A general quantitative theory of forest structure and dynamics

We present the first part of a quantitative theory for the structure and dynamics of forests at demographic and resource steady state. The theory uses allometric scaling relations, based on metabolism and biomechanics, to quantify how trees use resources, fill space, and grow. These individual-level traits and properties scale up to predict emergent properties of forest stands, including size–frequency distributions, spacing relations, resource flux rates, and canopy configurations. Two insights emerge from this analysis: (i) The size structure and spatial arrangement of trees in the entire forest are emergent manifestations of the way that functionally invariant xylem elements are bundled together to conduct water and nutrients up from the trunks, through the branches, to the leaves of individual trees. (ii) Geometric and dynamic properties of trees in a forest and branches in trees scale identically, so that the entire forest can be described mathematically and behaves structurally and functionally like a scaled version of the branching networks in the largest tree. This quantitative framework uses a small number of parameters to predict numerous structural and dynamical properties of idealized forests.

[1]  P. Richards,et al.  The Vegetation of Moraballi Creek, British Guiana: An Ecological Study of a Limited Area of Tropical Rain Forest. Part II , 1933 .

[2]  H. M. Steven,et al.  Forest Management , 2020, Nature.

[3]  G. E. Hutchinson,et al.  A Theoretical Ecological Model of Size Distributions Among Species of Animals , 1959, The American Naturalist.

[4]  K. Yoda,et al.  Self-thinning in overcrowded pure stands under cultivated and natural conditions (Intraspecific competition among higher plants. XI) , 1963 .

[5]  C. Donald,et al.  Competition Among Crop and Pasture Plants , 1963 .

[6]  T. Kira,et al.  A QUANTITATIVE ANALYSIS OF PLANT FORM-THE PIPE MODEL THEORY : II. FURTHER EVIDENCE OF THE THEORY AND ITS APPLICATION IN FOREST ECOLOGY , 1964 .

[7]  T. Schoener,et al.  Differences in Insect Abundance and Diversity Between Wetter and Drier Sites During a Tropical Dry Season , 1968 .

[8]  R. Macarthur.  Mathematical Ecology and Its Place among the Sciences. (Book Reviews: Geographical Ecology. Patterns in the Distribution of Species) , 1974 .

[9]  S. R. Kerr,et al.  Theory of Size Distribution in Ecological Communities , 1974 .

[10]  J. Diamond,et al.  Ecology and Evolution of Communities , 1976, Nature.

[11]  T. McMahon,et al.  Tree structures: deducing the principle of mechanical design. , 1976, Journal of theoretical biology.

[12]  J. Harper.  Population Biology of Plants , 1979 .

[13]  J. Damuth,et al.  Population density and body size in mammals , 1981, Nature.

[14]  D. Tilman.  Resource competition and community structure. , 1983, Monographs in population biology.

[15]  Mark Westoby,et al.  The Self-Thinning Rule , 1984 .

[16]  J. Roughgarden,et al.  A model for the dynamics of an annual plant population , 1985 .

[17]  J. Lawton,et al.  Fractal dimension of vegetation and the distribution of arthropod body lengths , 1985, Nature.

[18]  Jacob Weiner,et al.  How competition for light and nutrients affects size variability in Ipomoea tricolor populations , 1986 .

[19]  Jacob Weiner,et al.  Size variability and competition in plant monocultures , 1986 .

[20]  R. Norberg,et al.  Theory of Growth Geometry of Plants and Self-Thinning of Plant Populations: Geometric Similarity, Elastic Similarity, and Different Growth Modes of Plant Parts , 1988, The American Naturalist.

[21]  James S. Clark,et al.  Integration of ecological levels: individual plant growth, population mortality and ecosystem processes. , 1990 .

[22]  A. Lugo,et al.  Tropical forest biomass estimation from truncated stand tables. , 1992 .

[23]  S. Pacala,et al.  Forest models defined by field measurements: I. The design of a northeastern forest simulator , 1993 .

[24]  T. Kohyama,et al.  Size-structured tree populations in gap-dynamic forest-the forest architecture hypothesis for the stable coexistence of species , 1993 .

[25]  James H. Brown,et al.  A General Model for the Origin of Allometric Scaling Laws in Biology , 1997, Science.

[26]  James H. Brown,et al.  Allometric scaling of plant energetics and population density , 1998, Nature.

[27]  M Franco,et al.  The interspecific mass-density relationship and plant geometry. , 1998, Proceedings of the National Academy of Sciences of the United States of America.

[28]  James H. Brown,et al.  The fourth dimension of life: fractal geometry and allometric scaling of organisms. , 1999, Science.

[29]  James H. Brown,et al.  A general model for the structure and allometry of plant vascular systems , 1999, Nature.

[30]  R. Kobe.  LIGHT GRADIENT PARTITIONING AMONG TROPICAL TREE SPECIES THROUGH DIFFERENTIAL SEEDLING MORTALITY AND GROWTH , 1999 .

[31]  Karl J. Niklas,et al.  Invariant scaling relations across tree-dominated communities , 2001, Nature.

[32]  K J Niklas,et al.  Invariant scaling relationships for interspecific plant biomass production rates and body size , 2001, Proceedings of the National Academy of Sciences of the United States of America.

[33]  Toshihiro Yamada,et al.  Dynamic steady state of patch-mosaic tree size structure of a mixed dipterocarp forest regulated by local crowding , 2001, Ecological Research.

[34]  Karl J Niklas,et al.  Global Allocation Rules for Patterns of Biomass Partitioning in Seed Plants , 2002, Science.

[35]  David A. Coomes,et al.  Disturbances prevent stem size‐density distributions in natural forests from following scaling relationships , 2003 .

[36]  Evan P. Economo,et al.  Scaling metabolism from organisms to ecosystems , 2003, Nature.

[37]  Karl J. Niklas,et al.  A general model for mass-growth-density relations across tree-dominated communities , 2003 .

[38]  J. Zak,et al.  Convergence across biomes to a common rain-use efficiency , 2004, Nature.

[39]  Geoffrey B. West,et al.  Effects of Body Size and Temperature on Population Growth , 2004, The American Naturalist.

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

[41]  H. Olff.  Effects of light and nutrient availability on dry matter and N allocation in six successional grassland species , 1992, Oecologia.

[42]  A. Kerkhoff,et al.  Ecosystem allometry: the scaling of nutrient stocks and primary productivity across plant communities. , 2006, Ecology letters.

[43]  C. Price,et al.  Scaling of mass and morphology in plants with minimal branching: an extension of the WBE model , 2006 .

[44]  Stephanie A. Bohlman,et al.  Testing metabolic ecology theory for allometric scaling of tree size, growth and mortality in tropical forests. , 2006, Ecology letters.

[45]  Karl J. Niklas,et al.  Biological scaling: Does the exception prove the rule? , 2007, Nature.

[46]  Ian J. Wright,et al.  “Diminishing returns” in the scaling of functional leaf traits across and within species groups , 2007, Proceedings of the National Academy of Sciences.

[47]  Nathan G. Swenson,et al.  A general integrative model for scaling plant growth, carbon flux, and functional trait spectra , 2007, Nature.

[48]  Charles A Price,et al.  A general model for allometric covariation in botanical form and function , 2007, Proceedings of the National Academy of Sciences.

[49]  B. Enquist,et al.  On estimating the exponent of power-law frequency distributions. , 2008, Ecology.

[50]  Ethan P White,et al.  On the relationship between mass and diameter distributions in tree communities. , 2008, Ecology letters.

[51]  James H Brown,et al.  Extensions and evaluations of a general quantitative theory of forest structure and dynamics , 2009, Proceedings of the National Academy of Sciences.