DOI: 10.5061/dryad.1ns1rn92j
title: Data from: Coordination of bark and wood traits underlies forest-to-savanna evolutionary transitions

description: Aim: To test the hypothesis that adaptive shifts leading to the assembly
 of tropical savannas involved coordination between bark and wood traits,
 and to understand the underlying mechanisms. Location: Tropical South
 America. Taxon: Angiosperms (woody). Methods: We compiled data on three
 bark traits (total, inner, and outer relative bark thickness), wood
 density, maximum height, five secondary xylem traits, and on species’
 habitat information (light environment, climate, soil, and fire history)
 for Neotropical savanna, forest, and generalist species (biome groups). We
 tested for pairwise and multivariate associations among traits across
 species, and if biome group and habitat conditions explained species
 positions along the resulting strategy axes. Results: Traits covaried
 along four different axes. The first axis was consistent with a trade-off
 between fire (thick barks) and shade tolerance (low bark to diameter
 ratio, high vessel density) and contributed to differentiate the three
 biome groups according to preference for shaded environments. Forest
 species also differed from savanna and generalist species in a separate
 axis by being more resource acquisitive. Maximum height and wood density
 did not strongly trade-off with bark thickness, although maximum height
 negatively covaried with relative outer bark thickness. Preference for
 shaded conditions was the main driver of variation in the two principal
 strategy axes, but temperature, fire and soil sand content also explained
 differences in plant stature between savanna and generalist species. Main
 conclusions: Allocation to bark is constrained by trade-offs with wood,
 opposing shade-tolerant and acquisitive forest species to fire-resistant
 and conservative savanna species. Rather than a single strategy axis,
 three axes are necessary to understand the functional differences among
 savanna, forest, and generalist species. Because two of these axes are
 controlled by light availability, the associated traits tend to covary in
 space and time, but not across species.
description: This dataset was
 compiled from 83 published studies conducted in the following tropical
 South America biomes: moist forest, dry forest and savannas (including
 flooded savannas). We compiled data on bark thickness divided by stem
 diameter (i.e., relative bark thickness; relBT), discriminating between
 the inner (relIBT) and the outer parts (relOBT), vessel diameter (VDi;
 i.e. vessel lumen diameter, excluding the walls), vessel density (VDe;
 i.e., quantity of vessels per millimetre square), intervessel pit diameter
 (IPD; i.e., the size of pits that link the vessels horizontally), vessel
 element length (VEL), fiber wall thickness (FWT), wood density (WD) and
 maximum tree height (HMax). We classified species as savanna (s), forest
 (f), and generalist (g) species (biome groups). The dataset includes both
 individual and species level information, depending on how the data were
 reported in the literature studies. Literature sources can be found in the
 published article. We also searched the Global
 Biodiversity Information Facility (GBIF) for occurrence data to the subset
 of species for which we had a full trait set (doi: 10.15468/dl.tf5yvv) in
 order to obtain environmental data for these species (the GBIF data is not
 included in this dryad submission). Mean principal component axes and
 environmental information per species are presented for the species with
 available occurrence information (63 species). We excluded occurrence data
 from outside of Brazil (the main region represented in our dataset). For
 the remaining occurrences, we obtained data on mean annual precipitation
 (map), rainfall seasonality (rs), mean annual temperature (mat), fire
 frequency (fire), soil sand content (sand) and soil cation exchange
 capacity (cec). We also created an index for the preference of species for
 shaded environments (shade): a value of 1 was attributed to the least
 shade tolerant species (i.e., savanna species), 3 to the most shade
 adapted species (i.e., forest species) and 2 to generalist species.
 Details can be found in the published article.
description: # Data from: Coordination of bark and wood traits underlies
 forest-to-savanna evolutionary transitions
 [https://doi.org/10.5061/dryad.1ns1rn92j](https://doi.org/10.5061/dryad.1ns1rn92j) This dataset includes functional trait data compiled from 83 published studies conducted in the following tropical South America biomes: moist forest, dry forest and savannas (including flooded savannas). The traits included are bark thickness divided by stem diameter (i.e., relative bark thickness; relBT), discriminating between the inner (relIBT) and the outer parts (relOBT), vessel diameter (VDi; i.e. vessel lumen diameter, excluding the walls), vessel density (VDe; i.e., quantity of vessels per millimetre square), intervessel pit diameter (IPD; i.e., the size of pits that link the vessels horizontally), vessel element length (VEL), fiber wall thickness (FWT), wood density (WD) and maximum tree height (HMax). We classified species as savanna (s), forest (f), and generalist (g) species (biome groups). The dataset includes both individual and species level information, depending on how the data were reported in the literature studies. Literature sources can be found in the published article. We also include mean species scores derived from a principal component analyses on species mean trait values and environmental data for species with a full trait set and available occurrence data in GBIF. ## Description of the data and file structure The first four spreadsheets are Raw Data for bark traits (Bark Traits Raw), secondary xylem traits (Sec Xyl Traits Raw), maximum height (Max Height Raw) and wood density (Wood Dens Raw) and include taxonomic information for all traits (species, genus and family). Reference source (ref) are provided for all traits except maximum height, for which data was obtained from a single reference source. Number of individuals sampled in each study are provided for bark and secondary xylem traits, as the information was not available for other traits. Bark and secondary xylem traits are presented either as individual values or species level means, depending on data availability from the reference source. In many cases, wood density data are presented as means from several references (as this was the format available from the original dataset). The fifth spreadsheet (Data Environmental Analyses) contain mean species scores derived from a principal component analyses on plant functional traits, mean environmental data and mean geographic coordinates for each species. Mean environmental data and geographic coordinates were obtained by mean aggregating the values across all occurrence points per species. In all sheets, the biome at which species occur (biome) and taxonomic information are also reported. Missing data are coded as NA, when a study did not measure a particular trait. Data use: The data can be used in functional trait studies interested in both interspecific and intraspecific trait variability, and plant strategies. Abreviation keys and descriptions relBT: relative bark thickness (proportion of diameter). Description: Bark thickness (mm) divided by stem diameter (mm). relIBT: relative inner bark thickness (proportion of diameter). Description: the thickness of the inner bark part (mm) divided by stem diameter (mm). relOBT: relative outer bark thickness (proportion of diameter). Description: the thickness of the outer bark part (mm) divided by stem diameter (mm). VEL: Vessel Element Length (µm). Description: The length of vessel elements in the vessel. IPD: Intervessel Pit Diameter (µm). Description: The size of pits that link the vessels horizontally. VDi: Vessel Diameter (μm). Description: Vessel lumen diameter, excluding the walls FWT: Fibre Wall Thickness (µm). Description: the thickness of fibre walls. VDe: Vessel Density (mm-2). Description: quantity of vessels per millimetre square. HMax: Maximum Plant Height (m). Description: the maximum height observed for the species in the field. WD: Wood Density (g cm-3). Description: The mass of branches divided by its volume. mat: Mean annual temperature. Description: mean annual temperature from WorldClim2 biovars (Bio_01; Fick & Hijmans, 2017) mean aggregated by species across all occurrence points from GBIF. map: Mean annual precipitation. Description: mean annual precipitation from WorldClim2 biovars (Bio_12; Fick & Hijmans, 2017) mean aggregated by species across all occurrence points from GBIF. rs: Rainfall seasonality (index). Description: rainfall seasonality index from WorldClim2 biovars (Bio_15; Fick & Hijmans, 2017) mean aggregated by species across all occurrence points from GBIF. fire: fire frequency (1985-2020). Description: derived from the MAPBIOMES database (Alencar et al. 2020) mean aggregated by species across all occurrence points from GBIF. cec: cation exchange capacity (cmolc/kg). Description: derived from the SoilGrids250m database (Hengl et al. 2017) mean aggregated by species across all occurrence points from GBIF. sand: soil sand content (g/100g (%)). Description: derived from the SoilGrids250m database (Hengl et al. 2017) mean aggregated by species across all occurrence points from GBIF. shade: shade preference (index; 1: shade avoider; 2: generalist; 3: shade specialist). Classified according to Mendonça et al. (2008) biome: the biome at which the species ocurred (s: savanna; g: generalist; f: forest). Classified according to Mendonça et al. (2008) ni: number of individuals sampled. Description: number of individuals of the species sampled in the study. ref: reference sources. Description: reference source from which the data was obtained (listed in "Data sources"). Organ: plant organ at which secondary xylem traits were measured. PC1-4: principal component axes 1-4. Principal component axes resulting from principal component analyses of species with a full trait set. longitude: mean longitude of species occurrence points. latitide: mean latitude of species occurrence points. obs1: NA in cells refer to missing data, resulting from a lack of information in the literature for that specific trait to that specific species. ## Sharing/Access information Data sources Abe, N., Miatto, R. C., & Batalha, M. A. (2018). Relationships among functional traits define primary strategies in woody species of the Brazilian “cerrado”. *Revista Brasileira de Botanica*, *41*(2), 351–360. doi: 10.1007/s40415-018-0448-x Alencar, A. A., Conciani, D. E., Costa, D. P., Rosa, E. R., Martin, E. V., Hasenack, H., Martenexen, L. F. M., Shimbo, J., Rosa, M., Crusco, N., Ferreira, N. C., Ribeiro, N. V., Santos, S. M. B., Duverger, S. G., Azevedo, T., Piontekowski, V. J., Arruda, V. L. S., Silva, W. V., & Rocha, W. F. (2020). 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