Juan Carlos Licona

Does functional trait diversity predict above-ground biomass and productivity of tropical forests? Testing three alternative hypotheses

Summary 1. Tropical forests are globally important, but it is not clear whether biodiversity enhances carbon storage and sequestration in them. We tested this relationship focusing on components of functional trait biodiversity as predictors. 2. Data are presented for three rain forests in Bolivia, Brazil and Costa Rica. Initial above-ground biomass and biomass increments of survivors, recruits and survivors + recruits (total) were estimated for trees ≥10 cm d.b.h. in 62 and 21 1.0-ha plots, respectively. We determined relationships of biomass increments to initial standing biomass (AGBi), biomass-weighted community mean values (CWM) of eight functional traits and four functional trait variety indices (functional richness, functional evenness, functional diversity and functional dispersion). 3. The forest continuum sampled ranged from ‘slow’ stands dominated by trees with tough tissues and high AGBi ,t o‘fast’ stands dominated by trees with soft, nutrient-rich leaves, lighter woods and lower AGBi. 4. We tested whether AGBi and biomass increments were related to the CWM trait values of the dominant species in the system (the biomass ratio hypothesis), to the variety of functional trait values (the niche complementarity hypothesis), or in the case of biomass increments, simply to initial standing biomass (the green soup hypothesis). 5. CWMs were reasonable bivariate predictors of AGBi and biomass increments, with CWM specific leaf area SLA, CWM leaf nitrogen content, CWM force to tear the leaf, CWM maximum adult height Hmax and CWM wood specific gravity the most important. AGBi was also a reasonable predictor of the three measures of biomass increment. In best-fit multiple regression models, CWM Hmax was the most important predictor of initial standing biomass AGBi. Only leaf traits were selected in the best models for biomass increment; CWM SLA was the most important predictor, with the expected positive relationship. There were no relationships of functional variety indices to biomass increments, and AGBi was the only predictor for biomass increments from recruits. 6. Synthesis. We found no support for the niche complementarity hypothesis and support for the green soup hypothesis only for biomass increments of recruits. We have strong support for the biomass ratio hypothesis. CWM Hmax is a strong driver of ecosystem biomass and carbon storage and CWM SLA, and other CWM leaf traits are especially important for biomass increments and carbon sequestration.

Natural regeneration and liberation of timber species in logging gaps in two Bolivian tropical forests

One-year-old logging gaps were sampled in dry and humid selectively-logged Bolivian tropical forests to determine the density of commercial tree regeneration. The efficacy of liberation treatments designed to enhance the growth and survival of sapling regeneration was evaluated over a period of 2 years. Liberation treatments included manual cleaning of competing vegetation around commercial tree saplings occurring in logging gaps, or spraying competing vegetation near these saplings with one of two herbicides (2,4-D or glyphosate). Of the nine species harvested in each forest type, only two species in each type had relatively abundant regeneration in logging gaps (Hura crepitans L. and Schizolobium parahyba (Vell. Conc.) S.F. Blake in the humid forest and Centrolobium microchaete C. Martius ex. Benth. and Caesalpinia pluviosa DC. in the dry forest). After 1 year, liberation treatments significantly increased the diameter growth of saplings in the humid forest, but not during the second year (P=0.09). In the dry forest, liberation treatments did not affect the diameter of liberated plants. Liberation treatments did not significantly affect sapling survival or height growth in either forest. Treatment costs were relatively low (US

Climate and soil drive forest structure in Bolivian lowland forests

1–2.3 per gap), but time until return on investment is long (20–30-year cutting cycles). Adjustments to liberation treatments, such as season and intensity of application may increase effectiveness of treatments. Problems observed in this study with the recruitment of commercial tree regeneration in logging gaps suggests the need for site preparation treatments and more judicious seed tree retention.

Invasion of Non-Commercial Tree Species After Selection Logging in a Bolivian Tropical Forest

Climate is one of the most important factors determining variation in forest structure, but whether soils have independent effects is less clear. We evaluate how climate and soil independently affect forest structure, using 89 200 stems = 10 cm dbh from 220 1-ha permanent plots distributed along environmental gradients in lowland Bolivia. Fifteen forest structural variables, related to vertical structure (forest height and layering), horizontal structure (basal area, median and the 99th percentile of the stem diameter and size-class distribution) and density of life forms (tree, palm and liana), were evaluated. Environmental variables were summarized in four multivariate axes, related to rainfall, temperature, soil fertility and soil texture. Multiple regression indicates that all structural variables were affected by one or more of the environmental axes, but the explained variation was generally low (median R2 = 0.15). Rainfall and soil texture affected most forest structural variables (respectively 87% and 80%) and had qualitatively similar effects. This suggests that plant water availability, as determined by rainfall and soil water retention capacity, is the strongest driver of forest structure, whereas soil fertility was a weaker driver of forest structure, affecting 53% of the variables. Maximum forest height, palm density, total basal area and liana infestation showed the strongest responses to environmental variation (with R2 ranging from 0.31–0.82). Forest height, palm density and total basal area increased with plant water availability, while liana infestation decreased with plant water availability. Therefore, multiple rather than single environmental factors must be used to explain the structure of tropical forests.

Old-growth Neotropical forests are shifting in species and trait composition

Abstract Single-tree selection logging will likely result in a 4.3% loss in the relative abundance and a 4.1% loss in basal area of trees of commercial species in one cutting cycle due to their removal by harvesting combined with their potential recolonization of only 31% of logging gaps in a Bolivian tropical dry forest. Densities of the most valuable species, Amburana cearensis and Cedrela fissilis, were particularly reduced by logging. To sustain the current harvesting rate, uses need to be developed for more non-commercial species and/or silvicul-tural treatments employed that increase regeneration of commercial species and remove non-commercial species using timber stand improvement techniques.

Abiotic and biotic drivers of biomass change in a Neotropical forest

Tropical forests have long been thought to be in stable state, but recent insights indicate that global change is leading to shifts in forest dynamics and species composition. These shifts may be driven by environmental changes such as increased resource availability, increased drought stress, and/or recovery from past disturbances. The relative importance of these drivers can be inferred from analyzing changes in trait values of tree communities. Here, we evaluate a decade of change in species and trait composition across five old-growth Neotropical forests in Bolivia, Brazil, Guyana, and Costa Rica that cover large gradients in rainfall and soil fertility. To identify the drivers of compositional change, we used data from 29 permanent sample plots and measurements of 15 leaf, stem, and whole-plant traits that are important for plant performance and should respond to global change drivers. We found that forests differ strongly in their community-mean trait values, resulting from differences in soil fertility and annual rainfall seasonality. The abundance of deciduous species with high specific leaf area increases from wet to dry forests. The community-mean wood density is high in the driest forests to protect xylem vessels against drought cavitation, and is high in nutrient-poor forests to increase wood longevity and enhance nutrient residence time in the plant. Interestingly, the species composition changed over time in three of the forests, and the community-mean wood density increased and the specific leaf area decreased in all forests, indicating that these forests are changing toward later successional stages dominated by slow-growing, shade-tolerant species. We did not see changes in other traits that could reflect responses to increased drought stress, such as increased drought deciduousness or decreased maximum adult size, or that could reflect increased resource availability (CO2, rainfall, or nitrogen). Changes in species and trait composition in these forests are therefore most likely caused by recovery from past disturbances. These compositional changes may also lead to shifts in ecosystem processes, such as a lower carbon sequestration and “slower” forest dynamics.

Assessing the extent of “conflict of use” in multipurpose tropical forest trees: A regional view

Abiotic and biotic variables and growth, recruitment and mortality for 48 1-ha plots in a moist tropical forest in Bolivia

Beyond reduced-impact logging: Silvicultural treatments to increase growth rates of tropical trees

In the context of multiple forest management, multipurpose tree species which provide both timber and non-timber forest products (NTFP), present particular challenges as the potential of conflicting use for either product may be high. One key aspect is that the magnitude of conflict of use can be location specific, thus adding complexity to policy development. This paper focuses on the extent to which the potential for conflict of use in multipurpose tree species varies across the Amazonian lowland forests shared by Peru, Bolivia, Colombia, Ecuador and Venezuela, emphasizing the economic dimension of conflict. Based on a review of the current normative and regulatory aspects of timber and NTFP extraction in the five countries, the paper also briefly discusses the opportunities and constraints for harmonization of timber and NTFP management of multipurpose species across the region. It was found that about half of the 336 timber species reviewed across the five countries also have non-timber uses. Eleven timber species are multipurpose in all five countries: Calophyllum brasiliense, Cedrela odorata, Ceiba pentandra, Clarisia racemosa, Ficus insipida, Jacaranda copaia, Schefflera morototoni, Simarouba amara and Terminalia amazonia. Seven other multipurpose species occurred only in either Venezuela (Tabebuia impetiginosa, Spondias mombin, Pentaclethra macroloba, Copaifera officinalis, Chlorophora tinctoria, Carapa guianensis) or Ecuador (Tabebuia chrysantha). Four multipurpose tree species presented the highest potential of conflict of use across the region: Dipteryx odorata, Tabebuia serratifolia, Hymenaea courbaril and Myroxylon balsamum yet these were not evenly distributed across all five countries. None of the five studied countries have specific legislation to promote sustainable use of any of the multipurpose species reported here and thus mitigate potential conflict of use; nor documented management options for integration or else segregation of both their timber and NTFP values.