Second-growth tropical forests are an important global carbon sink. As current knowledge on biomass accumulation during secondary succession is heavily based on chronosequence studies, direct ...estimates of annual rates of biomass accumulation in monitored stands are largely unavailable. We evaluated the contributions of tree diameter increment, recruitment, and mortality to annual tree biomass change during succession for three groups of tree species: second-growth (SG) specialists, generalists, and old-growth (OG) specialists. We monitored six second-growth tropical forests that varied in stand age and two old-growth forests in northeastern Costa Rica. We monitored these over a period of 8 to 16 years. To assess rates of biomass change during secondary succession, we compared standing biomass and biomass dynamics between second-growth forest stages and old-growth forest, and evaluated the effect of stand age on standing biomass and biomass dynamics in second-growth forests.
Standing tree biomass increased with stand age during succession, whereas the rate of biomass change decreased. Biomass change was largely driven by tree diameter increment and mortality, with a minor contribution from recruitment. The relative importance of these demographic drivers shifted over succession. Biomass gain due to tree diameter increment decreased with stand age, whereas biomass loss due to mortality increased. In the age range of our second-growth forests, 10-41 years, SG specialists dominated tree biomass in second-growth forests. SG specialists, and to a lesser extent generalists, also dominated stand-level biomass increase due to tree diameter increment, whereas SG specialists largely accounted for decreases in biomass due to mortality.
Our results indicate that tree growth is largely driving biomass dynamics early in succession, whereas both growth and mortality are important later in succession. Biomass dynamics are largely accounted for by a few SG specialists and one generalist species,
Pentaclethra macroloba
. To assess the generality of our results, similar long-term studies should be compared across tropical forest landscapes.
In closed-canopy forests, gap formation and closure are thought to be major drivers of forest dynamics. Crown defoliation by insects, however, may also influence understory resource levels and thus ...forest dynamics. We evaluate the effect of a forest tent caterpillar outbreak on understory light availability, soil nutrient levels and tree seedling height growth in six sites with contrasting levels of canopy defoliation in a hardwood forest in northern lower Michigan. We compared resource levels and seedling growth of six hardwood species before, during and in the three years after the outbreak (2008-2012). Canopy openness increased strongly during the forest tent caterpillar outbreak in the four moderately and severely defoliated sites, but not in lightly defoliated sites. Total inorganic soil nitrogen concentrations increased in response to the outbreak in moderately and severely defoliated sites. The increase in total inorganic soil nitrogen was driven by a strong increase in soil nitrate, and tended to become stronger with increasing site defoliation. Seedling height growth increased for all species in the moderately and severely defoliated sites, but not in lightly defoliated sites, either during the outbreak year or in the year after the outbreak. Growth increases did not become stronger with increasing site defoliation, but were strongest in a moderately defoliated site with high soil nutrient levels. Growth increases tended to be strongest for the shade intolerant species Fraxinus americana and Prunus serotina, and the shade tolerant species Ostrya virginiana. The strong growth response of F. americana and P. serotina suggests that recurring forest tent caterpillar outbreaks may facilitate the persistence of shade intolerant species in the understory in the absence of canopy gaps. Overall, our results suggest that recurrent canopy defoliation resulting from cyclical forest insect outbreaks may be an additional driver of dynamics in temperate closed-canopy forests.
Over the last decade the field of tropical dendroecology has developed rapidly and major achievements have been made. We reviewed the advances in three main themes within the field. First, long ...chronologies for tropical tree species were constructed which allowed climate reconstructions, revealed sources of climatic variation and clarified climate-growth relations. Other studies combined tree-ring data and stable isotope (¹³C and ¹⁸O) measurements to evaluate the response of tropical trees to climatic variation and changes. A second set of studies assessed long-term growth patterns of individual trees throughout their life. These studies enhanced the understanding of growth trajectories to the canopy, quantified autocorrelated tree growth and yielded new estimates of tree ages. Such studies were also used to reconstruct the disturbance history of tropical forests. The last set of studies applied tree-ring data to growth models. Tree-ring data can replace diameter measurements from research plots, provide additional information to construct population models, improve timber yield models and validate model output. Based on our review, we propose two main directions for future research. (1) An evaluation of the causes and consequences of growth variation within and among trees and their relation to environmental variation. Studies evaluating this directly contribute to improved understanding of tropical tree ecology. (2) The simultaneous measurement of widths and stable isotope fractions in tree rings offers the potential to study responses of trees to climatic change. Given the major role of tropical forests in the global carbon cycle, knowing these responses is of high priority.
The achievement of international goals and national commitments related to forest conservation and management, climate change, and sustainable development requires credible, accurate, and reliable ...monitoring of stocks and changes in forest biomass and carbon. Most prominently, the Paris Agreement on Climate Change and the United Nations’ Sustainable Development Goals in particular require data on biomass to monitor progress. Unprecedented opportunities to provide forest biomass data are created by a series of upcoming space-based missions, many of which provide open data targeted at large areas and better spatial resolution biomass monitoring than has previously been achieved. We assess various policy needs for biomass data and recommend a long-term collaborative effort among forest biomass data producers and users to meet these needs. A gap remains, however, between what can be achieved in the research domain and what is required to support policy making and meet reporting requirements. There is no single biomass dataset that serves all users in terms of definition and type of biomass measurement, geographic area, and uncertainty requirements, and whether there is need for the most recent up-to-date biomass estimate or a long-term biomass trend. The research and user communities should embrace the potential strength of the multitude of upcoming missions in combination to provide for these varying needs and to ensure continuity for long-term data provision which one-off research missions cannot provide. International coordination bodies such as Global Forest Observations Initiative (GFOI), Committee on Earth Observation Satellites (CEOS), and Global Observation of Forest Cover and Land Dynamics (GOFC‐GOLD) will be integral in addressing these issues in a way that fulfils these needs in a timely fashion. Further coordination work should particularly look into how space-based data can be better linked with field reference data sources such as forest plot networks, and there is also a need to ensure that reference data cover a range of forest types, management regimes, and disturbance regimes worldwide.
Backgrounds and aims
Litter protects the underlying soil, depending on litterfall and decomposition, but dynamics of the standing litter stock in agroforestry systems remain poorly understood. We ...aimed to unravel effects of litter quality, temporal patterns, microclimate, and a possible home-field advantage (HFA) on standing litter dynamics across a land-use gradient.
Methods
We quantified litterfall, the standing litter stock, and microclimate during a year in (remnant) forest, cacao-based simple and complex agroforestry, cacao monocultures, and annual crops in a cacao producing area in Indonesia. We conducted a reciprocal litter transfer experiment, and tested decomposition rates of pruning residues. Standing litter stocks during the year were estimated from monthly litterfall and decomposition rates.
Results
Variation in litter quality influenced decomposition rates more strongly than variation in microclimate or HFA. Lower litter quality in complex agroforestry and in the cacao monoculture decreased the decay rate compared to simple agroforestry systems; mean litter residence time was over a year. Mixing high- and low-quality material in pruning residues modified the decomposition rate, soil C and N changes, offering options for targeted management of soil protection and nutrient release.
Conclusions
The seasonal patterns of litterfall and relatively slow decomposition rates supported permanence of the litter layer in all cacao production systems, protecting the underlying soil.
Trees forage for light through optimal leaf display. Effective leaf display is determined by metamer traits (i.e., the internode, petiole, and corresponding leaf), and thus these traits strongly ...co-determine carbon gain and as a result competitive advantage in a light-limited environment. We examined 11 metamer traits of sun and shade trees of 38 coexisting moist forest tree species and determined the relative strengths of intra- and interspecific variation. Species-specific metamer traits were related to two variables that represent important life history variation; the regeneration light requirements and average leaf size of the species. Metamer traits varied strongly across species and, in contrast to our expectation, showed only modest changes in response to light. Intra- and interspecific responses to light were only congruent for a third of the traits evaluated. Four traits, amongst which leaf size, specific leaf area (SLA), and leaf area ratio at the metamer level (LAR) showed even opposite intra- and interspecific responses to light. Strikingly, these are classic traits that are thought to be of paramount importance for plant performance but that have completely different consequences within and across species. Sun trees of a given species had small leaves to reduce the heat load, but light-demanding species had large leaves compared to shade-tolerants, probably to outcompete their neighbors. Shade trees of a given species had a high SLA and LAR to capture more light in a light-limited environment, whereas shade-tolerant species have well-protected leaves with a low SLA compared to light-demanding species, probably to deter herbivores and enhance leaf lifespan. There was a leaf-size-mediated trade-off between biomechanical and hydraulic safety, and the efficiency with which species can space their leaves and forage for light. Unexpectedly, metamer traits were more closely linked to leaf size than to regeneration light requirements, probably because leaf-size-related biomechanical and vascular constraints limit the trait combinations that are physically possible. This suggests that the leaf size spectrum overrules more subtle variation caused by the leaf economics spectrum, and that leaf size represents a more important strategy axis than previously thought.
The terrestrial forest carbon pool is poorly quantified, in particular in regions with low forest inventory capacity. By combining multiple satellite observations of synthetic aperture radar (SAR) ...backscatter around the year 2010, we generated a global, spatially explicit dataset of above-ground live biomass (AGB; dry mass) stored in forests with a spatial resolution of 1 ha. Using an extensive database of 110 897 AGB measurements from field inventory plots, we show that the spatial patterns and magnitude of AGB are well captured in our map with the exception of regional uncertainties in high-carbon-stock forests with AGB >250 Mg ha−1, where the retrieval was effectively based on a single radar observation. With a total global AGB of 522 Pg, our estimate of the terrestrial biomass pool in forests is lower than most estimates published in the literature (426–571 Pg). Nonetheless, our dataset increases knowledge on the spatial distribution of AGB compared to the Global Forest Resources Assessment (FRA) by the Food and Agriculture Organization (FAO) and highlights the impact of a country's national inventory capacity on the accuracy of the biomass statistics reported to the FRA. We also reassessed previous remote sensing AGB maps and identified major biases compared to inventory data, up to 120 % of the inventory value in dry tropical forests, in the subtropics and temperate zone. Because of the high level of detail and the overall reliability of the AGB spatial patterns, our global dataset of AGB is likely to have significant impacts on climate, carbon, and socio-economic modelling schemes and provides a crucial baseline in future carbon stock change estimates. The dataset is available at https://doi.org/10.1594/PANGAEA.894711 (Santoro, 2018).
Plant species composition and diversity are known to change across local gradients of light, moisture and nutrients, but ecologists still have a relatively limited understanding of how communities ...respond to multiple limiting resources.
We used a trait‐based approach to investigate how the functional composition and diversity of forest understorey plant communities change along gradients in light, soil moisture and nitrogen availability. We used a total of seven leaf, root and whole‐plant traits for 55–78 species, and estimated the effects of the three resources on the mean and dispersion of these traits in understorey plant communities across 50 forest sites.
Soil moisture and nitrogen availability (C/N ratio) both influenced plant community traits, but light availability (canopy openness) did not. Generally, increases in moisture and nitrogen both resulted in shifts towards more acquisitive resource use strategies, including greater leaf area, specific leaf area and maximum plant height, and lower leaf dry matter content, root dry matter content and rooting depth. Functional diversity of most traits also increased with increasing soil moisture and nitrogen. Although most traits varied with soil moisture on nitrogen‐poor sites, moisture did not influence of the distribution of any traits on nitrogen‐rich sites.
Synthesis. Independent co‐limitation of soil moisture and nitrogen appeared to influence the functional composition and diversity of understorey vegetation in our study area. The co‐occurrence of species with resource acquisitive and conservative strategies on nitrogen‐rich sites may make plant communities relatively resistant to changes to soil moisture. These results suggest that altered precipitation regimes under climate change could lead to greater changes in the composition and diversity of plant communities on nutrient‐poor soils than on nutrient‐rich soils.
Independent co‐limitation of soil moisture and nitrogen appeared to influence the functional composition and diversity of understorey vegetation in our study area. The co‐occurrence of species with resource acquisitive and conservative strategies on nitrogen‐rich sites may make plant communities relatively resistant to changes to soil moisture. These results suggest that altered precipitation regimes under climate change could lead to greater changes in the composition and diversity of plant communities on nutrient‐poor soils than on nutrient‐rich soils.
As countries advance in greenhouse gas (GHG) accounting for climate change mitigation, consistent estimates of aboveground net biomass change (∆AGB) are needed. Countries with limited forest ...monitoring capabilities in the tropics and subtropics rely on IPCC 2006 default ∆AGB rates, which are values per ecological zone, per continent. Similarly, research into forest biomass change at a large scale also makes use of these rates. IPCC 2006 default rates come from a handful of studies, provide no uncertainty indications and do not distinguish between older secondary forests and old‐growth forests. As part of the 2019 Refinement to the 2006 IPCC Guidelines for National Greenhouse Gas Inventories, we incorporate ∆AGB data available from 2006 onwards, comprising 176 chronosequences in secondary forests and 536 permanent plots in old‐growth and managed/logged forests located in 42 countries in Africa, North and South America and Asia. We generated ∆AGB rate estimates for younger secondary forests (≤20 years), older secondary forests (>20 years and up to 100 years) and old‐growth forests, and accounted for uncertainties in our estimates. In tropical rainforests, for which data availability was the highest, our ∆AGB rate estimates ranged from 3.4 (Asia) to 7.6 (Africa) Mg ha−1 year−1 in younger secondary forests, from 2.3 (North and South America) to 3.5 (Africa) Mg ha−1 year−1 in older secondary forests, and 0.7 (Asia) to 1.3 (Africa) Mg ha−1 year−1 in old‐growth forests. We provide a rigorous and traceable refinement of the IPCC 2006 default rates in tropical and subtropical ecological zones, and identify which areas require more research on ∆AGB. In this respect, this study should be considered as an important step towards quantifying the role of tropical and subtropical forests as carbon sinks with higher accuracy; our new rates can be used for large‐scale GHG accounting by governmental bodies, nongovernmental organizations and in scientific research.
As countries advance in greenhouse gas (GHG) accounting for climate change mitigation, consistent estimates of aboveground biomass change (∆AGB) in natural forests are needed. Countries with limited forest monitoring capabilities in the (sub)tropics rely on the 2006 IPCC guidelines for default ∆AGB rates. This study provides refined default ∆AGB rate estimates based on forest plot data in younger secondary forests, older secondary forests and old‐growth forests, located in 42 countries across (sub)tropical ecozones. These new estimates contribute towards the 2019 IPCC Refinement and can be used for large‐scale GHG accounting by governmental bodies, nongovernmental organizations and in scientific research.
The magnitude of the carbon sink in second-growth forests is expected to vary with successional biomass dynamics resulting from tree growth, recruitment, and mortality, and with the effects of ...climate on these dynamics. We compare aboveground biomass dynamics of dry and wet Neotropical forests, based on monitoring data gathered over 3–16 years in forests covering the first 25 years of succession. We estimated standing biomass, annual biomass change, and contributions of tree growth, recruitment, and mortality. We also evaluated tree species’ contributions to biomass dynamics. Absolute rates of biomass change were lower in dry forests, 2.3 and 1.9 Mg ha⁻¹ y⁻¹, after 5–15 and 15–25 years after abandonment, respectively, than in wet forests, with 4.7 and 6.1 Mg ha⁻¹ y⁻¹, in the same age classes. Biomass change was largely driven by tree growth, accounting for at least 48% of biomass change across forest types and age classes. Mortality also contributed strongly to biomass change in wet forests of 5–15 years, whereas its contribution became important later in succession in dry forests. Biomass dynamics tended to be dominated by fewer species in early-successional dry than wet forests, but dominance was strong in both forest types. Overall, our results indicate that biomass dynamics during succession are faster in Neotropical wet than dry forests, with high tree mortality earlier in succession in the wet forests. Long-term monitoring of second-growth tropical forest plots is crucial for improving estimates of annual biomass change, and for enhancing understanding of the underlying mechanisms and demographic drivers.
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