G. Bruce Williamson

Biomass resilience of Neotropical secondary forests

Land-use change occurs nowhere more rapidly than in the tropics, where the imbalance between deforestation and forest regrowth has large consequences for the global carbon cycle. However, considerable uncertainty remains about the rate of biomass recovery in secondary forests, and how these rates are influenced by climate, landscape, and prior land use. Here we analyse aboveground biomass recovery during secondary succession in 45 forest sites and about 1,500 forest plots covering the major environmental gradients in the Neotropics. The studied secondary forests are highly productive and resilient. Aboveground biomass recovery after 20 years was on average 122 megagrams per hectare (Mg ha−1), corresponding to a net carbon uptake of 3.05 Mg C ha−1 yr−1, 11 times the uptake rate of old-growth forests. Aboveground biomass stocks took a median time of 66 years to recover to 90% of old-growth values. Aboveground biomass recovery after 20 years varied 11.3-fold (from 20 to 225 Mg ha−1) across sites, and this recovery increased with water availability (higher local rainfall and lower climatic water deficit). We present a biomass recovery map of Latin America, which illustrates geographical and climatic variation in carbon sequestration potential during forest regrowth. The map will support policies to minimize forest loss in areas where biomass resilience is naturally low (such as seasonally dry forest regions) and promote forest regeneration and restoration in humid tropical lowland areas with high biomass resilience.

Measuring wood specific gravity…Correctly

The specific gravity (SG) of wood is a measure of the amount of structural material a tree species allocates to support and strength. In recent years, wood specific gravity, traditionally a foresters variable, has become the domain of ecologists exploring the universality of plant functional traits and conservationists estimating global carbon stocks. While these developments have expanded our knowledge and sample of woods, the methodologies employed to measure wood SG have not received as much scrutiny as SGs ecological importance. Here, we reiterate some of the basic principles and methods for measuring the SG of wood to clarify past practices of foresters and ecologists and to identify some of the prominent errors in recent studies and their consequences. In particular, we identify errors in (1) extracting wood samples that are not representative of tree wood, (2) differentiating wood specific gravity from wood density, (3) drying wood samples at temperatures below 100°C and the resulting moisture content complications, and (4) improperly measuring wood volumes. In addition, we introduce a new experimental technique, using applied calculus, for estimating SG when the form of radial variation is known, a method that significantly reduces the effort required to sample a trees wood.

Bioassays for allelopathy: Measuring treatment responses with independent controls

In bioassays for allelopathy, where responses to treatments are determined in conjunction with responses to independent controls, statistical comparisons among treatments require an index which measures each treatment response (T) in relation to its control response (C). The most commonly used index, the treatment-control ratio (T/C), exhibits two analytical problems. First, means ofT/C values are distorted upward when any of the individual values is greater than one, i.e., when stimulation occurs. Second, the distribution ofT/C values may not be normal and homoscedastic. We provide two alternative indices of response whose means do not exhibit the upward distortion ofT/C means. Then, the two indices are compared toT/C values in an empirical test for normality and homoscedasticity on a large bioassay data set. Results indicate that for this data set, one of the alternatives,RI, is clearly superior.RI is defined as 1 — (C/T) ifT≥C and asT/C — 1 ifT

Successional dynamics in Neotropical forests are as uncertain as they are predictable

Significance Although forest succession has been approached as a predictable process, successional trajectories vary widely, even among nearby stands with similar environmental conditions and disturbance histories. We quantified predictability and uncertainty during tropical forest succession using dynamical models describing the interactions among stem density, basal area, and species density over time. We showed that the trajectories of these forest attributes were poorly predicted by stand age and varied significantly within and among sites. Our models reproduced the general successional trends observed, but high levels of noise were needed to increase model predictability. These levels of uncertainty call into question the premise that successional processes are consistent over space and time, and challenge the way ecologists view tropical forest regeneration. Although forest succession has traditionally been approached as a deterministic process, successional trajectories of vegetation change vary widely, even among nearby stands with similar environmental conditions and disturbance histories. Here, we provide the first attempt, to our knowledge, to quantify predictability and uncertainty during succession based on the most extensive long-term datasets ever assembled for Neotropical forests. We develop a novel approach that integrates deterministic and stochastic components into different candidate models describing the dynamical interactions among three widely used and interrelated forest attributes—stem density, basal area, and species density. Within each of the seven study sites, successional trajectories were highly idiosyncratic, even when controlling for prior land use, environment, and initial conditions in these attributes. Plot factors were far more important than stand age in explaining successional trajectories. For each site, the best-fit model was able to capture the complete set of time series in certain attributes only when both the deterministic and stochastic components were set to similar magnitudes. Surprisingly, predictability of stem density, basal area, and species density did not show consistent trends across attributes, study sites, or land use history, and was independent of plot size and time series length. The model developed here represents the best approach, to date, for characterizing autogenic successional dynamics and demonstrates the low predictability of successional trajectories. These high levels of uncertainty suggest that the impacts of allogenic factors on rates of change during tropical forest succession are far more pervasive than previously thought, challenging the way ecologists view and investigate forest regeneration.

Carbon sequestration potential of second-growth forest regeneration in the Latin American tropics

Models reveal the high carbon mitigation potential of tropical forest regeneration. Regrowth of tropical secondary forests following complete or nearly complete removal of forest vegetation actively stores carbon in aboveground biomass, partially counterbalancing carbon emissions from deforestation, forest degradation, burning of fossil fuels, and other anthropogenic sources. We estimate the age and spatial extent of lowland second-growth forests in the Latin American tropics and model their potential aboveground carbon accumulation over four decades. Our model shows that, in 2008, second-growth forests (1 to 60 years old) covered 2.4 million km2 of land (28.1% of the total study area). Over 40 years, these lands can potentially accumulate a total aboveground carbon stock of 8.48 Pg C (petagrams of carbon) in aboveground biomass via low-cost natural regeneration or assisted regeneration, corresponding to a total CO2 sequestration of 31.09 Pg CO2. This total is equivalent to carbon emissions from fossil fuel use and industrial processes in all of Latin America and the Caribbean from 1993 to 2014. Ten countries account for 95% of this carbon storage potential, led by Brazil, Colombia, Mexico, and Venezuela. We model future land-use scenarios to guide national carbon mitigation policies. Permitting natural regeneration on 40% of lowland pastures potentially stores an additional 2.0 Pg C over 40 years. Our study provides information and maps to guide national-level forest-based carbon mitigation plans on the basis of estimated rates of natural regeneration and pasture abandonment. Coupled with avoided deforestation and sustainable forest management, natural regeneration of second-growth forests provides a low-cost mechanism that yields a high carbon sequestration potential with multiple benefits for biodiversity and ecosystem services.

Ceratiolin and other flavonoids from Ceratiola ericoides

Abstract Chemical analysis of ground aerial parts of Ceratiola ericoides yielded the two known dihydrochalcones angoletin and 2′,6′-dihydroxy-4-methoxy-3′,5′-dimethyldihydrochalcone, as well as 2′,4′-dihydroxychalcone. Furthermore, the known flavanones 7-hydroxyflavanone, 8-methylpinocembrin and 6,8-dimethylpinocembrin were isolated. Methanol extracts of ground leaves provided catechin, epicatechin and epicatechin-(4β → 8; 2β → 0 → 7)-epicatechin. From water washes of freshly harvested leaves a novel dihydrochalcone, ceratiolin, was isolated. The structures were inferred from NMR, mass spectral and chemical data, and the molecular structure of 6,8-dimethylpinocembrin was determined by single crystal X-ray analysis.

Radial variation in the wood specific gravity of Joannesia princeps: the roles of age and diameter

Two models are developed to illustrate how age or diameter may control the radial increases in wood specific gravity (SG), a feature common to lowland tropical trees. In the age-dependent model, trees of the same age produce new secondary xylem of the same SG regardless of their diameters, i.e., SG is dependent on age. In the radius-dependent model, trees of the same radius produce new secondary xylem of the same SG regardless of their ages. Then, predictions of the two models are tested on radial wood samples from the trunkwood of Joannesia princeps Vell., growing in a 17-year-old plantation in Espirito Santo, Brazil. For this cohort, tests of four predictions supported the age-dependent model over the radius-dependent model: final specific gravity was independent of radius (smaller trees did not have smaller final specific gravities), the slope of the radial increase in SG with tree radius was negatively dependent on tree radius (smaller trees had steeper slopes), the coefficient of variation (CV) of SG of the final wood was the same or less than the CV of SG of the initial wood, and the CV of SG of the final wood was much less than the CV of the tree radius. Thus, for plantation trees of the same age, the SG of wood produced is primarily a function of age, not radius of the tree. Forest trees show similar relationships but with the effects of age and radius confounded because tree ages are unknown.

Inhibition ofSchizachyrium scoparium (poaceae) by the allelochemical hydrocinnamic acid

Bare zones around shrubs in the Florida scrub indicate the possibility of allelopathy by shrubs controlling the distribution of grasses invading from adjacent sandhills. The allelochemical, hydrocinnamic acid, has been identified as a breakdown product of ceratiolin, which is released from the shrubCeratiola ericoides. Here, hydrocinnamic acid (HCA) was shown to have a strongly inhibitory effect on shoot and root biomass of the grassSchizachyrium scoparium in greenhouse bioassays lasting 4.5 months. Linear increases in the concentration of HCA from 0 to 200 ppm, applied biweekly, resulted in exponential decreases in root and shoot biomass at harvest. Plants grown at 200 ppm HCA had root and shoot biomasses 13% and 17% of controls, respectively. Concurrent investigation of reduced nutrient levels indicated greater inhibition by HCA in a reduced nitrogen (N) treatment and in a reduced potassium (K) treatment relative to HCA inhibition in the full nutrient treatment. The negative slopes of the regressions of log of biomass on HCA concentration were steepest in the reduced N and reduced K treatments. Root and shoot biomasses in reduced N treatments were 20–43% and 24–34% less than the respective biomasses in the full nutrient treatment. Comparable reductions in the reduced K treatment were as much as 19% and 10% for root and shoot biomasses, respectively. The effects of HCA in a reduced phosphorus (P) treatment and in a reduced P and K treatment were not significantly different from the effects of HCA in the full nutrient treatment. Extraction of the soils at harvest indicated no buildup of HCA at the end of the experiment. The sensitivity ofSchizachyrium scoparium to HCA in general and increased sensitivity under low N and low K solutions may be important in the Florida scrub community where levels of N and K are known to be low.

Allelopathic potential of menthofuran monoterpenes fromCalamintha ashei

A reversed-phase HPLC analysis was used to separate and quantify five menthofuran monoterpenes inCalamintha ashei leaf soaks and washes. (+)-Evodone and desacetylcalaminthone were the major constituents of both soaks and washes. Concentrations of (+)-evodone and desacetylcalaminthone were as high as 0.66 and 0.74 mM, respectively, in leaf soaks. The highest concentration of monoterpenes in leaf washes obtained by misting was 0.021 mM. Aqueous solubilities of the menthofurans were determined to exceed concentrations required for growth inhibition. Bioassays of individualCalamintha monoterpenes demonstrated effects on germination as low as 0.05 mM for (+)-evodone. An equimolar mixture of desacetylcalaminthone and (+)-evodone reducedRudbeckia hirta germination by 17% at a combined concentration of 0.025 mM forLeptochloa dubia. Confirmation of allelopathic effects byCalamintha ashei will require long-term bioassays ofCalamintha menthofurans on the growth of native sandhill species under conditions comparable to the harsh environment of the Florida scrub.

Reproductive Phenology of Central Amazon Pioneer Trees

This study characterizes the flowering and fruiting phenology of the 13 most common pioneer tree species in early successional forests of the Central Amazon. For each species, 30 individuals, 10 each in three secondary forests, were monitored monthly for four years at the Biological Dynamics of Forest Fragments Project, north of Manaus. Five species showed nearly continuous flowering and fruiting throughout the study, indicating that resources were available to pollinators and dispersers on a regular basis. The other eight species showed stronger seasonality in reproduction, seven of them annually, and one supra-annually. Overall, flowering was concentrated in the transition from the dry to the rainy season and fruiting was concentrated in the rainy season. There was no relationship between reproductive phenology and tree pollinator type or dispersal mode. Reproductive phenology was remarkably consistent year to year. The pioneer community showed a variety of phenological patterns but as a whole tended to be characterized by annual flowering and fruiting, either continuously or seasonally, thereby fitting generalizations of pioneer species relative to mature forest species.