Tree height–diameter allometry across the United States

Abstract

The relationship between tree height and diameter is fundamental in determining community and ecosystem structure as well as estimates of biomass and carbon storage. Yet our understanding of how tree allometry relates to climate and whole organismal function is limited. We used the Forest Inventory and Analysis National Program database to determine height–diameter allometries of 2,976,937 individuals of 293 tree species across the United States. The shape of the allometric relationship was determined by comparing linear and nonlinear functional forms. Mixed-effects models were used to test for allometric differences due to climate and floristic (between angiosperms and gymnosperms) and functional groups (leaf habit and shade tolerance). Tree allometry significantly differed across the United States largely because of climate. Temperature, and to some extent precipitation, in part explained tree allometric variation. The magnitude of allometric variation due to climate, however, had a phylogenetic signal. Specifically, angiosperm allometry was more sensitive to differences in temperature compared to gymnosperms. Most notably, angiosperm height was more negatively influenced by increasing temperature variability, whereas gymnosperm height was negatively influenced by decreasing precipitation and increasing altitude. There was little evidence to suggest that shade tolerance influenced tree allometry except for very shade-intolerant trees which were taller for any given diameter. Tree allometry is plastic rather than fixed and scaling parameters vary around predicted central tendencies. This allometric variation provides insight into life-history strategies, phylogenetic history, and environmental limitations at biogeographical scales.

References Aiba, S. I., and T. Kohyama. 1996. Tree species stratification in relation to allometry and demography in a warm-temperate rain forest. J. Ecol. 84:207–218. Baker, F. S. 1948. A revised tolerance table. J. Forest. 47:179–181. Banin, L., T. R. Feldpausch, O. L. Phillips, T. R. Baker, J. Lloyd, K. Affum-Baffoe, et al. 2012. What controls tropical forest architecture? Testing environmental, structural and floristic drivers. Glob. Ecol. Biogeogr. 21:1179–1190. Bechtold, W. A., andC. T. Scott (2005) The forest inventory and analysis plot design. The enhanced Forest Inventory and Analysis program - national sampling design and estimation procedures. Pp. 27–42 in W. A. Bechtold, P. L. Patterson, eds. USDA forest service general technical reports SRS-80. U.S. Department of Agriculture, Forest Service, Southern Research Station, Asheville, North Carolina, USA. Becker, P. 2000. Competition in the regeneration niche between conifers and angiosperms: Bond's slow seedling hypothesis. Funct. Ecol. 14:401–412. Bond, W. J. 1989. The tortoise and the hare: ecology of angiosperm dominance and gymnosperm persistence. Biol. J. Linn. Soc. Lond. 36:227–249. Brown, S., A. J. R. Gillespi, and A. E. Lugo. 1989. Biomass estimation methods for tropical forests with applications to forestry inventory data. For. Sci. 35:881–902. Brown, J. H., G. B. West, and B. J. Enquist. 2000. Scaling in biology: patterns and processes, causes and consequences. Pp. 167–198 in J. H. Brown, G. B. West, eds. Scaling in biology. Oxford Univ. Press, Oxford. Bullock, S. H. 2000. Developmental patterns of tree dimensions in a Neotropical deciduous forest. Biotropica 32:42–52. Burnham, K. P., and D. R. Anderson. 2002. Model selection and multimodel inference. A practical information-theoretic approach. Springer, New York, NY. Burns, R. M., B. H. Honkala, and Tech. Coords. 1990. Silvics of North America: 1. Conifers; 2. Hardwoods. Agriculture Handbook, Pp. 877. U.S. Department of Agriculture, Forest Service, Washington, DC. Choat, B., S. Jansen, T. J. Brodribb, H. Cochard, S. Delzon, R. Bhaskar, et al. 2012. Global convergence in the vulnerability of forests to drought. Nature 491:752–755. Coll, M., J. Penuelas, M. Ninyerola, X. Pons, and J. Carnicer. 2013. Multivariate effect gradients driving forest demographic responses in the Iberian Peninsula. For. Ecol. Manage. 303:195–209. Ducey, M. J. 2012. Evergreenness and wood density predict height-diameter scaling in trees of the northeastern United States. For. Ecol. Manage. 279:21–26. Dudley, R., and C. Gans. 1991. A critique of symmorphosis and optimality models in physiology. Physiol. Zool. 64:627–637. Enquist, B. J., J. H. Brown, and G. West. 1998. Allometric scaling of plant energetics and population density. Letters to Nature 395:163–165. ESRI 2014. ArcGIS Desktop: Release 10.2.2. Redlands, CA: Environmental Systems Research Institute. Feldpausch, T. R., L. Banin, O. L. Phillips, T. R. Baker, S. L. Lewis, C. A. Quesada, et al. 2011. Height-diameter allometry of tropical forest trees. Biogeosciences, 8:1081–1106. Givnish, T. J. 1982. On the adaptive significance of leaf height in forest herbs. Am. Nat. 120:353–381. Givnish, T. J. 1995. Plant stems: biomechanical adaptation for energy capture and influence on species distributions. Pp. 3–49. in B. L. Gartner, eds. Plant stems: physiology and functional morphology. Academic Press, San Diego, CA. Gómez-Aparicio, L., R. Garcia-Valdes, P. Ruiz-Benito, and M. A. Zavala. 2011. Disentangling the relative importance of climate, size, and competition on tree growth in Iberian forests: implications for forest management under global change. Glob. Change Biol. 17:2400–2414. Greenhill, G. 1881. Determination of the greatest height consistent with stability that a vertical pole or mast can be made, and the greatest height to which a tree of given proportions can grow. Proc. Camb. Philos. Soc. 4:65–73. Grime, J. P. 2002. Plant strategies, vegetation processes, and ecosystem properties. Wiley, Chichester, U.K. Hacke, U. G., J. S. Sperry, and J. Pitterman. 2005. Efficiency versus safety tradeoffs for water conduction in angiosperm vessels versus gymnosperm tracheids. Pp. 333–353 in N. M. Holbrook and M. A. Zwieniecki, eds. Vascular transport in plants. Elsevier Academic Press, London, U.K. Hara, T. 1988. Dynamics of size structure in plant populations. Trends Ecol. Evol. 3:129–133. Henry, H. A. L., and L. W. Aarssen. 1999. The interpretation of stem diameter-height allometry in trees: biomechanical constraints, neighbor effects or biased regression? Ecol. Lett. 3:89–97. Hijmans, R. J., S. E. Cameron, J. L. Parra, P. G. Jones, and A. Jarvis. 2005. Very high resolution interpolated climate surfaces for global land areas. Int. J. Climatol. 25:1965–1978. Huxley, J. S. 1932. Problems of relative growth. Mcthuen, London. Reprinted 1972, Dover Publications, New York, NY. Kempes, C. P., G. B. West, K. Crowell, and M. Girvan. 2011. Predicting maximum tree heights and other traits from allometric scaling and resource limitations. PLoS One 6:e20551. King, D. A. 1991. Tree size. Natl Geogr. Res. Explor. 7:342–351. King, D. A., S. J. Davies, and N. S. M. Noor. 2006. Growth and mortality are related to adult tree size in a Malaysian mixed dipterocarp forest. For. Ecol. Manage. 223:152–158. Kobe, R. K. 1999. Light gradient partitioning among tropical tree species through differential seedling mortality and growth. Ecology 80:187–201. Kohyama, T. 1993. Size-structured tree populations in gap-dynamic forest-the forest architecture hypothesis for the stable coexistence of species. J. Ecol. 81:131–143. Lefcheck, J., and J. S. Casallas. 2014. R2 for generalized linear mixed effects models. GitHub repository. Available at https://github.com/jslefche/rsquared.glmer.git Lines, E. R., M. A. Zavala, D. W. Purves, and D. A. Coomes. 2012. Predictable changes in aboveground allometry of trees along gradients of temperature, aridity and competition. Glob. Ecol. Biogeogr. 21:1017–1028. McCarthy, M. C., and B. J. Enquist. 2007. Consistency between an allometric approach and optimal partitioning theory in global patterns of plant biomass allocation. Funct. Ecol. 21:713–720. McMahon, T. A. 1973. Size and shape in biology. Science 179:1202–1204. Moles, A. T., D. I. Warton, L. Warman, N. G. Swenson, S. W. Laffan, A. E. Zanne, et al. 2009. Global patterns in plant height. J. Ecol. 97:923–932. Muller-Landau, H. C., S. C. Condit, S. A. Chave, S. Thomas, S. Bohlman, R. Bunyavejchewin, et al. 2006. Testing metabolic ecology theory for allometric scaling of tree size, growth and mortality in tropical forests. Ecol. Lett., 9:575–588. Nakagawa, S., and H. Schielzeth. 2013. A general and simple method for obtaining R² from generalized linear mixed-effects models. Methods Ecol. Evol. 4:133–142. Niinements, Ü., and F. Valladares. 2006. Tolerance to shade, drought, and waterlogging of temperate northern hemisphere trees and shrubs. Ecol. Monogr. 76:521–547. Niklas, K. J. 1993. The scaling of plant height: a comparison among major plant clades and anatomical grades. Ann. Bot. 72:165–172. Niklas, K. J. 1994. Plant allometry: the scaling of form and process. Univ. of Chicago Press, USA. Niklas, K. J. 2004. Plant allometry: is there a grand unifying theory? Biol. Rev. Camb. Philos. Soc. 79:871–889. Niklas, K. J., and H. C. Spatz. 2004. Growth and hydraulic (not mechanical) constraints govern the scaling of tree height and mass. Proc. Natl Acad. Sci. USA 101:15661–15663. Osunkoya, O. O., K. Omar-Ali, N. Amit, J. Dayan, D. S. Daud, and T. K. Sheng. 2007. Comparative height-crown allometry and mechanical design in 22 tree species of Kuala Belalong rainforest, Brunei, Borneo. Am. J. Bot. 94:1951–1962. Pinheiro, J., D. Bates, S. DebRoy, D. Sarkar, and the R Development Core Team. 2013. nlme: Linear and Nonlinear Mixed Effects Models. R package. Poorter, L. 2001. Light-dependent changes in biomass allocation and their importance for growth of rain forest tree species. Funct. Ecol. 15:113–123. Poorter, L., F. J. J. M. Bongers, F. J. Sterck, and H. Wöll. 2003. Architecture of 53 rain forest tree species differing in adult stature and shade tolerance. Ecology 84:602–608. Purves, D. W., J. W. Lichstein, and S. W. Pacala. 2007. Crown plasticity and competition for canopy space: a new spatially implicit model parameterized for 250 North American Tree Species. PLoS One 2:e870. R Core Team (2014) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Aus?tria. URL http://www.R-project.org/ Russo, S. E., S. K. Wiser, and D. A. Coomes. 2007. Growth–size scaling relationships of woody plant species differ from predictions of the metabolic ecology model. Ecol. Lett. 10:889–901. Ryan, M. G., N. Phillips, and B. J. Bond. 2006. The hydraulic limitation hypothesis revisited. Plant, Cell Environ. 29:367–381. Schwinning, S., and J. R. Ehleringer. 2001. Water use trade-offs and optimal adaptations to pulse-driven arid ecosystems. J. Ecol. 89:464–480. Swenson, N. G., and B. J. Enquist. 2007. Ecological and evolutionary determinants of a key plant functional trait: wood density and its community-wide variation across latitude and elevation. Am. J. Bot. 91:451–459. Swenson, N. G., and M. D. Weiser. 2010. Plant geography upon the basis of functional traits: an example from eastern North American trees. Ecology 91:2234–2241. Wang, J., N. E. Ives, and M. J. Lechowicz. 1992. The relation of foliar phenology to xylem embolism in trees. Funct. Ecol. 6:469–475. Wang, X., J. Fang, Z. Tang, and B. Zhu. 2006. Climatic control of primary forest structure and DBH-height allometry in Northeast China. For. Ecol. Manage. 234:264–274. Woudenberg, S. W., B. L. Conkling, B. M. O'Connell, E. B. LaPoint, J. A. Turner, and K. L. Waddell. 2010. The forest inventory and analysis database: database description and user's manual version 4.0 for Phase 2. United States Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fort Collins, Colorado, USA. Zuur, A. F., E. N. Ieno, N. J. Walker, A. A. Saveliev, and G. Smith. 2009. Mixed effects models and extensions in ecology with R. Springer, New York, NY.