Responses of transplanted native tree species to invasive alien grass removals in an abandoned cattle pasture in the Lacandon region, Mexico

We tested the early performance of 16 native early-, mid-, and late-successional tree species in response to four intensities of grass removal in an abandoned cattle pasture dominated by the introduced, invasive African grass, Cynodon plectostachyus, within the Lacandon rainforest region, southeast Mexico. The increase in grass removals significantly improved the performance of many species, especially of early-and mid-successional species, while performance of late-successional species was relatively poor and did not differ significantly among treatments. Good site preparation and at least one additional grass removal four months after seedling transplant were found to be essential; additional grass removals led to improved significantly performance of saplings in most cases. In order to evaluate the potential of transplanting tree seedlings successfully in abandoned tropical pastures, we developed a "planting risk index", combining field performance measurements and plantation cost estimations. Our results showed a great potential for establishing restoration plantings with many early-and mid-successional species. Although planting risk of late-successional species was considered high, certain species showed some possibilities of acclimation after 18 months and should be considered in future plantation arrangements in view of their long-term contributions to biodiversity maintenance and also to human welfare through delivery of ecosystem services. Conducting a planting risk analysis can help avoid failure of restoration strategies involving simultaneous planting of early-, mid-, and late-successional tree species. This in turn will improve cost-effectiveness of initial interventions in large-scale, long-term restoration programs.

Improving Planting Stocks for the Brazilian Atlantic Forest Restoration through Community-Based Seed Harvesting Strategies

High-diversity reforestation can help jumpstart tropical forest restoration, but obtaining viable seedlings is a major constraint: if nurseries do not offer them, it is hard to plant all the species one would like. From 2007 to 2009, we investigated five different seed acquisition strategies employed by a well-established tree nursery in southeastern Brazil, namely (1) in-house seed harvesters; (2) hiring a professional harvester; (3) amateur seed harvesters; or (4) a seed production cooperative, as well as (5) participating in a seed exchange program. In addition, we evaluated two strategies not dependent on seeds: harvesting seedlings from native tree species found regenerating under Eucalyptus plantations, and in a native forest remnant. A total of 344 native tree and shrub species were collected as seeds or seedlings, including 2,465 seed lots. Among these, a subset of 120 species was obtained through seed harvesting in each year. Overall, combining several strategies for obtaining planting stocks was an effective way to increase species richness, representation of some functional groups (dispersal syndromes, planting group, and shade tolerance), and genetic diversity of seedlings produced in forest tree nurseries. Such outcomes are greatly desirable to support high-diversity reforestation as part of tropical forest restoration. In addition, community-based seed harvesting strategies fostered greater socioeconomic integration of traditional communities in restoration projects and programs, which is an important bottleneck for the advance of ecological restoration, especially in developing countries. Finally, we discuss some of the limitations of the various strategies for obtaining planting stocks and the way forward for their improvement.

Emissions of volatile organic compounds from tropical savanna vegetation and biomass burning

This dissertation focuses on characterizing the emissions of volatile organic compounds (VOCs) from grasses and young trees, and the burning of biomass mainly from Africa and Indonesia. The measurements were performed with a proton-transfer-reaction mass spectrometer (PTR-MS). The biogenic emissions of tropical savanna vegetation were studied in Calabozo (Venezuela). Two field campaigns were carried out, the first during the wet season (1999) and the second during the dry season (2000). Three grass species were studied: T. plumosus, H. rufa and A. canescens, and the tree species B. crassifolia, C. americana and C. vitifolium. The emission rates were determined with a dynamic plant enclosure system. In general, the emissions increased exponentially with increasing temperature and solar radiation. Therefore, the emission rates showed high variability. Consequently, the data were normalized to a standard temperature of 30°C, and standard emission rates thus determined allowed for interspecific and seasonal comparisons. The range of average daytime (10:00-16:00) emission rates of total VOCs measured from green (mature and young) grasses was between 510-960 ngC/g/h. Methanol was the primary emission (140-360 ngC/g/h), followed by acetaldehyde, butene and butanol and acetone with emission rates between 70-200 ngC/g/h. The emissions of propene and methyl ethyl ketone (MEK) were <80 ngC/g/h, and those of isoprene and C5-alcohols were between 10-130 ngC/g/h. The oxygenated species represented 70-75% of the total. The emission of VOCs was found to vary by up to a factor of three between plants of the same species, and by up to a factor of two between the different species. The annual source of methanol from savanna grasses worldwide estimated in this work was 3 to 4.4 TgC, which could represent up to 12% of the current estimated global emission from terrestrial vegetation. Two of the studied tree species, were isoprene emitters, and isoprene was also their primary emission (which accounted for 70-94% of the total carbon emitted) followed by methanol and butene + butanol. The daytime average emission rate of isoprene measured in the wet season was 27 mgC/g/h for B. crassifolia, and 123 mgC/g/h for C. vitifolium. The daytime emissions of methanol and butene + butanol were between 0.3 and 2 mgC/g/h. The total sum of VOCs emission measured during the day in the wet season was between 30 and 130 mgC/g/h. In the dry season, in contrast, the methanol emissions from C. vitifolium saplings –whose leaves were still developing– were an order of magnitude higher than in the wet season (15 mgC/g/h). The isoprene emission from B. crassifolia in the dry season was comparable to the emission in the wet season, whereas isoprene emission from C. vitifolium was about a factor of three lower (~43 mgC/g/h). Biogenic emission inventories show that isoprenoids are the most prominent and best-studied compounds. The standard emission rates of isoprene and monoterpenes of the measured savanna trees were in the lower end of the range found in the literature. The emission of other biogenic VOCs has been sparsely investigated, but in general, the standard emissions from trees studied here were within the range observed in previous investigations. The biomass burning study comprised the measurement of VOCs and other trace-gas emissions of 44 fires from 15 different fuel types, primarily from Africa and Indonesia, in a combustion laboratory. The average sum of emissions (excluding CO2, CO and NO) from African fuels was ~18 g(VOC)/kg. Six of the ten most important emissions were oxygenated VOCs. Acetic acid was the major emission, followed by methanol and formaldehyde. The emission of methane was of the same order as the methanol emission (~5 g/kg), and that of nitrogen-containing compounds was ~1 g/kg. An estimate of the VOC source from biomass burning of savannas and grasslands worldwide suggests that the sum of emissions is about 56 Tg/yr, of which 34 Tg correspond to oxygenated VOCs, 14 Tg to unsaturated and aromatic compounds, 5 Tg to methane and 3 Tg to N-compounds. The estimated emissions of CO, CO2 and NO are 216, 5117 and 9.4 Tg/yr, respectively. The emission factors reported here for Indonesian fuels are the first results of laboratory fires using Indonesian fuels. Acetic acid was the highest organic emission, followed by acetol, a compound not previously reported in smoke, methane, mass 97 (tentatively identified as furfural, dimethylfuran and ethylfuran), and methanol. The sum of total emissions of Indonesian fuels was 91 g/kg, which is 5 times higher than the emissions from African fuels. The results of this study reinforces the importance of oxygenated compounds. Due to the vast area covered by tropical savannas worldwide, the biogenic and biomass burning emission of methanol and other oxygenated compounds may be important for the regional and even global tropospheric chemistry.

Island Invasion by a Threatened Tree Species: Evidence for Natural Enemy Release of Mahogany (Swietenia macrophylla) on Dominica, Lesser Antilles

Despite its appeal to explain plant invasions, the enemy release hypothesis (ERH) remains largely unexplored for tropical forest trees. Even scarcer are ERH studies conducted on the same host species at both the community and biogeographical scale, irrespective of the system or plant life form. In Cabrits National Park, Dominica, we observed patterns consistent with enemy release of two introduced, congeneric mahogany species, Swietenia macrophylla and S. mahagoni, planted almost 50 years ago. Swietenia populations at Cabrits have reproduced, with S. macrophylla juveniles established in and out of plantation areas at densities much higher than observed in its native range. Swietenia macrophylla juveniles also experienced significantly lower leaf-level herbivory (~3.0%) than nine co-occurring species native to Dominica (8.4–21.8%), and far lower than conspecific herbivory observed in its native range (11%–43%, on average). These complimentary findings at multiple scales support ERH, and confirm that Swietenia has naturalized at Cabrits. However, Swietenia abundance was positively correlated with native plant diversity at the seedling stage, and only marginally negatively correlated with native plant abundance for stems ≥1-cm dbh. Taken together, these descriptive patterns point to relaxed enemy pressure from specialized enemies, specifically the defoliator Steniscadia poliophaea and the shoot-borer Hypsipyla grandella, as a leading explanation for the enhanced recruitment of Swietenia trees documented at Cabrits.

Effects of experimental nitrogen additions on plant diversity in an old-growth tropical forest

Response of plant biodiversity to increased availability of nitrogen (N) has been investigated in temperate and boreal forests, which are typically N-limited, but little is known in tropical forests. We examined the effects of artificial N additions on plant diversity (species richness, density and cover) of the understory layer in an N saturated old-growth tropical forest in southern China to test the following hypothesis: N additions decrease plant diversity in N saturated tropical forests primarily from N-mediated changes in soil properties. Experimental additions of N were administered at the following levels from July 2003 to July 2008: no addition (Control); 50 kg N ha−1 yr−1 (Low-N); 100 kg N ha−1 yr−1 (Medium-N), and 150 kg N ha−1 yr−1 (High-N). Results showed that no understory species exhibited positive growth response to any level of N addition during the study period. Although low-to-medium levels of N addition (≤100 kg N ha−1 yr−1) generally did not alter plant diversity through time, high levels of N addition significantly reduced species diversity. This decrease was most closely related to declines within tree seedling and fern functional groups, as well as to significant increases in soil acidity and Al mobility, and decreases in Ca availability and fine-root biomass. This mechanism for loss of biodiversity provides sharp contrast to competition-based mechanisms suggested in studies of understory communities in other forests. Our results suggest that high-N additions can decrease plant diversity in tropical forests, but that this response may vary with rate of N addition.

Seedling survival and growth of three ectomycorrhizal caesalpiniaceous tree species in a Central African rain forest

Tree recruitment is determined in part by the survivorship and growth of seedlings. Two seedling cohorts of the three most abundant caesalpiniaceous species forming groves at Korup, Cameroon, were followed from 1995/1997 to 2002, to investigate why Microberlinia bisulcata, the most abundant species, currently has very few recruits compared with Tetraberlinia korupensis and T. bifoliolata. Numbers of seedlings dying, and the heights and leaf numbers of survivors, were recorded on 30 occasions. Survivorship after 2.5 y was 30% for M. bisulcata and 59% for the similar Tetraberlinia spp. together. After 7 y the corresponding values were 4 and 21%. Growth of all species was slow for the first 4 y; but survivors of T. korupensis became 63% taller, as the other species stagnated, by 7 y. The poor recruitment of M. bisulcata was the result of its very low seedling survival. Within species, the tallest seedlings of M. bisulcata and T. bifoliolata, but medium-height ones of T. korupensis, survived longest. This was likely due to higher root allocation in T. korupensis. Seedling dynamics of M. bisulcata and T. korupensis over 7 y accorded well with relative abundances of adult trees; T. bifoliolata is predicted to recruit later.

On the detection of dynamic responses in a drought-perturbed tropical rainforest in Borneo

The dynamics of aseasonal lowland dipterocarp forest in Borneo is influenced by perturbation from droughts. These events might be increasing in frequency and intensity in the future. This paper describes drought-affected dynamics between 1986 and 2001 in Sabah, Malaysia, and considers how it is possible, reliably and accurately, to measure both coarse- and fine-scale responses of the forest. Some fundamental concerns about methodology and data analysis emerge. In two plots forming 8 ha, mortality, recruitment, and stem growth rates of trees ≥10 cm gbh (girth at breast height) were measured in a ‘pre-drought’ period (1986–1996), and in a period (1996–2001) including the 1997–1998 ENSO-drought. For 2.56 ha of subplots, mortality and growth rates of small trees (10–<50 cm gbh) were found also for two sub-periods (1996–1999, 1999–2001). A total of c. 19 K trees were recorded. Mortality rate increased by 25% while both recruitment and relative growth rates increased by 12% for all trees at the coarse scale. For small trees, at the fine scale, mortality increased by 6% and 9% from pre-drought to drought and on to ‘post-drought’ sub-periods. Relative growth rates correspondingly decreased by 38% and increased by 98%. Tree size and topography interacted in a complex manner with between-plot differences. The forest appears to have been sustained by off-setting elevated tree mortality by highly resilient stem growth. This last is seen as the key integrating tree variable which links the external driver (drought causing water stress) and population dynamics recorded as mortality and recruitment. Suitably sound measurements of stem girth, leading to valid growth rates, are needed to understand and model tree dynamic responses to perturbations. The proportion of sound data, however, is in part determined by the drought itself.

Spectral biases in tree-ring climate proxies

External forcing and internal dynamics result in climate system variability ranging from sub-daily weather to multi-centennial trends and beyond1, 2. State-of-the-art palaeoclimatic methods routinely use hydroclimatic proxies to reconstruct temperature (for example, refs 3, 4), possibly blurring differences in the variability continuum of temperature and precipitation before the instrumental period. Here, we assess the spectral characteristics of temperature and precipitation fluctuations in observations, model simulations and proxy records across the globe. We find that whereas an ensemble of different general circulation models represents patterns captured in instrumental measurements, such as land–ocean contrasts and enhanced low-frequency tropical variability, the tree-ring-dominated proxy collection does not. The observed dominance of inter-annual precipitation fluctuations is not reflected in the annually resolved hydroclimatic proxy records. Likewise, temperature-sensitive proxies overestimate, on average, the ratio of low- to high-frequency variability. These spectral biases in the proxy records seem to propagate into multi-proxy climate reconstructions for which we observe an overestimation of low-frequency signals. Thus, a proper representation of the high- to low-frequency spectrum in proxy records is needed to reduce uncertainties in climate reconstruction efforts.

Current and future variations of nutrient depositions and influences on tree growth

Current nutrient deposition shows episodic variations which likely may impact the local nutrient cycle at the RBSF. Comparing analyses of deposition data during present-day atmospheric circulation and phases of high biomass burning in the Amazon, characteristic relationships between remote emissions and local deposition are determined. By using projections drawn from the special report on emission scenarios (SRES) in combination with a trajectory modeling tool, future nutrient deposition conditions of the mountain ecosystem are assessed. Observations of relations between climatic variables, current time series of nutrient deposition, and tree growth point to an impact of the remote fertilization effect of atmospheric matters, emitted primarily by human activities like biomass burning and agricultural and industrial sources. The increasing emissions in the future may have adverse effects on the ecosystem in the long run.

The dating of dipterocarp tree rings: establishing a record of carbon cycling and climatic change in the tropics

In a first step to obtain a proxy record of past climatic events (including the El Ni (n) over tildeo-Southern Oscillation) in the normally aseasonal tropical environment of Sabah, a radial segment from a recently fallen dipterocarp (Shorea Superba) was radiocarbon dated and subjected to carbon isotope analysis. The high-precision radiocarbon results fell into the ambiguous modern plateau where several calibrated dates can exist for each sample. Dating was achieved by wiggle matching using a Bayesian approach to calibration. Using the defined growth characteristics of Shorea superba, probability density distributions were calculated and improbable dates rejected. It was found that the tree most likely started growing around AD 1660-1685. A total of 173 apparent growth increments were measured and, therefore, it could be determined that the tree formed one ring approximately every two years. Stable carbon isotope values were obtained from resin-extracted wholewood from each ring. Carbon cycling is evident in the `juvenile effect', resulting from the assimilation of respired carbon dioxide and lower light levels below the canopy, and in the `anthropogenic effect' caused by increased industrial activity in the late-nineteenth and twentieth centuries. This study demonstrates that palaeoenvironmental information can be obtained from trees growing in aseasonal environments, where climatic conditions prevent the formation of well-defined annual rings.

Aluminum toxicity in a tropical montane forest ecosystem in southern Ecuador

Aluminum phytotoxicity frequently occurs in acid soils (pH < 5.5) and was therefore discussed to affect ecosystem functioning of tropical montane forests. The susceptibility to Al toxicity depends on the sensitivity of the plant species and the Al speciation in soil solution, which can vary highly depending e.g., on pH, ionic strength, and dissolved organic matter. An acidification of the ecosystem and periodic base metal deposition from Saharan dust may control plant available Al concentrations in the soil solutions of tropical montane rainforests in south Ecuador. The overall objective of my study was to assess a potential Al phytotoxicity in the tropical montane forests in south Ecuador. For this purpose, I exposed three native Al non-accumulating tree species (Cedrela odorata L., Heliocarpus americanus L., and Tabebuia chrysantha (Jacq.) G. Nicholson) to increased Al concentrations (0 – 2400 μM Al) in a hydroponic experiment, I established dose-response curves to estimate the sensitivity of the tree species to increased Al concentrations, and I investigated the mechanisms behind the observed effects induced by elevated Al concentrations. Furthermore, the response of Al concentrations and the speciation in soil solution to Ca amendment in the study area were determined. In a final step, I assessed all major Al fluxes, drivers of Al concentrations in ecosystem solutions, and indicators of Al toxicity in the tropical montane rainforest in Ecuador in order to test for indications of Al toxicity. In the hydroponic experiment, a 10 % reduction in aboveground biomass production occurred at 126 to 376 μM Al (EC10 values), probably attributable to decreased Mg concentrations in leaves and reduced potosynthesis. At 300 μM Al, increased root biomass production of T. chrysantha was observed. Phosphorus concentrations in roots of C. odorata and T. chrysantha were significantly highest in the treatment with 300 μM Al and correlated significantly with root biomass, being a likely reason for stimulated root biomass production. The degree of organic complexation of Al in the organic layer leachate, which is central to plant nutrition because of the high root density, and soil solution from the study area was very high (mean > 99 %). The resulting low free Al concentrations are not likely to affect plant growth, although the concentrations of potentially toxic Al3+ increased with soil depth due to higher total Al and lower dissolved organic matter concentrations in soil solutions. The Ca additions caused an increase of Al in the organic layer leachate, probably because Al3+ was exchanged against the added Ca2+ ions while pH remained constant. The free ion molar ratios of Ca2+:Al3+ (mean ratio ca. 400) were far above the threshold (≤ 1) for Al toxicity, because of a much higher degree of organo-complexation of Al than Ca. High Al fluxes in litterfall (8.8 – 14.2 kg ha−1 yr−1) indicate a high Al circulation through the ecosystem. The Al concentrations in the organic layer leachate were driven by the acidification of the ecosystem and increased significantly between 1999 and 2008. However, the Ca:Al molar ratios in organic layer leachate and all aboveground ecosystem solutions were above the threshold for Al toxicity. Except for two Al accumulating and one non-accumulating tree species, the Ca:Al molar ratios in tree leaves from the study area were above the Al toxicity threshold of 12.5. I conclude that toxic effects in the hydroponic experiment occurred at Al concentrations far above those in native organic layer leachate, shoot biomass production was likely inhibited by reduced Mg uptake, impairing photosynthesis, and the stimulation of root growth at low Al concentrations can be possibly attributed to improved P uptake. Dissolved organic matter in soil solutions detoxifies Al in acidic tropical forest soils and a wide distribution of Al accumulating tree species and high Al fluxes in the ecosystem do not necessarily imply a general Al phytotoxicity.

Effects of elevation and land use on the biomass of trees, shrubs and herbs at Mount Kilimanjaro

The protection and sustainable management of forest carbon stocks, particularly in the tropics, is a key factor in the mitigation of global change effects. However, our knowledge of how land use and elevation affect carbon stocks in tropical ecosystems is very limited. We compared aboveground biomass of trees, shrubs and herbs for eleven natural and human-influenced habitat types occurring over a wide elevation gradient (866–4550 m) at the world's highest solitary mountain, Mount Kilimanjaro. Thanks to the enormous elevation gradient, we covered important natural habitat types, e.g., savanna woodlands, montane rainforest and afro-alpine vegetation, as well as important land-use types such as maize fields, grasslands, traditional home gardens, coffee plantations and selectively logged forest. To assess tree and shrub biomass with pantropical allometric equations, we measured tree height, diameter at breast height and wood density and to assess herbaceous biomass, we sampled destructively. Among natural habitats, tree biomass was highest at intermediate elevation in the montane zone (340 Mg ha−1), shrub biomass declined linearly from 7 Mg ha−1 at 900 m to zero above 4000 m, and, inverse to tree biomass, herbaceous biomass was lower at mid-elevations (1 Mg ha−1) than in savannas (900 m, 3 Mg ha−1) or alpine vegetation (above 4000 m, 6 Mg ha−1). While the various land-use types dramatically decreased woody biomass at all elevations, though to various degrees, herbaceous biomass was typically increased. Our study highlights tropical montane forest biomass as important aboveground carbon stock and quantifies the extent of the strong aboveground biomass reductions by the major land-use types, common to East Africa. Further, it shows that elevation and land use differently affect different vegetation strata, and thus the matrix for other organisms.

Simulating Carbon Stocks and Fluxes of an African Tropical Montane Forest with an Individual-Based Forest Model

Tropical forests are carbon-dense and highly productive ecosystems. Consequently, they play an important role in the global carbon cycle. In the present study we used an individual-based forest model (FORMIND) to analyze the carbon balances of a tropical forest. The main processes of this model are tree growth, mortality, regeneration, and competition. Model parameters were calibrated using forest inventory data from a tropical forest at Mt. Kilimanjaro. The simulation results showed that the model successfully reproduces important characteristics of tropical forests (aboveground biomass, stem size distribution and leaf area index). The estimated aboveground biomass (385 t/ha) is comparable to biomass values in the Amazon and other tropical forests in Africa. The simulated forest reveals a gross primary production of 24 tcha-1yr-1. Modeling above- and belowground carbon stocks, we analyzed the carbon balance of the investigated tropical forest. The simulated carbon balance of this old-growth forest is zero on average. This study provides an example of how forest models can be used in combination with forest inventory data to investigate forest structure and local carbon balances.

Using plant functional traits and phylogenies to understand patterns of plant community assembly in a seasonal tropical forest in Lao PDR

Plant functional traits reflect different evolutionary responses to environmental variation, and among extant species determine the outcomes of interactions between plants and their environment, including other plant species. Thus, combining phylogenetic and trait-based information can be a powerful approach for understanding community assembly processes across a range of spatial scales. We used this approach to investigate tree community composition at Phou Khao Khouay National Park (18°14’-18°32’N; 102°38’- 102°59’E), Laos, where several distinct forest types occur in close proximity. The aim of our study was to examine patterns of plant community assembly across the strong environmental gradients evident at our site. We hypothesized that differences in tree community composition were being driven by an underlying gradient in soil conditions. Thus, we predicted that environmental filtering would predominate at the site and that the filtering would be strongest on sandier soil with low pH, as these are the conditions least favorable to plant growth. We surveyed eleven 0.25 ha (50x50 m) plots for all trees above 10 cm dbh (1221 individual trees, including 47 families, 70 genera and 123 species) and sampled soils in each plot. For each species in the community, we measured 11 commonly studied plant functional traits covering both the leaf and wood economic spectrum traits and we reconstructed a phylogenetic tree for 115 of the species in the community using rbcL and matK sequences downloaded from Genebank (other species were not available). Finally we compared the distribution of trait values and species at two scales (among plots and 10x10m subplots) to examine trait and phylogenetic community structures. Although there was strong evidence that an underlying soil gradient was determining patterns of species composition at the site, our results did not support the hypothesis that the environmental filtering dominated community assembly processes. For the measured plant functional traits there was no consistent pattern of trait dispersion across the site, either when traits were considered individually or when combined in a multivariate analysis. However, there was a significant correlation between the degree of phylogenetic dispersion and the first principle component axis (PCA1) for the soil parameters.Moreover, the more phylogenetically clustered plots were on sandier soils with lower pH. Hence, we suggest that the community assembly processes across our sitemay reflect the influence ofmore conserved traits that we did not measure. Nevertheless, our results are equivocal and other interpretations are possible. Our study illustrates some difficulties in combining trait and phylogenetic approaches that may result from the complexities of integrating spatial and evolutionary processes that vary at different scales.

Mapping tree density at a global scale

The global extent and distribution of forest trees is central to our understanding of the terrestrial biosphere. We provide the first spatially continuous map of forest tree density at a global scale. This map reveals that the global number of trees is approximately 3.04 trillion, an order of magnitude higher than the previous estimate. Of these trees, approximately 1.39 trillion exist in tropical and subtropical forests, with 0.74 trillion in boreal regions and 0.61 trillion in temperate regions. Biome-level trends in tree density demonstrate the importance of climate and topography in controlling local tree densities at finer scales, as well as the overwhelming effect of humans across most of the world. Based on our projected tree densities, we estimate that over 15 billion trees are cut down each year, and the global number of trees has fallen by approximately 46% since the start of human civilization.