Forest Age Rivals Climate to Explain Reproductive Allocation Patterns in Forest Ecosystems Globally

Rachel E. Ward; Huanyuan Zhang-Zheng; Kate Abernethy; Stephen Adu-Bredu; Luzmilla Arroyo; Andrew Bailey; Jos Barlow; Erika Berenguer; Liana Chesini-Rossi; Percival Cho; Cecilia A.L. Dahlsjö; Eder Carvalho das Neves; Bianca de Oliveira Sales; William Farfan-Rios; Joice Nunes Ferreira; Renata Freitag; Cécile Girardin; Walter Huaraca Huasco; Carlos A. Joly; Yadvinder Malhi; Beatriz Marimon; Ben Hur Marimon Junior; Alexandra C. Morel; Helene C. Muller-Landau; Karine da Silva Peixoto; Simone Reis; Terhi Riutta; Norma Salinas; Marina Seixas; Miles R. Silman; Lara M. Kueppers

doi:10.1111/ele.70191

Forest Age Rivals Climate to Explain Reproductive Allocation Patterns in Forest Ecosystems Globally

Rachel E. Ward (Lead / Corresponding author)
, Huanyuan Zhang-Zheng
, Kate Abernethy
, Stephen Adu-Bredu
, Luzmilla Arroyo
, Andrew Bailey
, Jos Barlow
, Erika Berenguer
, Liana Chesini-Rossi
, Percival Cho
, Cecilia A.L. Dahlsjö
, Eder Carvalho das Neves
, Bianca de Oliveira Sales
, William Farfan-Rios
, Joice Nunes Ferreira
, Renata Freitag
, Cécile Girardin
, Walter Huaraca Huasco
, Carlos A. Joly
, Yadvinder Malhi

Beatriz Marimon, Ben Hur Marimon Junior, Alexandra C. Morel, Helene C. Muller-Landau, Karine da Silva Peixoto, Simone Reis, Terhi Riutta, Norma Salinas, Marina Seixas, Miles R. Silman, Lara M. Kueppers

Energy Environment and Society

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Abstract

Forest allocation of net primary productivity (NPP) to reproduction (carbon required for flowers, fruits, and seeds) is poorly quantified globally, despite its critical role in forest regeneration and a well-supported trade-off with allocation to growth. Here, we present the first global synthesis of a biometric proxy for forest reproductive allocation (RA) across environmental and stand age gradients from a compiled dataset of 824 observations across 393 sites. We find that ecosystem-scale RA increases ~60% from boreal to tropical forests. Climate shows important non-linear relationships with RA, but is not the sole predictor. Forest age effects are comparable to climate in magnitude (MAT: ß = 0.24, p = 0.021; old growth forest: ß = 0.22, p < 0.001), while metrics of soil fertility show small but significant relationships with RA (soil pH: ß = 0.07, p = 0.001; soil N: ß = −0.07, p = 0.001). These results provide strong evidence that ecosystem-scale RA is mediated by climate, forest age, and soil conditions, and is not a globally fixed fraction of positive NPP as assumed by most vegetation and ecosystem models. Our dataset and findings can be used by modellers to improve predictions of forest regeneration and carbon cycling.

Original language	English
Article number	e70191
Journal	Ecology letters
Volume	28
Issue number	8
DOIs	https://doi.org/10.1111/ele.70191
Publication status	Published - 25 Aug 2025

Keywords

climate
ecosystem modelling
forest age
forest ecosystems
forest regeneration
reproductive allocation
soil fertility

ASJC Scopus subject areas

Ecology, Evolution, Behavior and Systematics

UN SDGs

This output contributes to the following UN Sustainable Development Goals (SDGs)

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10.1111/ele.70191Licence: CC BY

Final published version
This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium,provided the original work is properly cited.© 2025 The Author(s). Ecology Letters published by John Wiley & Sons Ltd. This article has been contributed to by U.S. Government employees and their work is in thepublic domain in the USA.
Final published version, 4.67 MBLicence: CC BY

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Ward, R. E., Zhang-Zheng, H., Abernethy, K., Adu-Bredu, S., Arroyo, L., Bailey, A., Barlow, J., Berenguer, E., Chesini-Rossi, L., Cho, P., Dahlsjö, C. A. L., das Neves, E. C., de Oliveira Sales, B., Farfan-Rios, W., Ferreira, J. N., Freitag, R., Girardin, C., Huaraca Huasco, W., Joly, C. A., ... Kueppers, L. M. (2025). Forest Age Rivals Climate to Explain Reproductive Allocation Patterns in Forest Ecosystems Globally. Ecology letters, 28(8), Article e70191. https://doi.org/10.1111/ele.70191

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title = "Forest Age Rivals Climate to Explain Reproductive Allocation Patterns in Forest Ecosystems Globally",

abstract = "Forest allocation of net primary productivity (NPP) to reproduction (carbon required for flowers, fruits, and seeds) is poorly quantified globally, despite its critical role in forest regeneration and a well-supported trade-off with allocation to growth. Here, we present the first global synthesis of a biometric proxy for forest reproductive allocation (RA) across environmental and stand age gradients from a compiled dataset of 824 observations across 393 sites. We find that ecosystem-scale RA increases \textasciitilde{}60\% from boreal to tropical forests. Climate shows important non-linear relationships with RA, but is not the sole predictor. Forest age effects are comparable to climate in magnitude (MAT: {\ss} = 0.24, p = 0.021; old growth forest: {\ss} = 0.22, p < 0.001), while metrics of soil fertility show small but significant relationships with RA (soil pH: {\ss} = 0.07, p = 0.001; soil N: {\ss} = −0.07, p = 0.001). These results provide strong evidence that ecosystem-scale RA is mediated by climate, forest age, and soil conditions, and is not a globally fixed fraction of positive NPP as assumed by most vegetation and ecosystem models. Our dataset and findings can be used by modellers to improve predictions of forest regeneration and carbon cycling.",

keywords = "climate, ecosystem modelling, forest age, forest ecosystems, forest regeneration, reproductive allocation, soil fertility",

author = "Ward, \{Rachel E.\} and Huanyuan Zhang-Zheng and Kate Abernethy and Stephen Adu-Bredu and Luzmilla Arroyo and Andrew Bailey and Jos Barlow and Erika Berenguer and Liana Chesini-Rossi and Percival Cho and Dahlsj{\"o}, \{Cecilia A.L.\} and \{das Neves\}, \{Eder Carvalho\} and \{de Oliveira Sales\}, Bianca and William Farfan-Rios and Ferreira, \{Joice Nunes\} and Renata Freitag and C{\'e}cile Girardin and \{Huaraca Huasco\}, Walter and Joly, \{Carlos A.\} and Yadvinder Malhi and Beatriz Marimon and \{Marimon Junior\}, \{Ben Hur\} and Morel, \{Alexandra C.\} and Muller-Landau, \{Helene C.\} and Peixoto, \{Karine da Silva\} and Simone Reis and Terhi Riutta and Norma Salinas and Marina Seixas and Silman, \{Miles R.\} and Kueppers, \{Lara M.\}",

note = "Publisher Copyright: {\textcopyright} 2025 The Author(s). Ecology Letters published by John Wiley \& Sons Ltd. This article has been contributed to by U.S. Government employees and their work is in the public domain in the USA.",

year = "2025",

month = aug,

day = "25",

doi = "10.1111/ele.70191",

language = "English",

volume = "28",

journal = "Ecology letters",

issn = "1461-023X",

publisher = "Wiley-Blackwell",

number = "8",

}

Ward, RE, Zhang-Zheng, H, Abernethy, K, Adu-Bredu, S, Arroyo, L, Bailey, A, Barlow, J, Berenguer, E, Chesini-Rossi, L, Cho, P, Dahlsjö, CAL, das Neves, EC, de Oliveira Sales, B, Farfan-Rios, W, Ferreira, JN, Freitag, R, Girardin, C, Huaraca Huasco, W, Joly, CA, Malhi, Y, Marimon, B, Marimon Junior, BH, Morel, AC, Muller-Landau, HC, Peixoto, KDS, Reis, S, Riutta, T, Salinas, N, Seixas, M, Silman, MR & Kueppers, LM 2025, 'Forest Age Rivals Climate to Explain Reproductive Allocation Patterns in Forest Ecosystems Globally', Ecology letters, vol. 28, no. 8, e70191. https://doi.org/10.1111/ele.70191

TY - JOUR

T1 - Forest Age Rivals Climate to Explain Reproductive Allocation Patterns in Forest Ecosystems Globally

AU - Ward, Rachel E.

AU - Zhang-Zheng, Huanyuan

AU - Abernethy, Kate

AU - Adu-Bredu, Stephen

AU - Arroyo, Luzmilla

AU - Bailey, Andrew

AU - Barlow, Jos

AU - Berenguer, Erika

AU - Chesini-Rossi, Liana

AU - Cho, Percival

AU - Dahlsjö, Cecilia A.L.

AU - das Neves, Eder Carvalho

AU - de Oliveira Sales, Bianca

AU - Farfan-Rios, William

AU - Ferreira, Joice Nunes

AU - Freitag, Renata

AU - Girardin, Cécile

AU - Huaraca Huasco, Walter

AU - Joly, Carlos A.

AU - Malhi, Yadvinder

AU - Marimon, Beatriz

AU - Marimon Junior, Ben Hur

AU - Morel, Alexandra C.

AU - Muller-Landau, Helene C.

AU - Peixoto, Karine da Silva

AU - Reis, Simone

AU - Riutta, Terhi

AU - Salinas, Norma

AU - Seixas, Marina

AU - Silman, Miles R.

AU - Kueppers, Lara M.

N1 - Publisher Copyright: © 2025 The Author(s). Ecology Letters published by John Wiley & Sons Ltd. This article has been contributed to by U.S. Government employees and their work is in the public domain in the USA.

PY - 2025/8/25

Y1 - 2025/8/25

N2 - Forest allocation of net primary productivity (NPP) to reproduction (carbon required for flowers, fruits, and seeds) is poorly quantified globally, despite its critical role in forest regeneration and a well-supported trade-off with allocation to growth. Here, we present the first global synthesis of a biometric proxy for forest reproductive allocation (RA) across environmental and stand age gradients from a compiled dataset of 824 observations across 393 sites. We find that ecosystem-scale RA increases ~60% from boreal to tropical forests. Climate shows important non-linear relationships with RA, but is not the sole predictor. Forest age effects are comparable to climate in magnitude (MAT: ß = 0.24, p = 0.021; old growth forest: ß = 0.22, p < 0.001), while metrics of soil fertility show small but significant relationships with RA (soil pH: ß = 0.07, p = 0.001; soil N: ß = −0.07, p = 0.001). These results provide strong evidence that ecosystem-scale RA is mediated by climate, forest age, and soil conditions, and is not a globally fixed fraction of positive NPP as assumed by most vegetation and ecosystem models. Our dataset and findings can be used by modellers to improve predictions of forest regeneration and carbon cycling.

AB - Forest allocation of net primary productivity (NPP) to reproduction (carbon required for flowers, fruits, and seeds) is poorly quantified globally, despite its critical role in forest regeneration and a well-supported trade-off with allocation to growth. Here, we present the first global synthesis of a biometric proxy for forest reproductive allocation (RA) across environmental and stand age gradients from a compiled dataset of 824 observations across 393 sites. We find that ecosystem-scale RA increases ~60% from boreal to tropical forests. Climate shows important non-linear relationships with RA, but is not the sole predictor. Forest age effects are comparable to climate in magnitude (MAT: ß = 0.24, p = 0.021; old growth forest: ß = 0.22, p < 0.001), while metrics of soil fertility show small but significant relationships with RA (soil pH: ß = 0.07, p = 0.001; soil N: ß = −0.07, p = 0.001). These results provide strong evidence that ecosystem-scale RA is mediated by climate, forest age, and soil conditions, and is not a globally fixed fraction of positive NPP as assumed by most vegetation and ecosystem models. Our dataset and findings can be used by modellers to improve predictions of forest regeneration and carbon cycling.

KW - climate

KW - ecosystem modelling

KW - forest age

KW - forest ecosystems

KW - forest regeneration

KW - reproductive allocation

KW - soil fertility

UR - https://www.scopus.com/pages/publications/105014522858

U2 - 10.1111/ele.70191

DO - 10.1111/ele.70191

M3 - Review article

C2 - 40853682

AN - SCOPUS:105014522858

SN - 1461-023X

VL - 28

JO - Ecology letters

JF - Ecology letters

IS - 8

M1 - e70191

ER -

Forest Age Rivals Climate to Explain Reproductive Allocation Patterns in Forest Ecosystems Globally

Abstract

Keywords

ASJC Scopus subject areas

UN SDGs

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