Forests are major components of the global carbon cycle, providing substantial feedback to atmospheric greenhouse gas concentrations1. Our ability to understand and predict changes in the forest carbon cycle—particularly net primary productivity and carbon storage—increasingly relies on models that represent biological processes across several scales of biological organization, from tree leaves to forest stands2,3. Yet, despite advances in our understanding of productivity at the scales of leaves and stands, no consensus exists about the nature of productivity at the scale of the individual tree4,5,6,7, in part because we lack a broad empirical assessment of whether rates of absolute tree mass growth (and thus carbon accumulation) decrease, remain constant, or increase as trees increase in size and age. Here we present a global analysis of 403 tropical and temperate tree species, showing that for most species mass growth rate increases continuously with tree size. Thus, large, old trees do not act simply as senescent carbon reservoirs but actively fix large amounts of carbon compared to smaller trees; at the extreme, a single big tree can add the same amount of carbon to the forest within a year as is contained in an entire mid-sized tree. The apparent paradoxes of individual tree growth increasing with tree size despite declining leaf-level8,9,10 and stand-level10 productivity can be explained, respectively, by increases in a tree’s total leaf area that outpace declines in productivity per unit of leaf area and, among other factors, age-related reductions in population density. Our results resolve conflicting assumptions about the nature of tree growth, inform efforts to undertand and model forest carbon dynamics, and have additional implications for theories of resource allocation11 and plant senescence12.
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We thank the hundreds of people who have established and maintained the forest plots and their associated databases; M. G. Ryan for comments on the manuscript; C. D. Canham and T. Hart for supplying data; C. D. Canham for discussions and feedback; J. S. Baron for hosting our workshops; and Spain’s Ministerio de Agricultura, Alimentación y Medio Ambiente (MAGRAMA) for granting access to the Spanish Forest Inventory Data. Our analyses were supported by the United States Geological Survey (USGS) John Wesley Powell Center for Analysis and Synthesis, the USGS Ecosystems and Climate and Land Use Change mission areas, the Smithsonian Institution Global Earth Observatory—Center for Tropical Forest Science (CTFS), and a University of Nebraska-Lincoln Program of Excellence in Population Biology Postdoctoral Fellowship (to N.G.B.). In addition, X.W. was supported by National Natural Science Foundation of China (31370444) and State Key Laboratory of Forest and Soil Ecology (LFSE2013-11). Data collection was funded by a broad range of organizations including the USGS, the CTFS, the US National Science Foundation, the Andrews LTER (NSF-LTER DEB-0823380), the US National Park Service, the US Forest Service (USFS), the USFS Forest Inventory and Analysis Program, the John D. and Catherine T. MacArthur Foundation, the Andrew W. Mellon Foundation, MAGRAMA, the Council of Agriculture of Taiwan, the National Science Council of Taiwan, the National Natural Science Foundation of China, the Knowledge Innovation Program of the Chinese Academy of Sciences, Landcare Research and the National Vegetation Survey Database (NVS) of New Zealand, the French Fund for the Global Environment and Fundación ProYungas. This paper is a contribution from the Western Mountain Initiative, a USGS global change research project. Any use of trade names is for descriptive purposes only and does not imply endorsement by the USA government.
The authors declare no competing financial interests.
Fitted model parameters for each species have been deposited in USGS’s ScienceBase at http://dx.doi.org/10.5066/F7JS9NFM.
Extended data figures and tables
Bars show the percentage of species with mass growth rates that increase with tree mass for each bin; black shading indicates percentage significant at P ≤ 0.05. Tree masses increase with bin number. a, Species fitted with one bin (165 species); b, Species fitted with two bins (139 species); c, Species fitted with three bins (56 species); and d, Species fitted with four bins (43 species).
Trees with growth rates ≤ 0 were dropped from the analysis, reducing the number of species meeting our threshold sample size for analysis. a, Africa (33 species); b, Asia (123 species); c, Australasia (22 species); d, Central and South America (73 species); e, Europe (41 species); and f, North America (89 species). Trunk diameters are approximate values for reference, based on the average diameters of trees of a given mass.
Extended Data Figure 3 Aboveground mass growth rates for 41 tree species in the absence of competition.
The ‘+’ or ‘−’ symbol preceding each species code indicates, respectively, species with mass growth rates that increased continuously with tree size or species with mass growth rates that declined in the largest trees. Sources of the diameter growth equations used to calculate mass growth were: a, ref. 45; b, ref. 46; c, ref. 48; d, ref. 47; and e, ref. 49. ABAM, Abies amabilis; ABBA, Abies balsamea; ABCO, Abies concolor; ABLA, Abies lasiocarpa; ABMA, Abies magnifica; ACRU, Acer rubrum; ACSA, Acer saccharum; BEAL, Betula alleghaniensis; BELE, Betula lenta; BEPA, Betula papyrifera; CADE, Calocedrus decurrens; CASA, Castanea sativa; FAGR, Fagus grandifolia; FASY, Fagus sylvatica; FRAM, Fraxinus americana; JUTH, Juniperus thurifera; PIAB, Picea abies; PICO, Pinus contorta; PIHA, Pinus halepensis; PIHY, Picea hybrid (a complex of Picea glauca, P. sitchensis and P. engelmannii); PILA, Pinus lambertiana; PINI, Pinus nigra; PIPINA, Pinus pinaster; PIPINE, Pinus pinea; PIRU, Picea rubens; PIST, Pinus strobus; PISY, Pinus sylvestris; PIUN, Pinus uncinata; POBA, Populus balsamifera ssp. trichocarpa; POTR, Populus tremuloides; PRSE, Prunus serotina; QUFA, Quercus faginea; QUIL, Quercus ilex; QUPE, Quercus petraea; QUPY, Quercus pyrenaica; QURO, Quercus robar; QURU, Quercus rubra; QUSU, Quercus suber; THPL, Thuja plicata; TSCA, Tsuga canadensis; and TSHE, Tsuga heterophylla.
a, The allometric equation for moist tropical forests34—used for the majority of tree species—shows no evident systematic bias in predicted aboveground dry mass, M, relative to trunk diameter (n = 1,504 trees). b, In contrast, our simplest form of allometric equation—used for 22% of our species and here applied to nine temperate species—shows an apparent bias towards overestimating M for large trees (n = 1,358 trees). c, New allometries that we created for the nine temperate species removed the apparent bias in predicted M.
Extended Data Figure 5 Estimated mass growth rates of the nine temperate species of Extended Data Fig. 4.
Growth was estimated using the simplest form of allometric model [log(M) = a + blog(D)] (a) and our allometric models fitted with piecewise linear regression (b). Regardless of the allometric model form, all nine species show increasing G in the largest trees.
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Stephenson, N., Das, A., Condit, R. et al. Rate of tree carbon accumulation increases continuously with tree size. Nature 507, 90–93 (2014). https://doi.org/10.1038/nature12914
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