• Energy Research

Summary There are two theories about how allocation of metabolic products occurs. The allometric biomass partitioning theory (APT) suggests that all plants follow common allometric scaling rules. The optimal partitioning theory (OPT) predicts that plants allocate more biomass to the organ capturing the most limiting resource. Whole‐plant harvests of mature and juvenile tropical deciduous trees, evergreen trees, and lianas and model simulations were used to address the following knowledge gaps: (1) Do mature lianas comply with the APT scaling laws or do they invest less biomass in stems compared to trees? (2) Do juveniles follow the same allocation patterns as mature individuals? (3) Is either leaf phenology or life form a predictor of rooting depth? It was found that: (1) mature lianas followed the same allometric scaling laws as trees; (2) juveniles and mature individuals do not follow the same allocation patterns; and (3) mature lianas had shallowest coarse roots and evergreen trees had the deepest. It was demonstrated that: (1) mature lianas invested proportionally similar biomass to stems as trees and not less, as expected; (2) lianas were not deeper‐rooted than trees as had been previously proposed; and (3) evergreen trees had the deepest roots, which is necessary to maintain canopy during simulated dry seasons.

Summary Censuses of tropical forest plots reveal large variation in biomass and plant composition. This paper evaluates whether such variation can emerge solely from realistic variation in a set of commonly measured soil chemical and physical properties. Controlled simulations were performed using a mechanistic model that includes forest dynamics, microbe‐mediated biogeochemistry, and competition for nitrogen and phosphorus. Observations from 18 forest inventory plots in Guanacaste, Costa Rica were used to determine realistic variation in soil properties. In simulations of secondary succession, the across‐plot range in plant biomass reached 30% of the mean and was attributable primarily to nutrient limitation and secondarily to soil texture differences that affected water availability. The contributions of different plant functional types to total biomass varied widely across plots and depended on soil nutrient status. In Central America, soil‐induced variation in plant biomass increased with mean annual precipitation because of changes in nutrient limitation. In Central America, large variation in plant biomass and ecosystem composition arises mechanistically from realistic variation in soil properties. The degree of biomass and compositional variation is climate sensitive. In general, model predictions can be improved through better representation of soil nutrient processes, including their spatial variation.

Most measurements and models of forest carbon cycling neglect the carbon flux associated with the turnover of branch biomass, a physiological process quantified for other organs (fine roots, leaves, and stems). Synthesizing data from boreal, temperate, and tropical forests (184,815 trees), we found that including branch turnover increased empirical estimates of aboveground wood production by 16% (equivalent to 1.9 Pg Cy −1 globally), of similar magnitude to the observed global forest carbon sinks. In addition, reallocating carbon to branch turnover in model simulations reduced stem wood biomass, a long-lasting carbon storage, by 7 to 17%. This prevailing neglect of branch turnover suggests widespread biases in carbon flux estimates across global datasets and model simulations. Branch litterfall, sometimes used as a proxy for branch turnover, ignores carbon lost from attached dead branches, underestimating branch C turnover by 38% in a pine forest. Modifications to field measurement protocols and existing models are needed to allow a more realistic partitioning of wood production and forest carbon storage.

AbstractSensitivity of forest mortality to drought in carbon‐dense tropical forests remains fraught with uncertainty, while extreme droughts are predicted to be more frequent and intense. Here, the potential of temporal autocorrelation of high‐frequency variability in Landsat Enhanced Vegetation Index (EVI), an indicator of ecosystem resilience, to predict spatial and temporal variations of forest biomass mortality is evaluated against in situ census observations for 64 site‐year combinations in Costa Rican tropical dry forests during the 2015 ENSO drought. Temporal autocorrelation, within the optimal moving window of 24 months, demonstrated robust predictive power for in situ mortality (leave‐one‐out cross‐validation R2 = 0.54), which allows for estimates of annual biomass mortality patterns at 30 m resolution. Subsequent spatial analysis showed substantial fine‐scale heterogeneity of forest mortality patterns, largely driven by drought intensity and ecosystem properties related to plant water use such as forest deciduousness and topography. Highly deciduous forest patches demonstrated much lower mortality sensitivity to drought stress than less deciduous forest patches after elevation was controlled. Our results highlight the potential of high‐resolution remote sensing to “fingerprint” forest mortality and the significant role of ecosystem heterogeneity in forest biomass resistance to drought.

AbstractRapid warming and changes in water availability at high latitudes alter resource abundance, tree competition, and disturbance regimes. While these changes are expected to disrupt the functioning of boreal forests, their ultimate implications for forest composition are uncertain. In particular, recent site‐level studies of the Alaskan boreal forest have reported both increases and decreases in productivity over the past few decades. Here, we test the idea that variations in Alaskan forest growth and mortality rates are contingent on species composition. Using forest inventory measurements and climate data from plots located throughout interior and south‐central Alaska, we show significant growth and mortality responses associated with competition, midsummer vapor pressure deficit, and increased growing season length. The governing climate and competition processes differed substantially across species. Surprisingly, the most dramatic climate response occurred in the drought tolerant angiosperm species, trembling aspen, and linked high midsummer vapor pressure deficits to decreased growth and increased insect‐related mortality. Given that species composition in the Alaskan and western Canadian boreal forests is projected to shift toward early‐successional angiosperm species due to fire regime, these results underscore the potential for a reduction in boreal productivity stemming from increases in midsummer evaporative demand.

AbstractThe strength and persistence of the tropical carbon sink hinges on the long‐term responses of woody growth to climatic variations and increasing CO2. However, the sensitivity of tropical woody growth to these environmental changes is poorly understood, leading to large uncertainties in growth predictions. Here, we used tree ring records from a Southeast Asian tropical forest to constrain ED2.2‐hydro, a terrestrial biosphere model with explicit vegetation demography. Specifically, we assessed individual‐level woody growth responses to historical climate variability and increases in atmospheric CO2 (Ca). When forced with historical Ca, ED2.2‐hydro reproduced the magnitude of increases in intercellular CO2 concentration (a major determinant of photosynthesis) estimated from tree ring carbon isotope records. In contrast, simulated growth trends were considerably larger than those obtained from tree rings, suggesting that woody biomass production efficiency (WBPE = woody biomass production:gross primary productivity) was overestimated by the model. The estimated WBPE decline under increasing Ca based on model‐data discrepancy was comparable to or stronger than (depending on tree species and size) the observed WBPE changes from a multi‐year mature‐forest CO2 fertilization experiment. In addition, we found that ED2.2‐hydro generally overestimated climatic sensitivity of woody growth, especially for late‐successional plant functional types. The model‐data discrepancy in growth sensitivity to climate was likely caused by underestimating WBPE in hot and dry years due to commonly used model assumptions on carbon use efficiency and allocation. To our knowledge, this is the first study to constrain model predictions of individual tree‐level growth sensitivity to Ca and climate against tropical tree‐ring data. Our results suggest that improving model processes related to WBPE is crucial to obtain better predictions of tropical forest responses to droughts and increasing Ca. More accurate parameterization of WBPE will likely reduce the stimulation of woody growth by Ca rise predicted by biosphere models.

AbstractEnhancement of net primary production (NPP) in forests as atmospheric [CO2] increases is likely limited by the availability of other growth resources. The Duke Free Air CO2 Enrichment (FACE) experiment was located on a moderate‐fertility site in the southeastern US, in a loblolly pine (Pinus taeda L.) plantation with broadleaved species growing mostly in mid‐canopy and understory. Duke FACE ran from 1994 to 2010 and combined elevated [CO2] (eCO2) with nitrogen (N) additions. We assessed the spatial and temporal variation of NPP response using a dataset that includes previously unpublished data from 6 years of the replicated CO2 × N experiment and extends to 2 years beyond the termination of enrichment. Averaged over time (1997–2010), NPP of pine and broadleaved species were 38% and 52% higher under eCO2 compared to ambient conditions. Furthermore, there was no evidence of a decline in enhancement over time in any plot regardless of its native site quality. The relation between spatial variation in the response and native site quality was suggested but inconclusive. Nitrogen amendments under eCO2, in turn, resulted in an additional 11% increase in pine NPP. For pine, the eCO2‐induced increase in NPP was similar above‐ and belowground and was driven by both increased leaf area index (L) and production efficiency (PE = NPP/L). For broadleaved species, coarse‐root biomass production was more than 200% higher under eCO2 and accounted for the entire production response, driven by increased PE. Notably, the fraction of annual NPP retained in total living biomass was higher under eCO2, reflecting a slight shift in allocation fraction to woody mass and a lower mortality rate. Our findings also imply that tree growth may not have been only N‐limited, but perhaps constrained by the availability of other nutrients. The observed sustained NPP enhancement, even without N‐additions, demonstrates no progressive N limitation.