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ACKNOWLEDGMENTS
We thank H. Okur and H. B. de Aguiar for helpful discussions.
S.R. thanks the Julia Jacobi Foundation.Funding:This work was
supported by the Swiss National Science Foundation (grant
200021-182606-1 to S.P., S.K., B.R., and S.R.).Author
contributions:S.P., S.K., and B.R. performed the experiments.
S.P., S.R., S.K., B.R., and A.H. interpreted the data. A.H. performed
the simulations. S.R. conceived and supervised the study. S.R.,
S.P., A.H., B.R., and S.K. wrote the manuscript.Competing
interests:The authors declare no competing interests.Data and
materials availability:All data in the manuscript and
supplementary materials are available through Zenodo ( 36 ).SUPPLEMENTARY MATERIALS
science.org/doi/10.1126/science.abj3007
Materials and Methods
Supplementary Text
Figs. S1 to S7
Table S1
References ( 37 – 60 )
4 May 2021; accepted 19 October 2021
10.1126/science.abj3007FOREST ECOLOGY
Multidimensional tropical forest recovery
Lourens Poorter^1 *, Dylan Craven^2 , Catarina C. Jakovac1,3, Masha T. van der Sande^1 , Lucy Amissah^4 ,
Frans Bongers^1 , Robin L. Chazdon5,6, Caroline E. Farrior^7 , Stephan Kambach^8 ,JorgeA.Meave^9 ,
Rodrigo Muñoz1,9, Natalia Norden^10 , Nadja Rüger8,11,12, Michiel van Breugel13,14,15,
Angélica María Almeyda Zambrano^16 , Bienvenu Amani^17 , José Luis Andrade^18 , Pedro H. S. Brancalion^19 ,
Eben N. Broadbent^20 , Hubert de Foresta^21 , Daisy H. Dent12,22, Géraldine Derroire^23 , Saara J. DeWalt^24 ,
Juan M. Dupuy^18 , Sandra M. Durán25,26, Alfredo C. Fantini^27 , Bryan Finegan^28 , Alma Hernández-Jaramillo^29 ,
José Luis Hernández-Stefanoni^18 , Peter Hietz^30 , André B. Junqueira^31 , Justin Kassi N’dja^32 ,
Susan G. Letcher^33 , Madelon Lohbeck1,34, René López-Camacho^35 , Miguel Martínez-Ramos^36 ,
Felipe P. L. Melo^37 , Francisco Mora^36 , Sandra C. Müller^38 ,AnnyE.N’Guessan^32 , Florian Oberleitner^39 ,
Edgar Ortiz-Malavassi^40 , Eduardo A. Pérez-García^9 , Bruno X. Pinho^37 , Daniel Piotto^41 ,
Jennifer S. Powers42,43, Susana Rodríguez-Buriticá^10 , Danaë M. A. Rozendaal44,45, Jorge Ruíz^46 ,
Marcelo Tabarelli^37 , Heitor Mancini Teixeira44,47,48, Everardo Valadares de Sá Barretto Sampaio^49 ,
Hans van der Wal^50 , Pedro M. Villa51,52, Geraldo W. Fernandes^53 , Braulio A. Santos^54 , José Aguilar-Cano^10 ,
Jarcilene S. de Almeida-Cortez^55 , Esteban Alvarez-Davila^56 , Felipe Arreola-Villa^36 , Patricia Balvanera^36 ,
Justin M. Becknell^57 , George A. L. Cabral^37 , Carolina Castellanos-Castro^10 ,BenH.J.deJong^58 ,
Jhon Edison Nieto^10 , Mário M. Espírito-Santo^59 , Maria C. Fandino^60 , Hernando García^10 ,
Daniel García-Villalobos^10 , Jefferson S. Hall^13 , Alvaro Idárraga^61 , Jaider Jiménez-Montoya^62 ,
Deborah Kennard^63 , Erika Marín-Spiotta^64 , Rita Mesquita^65 ,YuleR.F.Nunes^59 , Susana Ochoa-Gaona^58 ,
Marielos Peña-Claros^1 , Nathalia Pérez-Cárdenas^36 , Jorge Rodríguez-Velázquez^36 ,
Lucía Sanaphre Villanueva66,18, Naomi B. Schwartz^67 , Marc K. Steininger^68 , Maria D. M. Veloso^59 ,
Henricus F. M. Vester^69 , Ima C. G. Vieira^70 , G. Bruce Williamson65,71, Kátia Zanini^38 , Bruno Hérault72,73,74
Tropical forests disappear rapidly because of deforestation, yet they have the potential to regrow
naturally on abandoned lands. We analyze how 12 forest attributes recover during secondary succession
and how their recovery is interrelated using 77 sites across the tropics. Tropical forests are highly
resilient to low-intensity land use; after 20 years, forest attributes attain 78% (33 to 100%) of their
old-growth values. Recovery to 90% of old-growth values is fastest for soil (<1 decade) and plant
functioning (<2.5 decades), intermediate for structure and species diversity (2.5 to 6 decades), and
slowest for biomass and species composition (>12 decades). Network analysis shows three independent
clusters of attribute recovery, related to structure, species diversity, and species composition.
Secondary forests should be embraced as a low-cost, natural solution for ecosystem restoration,
climate change mitigation, and biodiversity conservation.
T
ropical forests are converted at alarm-
ing rates to other land uses ( 1 ), yet they
also have the potential to regrow natu-
rally on abandoned agricultural fields and
pastures. Widespread land abandonment
because of fertility loss, migration, or alter-
native livelihood options has led to a rapid
increase in the extent of regrowing forests.
Currently, regrowth covers as much as 28%
(2.4 million km^2 ) of the neotropics alone ( 2 ).
Regrowing secondary forests (SFs) form a
large and important component of human-
modified tropical landscapes and have the
potential to play a key role in biodiversity
conservation ( 3 ), climate change mitigation
( 2 ), and landscape restoration ( 4 ). A holistic,
quantitative understanding of the recovery
of multiple SF functions is needed to inform
and design effective policies that benefit na-
ture and people from local to global scales.
In this study, we assess the resilience of 12
forest attributes to recover from agricultureand pasture use. Resilience is the ability of
a system to absorb disturbances and return
to its previous state ( 5 ). Resilience is driven
by two underlying components: the ability
to resist disturbance and the ability to re-
cover after disturbance ( 6 ). We defined“re-
sistance”as the difference between the value
of the forest attribute at the start of suc-
cession and the average old-growth forest
(OGF) values [compare ( 7 )], which reflects
the combined legacies of previous forest and
previous land use, and“recovery”as the abil-
ity to return to OGF attribute values after
succession. Succession is defined as a change
in vegetation structure, species composition
(SC), and ecosystem functioning over time
after a disturbance ( 8 ). Secondary succession
occurs on previously vegetated lands when
a disturbance removes most of the above-
ground vegetation and can proceed at fast
rates due to legacy effects of previous forest
or previous land use, such as a developed
soil, seed bank, remnant trees, and resprout-
ing stumps. Successional pathways are, to
some extent, predictable but, because of local
stochastic factors, are also, to some extent,
uncertain ( 9 ).
Most successional theories have focused on
specific ecosystem attributes, such as SC ( 10 ),
species richness (SR) ( 11 ), forest structure ( 12 ),
or soils ( 13 ), but they have rarely been con-
ceptually integrated and assessed together.
Recovery of these different ecosystem attrib-
utes (i.e., dimensions) is likely to depend on
one another. For example, rapid recovery of
biomass may lead to high litter production
and decomposition and, hence, rapid recovery
of soil organic carbon.
Chronosequence studies allow us to infer
long-term trends in forest recovery by com-
paring forests with similar land-use history
that differ in age since agricultural or pas-
ture abandonment. Single-site studies have
assessed the recovery of multiple attributes
[summarized in ( 14 )], and several synthetic
analyses have assessed the recovery of single
attributes. They have found that ecosystem
functioning, such as nitrogen fixation, re-
covers fast [in about three decades ( 15 )],1370 10 DECEMBER 2021•VOL 374 ISSUE 6573 science.orgSCIENCE
RESEARCH | RESEARCH ARTICLES