Frans Bongers - Academia.edu (original) (raw)

Papers by Frans Bongers

Journal of Tropical Ecology, 1999

Forest Ecology and Management, Mar 1, 2004

Forest Ecology and Management

University of Exeter, 2021

ABSTRACT: The sensitivity of tropical forest carbon to climate is a key uncertainty in predicting... more ABSTRACT: The sensitivity of tropical forest carbon to climate is a key uncertainty in predicting global climate change. Although short-term drying and warming are known to affect forests, it is unknown if such effects translate into long-term responses. Here, we analyze 590 permanent plots measured across the tropics to derive the equilibrium climate controls on forest carbon. Maximum temperature is the most important predictor of aboveground biomass (−9.1 megagrams of carbon per hectare per degree Celsius), primarily by reducing woody productivity, and has a greater rate of decline in the hottest forests (>32.2°C). Our results nevertheless reveal greater thermal resilience than observations of short-term variation imply. To realize the long-term climate adaptation potential of tropical forests requires both protecting them and stabilizing Earth's climate.

There is consensus that plant diversity and ecosystem processes are negatively affected by land-u... more There is consensus that plant diversity and ecosystem processes are negatively affected by land-use intensification (LUI), but, at the same time, there is empirical evidence that a large heterogeneity can be found in the responses. This heterogeneity is especially poorly understood in tropical ecosystems. We evaluated changes in community functional properties across five common land-use types in the wet tropics with different land-use intensity: mature forest, logged forest, secondary forest, agricultural land, and pastureland, located in the lowlands of Bolivia. For the dominant plant species, we measured 12 functional response traits related to their life history, acquisition and conservation of resources, plant domestication, and breeding. We used three single-trait metrics to describe community functional properties: community abundance-weighted mean (CWM) traits values, coefficient of variation, and kurtosis of distribution. The CWM of all 12 traits clearly responded to LUI. Overall, we found that an increase in LUI resulted in communities dominated by plants with acquisitive leaf trait values. However, contrary to our expectations, secondary forests had more conservative trait values (i.e., lower specific leaf area) than mature and logged forest, probably because they were dominated by palm species. Functional variation peaked at intermediate land-use intensity (high coefficient of variation and low kurtosis), which included secondary forest but, unexpectedly, also agricultural land, which is an intensely managed system. The high functional variation of these systems is due to a combination of how response traits (and species) are filtered out by biophysical filters and how management practices introduced a range of exotic species and their trait values into the local species pool. Our results showed that, at local scales and depending on prevailing environmental and management practices, LUI does not necessarily result in communities with more acquisitive trait values or with less functional variation. Instead of the widely expected negative impacts of LUI on plant diversity, we found varying responses of functional variation, with possible repercussions on many ecosystem services. These findings provide a background for actively mitigating negative effects of LUI while meeting the needs of local communities that rely mainly on provisioning ecosystem services for their livelihoods

Tropical forests are global centres of both biodiversity and carbon storage. Many tropical countr... more Tropical forests are global centres of both biodiversity and carbon storage. Many tropical countries aspire to protect forest to fulfil biodiversity and climate mitigation policy targets, but the conservation strategies needed to achieve these two functions depend critically on the tropical forest diversity-carbon relationship and this remains largely unexplored. Attempts to assess and understand this relationship in tropical forest ecosystems have been hindered by the scarcity of inventories where carbon storage in aboveground biomass and species identifications have been simultaneously and robustly quantified. Here, we compile a unique pan-tropical dataset of 360 plots located in old-growth closed-canopy forest, surveyed using standardised methods, allowing a multi-scale evaluation of the relationship between carbon storage and tree diversity. We find strongly contrasting variation in diversity and carbon among continents. Thus, on average, African forests have high carbon storage...

Nature Ecology & Evolution, 2021

Forest restoration increases global forest area and ecosystem services such as primary productivi... more Forest restoration increases global forest area and ecosystem services such as primary productivity and carbon storage. How tree species functional composition impacts the provisioning of these services as forests develop is sparsely studied. We used 10-year data from 478 plots with 191,200 trees in a forest biodiversity experiment in subtropical China to assess the relationship between community productivity and community-weighted mean (CWM) or functional diversity (FD) values of 38 functional traits. We found that effects of FD values on productivity became larger than effects of CWM values after 7 years of forest development and that the FD values also became more reliable predictors of productivity than the CWM values. In contrast to CWM, FD values consistently increased productivity across ten different species-pool subsets. Our results imply that to promote productivity in the long term it is imperative for forest restoration projects to plant multispecies communities with large functional diversity.

Frontiers in Forests and Global Change, 2020

Tropical Conservation Science, 2018

Forest Ecology and Management, 2017

Science, May 8, 2015

TROPICAL FORESTS ARE increasingly modified by human activities. Centuries of human–forest interac... more TROPICAL FORESTS ARE increasingly modified by human activities. Centuries of human–forest interactions have led to a diverse array of forest areas in different phases of succession. In recent decades, forest conversion to cattle pasture or agricultural fields, followed by land abandonment, has led to large areas of second-growth forest in the Amazon. These forests grow rapidly and sequester large amounts of carbon in their biomass, but they tend to be ignored, as most of the debate on the carbon balance of the Amazon basin tends to revolve around old-growth forests. For example, a recent study has shown that the net carbon uptake of Amazonian old-growth forests has declined by a third per decade from 1990 to 2010 ([ 1 ][1], [ 2 ][2]). When extrapolated over the whole Amazon basin, these results translate into a reduced role of intact tropical forests in climate change mitigation. This alarming conclusion, however, completely ignores the important role of regenerating forests as carbon sinks. For instance, in 2010 about 25% of formerly deforested areas in Para, Brazil, were occupied by second-growth forests. Although re-growing forests have lower carbon stocks (45 to 48% of old growth forest), their net carbon sequestration rate is up to 20 times higher (4.6 to 5.8 Mg carbon ha−1 year−1) ([ 3 ][3]) than old-growth forests ([ 1 ][1]). Additionally, about one-quarter of the forests in the Amazon basin are managed for timber production. Net carbon sequestration rates after timber extraction are high, and the application of reduced-impact logging techniques further increases carbon sequestration rates (2.8 Mg ha−1 year−1 compared with 0.5 for conventionally logged areas) ([ 4 ][4]). Consequently, it is essential to incorporate the carbon sequestration potential of second-growth, logged, and managed forests in future assessments of the Amazon basin as a global carbon sink. 1. [↵][5] 1. R. J. W. Brienen 2. et al ., Nature 519, 344 (2015). [OpenUrl][6][CrossRef][7][GeoRef][8][PubMed][9] 2. [↵][10] , Amazon rainforest ability to soak up carbon dioxide is falling (2015); . 3. [↵][11] 1. Y. Pan 2. et al ., Science 333, 988 (2011). [OpenUrl][12][Abstract/FREE Full Text][13] 4. [↵][14] 1. T. A. P. West, 2. E. Vidal, 3. F. E. Putz , Forest Ecol. Manage. 314, 59 (2014). [OpenUrl][15][CrossRef][16] [1]: #ref-1 [2]: #ref-2 [3]: #ref-3 [4]: #ref-4 [5]: #xref-ref-1-1 "View reference 1 in text" [6]: {openurl}?query=rft.jtitle%253DNature%26rft.stitle%253DNature%26rft.issn%253D0028-0836%26rft.aulast%253DBrienen%26rft.auinit1%253DR.%2BJ.%2BW.%26rft.volume%253D519%26rft.issue%253D7543%26rft.spage%253D344%26rft.epage%253D348%26rft.atitle%253DLong-term%2Bdecline%2Bof%2Bthe%2BAmazon%2Bcarbon%2Bsink%26rft_id%253Dinfo%253Adoi%252F10.1038%252Fnature14283%26rft_id%253Dinfo%253Apmid%252F25788097%26rft.genre%253Darticle%26rft_val_fmt%253Dinfo%253Aofi%252Ffmt%253Akev%253Amtx%253Ajournal%26ctx_ver%253DZ39.88-2004%26url_ver%253DZ39.88-2004%26url_ctx_fmt%253Dinfo%253Aofi%252Ffmt%253Akev%253Amtx%253Actx [7]: /lookup/external-ref?access_num=10.1038/nature14283&link_type=DOI [8]: /lookup/external-ref?access_num=2015036225&link_type=GEOREF [9]: /lookup/external-ref?access_num=25788097&link_type=MED&atom=%2Fsci%2F348%2F6235%2F642.3.atom [10]: #xref-ref-2-1 "View reference 2 in text" [11]: #xref-ref-3-1 "View reference 3 in text" [12]: {openurl}?query=rft.jtitle%253DScience%26rft_id%253Dinfo%253Adoi%252F10.1126%252Fscience.1201609%26rft_id%253Dinfo%253Apmid%252F21764754%26rft.genre%253Darticle%26rft_val_fmt%253Dinfo%253Aofi%252Ffmt%253Akev%253Amtx%253Ajournal%26ctx_ver%253DZ39.88-2004%26url_ver%253DZ39.88-2004%26url_ctx_fmt%253Dinfo%253Aofi%252Ffmt%253Akev%253Amtx%253Actx [13]: /lookup/ijlink/YTozOntzOjQ6InBhdGgiO3M6MTQ6Ii9sb29rdXAvaWpsaW5rIjtzOjU6InF1ZXJ5IjthOjQ6e3M6ODoibGlua1R5cGUiO3M6NDoiQUJTVCI7czoxMToiam91cm5hbENvZGUiO3M6Mzoic2NpIjtzOjU6InJlc2lkIjtzOjEyOiIzMzMvNjA0NS85ODgiO3M6NDoiYXRvbSI7czoyNDoiL3NjaS8zNDgvNjIzNS82NDIuMy5hdG9tIjt9czo4OiJmcmFnbWVudCI7czowOiIiO30= [14]: #xref-ref-4-1 "View reference 4 in text" [15]: {openurl}?query=rft.jtitle%253DForest%2BEcol.%2BManage.%26rft.volume%253D314%26rft.spage%253D59%26rft_id%253Dinfo%253Adoi%252F10.1016%252Fj.foreco.2013.11.022%26rft.genre%253Darticle%26rft_val_fmt%253Dinfo%253Aofi%252Ffmt%253Akev%253Amtx%253Ajournal%26ctx_ver%253DZ39.88-2004%26url_ver%253DZ39.88-2004%26url_ctx_fmt%253Dinfo%253Aofi%252Ffmt%253Akev%253Amtx%253Actx [16]: /lookup/external-ref?access_num=10.1016/j.foreco.2013.11.022&link_type=DOI

Journal of Tropical Ecology, Oct 20, 2006

Stand structure dynamics during early secondary forest succession were related to mortality, grow... more Stand structure dynamics during early secondary forest succession were related to mortality, growth and recruitment rates, and the dependence of these demographic processes on fallow age and initial stand structure attributes was evaluated. In 11 secondary tropical rainforest sites (1.5-19 y) in Chiapas, Mexico, one plot of 10 × 50 m was established. Diameter and height were measured for all trees ≥ 1 cm dbh, and their survival, growth and recruitment was monitored over a 2-y period. Changes in stand structure were especially fast in the first 5 y of succession, and decreased rapidly afterwards, which resulted from similar stand-level changes in relative mortality, growth and recruitment rates. Demographic processes were negatively related with initial stand basal area, but independent of initial tree density. Basal area was a better explanatory variable of the among-stand variability in these rates than fallow age. Results suggest that asymmetric competition and resulting patterns of tree-thinning are major driving forces determining secondary forest successional pathways. Fallow age per se is a compound variable reflecting community organization at a certain point along the successional axis, while community structure drives succession. Sudden mass mortality among dominant species in some stands showed that early secondary forest succession is not always a gradual and unidirectional process.

Ecology, Sep 29, 2017

Mechanisms of community assembly and ecosystem function are often analyzed using community-weight... more Mechanisms of community assembly and ecosystem function are often analyzed using community-weighted mean trait values (CWMs). We present a novel conceptual framework to quantify the contribution of demographic processes (i.e., growth, recruitment, and mortality) to temporal changes in CWMs. We used this framework to analyze mechanisms of secondary succession in wet tropical forests in Mexico. Seed size increased over time, reflecting a trade-off between colonization by small seeds early in succession, to establishment by large seeds later in succession. Specific leaf area (SLA) and leaf phosphorus content decreased over time, reflecting a trade-off between fast growth early in succession vs. high survival late in succession. On average, CWM shifts were driven mainly (70%) by growth of surviving trees that comprise the bulk of standing biomass, then mortality (25%), and weakly by recruitment (5%). Trait shifts of growing and recruiting trees mirrored the CWM trait shifts, and traits of dying trees did not change during succession, indicating that these traits are important for recruitment and growth, but not for mortality, during the first 30 yr of succession. Identifying the demographic drivers of functional composition change links population dynamics to community change, and enhances insights into mechanisms of succession.

Journal of Tropical Ecology, 1999

Forest Ecology and Management, Mar 1, 2004

Forest Ecology and Management

University of Exeter, 2021

ABSTRACT: The sensitivity of tropical forest carbon to climate is a key uncertainty in predicting... more ABSTRACT: The sensitivity of tropical forest carbon to climate is a key uncertainty in predicting global climate change. Although short-term drying and warming are known to affect forests, it is unknown if such effects translate into long-term responses. Here, we analyze 590 permanent plots measured across the tropics to derive the equilibrium climate controls on forest carbon. Maximum temperature is the most important predictor of aboveground biomass (−9.1 megagrams of carbon per hectare per degree Celsius), primarily by reducing woody productivity, and has a greater rate of decline in the hottest forests (>32.2°C). Our results nevertheless reveal greater thermal resilience than observations of short-term variation imply. To realize the long-term climate adaptation potential of tropical forests requires both protecting them and stabilizing Earth's climate.

There is consensus that plant diversity and ecosystem processes are negatively affected by land-u... more There is consensus that plant diversity and ecosystem processes are negatively affected by land-use intensification (LUI), but, at the same time, there is empirical evidence that a large heterogeneity can be found in the responses. This heterogeneity is especially poorly understood in tropical ecosystems. We evaluated changes in community functional properties across five common land-use types in the wet tropics with different land-use intensity: mature forest, logged forest, secondary forest, agricultural land, and pastureland, located in the lowlands of Bolivia. For the dominant plant species, we measured 12 functional response traits related to their life history, acquisition and conservation of resources, plant domestication, and breeding. We used three single-trait metrics to describe community functional properties: community abundance-weighted mean (CWM) traits values, coefficient of variation, and kurtosis of distribution. The CWM of all 12 traits clearly responded to LUI. Overall, we found that an increase in LUI resulted in communities dominated by plants with acquisitive leaf trait values. However, contrary to our expectations, secondary forests had more conservative trait values (i.e., lower specific leaf area) than mature and logged forest, probably because they were dominated by palm species. Functional variation peaked at intermediate land-use intensity (high coefficient of variation and low kurtosis), which included secondary forest but, unexpectedly, also agricultural land, which is an intensely managed system. The high functional variation of these systems is due to a combination of how response traits (and species) are filtered out by biophysical filters and how management practices introduced a range of exotic species and their trait values into the local species pool. Our results showed that, at local scales and depending on prevailing environmental and management practices, LUI does not necessarily result in communities with more acquisitive trait values or with less functional variation. Instead of the widely expected negative impacts of LUI on plant diversity, we found varying responses of functional variation, with possible repercussions on many ecosystem services. These findings provide a background for actively mitigating negative effects of LUI while meeting the needs of local communities that rely mainly on provisioning ecosystem services for their livelihoods

Tropical forests are global centres of both biodiversity and carbon storage. Many tropical countr... more Tropical forests are global centres of both biodiversity and carbon storage. Many tropical countries aspire to protect forest to fulfil biodiversity and climate mitigation policy targets, but the conservation strategies needed to achieve these two functions depend critically on the tropical forest diversity-carbon relationship and this remains largely unexplored. Attempts to assess and understand this relationship in tropical forest ecosystems have been hindered by the scarcity of inventories where carbon storage in aboveground biomass and species identifications have been simultaneously and robustly quantified. Here, we compile a unique pan-tropical dataset of 360 plots located in old-growth closed-canopy forest, surveyed using standardised methods, allowing a multi-scale evaluation of the relationship between carbon storage and tree diversity. We find strongly contrasting variation in diversity and carbon among continents. Thus, on average, African forests have high carbon storage...

Nature Ecology & Evolution, 2021

Forest restoration increases global forest area and ecosystem services such as primary productivi... more Forest restoration increases global forest area and ecosystem services such as primary productivity and carbon storage. How tree species functional composition impacts the provisioning of these services as forests develop is sparsely studied. We used 10-year data from 478 plots with 191,200 trees in a forest biodiversity experiment in subtropical China to assess the relationship between community productivity and community-weighted mean (CWM) or functional diversity (FD) values of 38 functional traits. We found that effects of FD values on productivity became larger than effects of CWM values after 7 years of forest development and that the FD values also became more reliable predictors of productivity than the CWM values. In contrast to CWM, FD values consistently increased productivity across ten different species-pool subsets. Our results imply that to promote productivity in the long term it is imperative for forest restoration projects to plant multispecies communities with large functional diversity.

Frontiers in Forests and Global Change, 2020

Tropical Conservation Science, 2018

Forest Ecology and Management, 2017

Science, May 8, 2015

TROPICAL FORESTS ARE increasingly modified by human activities. Centuries of human–forest interac... more TROPICAL FORESTS ARE increasingly modified by human activities. Centuries of human–forest interactions have led to a diverse array of forest areas in different phases of succession. In recent decades, forest conversion to cattle pasture or agricultural fields, followed by land abandonment, has led to large areas of second-growth forest in the Amazon. These forests grow rapidly and sequester large amounts of carbon in their biomass, but they tend to be ignored, as most of the debate on the carbon balance of the Amazon basin tends to revolve around old-growth forests. For example, a recent study has shown that the net carbon uptake of Amazonian old-growth forests has declined by a third per decade from 1990 to 2010 ([ 1 ][1], [ 2 ][2]). When extrapolated over the whole Amazon basin, these results translate into a reduced role of intact tropical forests in climate change mitigation. This alarming conclusion, however, completely ignores the important role of regenerating forests as carbon sinks. For instance, in 2010 about 25% of formerly deforested areas in Para, Brazil, were occupied by second-growth forests. Although re-growing forests have lower carbon stocks (45 to 48% of old growth forest), their net carbon sequestration rate is up to 20 times higher (4.6 to 5.8 Mg carbon ha−1 year−1) ([ 3 ][3]) than old-growth forests ([ 1 ][1]). Additionally, about one-quarter of the forests in the Amazon basin are managed for timber production. Net carbon sequestration rates after timber extraction are high, and the application of reduced-impact logging techniques further increases carbon sequestration rates (2.8 Mg ha−1 year−1 compared with 0.5 for conventionally logged areas) ([ 4 ][4]). Consequently, it is essential to incorporate the carbon sequestration potential of second-growth, logged, and managed forests in future assessments of the Amazon basin as a global carbon sink. 1. [↵][5] 1. R. J. W. Brienen 2. et al ., Nature 519, 344 (2015). [OpenUrl][6][CrossRef][7][GeoRef][8][PubMed][9] 2. [↵][10] , Amazon rainforest ability to soak up carbon dioxide is falling (2015); . 3. [↵][11] 1. Y. Pan 2. et al ., Science 333, 988 (2011). [OpenUrl][12][Abstract/FREE Full Text][13] 4. [↵][14] 1. T. A. P. West, 2. E. Vidal, 3. F. E. Putz , Forest Ecol. Manage. 314, 59 (2014). [OpenUrl][15][CrossRef][16] [1]: #ref-1 [2]: #ref-2 [3]: #ref-3 [4]: #ref-4 [5]: #xref-ref-1-1 "View reference 1 in text" [6]: {openurl}?query=rft.jtitle%253DNature%26rft.stitle%253DNature%26rft.issn%253D0028-0836%26rft.aulast%253DBrienen%26rft.auinit1%253DR.%2BJ.%2BW.%26rft.volume%253D519%26rft.issue%253D7543%26rft.spage%253D344%26rft.epage%253D348%26rft.atitle%253DLong-term%2Bdecline%2Bof%2Bthe%2BAmazon%2Bcarbon%2Bsink%26rft_id%253Dinfo%253Adoi%252F10.1038%252Fnature14283%26rft_id%253Dinfo%253Apmid%252F25788097%26rft.genre%253Darticle%26rft_val_fmt%253Dinfo%253Aofi%252Ffmt%253Akev%253Amtx%253Ajournal%26ctx_ver%253DZ39.88-2004%26url_ver%253DZ39.88-2004%26url_ctx_fmt%253Dinfo%253Aofi%252Ffmt%253Akev%253Amtx%253Actx [7]: /lookup/external-ref?access_num=10.1038/nature14283&link_type=DOI [8]: /lookup/external-ref?access_num=2015036225&link_type=GEOREF [9]: /lookup/external-ref?access_num=25788097&link_type=MED&atom=%2Fsci%2F348%2F6235%2F642.3.atom [10]: #xref-ref-2-1 "View reference 2 in text" [11]: #xref-ref-3-1 "View reference 3 in text" [12]: {openurl}?query=rft.jtitle%253DScience%26rft_id%253Dinfo%253Adoi%252F10.1126%252Fscience.1201609%26rft_id%253Dinfo%253Apmid%252F21764754%26rft.genre%253Darticle%26rft_val_fmt%253Dinfo%253Aofi%252Ffmt%253Akev%253Amtx%253Ajournal%26ctx_ver%253DZ39.88-2004%26url_ver%253DZ39.88-2004%26url_ctx_fmt%253Dinfo%253Aofi%252Ffmt%253Akev%253Amtx%253Actx [13]: /lookup/ijlink/YTozOntzOjQ6InBhdGgiO3M6MTQ6Ii9sb29rdXAvaWpsaW5rIjtzOjU6InF1ZXJ5IjthOjQ6e3M6ODoibGlua1R5cGUiO3M6NDoiQUJTVCI7czoxMToiam91cm5hbENvZGUiO3M6Mzoic2NpIjtzOjU6InJlc2lkIjtzOjEyOiIzMzMvNjA0NS85ODgiO3M6NDoiYXRvbSI7czoyNDoiL3NjaS8zNDgvNjIzNS82NDIuMy5hdG9tIjt9czo4OiJmcmFnbWVudCI7czowOiIiO30= [14]: #xref-ref-4-1 "View reference 4 in text" [15]: {openurl}?query=rft.jtitle%253DForest%2BEcol.%2BManage.%26rft.volume%253D314%26rft.spage%253D59%26rft_id%253Dinfo%253Adoi%252F10.1016%252Fj.foreco.2013.11.022%26rft.genre%253Darticle%26rft_val_fmt%253Dinfo%253Aofi%252Ffmt%253Akev%253Amtx%253Ajournal%26ctx_ver%253DZ39.88-2004%26url_ver%253DZ39.88-2004%26url_ctx_fmt%253Dinfo%253Aofi%252Ffmt%253Akev%253Amtx%253Actx [16]: /lookup/external-ref?access_num=10.1016/j.foreco.2013.11.022&link_type=DOI

Journal of Tropical Ecology, Oct 20, 2006

Stand structure dynamics during early secondary forest succession were related to mortality, grow... more Stand structure dynamics during early secondary forest succession were related to mortality, growth and recruitment rates, and the dependence of these demographic processes on fallow age and initial stand structure attributes was evaluated. In 11 secondary tropical rainforest sites (1.5-19 y) in Chiapas, Mexico, one plot of 10 × 50 m was established. Diameter and height were measured for all trees ≥ 1 cm dbh, and their survival, growth and recruitment was monitored over a 2-y period. Changes in stand structure were especially fast in the first 5 y of succession, and decreased rapidly afterwards, which resulted from similar stand-level changes in relative mortality, growth and recruitment rates. Demographic processes were negatively related with initial stand basal area, but independent of initial tree density. Basal area was a better explanatory variable of the among-stand variability in these rates than fallow age. Results suggest that asymmetric competition and resulting patterns of tree-thinning are major driving forces determining secondary forest successional pathways. Fallow age per se is a compound variable reflecting community organization at a certain point along the successional axis, while community structure drives succession. Sudden mass mortality among dominant species in some stands showed that early secondary forest succession is not always a gradual and unidirectional process.

Ecology, Sep 29, 2017

Mechanisms of community assembly and ecosystem function are often analyzed using community-weight... more Mechanisms of community assembly and ecosystem function are often analyzed using community-weighted mean trait values (CWMs). We present a novel conceptual framework to quantify the contribution of demographic processes (i.e., growth, recruitment, and mortality) to temporal changes in CWMs. We used this framework to analyze mechanisms of secondary succession in wet tropical forests in Mexico. Seed size increased over time, reflecting a trade-off between colonization by small seeds early in succession, to establishment by large seeds later in succession. Specific leaf area (SLA) and leaf phosphorus content decreased over time, reflecting a trade-off between fast growth early in succession vs. high survival late in succession. On average, CWM shifts were driven mainly (70%) by growth of surviving trees that comprise the bulk of standing biomass, then mortality (25%), and weakly by recruitment (5%). Trait shifts of growing and recruiting trees mirrored the CWM trait shifts, and traits of dying trees did not change during succession, indicating that these traits are important for recruitment and growth, but not for mortality, during the first 30 yr of succession. Identifying the demographic drivers of functional composition change links population dynamics to community change, and enhances insights into mechanisms of succession.