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Assembly of functional diversity in an oceanic island flora


Oceanic island floras are well known for their morphological peculiarities and exhibit striking examples of trait evolution1,2,3. These morphological shifts are commonly attributed to insularity and are thought to be shaped by the biogeographical processes and evolutionary histories of oceanic islands2,4. However, the mechanisms through which biogeography and evolution have shaped the distribution and diversity of plant functional traits remain unclear5. Here we describe the functional trait space of the native flora of an oceanic island (Tenerife, Canary Islands, Spain) using extensive field and laboratory measurements, and relate it to global trade-offs in ecological strategies. We find that the island trait space exhibits a remarkable functional richness but that most plants are concentrated around a functional hotspot dominated by shrubs with a conservative life-history strategy. By dividing the island flora into species groups associated with distinct biogeographical distributions and diversification histories, our results also suggest that colonization via long-distance dispersal and the interplay between inter-island dispersal and archipelago-level speciation processes drive functional divergence and trait space expansion. Contrary to our expectations, speciation via cladogenesis has led to functional convergence, and therefore only contributes marginally to functional diversity by densely packing trait space around shrubs. By combining biogeography, ecology and evolution, our approach opens new avenues for trait-based insights into how dispersal, speciation and persistence shape the assembly of entire native island floras.

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Fig. 1: Trait space of the native flora of Tenerife fits within the global trait space and is subject to similar constraints to the global trait space but is biased towards medium-statured species with intermediate trait values (shrubby functional hotspot).
Fig. 2: Imprint of biogeography and evolution on the trait space of Tenerife native flora.
Fig. 3: Contribution of biogeographical and evolutionary processes to the functional diversity of the native flora of Tenerife.

Data availability

The trait data and floristic information of the species are available in a Figshare repository ( Source data are provided with this paper.

Code availability

The analysis and data visualization performed in R software are available in a Figshare repository (


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M.P.B.B. and H.K. acknowledge funding from the German Research Foundation (DFG) Research Training Group 1644 ‘Scaling Problems in Statistics’, grant no. 152112243. D.C. acknowledges funding from the Agencia Nacional de Investigación y Desarrollo (Chile; FONDECYT Regular no. 1201347). We thank the Jardín Botánico Canario ‘Viera y Clavijo’ in Gran Canaria for allowing measurements in the seed bank as well as for the samples taken of rare species that were not possible to find in the field; N. Straßburger, M. V. Rodríguez, A. A. Diez, B. Bhattarai, M. Mulligan and W. Osterman for invaluable assistance in the field and in the laboratory; B. Ø. Hansen for his helpful programming advice; R. Barone for assisting with plant identification and fieldwork; and G. Ottaviani, P. Gaüzère and L. Valente for providing valuable comments that improved our work immensely.

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Authors and Affiliations



H.K. conceived the initial idea. M.P.B.B., D.C., P.W. and H.K. further developed the concepts and designed the research. M.P.B.B. collected and measured plant traits and performed the statistical analysis. D.C. and P.D. contributed to the statistical analysis. R.O. and J.M.F.-P. supported the fieldwork logistics and execution, that is, plant trait data collection and species identification. J.P. curated lineage information. S.D. supported trait data curation and writing the discussion. All authors contributed to the interpretation of the results and the writing of the paper.

Corresponding author

Correspondence to Martha Paola Barajas Barbosa.

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The authors declare no competing interests.

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Nature thanks Pierre Gaüzère, Gianluigi Ottaviani and Luis Valente for their contribution to the peer review of this work. Peer reviewer reports are available.

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Extended data figures and tables

Extended Data Fig. 1 Trait data overview and assessment.

a. Present empirical data and missing data distribution across 100% (n = 436) of Tenerife native seed plant species. b. Present empirical data and missing data distribution across 80% (n = 348, i.e., species with minimum of five trait values) of Tenerife native seed plant species, and c. across five species groups included in the main analysis. The missing trait data represented in black, were filled using phylogenetically informed imputation (see Methods). Percentages reported for each trait refer to the proportion of missing data. d. Density distribution of the empirical trait values only, and including imputed trait values for each trait. Percentages correspond to the proportion of species with imputed trait values.

Extended Data Fig. 2 Overview of the principal components explained variance of both island and global trait data, and hypervolume overlap.

a. Percentage of explained variance of the six dimensions of the principal component analysis (PCA) based on six plant functional traits of Tenerife’s native seed plants and seed plants from across the globe (348 Tenerife and 2,274 global species). b. Percentage of contributions of each trait to each dimension of the PCA. Trait values were z- and log-transformed prior to the PCA. c. Tenerife species projected onto the global spectrum of plant form and function21, contour lines indicating 0.5, 0.95 and 0.99 quantiles using kernel density estimation; PCA protection following74. Note that Tenerife native flora falls within the 99% isoline space of the global trait space. d. Estimated overlap, i.e., Sørensen index, between the island and global trait space estimated via null model (where n = 300 species randomly sampled, from island and global data separately, with replacement 999 times) based on hypervolume approach (see details in Methods). Null model 1 controls for a potential underrepresentation of shrubs in the global trait space by sampling the current proportions of growth form sensu72 (see Methods) and for the species richness difference between datasets. Null model 2 controls only for the species richness difference between both global and Tenerife datasets. In d. dots and error bars (not visible here as they are close to the mean) correspond to the mean values and 95% confidence intervals.

Extended Data Fig. 3 Overview of principal components analysis (PCA) for eight plant functional traits (n = 348 species) of Tenerife native seed plants.

a. Percentage of explained variance of the eight dimensions of the PCA. b. Percentage of contributions of each trait to each dimension of the PCA. Trait values were log- and scaled. c. Differences among the first and second principal component of the trait spaces of different biogeographical groups, Non-endemic native species (NEN), Macaronesian endemic species (MAC), Canary Islands endemic species (CE), Tenerife endemic species (TE) and cladogenetic species (CLA). Identical letters indicate no significant differences (Kruskal-Wallis text, P>0.05). Data are represented as boxplots where the middle line is the median, the lower and upper hinges correspond to the 25th and 75th percentiles, upper and lower whiskers extend from the hinge to the largest and lowest value, respectively, no further than 1.5 * IQR from the hinge.

Extended Data Fig. 4 Tenerife climatic representativity mapped on Whittaker biomes.

a. and b. field sites visited in 2017 and 2018 (n = 500), where plant material was collected to measure plant functional traits for this study.

Extended Data Fig. 5 Hot clades of endemic species of the Macaronesia (green), Tenerife (blue) and of native non-endemic species (black) are highlighted.

Hot nodes indicate clades with more species than expected at random. Node corresponds to standard effect size and node transparency is scaled by the number of sample trees in which a node was significantly overdispersed. Nodes include at least 10 species and occur in at least 50 sample trees. Note that the vast majority of hot nodes are among Tenerife endemic species.

Extended Data Fig. 6 Functional contribution and originality of the major radiated plant lineages present in Tenerife.

a. Functional contribution and b. functional originality for the 63 radiated lineages. Note that only Aeonium alliance and Polycarpaea lineages (marked with *) are significantly contributing to the expansion of Tenerife trait space, as their functional contribution or originality are significantly different from other species that do not belong to the lineage (results based on Kruskal-Wallis, P < 0.05). Each box plot in magenta colour correspond to a lineage; number of species on a lineage (i.e., n numbers) are the same in a-b. Each box plot in grey colour corresponds to all island species, except for species included in the corresponding compared lineage. In a–b data are represented as boxplots where the middle line is the median, the lower and upper hinges correspond to the 25th and 75th percentiles, upper and lower whiskers extend from the hinge to the largest and lowest value, respectively, no further than 1.5 * IQR from the hinge. c. Location of Aeonium alliance and Polycarpaea species in the island trait space; lineages extend it towards high values of leaf thickness. d. Number of radiated lineages present in Tenerife (31), Canary Islands (54) and Macaronesia (14). The 63 major lineages are composed of 195 species or 56% of all species included in the main analysis. The 63 lineages are nested within the Macaronesia, Canary Islands and Tenerife.

Extended Data Fig. 7 Sensitivity analyses using only cases (n = 237 species) with complete empirical data for all 8 traits.

a. Trait spaces for the native flora of Tenerife and dissected into five distinct species groups that illustrate the imprint of biogeography and evolution on the functional diversity of an oceanic island flora. b. Functional richness, functional dispersion and functional evenness were calculated using n-dimensional hypervolumes while controlling for the number of species richness for the five groups and for a null group in grey, where dots and error bars correspond to the mean values and 95% confidence intervals, based on null models (top). Functional contribution and originality of each group with respect to the rest of island species (bottom); data is represented as boxplots where the middle line is the median, the lower and upper hinges correspond to the 25th and 75th percentiles, upper and lower whiskers extend from the hinge to the largest and lowest value, respectively, no further than 1.5 * IQR from the hinge. See details in Methods.

Extended Data Table 1 Validation of trait values imputation
Extended Data Table 2 Phylogenetic signal of empirical trait values and phylogenetic signal of missing trait values

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Barajas Barbosa, M.P., Craven, D., Weigelt, P. et al. Assembly of functional diversity in an oceanic island flora. Nature 619, 545–550 (2023).

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