimen, the sample orientation and position relative to the sensor were varied with each radiance measurement (the sample was moved, the sensor remained stationary). Radiance from a white Spectralon® panel (Labsphere, North Sutton, NH, USA) placed in the box center was used as a standard to convert bark radiance measurements to percent reflectance. A final bark reflectance spectrum was an average of fifteen reflectance spectra from the specimen. The ASD spectrometer had 1-nm spectral sampling covering 350 to 2500 nm. I convolved leaf and bark 141 laboratory spectra to HYDICE band center positions (161 bands) using full-width, half-maximum information for each HYDICE band. For the seven study species, the bark library (Fig. 4.1b) contained 66 spectra: 15 BAEL, 9 CEPE, 10 DIPA, 8 HYME, 5 HYAL, 12 LEAM, and 7 TEOB. The leaf spectral library contained 152 spectra: 16 BAEL, 15 CEPE, 30 DIPA, 23 HYME, 30 HYAL, 14 LEAM, and 24 TEOB. Spectral metrics were calculated from these bark and leaf spectra. For spectral mixture analysis, I sought a more exhaustive bark spectral library and included spectra from the study species, 12 spectra from five other species (listed above), as well as 24 field spectra. Bark field spectra came from tree trunks, which were measured at LSBS at 9:30-10:00 am local time in August, 2002. The ASD FieldSpec sensor with 8° fore-optic was held about 1 m from the trunk and measured radiance was converted to reflectance using an in situ white Spectralon® calibration panel in full sun. 4.3.3. Calculation of spectral metrics 4.3.3.1. Narrowband, ratio-based indices Narrowband vegetation and water-absorption indices were calculated using formulas in Table 4.1 applied to reflectance spectra at tissue, pixel and crown scales for HYDICE bands. The HYDICE bands chosen for the indices were closest to those in the formulas presented in the literature. 142 4.3.3.2. Derivative-based metrics Derivative analysis was used to measure the wavelength position and magnitude of the blue edge (BE), green peak (GP), yellow edge (YE), red well (RW), red edge (RE), NIR water absorption edges (NE1 & NE2), and the SWIR1 edge (SE) (Table 4.3; Fig. 4.1a) for HYDICE bands. Derivatives for these features were calculated using a polynomial-fitting technique (Pu, Gong et al., 2003). Reflectance data were retrieved for all bands in the region covering a spectral feature (e.g., red edge) and a function was then fit to the wavelength and reflectance values according to a polynomial equation and a least-squares minimization: n ρ = α 0 + ∑ α i λi (1) i =1 where ρ represents the HYDICE reflectance values at λ wavelength bands within the absorption feature (Table 4.3), n is the polynomial order, and α represents the fitted polynomial constants. A continuous derivative spectrum for each feature was then calculated using the fitted polynomial function and used to estimate the wavelength of the local minima or maxima for the edge features, corresponding to the inflection point in the original reflectance domain. Similarly, the wavelength position at which the derivative was zero identified the GP and RW features, which corresponded to the peak and well in the reflectance domain, respectively. In addition, the magnitude of the derivative (i.e., reflectance slope) was retrieved at edge inflection wavelengths, while reflectance was retrieved at the GP and RW wavelengths (Gong et al., 2002). 143 Green vegetation metrics based on derivative area were calculated following methods outlined in Elvidge & Chen (1995) and Gong et al. (2002). First-order derivatives of spectra were calculated using a 3-point, Lagrangian interpolation and then the total area under the derivatives was calculated for the blue edge (BE-DArea; 491.0 -532.1 nm; SDB in Gong et al., 2002), yellow edge (YE-DArea; 549.6-581.9 nm; SDY in Gong et al., 2002) and the red well-red edge features (RWE-DArea; 626.7 to 798.2 nm; 1DL_DGVI in Elvidge & Chen, 1995). For the RWE feature, the first-derivative area was also calculated after normalizing the first-derivative spectra to the value at 626.7 nm (RWE-DNArea; 1DZ_DGVI in Elvidge & Chen, 1995). A second-order derivative was then calculated from normalized first-order spectra and the area calculated (RWE-2DNArea; 2DZ_DGVI in Elvidge & Chen, 1995). 4.3.3.3. Absorption-based metrics Using methods adapted from Pu, Ge et al., 2003, I calculated the maximum depth (D), wavelength position of maximum depth (λ), width (W), maximum depth x width area (A1) and asymmetry (As) for photosynthetic pigment (blue and red), water (NIR) and other biochemical (SWIR) absorption features (Table 4.3). Absorption feature area (A2) was also calculated as the sum of the depths measured at each band within the feature’s range (Table 4.3). I calculated EWT using a BeerLambert light-extinction model, with parameters fit to reflectance data between 865 and 1065 nm using a non-linear least-squares minimization routine (method detailed in Dennison et al., 2003). The spectrum of water absorption coefficients required by 144 this technique was interpolated to HYDICE wavelengths from data in Palmer and Williams (1974). 4.3.3.4. Spectral mixture analysis (SMA) fractions Pixel- and crown-scale spectra were unmixed using a three-endmember model composed of green vegetation (GV), non-photosynthetic vegetation (NPV) and photometric shade. The GV endmember was selected by plotting crown pixels in a red versus NIR scatter plot. Thirteen pixels with relatively low red and high NIR were averaged to form one GV image endmember, and nine pixels with high red and low NIR were averaged to form one NPV image endmember. A GV image endmember was used in SMA because it contained pronounced absorption features due to multiple-scattering within the canopy whereas laboratory-measured, leaf-scale spectra had weaker absorption features. The NPV image spectrum was not “pure,” but rather it was a mixture of spectral properties from NPV (e.g., bark) and GV (e.g., tree leaves, canopy epiphytes, moss) measured within the sensor’s instantaneous field of view (IFOV). A library of 102 field and laboratory spectra was used to select a relatively pure NPV endmember spectrum. I chose the NPV endmember from the library using the criteria that it yielded a low root-mean-square error (RMSE) and physically-reasonable fractions when unmixing the image NPV endmember with the image GV endmember (Roberts, Batista et al., 1998). The final NPV reference endmember was from a lichen-covered tree trunk acquired from a suspension bridge at LSBS. 145 4.3.4. Tree species classification techniques I explored the binary decision tree (DT) classifier (Breiman et al., 1984; Friedl & Brodley, 1997; Pal & Mather, 2003; Roberts et al., 2002) for ITC classification. Training data are used to build a DT framework composed of a sequence of binary tests (nodes) applied to the predictor variables (i.e., spectral metrics) that assign pixels to their final class at the end of the decision tree (terminal nodes). Decisiontree classifiers have an advantage over more traditional classifiers like the maximum likelihood classifier, in that they make no assumptions about the data distributions (i.e., they are non-parametric), can handle data with different scales, and can adapt to noise and non-linear relationships inherent in remote sensing data (Friedl & Brodley, 1997). Also, DT rules are explicitly defined and are often interpretable in terms of meaningful, physical factors that are important in discriminating classes. In this chapter, I used the “Tree” package in the R statistical environment for DT analysis (R Development Core Team, 2004; Tree v1.0-18, R v2.0). Parameters for growing trees were “mincut” of 5, “minsize” of 10, “mindev” of 0.001, and “deviance” as the criteria for splitting data into homogenous sets. Similar to methods in Chapter 3, two ITC species classification schemes were analyzed: crown-scale and pixel-majority. The crown-scale approach labeled ITC species using classified crown-scale spectra. Crown-scale reflectance spectra were first calculated from the average of all within-crown pixels (shaded and sunlit) which provided one spectrum per crown. Spectral metrics (e.g., NDVI, red-edge position, etc.) were calculated from crown-scale reflectance spectra. Crown-scale DT used a leave-one-out cross-validation approach. 146 An ITC was classified by removing the crown from the analysis and a DT was then built using the remaining crowns as training data (n=213). Initial analyses revealed that classification accuracy was higher when DTs were constructed using a balanced training data set of equal class sample size. Since each species class had a different number of crowns, balanced training data were acquired with a random sample with replacement of 50 crowns per class. No DT pruning was performed because low sample sizes kept the decision rules relatively simple. The DT was then used to classify the ITC that had been held out from the training set. This process of training and classification was repeated 50 times per ITC and the final ITC label was chosen using the majority class of the 50 DTs. Pixel-majority classification was also performed with a leave-one-out cross validation. In this case, each ITC was iteratively withdrawn from analysis and 2,000 pixels per species were randomly-selected without replacement from the remaining ITCs (n=213). For each class, the sampled pixels were then divided evenly into training and pruning data sets (i.e., 1,000 pixels each). Decision trees were automatically pruned with the R Prune.Tree routine, which constructs a nested sequence of decision sub-trees by snipping off the least important splits based on a cost-complexity parameter. Pruning data were dropped down each decision sub-tree and deviances were calculated between the predicted and response classes. The decision sub-tree producing the minimum deviance was chosen as the pruned decision tree. Within-crown pixels from the withheld ITC were then classified using the pruned decision tree and the ITC label was chosen using the majority class of the within-crown classified pixels. For each ITC withheld from training, the process of 147 decision tree construction, pruning and ITC majority-class labeling was done 50 times—with a new set of mutually-exclusive, 1,000 training and 1,000 pruning pixels per class randomly sampled each time. The final ITC label was then assigned using the majority class of the 50 crown labels for each ITC. 4.4. Results 4.4.1. Differences in spectral metrics among species A summary of the spectral metrics calculated in this chapter is presented in Table 4.4. Leaf-, bark-, pixel- and crown-scale summary statistics and tests are presented for ratio-based (Appendix 2.1), derivative-based (Appendix 2.2), and absorptionbased (Appendix 2.3) metrics, while spectral mixture analysis fractions are presented at pixel and crown scales (Appendix 2.4). I tested mean differences in spectral metrics among species with an ANOVA and I used a Tukey’s Post Hoc Honestly Significant Different procedure (Zar, 1996) for multiple comparison pair-wise tests of mean differences between two species. The relative statistical strength of metrics for distinguishing species was ranked primarily by the magnitude of the ANOVA F statistic and secondarily by the number of significant pair-wise differences (21 total). Out of 73 metrics calculated from laboratory leaf and bark spectra, 61 and 55 had significant differences in mean values among species, respectively (Appendix 2.12.3). At pixel and crown scales, 76 and 69 out of 77 of the spectral metrics showed significant differences in mean response among species (Appendix 2.1-2.4). 148 4.4.2. Photosynthetic pigment absorption metrics In laboratory leaf spectra, 94% of photosynthesis metrics showed significant differences among species. Metrics covering the red-edge feature—RVSI and REλ—were the most important of these metrics at this scale (Table 4.5, leaves). The blue absorption wavelength (Blue-λ), depth (Blue-D), and area (Blue-A2, Blue-A1) were the next high-ranking photosynthesis metrics (ranks 16, 19, 20, and 21, respectively). For laboratory bark spectra, 65% of the photosynthesis metrics had significant mean differences. As with leaf spectra, top-ranking metrics for bark were also concentrated in the red-edge region: RWE-DArea, RE-Mag and RVSI (Table 4.5, bark). Blue-edge derivative metrics, BE-DArea and BE-Mag, were more important than blue absorption-based metrics in bark spectra. At pixel scales, the red absorption wavelength (Red-λ) was the only photosynthesis metric that did not have a significant mean difference among species. The metrics SR, Red-A1, Red-A2 and ARVI were among the top-ten metrics for species discrimination at pixel scales (Table 4.5, pixels). The deciduous species DIPA and LEAM had the lowest SR values, while high-LAI species (CEPE, HYAL, TEOB) had the highest SR values (Fig. 4.2). Other highly-significant metrics included Red-As, Red-W, NDVI, PRI and Red-D (ranks 11, 12, 13, 14, and 16, respectively). Half of the top-ranking metrics at crown scales were photosynthesis metrics (Table 4.5, crowns). In particular, SR and YE-DArea had the highest ranks. The red absorption feature area, asymmetry and width were the next set of important metrics 149 for crown-scale species discrimination, followed by the vegetation indices ARVI, SAVI and EVI (ranks 14, 16, and 18, respectively). 4.4.3. Water absorption metrics In leaf spectra, seven out of the top-ten metrics encompassed some aspect of the NIR water absorption features (Table 4.5, leaves), which had mean locations at 994 and 1175 nm (NIR1-λ and NIR2-λ, Appendix 2.3). Important metrics describing these features were the area, depth and the down-sloping edge inflection magnitude (Table 4.5, leaves). All water metrics at leaf scales had significant differences among species except for WE- λ, and leaves were more strongly separated with NIR2 versus NIR1 absorption-based metrics. In contrast to leaves, there were only two top-ranking NIR water absorption metrics for separating bark spectra (Table 4.5, bark), although all water metrics had significant mean differences. The two indices, NDWI and WBI, were more important for distinguishing bark spectra than the other types of water metrics. At pixel scales, NDWI was the top-ranking metric, followed by NIR1 area metrics (Table 4.5, pixels; Fig. 4.2). Greater pixel-level water absorption was observed in high-LAI species, especially the broad-leaved species CEPE, HYAL and TEOB (Fig. 4.2; NDWI, NIR1-A1). The importance of water metrics decreased substantially at crown scales. The NDWI was ranked 9 (Table 4.5, crowns; Fig. 4.2), but all other water metrics were ranked 38 or greater out of all metrics (77 total). 150 4.4.4. Shortwave-infrared biochemical absorption metrics Only 8 out of 20 SWIR biochemical absorption metrics had significant mean differences at leaf scales. The SWIR3 region was the best of the three absorption features, while SWIR1 and SWIR2 absorption features were either not detected or had minor significance in both leaf and bark spectra (Appendix 2.3). For leaves, SWIR3 area (A1 and A2), depth, asymmetry and width had ranks 9, 11, 13, 17 and 18 out of all 73 metrics. All of these absorption metrics had top-ten ranks for distinguishing species with bark spectra (Table 4.5; bark). In contrast to leaves and bark, the SWIR1 and SWIR2 metrics had high rankings at the pixel and crown scales. The SWIR1-As and SWIR1-W metrics had ranks 8 and 9 out of 77 metrics at pixel scales (Table 4.5, pixels). The SWIR2 absorption feature area (A1 and A2), SWIR1-As and SWIR1-W had the strongest statistical separation at crown scales (Table 4.5; crowns). At both pixel and crown scales, the high-LAI species CEPE, HYAL and TEOB had low SWIR2-A1 (Fig. 4.2). In contrast, low LAI species (DIPA, LEAM) to moderate LAI species (BAEL, HYME) had comparatively high SWIR2-A1 values. 4.4.5. Pixel- and crown-scale spectral mixture analysis fractions Mean values of SMA fractions from ITCs were roughly 40% GV, 15% NPV and 44% shade (+ 1% error; Appendix 2.4) at both pixel and crown scales. Canopies of deciduous DIPA and LEAM had relatively high fractions of NPV and low fractions of GV (Figs. 4.2 & 4.3). In contrast, leaf-on broad-leaf species CEPE, HYAL and TEOB had relatively high fractions of GV and low fractions of NPV. Figure 4.3 151 reveals that there was variability in the crown-level proportions of GV, NPV and shade among individuals of the same species (i.e., conspecific variability). For example, BAEL and HYME (fine compound leaves, moderate LAI) had some individuals with NPV and GV fractions similar to low-LAI crowns of DIPA and LEAM, while other individuals of BAEL and HYME had fractions that resembled the high-LAI crowns of broadleaf species HYAL and TEOB. As an index of nonphotosynthetic vegetation, NPV fractions were expected to correlate positively with bark spectral properties and negatively with leaf properties (e.g., chlorophyll absorption). This was indeed the case: NPV had negative linear correlations with SR, Red-A1, Red-A2 and ARVI photosynthesis metrics (i = -0.75 to -0.78) and NDWI and NIR1-A1 water absorption metrics (r = -0.71 and -0.32, respectively). To quantify the influence of illumination variation on spectral metrics, I calculated the Pearson’s linear correlation between each metric and the SMA shade fraction. Pixel-scale metrics had correlations (r) ranging from -0.85 (BE-DArea) to +0.60 (YE-DArea), with an average correlation of -0.09 (±0.30). Crown-scale metrics had correlations (r) ranging from -0.63 (BE-DArea) to +0.30 (NIR2-A1), with an average correlation of -0.02 (±0.23). 4.4.6. Decision-tree classification 4.4.6.1. Pixel-majority ITC classification ITC classification accuracies were below 69% for DT pixel-majority classifiers (Table 4.6). Including all 77 metrics produced the best overall accuracy (68.2%; Table 4.6). From the analysis of all 77 metrics, the 10 top-ranked metrics were 152 selected according to the number of times the metric was found in decision trees (Table 4.7; pixel scale). These 10 metrics were then applied in a separate DT analysis. This approach produced slightly less overall accuracy than using all 77 metrics (Z = 0.27, not significant at α=0.05; Congalton, 1991). Of the groups of spectral metrics considered, derivative-based metrics yielded the highest overall accuracy. However, the 10 top-ranked metrics included indices, absorption-based, derivative-based and SMA metrics (Table 4.7). In terms of the number of times a metric appeared in decision trees, species were distinguished with a wide variety of absorption features, including blue absorption (Blue-D), yellow edge (YE-DArea), red-well (RW-λ), red-edge (RVSI), NIR water absorption (NDWI, NIR1-A1), and SWIR biochemical absorption (SWIR1-W, SWIR2-A2). Even though they appeared many times in DTs, the ten top-ranked metrics had average node depths between 3 to 5 tiers. A metric that consistently splits data at a primary (root) node would have an average node depth of zero. Therefore, the most widely-used metrics were not necessarily forming primary splits, but also played decisive roles closer to the terminal nodes, where final species labels were determined. The lowest mean node depth was only 3.5 (ARVI), which indicates that no single metric consistently split the data at the root node. 4.4.6.2. Crown-scale ITC classification Overall ITC classification accuracies using crown-scale metrics were slightly higher than with the pixel-majority approach when considering absorption-based, SMA fractions and all metrics (Table 4.6). The best crown-scale accuracy achieved 153 was 70.1% with all 77 metrics (Table 4.8). The selection of the 10 top-ranked metrics (Table 4.7; Crown scale) slightly decreased overall accuracy relative to using all 77 metrics (Table 4.6; Z = 0.58, not significant at α=0.05). Top-ranked metrics were similar to those observed in pixel-scale DT analyses, except SWIR3-D (depth) was more influential at crown scales (Table 4.7; Crown-scale). The SWIR2A1 metric had a mean node depth of 0.7, which is the lowest depth of all other metrics and shows that the SWIR2 area was often the primary split separating species. An example decision tree using all 77 metrics showed that some individuals of high-LAI species (Fig. 4.4 A) were separated from moderate- and low-LAI species (Fig. 4.4 B & C) primarily with the SWIR2 area metric—lower SWIR2-A1, more LAI (Fig. 4.2). The YE-DArea was negative in species with higher LAI (Fig. 4.2; HYAL, HYME, TEOB) because derivative values were from a steeply down-sloping edge in reflectance spectra. Low YE-DArea values were used by the decision tree to separate moderate-LAI individuals (Fig. 4.4b) from individuals with lower LAI (Fig. 4.4c). Photosynthetic pigment, water and SWIR biochemical absorption features distinguished individuals of BAEL, LEAM and DIPA (Fig. 4.4c). These species had the most interspecific confusion (Table 4.8). For comparison with previous research in Chapter 3, I also present the overall accuracy achieved when using reflectance narrowband (hyperspectral) data in DTs instead of spectral metrics. With crown-scale reflectance spectra, a linear discriminant analysis (LDA) classifier had 92.1% overall accuracy using 30 optimally-selected bands and 61.2% accuracy with all 161 bands. I used the same 154 methods to select optimal spectral metrics for crown-scale LDA classification. The best classification had an overall accuracy of 84.1% (DIPA User’s = 85.5%, Producer’s = 87.7%) with the inclusion of 18 optimal metrics, 6% lower accuracy than with 30 reflectance bands (Z=2.60, significant at α=0.05). With the DT classification scheme used in this chapter, crown-scale overall accuracy was only 49.5% using either 30 or 161 bands (Table 4.6). Although the DT classifier was 20.6% more accurate when applied to spectral metrics over reflectance data, the LDA classifier applied to reflectance data outperformed the best metric-based DT classifier by 22.0% (Chapter 3; Z=6.17, significant at α=0.05). 4.5. Discussion 4.5.1. Crown structure and phenology detected by spectral mixture analysis fractions At pixel and crown scales, tropical tree spectra are mainly a mixture of GV (e.g., leaves, green bark, epiphytes, lianas), NPV (e.g., non-photosynthetic bark, fruits, flowers) and shade. The biophysical properties of the crown, such as leaf density, angles and clumping, determine the fractions of these components that form the mixed signal within an image pixel. Phenological variation in trees, such as leaf aging and leaf drop, drought stress, and flowering, will also affect the multi-temporal spectral signature of tree species and may help discriminate individual or assemblages of plant species (Blackburn & Milton, 1995; Key et al., 2001). In tropical forests, previous research has shown that individual deciduous trees with 155 low leaf cover have a clear spectral contrast with leaf-on trees in the visible and NIR regions of the spectrum (Bohlman & Lashlee, in press). As noted in Chapter 3, the DIPA and LEAM trees in our hyperspectral imagery were mainly deciduous (i.e., low crown-level LAI) at the time of image acquisition. Spectral mixture analysis fractions revealed that the DIPA and LEAM populations as a whole had crowns composed of relatively high percentages of NPV materials relative to other species (Fig. 4.3), which can be attributed to non-photosynthetic branch and trunk tissues exposed to the sensor. BAEL trees also had relatively high amounts of NPV. Although BAEL was expected to have high mature leaf cover during image acquisition (O’Brien, 2001), I attribute high NPV in some BAEL individuals to their architectural properties rather than deciduousness; BAEL trees have compound leaves with fine leaflets and the overall architectural arrangement of these leaves exposed conspicuous white bark when viewed from above by the sensor (see Fig. 3.2a in Chapter 3). ITCs with relatively low LAI, such as individual DIPA, were easily identified in both the hyperspectral reflectance imagery (Fig. 4.5; natural color) and the images of spectral metrics (Fig. 4.5; SMA fractions, NDWI, NDVI, SWIR2-A1). In the SMA false-color image, ITCs with low-LAI were clearly seen as groups of pixels with high NPV (red) in a matrix of high-GV canopy (green) and high-shade gaps (blue). 4.5.2. Species differences in photosynthetic-pigment absorption features My analyses highlighted the importance of the red-edge for species discrimination with leaf and bark spectra. These results are consistent with work by 156 Cochrane (2000), who also found this region useful for distinguishing tropical tree species with their leaf spectra. The shape of the red edge is heavily influenced by photon absorption and florescence from chlorophyll in tissues (Gates et al., 1965; Zarco-Tejada et al., 2000). Research has shown that high concentrations of chlorophyll in leaves widens and deepens red-well absorption and shifts the red-edge inflection wavelength (RE-λ) toward longer wavelengths (Blackburn, 1998). The RVSI and RE-λ were the strongest metrics for discriminating leaf spectra. The RVSI was designed to detect red-edge curve concavity or convexity relative to a baseline (e.g., continuum removal). The index was originally used to track vegetation communities through time, not spectral properties of tissues. I found that leaf-scale RVSI was positively correlated with RE-λ (r = +0.72), and so positive RSVI values were related to a red-shift in leaf RE-λ. In contrast to leaves, bark spectra had low to no chlorophyll and so there was relatively low red-well absorption (Fig. 4.1b), increased red-well reflectance (RWRefl), decreased slope of the red-edge (RE-Mag), and a shift in the red-edge position toward lower wavelengths—a blue-shift (Appendix 2.1 & 2.2). There were significant differences among species with bark red-edge metrics such as RE-Mag. Some bark spectral variation in the visible to red-edge regions is likely related to a strong ultraviolet-blue absorption wing from 400 to 900 nm created by lignin and tannins (Elvidge, 1990), while overlapping red-well absorption may be caused by algae, bryophytes, moss and lichen on the bark. I found that BAEL bark had the highest mean RE-Mag, while TEOB bark had the lowest mean. BAEL had a lightbrown to red bark color, and no evidence of surface chlorophyll from other 157 organisms; these properties gave BAEL bark relatively high red-well reflectance and a steep red-edge slope (Fig. 4.1b-BAEL). In contrast, TEOB bark samples were darker brown with green mottling (providing evidence of chlorophyll in the tissues) and their red-well reflectance was relatively low with a more shallow slope (Fig. 4.1b-TEOB). The RVSI of bark was not as strongly correlated with RE-λ (r = +0.38) as with leaf spectra, but was mostly related to red-edge slope, RE-Mag (r =+0.52). The red absorption feature also had potential for discriminating species at pixel and crown scales. The SR and red absorption area (Red-A1 and Red-A2) were among the top-ranking variables in tests of mean differences among species (Table 4.5). A typical pixel from a high-LAI species, such as HYAL, revealed low red-well reflectance due to chlorophyll absorption and high NIR reflectance due to multiple scattering among transmittive leaves (Fig. 4.6); and consequently, the metrics SR, NDVI and Red-A1 were relatively high. In contrast, a pixel from low-LAI DIPA had a higher concentration of bark in the sensor IFOV, which translated into relatively high red-well reflectance due to lower chlorophyll absorption and low NIR reflectance due to less photon scattering within the crown (Fig. 4.6); and thus, the metrics SR, NDVI and Red-A1 were relatively low. I found that the ANOVA rankings (Table 4.5) were not necessarily predictive of overall metric utility in DTs; for example, red-well wavelength position (RW-λ) was a key metric in DTs at both pixel and crown scales (Table 4.7), but was ranked 43 and 51 out 77 metrics based on pixel and crown F statistics, respectively (Appendices 2.1-2.4). This metric mainly separated BAEL from the other species 158 (Fig. 4.2, RW-λ). BAEL had the highest mean red-well reflectance relative to other species, and this spectral shape tended to shift the red-well position toward smaller wavelengths. The RW-λ was observed classifying a sub-set of BAEL in Figure 4.4 B, down at DT tier 3 (root tier is 0); and thus, RW-λ was important for lower-tier decisions, even though overall mean species differences were weak (i.e., low ANOVA F statistic). 4.5.3. Species differences in water absorption features The NIR1 and NIR2 water absorption features dominated top-ranking metrics for leaf spectra. Internal leaf structure, such as the air-cell wall interfaces in the spongy mesophyll, heavily scatters NIR photons and increases the expression of absorption features (Gates et al., 1965; Gausman, 1985). Species differences in water absorption depth, width and area may thus result from a combination of factors that affect internal leaf photon scattering and absorption, such as leaf thickness, water content and the distribution of air-cell wall interfaces. Bark water absorption metrics were more variable than those for leaves. For example, species standard deviations of NDWI ranged from 0.004 (DIPA) to 0.021 (HYME) for leaves while 0.032 (CEPE) to 0.118 (DIPA) for bark. This variability in bark spectra made species less distinct relative to leaves (in terms of F statistic and significant pairs). Bark laboratory specimens were cleared of large bryophytes and mosses, which when wet would further increase water absorption variability within species. In my laboratory data, much of the within-species bark variability comes from spectral differences between branch and trunk bark. 159 One striking example was DIPA. Branches sampled from young DIPA trees had smooth, green bark. Spectra from these branches had clear chlorophyll absorption features at 680 nm (Fig. 4.1c), and so they can not be considered as “pure” NPV. In contrast, trunk bark samples from mature DIPA crowns were woody, rough, and brown (i.e., nonphotosynthetic). The green bark spectra had more notable NIR water absorption features than the woody trunk bark (Fig. 4.1c). These data indicate that bark spectral properties can vary considerably within a population of tropical tree species. Williams (1991) found that first-year twig spectra for conifer and hardwood species showed evidence of chlorophyll and NIR water absorption, and the depth of these features decreased considerably in second-year twigs from the conifer species, which were more woody. It thus appears that bark chlorophyll and NIR water absorption features are correlated, likely because photosynthesis requires water. Considering all species, there were significant (p≤0.01) differences in means of NIR water absorption metrics between leaf and bark laboratory spectra. The ratiobased indices, WBI and NDWI, had significantly greater values in leaves than bark; however, absorption-based metrics such as EWT and area (NIR1 or NIR2) were significantly higher in bark relative to leaves. Bark NDWI tended to be negative because NIR reflectance at 862 nm was low relative to the absorption feature at 1239 nm, while the opposite trend was found in leaves. This finding is similar to Gao (1996), who found that the NDWI metric was negative in non-photosynthetic materials (e.g., dead grass). The larger NIR water absorption area in bark is seen clearly in Figure 4.1 when comparing BAEL, DIPA and HYAL mean spectra in bark (a) to leaves (c). Large NIR water absorption features, especially in the 1200 nm 160 region, are unusual in dry bark spectra from temperate regions (Elvidge, 1990; Roberts et al., 2004). However, my bark samples from tropical rain forest trees were not dried and evidently contained a great deal of moisture, especially in green bark. Bark water absorption not only manifests as broad NIR features but it also obscures SWIR biochemical absorption features that are otherwise prominent in drier bark material (Elvidge, 1990). The depth, width and area of NIR water absorption features tended to increase from the centimeter spatial scales of leaf and bark laboratory spectra to the 1.6-m scale of HYDICE image pixels (Appendix 2.3). The depth of these features can be partly attributed to poor radiometric calibration and signal-to-noise in the HYDICE sensor (discussed in Chapter 3). However, the three-dimensional stacking of leaves and branches within the crown, which increases the path length of photons and increases their multiple-scattering and absorption, is expected to accentuate tissuelevel biochemical and biophysical properties (Asner, 1998; Curran, 1989; Roberts et al., 2004). Water absorption metrics calculated at pixel and crown scales should thus respond to the combination of water content in GV and NPV tissues and the structural distribution of these tissues within the crown (Dennison et al., 2003; Gao, 1996; Roberts et al., 2004). The water absorption signal should be greater with more leaf area relative to branch area because leaves are more transmittive, thereby permitting more photon scattering. This effect can be observed in Figure 4.6, where a leaf-off DIPA pixel spectrum had lower NDWI and NIR1 and NIR2 water absorption area relative to a leaf-on HYAL pixel spectrum. Furthermore, the pixels within DIPA crowns formed dark ITCs in the NDWI image (Fig. 4.5). 161 In terms of the F statistic and number of significant paired comparisons, species differences in NIR water absorption metrics were greater at pixel scales than at leaf scales. This finding suggests that pixel-scale biophysical differences among species affect NIR water absorption in a way that amplifies species separability. This scaling effect may explain why several NIR water absorption metrics were among the primary variables for separating species in DTs at both pixel and crown scales (Table 4.7). Also, results indicate that both NIR1 (e.g., NIR1-A1, WBI) and NIR2 (e.g., NDWI) absorption features were equally important variables for species classification. 4.5.4. Species differences in shortwave infrared absorption features There were also strong differences among species in metrics covering the SWIR3 absorption feature in both leaf and bark spectra. The SWIR3 absorption feature results from a combination of overlapping absorptions by protein, nitrogen, starch, lignin, cellulose, and sugars (Curran, 1989; Elvidge, 1990). In HYDICE data, the deepest part of the feature had a mean wavelength of 2294 nm and 2308 nm for bark and leaves, respectively (SWIR3-λ, Appendix 2.3), which is a wavelength region predominantly associated with absorption from N-H and C=O bond stretching within the amino acids of proteins (Osborne & Fearn, 1986). As mentioned, the tails of strong water absorption features in the SWIR region centered at 1400 and 1940 nm can mask the expression of more subtle SWIR absorption features in green vegetation (Curran, 1989; Elvridge, 1990; Kokaly & Clark, 1999). There were +0.20 and -0.38 correlations between NIR2-A1 and SWIR3-A1 in leaf and bark 162 spectra, respectively. If the NIR2 absorption feature area is considered as an indicator of tissue water concentration, it appears that absorptions by bark biochemical constituents (e.g., lignin, cellulose, proteins) are negatively associated with water content. It is unclear why there is a weak positive correlation between leaf water absorption and SWIR3 biochemical absorption. Biochemical assays acquired at the time of spectral measurement would be necessary to establish conclusive links between species biochemistry and observed reflectance patterns (e.g., Kokaly & Clark, 1999). Although the SWIR3 absorption feature had strong statistical separation among species bark spectra, the importance of the feature diminished at pixel and crown scales. This may be due to low signal-to-noise in the HYDICE sensor and low reflectance in the SWIR3 region (Basedow et al., 1995; Chapter 3). In contrast to SWIR3, the SWIR1 and SWIR2 absorption features were detected in only 25% and 7% of laboratory leaf spectra, respectively, and in 70% and 15% of bark spectra, respectively. However, metrics incorporating the area, asymmetry and width of SWIR1 and SWIR2 features were among the top-ranked discriminatory metrics at pixel and crown scales. The SWIR1 feature is associated with lignin (primarily), starch, protein and nitrogen absorption and the SWIR2 feature is associated with protein and nitrogen (primarily), starch and cellulose absorption (Curran, 1989). One explanation why SWIR1 and SWIR2 metrics were important for species discrimination in pixel and crown spectra is that overlapping absorptions broadened due to multiple scattering within the crown, thereby permitting feature detection in pixel and crown-scale spectra. This SWIR absorption broadening should increase as 163 the fraction of bark tissues increases and as the fraction of leaves decreases. A lower fraction of GV should correlate with lower SWIR water absorption and allow the detection of bark SWIR1 and SWIR2 features. At crown scales, SWIR1-A2 and SWIR2-A2 had positive correlations with NPV (r = +0.65 and +0.74, respectively) and negative correlations with GV (r = -0.70 and -0.78, respectively), suggesting that crowns with lower LAI and more exposed NPV had more strongly expressed SWIR1 and SWIR2 features. Furthermore, low- to moderate-LAI DIPA, LEAM and BAEL had the highest mean SWIR2 area, while high-LAI CEPE, HYAL and TEOB had the lowest mean area (Fig. 4.2). This pattern is illustrated in Figure 4.6, where a typical DIPA pixel spectrum had low SWIR1 and SWIR2 area relative to a HYAL pixel. DIPA pixels formed bright ITCs in the SWIR2-A1 image (Fig. 4.5). This image also reveals a pattern of noise in SWIR2, seen as diagonal stripes. The absorption feature may have been more important at crown scales because averaging many within-crown pixels will reduce spurious spectral variability due to sensor noise. This reduction in variance can be seen as lower standard deviations at crown scales relative to pixel scales in Figure 4.2 (SWIR2-A1). 4.5.5. Individual tree crown classification with decision trees Despite the many statistically significant differences detected among species in the various spectral metrics, overall ITC classification accuracies were no greater than 71% using pixel-scale or crown-scale spectra. Some derivative-based and absorption-based metrics had null values when absorption features could not be detected, such as when calculating SWIR2 metrics when they were masked by water 164 absorption. These null values were not necessarily random values, and so they should be included in the classifier. An advantage of the DT classifier was that it could form splits on null values (coded as very negative numbers), whereas null values are excluded in a traditional classifier such as maximum likelihood. Overall pixel-majority DT classification accuracies were up to 12% lower when null values were excluded from the analysis, indicating that there was an advantage to using null values in DTs. The crown-scale DT classifier had better performance than the pixel-majority technique (Table 4.6). There are several reasons to explain higher crown-scale accuracy. Crown-scale DTs had fewer nodes than pixel-scale DTs used in pixelmajority classification because there was lower variability in crown-scale spectra (Appendix 2.1-2.4; Fig. 4.2), as they were averages of pixel-scale spectra. Despite pruning, pixel-scale DTs still had complex structures that made class decision rules more unstable. Furthermore, important crown-scale metrics changed less often in DT depth (i.e., low average node depth) relative to pixel-scale metrics. Again, lower crown-scale variability permitted more stable structural relationships among metrics in separating species, thereby leading to greater ITC classification accuracy. The best crown-scale accuracy was achieved by including all 77 metrics in decision trees. The top 10 metrics used in decision rules included 4 absorptionbased metrics, 2 derivative-based metrics, 3 indices, and 1 SMA fraction (Table 4.7). Red, NIR and SWIR features were all characterized by these top-ranking metrics. It thus appears that a wide array of information extraction techniques applied to the 165 whole 400-2500 nm spectrum should be considered when classifying tree species from hyperspectral imagery. The crown-scale DT classification using 77 metrics had an overall accuracy of 70.1%. The 80.2% Producer’s and 79.3% User’s accuracies for DIPA (Table 4.8) are also encouraging for mapping this species of high conservation value (see Chapter 3). Confusion of DIPA with BAEL and LEAM (Table 4.8) was likely because these species had low crown LAI at the time of image acquisition, which would result in similar fractions of GV and NPV tissues and comparable photon scattering environments. 4.5.6. Comparison with past research and recommendations for future research As found in Chapter 3, the analysis of crown-scale data provided better classification accuracy than using pixel-scale data. Both DT and LDA classifiers thus indicated that crown-scale hyperspectral data may be sufficient for tropical rain forest species classification at the canopy scale. Airborne hyperspectral sensors, such as AVIRIS and HyMap, are typically flown with altitudes that provide 4- to 20m pixels. Since canopy-level tree crowns in old-growth forest at La Selva generally have 20-27 m diameters (Chapter 2), a single crown could contain 1 to < 40 pixels, depending on crown size and sensor resolution. By delineating ITCs and averaging their 1.6-m pixels, crown-scale spectra in this research had minimal spectral mixing with neighboring vegetation and soil. These factors would be expected to diminish species signal and subsequent separability as pixel size increases. 166 Consistent with my results in this chapter, previous analyses found that optimally-selected bands using LDA were located across the VIS, NIR, and SWIR regions (Chapter 3). Several of the 20 most important reflectance bands were found in SWIR1, SWIR2, and SWIR3 absorption features; and thus, both DT and LDA classification schemes identified the SWIR region of the spectrum as important for species discrimination. I recommend that future research use imaging spectrometers that measure the full 400 to 2400 nm spectral range. Engineering improvements in sensor signal-to-noise in the SWIR region should lead to a greater ability to discriminate species with distinct phenology or structure. Overall accuracy was 22% greater when using optimal reflectance bands and LDA (Chapter 3) relative to spectral metrics and decision trees. LDA and reflectance bands had 6% higher accuracy than LDA with spectral metrics. These results indicate that future research should explore methods to exploit reflectance spectra rather than developing new spectral metrics. Why did LDA applied to optimal reflectance bands outperform the DT or LDA classifier with spectral metrics? One explanation is that illumination variation drives spectral differences among species, such as relatively low NIR reflectance from lowLAI DIPA relative to leaf-on HYAL. Illumination is best captured by reflectance spectra as opposed to spectral metrics, whose brightness variation is minimized from continuum removal, derivative analysis or band ratios. If illumination was the dominant factor in distinguishing species, I would have expected the shade fraction to be among the top metrics in ANOVA tests or in decision trees. Instead, the shade fraction was ranked at 60 and 59 out of the 77 metrics in terms of F statistics for 167 pixel and crown scales, respectively, and it was ranked at 56 and 28 for pixelmajority and crown-scale classifications in terms of times found in decision trees. Also, shade was not included as an optimal metric in LDA classification. These results suggest that illumination variation is not a principal factor in discriminating species. In general, spectral metrics were not well correlated with shade, indicating that species were mostly separated according to biochemical absorptions rather than illumination. However, of the top-ten metrics in decision trees, the NPV, RVSI and YE-DArea metrics were correlated to shade at pixel and crown scales (Table 4.7; r = -0.63 to +0.60). Illumination may thus account for some variation in metrics, but it does not appear that it aids species discrimination. My comparison of results with Chapter 3 indicates that LDA is a superior classifier, regardless of using spectral metrics or reflectance bands. LDA considers all predictor variables simultaneously to build a decision space using predictor variable covariance. In contrast, the decision structure of a DT is built one variable at time and decisions at terminal nodes are dependent upon those made at higher nodes. The DT is thus more susceptible to misclassification errors when data variability causes many different possible splits, as is the case when using highlyvariable spectra from ITCs in tropical rain forest. Despite better LDA performance, I found decision trees useful for both ranking the importance of spectral metrics and visualizing the decision rules in species discrimination. 168 4.6. Conclusions A major goal of this chapter was to assess airborne hyperspectral imagery for the species-level classification of individual tree crowns (ITCs) in a tropical rain forest. The spectral metrics used to train the decision-tree (DT) classifier were expected to capitalize on the detailed spectral information contained in the imagery. The overall accuracy of ITC classification was 22% lower than when using the LDA classifier (Chapter 3). Despite this weaker accuracy, the decision tree classifier was a useful tool for ranking important spectral metrics and for visualizing the hierarchical decision space used in the classification. Statistical tests and classifier variable-selection analyses revealed that there were detectable differences among species absorption features. Properties of the red chlorophyll, NIR water and SWIR biochemical absorption features were all found to differentiate species from centimeter scales of leaf and bark tissue to pixel or crown scales measured by the airborne spectrometer. Tree structure and phenology at the time of image acquisition were driving factors that influenced spectral separability of species. For example, low leaf area crowns had low NIR water absorption due to more exposed non-photosynthetic tissues and a relatively weak photon-scattering environment. Several spectral metrics explored in this chapter, such as the NPV fraction, NDWI, and SWIR2 area, revealed the immense potential of hyperspectral metrics for mapping deciduous crowns in tropical rain forest canopies. The capability to detect deciduous crowns could permit the study of species phenology across large spatial extents and may prove a useful tool for monitoring changes in tree phenology, stress or mortality due global climate change. 169 These results also have important implications for studies that seek to estimate tropical forest biochemical and biophysical parameters over broad spatial scales using spectral metrics derived from airborne or satellite imagery. Attempts have been made to estimate aboveground biomass using a positive correlation with metrics such as NDVI or other indices (Foody et al,. 2001; Thenkabail et al., 2004). Although leaf-off deciduous trees may represent a substantial fraction of standing live biomass, their unique crown spectra will influence metric values measured at coarse spatial scales. For example, the NDVI value in a 30 x 30-m Landsat pixel will go down with increasing fraction of deciduous crowns (e.g., Fig. 4.5-NDVI). The fractional abundance of deciduous trees within a coarser scale pixel should be considered in prediction models, especially if the imagery is acquired when a large proportion of trees are leaf-off. This generally occurs in the dry season when there is also the greatest opportunity to acquire cloud-free imagery. 170 Table 4.1. Formulas for narrow-band, ratio-based indices (ρ is reflectance at a specific wavelength in nm). Wavelengths chosen are the closest HYDICE wavelengths to the formulas in the cited literature. Vegetation Indices Simple Ratio Tucker, 1979 ρ798 Jordan, 1969 SR = —— Ρ679 Normalized Difference Vegetation Index ρ798 - ρ679 NDVI = ————— ρ798 + ρ679 Soil-Adjusted Vegetation Index 1.5 * ρ798 - ρ679 SAVI = ——————— ρ798 + ρ679 + 0.5 Turner, 1979 Rouse et al., 1973 Huete, 1988 Photochemical Reflectance Index ρ532 - ρ568 PRI = ————— ρ532 + ρ568 Gamon et al., 1997 Enhanced Vegetation Index ρ798 - ρ679 EVI = ————————————— 1 + ρ798 + 6 * ρ679 – 7.5 * ρ482 Huete et al., 2002 Atmospherically Resistant Vegetation Index ρ798 - 2 * ρ679 + ρ482 ARVI = —————————— ρ798+ 2 * ρ679 - ρ482 Kaufman & Tanre, 1992 Red-Edge Vegetation Stress Index ρ719 + ρ752 RVSI = ————— – ρ730 2 Merton, 1998 171 Table 4.1. (continued). Liquid Water Content Indices Water Band Index ρ902 WBI = —— ρ973 Peñuelas et al.,1997b Normalized Difference Water Index ρ862 - ρ1239 NDWI = —————— ρ862 + ρ1239 Gao, 1996 172 Table 4.2. Study tree species attributes (Adapted from O’Brien, 2001 and Frankie et al., 1974). Leaf cover is for late-March to early-April, and is what would be expected for the majority of individuals for each species based on available literature data and personal field observations. Tree species Code Leaf Leaf 3/30 [family or sub-family] Phenology Exchange Leaf Functional Timing Cover Group Balizia elegans BAEL Deciduous Annual High (Ducke) Barneby & Grimes [Mimosoideae] Ceiba pentandra CEPE Deciduous Annual High Gaertn. [Bombacaceae] Dipteryx panamensis DIPA Deciduous Annual Low (Pittier) Record & Mell [Papilionoideae] Hymenolobium mesoamericanum HYME Deciduous Sub-annual High Lima [Papilionoideae] Hyeronima alchorneoides HYAL Evergreen Continuous High Allemão [Euphorbiaceae] Lecythis ampla LEAM Deciduous Annual Low Miers [Lecythidaceae] Terminalia oblonga TEOB Evergreen Continuous High (Ruiz & Pav.) Steud. [Combretaceae] 173 174 Table 4.3. Definition of spectral features analyzed (See Fig. 4.1a). Minimum and maximum wavelengths are for HYDICE band centers and the number of bands within the range is controlled by HYDICE band spacing. Spectral Feature Abbrev. Min. λ Max. λ No. Polynomial (nm) (nm) Bands Order Derivative-based analyses (polynomial fitting) Blue Edge (up-sloping) BE 491.0 532.1 9 5 Green Peak GP 532.1 581.9 9 5 Yellow Edge (down-sloping) YE 549.6 581.9 6 5 Red Well RW 643.3 698.7 7 5 Red Edge (up-sloping) RE 679.3 751.5 8 5 NIR Water1 Edge (up-sloping) NE1 958.2 1075.3 9 3 NIR Water2 Edge (down-sloping) NE2 1105.1 1164.9 5 3 SWIR Edge (up-sloping) SE 1494.5 1638.0 12 5 Absorption-based analyses Blue Absorption Blue 460.5 537.7 17 n/a Red Absorption Red 588.9 751.5 19 n/a NIR Water Absorption 1 NIR1 902.0 1075.3 13 n/a NIR Water Absorption 2 NIR2 1105.1 1253.9 11 n/a SWIR Absorption 1 SWIR1 1650.5 1771.9 11 n/a SWIR Absorption 2 SWIR2 2045.8 2221.6 19 n/a SWIR Absorption 3 SWIR3 2240.2 2365.9 15 n/a 174 Table 4.4. Summary of hyperspectral metrics organized by methods (in bold) and spectral region and dominant absorption feature (in italics). Indices Absorption-based Derivative-based SMA Visible – Photosynthetic pigments SR Blue-λ,D,W,A1,A2,As BE-λ,Mag NDVI Red-λ,D,W,A1,A2,As GP-λ,Refl SAVI YE-λ,Mag PRI RW-λ,Refl EVI RE-λ,Mag ARVI BE-DArea RVSI YE-DArea RWE-DArea RWE-DNArea RWE-2DNArea Near infrared – Water WBI EWT NE1-λ,Mag NDWI NIR1-λ,D,W,A1,A2,As NE2-λ,Mag NIR2-λ,D,W,A1,A2,As Shortwave infrared – Other biochemicals SWIR1-λ,D,W,A1,A2,As SE-λ,Mag SWIR2-λ,D,W,A1,A2,As SWIR3-λ,D,W,A1,A2,As Full-spectrum – All absorption features GV NPV Shade RMSE λ = wavelength, Mag = derivative magnitude, Refl = percent reflectance, DArea = area under 1st derivative, DNArea = area under normalized 1st derivative, 2DNArea = area under 2nd derivative, D = depth, λ = wavelength, W=width, A1 = area calculated using width and depth, A2 = area calculated using tabulation, As = Asymmetry. 175 Table 4.5. Ranking of metrics based on F statistics and significant pair-wise comparisons. Rank Bark Leaves Pixels Crowns 1 SWIR3-λ RVSI NDWI SWIR2-A2 2 SWIR3-D NIR2-A1 NIR1-A1 SR 3 NDWI NIR2-A2 SR SWIR2-A1 4 SWIR3-As NIR2-D NIR1-A2 SWIR1-As 5 SWIR3-W NE2-Mag Red-A2 SWIR1-W 6 WBI RE-λ Red-A1 YE-DArea 7 RWE-DArea NIR1-A1 ARVI Red-A2 8 RE-Mag NIR1-A2 SWIR1-As Red-As 9 SWIR3-A2 SWIR3-A1 SWIR1-W NDWI 10 RVSI NIR1-D NPV Red-W 176 177 Table 4.6. Accuracy of individual tree crown classification using leave-one-out crossvalidation and a decision tree classifier. Pixel-majority: pixel metrics were classified and then aggregated into crown objects using a majority class rule. Crown-scale: crown-scale metrics classified. Pixel-majority Crown-scale Vars Species DIPA Species DIPA Metrics Overall User/Producer Overall User/Producer Accuracy Accuracy Accuracy Accuracy Indices 9 53.3 76.2 / 59.3 49.5 68.5 / 61.7 Derivative-based 21 65.4 89.3 / 61.7 56.5 78.3 / 66.7 Absorption-based 43 62.6 0.0 / 67.9 65.9 81.1 / 74.1 SMA fractions 4 44.4 78.6 / 27.2 46.7 64.7 / 40.7 All metrics 77 68.2 82.4 / 69.1 70.1 79.3 / 80.2 Top-ranked metricsa 10 67.3 75.0 / 70.4 67.3 75.0 / 70.4 Reflectance bandsb 30 55.6 87.5 / 34.6 49.5 64.7 / 54.3 Reflectance bands 161 54.2 89.7 / 32.1 49.5 66.7 / 61.7 a Top-ranked metrics were those most common in trees when using all metrics. b Selected using step-wise linear discriminant analysis (Chapter 3). 177 Table 4.7. Ten top-ranked spectral metrics for decision trees (DTs) at pixel and crown scales. Metrics are ranked by times found in DTs. Shade corr. is the Pearson’s linear correlation of the metric with the SMA shade fraction. There were a total of 10,700 DTs built at each scale of analysis, and some metrics appeared more than once in DTs. Pixel scale Rank Metric Times in DTs Shade Corr. 1 RW-λ 95961 -0.02 2 NDWI 81305 0.18 3 NIR1-A1 76031 0.09 4 SWIR1-W 44616 -0.05 5 NPV 41137 -0.52 6 RVSI 40241 -0.63 7 SWIR2-A2 34957 0.02 8 SWIR1-As 33623 -0.05 9 YE-DArea 32499 0.60 10 Blue-D 30283 -0.36 Crown Scale Rank Metric Times in DTs Shade Corr. 1 RW-λ 21486 0.21 2 RWI 17004 0.18 3 NDWI 16618 0.09 4 SWIR3-D 14372 0.17 5 YE-DArea 13030 0.28 6 NPV 12128 -0.42 7 SWIR2-A2 9836 -0.16 8 RVSI 9800 -0.36 9 SWIR2-A1 9531 -0.15 10 SWIR1-W 7056 0.12 178 179 Classification Table 4.8. Error matrix for crown-scale classification using all spectral metrics (Kappa = 0.62). Species BAEL CEPE DIPA HYAL HYME LEAM TEOB Total Producer’s BAEL 11 8 2 6 2 29 37.9% CEPE 4 2 2 2 10 40.0% DIPA 5 3 65 2 1 5 81 80.2% Field Reference HYAL HYME LEAM 2 2 1 1 1 1 6 28 2 1 8 14 2 34 14 21 82.4% 57.1% 66.7% 179 TEOB 4 1 20 25 80.0% Total 21 13 82 37 16 21 24 214 User’s 52.4% 30.8% 79.3% 75.7% 50.0% 66.7% 83.3% 70.1% Percent Refelctance 0.8 A 0.7 0.6 RE 0.5 0.4 GP 0.3 Blue BE YE 0.2 0.1 NIR1 NE2 NE1 NIR2 SWIR1 SE SWIR2 SWIR3 BAEL CEPE DIPA HYAL HYME LEAM TEOB RW/Red 0 350 600 850 1100 1350 1600 1850 2100 2350 Wavelength (nm) Percent Refelctance 0.8 B 0.7 NIR1 NIR2 SWIR1 0.6 SWIR2 SWIR3 RE 0.5 0.4 RW 0.3 0.2 BAEL CEPE DIPA HYAL HYME LEAM TEOB 0.1 0 350 600 850 1100 1350 1600 1850 2100 2350 Wavelength (nm) Percent Reflectance 0.8 0.7 C DIPA 0.6 0.5 Trunk 0.4 Branch 0.3 0.2 0.1 0 350 600 850 1100 1350 1600 1850 2100 2350 Wavelength (nm) Figure 4.1. Laboratory spectra for study species. A) Mean spectra of leaf samples, B) Mean spectra of bark samples, C) Dipteryx (DIPA) trunk and branch bark spectra. All spectra are convolved to HYDICE bands. 180 1.0 0.8 0.8 0.6 0.6 GV NPV 1.0 0.4 0.2 0.4 0.2 0.0 0.0 BAEL CEPE DIPA HYAL HYME LEAM TEOB BAEL 0.14 NIR1-A1 NDWI DIPA HYAL HYME LEAM TEOB 20.0 0.10 0.06 0.02 15.0 10.0 5.0 -0.02 -0.06 0.0 BAEL CEPE DIPA HYAL HYME LEAM TEOB BAEL CEPE DIPA 669.0 25.0 668.0 20.0 667.0 15.0 SR RW-λ CEPE 666.0 665.0 HYAL HYME LEAM TEOB 10.0 5.0 664.0 0.0 BAEL CEPE DIPA HYAL HYME LEAM TEOB BAEL CEPE DIPA HYAL HYME LEAM TEOB -0.001 8.0 YE-DArea SWIR2-A1 10.0 6.0 4.0 2.0 0.0 -0.006 -0.011 -0.016 BAEL CEPE DIPA HYAL HYME LEAM TEOB BAEL CEPE DIPA HYAL HYME LEAM TEOB Figure 4.2. Species mean (bars) and standard deviation (error bars) for selected spectral metrics calculated from pixel-scale (white; n=300) and crown-scale spectra (gray; n=214). 181 NPV Species BAEL CEPE DIPA HYAL HYME LEAM TEOB GV Shade Figure 4.3. Spectral mixture analysis (SMA) fractions of green photosynthetic vegetation (GV), non-photosynthetic vegetation (NPV) and shade for individual tree crowns. Ternary diagram values are the individual tree crown fractions from crown-scale spectra. 182 SWIR2-A1 < 3.7 | YE-DArea < -0.007 SWIR2-A2 < 1.4 183 NDWI < 0.09 YE-DArea < -0.01 SWIR3-D < 0.06 NPV < 0.09 PA DI AM LE PA DI L PA EL BA EVI < 0.084 SE-Mag < 0.0007 SAVI < 0.49 E BA B NE2-λ < 1114.23 DI HY ME BA EL HY ME NIR1-As < 0.93 SWIR1-D < 0.07 AM LE PE CE PA DI L E A RWI < 0.10 SWIR2-W < 97.74 NDWI < -0.01 RW-λ < 666.2 BA PA DI ME HY TEOB TEOB HYAL NDWI < 0.02 CEPE HY AL RWI < 1.1 HYAL BA EL SWIR1-W < 39.6 C Figure 4.4. An example decision tree constructed to classify tree species with all 77 crown-scale spectral metrics. Species codes are listed in Table 4.2. Node depth is scaled to the change in deviance relative to the parent node. 183 Color SMA DIPA HYAL NDWI NDVI SWIR2-A1 HYAL GV = 54% NPV = 0% Shade = 46% DIPA GV = 21% NPV = 33% Shade = 46% NDWI = 0.08 NDVI = 0.91 SWIR2-A1 = 4.1 NDWI = 0.01 NDVI = 0.69 SWIR2-A1 = 8.9 Figure 4.5. A 300 x 300-m section of hyperspectral data and derived metrics over individual Dipteryx (DIPA) and Hyeronima (HYAL) trees, delineated with yellow polygons. The natural color image displays reflectance bands 482, 550, and 679 nm. The SMA fraction image displays (Red = NPV, Green = GV, Blue = Shade). High values of NDWI, NDVI and SWIR2-A1 are white. Mean values of SMA fractions, NDWI, NDVI and SWIR2-A1 are shown for the DIPA and HYAL individuals. 184 Percent Reflectance 1 0.9 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0 DIPA HYAL 350 600 850 1100 1350 1600 1850 2100 2350 Wavelength (nm) Metric GV NPV Shade NDVI SR RVSI Red-A1 YE-DArea NDWI NIR1-A1 NIR2-A1 SWIR1-A1 SWIR2-A1 DIPA 19.9 61.2 18.9 58.3 3.8 1.1 49.9 0.39 -1.8 7.3 11.8 4.1 6.1 HYAL 82.1 12 16.7 89.9 18.9 2.7 101.8 -1.53 7.8 10.3 14.9 2.9 1.4 Figure 4.6. A reflectance spectrum and associated metrics for a single pixel from the Dipteryx (DIPA) and Hyeronima (HYAL) crowns delineated in Fig. 4.5. Values for SMA fractions, NDVI, RVSI, YE-DArea and NDWI are multiplied by 100. 185 CHAPTER 5: Classification of tree species with multiple endmember spectral mixture analysis 5.1. Introduction Species-level mapping of individual tree crowns (ITCs) using remote sensing technology has immense potential for extending our understanding and monitoring of species distributions and organization in the face of global climate change and increased pressure on forest resources (Chapter 3; Gougeoun & Leckie, 2003). A variety of spaceborne and airborne sensors offer high spatial resolution (<4 m), multispectral digital imagery in which ITCs are readily delineated by manual or automated methods. Some progress has been made toward automated, digital classification of ITC species in relatively species-poor temperate and boreal forests (McGraw et al., 1998; Gougeon & Leckie, 2003). However, my analyses in species-rich tropical rain forest (TRF) indicate that multispectral data are insufficient for discriminating even a limited number of tree species (Chapter 3). Spectral variability among species is greatly limited due to dominant biochemical controls on photon absorption by chlorophyll and water in tissues (Price, 1992; Cochrane, 2000). Subtle species differences in tropical tree spectra do exist at leaf to crown scales, but harnessing this information requires the higher spectral resolution offered by imaging spectrometers, or hyperspectral sensors (Cochrane, 2000). The spectral properties of materials within the sensor’s instantaneous field of view (IFOV) mix to create the spectrum within an image pixel. In tree canopies, the 186 dominant materials of concern are photosynthetic leaf and non-photosynthetic bark tissues, which have strongly contrasting spectral properties (Asner, 1998; Roberts et al., 2004; Williams, 1991). Variation in pixel spectra among and within species is largely related to differences in relative proportions of leaf and bark tissues within the sensor IFOV, as controlled by crown architecture and phenology. Leaf and bark spectra may also mix with the spectral characteristics of soil, understory vegetation, neighboring trees, strangler trees, lianas, epiphytes, lichen, or moss. The extent to which these factors can be considered noise or signal is related to species specificity. For example, lichens may only affect trees with a certain bark texture, and so lichens may add a unique spectral component to a species or functional type. Complicating matters, individual trees within a population may exhibit a site-specific spectral response to nutrient levels, water balance, light conditions, and herbivory. These factors increase conspecific (within-species) variability and make automated TRF tree species discrimination more challenging. One promising technique for the analysis of mixed pixels is Spectral Mixture Analysis (SMA), which models image spectra as a linear combination of two or more dominant or “pure” spectral components, or endmembers (reviewed in Keshava & Mustard, 2002). Endmembers are selected from a library of field, laboratory, image-derived or “virtual” spectra. A shade endmember is generally used to account for spectral illumination variation. Other bright endmember(s) represent major spectral components, such as green photosynthetic vegetation (GV: leaves, photosynthetic bark), non-photosynthetic vegetation (NPV: litter, bark, branches) and soil (Roberts et al., 1993). 187 Mixed reflectance spectra (ρ’λ) are modeled as the sum of the fractional abundance of each endmember according to the following formula: n ρ λ = ∑ f i ∗ ρ iλ + ε λ , (1) i =1 where n is the number of endmembers in the mixing model, fi is the fractional abundance of each endmember, ρiλ is the reflectance spectrum of each endmember, and ελ is a residual error spectrum. SMA output is the fractional abundance of each endmember, and all fractions are generally constrained so that they sum to 100%. The root mean squared error (RMSE) of the model fit can by calculated according to the formula: b RMSE = (ε λ ) ∑ λ 2 =1 (2) b where b is the number of bands. The model solution is strengthened with hyperspectral data over multispectral data because there are more high-signal bands than model endmembers. A single linear-mixture model with two to three endmembers will not adequately model every pixel in a spectrally-complex image (Dennison & Roberts, 2003a; Roberts, Gardner et al., 1998), such as from a TRF canopy. Multiple Endmember Spectral Mixture Analysis (MESMA) tackles this problem by automatically selecting endmembers on a per-pixel basis from a library of spectrally-diverse endmembers (Dennison & Roberts, 2003a; Roberts, Gardner et al., 1998). 188 MESMA is also a promising classifier. In this mode, the class of the spectrum being unmixed is determined based on the class label of the optimally-assigned, nonshade endmember in the SMA modeling process (e.g., bright endmember from a particular species). Although there have been no tropical applications involving MESMA classification, it has been used to map dominant chaparral vegetation types in southern California, USA (Dennison & Roberts, 2003a; Roberts, Gardner et al., 1998; Roberts, Dennison et al., 2003), soils (Okin et al., 2001), and snow cover and grain size (Painter et al., 1998). Best results (88.6% overall accuracy) were achieved when using a regionally-specific image endmember library and when assessing accuracy at the scale of a vegetation patch rather than at the image pixel scale (Dennison & Roberts, 2003a). Conducting MESMA with a large set of candidate endmembers can be computationally intensive when applied to a high spatial and spectral resolution image because multiple models must be evaluated on a per-pixel basis. Methods are thus necessary to reduce the endmember library to contain only those spectra that are pertinent to species discrimination. In particular, the spectral library should include those endmembers that specialize in modeling conspecific spectral variability, while excluding more generalist endmember spectra that model a broad range of pixels from other species or materials within the scene. One approach to MESMA endmember selection includes using expert knowledge of spectral mixtures within a scene to guide the collection of a limited number of pure field or image spectra that produce known fractional abundance (Okin et al., 2001; Painter et al., 1998). Bateson et al. (2000) developed a spectrally-diverse library of image endmembers by 189 including bundles of spectra arrayed in the convex hull of spectral space. Asner and Heidebrecht (2002) conducted SMA in an arid landscape using endmember bundles simulated with Monte Carlo sampling of field spectra. Roberts, Gardner et al. (1998) explored an alternative method where a large spectral library of reference endmembers was searched using a maximal coverage procedure to find endmembers that both mapped a large spatial area in the image while had minimal model overlap. Finally, Dennison and Roberts (2003a) developed an automated technique that selected endmembers that best modeled other spectra in an image-derived spectral library. The single candidate endmember that produced the minimum endmember average RMSE (EAR) for models within its class was selected to represent the class in MESMA classification. This technique was later expanded to include two optimal endmembers per class for multi-temporal characterization of chaparral vegetation (Dennison & Roberts, 2003b). Despite its ability to incorporate within-class variability, researchers have noted that MESMA, as a classifier, suffers from many of the same problems that plague other classifiers—high conspecific variability, lack of sharp interspecific (among species) spectral contrast, mixing with background material, and sensor noise (Okin et al., 2001; Roberts, Gardner et al., 1998). Furthermore, it is assumed in SMA that the non-linear mixing of photons among spectral components is minimal. Nonlinear mixing does occur in vegetation mainly because leaves permit light transmission and photon multiple-scattering, especially in NIR wavelengths (Roberts et al., 1993). Non-linear mixing will tend to increase fraction estimation error and 190 possibly decrease MESMA classification accuracy by selecting the wrong endmember as optimal in a linear mixing solution. The objective of this chapter was to assess MESMA for the species-level classification of ITCs in a tropical rain forest. Starting with a large spectral library of image and laboratory spectra, I developed a new automated approach to select optimal endmembers for two- and three-endmember MESMA models. In particular, the method sought to find a parsimonious set of endmember spectra that could simultaneously model the greatest number of conspecific spectra with little overlap, while modeling the fewest spectra from other species (extraspecific). 5.2. Methods 5.2.1. Canopy-emergent trees My analyses focused on the classification of canopy emergent individuals of seven tree species (Table 4.1). Chapter 3 provides details about the species and number of study trees and how their ITC polygons were digitized on the HYDICE hyperspectral imagery. As explained in Chapter 3, some overstory tree species are completely deciduous, generally beginning in the first dry season, while others are evergreen and continuously flush small amounts of leaves throughout the year (Table 4.1). Hyperspectral imagery was acquired on March 30, 1998 (Chapter 1), at the end of the first dry season, and all study trees were expected to have high mature leaf cover except DIPA and LEAM (summarized in Table 4.1). However, BAEL and HYME had relatively fine compound leaves and so their mature leaf cover was expected to have a lower LAI relative to the broadleaf species CEPE, HYAL and 191 TEOB. I thus expected LAI for the study species to be low for DIPA and LEAM, moderate for BAEL and HYME, and high for CEPE, HYAL and TEOB. 5.2.2. Laboratory bark spectra Pure NPV endmembers (i.e., bark) were difficult to locate in the hyperspectral image because these components rarely occupied a whole image pixel, and so laboratory-measured endmembers were used in 2EM+Shade models (see Section 5.2.4). Bark specimens from the 7 study species were sampled in the station vicinity and their reflectance was measured in a laboratory dark room with an ASD FieldSpec spectrometer (Analytical Spectral Devices, Boulder, CO, USA), which has 1-nm spectral sampling covering 350 to 2500 nm (detailed methods in Chapter 4). These ASD bark spectra were convolved to HYDICE band center positions (161 bands) using full-width, half-maximum information for each HYDICE band. The final laboratory library contained 66 bark spectra. DIPA had considerable withinspecies variation due to some specimens having green bark (Chapter 4; Fig. 5.1). 5.2.3. Scales of analysis As in Chapters 3 and 4, two ITC species classification schemes were analyzed: crown-scale and pixel-majority. The crown-scale approach labeled ITC species using classified crown-scale spectra. Crown-scale reflectance spectra were calculated as the mean of all within-crown pixels, providing one spectrum per crown, and MESMA was then applied directly to crown-scale spectra. The pixelmajority classification scheme applied the MESMA classifier to pixel-scale 192 reflectance spectra, and the ITC species label was assigned to the class representing the majority of classified pixels within the each crown. 5.2.4. Multiple endmember spectral mixture analysis (MESMA) classifier The MESMA classifier was evaluated using 1) multiple two-endmember models composed of an image spectrum and photometric shade (1EM+Shade), and 2) with multiple three-endmember models composed of a GV image spectrum, a NPV (bark) laboratory spectrum, and photometric shade (2EM+Shade). Photometric shade is a spectrum of zeros. Prior to MESMA, endmembers were selected for each species from a spectral library. The goal of the selection process was to find a set of endmembers from the library that modeled the maximum number of conspecific spectra while modeling the minimal number of extraspecific spectra. For 1EM+Shade MESMA, the spectral library included 300 pixel spectra per species from study crowns in the image (7 species x 300 pixels = 2100 total pixels). Pixels were drawn randomly from every crown without regard to their brightness, but sampling from any given crown was terminated if it had 40 or fewer available pixels. The sample size of 300 pixels per species was arbitrary yet ensured that crowns had a large number of independent test pixels after samples were removed from the classifier. Endmember selection proceeded as follows: every pixel spectrum was used to unmix every other pixel spectrum in the spectral library. Each model was considered valid according to the following constraints. The sum of SMA fractions was constrained to sum to 100%; however, valid models could have non-shade 193 endmember fractions between -6% and 106%. This fraction constraint was empirically determined by Halligan (2002) to allow for model error while still providing acceptable MESMA classification accuracy. Highly shaded pixels can have low signal-to-noise and can produce unreliable models; and thus, I imposed a constraint that the modeled shade fraction be less than 80% (Roberts, Gardner et al., 1998). Model fit was also considered unacceptable if overall error (RMSE) was greater than 2.5% (Roberts, Gardner et al., 1998). Some spectra could have spurious features in model residual spectra, such as spikes due to poor sensor calibration or extreme illumination-view geometry, yet still yield acceptable RMSE. To distinguish between spectra with these erroneous features from residuals resulting from the presence or absence of an absorption feature in the candidate endmember, I imposed a constraint that no seven contiguous bands have reflectance above 2.5% (Roberts, Gardner et al., 1998). Since HYDICE band spacing was variable across the full spectral range, this 7-band threshold applied to a spectral region between 23nm to 90-nm wide. Any given candidate spectrum could be modeled by multiple spectra from its class (e.g., conspecific in the case of species) as well as with spectra from other classes (e.g., extraspecific). For each species, I sought endmembers that had the highest percentage of successful conspecific models and the lowest percentage of successful extraspecific models. This was accomplished by finding the candidate endmember within each class that maximized the COunt-Based Index (COBI) according to Equations 3 through 5: 194 n COBI con = COBI extra ∑ Mod i =1 n i Mod = 0: not modeled Mod = 1: modeled (3) p t ⎛ o ⎜ ∑ Mod j ∑ Mod k Mod l ∑ ⎜ j =1 k =1 l =1 + + + ... ⎜ o p t ⎜ =⎝ c −1 ⎞ ⎟ ⎟ ⎟ ⎟ ⎠ (4) COBI = COBI con − COBI extra (5) where c is the number of extraspecific classes excluding the candidate species (i.e., c = 6) and Mod is a SMA model of the candidate endmember unmixing other n conspecific (i.e., n = 299 excluding itself) and o, p, … t spectra per extraspecific class. In this chapter, there were 6 extraspecific classes, each with 300 spectra, and so o, p, q, r, s, and t were equal to 300. The candidate endmember was either successful in modeling the spectrum (Mod = 1), or not (Mod = 0). The endmember selection scheme is depicted in Figure 5.2 for three hypothetical species (A, B & C), each with 4 candidate endmember spectra. These candidate spectra (in columns) and photometric shade were used in two-endmember SMA to model spectra in rows. 195 Successful SMA models are indicated as black (conspecific) or gray (extraspecific) boxes, while unsuccessful models are white boxes. The COBI was calculated for each candidate endmember column (Eq. 3-5), and the best species endmember had the highest COBI. Ties among candidate endmembers with the same COBI were decided by selecting the endmember with the lowest endmember average RMSE (EAR; Dennison & Roberts, 2003a), calculated from all n conspecific models within the endmember’s column. In this example, Species A‘s candidate endmember #3 was selected because it had the highest COBI within its class (Fig. 5.2). Species B had endmembers with the least amount of extraspecific spectral similarity (positive COBI indices) while Species C had the most extraspecific spectral overlap (zero to negative COBI indices). Once an optimal endmember was chosen for its class, the spectrum and all spectra successfully-modeled were removed from the analysis and then COBIs were re-calculated for the remaining spectra. The new COBIs were then used to select another endmember for each class. By removing spectra modeled by the first selected endmember, the second-iteration endmember had reduced spectral similarity with the first-iteration endmember. This iterative-selection process was repeated for each class until COBI was zero or less, indicating that remaining candidate endmembers modeled a greater percentage of extraspecific spectra than conspecific spectra. Final endmembers were not necessarily pure GV spectra, but rather bright spectra that could model other unique mixtures within each species. For example, an endmember from a deciduous leaf-off crown could have been a bright NPV-dominated spectrum. 196 A similar COBI approach was used for selecting jointly-optimal endmembers for 2EM+Shade MESMA models composed of shade, GV and NPV. Bark spectral endmembers for each species were selected from the laboratory library of 66 spectra (Section 2.2.2). Even though this library included some green and presumably photosynthetic bark, I considered it a “pure” NPV spectral library. Candidate image endmembers representing GV were selected by first calculating the NIR minus red contrast in all 2100 pixels in the image spectral library and then selecting those spectra that were in the top 90th percentile for each class. This reduced library included 217 image spectra (31 pixels per species). Every combination of GV (image) and NPV (laboratory) spectra were matched within each species, forming 2046 2EM+Shade candidate models, which were then used to unmix the original library of 2100 image spectra. The COBI endmember selection scheme, as described for 1EM+Shade models, was then implemented for simultaneous selection of GV and NPV endmember pairs for each species. Pixel-majority MESMA classification of ITCs proceeded by modeling pixels within all crowns using 1EM+Shade or 2EM+Shade models. Pixels included in the image spectral library were excluded from the classification analyses. For 2EM+Shade models, only combinations of GV and NPV endmembers from the same species were considered (e.g., a GV image endmember from BAEL and an NPV laboratory endmember from BAEL). The best model for each pixel was chosen using the same constraints used in COBI endmember selection: non-shade fractions were between -6% and 106%, the shade fraction was less than 80%, no 7 contiguous model residuals were above 2.5% reflectance, and the model RMSE was less than 197 2.5%. If multiple models met these constraints for a given pixel, then the model with the lowest RMSE was selected. A species label was assigned to each pixel using the class of the best-fit model for the pixel. Pixels with no acceptable models were labeled as unclassified. For the pixel-majority ITC classification, the species label was assigned based on the majority class of classified pixels within each crown. Image and laboratory endmembers selected for pixel-scale MESMA were used in crown-scale MESMA classification. In this case, 1EM+Shade and 2EM+Shade models were used to model crown-scale spectra. 5.3. Results 5.3.1. Endmember selection A matrix of 2100 candidate endmembers (columns) modeling the same spectra (rows) using 1EM+Shade SMA is shown in Figure 5.3. Models meeting the constraints (Section 2.2.4) were colored black, and white otherwise. Spectra were sorted primarily by species and secondarily by spectral brightness based on their 800-nm reflectance, with spectra ranked from dark (left or up) to bright (right or down). As expected, the brighter candidate spectra within a species successfully modeled more spectra than darker spectra, producing within-species triangular patterns in the modeled/un-modeled matrix (Fig. 5.3). Part of this trend is due to the relatively liberal -6 to 106% fraction constraints on successful models. I expected that candidate endmembers from leaf-off deciduous species (DIPA and LEAM) would model spectra between those two species but not among leaf-on species because leaf-off crowns had distinct spectral properties such as lower NIR 198 reflectance and shortwave infrared absorption features from exposed NPV (Chapter 4). Contrary to what was expected, many deciduous candidate endmembers modeled spectra from across all species, regardless of phenology (Fig. 5.3). Also, most dark spectra of HYAL and HYME were successfully modeled by candidate endmembers from other species, suggesting potential confusion of HYAL and HYME by other species endmembers during actual MESMA classification (Fig. 5.3). However, Figure 5.3 only depicts spectra that meet model constraints (black dots), and does not take into account the RMSE selection criteria used for final model selection in MESMA. The first COBI-selected endmember per species modeled 41% (BAEL) to 81% (TEOB) of conspecific image spectra in the library (Table 5.1). A second and third COBI endmember modeled an additional 13% (LEAM) to 39% (BAEL) of conspecific spectra. The fourth and greater endmembers per species modeled a low percentage of conspecific spectra. Although training pixels came from every crown, the final set of endmembers for each species came from five (TEOB) to nine (DIPA) different crowns (Table 5.1, Crown ID). A spectrum’s 800-nm (near infrared) reflectance was used to gauge spectral brightness (Table 5.1). No deciduous leaf-off endmembers had 800-nm reflectance greater than 58.0%, while the leaf-on species endmembers covered a wider range of brightness (e.g., up to 86.6% for HYAL). A near-infrared reflectance of 86.6% is unusually bright for a plant image spectrum. The presence of extremely bright spectra in the image resulted mainly from poor radiometric calibration of the hyperspectral data and solar geometry that made the eastern sides of the crowns 199 relatively bright (HYDICE scene was acquired early in the morning at 7:55-8:27 am local time, 56.3° to 48.4° solar zenith, and 92° to 94° solar azimuth). Extremely bright endmembers are expected to properly model other pixels within the crown if their spectral mixtures vary only by brightness and not shape. Leaf-off DIPA image spectra in the library showed clear separation from a leafon species, TEOB (Fig. 5.4). Relative to DIPA, TEOB had more 680 nm and 1650 nm absorption (lower reflectance) due to chlorophyll and water, respectively. The scatter plots also depict the mixing lines connecting three COBI-selected endmembers per class (Fig. 5.4). These endmembers were not necessarily the brightest pixels in the library. Also, some DIPA spectra strayed into the TEOB spectral space determined by 680 and 800 nm bands, indicating the presence of green tissues in their spectral mixtures. Endmembers 3, 5, 8, and 9 for DIPA formed mixing lines in the TEOB 680-800 nm space (Fig. 5.4, EM3 is shown), and it was expected that these endmembers would be misclassified as TEOB or other leaf-on species in a MESMA classification. 5.3.2. Pixel-majority ITC classification When MESMA was applied to pixels with 1EM+Shade models, overall classification accuracy was 36.3% with one optimal endmember per species, and accuracy increased 4.4% with the inclusion of 5 to 9 COBI-selected endmembers per species class (Table 5.2). However, including multiple endmembers per class did not necessarily improve class Producer’s accuracies. For example, the best CEPE Producer’s accuracy included 5-9 endmembers per species, while the best LEAM 200 Producer’s accuracy included just one endmember per species. When all 2100 image spectra in the library (300 spectra per species) were included in 1EM+Shade MESMA, overall accuracy and most Producer’s accuracies were higher than when using COBI-selected endmembers. The best pixel-majority ITC classification using COBI-selected endmembers had 59.8% overall accuracy, with 1EM+Shade models with 3 endmembers per species (Table 5.3). With the 2EM+Shade endmember selection, COBI values were negative after selecting one pair of GV-NPV endmembers for HYAL, HYME and TEOB, while DIPA had up to 4 GV-NPV endmember pairs. The 1EM+Shade classification with 3 endmembers per species was 6.5% more accurate than the equivalent 2EM+Shade classification with 1-3 endmembers (not significant, Z=1.86, α=0.05;Congalton, 1991). Example ITCs for each species are shown for 1EM+Shade MESMA with 3 endmembers per species (Fig. 5.5). All ITCs in this example were correctly-labeled. Some ITCs had spectra with similar shapes but differed in brightness, and so a single endmember modeled most of the crown. For example, 96% of the pixels in the CEPE crown in Figure 5.5 were modeled by its third optimal endmember (EM3), which was relatively bright (Table 5.1). In contrast, the DIPA crown had 34% of its pixels modeled by EM1 and 19% of its pixels modeled by EM3 (Fig. 5.5). These two DIPA endmembers had NPV and GV spectral properties, respectively (Fig. 5.4). Only 48% of the HYME crown’s pixels were correctly classified, which may be a result of broad spectral overlap with endmembers from other species (Fig. 5.3, discussed in Section 5.3.1). However, the pixel-majority vote was HYME because 201 no other species had a greater number of pixels classified within the crown. The error matrix for this classification (Table 5.4) revealed misclassification of deciduous DIPA (mostly leaf-off) individuals with BAEL individuals, a leguminous species with fine compound leaves (Chapter 3). Some BAEL individuals were also mapped as HYME, another leguminous, composite-leaved species. Crowns of TEOB were confused with HYAL, which were both evergreen species with broad leaves. When all 2100 image spectra in the library (300 spectra per species) were included in 1EM+Shade models, pixel-majority MESMA accuracy increased dramatically to 90.2% (Tables 5.3 and 5.5). Much of the confusion in the classification was with DIPA crowns classified as BAEL, as seen with MESMA and 3 endmembers per species. Overall accuracy did not significantly improve (Z=0.43) with 2EM+Shade models using 2100 image spectra matched by species with 66 laboratory bark spectra (19,800 unique models; Table 5.3, pixel-majority). 5.3.3. Crown-scale ITC classification In general, overall accuracies at the crown-scale were lower than with the pixelmajority technique (Table 5.3). The best COBI-based MESMA accuracy was only 54.7% with 1EM+Shade models composed of three image endmembers per species. When including all 2100 image spectra in 1EM+Shade models, overall accuracy reached 67.3%. 202 5.3.4. Model constraints Roberts, Gardner et al. (1998) implemented MESMA with model fractions constrained from -1% to 101%. The reference spectral library used in that study had high radiometric quality and tended to be bright relative to image spectra, which include shadowing in the IFOV. Working in a temperate forest landscape, Halligan (2002) found that MESMA classification accuracy improved when fraction constraints were relaxed to accommodate extremely bright or dark endmembers in an image spectral library. Concurring with those results, I also found that fraction constraints of -1% to 101% on COBI and MESMA produced weaker classification accuracies than those presented for -6% to 106% fraction constraints (data not shown). 5.4. Discussion 5.4.1. Count-based index (COBI) selection of image endmembers The primary goal of the COBI endmember selection scheme was to find the set of spectra that model the most within-species spectra with minimal ability to model spectra from other species. In other words, I sought endmembers from the spectral library that were conspecific ”specialists” as opposed to interspecific “generalists” (Roberts, Gardner et al., 1998). It was apparent from Figure 5.3 that this objective would be difficult to achieve for this forest type since most bright candidate endmembers were generalists, regardless of species. This is because vegetation spectra had similar spectral shapes due to dominant biochemical controls on photon absorption within tissues (Price, 1992). 203 Despite this broad spectral overlap among spectra, the COBI technique was successful in selecting at least one specialist endmember for each species. The firstiteration endmembers modeled a large proportion of conspecific spectra in the library (41% to 81%). Species-level differences in pixel- and crown-scale spectra are related to crown structure and phenology, which controls the spectral mixing of leaves, bark and shadows within the sensor IFOV (Asner, 1998; Roberts, Ustin et al., 2004). Specialist endmembers should thus respond to unique aspects of crown structure and phenology for each species. 5.4.2. MESMA classification With one COBI endmember per species, MESMA was able to correctly classify a third of ITC pixels. About half of ITCs were correctly classified with the pixelmajority and crown-scale classifiers (1EM+Shade models). These results indicate that there are unique spectral properties of species at both pixel and crown scales that can be discriminated by MESMA with one image endmember per species. Despite this encouraging finding, the overall accuracy of the final ITC classification was still too low for operational use. Including up to three COBIselected endmembers in 1EM+Shade MESMA increased pixel-scale accuracy, translating into improved pixel-majority accuracy. Evidently, the 2 to 3 endmembers identify other unique spectra for species at pixel and crown scales. For example, the first three DIPA endmembers included a spectrum with characteristics of NPV (EM2), one spectrum similar to a leaf-on TEOB spectrum (i.e., high GV, EM3), and a spectrum in between these two spectra (EM1). Additional endmembers 204 (4+) tended to decrease overall accuracy, likely a result of additional endmembers having spectral overlap with extraspecific spectra, which leads to increased class confusion in MESMA. MESMA with 1EM+Shade models generally had higher overall accuracies than with 2EM+Shade models. Relative to 1EM+Shade models, including one laboratory and image spectrum per species in 2EM+Shade models decreased overall accuracy by 2% with pixel-majority while 11% with crown-scale analysis. I concluded that including laboratory NPV spectra in models does not help discriminate species. One explanation is that laboratory NPV spectra may differ substantially from image NPV spectra due to differences in measurement scale, scattering environments, atmospheric contamination, and radiometric calibration. In this situation, laboratory NPV spectra would be too dissimilar from image NPV to be effective in the mixture models with image GV endmembers. Unfortunately, pure NPV endmembers were difficult to acquire for leaf-on species, and even NPV spectra from leaf-off species had distinct chlorophyll signals (i.e., red absorption well) from internal photon scattering and mixing with epiphytes, lianas, mosses or lichen on branches. In contrast, image endmembers included the whole range of GV-dominated to NPVdominated spectral mixtures and incorporated sensor calibration artifacts and nonlinear mixing from photon scattering that is absent in laboratory spectra. My results indicate that, although they may not be “pure”, image endmembers provide a considerable level of species discrimination at pixel and crown scales. When mixed pixels are modeled with slightly-mixed GV and NPV endmembers, there may be a variety of models that have adequate fits. In this case, the fractional abundance of 205 NPV relative to GV in a crown may help distinguish an ITC species, rather than the endmembers’ species label. Regardless of MESMA model complexity, the overall accuracies with COBI endmembers were lower than the 86% (Chapter 3) and 68% (Chapter 4) overall accuracies with pixel-majority linear discriminant analysis (LDA) and decision tree (DT) classifiers, respectively. MESMA applied to crown-scale spectra had even lower accuracies relative to those using LDA and DT classifiers (Chapters 3 & 4). I was able to boost MESMA accuracy to a relatively high level (90%) only by including the full library of image spectra as potential 1EM+Shade models. However, one caveat with this result is that endmember-selection pixels and testing pixels came from the same crowns. Although the endmember-selection pixels were excluded from ITC classification analyses, there were still spatial autocorrelation effects—pixels from the same crown have similar spectral properties. It is likely that the accuracies reported for the full-library MESMA are optimistic due to a lack of complete independence between library and testing data. This bias was negligible when considering only nine or fewer endmembers per species because ITCs contained 41 to 662 pixels, no crown had fewer than 40 testing pixels, and no more than 2 endmembers came from the same crown (Table 5.1). In Chapters 3 and 4, DT and LDA analysis were performed with a cross-validation approach and so classifiers did not have autocorrelated spectra in training sets. 206 5.4.3. Recommendations for future research Despite the potential bias when using the full spectral library to classify ITCs, it appears that many more than five endmembers per species are necessary to adequately use MESMA as an ITC classifier in this TRF environment. These results suggest that future research using MESMA for species classification should focus on methods to increase conspecific endmember variability while minimizing extraspecific endmember similarity. However, increasing the number of mixture models does have a computational cost. For example, in the per-pixel analysis of 2100 1EM+Shade models using IDL code (RSI, Inc., Boulder, CO, USA) running on a 3.2 GHz Intel Pentium 4 processor, 35,043 ITC pixels were processed in 3 hrs, 14 min (181 pixels/min). The 19,800 2EM+Shade models took 29 hrs and 45 min. to process (20 pixels/min). To process the entire HYDICE mosaic of 3,774,300 pixels would have taken 14.5 and 133.5 days for evaluating all 1EM+Shade and 2EM+Shade models, respectively. In an application over a large spatial extent, segmenting the hyperspectral image into ITCs of interest (e.g., Gougeon & Leckie, 2003) would greatly improve processing time by excluding canopy gaps or nontarget vegetation from the modeling process. As with all remote sensing applications, limitations on processing time are also expected to decrease as computer processing speed increases per unit cost according to Moore’s Law (Moore, 1998). I have identified two improvements for the COBI selection technique that may be useful in future studies. For one, the extraspecific COBI formula (Equation 4) was implemented with an equal class weighting scheme. Weights could be 207 multiplied by each summation term in Equation 4 to influence the selection of certain types of endmembers. For example, if more NPV-dominated endmembers were desired for leaf-off species, then high weights could be applied to leaf-on species in Equation 4, which would select against GV-dominated endmembers by lowering their COBI values (Eq. 5). Another COBI improvement involves how model strength is evaluated. Just because a candidate endmember successfully models extraspecific spectra in the library, the endmember may still have stronger model fits with its conspecific spectra. The COBI formula could be modified to favor candidates with stronger conspecific over extraspecific fits, possibly by applying a weighting scheme to RMSE, residuals and fractions from model outputs. 5.5. Conclusions A major goal of this chapter was to assess airborne hyperspectral imagery for the species-level classification of ITCs in a tropical rain forest. The MESMA classifier explored in this chapter was expected to have optimal performance with hyperspectral over multispectral imagery of equal spatial resolution. I found that MESMA has potential as a classifier, providing 90% overall accuracy, but only when presented with a library of 2100 model endmembers (300 endmembers per species). Processing this quantity of models on a per-pixel basis greatly increases processing time for large study extents. The requirement for 300 endmembers per tree species suggests that their crowns have many unique spectral signatures, probably due to spectral mixing. For 208 MESMA to be a more useful classifier, the number of candidate endmembers will need to be constrained to those that encompass the unique spectral shapes that make species distinct. The endmember selection technique presented in this chapter provides one method for finding the optimal set of endmembers. For example, MESMA with one optimal endmember from each species discriminated ITC species with 49% and 50% overall accuracy, depending on pixel-majority or crown-scale evaluation, respectively. Although my count-based MESMA classification accuracy was not adequate for species-level mapping, the technique may yield more physically-accurate fractions over standard SMA with single GV and NPV endmembers. Such improvements could greatly benefit applications involving land-use classification, estimation of deciduous and evergreen components of forest canopy, or linking fractions to biophysical parameters, such as aboveground biomass and percent cover. In this context, the proper matching of endmembers to their respective image pixels would yield optimal fraction estimates, a process that may benefit from another type of classifier. I found LDA to be a stronger classifier for my study species (Chapter 3). One approach to estimating endmember fractions would be to apply the LDA classifier to pixels, perform MESMA using only endmembers for each pixel’s class, and then select the best fractions based on model parameters (e.g., lowest RMSE). 209 Table 5.1. Summary statistics of optimal endmembers selected by the countbased index (COBI) and 2100 training image spectra (Section 5.2.4). Each row is a selected candidate endmember (EM). Columns are the species code, selection rank, crown identifier, reflectance at 800 nm, count of valid conspecific models, percent of all conspecific models, COBI * 100 (Eq. 3), and average RMSE*100 (EAR). Species EM # Crown ID 800 nm Count Percent COBI EAR BAEL 1 106 66.0 124 41.3 25 20 BAEL 2 177 55.4 47 15.7 13 23 BAEL 3 166 64.1 70 23.3 18 21 BAEL 4 61 60.3 29 9.7 40 27 BAEL 5 35 63.5 12 4.0 30 23 BAEL 6 149 24.8 5 1.7 22 56 BAEL 7 126 73.6 6 2.0 31 30 BAEL 8 106 63.5 2 0.7 27 39 CEPE 1 116 50.5 155 51.7 31 17 CEPE 2 93 64.8 61 20.3 16 23 CEPE 3 204 74.5 22 7.3 26 35 CEPE 4 10 35.9 20 6.7 16 35 CEPE 5 173 45.6 19 6.3 21 22 CEPE 6 113 73.9 7 2.3 28 25 CEPE 7 10 34.6 2 0.7 10 57 CEPE 8 110 28.8 2 0.7 11 26 CEPE 9 116 44.4 3 1.0 12 26 DIPA 1 136 40.5 178 59.3 37 16 DIPA 2 107 33.7 38 12.7 26 29 DIPA 3 103 41.5 28 9.3 18 23 DIPA 4 154 44.8 11 3.7 12 30 DIPA 5 5 49.2 7 2.3 11 28 DIPA 6 97 40.7 6 2.0 13 46 DIPA 7 39 43.9 5 1.7 9 29 DIPA 8 14 54.3 10 3.3 8 27 DIPA 9 62 48.8 3 1.0 13 30 210 Table 5.1. (continued). HYME HYME HYME HYME HYME HYME HYME HYME HYAL HYAL HYAL HYAL HYAL HYAL LEAM LEAM LEAM LEAM LEAM LEAM LEAM LEAM LEAM TEOB TEOB TEOB TEOB TEOB 1 2 3 4 5 6 7 8 1 2 3 4 5 6 1 2 3 4 5 6 7 8 9 1 2 3 4 5 45 128 147 171 185 186 121 212 202 145 187 200 211 19 184 168 153 179 161 179 168 142 135 89 79 87 92 85 79.0 45.7 70.3 36.3 48.0 58.4 67.0 64.7 69.3 72.4 80.0 86.6 43.0 77.5 46.2 11.4 58.1 34.2 44.3 57.8 12.9 45.3 43.1 65.4 70.0 27.7 78.3 61.6 197 51 17 16 6 4 2 2 226 39 8 13 7 2 208 13 26 18 14 4 3 7 2 242 34 13 5 2 211 65.7 17.0 5.7 5.3 2.0 1.3 0.7 0.7 75.3 13.0 2.7 4.3 2.3 0.7 69.3 4.3 8.7 6.0 4.7 1.3 1.0 2.3 0.7 80.7 11.3 4.3 1.7 0.7 48 25 24 25 26 19 20 14 45 22 19 29 37 27 48 14 18 26 18 18 17 17 25 41 37 45 42 32 14 15 20 21 30 25 25 31 13 16 25 17 15 26 14 71 20 28 24 28 63 26 26 12 14 19 15 18 Table 5.2. Pixel-scale percent producer’s accuracy and overall accuracy using Multiple Endmember Spectral Mixture Analysis (MESMA) and models of 1 to 9 image endmembers per species and photometric shade (1EM+Shade). Numbers are percentages except n, the total of non-training pixels per class (total n = 35043). EM / specie Overall s BAEL CEPE DIPA HYAL HYME LEAM TEOB Accuracy 3752 2691 16115 4859 2317 2561 2748 n 1 25.7 33.4 34.3 52.4 44.8 20.3 51.0 36.3 2 32.4 47.6 41.8 34.5 48.8 20.2 39.5 39.9 3 35.6 52.8 49.5 30.6 50.8 25.2 38.2 44.5 4 28.8 52.7 45.4 31.5 52.4 21.8 39.1 42.0 5-9 30.1 65.7 42.3 30.6 48.5 25.4 30.7 40.7 300 55.6 60.6 48.2 59.0 53.7 46.3 74.5 53.7 212 Table 5.3. Overall ITC classification accuracy using Multiple Endmember Spectral Mixture Analysis (MESMA). 1EM+Shade models included photometric shade and 1 image endmember. 2EM+Shade models included photometric shade, 1 image endmember and 1 laboratory endmember (see Section 5.2.4). No. No. 2EM+Shade EMs per Models 1EM+Shade EMs per Model Models species Models species s EM1 EM1 EM2 Pixel-majority 1 7 48.6 1 1 7 46.7 2 14 53.7 1-2 1-2 19 55.1 3 21 59.8 1-3 1-3 34 53.3 4 28 54.7 1-4 1-4 41 53.3 5-9 54 51.9 n/a n/a n/a n/a 300 2100 90.2 300 5-15 19800 91.6 Crown-scale 1 7 50.0 1 1 7 39.3 2 14 52.8 1-2 1-2 19 52.3 3 21 54.7 1-3 1-3 34 51.4 4 28 53.3 1-4 1-4 41 51.4 5-9 54 52.8 n/a n/a n/a n/a 300 2100 67.3 300 5-15 19800 76.6 213 Classification 214 Table 5.4. Error matrix of pixel-majority classification using 3 COBI-selected image endmembers per species in 1EM+Shade models (Kappa = 0.49). Field Reference Species BAEL CEPE DIPA HYAL HYME LEAM TEOB BAEL 16 12 1 4 1 CEPE 6 1 8 4 DIPA 5 1 60 2 10 1 HYAL 2 2 1 22 1 9 HYME 4 2 3 9 1 LEAM 2 1 6 TEOB 1 3 1 2 9 Total 29 10 81 34 14 21 25 Producer’s 55.2% 60.0% 74.1% 64.7% 64.3% 28.6% 36.0% 214 Total 34 19 79 37 19 9 16 214 User’s 47.1% 31.6% 76.0% 59.5% 47.4% 66.7% 56.3% 59.8% Classification 215 Table 5.5. Error matrix of pixel-majority classification using 300 image endmembers per species in 1EM+Shade models (Kappa = 0.88). Field Reference Species BAEL CEPE DIPA HYAL HYME LEAM TEOB Total User’s BAEL 28 9 1 38 73.7% CEPE 10 1 11 90.9% DIPA 67 1 68 98.5% HYAL 2 32 34 94.1% HYME 1 12 13 92.3% LEAM 2 19 21 90.5% TEOB 2 1 1 25 29 86.2% Total 29 10 81 34 14 21 25 214 Producer’s 96.6% 100.0% 82.7% 94.1% 85.7% 90.5% 100.0% 90.2% 215 . BAEL (N =15) Reflectance 80% 60% 60% 40% 40% 20% 20% 0% 0% 350 850 Reflectance 1350 1850 2350 DIPA (N = 10) 80% 350 850 60% 60% 40% 40% 20% 20% 1350 1850 2350 HYAL (N = 5) 80% 0% 0% 350 850 1350 1850 2350 HYME (N = 8) 80% Reflectance CEPE (N = 9) 80% 350 850 60% 40% 40% 20% 20% 0% 1850 2350 LEAM (N = 12) 80% 60% 1350 0% 350 850 1350 1850 2350 350 850 1350 1850 2350 Wavelength (nm) Reflectance 80% TEOB (N = 7) 60% 40% 20% 0% 350 850 1350 1850 2350 Wavelength (nm) Figure 5.1. Bark mean (bold line) and standard deviation (±1 S.D., thin line) of simulated HYDICE reflectance for each species. 216 Candidate Endmember Spectra Modeled Spectra Sp. A Sp. C Sp. B 4 3 2 1 4 3 2 1 4 3 2 1 Sp. C Sp. B Sp. A 1 2 3 4 1 2 3 4 1 2 3 4 COBI 0.0 -0.50 0.25 0.13 0.0 0.25 0.50 0.75 -1.0 -0.75-0.50-0.25 EXAMPLE: Species A, candidate endmember #3 COBIcon = 2/4 = 0.50 COBIextra = (1/4 + 1/4)/2 = 0.25 COBI = COBIcon – COBIextra = 0.50 – 0.25 = 0.25 Figure 5.2. Hypothetical depiction of COBI endmember selection. Four candidate endmember spectra (columns) from three species (A, B & C) are randomly selected from the image pixels. Each candidate endmember is then used with photometric shade to model every other spectrum (rows) in a spectral mixture analysis. Successful models (defined in Section 5.2.4) are black (conspecific) and gray (extraspecific) boxes with values of 1, while unsuccessful models are white boxes with values of 0. The COBI is calculated using successful models in columns (see formula in Section 5.2.4). The best species endmember has the highest COBI. 217 Candidate Endmember Spectra TEOB LEAM HYAL HYME DIPA CEPE BAEL Modeled Spectra BAEL CEPE DIPA HYME HYAL LEAM TEOB 2100 spectra Dark Bright 300 pixels Figure 5.3. Matrix of spectral mixture analysis counts used in count-based COBI endmember selection (Section 5.2.4). A total of 2100 randomly-selected pixels (300 per species) were used to model the same spectra. Valid models were marked as black points and blank otherwise. Within a species, spectra were sorted from dark (left) to bright (right) according to their 800-nm reflectance; and thus, brighter candidate endmembers successfully model more spectra. 218 90% TEOB EM1 Reflectance (800 nm) 80% DIPA EM3 70% DIPA EM1 60% 50% 40% 30% 20% 10% 0% 0% 2% 4% 6% 8% 10% 12% Reflectance (680 nm) 40% DIPA EM1 Reflectance (1650 nm) 35% 30% 25% 20% TEOB EM1 15% 10% 5% 0% 0% 10% 20% 30% 40% 50% 60% 70% 80% Reflectance (800 nm) Figure 5.4. A) Near-infrared vs. red scatter plot for deciduous Dipteryx (DIPA; closed circles) and evergreen Terminalia (TEOB; open circles) pixels from the spectral library. Three candidate endmembers selected by the COBI selection scheme are plotted as vectors (DIPA – black, TEOB – gray) connecting to photometric shade. B) A near-infrared vs. shortwave-infrared scatter plot for the same spectra and endmembers. 219 90% 800 nm Classified 800 nm BAEL HYME CEPE LEAM DIPA TEOB HYAL Classified EM other species EM1 EM2 EM3 Background or not modeled Figure 5.5. Correctly-labeled crowns for the seven study species using image endmember and shade (1EM+Shade) MESMA with 3 optimal endmembers per species. The 800-nm reflectance is shown for each crown. The endmembers selected by the MESMA classifier are shown for the classified images. Conspecific endmembers are gray-scale (1 through 3), while white pixels were modeled by extraspecific endmembers. The background and non-modeled pixels are black. 220 CHAPTER 6: Comparison of lidar and hyperspectral data for tree classification 6.1. Introduction Tree structure from leaf to crown scales can vary greatly among species, and these differences are a vital tool for visual interpretation of aerial photography. Through a complex visual and cognitive process, photo-interpreters discriminate individual tree crown (ITC) species using crown color and structural properties, such as branch architecture, canopy position, contour shape, size, foliage cover and texture (Fournier et al., 1995; Herwitz et al., 1998; Myers & Benson, 1981; Trichon, 2001). Visual interpretation of fine spatial scale image objects, such as trees, is a time consuming, costly and often inconsistent process when dealing with photographs spanning large areas, and so computer-based automated techniques for crown detection, delineation and species classification are needed (Gougeon & Leckie, 2003; Leckie, Gougeon, Hill et al., 2003). However, training a computer to discriminate tree species using remotely-sensed digital data is a challenging task. Optical sensors provide digital color information, and in the case of full-range hyperspectral sensors (i.e., imaging spectrometers, 400-2500 nm), the spectral information recorded by the sensor surpasses that available from human vision. With high spatial resolution imagery (< 5 m), spectral information alone may be adequate for automated tree species classification (Carleer & Wolff, 2004; Gougeon, 221 1995; Leckie et al., 2005; Wang et al., 2004; Xiao et al., 2004), especially if species have distinct phenology (i.e., leaf cover, flowering) at the time of image acquisition (Chapter 3) or through multiple image dates (Key et al., 2001). Digital optical imagery also encodes properties of crown structure. In essence, crown structure refers to the size and three-dimensional arrangement of its components (i.e., leaves, trunk and branches). Because these components have distinct absorptive, transmittive and reflective properties (e.g., chlorophyll and water content in leaves, lignin and cellulose in bark) and block light to create within-crown shadows, there is spectral variation across the crown (Kimes, 1983; Sandmeier et al., 1998). If pixel spatial resolution is finer than the scale of a crown, then the spatial arrangement of pixel spectra (e.g., texture) may respond to aspects of crown structure. For example, the shadows cast on one side of a crown due to illumination geometry is related to the crown’s overall structure, and shadows and exposed bark within the crown may create finer-scale spectral variation. Few studies have used pixel spatial information for automated ITC species discrimination. Meyer et al. (1996) reported that a standard deviation metric calculated from a near-infrared (NIR) band improved classification accuracy of conifers and hardwoods over that achieved with spectral data alone. At patch or stand scales, however, there is inconclusive evidence that image texture greatly improves forest composition classification (Franklin et al., 2000; Leckie, Gougeon, Walsworth et al., 2003, Wang et al., 2004; although see Franklin et al., 2001; Zhang, et al., 2004). Small-footprint lidar sensors record the three-dimensional height distribution of surface materials (Lefsky et al., 2002), and so these sensors are ideal for quantifying 222 ITC structure. Several studies have shown that ITC location, shape and area can be automatically described with algorithms applied to lidar-derived digital canopy models, or DCMs (Brandtberg et al., 2003; Holmgren & Persson, 2004; Leckie, Gougeon, Hill et al., 2003; Persson et al., 2002). Various crown structure metrics can be calculated from the height distribution of within-crown DCM cells or xyz points, as well as shape and area metrics from the delineated ITC polygons. It has been shown that lidar-based metrics are strongly correlated to field-measured ITC height and diameter (Brandtberg et al., 2003; Holmgren & Persson, 2004; Persson et al., 2002; Popescu et al., 2003; and Chapter 2). The properties of crown structure described by lidar metrics are also useful for tree species discrimination. Working with three hardwood species in leaf-off conditions, Brandtberg et al. (2003) calculated mean, standard deviation, skewness, and kurtosis from the histograms of lidar height (normalized to crown maximum height) and NIR reflectance from automatically-segmented ITCs. All metrics had highly significant differences among the study species due to species-level variation in vertical structure and branch distribution. The study achieved an overall accuracy of 60% when using all variables in a linear discriminant analysis (LDA) classification scheme. Holmgren and Persson (2004) classified two conifer species with 95% accuracy using a suite of lidar-derived crown shape and NIR reflectance intensity measurements. The proportion of first lidar returns and the standard deviation of intensity within the crown were the most important variables in the classification. Increased variance in laser-return intensity was thought to be caused by gaps within the crown. 223 Chapters 3 through Chapter 5 focused on spectral-based discrimination of ITC species with hyperspectral and multispectral data. The objective of this chapter is to extend these analyses to include lidar-derived, crown structure metrics calculated from the DCM (Chapter 1). Similar to methods in Chapters 3 and 4, I investigated decision trees and linear discriminant analysis for species classification. 6.2. Methods 6.2.1. Canopy-emergent trees Analyses focused on the classification of canopy emergent individuals of six target species and 18 non-target species (Table 6.1). Emergent trees with large, exposed crowns provided a large sample of pixels that were less influenced by spectral shadowing or scattering by neighboring trees and they were relatively easy to locate in the hyperspectral and lidar data. In my previous Chapters, the species Ceiba pentandra (CEPE) was included as a target species. For this Chapter, I excluded this species as a target species because there were only 6 individuals within both the hyperspectral and lidar data sets. As explained in Chapter 3, some overstory tree species are completely deciduous, generally beginning in the first dry season, while others are evergreen and continuously flush small amounts of leaves throughout the year. The hyperspectral imagery was acquired in March, at the end of the first dry season, and the target trees DIPA and LEAM were expected to be leaf-off (Table 6.1). The species HYAL and TEOB had evergreen crowns with broad leaves, while the species BAEL and HYME had fine, sparsely-distributed compound leaves that gave 224 them more intermediate leaf area. For the hyperspectral image, I thus expected leaf area index (LAI) for my target species to be low for DIPA and LEAM, moderate for BAEL and HYME, and high for HYAL and TEOB. Of the non-target species, I did not expect any leaf-off species at the time of the hyperspectral image acquisition (Table 6.1). The lidar data were acquired in mid-September, during the second and smaller dry season (Section 2.1.1). O’Brien (2001) found that of the HYME individuals studied, roughly 60% of the crowns had >75% mature leaf cover in September, 1997 (Table 6.1). With visual verification using videography acquired at the time of the lidar flight, in contrast, I found that all HYME individuals were leafoff (Fig. 6.2). Other leaf-off trees included INAL and PTOF (Table 6.1; Fig. 6.2INAL). Although BAEL individuals have leaves in September (O’Brien, 2001; verified in videography), their leaves are sparse and finely compound giving them a low-LAI crown structure (Fig. 6.2-BAEL; Chapter 3). In this chapter, I was particularly interested in mapping large Dipteryx (DIPA) trees (Fig. 6.2). The population of large individuals of this species is in decline across the Sarapiquí region due to deforestation. An important ecological function of large individuals of this species is to provide seeds and nesting cavities for the endangered Great green macaw (Ara ambigua) (pers. comm., Powell 2001). Remote sensing technology that can identify large Dipteryx crowns may contribute to macaw conservation efforts by providing a rapid and cost-effective means to map their habitat and migration corridors across the region. The 2-dimensional area of the tree crowns were manually digitized over the HYDICE imagery (see Chapter 3). As in previous studies, I refer to digitized crown 225 polygons as individual tree crowns (ITCs). The HYDICE runs were orthorectified using prominent tree crowns in the DCM as spatial control points (Chapter 3). Some mismatch in spatial location still existed between the lidar and hyperspectral data sets after HYDICE orthorectification. To minimize this error, ITCs that were digitized on HYDICE imagery were overlaid on the DCM and visually repositioned to align with the ITC apparent in the lidar data, thus creating a second ITC layer. There were also slight differences in crown shape between data sets due to differences in spatial resolution, phenology and other architectural changes (e.g. branch fall) between September, 1997 and March, 1998. For some ITCs in the lidar layer, polygon shapes were modified to conform to the crown shape observed in the DCM. 6.2.2. Lidar metrics I used DCM cells from ITCs to calculate a suite of lidar metrics that characterize crown structure. The first set of metrics quantified crown height and size (Table 6.2). Crown area was the total 2-dimensional area of each ITC (Crownarea). Crown width was calculated as the maximum 2D distance between pairs of cells within the ITC (Crownwidth). Maximum crown height (Maxheight) was the highest DCM cell within an ITC (Table 6.2). I determined crown base height using an automated method outlined in Holmgren and Persson (2004). First, a height-profile histogram with 0.5-m bins was calculated for DCM cells within the ITC (maximum bin was 54.5 to 55.0 m). Next, those bins whose count was <1% of the total ITC cell count where coded as 0, and 1 226 otherwise. The influence of understory vegetation was minimized by passing a 9point median filter along the histogram. The crown base height (Baseheight; Table 6.2) was then set as the first height bin with a value of 1 when moving along the histogram from the 0.0-0.5 height bin upward. Applied in a conifer-dominated forest in Sweden, this technique estimated crown base height with a 0.84 linear correlation (RMSE = 2.82 m), with an overestimation of 0.75 m. Crown height was calculated as: Crownheight = Maxheight – Baseheight (1) Another suite of metrics was calculated from the relative height of DCM cells within each crown. Those ITC cells that had a height value less than the crown base height were removed from the calculation, thereby filtering out cells that did not belong to the crown’s 3-dimensional volume (i.e., heights from understory vegetation). Relative height was calculated by dividing each of the remaining ITC cells by the ITC’s maximum height (Maxheight). The median (Relmed), mean (Relmean), mean 10th-percentile (Rel10perc), mean 90th-percentile (Rel90perc), standard deviation (Relstdev), kurtosis (Relkurtosis), and skewness (Relskewness) were calculated from the relative-height cells within the ITC (Table 6.2; sensu Holmgren & Persson, 2004). Finally, I explored omnidirectional variograms to quantify the spatial properties of relative heights within ITCs (Goovaerts, 1997). Empirical variograms for each ITC were calculated by treating the 33-cm DCM cells as xyz points, where x and y 227 were the 2D spatial location and z was the relative height. In calculated variograms, cells that were lower than the relative base height (i.e., not in the crown volume) were set to the relative base height value. Variograms were calculated with 0.5, 1.5, 3.0 and 4.5-m lags, with a 0.25-m distance tolerance around each lag. The semivariance values at each lag were then used in ITC classification analyses (Table 6.2). 6.2.3. Hyperspectral metrics A suite of hyperspectral metrics were calculated for each ITC from its crownscale spectrum (see Chapter 4 for details). Metrics targeted key photosynthetic pigment, water or other biochemical absorption features and included narrowband indices, derivative-based metrics, absorption-based metrics, and Spectral Mixture Analysis (SMA) fractions (Table 4.4). 6.2.4. Tree species classification techniques I investigated two classifiers for ITC species classification: linear discriminant analysis (LDA: Chapter 3) and decision trees (DT: Chapter 4). In Chapter 3, I used LDA to classify ITC species with optimally-selected HYDICE reflectance bands. Bands were selected using forward-stepwise selection based on discriminant analysis. This method was implemented using the SAS STEPDISC procedure (SAS Institute Inc., Cary, NC, USA). I found that the first 30 significant bands (α = 0.05) from throughout the whole VIS to SWIR spectrum produced the best classification 228 accuracy for seven target species. In Chapter 4, I used decision trees to classify species using the 77 spectral metrics. Similar to methods in Chapters 3 and 4, classifiers were applied to ITCs using a leave-one-out approach. That is, each ITC was classified by holding it out for testing, while all remaining crowns were used for classifier training. Several variable bundles were analyzed separately to assess the relative benefits of lidar and hyperspectral information for species identification. These variable bundles were: 1. Reflectance bands for crown-scale spectra (“reflectance bands”). 2. The 77 spectral metrics from Chapter 4 for crown-scale spectra (“spectral metrics”; Table 4.4). 3. The 17 structural metrics derived from lidar data (“lidar metrics”; Table 6.2). 4. Combined set of variables from lidar (4) and reflectance bands (1) datasets (“lidar + reflectance bands”). 5. Combined set of variables from lidar (4) and spectral metrics (2) datasets (“lidar + spectral metrics”). Three different levels of classification were analyzed: 6 target species (BAEL, DIPA, HYAL, HYME, LEAM, TEOB), 6 target species and an “other” (Other) species class, and DIPA and Other species. For LDA, each variable bundle (1 through 3) was first submitted to a STEPDISC procedure (Chapter 3) with a 95% confidence criteria (α=0.05) for each classification level. 229 Selected variables in bundles 1 and 2 where then combined with selected variables in bundle 3 (lidar) to create variable bundles 4 and 5. In contrast to LDA, all variables within each bundle were used in DT classification. This is because DTs have the ability to rank the importance of variables. If a variable is not useful for discriminating species, it will be left out of the decision rules. Decision tree classification was implemented using methods detailed in Chapter 4. As with LDA, the DT classifier was applied with leave-one-out cross-validation using code in the R statistical package (R Development Core Team, 2004) that interfaced with the Tree package (Tree v1.0-18, R v2.0). Parameters for growing trees were “mincut” of 5, “minsize” of 10, “mindev” of 0.001, and “deviance” as the criteria for splitting data into homogenous sets. Each ITC was classified 50 times using 50 samples per species (with replacement) from the training set, and the final ITC label was chosen using the majority class from the 50 DTs (see Chapter 4). 6.3. Results & Discussion 6.3.1. Differences in lidar metrics among species There were significant differences among means of the six target species for 15 out of 17 lidar metrics (Table 6.3, ANOVA). Maximum tree height (Maxheight), crown base height (Baseheight) and crown height (Crownheight) were top-ranked metrics (Table 6.3; Fig. 6.3). Average tree height was 42 m for the six target species. Individuals of TEOB were the tallest of the six species while BAEL trees were the shortest, and variance was relatively low within each species (Fig. 6.3). 230 The semivariance at lag distances 0.5 and 1.5 m had moderate utility for species discrimination (Table 6.3). The species HYME had dramatically higher semivariance than the other species for 0.5 lags (Fig. 6.3). I observed the same relative pattern in the other lags. There was greater laser penetration into leaf-off HYME crowns relative to crowns from the other leaf-on target species. This effect can be observed in Figure 6.4 for the HYME example crown, where relatively high branches (brighter pixels) are juxtaposed with low, within-crown gaps (dark pixels) due to low leaf cover. These sharp contrasts in height over short distances created relatively high semivariance at all four lags (Table 6.4). In contrast, leaf-on crowns had relatively smooth surfaces (Figs. 2 & 4, DIPA and LEAM) due to less laser penetration, and semivariance values were relatively low (Fig. 6.3, all species except HYME; Table 6.4, DIPA and LEAM). The standard deviation metrics (Relstdev) did not capture the structural differences between leaf-on and leaf-off ITCs as clearly as semivariance (Fig. 6.3; Table 6.3). Individuals of TEOB were generally the tallest of the target species, while DIPA and HYME had the widest and largest crowns (Fig. 6.3, Maxheight, Crownwidth, Crownarea). Individuals of HYME also had very deep crowns (Fig. 6.3, large Crownheight values). This is because their base heights were relatively low while their maximum heights were relatively high (see Eq. 1). As with semivariance discussed above, the sharp contrast in HYME crown height is related to the leaf-off state of the species during the lidar flight. High laser penetration into the crown shifted more DCM cell counts into the lower bins of the crown-height histogram, which tended to lower the estimated crown base height (Fig. 6.4). Providing that the 231 maximum crown height is unaffected by leaf phenology (i.e., the laser detects the top-most branch), then lower leaf cover will tend to increase crown height as base height goes down (i.e., deeper into crown). In contrast, height profiles from leaf-on species such as DIPA and LEAM tended to have most cell counts in the upper-height bins due to relatively low laser penetration (Fig. 6.4); and therefore, their crown base heights were shallow relative to their maximum crown heights; and subsequently, their calculated crown heights were relatively low (Fig. 6.4; Table 6.4). I do not have field data to assess how well base or crown heights were predicted. However, what is important for my analyses is that metrics detect species differences in crown structure as mediated by leaf phenology. The Crownheight metric thus appears quite useful for this purpose. Leaf-off HYME crowns had the lowest (more negative) mean Relkurtosis and highest (less negative) mean Relskewness values (Fig. 6.3). Negative kurtosis indicates a platykurtic distribution of heights, where there are more values than would be expected for a normal distribution (Zar, 1996). For HYME, kurtosis was exaggerated by deeper laser penetration that resulted in more lower-crown versus upper-crown heights (Fig. 6.4 & Table 6.4: HYME). Negative skewness values indicate a distribution that is skewed to the left, with a longer tail toward the relative base height. For the three example crowns in Figure 6.4, laser penetration created a distribution that was less skewed for HYME and DIPA individuals relative to the LEAM individual, which had a strong negative Relskewness (Table 6.4). 232 6.3.2. Decision-tree classification When considering 6 target species, lidar metrics alone (bundle 3) produced a 41.1% overall accuracy. The lidar and spectral metrics (bundle 5) produced the highest overall accuracy, at 69.6%. However, this was only 2.0% more accurate than with spectral metrics alone (Table 6.5; Z = 0.51, not significant; Congalton, 1991). The full-spectrum reflectance bands with and without lidar (bundles 1 & 4) had poorer classification accuracy than when using spectral metrics. I next included an Other species class in the decision trees. This class encompassed all 41 non-target ITCs (18 different species; Table 6.1) and was expected to have more variation in predictor variables relative to target species. With these 7 classes (6 target species + Other class), again the highest accuracy achieved was with lidar metrics combined with spectral metrics (Table 6.5). This overall accuracy, 62.9%, was a 3.6% increase over not using lidar metrics (Z = 0.76, not significant). The Producer’s accuracy (omission error) and User’s accuracies (commission error) of the Other class for all classifications were below 50%, indicating a high level of confusion among non-target and target species. The decision tree was next limited to only DIPA (n=86) and the Other-species class, now comprising ITCs from all non-DIPA species (n=162; Table 6.1). The classification that included only lidar metrics (bundle 3) produced an overall accuracy of 66.9% (Table 6.5). Overall accuracy improved by 17.4% when the analysis was limited to spectral metrics (bundle 2), and DIPA User’s accuracy was 72.8%. Combining lidar metrics with the spectral metrics (bundle 5) did not improve the classification accuracy. 233 These results are similar to those in Chapter 4, in which spectral metrics outperformed reflectance bands in DT classification. The analysis focusing on 6 target species is similar to our previous analysis of 7 species, which achieved an overall accuracy of 70.1%. Overall accuracy with the 6 species was less (67.6%), even though it did not include CEPE, which was poorly classified in the sevenspecies analysis. This discrepancy is likely due to the difference in ITC data between the studies (this study had more DIPA and BAEL, and less LEAM and TEOB). My analyses with the Other class and the addition of lidar metrics are new for this study. In general, accuracies from decision trees are disappointing, and were only acceptable for the DIPA vs. Other classification with spectral metrics. 6.3.3. Linear discriminant analysis classification There were five selected lidar metrics for discriminating the 6 target species: Maxheight, Relmean, Crownarea, Relperc90, and Relkurtosis. The Crownarea, Relperc90 and Relkurtosis metrics had relatively low ranks in the ANOVA results (Table 6.3). This is because the LDA selection finds the combination of variables that best discriminate classes (Tabachnick & Fidell, 1989), while ANOVA assesses whether groups have different means in a single response variable. The stepwiseselected lidar metrics (bundle 3) classified the 6 target species with only 45.9% overall accuracy (Table 6.6). However, this was 4.8% more accurate than when using all lidar metrics in decision trees (Table 6.5). The 34 selected reflectance bands (bundle 1) produced the highest accuracy for the 6 target species (Table 6.6), 234 while classification accuracy was 0.5% less with the addition of the 5 lidar metrics (Z = 0.01, not significant). With the 34 reflectance bands, there was confusion between BAEL and DIPA, and HYME was confused with LEAM, DIPA, and BAEL (Table 6.7). As explained in Chapter 4, leaf-off DIPA and LEAM were confused with BAEL and HYME in the hyperspectral imagery because BAEL and HYME had low LAI and exposed bark due to their compound leaves, giving them spectral properties similar to each other and to leaf-off trees (e.g., Fig. 6.2, BAEL). The addition of lidar metrics decreased confusion between DIPA and BAEL, and between DIPA and HYME. The structural properties of leaf-off (HYME) and low LAI (BAEL) crowns captured by the lidar metrics thus aided in discriminating these species from DIPA, which had lidarstructure properties of a high-LAI crown (Figs. 2 & 4). For HYME, the addition of lidar data improved Producer’s and User’s accuracies by 7.1% and 10.0%, respectively. However, the confusion among species that had high-LAI in lidar (DIPA, HYAL and LEAM) tended to increase because these species had similar crown structure (Figs. 2 & 4). The addition of lidar may thus provide key structural information for discriminating a particular species with distinct phenology (i.e., leafoff in this case), yet it may add to confusion among species with similar structural properties. When the Other species class was included in the LDA classifier, the best accuracy was 81.9% with 41 reflectance bands and 3 lidar metrics (Table 6.6). Overall accuracy was only 0.4% lower without the lidar metrics (Z=0.01, not significant). Lidar metrics alone could only classify species with 36.7% overall 235 accuracy. In general, the User’s and Producer’s accuracy of the DIPA and Other classes were higher with LDA relative to the DT classifier. The final LDA analysis focused on classifying DIPA from the other species, as was done with the DT analysis (Section 3.2). Two significant lidar metrics (bundle 3), Baseheight and Crownarea, were able to classify these two classes with 68.5% overall accuracy (Table 6.6). However, DIPA Producer’s accuracy was very low. In contrast to the other LDA classifications, spectral metrics and lidar metrics (bundle 5) produced the highest overall accuracy, at 90.7%, and DIPA Producer’s and User’s accuracies were 86.0% and 87.1%, respectively. Removing the 2 lidar metrics only lowered the accuracy by 0.04% (not significant) and caused DIPA Producer’s and User’s accuracy to decrease 1.1% and 0.2%, respectively (Table 6.6). The 9 significant spectral metrics from LDA stepwise selection were (in order of importance): Red-A2, ARVI, YE-DArea, NIR2-λ, SR, RE-λ, NE1-Mag, NPV, and SWIR3-A2. There were 11 ITCs committed to the DIPA class (Table 6.8), which decreased User’s accuracy. These crowns included 3 BAEL, 2 CEPE, 1 HYAL, 1 HYME, and 4 LEAM. Another approach to map DIPA crowns was to classify the target species with the Other species class, and then combine the classified non-DIPA target species into one class with the other species. However, this approach did not substantially improve the Producer’s and User’s accuracy of DIPA (Table 6.6). I therefore concluded that the best method for discriminating DIPA from other species was to focus the LDA classifier on two class—DIPA vs. all other species. 236 I also experimented with a probability threshold (see Chapter 3) on the LDA classifier with spectral metrics (bundle 2). In this classification scheme, those ITCs that had a DIPA class probability less than the threshold were left in the Other class. Probability thresholds included 50, 60, 70, 80 and 90 percent. As I increased the probability threshold, DIPA Producer’s accuracy declined because questionable ITCs (i.e., low DIPA probability) were omitted from the DIPA class (Fig. 6.5). However, the stricter probability thresholds tended to increase the DIPA User’s accuracy because fewer non-DIPA crowns were committed to the DIPA class. Overall accuracy remained relatively stable up to the 70% probability threshold. This threshold gave high DIPA User’s accuracy (94.1%; Table 6.9), ensuring that those DIPA in the map were likely to be DIPA in the field. The DIPA Producer’s accuracy dropped to 74.4% with the seventy-percent threshold (Table 6.9), indicating that not all DIPA were mapped (increased omission error). The misclassified DIPA crowns included 2 CEPE, 1 HYME and 1 LEAM. To show the potential of remote sensing for ecological applications, I made a final map of DIPA crowns using the seventy-percent threshold (Fig. 6.6). When compared to the DTM derived from the lidar data (Chapter 2), it can be seen that DIPA were distributed on the edges of low-elevation landforms (old alluvial terraces) and away from water drainages. Similar results are described in Clark et al. (1998), which is not surprising since many of the DIPA crown locations came from data in that study. A true depiction of DIPA crown distributions would be to delineate every tree crown in the HYDICE image through automated means and then classify the crowns as DIPA or Other. The non-random distribution of DIPA in 237 Figure 6.6 is intriguing from both an ecological and remote sensing perspective. Many TRF species have non-random distributions relative to soil water and nutrient gradients (Clark et al., 1998; Condit et al., 2002; Tuomisto, Ruokolainen et al., 2003). If these factors can be mapped economically with remote sensing, then they may provide additional information for species classification (e.g., Franklin, 1998). For example, the lidar-derived DTM could be used to model soil moisture, which could then be incorporated into the classification scheme. However, such an analysis is beyond the scope of this study. 6.4. Conclusions Species leaf phenology is a dominant factor in discriminating tropical rain forest ITCs with either hyperspectral or lidar data. Chapters 3 through Chapters 5 described how ITC leaf cover affected the spectral response of my target species. Here I focused my attention on the contribution of lidar data to species discrimination. Lidar sensors measure the 3-dimensional height distribution of materials within the crown, and thus lidar metrics can be used to quantify crown structure. Deciduous leaf-off species had distinct architectural properties relative to those species with fully-flushed crowns, and these differences were observed in several lidar metrics, such as semivariance and crown height. I found that lidar data alone was insufficient to adequately classify the study species with neither the DT nor LDA classifier. This is likely because the lidar data were acquired in the second dry season (September), when leaf phenological variation among species was relatively low. Combined with the hyperspectral data 238 in the LDA classifier, lidar metrics were mostly useful for improving the accuracy of Hymenolobium, a species that was leaf-on in the hyperspectral data, yet leaf-off in the lidar data; however, the leaf-on species tended to have similar crown architecture, as measured by lidar metrics, and their inter-species confusion increased relative to that from hyperspectral data. In general, there is much more variation in leaf phenology in the primary dry season, between January and April (Table 6.1), when the hyperspectral data were acquired. It is unclear if lidar data acquired in the primary dry season would have improved ITC species discrimination. I conclude that hyperspectral data alone were generally adequate for classifying the target species. In particular, Dipteryx could be classified with 84.9% Producer’s and 86.9% User’s accuracy with LDA and optimal reflectance bands. However, my classification methods require crown-scale spectra from segmented crowns (i.e., polygons). Operational ITC classification will require an automated crown delineation algorithm (Chapter 1). Lidar data may be most useful for crown delineation (Brandtberg et al., 2003; Leckie, Gougeon, Hill et al., 2003; Persson et al., 2002), perhaps in combination with optical data to help distinguish near crowns of similar height. However, lidar-based crown delineation will need to accommodate the variable heights within leaf-off crowns (e.g., Brandtberg et al., 2003). Other types of ecological applications may benefit from the biochemical and structural information offered by hyperspectral and lidar sensors, respectively. One example is estimating aboveground biomass. Lidar data can provide relatively accurate estimates of total aboveground biomass (Drake, Dubayah, Clark, et al., 239 2002), or carbon stocks, across a tropical landscape. Hyperspectral data responds to changes in canopy biochemistry associated with leaf phenology, and thus it may be useful for estimating carbon flux, such as the percentage of canopy leaf turnover through time. 240 Table 6.1. Individual tree crown species and counts. Bold indicates target species. Tree phenology was unknown, evergreen or deciduous (months). For deciduous species, the months indicate when a large percentage of individuals have low leaf area (e.g., leaf-off). For some species, phenology was estimated using data on another species of the same genus. Species Name Albizia sp. Balizia elegans Carapa nicaraguensis Cecropia insignis Cedrela odorata Ceiba pentandra Dipteryx panamensis Dussia macroprophyllata Hyeronima alchorneoides Hymenolobium mesoamericanum Inga alba Lecythis ampla Luehea seemannii Minquartia guianensis Ocotea hartshorniana Pentaclethra macroloba Pterocarpus officinalis Sacoglottis trichogyna Simarouba amara Stryphnodendron microstachyium Tachigali costaricensis Terminalia oblonga Vochysia ferruginea Vochysia guatemalensis a O’Brien, 2001. b Frankie et al., 1974. c Visual verification from videography Abbrevatio n ALSP BAEL CANI CEIN CEOD CEPE DIPA DUMA HYAL HYME INAL LEAM LUSE MIGU OCHA PEMA PTOF SATR SIAM STMI TACO TEOB VOFE VOGU 241 Phenology Count 1 41 5 1 1 6 86 2 26 14 3 17 1 1 1 7 1 2 3 3 1 23 1 1 Unknown Novac Evergreenbc Evergreenbc Jan-Febbc Jan-Marbc Mar-Mayac Janbc Evergreenbc May-Febac Septc Mar-Juneac Maybc Evergreenbc Evergreenb Evergreenbc Septc Evergreenbc Evergreenabc Jan-Febbc Unknownc Evergreenbc Evergreenbc Evergreenbc Table 6.2. Summary of lidar metrics organized by methods. Height & Size Crownarea (m2) Crownwidth (m) Maxheight (m) Baseheight (m) Crownheight (m) Relbaseheight Relmedian Relmean Relperc10 Relperc90 Relstdev Relkurtosis Relskewness Semivariance Lag_0.5 Lag_1.5 Lag_3.0 Lag_4.5 242 Table 6.3. Lidar metric ANOVA results for 6 target species (BAEL, DIPA, HYAL, HYME, LEAM and TEOB), total individual tree crown n=207. Metrics are ranked primarily by F statistic and secondarily by the number of significant differences between species pairs (15 total comparisons). Significance levels for ANOVA F statistic: ns = not significant, * = p≤0.05, ** = p≤0.01, *** = p≤0.001, **** = p≤0.0001. Bold indicates those metrics used in the LDA classification. Rank Metric Mean Std. Dev. Sig. Pairs F 1 Maxheight 42.0 6.0 13.2 **** 7 2 Baseheight 27.4 5.3 10.1 **** 6 3 Crownheight 14.6 4.6 8.9 **** 6 4 Relmean 0.85 0.05 8.4 **** 5 5 Lag_0.5 0.005 0.004 8.2 **** 5 6 Relmedian 0.86 0.05 8.2 **** 5 7 Lag_1.5 0.007 0.006 7.8 **** 5 8 Relperc10 0.69 0.09 6.9 **** 5 9 Lag_3.0 0.008 0.006 6.6 **** 5 10 Relbaseheight 0.65 0.09 6.5 **** 5 11 Lag_4.5 0.010 0.007 5.9 **** 4 12 Relstdev 0.082 0.027 5.8 **** 5 13 Crownarea 372.0 181.6 3.6 ** 1 14 Crownwidth 24.7 6.5 3.1 ** 1 15 Relskewness -0.47 0.34 3.1 ** 2 16 Relperc90 0.97 0.03 2.3 ns 0 17 Relkurtosis -0.44 1.00 0.7 ns 0 243 Table 6.4. Lidar metrics for example individual tree crowns shown in Fig. 6.3. Species abbreviations in Table 6.1. Lag semivariance is multiplied by 100. LEAM DIPA HYME Metric Tree #91 Tree #96 Tree #97 Crownarea 360.9 620.1 958.3 Crownwidth 24.4 29.8 38.0 Maxheight 42.7 48.6 50.9 Baseheight 28.5 32.5 24.0 Crownheight 14.2 16.1 26.9 Relbaseheight 0.67 0.67 0.47 Relmean 0.85 0.84 0.73 Relperc90 0.96 0.95 0.92 Relperc10 0.71 0.70 0.51 Relstdev 0.07 0.07 0.12 Relkurtosis -0.52 -0.58 -0.70 Relskewness -0.50 -0.33 -0.35 Relmedian 0.86 0.85 0.76 Lag_0.5 0.24 0.50 1.28 Lag_1.5 0.34 0.62 1.67 Lag_3.0 0.46 0.72 1.89 Lag_4.5 0.57 0.80 2.04 Number of DCM pixels 3239 5570 8643 244 Table 6.5. Decision tree classification accuracy. Kappa variance multiplied by 100. Sensor combination reflectance bands spectral metrics lidar metrics lidar + reflectance bands lidar + spectral metrics 245 reflectance bands spectral metrics lidar metrics lidar + reflectance bands lidar + spectral metrics reflectance bands spectral metrics lidar metrics lidar + reflectance bands lidar + spectral metrics No. DIPA DIPA Other Other Vars Prod User Prod User 6 Target Species 161 58.1 71.4 n/a n/a 77 77.9 82.7 n/a n/a 17 48.8 67.7 n/a n/a 178 66.3 69.5 n/a n/a 94 81.7 77.9 n/a n/a 6 Target and Other Species 161 57.0 68.1 26.8 47.8 77 73.3 79.7 34.1 45.2 17 51.2 65.7 19.5 26.7 178 64.0 71.4 34.1 42.4 94 76.7 77.6 39.0 48.5 DIPA and Other 161 81.4 62.5 74.1 88.2 77 87.2 72.8 82.7 92.4 17 67.4 51.8 66.7 79.4 178 87.2 62.0 71.6 91.3 94 86.0 73.3 83.3 91.8 245 Overall Kappa Kappa Var. 55.1 67.6 41.1 58.0 69.6 0.42 0.57 0.25 0.45 0.60 0.19 0.17 0.18 0.20 0.17 48.4 59.3 35.5 52.8 62.9 0.37 0.50 0.21 0.42 0.54 0.14 0.14 0.12 0.14 0.14 76.6 84.3 66.9 77.0 84.3 0.52 0.67 0.32 0.54 0.67 0.31 0.24 0.39 0.29 0.24 246 Table 6.6. Linear discriminant analysis classification accuracy. Bands were selected using stepwise linear discriminant analysis with statistical significance set at α =0.05. Kappa variance multiplied by 100. No. DIPA DIPA Other Other Kappa Sensor combination Vars Prod User Prod User Overall Kappa Var. 6 Target Species reflectance bands 34 90.7 87.6 n/a n/a 88.9 0.85 0.09 spectral metrics 24 86.0 91.4 n/a n/a 83.6 0.78 0.12 lidar metrics 5 47.7 72.1 n/a n/a 45.9 0.22 0.26 lidar + reflectance bands 39 89.5 90.6 n/a n/a 88.4 0.85 0.09 lidar + spectral metrics 29 83.7 91.1 n/a n/a 83.6 0.78 0.12 6 Target and Other Species reflectance bands 41 86.0 84.1 61.0 67.6 81.5 0.77 0.10 spectral metrics 27 81.4 84.3 51.2 67.7 76.6 0.71 0.11 lidar metrics 3 76.7 39.5 17.1 23.3 36.7 0.12 0.21 lidar + reflectance bands 44 84.9 86.9 63.4 72.2 81.9 0.77 0.09 lidar + spectral metrics 30 83.7 85.7 53.7 64.7 79.8 0.75 0.10 DIPA and Other reflectance bands 6 80.2 83.1 91.4 89.7 87.5 0.72 0.22 spectral metrics 9 84.9 86.9 93.2 92.1 90.3 0.79 0.17 lidar metrics 2 26.7 60.5 90.7 70.0 68.5 0.20 0.01 lidar + reflectance bands 8 77.9 82.7 91.4 88.6 86.7 0.70 0.23 lidar + spectral metrics 11 86.0 87.1 93.2 92.6 90.7 0.79 0.17 246 Classification Table 6.7. Error matrix for LDA classification with 34 reflectance bands without and with 5 lidar metrics. Classification 247 Species BAEL DIPA HYAL HYME LEAM TEOB Total Prod. Species BAEL DIPA HYAL HYME LEAM TEOB Total Prod. Reflectance Bands Field Reference HYAL HYME LEAM TEOB 1 2 3 2 24 9 1 1 14 23 26 14 17 23 92.3% 64.3% 82.4% 100.0% BAEL 36 4 1 41 87.8% DIPA 5 78 1 2 86 90.7% BAEL 36 3 2 41 87.8% Reflectance Bands + Lidar Metrics Field Reference DIPA HYAL HYME LEAM TEOB 4 1 2 77 2 3 2 23 10 3 2 14 23 86 26 14 17 23 89.5% 88.5% 71.4% 82.4% 100.0% 247 Total 42 89 26 10 17 23 207 87.8% User 85.7% 87.6% 92.3% 90.0% 82.4% 100.0% Total 43 85 27 10 19 23 207 User 83.7% 90.6% 85.2% 100.0% 73.7% 100.0% 88.9% 88.4% Classification Table 6.8. Error matrix for Dipteryx individual tree crown classification using 9 stepwise-selected spectral metrics and LDA with no threshold (Kappa = 0.79). Reference Species DIPA Other Total User’s DIPA 73 11 84 86.9% Other 13 151 164 92.1% Total 86 162 248 Producer’s 84.9% 93.2% 90.3% Classification Table 6.9. Error matrix for Dipteryx individual tree crown classification using 9 stepwise-selected spectral metrics and LDA with a 70-percent threshold (Kappa = 0.76). Reference Species DIPA Other Total User’s DIPA 64 4 68 94.1% Other 22 158 180 87.8% Total 86 162 248 Producer’s 74.4% 97.5% 90.5% 248 Figure 6.1. The La Selva Biological Station study site and extent of HYDICE hyperspectral and FLI-MAP lidar datasets. The 248 study crowns are labeled with black polygons. 249 BAEL DIPA Tree#108 Tree#58 HYAL HYME Tree#56 Tree#247 LEAM TEOB Tree#127 Tree#25 CEPE INAL Tree#162 Tree#197 Figure 6.2. Example individual tree crowns from color videography acquired simultaneously with the lidar data. Video time codes are UTC. Species codes are listed in Table 6.1. 250 40 50 Baseheight (m) Maxheight (m) 60 40 30 20 10 0 30 20 10 0 BAEL DIPA HYAL HYME LEAM TEOB BAEL DIPA HYAL HYME LEAM TEOB Crownarea (m 2) Crownheight (m) 30 25 20 15 10 5 0 700 600 500 400 300 200 100 0 35 30 25 20 15 10 5 0 BAEL DIPA HYAL HYME LEAM TEOB 1.0 0.8 Relmean Crownwidth (m) BAEL DIPA HYAL HYME LEAM TEOB 0.6 0.4 0.2 0.0 BAEL DIPA HYAL HYME LEAM TEOB 0.15 2.0 1.5 Relstdev Lag_0.5 Semivariance BAEL DIPA HYAL HYME LEAM TEOB 1.0 0.5 0.10 0.05 0.00 0.0 BAEL DIPA HYAL HYME LEAM TEOB BAEL DIPA HYAL HYME LEAM TEOB BAEL DIPA HYAL HYME LEAM TEOB 0.00 -0.5 -0.25 -1.0 -1.5 -2.0 Relskewness Relkurtosis BAEL DIPA HYAL HYME LEAM TEOB 0.0 -0.50 -0.75 -1.00 Figure. 6.3. Species mean (bars) and standard deviation (error bars) for selected lidar metrics calculated from target-species tree crowns (n=207). Species codes are listed in Table 6.1. Semivariance is multiplied by 100. 251 LEAM Tree #91 DIPA Tree #96 HYME Tree #97 Height (m) DIPA – Tree # 96 Height (m) 50 48.6 m 40 16.1 m 50.0 25 Meters 0.0 32.5 m 30 20 Maxheight HYME – Tree #97 10 50 100 150 200 250 300 Height (m) Count LEAM – Tree #91 50.9 m 40 26.9 m 30 24.0 m 20 50 42.7 m Height (m) 40 Crownheight 0 50 Baseheight 10 14.2 m 30 28.5 m 0 50 100 150 200 250 300 Count 20 10 0 50 100 150 200 250 300 Count Figure 6.4. A subset of the digital canopy model (DCM) with three example individual tree crowns. Graphs depict the vertical height profile for each crown with count of DCM pixels in each height 0.5-m height bin. The horizontal dashed line in each graph is the maximum crown height and the solid line is the base height (Baseheight), as calculated by the automated algorithm. The value between the bars is the crown height (Eq. 1). 252 100 90 Percent 80 70 DIPA User's 60 DIPA Producer's 50 Other User's 40 Other Producer's 30 Overall 20 10 0 None 50% 60% 70% 80% 90% LDA Probability Threshold Figure 6.5. Overall accuracy and User’s and Producer’s accuracy for DIPA (Dipteryx) and other species (Other) with change in linear discriminant analysis probability threshold. Figure 6.6. Map of DIPA (Dipteryx) crowns using the LDA, 70-percent threshold classifier (Table 6.9). Four DIPA crowns were incorrectly classified. 253 CHAPTER 7: Conclusions 7.1. Summary of research Overview The primary goal of this research was to assess two types of emerging remote sensing technology, hyperspectral and lidar sensors, for the automated, species-level mapping of individual tropical rain forest trees. Such maps will be useful for broadscale forest inventory, prioritization for protection, long-term monitoring, management and ecological analyses. The study was conducted in a species-rich tropical wet forest in Costa Rica and focused on emergent individuals of 7 out of more than 400 tree species. The findings presented here are therefore not exhaustive, but rather represent a first attempt to conduct automated species classification with these new types of remotely-sensed data. The hyperspectral measurements respond to the biochemical and structural properties of crowns, while the lidar measurements respond to crown structure. The central question guiding this research was: can species could be discriminated based on their spectral or structural properties? Does the combined biochemical and structural information offered by both lidar and hyperspectral sensors improve species classification accuracy? To answer these questions, I outlined several objectives in Section 1.2: 254 1. Develop hyperspectral techniques for classifying tropical rain forest tree species Five different hyperspectral classification techniques were investigated (Chapters 3-5). The linear discriminant analysis produced the best accuracy when presented with an optimal set of 30 reflectance bands selected from across the full 400 to 2500 nm spectrum. This technique can be implemented using existing statistical software packages. 2. Identify the optimal spectral regions and spatial scale for species discrimination Initial species classification analyses using hyperspectral data spanned leaf, pixel to crown scales. Although the leaf scale had accuracies reaching 100%, it was not a realistic assessment of the technology in an operational setting because spectra lacked atmospheric noise and spectral mixing. At operational scales, species were generally best discriminated using crown-scale relative to pixel-scale spectra, indicating that very high spatial resolution imagery is not needed for species discrimination. Leaf phenology was a major factor that caused species-level differences in spectral absorption features. The visible, near-infrared and shortwave infrared regions were all useful for detecting this phenological spectral variation. 3. Evaluate the importance of lidar-derived crown structure information for species discrimination Many properties of crown structure measured from the lidar dataset were influenced by tree leaf phenology. However, most species were leaf-on at the time of the lidar acquisition, and there was relatively low variation in lidar-derived 255 metrics among leaf-on species for adequate species discrimination. The hyperspectral data, which responded to biochemical and structural properties of crowns at a time of high phenological contrast, were sufficient for distinguishing species. 4. Assess lidar technology for ecological analyses of tropical rain forests Although lidar-derived crown structure was not sufficient for species discrimination, the technology did prove useful for other types of ecological analysis. The terrain surface, generated as a by-product of vegetation analyses, had remarkable detail and impressive accuracy, even below structurally-complex oldgrowth forest. The lidar sensor was also promising for scaling plot-scale forest structure parameters (e.g., height) to landscape scales. Chapter 2 summary Chapter 2 involves the pre-processing of the small-footprint lidar dataset for later analyses of individual tree crown (ITC) structure at LSBS (Chapter 6). A fullyautomated, local-minima algorithm was developed to separate lidar ground returns from overlying vegetation returns in the original lidar height surface. The IDW and OK geostatistical techniques were then used for interpolating a sub-canopy DTM. OK was determined to be a superior interpolation scheme because it smoothed finescale variance created by spurious understory heights in the ground-point dataset. The final DTM had a strong linear-correlation of 1.00 and a RMSE of 2.29 m when compared against 3859 well-distributed ground-survey points. In old-growth forests, 256 RMSE on steep slopes was 0.67-m greater than on flat slopes. On flatter slopes, variation in vegetation complexity associated with land-use caused highly significant differences in DTM error distribution across the landscape. The highest DTM accuracy observed in this study was on flat, open-canopy areas with relatively smooth surfaces. Lidar ground-retrieval was complicated by dense, multi-layered evergreen canopy in old-growth forests, causing DTM overestimation. A DCM was calculated by subtracting the DTM from the original lidar surface. Individual and plot-scale heights were estimated from DCM metrics and compared to field data measured using similar spatial supports and metrics. For old-growth forest emergent trees and isolated pasture trees, individual tree heights were underestimated and had 3.67 and 2.33-m mean absolute error, respectively. Linearregression models explained 51% (4.15-m RMSE) and 95% (2.41-m RMSE) of the variance, respectively. It was determined that improved elevation and field-height estimation in pastures explained why individual pasture trees could be estimated more accurately than old-growth trees. Mean height of tree stems in 32 young plantation plots (0.38 to 18.53-m tall) was estimated with a mean absolute error of 0.90 m (r2=0.97; 1.08-m model RMSE) using the mean of lidar returns in the plot. As in other small-footprint lidar studies, plot mean height was underestimated; however, the plot-scale results from this analysis had stronger linear models than previously-reported models for temperate-zone conifer and deciduous hardwoods. 257 Chapter 3 summary Chapter 3 investigates the utility of high spectral and spatial resolution imagery for the automated species-level classification of ITCs in the LSBS old-growth forest. Laboratory spectrometer and airborne reflectance spectra (161 bands, 437-2434 nm) were acquired from seven species of emergent trees. Analyses focused on leaf-, pixel- and crown-scale spectra. The spectral regions and factors that most influenced spectral separability among species were reviewed. Next, spectral-based species classification was performed using traditional classifiers, spectral angle mapper (SAM), maximum likelihood (ML) and linear discriminant analysis (LDA), in combination with a stepwise band selection procedure. Optimal regions of the spectrum for species discrimination varied with scale. However, near-infrared (7001327 nm) bands were consistently important regions across all scales. Bands in the visible region (437-700 nm) and shortwave infrared (1994-2435 nm) were more important at pixel and crown scales. Overall classification accuracy decreased from leaf scales measured in the laboratory to pixel and crown scales measured from the airborne sensor. The highest crown-scale ITC accuracy was 92% with LDA and 30 bands. Producer’s accuracies ranged from 70% to 100% and User’s accuracies ranged from 81% to 100%. The SAM classifier performed poorly at all scales and spectral regions of analysis. ITCs were also classified using a pixel-majority approach in which crown species labels were assigned according to the majority class of classified pixels within a crown. An overall accuracy of 86% was achieved with a pixel-majority LDA classifier applied to 30 bands of data. Pixel-majority and crown-scale ITC 258 classifications were significantly more accurate with 10 narrow-bands relative to accuracies achieved with simulated multispectral, broadband data. Chapter 4 summary Chapter 4 approaches ITC classification with a less conventional technique. Classification variables included hyperspectral metrics that responded to crown structure and absorption features from photosynthetic pigments, water and other biochemicals. The metrics included narrowband indices, derivative-based metrics, absorption-based metrics and spectral mixture analysis fractions that were calculated from spectra acquired at tissue (leaf, bark), pixel, and crown scales. Differences in metrics among species were ranked using statistical tests. Leaf and pixel-scale spectra were best discriminated by near-infrared water absorption features while bark and crown-scale spectra were better distinguished by shortwave infrared biochemical absorption features. Differences in spectral metrics among species at pixel and crown scales were largely dependent on tree leaf phenology and structure, which controlled the relative amounts of leaf and bark tissues within a crown. ITC species were classified with a decision tree (DT) classifier using within-crown pixel spectra and a pixel-majority classification rule or with crown-scale spectra. The best classification scheme was with crown-scale metrics and had 70.1% overall accuracy. Hyperspectral metrics and DT classifiers were instructive for identifying key spectral reflectance properties for tropical tree species discrimination. In terms of number of times appearing in DTs, important metrics characterized absorption 259 features across the whole 437-2435-nm spectrum, but particularly in the shortwave infrared region (1467-2435 nm). Chapter 5 summary Chapter 5 explores multiple endmember spectral mixture analysis (MESMA) for classifying ITCs. To increase computational efficiency of MESMA, an automated technique was developed to select optimal endmembers for each species from image and laboratory libraries. Candidate endmembers were used to model other spectra in the library, and endmembers that maximized within-species modeling capability where chosen over more generalist endmembers. The species of ITCs were determined by applying the MESMA classifier to crown-scale spectra or to pixel-scale spectra, followed by a pixel-majority classification. MESMA classification with two- and three-endmember models was explored. With a single endmember per species and a shade endmember in twoendmember models, MESMA could discriminate ITC species with 48.6% and 50.0% with crown-scale and pixel-majority classifications, respectively. Increasing classification accuracy required the addition of endmembers to accommodate additional within-species spectral variability likely caused by spectral mixtures. Pixel-majority overall accuracy reached 90.2% and 91.6% when including the full spectral library as potential two- and three-endmember models, respectively. However, the processing time needed to evaluate this large number of models was determined to be too restrictive for operational use. The ability of MESMA to detect 260 unique spectral mixtures for each species is an encouraging result, and several avenues for future research were identified. Chapter 6 summary Spectral and structural properties of tree crowns are important for visual discrimination of species in aerial photography. It was expected that these properties would be useful for computer-based species classification with remotely-sensed data. Chapter 6 focuses on using hyperspectral and lidar data to classify the species of ITCs in tropical rain forest. Crown-scale spectral data from the HYDICE sensor combined with structural data from the FLI-MAP lidar sensor were used in classification analyses. There were significant differences in the majority of lidarderived metrics among the study tree species, indicating that species have unique structural properties. Crown leaf cover, especially in deciduous leaf-off trees, was the primary factor controlling variation in lidar metrics. Following methods in Chapters 3 and 4, the LDA and DT classifiers were used for discriminating tree species, but at three levels of class aggregation. The best classifier was LDA at all three levels of aggregation. Overall accuracy with spectral data was 88.9% when classifying 6 target species, 81.5% for target species with an other-species class, and 90.3% when discriminating a species of conservation interest, Dipteryx, from other species. In contrast, overall accuracy with lidar data alone was 45.9%, 36.7% and 68.1% for these same levels of classification, respectively. The addition of lidarderived structure information to the classifier did not improve classification accuracies. However, lidar did help increase the accuracy of a particular species 261 with distinct leaf-off phenology at the cost of additional confusion among leaf-on species. 7.2. Conclusions and recommendations The main objective this research was to assess two types of emerging remote sensing technology, hyperspectral and lidar sensors, for the discrimination of tropical rain forest tree species. The hyperspectral data contained information on the biochemical and structural properties of crowns, while the lidar data contained structural information. I hypothesized that these two datasets combined would allow greater species classification accuracy than either dataset alone. I did not find that the structural information from lidar to be useful for species classification. Considerable effort was required to prepare the lidar dataset for analysis of individual tree crown structure (Chapter 2), and improvement in species classification accuracy with lidar was negligible or absent (Chapter 6). However, a byproduct of the lidar pre-processing, the DTM, had impressive detail and was quite accurate, even under dense rain forest canopy (Chapter 2). For comparison, the 3800+ survey points took weeks to measure in the field and still did not provide the same level of terrain detail as the FLI-MAP sensor, which was flown in just 2 days. Lidar thus proved itself as a powerful new technology for generating DTMs in tropical landscapes. Although large-footprint sensors can cover larger areas in the same amount of time, my analyses showed that small-footprint sensors offer greater DTM detail and accuracy. There are no commercially-available large-footprint sensors at this time, and small-footprint systems are still prohibitively expensive for 262 campaigns covering a whole region, such as a large watershed. However, costs are expected to decline as the technology matures. As a reference, bids for a 2005 smallfootprint, multi-return lidar acquisition at LSBS and vicinity covering an area approximately 6,000 ha were priced at US$95,000 or more. Roughly half of this cost is for mobilizing the equipment and team from North America, and so costs should drop as the technology spreads to other regions of the world. The DCM has many uses in ecological applications that were not fully explored in this research. I found that plot-scale estimates of forest height were more accurate than ITC height estimates (Chapter 2), and since stand height has an allometric relationship to aboveground biomass, it is expected that fine-scale, lidar-derived DCMs can be used to predict carbon stocks and forest structure parameters (Lim et al., 2003; Popescu et al., 2003). Such a strong relationship to plantation and oldgrowth forest structure has been found for a large-footprint system at LSBS (Drake, Dubayah, Clark et al., 2002). However, as with the DTM, a small-footprint sensor provides a more detailed perspective on forest structure that may be more appropriate for certain kinds of ecological research, such as tracking tree-fall gap dynamics, identifying wildlife habitat, or scaling point-based data (e.g., tower eddyflux carbon measurements) to a broader scale. Hyperspectral data were found to be critical for remote classification of TRF tree species. Species were relatively easy to distinguish based on their leaf-scale reflectance properties measured in the laboratory; however, accuracy decreased when using the airborne hyperspectral data (Chapter 3). The importance of spectral regions also varied with scale. Therefore, studies that classify laboratory reflectance 263 and infer crown-level species separability (e.g., Cochrane, 2000) should be interpreted with caution. A more realistic test of species separability is to use data from airborne or spaceborne imaging spectrometers, with all of the associated variability introduced by poor radiometric calibration, atmospheric contamination and illumination geometry. There are two general findings from my research that are encouraging for operational classification of TRF tree species. First, the elaborate methods and labor involved in implementing the DT classifier with hyperspectral metrics or the MESMA classifier with optimal endmembers did not translate into improved species classification accuracy relative to using reflectance bands with LDA, a more traditional classifier found in many statistical packages. Another important finding was that crown-scale spectra provided more accurate ITC species classification, with both LDA and DT classifiers, than the pixel-majority approach, and there was no advantage in isolating sunlit crown spectra from the entire crown spectra. These results suggest that image spatial resolution does not need to be extremely fine for species discrimination, as long as pixels are not so coarse that they mix crown radiance with background radiance (e.g., canopy gaps, other trees). This research investigated hyperspectral imagery with 1.6-m pixels, and there may be an optimum spatial resolution greater than 1.6-m for species discrimination. However, one advantage of this high resolution imagery was that it allowed the accurate delineation of crowns, thereby minimizing spectral mixing with the background. As mentioned, coarser spatial resolution imagery will not allow such detailed delineation of crown boundaries. 264 Chapters 3 and 4 both identified the SWIR region as important for species discrimination. Most analyses of tropical leaf spectra have covered only the VIS and NIR regions (Cochrane, 2000; Fung et al., 1998; Poorter et al., 1995), likely because hyperspectral sensors that include SWIR are generally more expensive. Results in Chapter 3 indicate that a multispectral sensor with 6 SWIR bands (ASTER) tended to increase accuracy over sensors with two broad bands (Landsat ETM+) or no bands (IKONOS) in SWIR. In addition, Chapter 3 and 4 both showed that fullspectrum information—either with optimally-selected bands or in absorption-feature metrics—provided optimal species discrimination. One advantage of hyperspectral data is they are over-sampled, and information that does not optimize separation among species can be discarded. Leaf phenology was found to be an important consideration in mapping TRF tree species. In the hyperspectral image, deciduous Dipteryx and Lecythis trees in near leaf-off conditions had distinct volume-scattering and spectral mixing properties that influenced the selection of reflectance bands (Ch. 3), spectral metrics (Ch. 4), and endmembers (Ch. 5). It is unclear from my research whether classification accuracies would have changed if the imagery had been flown in the wet season, when Dipteryx and Lecythis have fully-flushed crowns. Also, phenological events such as senescence before leaf drop or flowering produce changes in reflectance spectra that may be amplified in leaf-on conditions due to volumetric scattering. If these changes in spectral reflectance occur synchronously in all individuals of the target population and do not overlap in time with other species, then image acquisition may be timed to maximize classification accuracy for a particular target 265 species. For example, Dipteryx has pink flowers and is known to flower with a peak between May and August when trees have leaves (Frankie et al., 1974; Newstrom et al., 1994; O’Brien, 2001). However, TRF tree phenology is complex, little understood, and difficult to generalize; at LSBS, Ceiba (CEPE) flower in the dry season when trees are leafless, Terminalia (TEOB) populations have variation in annual flowering intensity, and some overstory trees in the LSBS forest have asynchronous flowering among individuals of the same species (Frankie et al., 1974). In the lidar data, Hymenolobium was leaf-off and had drastically different structural properties relative to leaf-on species (Ch. 6). Many of the leaf-on species in the lidar data had similar structural attributes due to low laser penetration. Classification accuracy may have benefited from the addition of lidar if the data had been acquired simultaneously with the hyperspectral data in the driest season, when phenological differences were more pronounced and the laser could penetrate deeper into the crowns of other leaf-off species. 7.3. Directions for future research It is clear from Chapters 3-6 that studies seeking to discriminate TRF tree species with remote sensing technology should focus efforts on hyperspectral data analysis. Since hyperspectral imagery is typically acquired at spatial scales > 1.6 m, there is a need for a sensitivity analysis to determine the trade-offs between pixel resolution, ability to detect crown position and shape, spectral mixing with background materials, and classification accuracy. 266 A limitation of my hyperspectral analyses is that they infer the phenological state and associated biophysical properties of tree species based on field data and observations not directly linked to the study ITCs. I recommend that future studies acquire ITC-level estimates of crown structure (e.g., leaf and branch area) and other biological information, such as liana cover, epiphyte cover, and flower density through either field observation or interpretation of very high spatial resolution imagery. Such data acquired at the time of image acquisition will permit a deeper understanding of the conspecific and interspecific spectral variation among ITCs that is expressed in canopy-level reflectance spectra or spectral metrics. Lidar analyses suggest that TRF tree crowns are not easily distinguished based on crown structure. However, besides measuring crowns when phenological variation was relatively low, my analyses were also constrained by first-return lidar data. The FLI-MAP sensor was state-of-the-art in 1997, but since then there has been a steady improvement in lidar technology that has permitted higher post density and multiple-return recording. For example, an ALTM 3100 lidar sensor (Optech Inc., Toronto, Canada) scheduled to fly over LSBS in July, 2005 has the capability to record 4 returns, including a last return. As discussed in Chapter 2, I expect that ground retrieval will be more accurate with this last-return data; and consequently, the DCM should also have improved vertical accuracy. In my analyses, I did not have access to the original xyz point dataset from the FLI-MAP lidar acquisition. Instead, the DSM data were interpolated from the xyz points, and some detail of crown structure was lost in the interpolation process. The ALTM 3100 dataset includes xyz data as part of the delivered products. With multiple returns and point 267 data, it is likely that the ALTM 3100 data will contain a richer representation of ITC internal structure. This improved detail may translate into greater species separability, especially if metrics were calculated directly from the point data, rather than through an interpolated surface (for example, see Brandtberg et al., 2003; Holmgren & Persson, 2003). I found that videography, even with its poor image quality, provides a useful aerial perspective for assessing ITC phenological state in the lidar data. The lidar campaign at LSBS will also acquire simultaneous, 18-cm orthorectified color imagery (ALTM 4k x 4k Digital Camera, Optech Inc., Toronto, Canada). ITC properties that are difficult to measure from the ground, such as leaf phenology, liana coverage, and epiphyte load, should be visually-interpretable in the aerial view afforded by 18-cm imagery, providing an unprecedented opportunity to investigate how these factors affect variability in lidar metrics. Since the digital data will include 3 multispectral bands (blue, green, red), there are also opportunities to investigate how crown structure, as measured by the lidar, affects spectral response and texture in imagery from optical sensors. Although hyperspectral sensors have immense potential for tree species discrimination, one major limitation encountered in this research was a lack of representative trees for each species; I only analyzed 7 out of 400+ tree species at LSBS. Given natural variability within populations and individuals of TRF tree species, I do not expect that hyperspectral and lidar technology are capable of discriminating all tree species in the forest canopy. Auxiliary variables may be needed to constrain classification rules, such as with the inclusion of soil moisture 268 variables from a lidar-derived DTM (Chapter 6). However, I have demonstrated that with hyperspectral imagery, certain species of interest may be discernable from the canopy-matrix of non-target species, especially if the target species have sharp phenological contrast. The capability of hyperspectral imagery to map target species thus permits remote and systematic monitoring of long-term changes in key-stone, endemic, rare or commercial tree species caused by factors, such as selective logging and climatic change, which are otherwise undetectable by coarse-resolution multispectral sensors. Higher classification accuracy may be found by grouping ITCs into functional types rather than individual species. Tree functional types may be based on common growth form, metabolism, water balance or disturbance properties (Box, 1996; Nobel & Gitay, 1996), and not all functional types have a physical manifestation (e.g. structure, chemistry) which can be detected by remote sensing. Deciduousness is an important tropical tree functional type that responds to climate, such as changes in regional precipitation patterns (Condit et al., 1996; Condit et al., 2000). Knowledge of the proportion of deciduous crowns in a canopy can be used to calibrate remote sensing estimates of forest parameters, such as aboveground biomass, productivity, or chlorophyll content (Condit et al., 2000). Leaf phenology of deciduous and evergreen species was a factor driving species separability in this research, and I expect that high spatial resolution hyperspectral and lidar data would be useful for mapping crowns into deciduous and evergreen classes. One potential application is to use a detailed map of deciduous and evergreen ITCs to calculate the 269 proportion of canopy occupied by deciduous trees. This information could then be used as reference data for studying how canopy-level deciduousness affects the spectral response of coarser spatial resolution imagery, such as from the Moderate Resolution Imaging Spectroradiometer (MODIS) with 250-m to 1000-m resolution pixels. Hyperspectral and lidar technology also have great potential for the remote sensing of aboveground biomass (AGB), a variable vital for assessing carbon stocks and flux at broad scales and for input into biogeochemical models (Hall et al., 1995). Tropical forests contain a large proportion of terrestrial carbon, and consequently have the greatest potential to increase atmospheric carbon dioxide from deforestation (Dixon et al, 1994), yet remote sensing AGB estimates from optical and synthetic aperture radar sensors tend to saturate in older secondary and old-growth TRF forests (Huete et al., 2002; Imhoff, 1995; Luckman et al., 1998; Steininger, 1996; Steininger, 2000). Drake, Dubayah, Clark et al. (2002) showed that lidar metrics applied to large-footprint waveforms have immense potential for estimating AGB over a wide range of tropical forest conditions without saturation. Small-footprint lidar has also been used to assess AGB, volume and tree height at stand scales for commercial forestry applications (Holmgren et al., 2003; Lim et al., 2003; Nilsson, 1996), and the plot-scale analysis of tree height in Chapter 2 could be extended to estimate biomass of plantation, secondary or old-growth forests. In tropical forests, a large proportion of plot-level AGB is explained by the largest trees (Brown & Lugo, 1992; Clark & Clark, 2000; Nascimento and Laurance, 270 2002). One advantage of small-footprint lidar data is that large ITCs can be detected in the canopy, and then AGB may be estimated for each tree from crown-level lidar metrics. These ITC estimates of AGB can then be aggregated to calculate plot- or stand-scale AGB. For example, Popescu et al. (2003) found that ITC diameter information from small-footprint lidar improved stand-scale AGB estimates for mixed conifer forests. With the ITC approach to AGB estimation, high spatial multispectral or hyperspectral imagery may be useful to calibrate AGB regression models to account for species differences in LAI, such as through the incorporation of the GV or NPV fractions from spectral mixture analysis. Furthermore, species composition explains a broad gradient of variation in AGB across Amazonian tropical forests due to taxonomic differences in wood specific gravity (Baker et al., 2004), yet the variable is generally not used when calculating AGB from allometric equations. Perhaps hyperspectral imagery can be most beneficial for biomass estimation if it is used to classify ITCs by wood specific gravity functional types. Such a classification could be developed from a reclassification of species-level (or genera-level) ITC maps, such as those found in this research. However, if wood specific gravity is related to deciduousness (Borchert, 1994; Choat et al., 2005), then it may be best mapped indirectly from phenology functional classes. This research also provides a foundation for applications which seek to estimate species richness across broad spatial scales. Several studies have shown that the spatial and temporal properties of NDVI may useful for predicting plant species richness (Bawa et al., 2002; Fairbanks & McGwire, 2004; Gould, 2000; Oindo & 271 Skidmore, 2002). NDVI was used as a predictor variable because it is linked to ecosystem net primary productivity and biomass as well as because red-well and NIR bands needed to compute NDVI are available in multispectral satellite sensors. Other spectral metrics may be more useful; for example, a metric that incorporates SWIR information should respond to species richness by detecting canopy variability in leaf phenology. The lidar DCM may also provide information on canopy structure, such as size and distribution of canopy gaps, which could be related to species richness (Denslow, 1995). This research encompasses an initial exploration of yet immature hyperspectral and lidar technology for future ecological applications in tropical forests. Considerable additional research is needed with different species, temporal variation, and sensors to confirm and extend my results. Despite the global importance of tropical rain forests, there is a major lack of hyperspectral and lidar data from tropical landscapes, mainly due to cost constraints and large distance between tropical study sites and major research institutions. 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International Journal of Remote Sensing, 25(4), 859-865. 305 APPENDIX I: List of Acronyms AGB Above-Ground Biomass ARVI Atmospherically Resistant Vegetation Index ASTER Advanced Spaceborne Thermal Emission & Reflection Radiometer BE Blue Edge COBI COunt-Based Index DCM Digital Canopy Model DSM Digital Surface Model DT Decision Tree DTM Digital Terrain Model EAR Endmember Average Root mean square error ETM Enhanced Landsat Thematic Mapper EVI Enhanced Vegetation Index EWT Equivalent Water Thickness FOV Field of View GP Green Peak GV Green Vegetation HYDICE HYperspectral Digital Imagery Collection Experiment IDW Inverse Distance Weighted ITC Individual Tree Crown 306 LAI Leaf Area Index LDA Linear Discriminant Analysis LIDAR LIght Detection And Ranging LSBS La Selva Biological Station MAE Mean Absolute Error MESMA Multiple Endmember Spectral Mixture Analysis ML Maximum Likelihood MODIS MODerate Resolution Imaging Spectroradiometer NASA National Aeronautics and Space Administration NDVI Normalized Difference Vegetation Index NDWI Normalized Difference Water Index NE Near-infrared water Edge NIR Near InfraRed NPMANOVA Non-Parametric Multivariate Analysis of Variance NPV Non-photosynthetic Vegetation PRI Photochemical Reflectance Index OK Ordinary Kriging RE Red Edge RMSE Root Mean Square Error RVSI Red-edge Vegetation Stress Index RW Red Well SAM Spectral Angle Mapper 307 SAR Synthetic Aperature Radar SAVI Soil-Adjusted Vegetation Index SE Shortware-infrared Edge SMA Spectral Mixture Analysis SqrtMAE Square-Root Transformation of the Mean Absolute Error SR Simple Ratio SWIR ShortWave InfraRed TRF Tropical Rain Forest VIS Visible WBI Water Band Index 308 APPENDIX II: Summary of spectral metrics (Chapter 4) 309 Appendix 2.1. Global mean and standard deviation (in parentheses) of narrow-band, ratio-based indices for the seven study species at different scales of measurement. Significant differences among species means were tested with an ANOVA test. Pair-wise multiple comparisons between species means tested with the Tukey’s Honestly Significant Difference method (p≤0.05). Significance levels for ANOVA F statistic: ns = not significant, * = p≤0.05, ** = p≤0.01, *** = p≤0.001, **** = p≤0.0001. Bark n=66; Leaf n=152; Pixel n=2100; Crown n=214. Mean Mean Mean Mean F F F F Metric (S.D.) Pairs (S.D.) Pairs (S.D.) Pairs (S.D.) Pairs Vegetation Indices SR 2.37 1.6 ns 9.59 8.1 **** 14.43 244.8 **** 12.87 64.3 **** (0.88) 0 (3.96) 9 (6.02) 19 (5.19) 15 NDVI 0.38 2.3 * 0.78 4.7 *** 0.84 173.4 **** 0.83 36.9 **** (0.12) 1 (0.09) 5 (0.08) 18 (0.06) 13 SAVI 0.28 6.1 **** 0.57 7.4 **** 0.54 64.1 **** 0.54 39.1 **** (0.10) 6 (0.08) 6 (0.15) 16 (0.09) 12 PRI -0.38 2.3 * -0.78 4.7 *** -0.84 173.4 **** -0.83 36.9 **** (0.12) 1 (0.09) 5 (0.08) 18 (0.06) 13 EVI 0.05 4.8 *** 0.10 7.3 **** 0.09 64.5 **** 0.09 38.9 **** (0.02) 4 (0.02) 7 (0.03) 15 (0.02) 12 309 310 Appendix 2.1. (continued) Mean F Metric (S.D.) Pairs ARVI 0.20 1.5 ns (0.14) 0 RVSI 0.006 9.1 **** (0.00) 10 Mean Mean F F (S.D.) Pairs (S.D.) Pairs 0.74 2.9 ** 0.77 197.1 **** (0.10) 0 (0.10) 18 0.001 8.1 **** 0.015 78.8 **** (0.01) 9 (0.00) 18 Liquid Water Content Indices WBI 0.94 14.0 **** 1.01 16.0 **** 1.09 155.5 **** (0.09) 10 (0.02) 9 (0.09) 18 NDWI -0.14 14.8 **** 0.02 14.5 **** 0.03 280.0 **** (0.12) 10 (0.03) 9 (0.05) 18 Feature abbreviations listed in Table 4.4. 310 Mean (S.D.) 0.76 (0.09) 0.014 (0.00) F Pairs 42.8 **** 12 31.0 **** 14 1.09 (0.07) 0.03 (0.04) 19.1 **** 12 48.2 **** 14 311 Appendix 2.2. Global mean and standard deviation (in parentheses) of derivative-based metrics for the seven study species at different scales of measurement. Tests are the same as in Appendix 2.1. Bark Leaves Pixels Crowns Mean Mean Mean Mean F F F F Metric (S.D.) Pairs (S.D.) Pairs (S.D.) Pairs (S.D.) Pairs Derivative Inflection Wavelength Position (nm) BE-λ 518.7 2.3 * 521.8 4.9 *** 519.9 16.3 **** 519.9 9.3 **** (3.3) 1 (1.7) 4 (2.4) 8 (1.2) 5 GP-λ 552.4 5.9 **** 555.8 39.0 *** 554.9 10.3 **** n/d n/d (4.1) 4 (12.3) 14 (11.0) 5 YE-λ 572.8 4.9 *** 572.0 1.1 ns 570.5 3.0 ** 570.4 0.6 ns (4.1) 6 (2.4) 0 (6.5) 2 (5.0) 0 RW-λ 663.7 1.4 ns 669.5 5.5 **** 665.9 44.2 **** 666.0 11.2 **** (2.1) 0 (6.5) 7 (1.8) 14 (1.0) 7 RE-λ 692.7 3.3 ** 712.1 28.7 **** 725.2 58.4 **** 724.7 9.0 **** (8.9) 1 (7.5) 10 (4.4) 15 (3.2) 6 NE1-λ 1028.2 2.7 * 1030.6 3.1 ** 996.0 13.9 **** 992.9 3.2 ** (25.9) 1 (25.9) 2 (17.0) 10 (5.4) 1 NE2-λ 1144.3 2.9 * 1143.1 2.1 ns 1130.2 5.0 **** 1127.6 0.2 ns (5.8) 1 (3.4) 0 (8.5) 3 (8.4) 0 SE-λ 1498.0 2.8 * 1496.1 7.2 **** 1522.0 36.6 **** 1512.8 12.1 **** (9.8) 3 (3.8) 6 (21.1) 14 (16.7) 9 311 312 Appendix 2.2. (continued) Derivative Inflection Magnitude (x 1000) or Percent Reflectance BE-Mag 0.65 8.1 **** 1.72 5.0 *** 0.88 31.0 **** 0.84 (0.27) 6 (0.88) 3 (0.38) 12 (0.21) GP-Refl 93.79 3.0 ** 42.85 13.3 **** 42.81 n/d n/d (35.1 2 (18.6) 10 (8.9) YE-Mag 0.27 4.7 *** -1.27 1.8 ns -0.49 41.2 **** -0.37 (0.20) 5 (1.18) 0 (0.31) 15 (0.18) RW-Refl 138.2 3.7 ** 51.67 2.9 * 25.40 51.5 **** 26.51 (47.7) 1 (27.5) 2 (13.3) 16 (9.4) RE-Mag 2.92 11.5 **** 7.72 8.5 **** 5.34 70.7 **** 5.03 (1.63) 8 (1.48) 6 (2.49) 15 (1.50) NE1-Mag 0.93 6.3 **** 0.24 9.8 **** 1.32 86.6 **** 1.27 (0.41) 6 (0.14) 6 (0.64) 17 (0.42) NE2-Mag -1.00 2.9 * -0.62 32.5 **** -2.91 56.1 **** -2.38 (1.30) 2 (0.22) 15 (1.62) 14 (1.01) SE-Mag 1.09 2.9 * 1.22 9.3 **** 0.91 15.3 **** 0.93 (0.25) 1 (0.20) 7 (0.43) 12 (0.21) 312 18.0 **** 8 4.6 *** 5 26.6 **** 10 19.1 **** 11 40.0 **** 12 17.6 **** 12 15.5 **** 10 5.9 **** 5 Appendix 2.2. (continued) 313 Derivative-based Area (x 100) BE2.30 8.1 **** 4.31 4.9 *** 2.35 22.4 **** 2.28 12.0 **** DArea (0.96) 6 (2.21) 3 (0.99) 12 (0.53) 7 YE1.22 6.0 **** -1.95 2.9 * -0.70 116.2 **** -0.63 49.7 **** DArea (0.61) 7 (1.25) 1 (0.51) 16 (0.37) 13 RWE21.12 13.3 **** 36.08 6.6 **** 33.52 56.1 **** 32.19 32.8 **** DArea (9.06) 11 (6.95) 5 (14.3) 15 (7.99) 12 RWE13.76 4.1 ** 38.88 6.9 **** 35.22 53.7 **** 33.77 31.5 **** DNArea (9.57) 4 (7.98) 6 (15.4) 15 (8.61) 12 RWE0.22 8.3 **** 0.06 8.0 **** 0.05 41.5 **** 0.05 9.4 **** 2DNArea (0.14) 7 (0.10) 6 (0.05) 14 (0.03) 7 Feature abbreviations listed in Table 4.4. λ = wavelength, Mag = derivative magnitude, Refl = percent reflectance, DArea = area under 1st derivative, DNArea = area under normalized 1st derivative, 2DNArea = area under 2nd derivative 313 314 Appendix 2.3. Global mean and standard deviation (in parentheses) of absorption-based metrics for the seven study species at different scales of measurement. Tests are the same as in Appendix 2.1. Bark Leaves Pixels Crowns Mean Mean Mean Mean F F F F Metrics (S.D.) Pairs (S.D.) Pairs (S.D.) Pairs (S.D.) Pairs Equivalent Water Thickness EWT 0.07 3.7 ** 0.04 14.8 **** 0.31 160.2 **** 0.30 15.8 **** (0.11) 3 (0.03) 12 (0.17) 16 (0.13) 12 Maximum Depth (% Reflectance) Blue-D 0.06 1.6 ns 0.32 11.4 **** 0.41 73.0 **** 0.36 35.6 **** (0.03) 0 (0.14) 10 (0.14) 16 (0.10) 11 Red-D 0.29 2.4 * 0.80 4.7 *** 0.83 162.3 **** 0.82 36.5 **** (0.13) 1 (0.08) 4 (0.09) 18 (0.07) 13 NIR1-D 0.03 2.9 * 0.02 20.0 **** 0.13 165.9 **** 0.12 14.7 **** (0.04) 2 (0.01) 11 (0.05) 16 (0.05) 14 NIR2-D 0.05 2.7 * 0.04 47.3 **** 0.17 145.0 **** 0.17 15.1 **** (0.05) 2 (0.01) 15 (0.07) 16 (0.06) 12 SWIR1-D 0.01 3.3 * 0.06 35.9 **** 0.06 9.1 **** (0.01) 1 n/d n/d (0.02) 15 (0.02) 4 SWIR2-D 0.07 36.0 **** 0.06 45.1 **** n/d n/d n/d n/d (0.03) 14 (0.02) 13 SWIR3-D 0.05 20.9 **** 0.03 15.9 **** 0.10 10.5 **** 0.07 22.3 **** (0.03) 11 (0.02) 011 (0.06) 10 (0.01) 12 314 Appendix 2.3. (continued). Blue-λ Red-λ NIR1-λ 315 NIR2-λ SWIR1-λ 495.8 (2.9) 672.3 (4.1) 990.0 (9.9) 1177.3 (12.1) 1727.5 (10.3) 1.1 ns 0 3.6 ** 1 3.1 ** 3 4.0 ** 3 4.5 ** 4 n/d 2293.7 (18.8) n/d 22.2 **** 11 SWIR2-λ SWIR3-λ Wavelength (nm) 498.1 12.3 **** 492.8 (5.6) 10 (7.0) 678.6 4.1 *** 670.0 (2.4) 5 (0.9) 994.1 2.9 * 958.0 (14.5) 1 (12.1) 1174.5 3.1 ** 1168.6 (9.1) 0 (12.9) 1749.3 n/d n/d (3.8) 2103.8 n/d n/d (27.6) 2307.9 2.4 * 2313.9 (24.0) 1 (23.6) 315 5.1 **** 6 0.6 ns 0 16.0 **** 10 8.3 **** 6 16.9 **** 11 26.3 **** 14 10.5 **** 11 495.8 (2.5) 669.9 (0.0) 957.7 (8.6) 1169.4 (9.6) 1748.4 (1.1) 2097.9 (21.1) 2299.2 (7.4) 2.7 * 1 3.7 ** 4 2.8 * 1 4.6 *** 4 1.8 ns 1 9.3 **** 9 1.5 ns 0 Appendix 2.3. (continued). Blue-W 316 44.1 (2.9) Red-W 73.6 (11.0) NIR1-W 49.9 (17.5) NIR2-W 74.0 (4.2) SWIR1-W 26.8 (12.0) SWIR2-W n/d SWIR3-W 57.1 (20.5) Blue-A1 Blue-A2 Red-A1 Red-A2 NIR1-A1 2.5 (1.1) 2.5 (1.1) 22.3 (13.0) 22.5 (12.2) 2.1 (3.1) 1.2 ns 0 3.2 ** 3 6.9 **** 6 3.8 ** 3 3.0 * 1 45.8 (3.8) 101.6 (9.1) 61.9 (15.8) 73.0 (3.1) Width (nm) 6.6 **** 7 5.7 **** 5 4.7 *** 3 3.9 ** 3 n/d n/d n/d 14.8 **** 9 n/d 51.7 (28.1) 1.9 ns 1 1.9 ns 1 2.2 ns 1 2.3 * 1 3.3 ** 2 14.8 (7.3) 14.2 (6.9) 81.4 (13.2) 76.9 (12.1) 1.2 (0.6) n/d 11.5 **** 8 Area 11.1 **** 10 11.4 **** 10 4.5 *** 4 4.3 *** 4 25.8 **** 11 316 46.4 (3.4) 109.2 (5.8) 63.6 (10.0) 74.2 (12.1) 44.1 (9.4) 69.1 (29.8) 50.3 (25.0) 20.7 **** 12 179.3 **** 18 31.6 **** 15 5.7 **** 6 196.4 **** 18 158.4 **** 17 19.5 **** 13 47.2 (1.4) 108.1 (4.7) 65.5 (7.0) 77.5 (9.2) 45.7 (7.4) 78.5 (22.7) 65.1 (12.5) 23.4 **** 9 46.9 **** 12 3.1 ** 2 0.4 ns 0 59.4 **** 14 38.4 **** 13 0.3 ns 0 19.0 (6.5) 18.1 (6.1) 91.3 (13.3) 85.5 (12.4) 8.2 (3.5) 90.9 **** 17 75.2 **** 17 199.6 **** 18 206.9 **** 18 246.8 **** 18 17.3 (5.1) 16.7 (4.9) 89.0 (11.4) 83.3 (10.7) 8.1 (3.1) 36.3 **** 11 36.6 **** 11 46.8 **** 13 49.0 **** 13 18.8 **** 14 317 Appendix 2.3. (continued). NIR1-A2 1.9 3.6 ** (3.3) 3 NIR2-A1 3.8 2.7 * (3.6) 2 NIR2-A2 3.7 2.8 * (3.5) 2 SWIR10.4 2.4 * A1 (0.4) 0 SWIR10.2 3.3 * A2 (0.7) 1 SWIR2A1 n/d n/d SWIR2A2 n/d n/d SWIR32.7 5.5 *** A1 (1.4) 3 SWIR33.1 10.7 **** A2 (1.5) 9 1.2 (0.7) 2.6 (0.9) 2.5 (0.8) 20.8 **** 11 58.4 **** 15 56.8 **** 15 n/d n/d n/d n/d n/d n/d n/d 1.9 (1.6) 1.6 (1.7) n/d 20.6 **** 11 17.3 **** 11 317 8.3 (3.9) 13.0 (5.5) 12.9 (5.5) 2.9 (1.3) 2.7 (1.6) 4.9 (3.2) 3.7 (3.6) 4.8 (3.4) 4.1 (5.0) 221.8 **** 16 160.7 **** 16 160.3 **** 16 79.0 **** 19 72.9 **** 19 102.5 **** 14 144.2 **** 16 8.9 **** 9 4.7 **** 4 8.2 (3.4) 12.8 (4.4) 12.7 (4.4) 3.0 (1.0) 2.9 (1.2) 4.7 (2.3) 4.0 (2.6) 4.3 (1.3) 4.1 (1.1) 18.7 **** 14 16.4 **** 13 16.2 **** 13 21.6 **** 7 21.3 **** 7 62.6 **** 13 67.4 **** 13 8.4 **** 5 20.2 **** 11 Appendix 2.3. (continued). Asymmetry (x 100) 6.6 **** 91.0 20.4 **** 90.9 22.6 **** 7 (0.6) 12 (0.3) 9 Red-As 5.1 **** 84.7 184.1 **** 84.8 48.3 **** 3 (0.8) 18 (0.6) 12 NIR1- As 4.9 *** 93.6 28.1 **** 93.4 2.7 * 3 (1.0) 14 (0.7) 2 NIR2- As 3.7 ** 93.9 6.0 **** 93.6 0.5 ns 3 (1.0) 6 (0.8) 0 SWIR197.5 196.6 **** 97.4 59.5 **** As n/d n/d (0.5) 18 (0.4) 14 SWIR296.8 159.5 **** 96.4 38.9 **** As n/d n/d n/d n/d (1.4) 17 (1.0) 13 SWIR397.5 14.8 **** 97.8 11.6 **** 97.9 19.9 **** 97.2 0.4 ns As (0.9) 9 (1.2) 8 (1.1) 13 (0.5) 0 Feature abbreviations listed in Table 4.4. n/d = not detected. D = depth, λ = wavelength, W=width, A1 = area calculated using width and depth, A2 = area calculated using tabulation, As = Asymmetry Blue-As 318 91.5 (0.5) 89.5 (1.5) 95.1 (1.7) 93.9 (0.3) 98.5 (0.7) 1.2 ns 0 3.3 ** 3 6.8 **** 6 3.6 ** 2 3.0 * 1 91.2 (0.7) 85.7 (1.2) 94.0 (1.5) 94.0 (0.2) 318 Appendix 2.4. Global mean and standard deviation (in parentheses) of Spectral Mixture Analysis (SMA) fractions and model error among the study species (3 endmember model) at pixel and crown scales of measurement. Tests are the same as in Appendix 2.1. Pixels Crowns Fraction Mean No. Mean No. (S.D.) Pairs (S.D.) Pairs F F 41.3 76.7 39.3 37.5 GV (18.4) **** 17 (11.4) **** 12 15.1 189.4 17.2 43.8 NPV (14.9) **** 18 (11.6) **** 14 43.6 19.4 43.5 8.1 Shade (20.9) **** 11 (9.2) **** 7 1.2 23.3 1.0 5.2 RMSE (0.5) **** 13 (0.3) **** 3 GV = % green vegetation, NPV= % non-photosynthetic vegetation, Shade = % photometric shade, RMSE=root mean square error, units percent reflectance. 319
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