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Wet and dry tropical forests show opposite successional pathways in wood density but converge over time


Tropical forests are converted at an alarming rate for agricultural use and pastureland, but also regrow naturally through secondary succession. For successful forest restoration, it is essential to understand the mechanisms of secondary succession. These mechanisms may vary across forest types, but analyses across broad spatial scales are lacking. Here, we analyse forest recovery using 1,403 plots that differ in age since agricultural abandonment from 50 sites across the Neotropics. We analyse changes in community composition using species-specific stem wood density (WD), which is a key trait for plant growth, survival and forest carbon storage. In wet forest, succession proceeds from low towards high community WD (acquisitive towards conservative trait values), in line with standard successional theory. However, in dry forest, succession proceeds from high towards low community WD (conservative towards acquisitive trait values), probably because high WD reflects drought tolerance in harsh early successional environments. Dry season intensity drives WD recovery by influencing the start and trajectory of succession, resulting in convergence of the community WD over time as vegetation cover builds up. These ecological insights can be used to improve species selection for reforestation. Reforestation species selected to establish a first protective canopy layer should, among other criteria, ideally have a similar WD to the early successional communities that dominate under the prevailing macroclimatic conditions.

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Data availability

The dataset on CWM WD values and WD ranges for the plots from 49 sites (Fig. 1) is available from the Data Archiving and Networked Services repository at For one other site, data are available on request. The data used to produce Fig. 2 can be found in Supplementary Table 1.


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This paper is a product of the 2ndFOR collaborative research network on secondary forests ( We thank the owners of the secondary forest sites and local communities for access to their forests, everyone who established and measured the plots, and the institutions and funding agencies that supported them. We thank the following agencies for financial support: the Netherlands Organisation for Scientific research (NWO-ALW.OP241 and NWO-ALW 863.15.017), Wageningen University and Research (INREF FOREFRONT and Terra Preta programmes, and the Strategic Investment Programme Resilience), a Macquarie University Visiting Researcher grant to L.P., the Biological Dynamics of Forest Fragments Project, CIFOR, COLCIENCIAS grant 1243-13-16640, CONACYT PhD grant 169510, Conselho Nacional de Desenvolvimento Científico e Tecnológico (grant numbers 481576/2009-6, 304817/2015-5, 309874/2015-7, 306375/2016-8 and 308471/2017-2), Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (grant number 88881.064976/2014-01), the German Academic Exchange Service, FAPEMIG, the Garden Club of America Award in Tropical Botany, the Global Environment Facility (grant VEN/SGP/2010-2015), ICETEX, Instituto Nacional de Investigaciones Agrícolas, Instituto Nacional de Ciência e Tecnologia dos Serviços Ambientais da Amazônia, the International Climate Initiative of the German Federal Ministry for the Environment, Nature Conservation, Building and Nuclear Safety, the Lewis and Clark Fund for Exploration and Field Research, NASA ROSES grant NNH08ZDA001N-TE, the Norwegian Agency for Development Cooperation, NUFFIC, Yale-NUS College (grant R-607-265-054-121), FOMIX-Yucatan (YUC-2008-C06-108863), OTS and the Christiane and Christopher Tyson Fellowship, PAPIIT-UNAM (218416 and 211114), the Ronald Bamford Endowment, the São Paulo Research Foundation (grant numbers 2011/06782-5 and 2014/14503-7), SENACYT grant COL10-052, SEP-CONACYT (CB-2009-128136 and CB 2015- 255544), Stichting Het Kronendak, Tropenbos International, the United Nations Development Program (Venezuela) and the US National Science Foundation (DEB-0639114, DEB-1147434, DEB-0424767, DEB-0639393, DEB-1147429, DEB- 1110722, NSF-0946618 and NSF-9208031). This is publication number 752 in the Technical Series of the Biological Dynamics of Forest Fragments Project BDFFP-INPA-SI and publication number 5 from 2ndFOR.

Author information

L.P. and M.W. conceived the idea. L.P. and D.M.A.R. coordinated the data compilations. D.M.A.R. carried out the data analysis. L.P., D.M.A.R., F.B. and M.W. contributed to the analytical tools used in the analysis. E.N.B. and A.M.A.Z. made the map. L.P. wrote the paper. All co-authors collected field data, discussed the results, suggested further analyses and commented on the manuscript.

Correspondence to Lourens Poorter.

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Fig. 1: Recovery of WD values and WD variation with time since abandonment in Neotropical secondary forest sites.
Fig. 2: CWM WD5y and successional changes in CWM WD as a function of CWA.