Classification of Tree Functional Types in a Megadiverse Tropical Mountain Forest from Leaf Optical Metrics and Functional Traits for Two Related Ecosystem Functions

Few plant functional types (PFTs) with fixed average traits are used in land surface models (LSMs) to consider feedback between vegetation and the changing atmosphere. It is uncertain if highly diverse vegetation requires more local PFTs. Here, we analyzed how 52 tree species of a megadiverse mountain rain forest separate into local tree functional types (TFTs) for two functions: biomass production and solar radiation partitioning. We derived optical trait indicators (OTIs) by relating leaf optical metrics and functional traits through factor analysis. We distinguished four OTIs explaining 38%, 21%, 15%, and 12% of the variance, of which two were considered important for biomass production and four for solar radiation partitioning. The clustering of species-specific OTI values resulted in seven and eight TFTs for the two functions, respectively. The first TFT ensemble (P-TFTs) represented a transition from low to high productive types. The P-TFT were separated with a fair average silhouette width of 0.41 and differed markedly in their main trait related to productivity, Specific Leaf Area (SLA), in a range between 43.6 to 128.2 (cm2/g). The second delineates low and high reflective types (E-TFTs), were subdivided by different levels of visible (VIS) and near-infrared (NIR) albedo. The E-TFTs were separated with an average silhouette width of 0.28 and primarily defined by their VIS/NIR albedo. The eight TFT revealed an especially pronounced range in NIR reflectance of 5.9% (VIS 2.8%), which is important for ecosystem radiation partitioning. Both TFT sets were grouped along elevation, modified by local edaphic gradients and species-specific traits. The VIS and NIR albedo were related to altitude and structural leaf traits (SLA), with NIR albedo showing more complex associations with biochemical traits and leaf water. The TFTs will support LSM simulations used to analyze the functioning of mountain rainforests under climate change.

[1]  W. R. Philipson,et al.  Field reflectance calibration with grey standard reflectors , 1989 .

[2]  Niro Higuchi,et al.  Species Spectral Signature: Discriminating closely related plant species in the Amazon with Near-Infrared Leaf-Spectroscopy , 2013 .

[3]  Marco Landi,et al.  Multiple functional roles of anthocyanins in plant-environment interactions. , 2015 .

[4]  J. Camargo,et al.  Near Infrared Spectroscopy Facilitates Rapid Identification of Both Young and Mature Amazonian Tree Species , 2015, PloS one.

[5]  Y. Li,et al.  Variation in leaf chlorophyll concentration from tropical to cold-temperate forests: Association with gross primary productivity , 2018 .

[6]  Ke Zhang,et al.  Ecosystem heterogeneity determines the ecological resilience of the Amazon to climate change , 2015, Proceedings of the National Academy of Sciences.

[7]  B. Luz,et al.  Attenuated total reflectance spectroscopy of plant leaves: a tool for ecological and botanical studies. , 2006, The New phytologist.

[8]  Sean C. Thomas,et al.  Increasing carbon storage in intact African tropical forests , 2009, Nature.

[9]  Jürgen Homeier,et al.  Is tropical montane forest heterogeneity promoted by a resource-driven feedback cycle? Evidence from nutrient relations, herbivory and litter decomposition along a topographical gradient , 2015 .

[10]  T. Hickler,et al.  A research framework for projecting ecosystem change in highly diverse tropical mountain ecosystems , 2021, Oecologia.

[11]  Nathan J B Kraft,et al.  Functional traits and the growth-mortality trade-off in tropical trees. , 2010, Ecology.

[12]  D. Horler,et al.  The red edge of plant leaf reflectance , 1983 .

[13]  S. Güsewell N : P ratios in terrestrial plants: variation and functional significance. , 2004, The New phytologist.

[14]  Dar A. Roberts,et al.  Species-Level Differences in Hyperspectral Metrics among Tropical Rainforest Trees as Determined by a Tree-Based Classifier , 2012, Remote. Sens..

[15]  A. Formaggio,et al.  Discrimination of sugarcane varieties in Southeastern Brazil with EO-1 Hyperion data , 2005 .

[16]  Anatoly A. Gitelson,et al.  Non‐destructive detection of water stress and estimation of relative water content in maize , 2009 .

[17]  D. Sims,et al.  Relationships between leaf pigment content and spectral reflectance across a wide range of species, leaf structures and developmental stages , 2002 .

[18]  J. Peñuelas,et al.  Remote sensing of nitrogen and lignin in Mediterranean vegetation from AVIRIS data: Decomposing biochemical from structural signals , 2002 .

[19]  Roberta E. Martin,et al.  Tropical forest leaves may darken in response to climate change , 2018, Nature Ecology & Evolution.

[20]  P. Reich,et al.  From tropics to tundra: global convergence in plant functioning. , 1997, Proceedings of the National Academy of Sciences of the United States of America.

[21]  O. Phillips,et al.  The variation of productivity and its allocation along a tropical elevation gradient: a whole carbon budget perspective. , 2017, The New phytologist.

[22]  Mui Lay,et al.  Reflectance properties and physiological responses of Salicornia virginica to heavy metal and petroleum contamination. , 2005, Environmental pollution.

[23]  J. Abadía,et al.  Photosystem II efficiency in low chlorophyll, iron-deficient leaves , 1999, Plant and Soil.

[24]  P. Reich The world‐wide ‘fast–slow’ plant economics spectrum: a traits manifesto , 2014 .

[25]  Rasmus Houborg,et al.  Thermal-based modeling of coupled carbon, water, and energy fluxes using nominal light use efficiencies constrained by leaf chlorophyll observations , 2014 .

[26]  Roberta E. Martin,et al.  Solar radiation and functional traits explain the decline of forest primary productivity along a tropical elevation gradient. , 2017, Ecology letters.

[27]  Susan L Ustin,et al.  Remote sensing of plant functional types. , 2010, The New phytologist.

[28]  Ben Somers,et al.  Optical trait indicators for remote sensing of plant species composition: Predictive power and seasonal variability , 2017 .

[29]  L. Başayiğit,et al.  Analysis of VNIR reflectance for prediction of macro and micro nutrient and chlorophyll contents in apple trees (Malus communis). , 2009 .

[30]  P. Fabian,et al.  Complex topography influences atmospheric nitrate deposition in a neotropical mountain rainforest , 2013 .

[31]  E. Garnier,et al.  Evidence for a ‘plant community economics spectrum’ driven by nutrient and water limitations in a Mediterranean rangeland of southern France , 2012 .

[32]  Yingjie Tian,et al.  A Comprehensive Survey of Clustering Algorithms , 2015, Annals of Data Science.

[33]  P. Reich,et al.  The Evolution of Plant Functional Variation: Traits, Spectra, and Strategies , 2003, International Journal of Plant Sciences.

[34]  Fabian Ewald Fassnacht,et al.  Differentiating plant functional types using reflectance: which traits make the difference? , 2018, Remote Sensing in Ecology and Conservation.

[35]  Chaoyang Wu,et al.  Estimating chlorophyll content from hyperspectral vegetation indices : Modeling and validation , 2008 .

[36]  G. A. Blackburn,et al.  Quantifying Chlorophylls and Caroteniods at Leaf and Canopy Scales: An Evaluation of Some Hyperspectral Approaches , 1998 .

[37]  D. Roberts,et al.  Estimation of tropical rain forest aboveground biomass with small-footprint lidar and hyperspectral sensors , 2011 .

[38]  Jörg Bendix,et al.  Modeling tropical montane forest biomass, productivity and canopy traits with multispectral remote sensing data , 2019, Remote Sensing of Environment.

[39]  Factors controlling the productivity of tropical Andean forests: climate and soil are more important than tree diversity , 2021 .

[40]  Benjamin L Turner,et al.  Leaf nitrogen to phosphorus ratios of tropical trees: experimental assessment of physiological and environmental controls. , 2010, The New phytologist.

[41]  V. Pillar,et al.  An improved method for searching plant functional types by numerical analysis , 2003 .

[42]  Benoit Rivard,et al.  Variability in leaf optical properties of Mesoamerican trees and the potential for species classification. , 2006, American journal of botany.

[43]  J. Chave,et al.  Towards a Worldwide Wood Economics Spectrum 2 . L E a D I N G D I M E N S I O N S I N W O O D F U N C T I O N , 2022 .

[44]  Benoit Rivard,et al.  Classification of tree species based on longwave hyperspectral data from leaves, a case study for a tropical dry forest , 2018, Int. J. Appl. Earth Obs. Geoinformation.

[45]  A. Gitelson,et al.  Three‐band model for noninvasive estimation of chlorophyll, carotenoids, and anthocyanin contents in higher plant leaves , 2006 .

[46]  Shixiao Yu,et al.  Linking Aboveground Traits to Root Traits and Local Environment: Implications of the Plant Economics Spectrum , 2019, Front. Plant Sci..

[47]  Bin Chen,et al.  Leaf chlorophyll content as a proxy for leaf photosynthetic capacity , 2017, Global change biology.

[48]  P. Alton How useful are plant functional types in global simulations of the carbon, water, and energy cycles? , 2011 .

[49]  Delbert Dueck,et al.  Clustering by Passing Messages Between Data Points , 2007, Science.

[50]  T. Nauss,et al.  Optical Properties of selected plants from a tropical mountain ecosystem : traits for plant functional types to parametrize a land surface model , 2011 .

[51]  André Große-Stoltenberg,et al.  Field Spectroscopy in the VNIR-SWIR Region to Discriminate between Mediterranean Native Plants and Exotic-Invasive Shrubs Based on Leaf Tannin Content , 2015, Remote. Sens..

[52]  A. Fries,et al.  Catchment precipitation processes in the San Francisco valley in southern Ecuador: combined approach using high-resolution radar images and in situ observations , 2014, Meteorology and Atmospheric Physics.

[53]  E. Mitchard The tropical forest carbon cycle and climate change , 2018, Nature.

[54]  Bruno Hérault,et al.  What drives long-term variations in carbon flux and balance in a tropical rainforest in French Guiana? , 2018 .

[55]  Luiz E. O. C. Aragão,et al.  Net primary productivity allocation and cycling of carbon along a tropical forest elevational transect in the Peruvian Andes , 2010 .

[56]  I. Noble,et al.  Deriving functional types for rain-forest trees , 1999 .

[57]  B. Schaffer,et al.  Evaluation of reflectance spectroscopy indices for estimation of chlorophyll content in leaves of a tropical tree species , 2012, Photosynthetica.

[58]  P. Rousseeuw Silhouettes: a graphical aid to the interpretation and validation of cluster analysis , 1987 .

[59]  Jörg Bendix,et al.  Natural or anthropogenic? On the origin of atmospheric sulfate deposition in the Andes of southeastern Ecuador , 2014 .

[60]  M. Paul,et al.  Sink regulation of photosynthesis. , 2001, Journal of experimental botany.

[61]  Environmental Filtering and Positive Plant Litter Feedback Simultaneously Explain Correlations Between Leaf Traits and Soil Fertility , 2015, Ecosystems.

[62]  Sophie Graefe,et al.  Elevation effects on the carbon budget of tropical mountain forests (S Ecuador): the role of the belowground compartment , 2011 .

[63]  S. Wand,et al.  Anthocyanins in vegetative tissues: a proposed unified function in photoprotection. , 2002, The New phytologist.

[64]  Sebastian Schmidtlein,et al.  Radiative transfer modelling reveals why canopy reflectance follows function , 2019, Scientific Reports.

[65]  B. Thies,et al.  Regionalization of wind-speed data to analyse tree-line wind conditions in the eastern Andes of southern Ecuador , 2015 .

[66]  R. Leigh,et al.  Calcium delivery and storage in plant leaves: exploring the link with water flow. , 2011, Journal of experimental botany.

[67]  Roberta E. Martin,et al.  Spectroscopy of canopy chemicals in humid tropical forests , 2011 .

[68]  P. Townsend,et al.  Foliar functional traits from imaging spectroscopy across biomes in eastern North America. , 2020, The New phytologist.

[69]  P. Curran Remote sensing of foliar chemistry , 1989 .

[70]  Sean C. Thomas,et al.  The worldwide leaf economics spectrum , 2004, Nature.

[71]  Sandra Lavorel,et al.  Plant functional types: are we getting any closer to the Holy Grail? , 2007 .

[72]  C. Tucker Red and photographic infrared linear combinations for monitoring vegetation , 1979 .

[73]  Steven F. Oberbauer,et al.  Leaf optical properties along a vertical gradient in a tropical rain forest canopy in Costa Rica. , 1995 .

[74]  Mark Westoby,et al.  EVOLUTIONARY DIVERGENCES IN LEAF STRUCTURE AND CHEMISTRY, COMPARING RAINFALL AND SOIL NUTRIENT GRADIENTS , 1999 .

[75]  Xinguo Li,et al.  The Significance of Calcium in Photosynthesis , 2019, International journal of molecular sciences.

[76]  Zong-Liang Yang,et al.  Technical description of version 4.5 of the Community Land Model (CLM) , 2013 .

[77]  A. Gitelson,et al.  Relationships between gross primary production, green LAI, and canopy chlorophyll content in maize: Implications for remote sensing of primary production , 2014 .

[78]  J. Peñuelas,et al.  The reflectance at the 950–970 nm region as an indicator of plant water status , 1993 .

[79]  J. Bendix,et al.  Cloud occurrence and cloud properties in Ecuador , 2006 .

[80]  Ulrich Bodenhofer,et al.  APCluster: an R package for affinity propagation clustering , 2011, Bioinform..

[81]  W. Kustas,et al.  Satellite retrievals of leaf chlorophyll and photosynthetic capacity for improved modeling of GPP , 2013 .

[82]  Susan L. Ustin,et al.  Leaf spectral clusters as potential optical leaf functional types within California ecosystems , 2016 .

[83]  C. Jordan Derivation of leaf-area index from quality of light on the forest floor , 1969 .

[84]  Yu-long Zheng,et al.  Specific leaf area relates to the differences in leaf construction cost, photosynthesis, nitrogen allocation, and use efficiencies between invasive and noninvasive alien congeners , 2008, Planta.

[85]  B. R. Smith,et al.  Aluminum-induced effects on Photosystem II photochemistry in citrus leaves assessed by the chlorophyll a fluorescence transient. , 2008, Tree physiology.

[86]  S. Ollinger Sources of variability in canopy reflectance and the convergent properties of plants. , 2011, The New phytologist.

[87]  Jörg Bendix,et al.  Rainfall distribution in the Andes of southern Ecuador derived from blending weather radar data and meteorological field observations , 2011 .