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Old and ancient trees are life history lottery winners and vital evolutionary resources for long-term adaptive capacity


Trees can live for many centuries with sustained fecundity and death is largely stochastic. We use a neutral stochastic model to examine tree demographic patterns that emerge over time, across a range of population sizes and empirically observed mortality rates. A small proportion of trees (~1% at 1.5% mortality) are life-history ‘lottery winners’, achieving ages >10–20× the median age. Maximum age increases with bigger populations and lower mortality rates. One-quarter of trees (~24%) achieve ages that are three to four times greater than the median age. Three age classes (mature, old and ancient) contribute unique evolutionary diversity across complex environmental cycles. Ancient trees are an emergent property of forests that requires many centuries to generate. They radically change variance in generation time and population fitness, bridging centennial environmental cycles. These life-history ‘lottery’ winners are vital to long-term forest adaptive capacity and provide invaluable data about environmental history and individual longevity. Old and ancient trees cannot be replaced through restoration or regeneration for many centuries. They must be protected to preserve their invaluable diversity.

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Fig. 1: Maximum tree and forest climax age, given different population sizes and annual stochastic mortality rates.
Fig. 2: Mean age and standard deviation of three age classes, given different mortality rates and population sizes, across 25 replicates.
Fig. 3: Mean and absolute maximum age achieved across 25 replicates over 15,000 years, given different population sizes and annual mortality.
Fig. 4: Population demographics in relation to changing environmental selection pressure (Supplementary Fig. 4), given an annual mortality rate of 1.5%.
Fig. 5: Major mechanisms evolved by ancient trees to defy ageing.
Fig. 6: Canopy rejuvenation and seed production in old trees.

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Data availability

The data used in the analyses were generated from the code provided. The authors strongly recommend that the data be generated directly from the code to demonstrate the reliability of the methods and to verify that the results are the same in different computing environments and settings. Computation time to generate the dataset was less than 8 h on a MacBook Pro with 2.5 Ghz Dual-Core Intel Core i7 processor and 16 GB of RAM. The original dataset used in this publication is available upon request from the corresponding author.

Code availability

The models and analyses were written in the Mathematica 12 programming environment. A notebook containing the simulation model and the code used to produce the results and figures has been uploaded to the publication website. Code can also be provided upon request by the corresponding author.


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C.H.C. thanks J. R. Harting for discussions about model development during the very early stages of this project. Research in S.M.-B. laboratory is supported by the PID2019-110335GB-I00 /AEI grant from the Spanish Government and through an ICREA Academia award and 2017 SGR 980 grant from the Catalan Government. G.P.’s research on sessile oak and beech longevity was funded respectively by Aspromonte and Pollino National Parks.

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C.H.C. and S.M.-B. conceived the idea. C.H.C. conceived of and wrote the simulation models, performed the analysis and wrote the manuscript. G.P. and S.M.-B. helped improve the analysis and wrote the manuscript.

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Correspondence to Charles H. Cannon.

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Nature Plants thanks Josep Penuelas and the other, anonymous, reviewer(s) for their contribution to the peer review of this work.

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Supplementary Figs. 1–3.

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Cannon, C.H., Piovesan, G. & Munné-Bosch, S. Old and ancient trees are life history lottery winners and vital evolutionary resources for long-term adaptive capacity. Nat. Plants 8, 136–145 (2022).

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