Jeannine Cavender-Bares John A. Gamon Philip A. Townsend *Editors*

# Remote Sensing of Plant Biodiversity

Remote Sensing of Plant Biodiversity

Jeannine Cavender-Bares • John A. Gamon Philip A. Townsend Editors

# Remote Sensing of Plant Biodiversity

*Editors* Jeannine Cavender-Bares Ecology, Evolution & Behavior University of Minnesota Saint Paul, MN, USA

Philip A. Townsend Department of Forest and Wildlife Ecology University of Wisconsin–Madison Madison, WI, USA

John A. Gamon University of Nebraska–Lincoln, School of Natural Resources Lincoln, NE, USA

University of Alberta, Departments of Earth & Atmospheric Sciences and Biological Sciences Edmonton, AB, Canada

#### ISBN 978-3-030-33156-6 ISBN 978-3-030-33157-3 (eBook) https://doi.org/10.1007/978-3-030-33157-3

© The Editor(s) (if applicable) and The Author(s) 2020

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# **Foreword**

At last, here it is. For some time now, the world has needed a text providing both a new theoretical foundation and practical guidance on how to approach the challenge of biodiversity decline in the Anthropocene. This is a global challenge demanding global approaches to understand its scope and implications. Until recently, we have simply lacked the tools to do so. We are now entering an era in which we can realistically begin to understand and monitor the multidimensional phenomenon of biodiversity at a planetary scale. This era builds upon three centuries of scientific research on biodiversity at site to landscape levels, augmented over the past two decades by airborne research platforms carrying spectrometers, lidars, and radars for larger-scale observations. Emerging international networks of fine-grain in-situ biodiversity observations complemented by space-based sensors offering coarsergrain imagery—but global coverage—of ecosystem composition, function, and structure together provide the information necessary to monitor and track change in biodiversity globally.

This book is a road map on how to observe and interpret terrestrial biodiversity across scales through plants—primary producers and the foundation of the trophic pyramid. It honors the fact that biodiversity exists across different dimensions, including both phylogenetic and functional. Then, it relates these aspects of biodiversity to another dimension, the spectral diversity captured by remote sensing instruments operating at scales from leaf to canopy to biome. The biodiversity community has needed a Rosetta Stone to translate between the language of satellite remote sensing and its resulting spectral diversity and the languages of those exploring the phylogenetic diversity and functional trait diversity of life on Earth. By assembling the vital translation, this volume has globalized our ability to track biodiversity state and change. Thus, a global problem meets a key component of the global solution.

The editors have cleverly built the book in three parts. Part 1 addresses the theory behind the remote sensing of terrestrial plant biodiversity: why spectral diversity relates to plant functional traits and phylogenetic diversity. Starting with first principles, it connects plant biochemistry, physiology, and macroecology to remotely

sensed spectra and explores the processes behind the patterns we observe. Examples from the field demonstrate the rising synthesis of multiple disciplines to create a new cross-spatial and spectral science of biodiversity.

Part 2 discusses how to implement this evolving science. It focuses on the plethora of novel in-situ, airborne, and spaceborne Earth observation tools currently and soon to be available while also incorporating the ways of actually making biodiversity measurements with these tools. It includes instructions for organizing and conducting a field campaign. Throughout, there is a focus on the burgeoning field of imaging spectroscopy, which is revolutionizing our ability to characterize life remotely.

Part 3 takes on an overarching issue for any effort to globalize biodiversity observations, the issue of scale. It addresses scale from two perspectives. The first is that of combining observations across varying spatial, temporal, and spectral resolutions for better understanding—that is, what scales and how. This is an area of ongoing research driven by a confluence of innovations in observation systems and rising computational capacity. The second is the organizational side of the scaling challenge. It explores existing frameworks for integrating multi-scale observations within global networks. The focus here is on what practical steps can be taken to organize multi-scale data and what is already happening in this regard. These frameworks include essential biodiversity variables and the Group on Earth Observations Biodiversity Observation Network (GEO BON).

This book constitutes an end-to-end guide uniting the latest in research and techniques to cover the theory and practice of the remote sensing of plant biodiversity. In putting it together, the editors and their coauthors, all preeminent in their fields, have done a great service for those seeking to understand and conserve life on Earth—just when we need it most. For if the world is ever to construct a coordinated response to the planetwide crisis of biodiversity loss, it must first assemble adequate—and global—measures of what we are losing.

Woody Turner Earth Science Division NASA Headquarters, Washington, DC, USA

# **Contents**

#### Contents

# **About the Authors**

**Christiana Ade** is a PhD student in the Civil and Environmental Engineering Department at the University of California Merced. She is passionate about using remote sensing to study changes in wetland vegetation diversity and water quality, and evaluate how these changes may impact key Earth system cycles.

**Julián Aguirre-Santoro** is an Assistant Professor at the Natural Sciences Institute of the National University of Colombia. His research focuses on taxonomy, phylogenetics, biogeography, and macroevolution of Neotropical plant lineages.

**Doreen Boyd** is a Professor in the School of Geography, University of Nottingham, UK. Her research focuses include active and passive remote sensing, ecology and conservation, and human rights intersections with environmental change.

**Betsabe de la Barreda-Bautista** is a research associate at the School of Geography, University of Nottingham (UK), working on remote sensing for environmentalhuman dynamics, principally monitoring how vegetation responds to climatological and anthropogenic events.

**Nicholas Basinger** is an Assistant Professor of Weed Science at the University of Georgia. His research focuses on weed ecology, biology, and diversification of the agroecosystem and non-cropland ecosystems to reduce the impacts of weedy species.

**Erik Bolch** is a Master's student in Environmental Systems at the University of California Merced. His research focuses utilizing UAS imaging spectroscopy to study biodiversity and invasive plants.

**Jan Bumberger** heads the working group "Environmental Sensor and Information Systems" and is the scientific coordinator for research data management at the Helmholtz Centre for Environmental Research – UFZ, Germany. His research focuses on scalable sensor network technologies as well as the calibration and validation of remote sensing data with broadband electromagnetic and optical spectral methods.

**Ana Carnaval** was born and raised in Brazil. She received her Master's degree in Zoology from the Museu Nacional do Rio de Janeiro and has a PhD in Evolutionary Biology from The University of Chicago. She is an Associate Professor at the City College of CUNY, and her research focuses on the historical biogeography of the Atlantic forest fauna and flora.

**Jeannine Cavender-Bares** is a Distinguished McKnight University Professor in the Department of Ecology, Evolution and Behavior at the University of Minnesota. Her research focuses on the ecology and evolution of plant function, applying phylogenetics and spectral data to community ecology, and remote sensing of biodiversity.

**Jennifer Costanza** is a Research Assistant Professor in the Department of Forestry and Environmental Resources at North Carolina State University. Her research focuses on forest dynamics, the ecological effects of global change, land change modeling, and disturbance ecology.

**Kyla Dahlin** is an Assistant Professor in the Department of Geography, Environment, and Spatial Sciences and the Ecology, Evolutionary Biology, and Behavior Program at Michigan State University. Her research aims to better understand and quantify ecosystem processes and disturbance responses through the application of emerging technologies, including air- and spaceborne remote sensing, spatial statistics, and process-based modeling.

**Néstor Fernández** is a conservation biology researcher interested in the advancement of biodiversity conservation solutions through the integration of multiple sources of earth observations in order to quantify and model changes in terrestrial ecosystem dynamics and in species distributions.

**Andrew O. Finley** is a Professor at the Departments of Forestry and Geography, Environment, and Spatial Sciences at Michigan State University. His research interests lie in developing methodologies for monitoring and modeling environmental processes, Bayesian statistics, spatial statistics, and statistical computing.

**Keith Gaddis** is the Senior Support Scientist for the National Aeronautics and Space Administration's Biological Diversity and Ecological Forecasting Programs. He is an ecologist and biogeographer by training with expertise using remote sensing and genetics to address questions in ecology, evolution, and conservation biology.

**John Gamon** is a Professor in Earth and Atmospheric Sciences and Biological Sciences at the University of Alberta, and also conducts airborne remote sensing research in vegetation biodiversity and productivity at the School of Natural Resources at the University of Nebraska, Lincoln. His interests include ecophysiology, ecosystem metabolism, biodiversity, ecoinformatics, and sustainability.

**Gary Geller** has a PhD in Biology from UCLA where he studied the interaction of plants with their environment, focusing on modeling plant architecture. At NASA/ JPL he combines that experience with system engineering and has worked with the NASA Ecological Forecasting program for the last 15 years, including two years seconded to the Group on Earth Observations in Geneva.

**Hamed Gholizadeh** is an Assistant Professor in the Geography Department at Oklahoma State University. His research focuses on using proximal, airborne, and spaceborne remote sensing to study biodiversity.

**Renato Goldenberg** is a Full Professor in the Department of Botany at Universidade Federal do Paraná, Brazil. His research focuses on systematics of flowering plants.

**John Grady** was a postdoctoral fellow at Michigan State University and Bryn Mawr College at the time of writing this chapter and is now a postdoctoral fellow at the National Great Rivers Research and Educational Center. He is broadly interested in how traits affect interactions and emergent features of ecology.

**Ryan P. Hanavan** is a Research Entomologist and the Remote Sensing Program Manager with the Forest Health Assessment and Applied Sciences Team with the US Forest Service. His research focuses on early detection remote sensing techniques for insect and disease threats.

**Erin Hestir** is an Associate Professor in Civil and Environmental Engineering at the University of California Merced. Her research focuses on aquatic ecosystems under threat from competing pressures to meet societal needs for water and food security while sustaining biodiversity and other ecosystem services.

**Marco Heurich** is Head of the Department of Visitor Management and National Park Monitoring of Bavarian Forest National Park and Associate Professor of Wildlife Ecology at the University of Freiburg. His research focuses on exploring movement ecology and biodiversity by using a variety of remote sensing techniques from camera traps to satellites.

**Martina Hobi** is a Group Leader in the Forest Resources and Management Research Unit at the Swiss Federal Institute for Forest, Snow, and Landscape Research WSL. She specializes in the analysis of forest dynamical processes using a combination of terrestrial, dendroecological, and remote sensing data.

**Sarah Hobbie** is a Distinguished McKnight University Professor in the Department of Ecology, Evolution and Behavior at the University of Minnesota. She studies global change impacts on ecosystems, urban ecosystem ecology, and human impacts on urban water resources.

**Stéphane Jacquemoud** is a Professor of Remote Sensing in the Institut de Physique du Globe de Paris at the University of Paris. His research focuses include remote sensing of natural surfaces, leaf and soil radiative transfer models, and imaging spectroscopy.

**Miriam Kaehler** has worked as a visiting scholar at the Institute of Systematic Botany at the New York Botanical Garden, and as a post-doc research fellow at Universidade de São Paulo and Universidade Federal do Paraná, both in Brazil. Her research focuses on molecular systematics and taxonomy of Bignoniaceae, as well as plant inventory.

**Shruti Khanna** is a Senior Environmental Scientist at the California Department of Fish and Wildlife. Her research focuses on understanding invasion ecology of nonnative plants through remote sensing.

**Doug King** is a Distinguished Research Professor in the Department of Geography and Environmental Studies, Carleton University, Ottawa, Canada. His research focuses on high-resolution UAV to satellite scale imaging for vegetation composition, structure, productivity and health modeling, mapping, and monitoring in forest, wetland, agriculture, and tundra environments.

**Mayara Krasinski Caddah** is a Professor in the Department of Botany at the Federal University of Santa Catarina, Brazil. Her research focuses on plant taxonomy and evolution, applying phylogenetics and population genetics to genera and species delimitation.

**Daniel Kükenbrink** is a PhD candidate at the Remote Sensing Laboratories of the University of Zurich. Three-dimensional forest reconstruction and the radiative transfer in forest ecosystems are among his main research interests.

**Andrew Latimer** is a Professor in the Department of Plant Sciences at the University of California Davis. He studies how plant populations and communities respond to change, including sudden, major disturbance such as fire and drought, as well as more gradual changes in climate.

**Angela Lausch** is head of the working group functional landscape ecology and Remote Sensing at the Helmholtz Centre for Environmental Research – UFZ, and Privat Dozent (PD) at the Humboldt University to Berlin. She uses hyperspectral remote sensing on airborne, drone, and laboratory platforms at the UFZ. Her research focuses on remote sensing, trait and functional ecology, bio- and geodiversity, data science, linked open data, and the semantic web.

**Reik Leiterer** is a Senior Scientist at the University of Zurich, Switzerland. His aim is to enable more robust and operational environmental monitoring using datadriven methods.

**Lúcia Lohmann** is a Professor in the Department of Botany at the University of São Paulo, Brazil. Her research is highly integrative, combining components of classic taxonomy, phylogenetics, molecular biology, ecology, evolution, and biogeography to address questions associated with the assembly and evolution of the Neotropical biota.

**Mike Madritch** is a Professor in the Department of Biology at Appalachian State University. His research focuses on linking aboveground plant communities with belowground microbial communities and ecosystem functioning.

**Paul Magdon** is a Senior Scientist at the Chair of Forest Inventory and Remote Sensing at the University of Göttingen, Germany. In his research he addresses methodological questions, relevant for the development of remote sensing–assisted forest and biodiversity monitoring systems.

**Sparkle Malone** is an Assistant Professor in the Department of Biological Sciences at Florida International University. Her primary research focus is to improve our understanding of how climate and disturbance regimes influence spatial and temporal variability in ecosystem structure and function using remote sensing, eddy covariance, and spatial and temporal models.

**Roberta Martin** is an Associate Professor in the School for Geographical Sciences and Urban Planning and the Center for Global Discovery and Conservation Science at the Arizona State University. Her research incorporates field and laboratory studies of trees and corals with aircraft- and satellite-based maps to understand the biodiversity and health of forests and reefs.

**Kyle C. McDonald** is Terry Elkes Professor in the Earth and Atmospheric Sciences Department at the City College of New York, City University of New York. Professor McDonald studies terrestrial ecosystems with satellite-borne remote sensing instruments.

**Jose Eduardo Meireles** is an Assistant Professor in the School of Biology and Ecology at the University of Maine. His research focuses on plant phylogenetics, population genetics, and evolutionary ecology.

**Fabián A. Michelangeli** is a Curator in the Institute of Systematic Botany at The New York Botanical Garden. His research focuses on tropical plant taxonomy, systematics, and their evolution.

**Felix Morsdorf** is a Group Leader in the Geography Department at the University of Zurich. He uses laser scanners, drones, and physical modeling to derive diversityrelated metrics of vegetation structure.

**Scott Ollinger** is a Professor in the Department of Natural Resources and the Environment at the University of New Hampshire. His research interests span a variety of topics within the fields of ecology and biogeochemistry including carbon and nitrogen cycling, forest productivity and succession, plant-soil interactions, remote sensing, ecosystem modeling, and the effects of multiple environmental stressors on forests.

**Brian O'Meara** studies macroevolution by developing and applying phylogenetic models. He is a Professor in the Department of Ecology and Evolutionary Biology at the University of Tennessee, Knoxville.

**Stephanie Pau** is an Associate Professor in the Department of Geography at Florida State University. Her research focuses on the diversity, structure, and function of forest and savanna systems, often combining field surveys with remote sensing imagery and statistical modeling.

**Ryan Pavlick** is a Research Technologist at the Jet Propulsion Laboratory, California Institute of Technology; working on observing and understanding Earth's biodiversity, terrestrial ecosystems and carbon cycle from space.

**Andrea Paz** has an MSc in Biology and is a PhD candidate at the City University of New York, USA. She is interested in understanding the influence of environments in determining species distributions and diversity patterns in the Neotropics.

**Henrique Miguel Pereira** is a Professor of Biodiversity Conservation at the German Centre for Integrative Biodiversity Research (iDiv) at the Martin-Luther University Halle-Wittenberg and the co-chair of the Group on Earth Observations Biodiversity Observation Network. His research uses a combination of field work, long-term monitoring, and modeling to understand the drivers of global biodiversity change.

**Jesús N. Pinto-Ledezma** is a Grand Challenge in Biology Postdoctoral Fellow in the Department of Ecology, Evolution and Behavior at the University of Minnesota. He works with both macroecological and evolutionary theory and methods with the aim of understanding the processes that underlie biodiversity patterns at different temporal and spatial scales.

**Erika Podest** is a scientist with the Carbon Cycle and Ecosystems Group at NASA's Jet Propulsion Laboratory in Pasadena, California. Her research focuses on the use of microwave remote sensing for characterizing wetland inundation dynamics and vegetation growing season length in the northern high latitudes.

**Jennifer Pontius** is a Research Ecologist with the US Forest Service and Director of the Environmental Science Program at the University of Vermont. Her research focuses on developing novel remote sensing methods to assess forest health, structure, and productivity to better inform the management of forested ecosystems.

**Quentin Read** was a postdoctoral fellow at Michigan State University and Bryn Mawr College at the time of writing this chapter and is now a postdoctoral fellow at the National Socio-Ecological Synthesis Center. A community ecologist by training, he is now working on quantifying the environmental and ecological impacts of food waste in the United States.

**Martin Reader** is a PhD student in the Department of Geography at the University of Zurich. He is examining the impacts of human modification on river delta biodiversity and ecosystem services.

**Sydne Record** is an Associate Professor in the Department of Biology at Bryn Mawr College. Her research focuses on modeling the past, present, and future states of the natural world to better understand the services that ecosystems afford to society.

**Marcelo Reginato** is a Professor in the Department of Botany at the Universidade Federal do Rio Grande do Sul, Brazil. His research focuses include plant systematics, biogeography, and macroevolution.

**Duccio Rocchini** is Professor in Biology and Ecology at the University of Trento, Italy. His research focuses on biodiversity analysis at multiple spatial scales, ecological informatics, plant community ecology, remote sensing, spatial ecology, spatio-ecological modeling by open-source software, and species distribution modeling.

**Maria J. Santos** is a Professor of Earth System Science in the Department of Geography, University of Zurich. Her research focuses on understanding coupled social-ecological systems and their impacts on biodiversity and ecosystem services, using field, archival, and remote sensing data, GIS, and statistical modeling.

**Michael E. Schaepman** is a Professor of Remote Sensing and Vice President of Research at the University of Zurich, Switzerland. Following a focus on optical engineering and designing imaging spectrometers, he moved on to signal processing and radiative transfer modeling and is currently working on remotely sensing ecological genomics.

**Gabriela Schaepman-Strub** is a Professor of Earth System Science in the Department of Evolutionary Biology and Environmental Studies at the University of Zurich, Switzerland. Her research focuses on radiation-vegetation interaction from leaf to canopy scale, Arctic biodiversity and ecosystem functions, and land surface energy fluxes under climate change.

**David Schimel** is a Senior Research Scientist at the Jet Propulsion Laboratory, California Institute of Technology; leading research focused on interactions between climate and ecosystems, combining models and observations. His team is working on satellite missions that involve spectroscopic detection of carbon cycle dynamics and biodiversity on Earth.

**Fabian Schneider** is a postdoc at the NASA Jet Propulsion Laboratory, California Institute of Technology. His research interests lie in remote sensing of plant functional traits, biodiversity, and ecosystem functioning to improve our understanding of the interactions between plant communities and global change.

**Franziska Schrodt** is an Assistant Professor in the School of Geography, University of Nottingham (UK). She combines ecological data on (functional, phylogenetic, and taxonomic) biodiversity and geodiversity with novel data sources (e.g., remotely sensed) and analysis techniques (e.g., machine learning) to further develop our understanding of ecosystem dynamics and potential impacts on human well-being.

**Karsten Schulz** is Professor of Hydrology and Integrated Water Management at the University of Natural Resources and Life Sciences Vienna, Austria. By integrating information from various remote sensing platforms, his research aims to improve the prediction and understanding of water, energy, and solute fluxes in the soilplant-atmosphere continuum at different spatial scales.

**Anna K. Schweiger** is a Postdoctoral Associate at the Institut de recherche en biologie végétale, Université de Montréal. Anna is currently particularly interested in developing theory and methods for using spectra of plants and spectral diversity of landscapes to study plant community composition and the ecosystem benefits of biodiversity.

**Shawn P. Serbin** is a Scientist in the Environmental and Climate Sciences Department at Brookhaven National Laboratory, Upton, New York. Shawn's research aims to improve the understanding of the processes driving the fluxes and pools of carbon, water, and energy in terrestrial ecosystems and utilizes a range of tools including remote sensing to develop novel scaling methods and model-data synthesis approaches.

**Philip Townsend** is a Distinguished Professor in the Department of Forest and Wildlife Ecology at the University of Wisconsin, Madison. His research focuses include physiological remote sensing, imaging spectroscopy, forest ecosystem ecology, and watershed hydrology.

**Woody Turner** has spent a career promoting the use of satellite imagery to answer questions about the distribution and abundance of life on Earth and providing natural resource managers with remote sensing solutions to their day-to-day challenges.

**Susan Ustin** is a Distinguished Professor in the Department of Land, Air and Water Resources at the University of California Davis. Her research focuses include remote sensing of plant functional properties, ecosystem processes, imaging spectroscopy, and mapping of plant species and biodiversity.

**Ran Wang** is a Research Assistant Professor at the University of Nebraska – Lincoln, and visiting scientist at University of Alberta, where he received his PhD. He uses proximal, airborne, and satellite remote sensing to study vegetation biodiversity and productivity.

**Chris Williams** is the Geospatial Analysis Lead at the British Geological Survey. His background is in glaciology, terrain and environmental data analysis, and geoenvironmental product development.

**Adam Wilson** is an Assistant Professor in the Geography Department and the Department of Environment and Sustainability at the University at Buffalo. His research investigates the spatial patterns and processes of biodiversity and ecosystem function using remote sensing and field observations together with mechanistic and statistical modeling to understand how ecosystems change through space and time.

**Phoebe Zarnetske** is an Associate Professor in the Department of Integrative Biology at Michigan State University. Her research uses a combination of observational data, experiments, and modeling to connect observed patterns of biodiversity and community composition with underlying mechanisms.

**Brian Zutta** is a researcher at Universidad Alas Peruanas and an assistant project scientist at the University of California Los Angeles. He recently coordinated the development of the national measurement, reporting and verification and forest monitoring system for the National Forest Conservation Program (PNCB) of the Ministry of the Environment (MINAM) in Lima, Peru.

# **About the Editors**

**Jeannine Cavender-Bares** is a Distinguished McKnight University Professor in the Department of Ecology, Evolution and Behavior at the University of Minnesota. She earned a Master's at the Yale School of Forestry and Environmental Studies and a PhD in Biology at Harvard University. Her research focuses on the ecology and evolution of plant function, applying phylogenetics and spectral data to community ecology, and remote sensing of biodiversity. She is committed to advancing international efforts for global monitoring and assessment of biodiversity and ecosystem services to aid management efforts towards sustainability.

**John A. Gamon** is a Professor in the Departments of Earth and Atmospheric Sciences and Biological Sciences at the University of Alberta. He also conducts research in Quantitative Remote Sensing at the Center for Advanced Land Management Information Technologies (CALMIT) in the School of Natural Resources at the University of Nebraska, Lincoln. He earned his Master's and his PhD in Botany at the University of California, Davis, and did his postdoctoral research on Remote Sensing and Ecophysiology at the Carnegie Institution in Stanford, CA. His research focuses include photosynthesis, ecosystem function, productivity, biodiversity, ecoinformatics, and sustainability.

**Philip A. Townsend** is a Distinguished Professor in the Department of Forest and Wildlife Ecology at the University of Wisconsin, Madison. He earned his PhD in Geography at the University of North Carolina, Chapel Hill. His research focuses include physiological remote sensing, imaging spectroscopy, ecosystem ecology, and watershed hydrology.

# **Chapter 1 The Use of Remote Sensing to Enhance Biodiversity Monitoring and Detection: A Critical Challenge for the Twenty-First Century**

**Jeannine Cavender-Bares, John A. Gamon, and Philip A. Townsend**

# **1.1 Introduction**

Improved detection and monitoring of biodiversity is critical at a time when Earth's biodiversity loss due to human activities is accelerating at an unprecedented rate. We face the largest loss of biodiversity in human history, a loss which has been called the "sixth mass extinction" (Leakey 1996; Kolbert 2014), given that its magnitude is in proportion to past extinction episodes in Earth history detectable from the fossil record. International efforts to conserve biodiversity (United Nations 2011) and to develop an assessment process to document changes in the status and trends of biodiversity globally through the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (Díaz et al. 2015) have raised awareness about the critical need for continuous monitoring of biodiversity at multiple spatial scales across the globe. Biodiversity itself—the variation in life found among ecosystems and organisms at any level of biological organization—cannot practically be observed everywhere. However, if habitats, functional traits, trait diversity, and the spatial turnover of plant functions can be remotely sensed, the potential exists to globally inventory the diversity of habitats and traits associated with terrestrial biodiversity. To face this challenge, there have been recent calls for

J. Cavender-Bares (\*)

Department of Ecology, Evolution & Behavior, University of Minnesota, Saint Paul, MN, USA e-mail<EMAIL_ADDRESS>
J. A. Gamon Department of Earth & Atmospheric Sciences, University of Alberta, Edmonton, AB, Canada

Department of Biological Sciences, University of Alberta, Edmonton, AB, Canada

CALMIT, School of Natural Resources, University of Nebraska – Lincoln, Lincoln, NE, USA

P. A. Townsend

Department of Forest and Wildlife Ecology, University of Wisconsin, Madison, WI, USA

a global biodiversity monitoring system (Jetz et al. 2016; Proença et al. 2017; The National Academy of Sciences 2017). A central theme of this volume is that remote sensing (RS) will play a key role in such a system.

## **1.2 Why a Focus on Plant Diversity?**

 Plants—collectively called vegetation—regulate the flow of critical biogeochemical cycles, including those for water, carbon, and nitrogen. They affect soil chemistry and other properties, which in turn affect the productivity and structure of ecosystems. Given the importance of plant diversity for providing the ecosystem services on which humans depend—including food production and the regulating services that maintain clean air and freshwater supply (Millennium Ecosystem Assessment 2005; IPBES 2018a)—it is critical that we monitor and understand plant biodiversity from local to global scales, encompassing genetic variation within and among species to the entire plant tree of life (Cavender-Bares et al. 2017; Jetz et al. 2016; Turner 2014).

Of the 340,000 known seed plants on Earth and the 60,000 known tree species (Beech et al. 2017), 1 out of every 5 seed plants and 1 out of every 6 tree species are threatened (Kew Royal Botanic Gardens 2016). Vulnerability to threats ranging from climate change to disease varies among species and lineages because of evolved differences in physiology and spatial proximity to threats. Across all continents, the largest threat to terrestrial biodiversity is land use change due to farming and forestry, while climate change, fragmentation, and disease loom as everincreasing threats (IPBES 2018a; b; c; d). Many plant species are at risk for extinction due to a combination of global change factors, including drought stress, exotic species, pathogens, land use change, altered disturbance regimes, application of chemicals, and overexploitation.

## **1.3 The Promise of Remote Sensing to Detect Plant Diversity**

Different plants have evolved to synthesize different mixes of chemical compounds arranged in contrasting anatomical forms to support survival and growth. In addition, the structures of plant canopies correspond to different growth strategies in response to climate, environment, or disturbance. Differences among individual plants, populations, and lineages result from contrasting evolutionary histories, genetic backgrounds, and environmental conditions. Because these differences are readily expressed in aboveground physiology, biochemistry, and structure, many of these properties can be detected using spectral reflectance from leaves and plant canopies (Fig. 1.1). Plant pigments absorb strongly electromagnetic radiation in the

**Fig. 1.1** (**a**) The chemical, structural, and anatomical attributes of plants influence the way they interact with electromagnetic energy to generate spectral reflectance profiles that reveal information about plant function and are tightly coupled to their evolutionary origins in the tree of life. (Adapted from Cavender-Bares et al. 2017.) Imaging spectroscopy offers the potential to remotely detect patterns in diversity and chemical composition and vegetation structure that inform our understanding of ecological processes and ecosystem functions. Examples are shown from the Cedar Creek Ecosystem Science Reserve long-term biodiversity experiment. (**b**) The image cube (0.5 m × 1 m) at 1 mm spatial resolution (400–1000 nm) detects sparse vegetation early in the season in which individual plants can be identified. The "Z-dimension" (spectral dimension) illustrates different spectral reflectance properties for different scene elements, including different species. At this spatial resolution, plant diversity is predicted from remotely sensed spectral diversity (Wang et al. 2018). (**c**) AVIRIS NextGen false color image of the full experiment at 1 m spatial resolution (400–2500 nm). Each square is a 9 × 9 m plot with a different plant composition and species richness. Wang et al. (2019) mapped chemical composition and a suite of other functional traits and their uncertainties in all of the experimental plots. By combining spectral data at different scales, proximal and remote imagery can be used to examine the scale dependence of the spectral diversity–biodiversity relationship in detail (e.g., Wang et al. 2018; Gamon et al. Chap. 16)

visible wavelengths (400–700 nm), while other chemical compounds and structural attributes of plants that tend to be conserved through evolutionary history affect longer wavelengths. The patterns of light absorbed, transmitted, and reflected at different wavelengths from vegetation reveal leaf and canopy surface properties, tissue chemistry, and anatomical structures and morphological attributes of leaves, whole plants, and canopies. Thus, technological advances for assessing optical properties of plants provide profound opportunities for detecting functional traits of organisms at different levels of biological organization. These advances are occurring at multiple spatial scales, with technologies ranging from field spectrometers and airborne systems to emerging satellite systems. As a consequence, there is high potential to detect and monitor plant diversity—and other forms of diversity—across a range of spatial scales, and to do so iteratively and continuously, particularly if multiple methods can be properly coordinated.

Calls for a global biodiversity observatory (Fig. 1.2) that can detect and monitor functional plant diversity from space (Jetz et al. 2016; Proença et al. 2017; The National Academy of Sciences 2017; Geller et al., Chap. 20) have been met with widespread support. Forthcoming satellite missions, including the Surface Biology and Geology (SBG) mission in planning stages at the US National Aeronautics and Space Administration (NASA) and related missions in Europe and Japan (Schimel et al., Chap. 19), will make unprecedented spectroscopic data available to scientists, management communities, and decision-makers, but at relatively coarse spatial scales. At the same time, rapid progress is being made with field spectroscopy using unmanned aerial vehicles (UAVs) and other airborne platforms that are offering novel ways to use RS to advance our understanding of the linkages between optical (e.g., spectral or structural) diversity and multiple dimensions of biodiversity (e.g., species, functional, and phylogenetic diversity) at finer spatial scales (Fig. 1.1). These advances present a timely and tremendously important opportunity to detect changes in the Earth's biodiversity over large regions of the planet. One can fairly ask whether user communities are ready to make use of the data. Effective interpretation and application of remotely sensed data to determine the status and trends of plant biodiversity and plant functions across the tree of life—with linkages to all other living organisms—requires integration across vastly different knowledge arenas. Critically, it requires integration with in-situ direct and indirect measures of species distributions, their evolutionary relationships, and their functions. Approaches for integration are the primary focus of this book.

A central requirement to advance monitoring of biodiversity at the global scale is to decipher the sources of variation that contribute to spectral variation, both from a biological perspective and from a physical perspective. Distinct fields of biology have developed a range of methodologies for understanding plant ecological and evolutionary processes that underlie these sources of variation. Similarly, radiative transfer models have been developed largely based on the physics of light interacting with vegetation canopy elements and the atmosphere. These models have yet to capture the full range of plant traits, often preferring to represent "average" vegetation conditions for a region instead of the variation present, so are not yet ready for the task. All of these methods have unique approaches to analyzing complex, multidimensional data sets, and neither the analytical approaches nor the data structures have been brought together in a systematic or comprehensive manner. A common language among disciplines (including biology, physiological ecology, landscape ecology, genetics, phylogenetics, geography, spectroscopy, and radiative transfer) related to the RS of biodiversity is currently lacking. This book provides a framework for how biodiversity, focusing particularly on plants, can be detected using proximally and remotely sensed hyperspectral data (with many contiguous spectral bands) and other tools, such as lidar (with its ability to detect structure). The chapters in this book present a range of perspectives and approaches on how RS can be

**Fig. 1.2** An envisioned global biodiversity observatory in which remotely sensed high spectral resolution spectroscopic data from satellites is integrated with biodiversity observations through natural history studies, phylogenetic systematics, functional trait measurements, and species distribution data. The figure is adapted from Jetz et al. 2016 based on the National Center for Ecological Analysis and Synthesis Working Group "Prospects and priorities for satellite monitoring of global terrestrial biodiversity"

integrated to detect and monitor the status and trends of plant diversity, as well the biodiversity of other organisms and life processes that depend on plants above- and belowground. Biological and computational experts from three disciplines—RS and leaf optics, plant functional biology, and systematics—present insights to advance our understanding of how to link spectral and other kinds of RS data with functional traits, species distributions, and the tree of life for biodiversity detection. The authors detail the approaches and conditions under which efforts to detect plant biodiversity are likely to succeed, being explicit about the advantages and disadvantages of each. A theme running through many chapters is the challenge of moving across spatial scales from the leaf level to the canopy, ecosystem, and global scale. We provide a glossary that allows a common language across disciplines to emerge.

Here we explore the prospects for integrating components from each of these fields to remotely detect biodiversity and articulate the major challenges in our ability to directly link spectral data of vegetation to species diversity, functional traits, phylogenetic information, and functional biodiversity at the global scale. RS offers the potential to fill in data gaps in biodiversity knowledge locally and globally, particularly in remote and difficult-to-access locations, and can help define the larger spatial and temporal background needed for more focused and effective local or regional studies. It also may increase the likelihood of capturing temporal variation, and it allows monitoring of biodiversity at different spatial scales with different platforms and approaches. In essence, it provides the context within which changing global biodiversity patterns can be understood. The concept of "optical surrogacy" (Magurran 2013)—in which the linkage of spectral measurements to associated patterns and processes is used—may be useful in predicting ecosystem processes and characteristics that themselves are not directly observable (Gamon 2008; Madritch et al. 2014; Fig. 1.3). In a broad sense, such relationships between various expressions of biodiversity and optical (spectral) diversity provide a fundamental principle for "why RS works" as a metric of biodiversity and why so many different methods at different scales can provide useful information.

## **1.4 The Contents of the Book**

The first section of the book presents the potential and basis for direct and indirect remote detection of biodiversity.

Cavender-Bares et al. (Chap. 2) present an overview of biodiversity itself, including the in-situ methods and metrics for measuring biodiversity, particularly plant diversity. The chapter provides a layperson's overview of the elements, methods, and metrics for detecting and analyzing biodiversity and points to the potential of spectral data, collected at multiple spatial and biological scales, to enhance the study of biodiversity. In doing so, it bridges an ecological and evolutionary understanding of the diversity of life, considering both its origins and consequences.

Serbin and Townsend (Chap. 3) describe various approaches for measuring plant and ecosystem function using spectroscopy, providing both the historical development of past advances and the potential of these approaches looking forward. The chapter explains why we are able to retrieve functional traits from spectra, which traits can be retrieved, and where spectra show features important for different aspects of plant function. It also raises the challenge of scaling plant function from leaves to canopies and landscapes.

**Fig. 1.3** Optical methods for detecting the functional, structural, and chemical components of vegetation, which are tightly coupled to the genetic and phylogenetic backgrounds of plants, are linked to belowground processes and the structure and function of microbial communities

Morsdorf et al. (Chap. 4) then present the Laegeren forest site in Switzerland as a virtual laboratory. They demonstrate how spectroscopy can be operationalized for RS of functional diversity to explain plant biodiversity patterns and ecosystem functions. The Laegeren site is one of the best-studied sites in the world for this purpose and is used as a case study to explain ground truthing and what can be learned from landscape-level detection of functional diversity.

Martin (Chap. 5) summarize the experiences with "spectronomics"—a framework aimed at integrating chemical, phylogenetic, and spectral RS data—using airborne imagery to detect forest composition and function in wet tropical forests. In these vast, largely inaccessible landscapes that harbor enormous taxonomic variation, approaches that rely solely on field-based observations are infeasible, illustrating an essential role for RS. As pioneers in using spectroscopy to detect plant chemistry, function, and biodiversity in tropical forests around the world, these researchers highlight some of the major lessons they have learned.

Pontius et al. (Chap. 6) consider how biodiversity can be protected given current threats to forest and vegetation conditions and present approaches for detailed and accurate detection of forest disturbance and decline. They review current techniques used to assess and monitor forest ecosystem condition and disturbance and outline a general approach for earlier, more detailed and accurate decline assessment. They also discuss the importance of engaging land managers, practitioners, and decisionmakers in these efforts to ensure that the products developed can be utilized by stakeholders to maximize their impact.

Meireles et al. (Chap. 7) provide a framework to explain how spectral reflectance data from plants is tightly coupled to the tree of life and demonstrate how spectra can reveal evolutionary processes in plants. They clarify that many spectral features in plants are inherited and are thus very similar among close relatives—in other words, they are highly phylogenetically conserved. Simulations reveal that spectral information of plants appears to follow widely used evolutionary models, making it possible to link plant spectra to the tree of life in a predictive manner. As a consequence, methods developed in evolutionary biology to understand the tree of life can now benefit the RS community. The chapter provides evidence that evolutionary lineages may be easier than individual species to detect through RS methods, particularly if they are combined with other approaches for estimating which species and lineages have the potential to be present in a given location. A caveat is that spatial resolution of satellite spectral data will limit such inferences, but leaf- and canopy-level spectra (obtainable from proximal and airborne sensing) can contribute enormously to our understanding of these fundamental links between spectral patterns and gene sequences.

Madritch et al. (Chap. 8) link aboveground plant biodiversity and productivity to belowground processes. They explain the functional mechanisms—which can be revealed by remotely sensed spectral data—that influence interactions of plant hosts with insects and soil organisms, in turn influencing ecosystem functions, such as decomposition and nutrient cycling. The chapter provides an example of using the concept of surrogacy, in which the biochemical linkage of spectral measurements to associated patterns and processes aboveground is used to provide estimates of soil and microbial processes belowground that are not directly observable via RS.

The next three chapters focus on linking satellite-based remotely sensed data to biodiversity prediction. Pinto-Ledezma and Cavender-Bares (Chap. 9) present an example of how currently available satellite-based RS products can be used to generate next-generation species distribution models to predict where species and lineages are likely occur and the habitats they may have access to and persist in under altered climates in the future. They compare RS-based methods for generating predictive models with widely used approaches that use meteorologically derived climate variables. They demonstrate the advantages of RS-based models in regions where meteorological data is only sparsely available. Such predictive modeling that harnesses species occurrence data and temporal information about the biotic environment may make spectral methods of species and evolutionary lineage detection more tractable.

Building on the availability of satellite RS data with near-global coverage to predict biodiversity, Record et al. (Chap. 10) explore how RS illuminates the relationship between biodiversity and geodiversity—the variety of abiotic features and processes that provide the template for the development of biodiversity. They introduce a variety of globally available geodiversity measures and examine how they can be combined with biodiversity data to understand how biodiversity responds to geodiversity. The authors use the analogy of the "stage" that defines the patterns of life to some

degree, often measured as habitat heterogeneity, a key driver of species diversity. They illustrate the approach by examining the relationship between biodiversity and geodiversity with tree biodiversity data from the US Forest Inventory and Analysis Program and geodiversity data from remotely sensed elevation from the Shuttle Radar Topography Mission (SRTM). In doing so, they outline the challenges and opportunities for using RS to link biodiversity to geodiversity.

Paz et al. (Chap. 11) present an approach for using RS data to predict patterns of plant diversity and endemism in the tropics within the Brazilian Atlantic rainforest. They examine how RS environmental data from tropical regions can be used to support biodiversity prediction at multiple spatial, temporal, and taxonomic scales.

Bolch et al. (Chap. 12) summarize the range of approaches that can be used to optimize detection of invasive alien species (IAS), which pose severe threats to biodiversity. These approaches emphasize the ability to detect individual plant species that have distinct functional properties. The chapter presents current RS capabilities to detect and track invasive plant species across terrestrial, riparian, aquatic, and human-modified ecosystems. Each of these systems has a unique set of issues and species assemblages with its own detection requirements. The authors examine how RS data collection in the spectral, spatial, and temporal domains can be optimized for a particular invasive species based on the ecosystem type and image analysis approach. RS approaches are enhancing studies of the invasion processes and enabling managers to monitor invasions and predict the spread of IAS.

The next three chapters of the book explore how components of diversity can be detected spectrally and remotely with a focus on optical detection methods and technical challenges. Lausch et al. (Chap. 13) delve into the complexity of monitoring vegetation diversity and explain how no single monitoring approach is sufficient on its own. The chapter introduces the range of Earth observation (EO) techniques available for assessing vegetation diversity, covering close-range EO platforms, spectral approaches, plant phenomics facilities, ecotrons, wireless sensor networks (WSNs), towers, air- and spaceborne EO platforms, UAVs, and approaches that integrate air- and spaceborne EO data. The chapter presents the challenges with these approaches and concludes with recommendations and future directions for monitoring vegetation diversity using RS.

Ustin and Jacquemoud (Chap. 14) provide the physical basis for detecting the optical properties of leaves based on how they modify the absorption and scattering of energy to reveal variation in function. The chapter provides considerable detail on how the combination of absorption and scattering properties of leaves together creates the shape of their reflectance spectrum. It also reviews and summarizes the most common interactions between leaf properties and light and the physical processes that regulate the outcomes of these interactions.

Schweiger (Chap. 15) describes a set of best practices for planning field campaigns and collecting and processing data, focusing on spectral data of terrestrial plants collected across various levels of measurements, from leaf to canopy to airborne. These approaches also generally apply to RS of aquatic systems, soil, and the atmosphere and to active RS systems, such as lidar, thermal, and satellite data collection. Schweiger discusses how goals for data collection can be broadly classified into model calibration, model validation, and model interpretation.

The final chapters of the book move to the issues of temporal and spectral scale and integration across scales. Gamon et al. (Chap. 16) present a thorough examination of the challenges in spectral methods for detecting biodiversity posed by issues of spatial, temporal, and spectral dimensions of scale. They explain why the size of the organism relative to the pixel size of detection has consequences for spectral detection of different components of biodiversity and draw on a rich history of literature on scaling effects, including geostatistical approaches for sampling across spatial scales. The chapter emphasizes the importance of developing biodiversity monitoring systems that are "scale-aware" as well as the value of an integrated, multi-scale sampling approach.

Schrodt et al. (Chap. 17) outline how environmental and socioeconomic data can be integrated with biodiversity and RS data to expand knowledge of ecosystem functioning and inform biodiversity conservation decisions. They present the concepts, data, and methods necessary to assess plant species and ecosystem properties across spatial and temporal scales and provide a critical discussion of the major challenges.

Fernández et al. (Chap. 18) provide a framework for understanding Essential Biodiversity Variables (EBVs) to integrate in-situ biodiversity observations and RS through modeling. They argue that open and reproducible workflows for data integration are critical to ensure traceability and reproducibility to allow each EBV to be updated as new data and observation systems become available. The chapter makes the case that the development of a globally coordinated system for biodiversity monitoring will require the mobilization of and integration of in-situ biodiversity data not yet publicly available with emerging RS technologies, novel biodiversity models, and informatics infrastructures.

Schimel et al. (Chap. 19) discuss the prospects and pitfalls for RS of biodiversity at the global scale, focusing on imaging spectroscopy and NASA's Surface Biology and Geology mission concept.

Finally, Geller et al. (Chap. 20) provide an epilogue to the book and present a vision for a global biodiversity monitoring system that is flexible and accessible to a range of user communities. Such a system will require a coordinated effort among space agencies, the RS community, and biologists to bring information about the status and trends in biodiversity, ecosystem functions, and ecosystem services together so that different data streams inform each other and can be integrated. The chapter explains that the Group on Earth Observations Biodiversity Observation Network (GEO BON), the International Long Term Ecological Research Site (ILTER) network, the US National Ecological Observatory Network (NEON), and a variety of sponsors and other organizations are working to enhance coordination and to develop guidelines and standards that will serve this vision.

Indeed, a rapidly advancing global movement has emerged with a shared vision to develop the capacity to monitor the status and trends in the Earth's biodiversity. The authors of this book have sought to contribute to that shared vision through their varied perspectives and experiences. Collectively, the chapters present a range of approaches and knowledge that can transform the ability of humanity to detect and interpret the changing functional biodiversity of planet Earth.

# **1.5 The Origins of the Book**

Before closing, we offer a note of acknowledgment on how this book came into existence. The editors, who themselves have contrasting backgrounds spanning several disciplines, were collaboratively funded, starting in 2013, by the US National Science Foundation (NSF) and the NASA Dimensions of Biodiversity program on the project *Linking remotely sensed optical diversity to genetic, phylogenetic and functional diversity to predict ecosystem processes* (DEB-1342872,1342778). We worked together in several field sites for 5 years to advance our own understanding approaches for remote detection of plant biodiversity. The importance and high potential for rapid advances, as well as the need for the involvement of numerous experts, were obvious from the start. With support from the National Institute for Mathematical and Biological Synthesis (NIMBioS) for the working group on Remote Sensing of Biodiversity, we brought together many of the experts represented within the book, including a symposium in the fall of 2018 where authors shared their work and provided feedback to each other. Further interactions were fostered by the National Center for Ecological Analysis and Synthesis (NCEAS), annual meetings of the NASA Biological Diversity and Ecological Forecasting Program, the Keck Institute for Space Studies, the NSF Research Coordination Network on Biodiversity across scales, and bioDISCOVERY, an international research program fostering collaborative interdisciplinary activities on biodiversity and ecosystem science. bioDISCOV-ERY, which is part of Future Earth and is hosted and supported by the University Zürich's University Research Priority Program on Global Change and Biodiversity, provided generous financial support for the editing process. Additional financial support was provided by the NSF RCN (DEB: 1745562) and the Keck Institute for Space Studies. Mary Hoff served as technical editor for all of the chapters in the book. We express gratitude to all of these institutions and our many colleagues who contributed to the conception of this work along the way.

## **References**

**Open Access** This chapter is licensed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license and indicate if changes were made.

The images or other third party material in this chapter are included in the chapter's Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the chapter's Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder.

# **Chapter 2 Applying Remote Sensing to Biodiversity Science**

**Jeannine Cavender-Bares, Anna K. Schweiger, Jesús N. Pinto-Ledezma, and Jose Eduardo Meireles**

A treatment of the topic of biodiversity requires consideration of what biodiversity is, how it arises, what drives its current patterns at multiple scales, how it can be measured, and its consequences for ecosystems. Biodiversity science, by virtue of its nature and its importance for humanity, intersects evolution, ecology, conservation biology, economics, and sustainability science. These realms then provide a basis for discussion of how remote detection of biodiversity can advance our understanding of the many ways in which biodiversity is studied and impacts humanity. We start with a discussion of how biodiversity has been defined and the ways it has been quantified. We briefly discuss the nature and patterns of biodiversity and some of the metrics for describing biodiversity, including remotely sensed spectral diversity. We discuss how the historical environmental context at the time lineages evolved has left "evolutionary legacy effects" that link Earth history to the current functions of plants. We end by considering how remote sensing (RS) can inform our understanding of the relationships among ecosystem services and the trade-offs that are often found between biodiversity and provisioning ecosystem services.

Institut de recherche en biologie végétale, Université de Montréal, Montréal, QC, Canada

J. E. Meireles Department of Ecology, Evolution and Behavior, University of Minnesota, Saint Paul, MN, USA

School of Biology and Ecology, University of Maine, Orono, ME, USA

J. Cavender-Bares (\*) · J. N. Pinto-Ledezma Department of Ecology, Evolution and Behavior, University of Minnesota, Saint Paul, MN, USA e-mail<EMAIL_ADDRESS>
A. K. Schweiger Department of Ecology, Evolution and Behavior, University of Minnesota, Saint Paul, MN, USA

## **2.1 What Is Biodiversity?**

Biodiversity encompasses the totality of variation in life on Earth, including its ecosystems, the species generated through evolutionary history across the tree of life, the genetic variation within them, and the vast variety of functions that each organism, species, and ecosystem possess to access and create resources for life to persist. Changes in the Earth's condition, including the actions of humanity, have consequences for the expression of biodiversity and how it is changing through time.

## **2.2 The Hierarchical Nature of Biodiversity**

Since Darwin (1859), we have understood that biodiversity is generated by a process of descent with modification from common ancestors. As a result, biological diversity is organized in a nested hierarchy that recounts the branching history of species (Fig. 2.1a). Individual organisms are nested within populations, which are nested within species and within increasingly deeper clades. This hierarchy ultimately represents the degree to which species are related to each other and often conveys when in time lineages split (Fig. 2.1a).

Evolution results in the accumulation of changes in traits that causes lineages to differ. The degree of trait divergence between taxa is expected to be proportional to the amount of time they have diverged from a common ancestor. As a consequence, distantly related taxa are expected to be phenotypically more dissimilar (Fig. 2.1b).

**Fig. 2.1** (**a**) The hierarchical organization of biodiversity. Species (triangles) are nested within phylogenetic lineages (clades) due to shared ancestry. All species within a lineage have common ancestor (filled circles). (**b**) Differences in the phenotypes (or trait values) of species (triangles) tend to increase with time since divergence from a common ancestor, shown by the orange circle. The divergence points at which species split are shown in the simulation, and the filled circles indicate the common ancestor of each lineage

Since spectral signals are integrated measures of phenotype, spectra should be more dissimilar among distantly related groups than among close relatives (Cavender-Bares et al. 2016b; McManus et al. 2016; Schweiger et al. 2018). This expectation can be seen from models of evolution in which traits change over time following a random walk (Brownian motion process; Fig. 2.1b) (O'Meara et al. 2006; Meireles et al., Chap. 7). In cases of convergent evolution—where natural selection causes distant relatives to evolve similar functions in similar environments—however, phenotypes can be more similar than expected under Brownian motion.

This hierarchy of life is relevant to RS of plant diversity because certain depths of the tree of life may be more accurately detected than others at different spatial resolutions and geographic regions. For example, it could be easier to detect deeper levels in the hierarchy (such as genera or families) in hyper-diverse communities than in shallow levels (such as species) because deep splits tend to have greater trait divergence. Meireles et al. (Chap. 7) further explain why and how phylogenetic information can be leveraged to detect plant diversity.

# **2.3 The Making of a Phenotype: Phylogeny, Genes, and the Environment**

The phenotype of an organism is the totality of its attributes, and it is quantified in terms of its myriad functions and traits. The phenotype of an organism is a product of the interaction between the information encoded in its genes—the genotype and the environment over the course of development. Understanding the relative influence of gene combinations, environmental conditions, and ontogenetic stage is an active area of investigation across different disciplines (Diggle 1994; Sultan 2000; Des Marais et al. 2013; Palacio-López et al. 2015).

Although genotypes often play a critical role in determining phenotypic outcomes, many processes can result in mismatches between genotype and phenotype. One of the most well documented of these processes is known as phenotypic plasticity—when organisms with the same genotype display different phenotypes, usually in response to different environmental conditions (Bradshaw 1965; Scheiner 1993; Des Marais et al. 2013). Plasticity can also result in distinct genotypes developing similar phenotypes when growing under the same environmental conditions.

A similar story can be told about the relationship of phenotypic similarity and phylogenetic relatedness. As we have seen earlier, closely related taxa are expected to be more similar to each other than distantly related taxa. However, convergent evolution can lead to plants from different branches of the tree of life to evolve very similar traits—such as succulents, which are found within both euphorbia and very distantly related cacti taxa.

The fact that phenotypes can be, but not necessarily are, directly related to specific genotypes and phylogenetic history should be considered when remotely sensing biodiversity. Only phenotypes can be remotely sensed directly. Genetic and phylogenetic information can only be inferred from spectra to the degree that absorption features of plant chemical or structural characteristics at specific wavelengths relate to phenotypic information. However, the effects plant traits have on spectra are only partially understood. Identifying the regions of the spectrum that are influenced by specific traits is complicated by overlapping absorption features and subtle differences in plant chemical, structural, morphological, and anatomical characteristics that simultaneously influence the shape of the spectral response (Ustin and Jacquemoud, Chap. 14).

## **2.4 Patterns in Plant Diversity**

One of the most intensively studied patterns in biodiversity is the latitudinal gradient, in which low-latitude tropical regions harbor more species, genera, and families of organisms than high-latitude regions. In particular, wet tropical areas tend to reveal higher diversity of organisms than colder and drier climates (Fig. 2.2). Humboldt (1817) documented these patterns quite clearly for plant diversity. Naturalists since then have sought to explain these patterns.

Tropical biomes have existed longer than more recent biomes, such as deserts, Mediterranean climates, and tundra, which expanded as the climate began to cool some 35 million years ago. Tropical biomes also cover more land surface area than other biomes. Tropical species thus have had more time and area (integrated over the time since their first appearance) for species to evolve and maintain viable populations (Fine and Ree 2006). Lineages that originally evolved in the tropics may also have been less able to disperse out of the tropics and to evolve new attributes adapted to cold or dry climates—due to phylogenetic conservatism—restricting their ability to diversify (Wiens and Donoghue 2004). However, not all lineages follow this latitudinal gradient. Ectomycorrhizal fungi, for example, show higher diversity at temperate latitudes, where they likely have higher tree host density (Tedersoo and Nara 2010). Moreover, other measures of diversity do not necessarily follow these patterns. Variation in functional attributes of species, for example, follow different patterns depending on the trait (Cavender-Bares et al. 2018; Echeverría-Londoño et al. 2018; Pinto-Ledezma et al. 2018b). Specific leaf area, one of the functional traits that is highly aligned with the leaf economic spectrum (discussed below), shows higher variation at high latitudes than low latitudes across the Americas. In contrast, seed size shows higher variation at low latitudes (Fig. 2.2b).

At regional scales, variation in the environment, as discussed by Record et al. (Chap. 10), sets the stage for variation in biodiversity because species have evolved to inhabit and can adapt to different environments, which allows them to partition resources and occupy different niches created by environmental variation. Thus, habitat diversity begets biodiversity, and remotely sensed measures of environmental variation have long been known to predict biodiversity patterns (Kerr et al. 2001).

Land area is another long-observed predictor of species diversity, first described for species within certain guilds on island archipelagoes (Diamond and Mayr 1976).

**Fig. 2.2** Map of the Americas showing plant species richness, phylogenetic diversity, and functional diversity. Species richness and phylogenetic and functional diversities were estimated based on available information from the Botanical Information and Ecology Network (BIEN) database (Enquist et al. 2016, https://peerj.com/preprints/2615/). Distribution of functional diversity (trait mean) for three functional traits (**d**–**f**) was log-transformed for plotting purposes. (**a**) Species richness; (**b**) phylogenetic diversity; (**c**) first principal component of functional trait means; (**d**) specific leaf area (mm2 /mg); (**e**) plant height (m); and (**f**) seed mass (mg). Diversity metrics were calculated from an estimated presence-absence matrix (PAM) for all vascular plant species at 1 degree spatial resolution (PAM dimension = 5353 pixels × 98,291 species) using range maps and predicted distributions. Functional diversity is based on the first principal component of a principal component analysis (PCA) of species means for the three functional traits

These observations led to the generalization that richness (number of species, *S*) increases with available land area (*A*), giving rise to the well-known species-area relationships, in which the log of species number increases linearly with the log of the area available:

$$
\log\left(S\right) = z \times \log\left(A\right) + c,\tag{2.1}
$$

or simply,

$$S = cA^{Z},\tag{2.2}$$

where *c* is the *y*-intercept of the log-log relationship and *z* is the slope.

# **2.5 Functional Traits, Community Assembly, and Evolutionary Legacy Effects on Ecosystems**

## *2.5.1 Functional Traits and the Leaf Economic Spectrum*

There is a long history of using functional traits to understand ecological processes, including the nature of species interactions, the assembly of species into ecological communities, and the resulting functions of ecosystems. Species with different functions are likely to have different performance in different environments and to use resources differently, allowing them to partition ecological niches. They are thus less likely to compete for the same resources, promoting their long-term coexistence. An increased focus on trait-based methodological approaches to understanding the relationship between species functional traits and the habitats or ecological niches was spurred by the formalization of the leaf economic spectrum (LES) (Wright et al. 2004). The LES shows that relationships exist among several key traits across a broad range of species and different climates (Reich et al. 1997; Wright et al. 2004) and that simple predictors, such as specific leaf area (SLA, or its reciprocal leaf mass per area, LMA) and leaf nitrogen content, represent a major axis of life history variation. This axis ranges from slow-growing ("conservative") species that tolerate low-resource environments to fast-growing ("acquisitive") species that perform well in high-resource environments (Reich 2014). Variations in relatively easy-to-measure plant traits are tightly coupled to hard-to-measure functions, such as leaf lifespan and growth rate, which reveal more about how a plant invests and allocates resources over time to survive in different kinds of environments. High correlations of functional traits provide strong evidence for trait coordination across the tree of life. The variation in plant function across all of its diversity is relatively constrained and can be explained by a few major axes of trait information (Díaz et al. 2015). Conveniently, traits such as SLA and N are readily detectable via spectroscopy. Other traits—such as leaf lifespan or photosynthetic rates—that are harder to measure but are correlated with these readily detectable traits can thus be inferred, permitting greater insight into ecological processes.

# *2.5.2 Plant Traits, Community Assembly, and Ecosystem Function*

Considerable evidence supports the perspective that plant traits influence how species sort along environmental gradients and are linked to abiotic environmental filters that prevent species without the appropriate traits from persisting in a given location. Traits thus influence the assembly of species in communities—and consequently, the composition, structure, and function of ecosystems. Variation in traits among individual plants and species within communities indicates differences in resource use strategies of plants, which have consequences for ecosystem functions, such as productivity and resistance to disturbance, disease, and extreme environmental conditions. Moreover, the distribution of plant traits within communities influences resource availability for other trophic levels, above- and belowground, which affects community structure and population dynamics in other trophic levels. A major goal of functional ecology is to develop predictive rules for the assembly of communities based on an understanding of which traits or trait combinations (e.g., the leaf-heightseed (LHS) plant ecology strategy, sensu Westoby 1998) are important in a given environment, how traits are distributed within and among species, and how those traits relate to mechanisms driving community dynamics and ecosystem function (Shipley et al. 2017). This predictive framework requires selecting relevant traits; describing trait variation and incorporating this variation into models; and scaling trait data to community- and ecosystem-level processes (Funk et al. 2017). Selecting functional traits for ecological studies is not trivial. Depending on the question, individual traits or trait combinations can be selected that contribute to a mechanistic understanding of the critical processes examined. One can distinguish *response traits*, which influence a species response to its environment, and *effect traits*, which influence ecosystem function (Lavorel and Garnier 2002). These may or may not be different traits. Disturbance or global change factors that influence whether a species can persist within a habitat or community based on its response traits may impact ecosystem functions in complex ways (Díaz et al. 2013). Plant traits are at the heart of understanding how the evolutionary past influences ongoing community assembly processes and ecosystem function (Fig. 2.3). Traits also influence species interactions, which contribute to continuing evolution. Remotely sensed plant traits, if detected and mapped (Serbin and Townsend, Chap. 3; Morsdorf et al., Chap. 4) at the appropriate pixel size and spatial extent (Gamon et al., Chap. 16), can provide a great deal of insight into these different processes (Fig. 2.4).

# *2.5.3 Phylogenetic, Functional, and Spectral Dispersion in Communities*

The rise of phylogenetics in community ecology was based on the idea that functional similarity due to shared ancestry should be predictive of environmental sorting and limiting similarity. These processes depend on physiological tolerances

**Fig. 2.3** Plant traits that have evolved over time influence how plants assemble into communities, which shapes ecosystem structure and function. Traits reflect biogeographic and environmental legacies and evolve in response to changing environments. They play a central role in ecological processes influencing the distribution of organisms and community assembly. A range of traits influence the way plants reflect light, such that many traits can be mapped continuously across large spatial extents with imaging spectroscopy. The remote detection of plant traits provides incredible potential to observe and understand patterns that reveal information about community assembly, changes in ecosystem function, and how legacies from the past shape community structure and ecosystem processes today. (Reprinted from Cavender-Bares et al. 2019, with permission)

in relation to environmental gradients and intensity of competition as a consequence of shared resource requirements (Webb 2000a, 2002). The underlying conceptual framework was formalized in terms of functional traits in individual case studies (Cavender-Bares et al. 2004; Verdu and Pausas 2007). The tendency to oversimplify the interpretation of phylogenetic patterns in communities, whereby phylogenetic overdispersion was equated with the outcome of competitive exclusion and phylogenetic clustering was interpreted as evidence for environmental sorting, led to a series of studies investigating the importance of scale (Cavender-Bares et al. 2006; Swenson et al. 2006) and the role of Janzen-Connell-type mechanisms, i.e., densitydependent mortality due to pathogens and predators (Gilbert and Webb 2007; Parker et al. 2015). Further developments revealed that the relationship between patterns and ecological processes is context-dependent—in particular, with respect to spatial scale (Emerson and Gillespie 2008; Cavender-Bares et al. 2009; Gerhold et al. 2015). Later studies revisited assumptions about the nature of competition and expected evolutionary and ecological outcomes (Mayfield and Levine 2010). Likewise, interpreting spectral dispersion will depend on the spatial resolution and pixel (grain) size of remotely sensed imagery relative to plant size (Marconi et al. 2019) as well as on the consideration of specific spectral regions and their functional importance. When traits and spectral regions are highly phylogenetically conserved (see Meireles et al., Chap. 7), trait, phylogenetic, and spectral data provide equivalent information. However, when some traits and spectral regions are

**Fig. 2.4** (**a**) Biological processes change with spatial and temporal scale as do the patterns they give rise to. (Adapted from Cavender-Bares et al. 2009.) Detection and interpretation of those patterns will shift with spatial resolution (pixel size) and extent (**b**–**e**). (**a**) At high spatial resolutions (1 cm pixel size)—that allow detection of individual herbaceous plants and their interactions—and relatively restricted spatial extents in which the abiotic environment is fairly homogeneous, spectral dissimilarity among pixels may indicate complementarity of contrasting functional types. (**b**) The grain size sufficient to detect species interactions is likely to shift with plant size. For example, the interactions of trees in the Minnesota oak savanna and their vulnerability to density-dependent diseases, such as oak wilt (*Bretziella fagacearum*), can be studied at a 1 m pixel size. (**c**) At somewhat larger spatial resolution (30 m pixel sizes) and extent, environmental sorting—which includes interactions of species with both the biotic and abiotic environments—may be detected by comparing spectral similarity of neighbors and comparing mapped functional traits to environmental variation. Images adapted from Singh et al. (2015). The ability to detect change through time may be especially important in understanding species interactions and ecological sorting processes in relation to the biotic and abiotic environment. (**d**) At the global scale, it may be possible to detect the evolutionary legacy effects. For instance, regions with similar climate and geology can differ in vegetation composition and ecosystem function as a consequence of differences in which lineages evolved in a given biogeographic region and their historical migration patterns. Shown are mapped values of %N and NPP based on Moderate Resolution Imaging Spectroradiometer (MODIS) data. (Adapted from Cavender-Bares et al. 2016a)

conserved, but others related to species interactions or with the abiotic environment vary considerably among close relatives, there is the potential to tease apart spectral signals that may relate to species interactions.

Spatial patterns of spectral similarity and dissimilarity also have the potential to provide meaningful information about ecological processes and the forces that dominate community assembly at a particular scale. For example, to the extent that spectral similarity of neighboring plants can be determined, high spectral similarity might indicate that functionally and/or phylogenetically similar individuals are sorting into the same environment, while spectral dissimilarity might indicate that quite distinct individuals are able to coexist if they exhibit complementarity by

**Fig. 2.5** Evolutionary legacy effects as a consequence of biogeographic origin. Two lineages are shown that have contrasting origins, one from the tropics and one from high latitudes. Both diversified and expanded to colonize intermediate latitudes such that their descendants sometimes cooccur. Lineages with ancestors from contrasting climatic environments likely differ in functional traits that reflect their origins and thus may assemble in contrasting microenvironments within the local communities where they co-occur. (Adapted from Cavender-Bares et al. 2016a)

partitioning resources. Approaches that use spectral detection of patterns that might be interpreted within this framework will need to pay close attention to the pixel-to-plant size ratio—or the grain size at which biological diversity varies (Gamon et al., Chap. 16; Serbin and Townsend, Chap. 3; Schimel et al., Chap. 19) as well as to the spatial extent at which density-dependent processes and environmental sorting pressures are strongest. Often these processes are expected to dominate at different spatial scales, such that competition and Janzen-Connell-type mechanisms operate at very local scales, while environmental sorting may be more important at landscape scales. Other factors, such as the geographic locations and environmental conditions under which lineages diversified, may impact spectral patterns of phylogenetic, functional trait, and spectral similarity at continental scales (Fig. 2.5). At the same time, spectral similarity will be driven by similar ecological forces, since both genetic and phylogenetic compositions, as well as environmental factors, drive phenotypic variation that can be spectrally detected.

## **2.6 Evolutionary Legacy Effects on Ecosystems**

Ecological communities are formed by resident species (incumbents) and colonizer species. Incumbents may have originated in the study region (or at least have had considerable time to adapt to their biotic and abiotic environment), whereas colonizers evolved elsewhere and subsequently dispersed into the region. However, the processes that determine species distributions and the assembly of ecological communities are complex. Species within communities experience unique combinations of evolutionary constraints and innovations due to legacies of their biogeographic origins and the environmental conditions in which they evolved (Cavender-Bares et al. 2016a; Pinto-Ledezma et al. 2018a). Historical contingencies play a role in which lineages can take advantage of opportunities to diversify following climate change or other disturbances and environmental transitions. The rate of species range expansion and contraction and the evolution of species functional traits that allow species to establish and persist in some regions or under particular environmental conditions but not elsewhere are shaped by biogeographic history (Moore et al. 2018). For example, when species from two distinct lineages—one that evolved in tropical climates and the other that evolved in temperate climates—colonize a new environment, they are predicted to persist in contrasting microhabitats as a consequence of niche conservatism (Ackerly 2003; Harrison 2010; Cavender-Bares et al. 2016a). These evolutionary legacies—collectively referred to as "historical factors" (Ricklefs and Schluter 1993)—operate at different spatial and temporal scales that leave their imprints on species current functional attributes and distributions and consequently on ecosystem function itself (Fig. 2.4c, Cavender-Bares et al. 2016a). RS approaches can help reveal how the deep past has influenced current biodiversity patterns and ecosystem function by decoupling climate and geological setting from ecosystem function. Current and forthcoming RS instruments (Lausch et al., Chap. 13; Schimel et al., Chap. 19) enable the monitoring of plant productivity, dynamics of vegetation growth, seasonal changes in chemical composition, and other ecosystem properties independently of climate and geology. These technologies thus provide opportunities to detect how biodiversity is sorted across the globe and to determine how variable ecosystem functions can be in the same geological and environmental setting. Both are important for developing robust predictive models of how lineages respond to current and future environmental conditions with important consequences for managing ecosystems in the Anthropocene.

## **2.7 Quantifying Multiple Dimensions of Biodiversity**

Several major dimensions of biodiversity have emerged in the literature that capture different aspects of the variation of life. *Taxonomic diversity* focuses on differences between species or between higher-order clades, such as genera or families. Estimating the numbers and/or abundances of different taxa across units of area captures this variation. *Phylogenetic diversity* captures the evolutionary distances between species or individuals, represented in terms of millions of years since divergence from a common ancestor or molecular distances based on accumulated mutations since divergence. *Functional diversity* focuses on the variation among species as a consequence of measured differences in their functional traits, frequently calculated as a multivariate metric but also calculated for individual trait variation. *Spectral diversity* captures the variability in spectral reflectance from vegetation (or from other surfaces), either measured and calculated among individual plants or, more commonly, calculated among pixels or among other meaningful spatial units.

Biodiversity metrics can have different components, including (1) taxonomic units; (2) abundance, frequency, or biomass of those units and their degree of evenness; and (3) the dispersion or distances between those units in trait, evolutionary, or spectral space. Myriad metrics quantify the major dimensions and components of diversity. Here we briefly describe several frequently used metrics; the equation for each metric and the source citation that provides the full details are given in Table 2.1.

**Table 2.1** A brief summary of metrics that are commonly used to estimate the diversity of different facets/dimensions of plant diversity

(continued)

**Table 2.1** (continued)

# *2.7.1 The Spatial Scale of Diversity: Alpha, Beta, and Gamma Diversity*

Diversity metrics are designed to capture biological variation at different spatial extents. Alpha diversity (*α*) represents the diversity within local communities, which are usually spatial subunits within a region or landscape. Whittaker first defined beta diversity (*β*) as the variation in biodiversity among local communities and gamma diversity (*γ*) as the total biodiversity in a region or a region's species pool (Whittaker 1960).

$$
\beta = \frac{\gamma}{a},
\tag{2.3}
$$

where *β* is beta diversity, *γ* gamma diversity, and *α* alpha diversity.

Other authors have defined beta diversity differently (see Tuomisto 2010), including using variance partitioning methods (Legendre and De Cáceres 2013).

## *2.7.2 Taxonomic Diversity*

Species richness is the number of species for a given area. It does not include abundance of individuals within species. However, the relative abundances, frequency, and biomass of species within a community matter in terms of capture rarity and evenness. Abundance-weighted metric, such as Simpson's diversity index (*D*), incorporates both richness and evenness. A set of indices based on Hill numbers—a unified standardization method for quantifying and comparing species diversity across samples, originally presented by Mark Hill (1973)—were refined by Chao et al. (2005, 2010). These are generalizable to all of the dimensions of diversity and consider the number of species and their relative abundances within a local community. Hill numbers require the specification of the diversity order (*q*), which determines the sensitivity of the metric to species relative abundance. Different orders of q result in different diversity measures; for example, *q* = 0 is simply species richness, *q* = 1 gives the exponential of Shannon's entropy index, and *q* = 2 gives the inverse of Simpson's concentration index.

# *2.7.3 Phylogenetic Diversity*

Phylogenetic diversity (PD) considers the extent of shared ancestry among species (Felsenstein 1985). For example, a plant community composed of two species that diverged from a common ancestor more recently is less phylogenetically diverse than a community of two species that diverged less recently. Faith's (1992) metric of PD sums the branch lengths among species within a community (from the root of the phylogeny to the tip). One feature of this metric is that it scales with species richness because as new species are added into the community, new branch lengths are also added. Other metrics were subsequently developed that calculate the mean evolutionary distances among species independently of the number of species [e.g., mean phylogenetic distance (MPD, Webb 2000b; Webb et al. 2002) or phylogenetic species variability (PSV), Helmus 2007]. Helmus (2007) developed two more phylogenetic diversity metrics that scale either with richness or by incorporating species abundances. Phylogenetic species richness (PSR) increases with the number of species, but reduces the effect of species richness proportionally to their degree of shared ancestry. Phylogenetic species evenness (PSE) is similar to PSV but includes abundances by adding individuals as additional tips descending from a single species node, with branch lengths of 0. Chao et al. (2010) defined the phylogenetic Hill number, *<sup>q</sup> D*(*T*), as the effective number of equally abundant and equally distinct lineages and phylogenetic branch diversity, *<sup>q</sup> PD*(*T*), as the effective total lineage length from the root node (i.e., the total evolutionary history of an assemblage) (Chao et al. 2014).

*Phylogenetic endemism* is another aspect of biodiversity that can be estimated from phylogenetic information and range maps of species (Faith et al. 2004). Phylogenetic endemism can be simply defined as the quantity of PD restricted to a given geographic area. This metric thus focuses on geographic areas, rather than on species, to discern areas of high endemism based on evolutionary history for conservation purposes.

## *2.7.4 Functional Diversity*

Widely used metrics of functional diversity consider the area or volume of trait space occupied by a community of species, the distances of each species to the center of gravity of those traits, and the trait distances between species (Mouillot et al. 2013). Functional attribute diversity (FAD) is a simple multivariate metric calculated as the sum of species pairwise distances of all measured continuous functional traits (Walker et al. 1999). Villeger et al. (2008) developed a series of functional diversity metrics that incorporate trait dispersion and distance among species as well as species abundances, including functional richness (FRic), functional divergence (FDiv), and functional evenness (FEve). Building on the framework of Villeger et al. (2008), Laliberté and Legendre (2010) developed functional dispersion (FDis), a functional diversity metric that is independent of species richness and can include species relative abundances (Table 2.1).

Scheiner's functional trait dispersion [*<sup>q</sup> D*(*TM*), Scheiner et al. 2017] calculates the effective number of species (or units) that are as distinct as the most distinct species (or unit) in that community. *<sup>q</sup> D*(*TM*) decomposes diversity estimates into three components: the number of units (*S*), functional evenness [*<sup>q</sup> E*(*T*), the extent to which units are equally dispersed], and mean dispersion [*M*', the average distance or the distinctiveness of these units]. Functional diversity measured as *<sup>q</sup> D*(*TM*) is maximized when there are more units in a community that are more equitably distributed (or less clumped) and more dispersed (or positioned further apart) in space. Like Chao's approach, *<sup>q</sup> D*(*TM*) includes Hill numbers (*<sup>q</sup>* ), which allow weighting of abundances: small and large q values emphasize rare and common species, respectively. Like many other biodiversity metrics, *<sup>q</sup> D*(*TM*) can be calculated from pairwise distances among species or individuals; thus, the metric can be applied to estimate different dimensions of biodiversity, including functional, phylogenetic (Scheiner 2012; Presley et al. 2014) and spectral components (Schweiger et al. 2018).

Briefly, functional trait dispersion [*<sup>q</sup> D*(*TM*)] is calculated as:

$$^{q}D\left(TM\right) = 1 + \left(S - 1\right) \times \,^{q}E\left(T\right) \times M\,\tag{2.4}$$

where:

*S* = species richness *E*(*T*) = trait evenness *M*' = trait dispersion *q* = Hill number

## *2.7.5 Spectral Diversity*

Like taxonomic, functional, and phylogenetic diversity, spectral diversity can be calculated in many different ways. Spectral alpha diversity metrics include the coefficient of variation of spectral indices (Oindo and Skidmore 2002) or spectral bands among pixels (Hall et al. 2010; Gholizadeh et al. 2018, 2019; Wang et al. 2018, the convex hull volume (Dahlin 2016) and the convex hull area (Gholizadeh et al. 2018) of pixels in spectral feature space, the mean distance of pixels from the spectral centroid (Rocchini et al. 2010), the number of spectrally distinct clusters or "spectral species" in ordination space (Féret and Asner 2014), and spectral variance (Laliberté et al. 2019). Schweiger et al. (2018) applied *<sup>q</sup> D*(*TM*) to species mean spectra and to individual pixels extracted at random from high-resolution proximal RS data. The second approach is independent of species identity and uses the same number of pixels per community for analysis. In this manner, the problem of diversity scaling with the number of species in a community is eliminated, and greater differences in reflectance spectra among pixels result in increased spectral diversity. Conceptually, spectral diversity metrics are versatile and can be tailored to match taxonomic or phylogenetic units, e.g., by using mean spectra for focal taxa, or to resemble functional diversity by selecting spectral bands that align with known absorption features for specific chemical traits or spectral indices that capture plant characteristics of known ecological importance. If measured at the appropriate scale (see Gamon et al. Chap. 16), spectral diversity can integrate the variation captured by other metrics of diversity and similarly predicts ecosystem function (Fig. 2.6).

**Fig. 2.6** (**a**) Aerial photo of the Cedar Creek long-term biodiversity experiment (BioDIV) (Courtesy of Cedar Creek Ecosystem Science Reserve). (**b**) Pairwise phylogenetic and (**c**) functional distances for the 17 most abundant prairie-grassland species in BioDIV are well-predicted by their spectral distances based on leaf-level spectral profiles (400–2500 nm). (**d**) Phylogenetic, (**e**) functional, and (**f**) leaf-level spectral diversities based on Scheiner's *<sup>q</sup> D*(*TM*) metric all predict ecosystem productivity in BioDIV. (**g**) Independent of information about species identities or their abundances, remotely sensed spectral diversity detected at high spatial resolution (1 mm) also predicts productivity. All graphs are redrawn from Schweiger et al. 2018. Species abbreviations in **b** and **c** are as follows: ACHMI = *Achillea millefolium* L., AMOCA = *Amorpha canescens* Pursh, ANDGE = *Andropogon gerardii* Vitman, ASCTU = *Asclepias tuberosa* L., KOEMA = *Koeleria macrantha* (Ledeb.) Schult., LESCA = *Lespedeza capitata* Michx., LIAAS = *Liatris aspera* Michx., LUPPE = *Lupinus perennis* L., MONFI = *Monarda fistulosa* L., PANVI = *Panicum virgatum* L., PASSMI = *Pascopyrum smithii* (Rydb.) Á. Löve, PETCA = *Petalostemum candidum* (Willd.), PETPU = *Petalostemum purpureum* (Vent.) Rydb., POAPR = *Poa pratensis* L., SCHSC = *Schizachyrium scoparium* Michx., SOLRI = *Solidago rigida* L., SORNU = *Sorghastrum nutans* (L.) Nash

## *2.7.6 Beta Diversity Metrics*

Whittaker's 1960 definition of beta diversity (Eq. 2.3) quantified the degree of differentiation among communities in relation to environmental gradients. Under this definition, beta diversity is defined as the ratio between regional (gamma) and local (alpha) diversities (Eq. 2.3) and measures the number of different communities in a region and the degree of differentiation between them (Whittaker 1960; Jost 2007). Indices such as Bray-Curtis dissimilarity and Jaccard and Sørensen indices evaluate similarity of communities based on the presence or abundance of species within them. Metrics of similarity used for species have been adapted for phylogenetic and functional trait distances (Bryant et al. 2008; Graham and Fine 2008; Kembel et al. 2010; Cardoso et al. 2014) and can equally be applied to spectral information (Gamon et al., Chap. 16).

While the ratio between regional and local communities provides a simple means to estimate beta diversity, there are many different ways to calculate taxonomic, functional, and phylogenetic beta diversity that can be grouped into pairwise and multiple-site metrics (reviewed in Baselga 2010). Notably, beta diversity can be partitioned into components that capture species replacement—the "turnover component"—caused by the exchange of species among communities and differences in the number of species, the "nestedness component," caused by differences in the number of species among communities. The turnover component can be interpreted as the difference between two community assemblages that contain contrasting subsets of species from a regional source pool, while the nestedness component represents the difference in species composition between two communities due to attrition of species in one assemblage relative to the other (Baselga 2010; Cardoso et al. 2014). Examining these different components of beta diversity for multiple dimensions of plant diversity provides a means to discern the role of historical and ongoing environmental sorting processes in the distribution of plant diversity at continental extents (Pinto-Ledezma et al. 2018b). In contrast to traditional diversity metrics, spectral diversity (alpha and beta) is only beginning to receive attention in biodiversity studies (Rocchini et al. 2018). Although different approaches have been proposed (Schmidtlein et al. 2007; Féret and Asner 2014; Rocchini et al. 2018; Laliberté et al. 2019), the estimation and mapping of dissimilarities in spectral composition (i.e., the variation among pixels) is similar to traditional estimations of beta diversity. For example, Laliberté et al. (2019) adapted the total community composition variance approach (Legendre and De Cáceres 2013) to estimate spectral diversity as spectral variance, partitioning the spectral diversity of a region (gamma diversity) into additive alpha and beta diversity components.

# **2.8 Links Between Plant Diversity, Other Trophic Levels, and Ecosystem Functions**

Plant diversity has consequences for other trophic levels, sometimes reducing herbivory on focal species (Castagneyrol et al. 2014), but also increasing the diversity of insects and their predators in an ecosystem (Dinnage et al. 2012; Lind et al. 2015). The distribution of plant traits within communities influences resource availability for other trophic levels above- and belowground, which affects community assembly and population dynamics across trophic levels. Diversity of neighbors surrounding focal trees can both increase and decrease pathogen and herbivore pressure on them (Grossman et al. 2019). Thus, while we know that plant diversity impacts other trophic levels, consistent rules across the globe that explain how and why these impacts occur remain elusive. An increasing number of studies reveal that plant diversity influences belowground microbial diversity and composition (Madritch et al. 2014; Cline et al. 2018). While these relationships are significant, they may explain limited variation given the number of other factors that influence microbial diversity and potentially due to a mismatch in sampling scales. Ultimately, it appears that chemical composition and productivity of aboveground components of ecosystems that can be remotely sensed are critical drivers of belowground processes, including microbial diversity (Madritch et al., Chap. 8).

Biodiversity loss is known to substantially decrease ecosystem functioning and ecosystem stability (Cardinale et al. 2011; O'Connor et al. 2017). Yet, the nature and scale of biodiversity-ecosystem function relationships remains a central question in biodiversity science. The issue is one that is ready to be tackled across scales using RS technology. The long-term biodiversity experiment at Cedar Creek Ecosystem Science Reserve (Tilman 1997) (Fig. 2.6), for example, has revealed the increasing effects of biodiversity on productivity over time (Reich et al. 2012) and that phylogenetic and functional diversity are highly predictive of productivity (Cadotte et al. 2008; Cadotte et al. 2009). Remotely sensed spectral diversity also predicts productivity (Sect. 2.9). Increased stability has also been linked to both higher plant richness (Tilman et al. 2006) and phylogenetic diversity (Cadotte et al. 2012) in this experiment. Tree diversity experiments show similar effects of increasing productivity with diversity (Tobner et al. 2016; Grossman et al. 2017) (Fig. 2.7), and these same trends emerge as the dominant pattern in forest plots globally

**Fig. 2.7** The Forest and Biodiversity (FAB) experiment at the Cedar Creek Ecosystem Science Reserve shows overyielding (**a**)—greater productivity than expected in species-rich communities compared to monocultures—also called the net biodiversity effect (NBE). Curves show 90% predictions from multiple linear regression models (yellow 2013–2014; blue 2014–2015). (Redrawn from Grossman et al. 2018.) Photos (**b**, **c**) show juvenile trees grown in mixtures with varying neighborhood composition. The first phase of the experiment, shown here, includes three 600 m2 blocks, each consisting of 49 plots (9.25 m2 ) planted in a grid with 0.5 m spacing

(Liang et al. 2016). Hundreds of rigorous biodiversity experiments have been designed and conducted to tease apart effects of changing numbers of species (richness) from effects of changing identities of species (composition) (O'Connor et al. 2017; Grossman et al. 2018; Isbell et al. 2018). Complementarity among diverse plant species that vary in their functional attributes and capture and respond to resources differently is the primary explanation for increasing productivity with diversity (Williams et al. 2017). Nevertheless, both the nature of biodiversity-ecosystem function (BEF) relationships and their causal mechanisms remain variable and scale dependent in natural systems. In the Nutrient Network global grassland experiments, in which communities have assembled naturally, the relationship between diversity and productivity is variable (Adler et al. 2011). In tropical forest plots around the globe, at spatial extents of 0.04 ha or less, the biodiversity-productivity relationship is strong. However, as scales increase to 0.25 or 1.0 ha, the relationship is no longer consistently positive and can frequently be negative (Chisholm et al. 2013). These varied relationships at contrasting spatial scales may result from nonlinear, hump-shaped relationships between biodiversity and ecosystem function across resource availability gradients as the nature of species interactions and their level of complementarity shift (Jaillard et al. 2014). RS methods—including imaging spectroscopy and LiDAR—that can detect both the diversity and the structure and function of ecosystems (Martin, Chap. 5; Atkins et al. 2018) can discern these relationships across spatial extents and biomes in natural systems. They thus have high potential to enhance our understanding of the scale and context dependence of linkages between biodiversity and ecosystem function (Grossman et al. 2018).

# **2.9 Incorporating Spectra into Relationships Between Biodiversity and Ecosystem Function**

Detection of spectral diversity, in particular, offers the potential to contribute to the quantification of BEF relationships at large scales (Schweiger et al. 2018) and is thus worth discussing in more detail. The variability captured by spectral diversity in a given ecosystem depends on the way the spectral diversity is calculated, as well as its spatial and spectral resolution (Sect. 2.7.5; Gamon et al., Chap. 16). From a functional perspective, spectral profiles measured at the leaf level depend on the chemical, structural, morphological, and anatomical characteristics of leaves (Ustin and Jacquemoud, Chap. 14). Variation in spectra and spectral diversity can be used to test hypotheses about how specific traits influence ecosystem function, community composition, and other characteristics of ecosystems, when using spectral bands or spectral indices with known associations with specific plant traits (Serbin and Townsend, Chap. 3). Moreover, spectral bands and indices can be weighted based on prior information about the relative contribution of individual traits to specific ecosystem characteristics. However, while the absorption features

of some chemical traits are known, the effects of other, particularly nonchemical, plant traits on spectra are less well understood, in part due to overlapping spectral features and challenges associated with accurately describing nonchemical traits (Ustin and Jacquemoud, Chap. 14). Using the full spectral profile of plants in spectral diversity calculations provides a means to integrate chemical, structural, morphological, and anatomical variation and to acknowledge the many ways plants differ from one another.

It is certainly more complicated to decipher the biological meaning of spectral diversity calculated from spectral profiles than from measures of biodiversity that are based on a specific set of plant traits or spectral bands or indices with known links to specific traits. However, the variance that is explained by models based on spectral profiles can be partitioned into known and unknown sources of variation. This provides a means to assess the relative contribution of traits with known spectral characteristics and traits that are less well understood spectrally or that are of yet-unrecognized importance. At the canopy level, when spectra are measured from a distance, the question of what spectra and spectral diversity represent is further complicated by the influences that plant architecture, soil, and other materials have on the spectral characteristics of image pixels (Wang et al. 2018; Gholizadeh et al. 2018). Again, the degree to which these characteristics matter for a particular ecosystem needs to be evaluated in the particular context of the study. Some ecosystem components such as shade, soil, rock, or debris, which influence remotely sensed spectra, are biologically meaningful because they influence light availability and microclimate and provide resources for other trophic levels.

The association between plant spectra and traits can be illustrated by plotting spectral distances against functional distances or dissimilarity, as illustrated using species from the Cedar Creek biodiversity experiment (Fig. 2.6d). Given that functional differences among species are expected to increase with evolutionary divergence time (Fig. 2.1b), positive relationships are also expected among spectral and phylogenetic distances. The observed associations among spectral, functional, and phylogenetic dissimilarity (Fig. 2.6a, b) allow biodiversity metrics based on any of these dimensions of biodiversity to explain a similar proportion of the total variability in aboveground productivity (Fig. 2.6c–e), which is known to increase with the functional diversity of the plant community in this system (Cadotte et al. 2009). The species in the biodiversity experiment at Cedar Creek are relatively functionally dissimilar and distantly related, such that spectral, functional, and phylogenetic diversity also predict species richness (not shown). One advantage of spectral diversity is that the metric can be calculated from remotely sensed image pixels without depending on information about the distribution and abundance of species in an area, their functional traits, or phylogenetic relationships (Schweiger et al. 2018). By extracting a random number of high-resolution image pixels in each plant community, Schweiger et al. (2018) found that remotely sensed spectral diversity explained the biodiversity effect on aboveground productivity about as well as spectral diversity calculated using leaf-level spectra (Fig. 2.6).

## **2.10 Links Between Biodiversity and Ecosystem Services**

Humans benefit from ecosystem functions and biodiversity. The benefits we derive from nature, often called ecosystem services, are a product of the biodiversity assembled over millions of years—and ecosystem properties of a given region, or the whole Earth (Daily 1997). Daily (1997) defines ecosystem services as "the conditions and processes through which natural ecosystems, and the species that make them up, sustain and fulfill human life." Ecosystem services, referred to by the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IPBES) as "nature's contributions to people" (Díaz et al. 2018), are a socioecological concept that emerged from the Millennium Ecosystem Assessment (2005) and include provisioning, regulating, supporting, and cultural services. Some ecosystem service categories include direct benefits of biodiversity—through the use and spiritual values that humans establish with elements of biodiversity and ecosystems—and indirect benefits through the contributions of biodiversity to critical regulating ecosystem functions. The diversities of functional traits of plants make up the primary productivity of life on Earth and are essential to the ecosystem services on which all life depends. Assessment of ecosystem services depends on understanding both the ecosystem functions on which ecosystem services are derived and how services are valued by humans (Schrodt et al., Chap. 17). Modeling efforts that incorporate remotely sensed data can be used to describe ecosystem functions and quantify the services they generate (Sharp et al. 2018). (For modeling tools that enable mapping and valuing ecosystem services, see https://naturalcapitalproject.stanford.edu/invest/.)

# **2.11 Trade-Offs Between Biodiversity and Ecosystem Services**

Biodiversity—as well as many regulating services to which biodiversity contributes and upon which it depends—frequently shows a negative trade-off with provisioning ecosystem services, such as agricultural production (Haines-Young and Potschin 2009). The nature of these trade-offs depends on the biophysical context, including the climate, soils, hydrology, and geology, and will differ among regions. A tradeoff curve represents the limits set by these biophysical constraints and can be thought of in economic terms as an "efficiency frontier" that sets the boundaries on possible combinations of biodiversity (or regulating services) and provisioning services (Polasky et al. 2008). Combinations above the curve are not possible; outcomes beneath the curve provide fewer total benefits than what is actually possible from the environment. Quantifying the biodiversity and ecosystem service potential from land and how they trade off are critical to efficient management of ecosystems. Current RS tools and forthcoming technologies are well-poised to decrease uncertainty in estimates of biodiversity—ecosystem service trade-offs—and can

Agricultural production

**Fig. 2.8** RS technologies that enable detection of biodiversity and ecosystem functions aid in modeling their trade-offs. (**a**) The "efficiency frontier," or biophysical constraints that limit biodiversity and crop production, depends on the specific climatic, historical, and resource context of the land area and on the growth or replenishment rate of the natural system. These constraints can vary among ecosystems (red vs black curves). (**b**) Where along the efficiency frontier we want to manage for depends on human values. The superimposed curves show isolines of equal utility (UA1–4 or UB1–4) for two different stakeholders (A and B) who have sharply different willingness to give up natural habitat for crop production and vice versa. Utility—or benefits to each stakeholder—increases moving from yellow to dark red. The two points at which the highest utility curve for each stakeholder intersects with the efficiency frontier represent the greatest feasible benefit to the stakeholder (points A and B). Often ecosystems are managed well below the efficiency frontier (green circle). RS may enable detection of components of biodiversity and ecosystem services at relevant spatial scales that can inform stakeholders about improved outcomes and aid negotiation among stakeholders. (**c**) Some trade-offs can have thresholds and tipping points that, once traversed, may result in a degraded alternative state. RS approaches that can aid in predicting uncertainties and temporal variability in trade-offs, indicated by thin blue lines, can help maximize ecosystem service benefits without overshooting thresholds that risk pushing the system into a degraded state. (Adapted from Cavender-Bares et al. 2015b)

contribute meaningfully to decision-making and resource management (Chaplin-Kramer et al. 2015; de Araujo Barbosa et al. 2015; Schrodt et al., Chap. 17).

Where along the efficiency frontier we wish to target our management efforts depends on human preferences. These can differ strongly among different stakeholders that have contrasting priorities (Cavender-Bares et al. 2015a, b). Distinguishing the biophysical limits of ecosystems from contrasting stakeholder preferences for what they want from ecosystems is a critical contribution to participatory processes that enable dialogue and progress toward sustainability (Cavender-Bares et al. 2015b; King et al. 2015). RS technologies that can enhance detection of biodiversity as well as both regulating and provisioning ecosystem services—and changes in these at multiple scales—can thus increase clarity in decision-making processes in the face of rapid global change (Fig. 2.8).

**Acknowledgments** We thank Shan Kothari for providing valuable feedback on an earlier version of the manuscript. The authors acknowledge funding from a NSF and NASA Dimensions grant (DEB 1342778), as well as the Cedar Creek NSF Long-Term Ecological Research grant (DEB 1234162) for support of some of the data presented here. We also thank the National Institute for Mathematical and Biological Synthesis (NIMBioS) working group on "Remotely Sensing Biodiversity," the Keck Institute for Space Studies working group "Unlocking a New Era in Biodiversity Science," and the NSF Research Coordination Network "Biodiversity Across Scales" (DEB 1745562).

## **References**

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# **Chapter 3 Scaling Functional Traits from Leaves to Canopies**

**Shawn P. Serbin and Philip A. Townsend**

# **3.1 Introduction**

Fossil energy use and land use change are the dominant drivers of the accelerating increase in atmospheric CO2 concentration and the principal causes of global climate change (IPCC 2018; IPBES 2018). Many of the observed and projected impacts of rising CO2 concentration and increased anthropogenic pressures on natural resources portend increasing risks to global terrestrial biomes, including direct impacts on biodiversity, yet the uncertainty surrounding the forecasting of biodiversity change, future climate, and the fate of terrestrial ecosystems by biodiversity and Earth system models (ESMs) is unacceptably high, hindering informed policy decisions at national and international levels (Jetz et al. 2007; Friedlingstein et al. 2014; Rice et al. 2018). As such, the impact of our changing climate and altered disturbance regimes on terrestrial ecosystems is a major focus of a number of disciplines, including the biodiversity, remote sensing (RS), and global change research communities.

Here we provide an overview of approaches to scale and map plant functional traits and diversity across landscapes with a focus on current approaches, leveraging on best practices provided by Schweiger (Chap. 15), benefits and issues with general techniques for linking and scaling traits and spectra, and other key considerations that need to be addressed when utilizing RS observations to infer plant functional traits across diverse landscapes.

S. P. Serbin (\*)

Brookhaven National Laboratory, Environmental and Climate Sciences Department, Upton, NY, USA e-mail<EMAIL_ADDRESS>
P. A. Townsend Department of Forest and Wildlife Ecology, University of Wisconsin, Madison, WI, USA

J. Cavender-Bares et al. (eds.), *Remote Sensing of Plant Biodiversity*, https://doi.org/10.1007/978-3-030-33157-3\_3

## *3.1.1 Plant Traits and Functional Diversity*

The importance of characterizing leaf and plant functional traits across scales is tied to the crucial role these traits play in mediating ecosystem structure, functioning, and resilience or response to perturbations (Lavorel and Garnier 2002; Reich et al. 2003; Wright et al. 2004; Reich 2014; Funk et al. 2017). The structural, biochemical, physiological, and phenological properties of plants regulate the growth and performance or fitness of plants and their ability to propagate or survive in diverse environments. As such, these traits are used to characterize the axes of variation that define broad plant functional types (PFTs), which in turn describe global vegetation patterns and properties (Ustin and Gamon 2010; Díaz et al. 2015), particularly in ESMs (Bonan et al. 2002; Wullschleger et al. 2014). Our focus here will be on leaf traits related to nutrition and defense that broadly fit within the concept of the leaf economics spectrum (LES, Wright et al. 2004), because these are most amendable to measurements using spectral methods. Other traits relating to reproductive strategies, hydraulics, physiology (though see Serbin et al. 2015), wood characteristics, etc. may be inferred from the traits described here, especially when combined with climate, soils, topography, or other data that generally are not directly detectable using RS.

Leaf nutritional properties and morphology are strong predictors of the photosynthetic capacity, plant growth, and biogeochemical cycling of terrestrial ecosystems (Aber and Melillo 1982; Green et al. 2003; Wright et al. 2004; Díaz et al. 2015). With respect to litter turnover and nutrient cycling, leaf traits that correspond to the distribution and magnitude of structural carbon and chemical compounds such as lignin and cellulose are used to infer the recalcitrant characteristics of canopy foliage (Madritch et al., Chap. 8). Capturing the spatial variation in these traits can therefore provide critical information on the nutrient cycling potential of ecosystems (Ollinger et al. 2002). On the other hand, leaf mass per area (LMA)—the ratio of a leaf's dry mass to its surface area—and its reciprocal, specific leaf area (SLA), correspond to a fundamental trade-off of leaf construction costs versus lightharvesting potential (Niinemets 2007; Poorter et al. 2009). The amount of foliar nitrogen within a leaf, on a mass (*N*mass, %) or area (*N*area, g/m2 ) basis, strongly regulates the photosynthetic capacity of leaves given its fundamental role in the lightharvesting pigments of leaves (chlorophyll a and b) and photosynthetic machinery, namely, the enzyme RuBisCo (Field and Mooney 1986; Evans and Clarke 2018). Other traits, such as the concentration or content of water and accessory pigments, are important indicators of plant health and stress (Ustin et al. 2009). Moreover, the covariation of traits is also a primary focus of ecological and biodiversity research given strong trade-offs defining different leaf form and function (Díaz et al. 2015). For example, across the spectrum of plant functional diversity (Wright et al. 2004), foliar nitrogen and LMA form a key axis of variation that describes end-members between "cheap" thinner, low-LMA leaves with high leaf nitrogen, higher photosynthetic rates and faster turnover versus thick, expensive leaves with high LMA, low nitrogen, slower turnover, and longer leaf life spans. Other traits with strong evidence for detection in the literature relate to plant allocation strategies (e.g., starch and sugar content) or defense compounds, such as phenolics (e.g., Asner et al. 2015; Kokaly and Skidmore 2015; Couture et al. 2016; Ely et al. 2019).