The global spectrum of plant form and function


Earth is home to a remarkable diversity of plant forms and life histories, yet comparatively few essential trait combinations have proved evolutionarily viable in today’s terrestrial biosphere. By analysing worldwide variation in six major traits critical to growth, survival and reproduction within the largest sample of vascular plant species ever compiled, we found that occupancy of six-dimensional trait space is strongly concentrated, indicating coordination and trade-offs. Three-quarters of trait variation is captured in a two-dimensional global spectrum of plant form and function. One major dimension within this plane reflects the size of whole plants and their parts; the other represents the leaf economics spectrum, which balances leaf construction costs against growth potential. The global plant trait spectrum provides a backdrop for elucidating constraints on evolution, for functionally qualifying species and ecosystems, and for improving models that predict future vegetation based on continuous variation in plant form and function.

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Figure 1: The volume in trait space occupied by vascular plant species is strongly constrained compared to theoretical null models.
Figure 2: The global spectrum of plant form and function.


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We are grateful to the many researchers who contributed to this study by making their data available, helping to check information, and/or providing comments at various stages. The study was supported by the TRY initiative on plant traits ( The TRY database is hosted at the Max Planck Institute for Biogeochemistry (Jena, Germany) and supported by DIVERSITAS/Future Earth, the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, and BACI (grant ID 640176). The study has also been partially supported by the following institutions and grants to S.Di.: Universidad Nacional de Córdoba and CONICET, FONCyT (PICT 554) and SECyT (Argentina), The Leverhulme Trust, UK, and Inter-American Institute for Global Change Research (IAI) SGP-CRA2015 (supported by US National Science Foundation grant GEO-1138881).

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Order in list of authors reflects overall participation in this article. S.Di., J.K. and S.L. designed the study. S.Di., J.K., J.H.C.C., I.J.W., S.L., M.K., C.W., E.G., I.C.P , M.W., H.P., P.B.R., A.T.M., J.D, A.N.G., A.E.Z., J.C., S.J.W., S.N.S., H.J., C.B., B.C., S.P., B. S. and D.K. contributed substantial amounts of data. S.Di., J.K., G.B., A.G. and V.F. constructed the database. S.Di., J.K., J.H.C.C., I.J.W., S.L., S.Dr., B.R., M.K., C.W., E.G., F.C., J.S.J., N.R., M.D.M. and L.D.G. carried out analyses. S.Di., J.K., J.H.C.C., I.J.W., S.L., M.K., C.W., I.C.P., M.W. and P.B.R. wrote the article with contributions from S.Dr., B.R., E.G., H.P., A.T.M., J.D., A.N.G., A.E.Z., J.C., S.J.W., S.N.S., H.J., C.B., B.C., S.P., B.S., DK, F.C., M.D.M. and L.D.G.

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Correspondence to Sandra Díaz.

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The authors declare no competing financial interests.

Extended data figures and tables

Extended Data Figure 1 Climatic and geographical coverage of the dataset.

ad, Green points, occurrence according to GBIF ( of species with information on all six traits (a, c) and at least one trait (b, d). Upper panels (a, b) show distribution in major climatic regions of the world; grey, MAP and MAT as in CRU0.5 degree climatology261; Biome classification according to ref. 262. Lower panels (c, d) show distribution in the global map (Robinson projection); grey, land surface. Maps based on the R package ‘maps’, accessed at The Comprehensive R Archive Network (

Extended Data Figure 2 Tests of the distribution of growth-forms (a) and major taxa (b) in trait space.

Woody and non-woody species differed significantly in their positions along PC1 but not along PC2. Angiosperms differed significantly from gymnosperms and pteridophytes in their positions along both axes; gymnosperms and pteridophytes differed in their position along PC1 but not along PC2 (see Methods for details of PCA analysis and a posteriori tests). Whiskers denote ± 3 s.d. from mean; n woody = 1,001; n non-woody = 1,209; n angiosperms = 2,120; n gymnosperms = 80; n pteridophytes = 14). Horizontal bars and dots within boxes indicate mean and median, respectively. Means with the same letter are not significantly different (Fisher’s least significant difference; P > 0.01).

Extended Data Figure 3 Selected bivariate relationships underlying the global spectrum of plant form and function, showing herbaceous (green) and woody (black) species separately.

See Extended Data Fig. 4 for standardized major axes statistics (slope, r2, sample size) of these and all other pairwise trait combinations.

Extended Data Figure 4 Bivariate relationships between the six traits that underlie the global spectrum of plant form and function.

The lower left portion of the matrix shows two-dimensional probability density distributions of bivariate trait relationships derived through kernel density estimation (see Methods). The colour gradient indicates regions of highest (red) to lowest (white) occurrence probability of trait combinations with contour lines indicating 0.5, 0.95 and 0.99 quantiles. The upper right portion contains standardized major axis (SMA)263 statistics (slope, r2, sample size n, and statistical significance, NS, P > 0.05; *0.05 > P > 0.01; **0.01 > P > 0.001; ***P < 0.001) for the corresponding relationships for all species (a), and for herbaceous (h) and woody species (w) separately. The diagonal displays the total sample sizes for each trait. For traits showing a strongly bimodal distribution, the all-species slope and correlation should be treated with caution. Pteridophytes show a discontinuous distribution in SM, but otherwise fall well within the general distribution of points; they represent less than 1% of the dataset, therefore including or excluding them does not significantly alter any of the relationships. SMAs were fitted using SMATR v.2 (

Extended Data Table 1 Principal component analyses (PCAs) of global plant trait data
Extended Data Table 2 Description and illustrative examples of species at different positions at the margin of the global spectrum of plant form and function

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Díaz, S., Kattge, J., Cornelissen, J. et al. The global spectrum of plant form and function. Nature 529, 167–171 (2016).

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