Variation in liana abundance and biomass along an elevational gradient in the tropical Atlantic Forest (Brazil)

Lianas play a key role in forest structure, species diversity, as well as functional aspects of tropical forests. Although the study of lianas in the tropics has increased dramatically in recent years, basic information on liana communities for the Brazilian Atlantic Forest is still scarce. To understand general patterns of liana abundance and biomass along an elevational gradient (0-1,100 m asl) of coastal Atlantic Forest, we carried out a standard census for lianas a parts per thousand yen1 cm in five 1-ha plots distributed across different forest sites. On average, we found a twofold variation in liana abundance and biomass between lowland and other forest types. Large lianas (a parts per thousand yen10 cm) accounted for 26-35% of total liana biomass at lower elevations, but they were not recorded in montane forests. Although the abundance of lianas displayed strong spatial structure at short distances, the present local forest structure played a minor role structuring liana communities at the scale of 0.01 ha. Compared to similar moist and wet Neotropical forests, lianas are slightly less abundant in the Atlantic Forest, but the total biomass is similar. Our study highlights two important points: (1) despite some studies have shown the importance of small-scale canopy disturbance and support availability, the spatial scale of the relationships between lianas and forest structure can vary greatly among tropical forests; (2) our results add to the evidence that past canopy disturbance levels and minimum temperature variation exert influence on the structure of liana communities in tropical moist forests, particularly along short and steep elevational gradients.

External drivers of biogeochemical cycles in a tropical montane forest in Ecuador

Successful conservation of tropical montane forest, one of the most threatened ecosystems on earth, requires detailed knowledge of its biogeochemistry. Of particular interest is the response of the biogeochemical element cycles to external influences such as element deposition or climate change. Therefore the overall objective of my study was to contribute to improved understanding of role and functioning of the Andean tropical montane forest. In detail, my objectives were to determine (1) the role of long-range transported aerosols and their transport mechanisms, and (2) the role of short-term extreme climatic events for the element budget of Andean tropical forest. In a whole-catchment approach including three 8-13 ha microcatchments under tropical montane forest on the east-exposed slope of the eastern cordillera in the south Ecuadorian Andes at 1850-2200 m above sea level I monitored at least in weekly resolution the concentrations and fluxes of Ca, Mg, Na, K, NO3-N, NH4-N, DON, P, S, TOC, Mn, and Al in bulk deposition, throughfall, litter leachate, soil solution at the 0.15 and 0.3 m depths, and runoff between May 1998 and April 2003. I also used meteorological data from my study area collected by cooperating researchers and the Brazilian meteorological service (INPE), as well as remote sensing products of the North American and European space agencies NASA and ESA. My results show that (1) there was a strong interannual variation in deposition of Ca [4.4-29 kg ha-1 a-1], Mg [1.6-12], and K [9.8-30]) between 1998 and 2003. High deposition changed the Ca and Mg budgets of the catchments from loss to retention, suggesting that the additionally available Ca and Mg was used by the ecosystem. Increased base metal deposition was related to dust outbursts of the Sahara and an Amazonian precipitation pattern with trans-regional dry spells allowing for dust transport to the Andes. The increased base metal deposition coincided with a strong La Niña event in 1999/2000. There were also significantly elevated H+, N, and Mn depositions during the annual biomass burning period in the Amazon basin. Elevated H+ deposition during the biomass burning period caused elevated base metal loss from the canopy and the organic horizon and deteriorated already low base metal supply of the vegetation. Nitrogen was only retained during biomass burning but not during non-fire conditions when deposition was much smaller. Therefore biomass burning-related aerosol emissions in Amazonia seem large enough to substantially increase element deposition at the western rim of Amazonia. Particularly the related increase of acid deposition impoverishes already base-metal scarce ecosystems. As biomass burning is most intense during El Niño situations, a shortened ENSO cycle because of global warming likely enhances the acid deposition at my study forest. (2) Storm events causing near-surface water flow through C- and nutrient-rich topsoil during rainstorms were the major export pathway for C, N, Al, and Mn (contributing >50% to the total export of these elements). Near-surface flow also accounted for one third of total base metal export. This demonstrates that storm-event related near-surface flow markedly affects the cycling of many nutrients in steep tropical montane forests. Changes in the rainfall regime possibly associated with global climate change will therefore also change element export from the study forest. Element budgets of Andean tropical montane rain forest proved to be markedly affected by long-range transport of Saharan dust, biomass burning-related aerosols, or strong rainfalls during storm events. Thus, increased acid and nutrient deposition and the global climate change probably drive the tropical montane forest to another state with unknown consequences for its functions and biological diversity.

Assessing land use and land cover change in tropical dry forest or Northern Chinandega, Nicaragus from 1985 to 2011

The objective of this study is to gain a quantitative understanding of land use and land cover change (LULCC) that have occurred in a rural Nicaraguan municipality by analyzing Landsat 5 Thematic Mapper (TM) images. By comparing the potential extent of tropical dry forest (TDF) with Landsat 5 TM images, this study analyzes the loss of this forest type on a local level for the municipality of San Juan de Cinco Pinos (63.5 km2) in the Department of Chinandega. Change detection analysis shows where and how land use has changed from 1985 to the present. From 1985 to 2011, nearly 15% of the TDF in San Juan de Cinco Pinos was converted to other land uses. Of the 1434.2 ha of TDF that was present in 1985, 1223.64 ha remained in 2011. The deforestation is primarily a result of agricultural expansion and fuelwood extraction. If current rates of TDF deforestation continue, the municipality faces the prospect of losing its forest cover within the next few decades.

Plurality of tree species responses to drought perturbation in Bornean tropical rain forest

Drought perturbation driven by the El Niño Southern Oscillation (ENSO) is a principal stochastic variable determining the dynamics of lowland rain forest in S.E. Asia. Mortality, recruitment and stem growth rates at Danum in Sabah (Malaysian Borneo) were recorded in two 4-ha plots (trees ≥ 10 cm gbh) for two periods, 1986–1996 and 1996–2001. Mortality and growth were also recorded in a sample of subplots for small trees (10 to <50 cm gbh) in two sub-periods, 1996–1999 and 1999–2001. Dynamics variables were employed to build indices of drought response for each of the 34 most abundant plot-level species (22 at the subplot level), these being interval-weighted percentage changes between periods and sub-periods. A significant yet complex effect of the strong 1997/1998 drought at the forest community level was shown by randomization procedures followed by multiple hypothesis testing. Despite a general resistance of the forest to drought, large and significant differences in short-term responses were apparent for several species. Using a diagrammatic form of stability analysis, different species showed immediate or lagged effects, high or low degrees of resilience or even oscillatory dynamics. In the context of the local topographic gradient, species’ responses define the newly termed perturbation response niche. The largest responses, particularly for recruitment and growth, were among the small trees, many of which are members of understorey taxa. The results bring with them a novel approach to understanding community dynamics: the kaleidoscopic complexity of idiosyncratic responses to stochastic perturbations suggests that plurality, rather than neutrality, of responses may be essential to understanding these tropical forests. The basis to the various responses lies with the mechanisms of tree-soil water relations which are physiologically predictable: the timing and intensity of the next drought, however, is not. To date, environmental stochasticity has been insufficiently incorporated into models of tropical forest dynamics, a step that might considerably improve the reality of theories about these globally important ecosystems.

The nitrogen cycle of tropical montane forest in Ecuador turns inorganic under environmental change

Water-bound nitrogen (N) cycling in temperate terrestrial ecosystems of the Northern Hemisphere is today mainly inorganic because of anthropogenic release of reactive N to the environment. In little-industrialized and remote areas, in contrast, a larger part of N cycling occurs as dissolved organic N (DON). In a north Andean tropical montane forest in Ecuador, the N cycle changed markedly during 1998–2010 along with increasing N deposition and reduced soil moisture. The DON concentrations and the fractional contribution of DON to total N significantly decreased in rainfall, throughfall, and soil solutions. This inorganic turn of the N cycle was most pronounced in rainfall and became weaker along the flow path of water through the system until it disappeared in stream water. Decreasing organic contributions to N cycling were caused not only by increasing inorganic N input but also by reduced DON production and/or enhanced DON decomposition. Accelerated DON decomposition might be attributable to less waterlogging and higher nutrient availability. Significantly increasing NO3-N concentrations and NO3-N/NH4-N concentration ratios in throughfall and litter leachate below the thick organic layers indicated increasing nitrification. In mineral soil solutions, in contrast, NH4-N concentrations increased and NO3-N/NH4-N concentration ratios decreased significantly, suggesting increasing net ammonification. Our results demonstrate that the remote tropical montane forests on the rim of the Amazon basin experienced a pronounced change of the N cycle in only one decade. This change likely parallels a similar change which followed industrialization in the temperate zone of the Northern Hemisphere more than a century ago.

Herbivores equalize the seedling height growth of three dominant tree species in an African tropical rain forest

Determining the impact of insect herbivores on forest tree seedlings and saplings is difficult without experimentation in the field. Moreover, this impact may be heterogeneous in time and space because of seasonal rainfall and canopy disturbances, or ‘gaps’, which can influence both insect abundance and plant performance. In this study we used fine netting to individually protect seedlings of Microberlinia bisulcata, Tetraberlinia bifoliolata and Tetraberlinia korupensis trees (Fabaceae = Leguminosae) from insects in 41 paired gap-understorey locations across 80 ha of primary rain forest (Korup, Cameroon). For all species, growth in height and leaf numbers was negligible in the understorey, where M. bisulcata had the lowest survival after c. 2 years. In gaps, however, all species responded positively with pronounced above-ground growth across seasons. When exposed to herbivores their seedling height growth was similar, but in the absence of herbivores, M. bisulcata significantly outgrew both Tetraberlinia species and matched their leaf numbers. This result suggests that insect herbivores might play an important role in maintaining species coexistence by mitigating sapling abundance of the more palatable M. bisulcata, which in gaps was eaten the most severely. The higher ratio in static leaf damage of control-to-caged M. bisulcata seedlings in gaps than understorey locations was consistent with the Plant Vigour Hypothesis. This result, however, did not apply to either Tetraberlinia species. For M. bisulcata and T. korupensis, but not T. bifoliolata (the most shade-tolerant species), caging improved relative seedling survival in the understory locations compared to gaps, providing restricted support for the Limiting Resource Model. Approximately 2.25 years after treatments were removed, the caged seedlings were taller and had more leaves than controls in all three species, and the effect remained strongest for M. bisulcata. We conclude that in this community the impact of leaf herbivory on seedling growth in gaps is strong for the dominant M. bisulcata, which coupled to a very low shade-tolerance contributes to limiting its regeneration. However, because gaps are common to most forests, insect herbivores may be having impacts upon functionally similar tree species that are also characterized by low sapling recruitment much more widely than currently appreciated. An implication for the restoration and management of M. bisulcata populations in forests outside of Korup is that physical protection from herbivores of new seedlings where the canopy is opened by gaps, or by harvesting, should substantially increase its subcanopy regeneration, and thus, too, its opportunities for adult recruitment.

Aluminum toxicity to tropical montane forest tree seedlings in southern Ecuador: response of biomass and plant morphology to elevated Al concentrations

In acid tropical forest soils (pH < 5.5) increased mobility of aluminum might limit aboveground productivity. Therefore, we evaluated Al phytotoxicity of three native tree species of tropical montane forests in southern Ecuador. An hydroponic dose-response experiment was conducted. Seedlings of Cedrela odorata L., Heliocarpus americanus L., and Tabebuia chrysantha (Jacq.) G. Nicholson were treated with 0, 300, 600, 1200, and 2400 mu M Al and an organic layer leachate. Dose-response curves were generated for root and shoot morphologic properties to determine effective concentrations (EC). Shoot biomass and healthy leaf area decreased by 44 % to 83 % at 2400 mu M Al, root biomass did not respond (C. odorata), declined by 51 % (H. americanus), or was stimulated at low Al concentrations of 300 mu M (T. chrysantha). EC10 (i.e. reduction by 10 %) values of Al for total biomass were 315 mu M (C. odorata), 219 mu M (H. americanus), and 368 mu M (T. chrysantha). Helicarpus americanus, a fast growing pioneer tree species, was most sensitive to Al toxicity. Negative effects were strongest if plants grew in organic layer leachate, indicating limitation of plant growth by nutrient scarcity rather than Al toxicity. Al toxicity occurred at Al concentrations far above those in native organic layer leachate.

Tree size and fecundity influence ballistic seed dispersal of two dominant mast-fruiting species in a tropical rain forest

Seed production, seed dispersal, and seedling recruitment are integral to forest dynamics, especially in masting species. Often these are studied separately, yet scarcely ever for species with ballistic dispersal even though this mode of dispersal is common in legume trees of tropical African rain forests. Here, we studied two dominant main-canopy tree species, Microberlinia bisulcata and Tetraberlinia bifoliolata (Caesalpinioideae), in 25 ha of primary rain forest at Korup, Cameroon, during two successive masting events (2007/2010). In the vicinity of c. 100 and 130 trees of each species, 476/580 traps caught dispersed seeds and beneath their crowns c. 57,000 pod valves per species were inspected to estimate tree-level fecundity. Seed production of trees increased non-linearly and asymptotically with increasing stem diameters. It was unequal within the two species’ populations, and differed strongly between years to foster both spatial and temporal patchiness in seed rain. The M. bisulcata trees could begin seeding at 42–44 cm diameter: at a much larger size than could T. bifoliolata (25 cm). Nevertheless, per capita life-time reproductive capacity was c. five times greater in M. bisulcata than T. bifoliolata owing to former’s larger adult stature, lower mortality rate (despite a shorter life-time) and smaller seed mass. The two species displayed strong differences in their dispersal capabilities. Inverse modelling (IM) revealed that dispersal of M. bisulcata was best described by a lognormal kernel. Most seeds landed at 10–15 m from stems, with 1% of them going beyond 80 m (<100 m). The direct estimates of fecundity significantly improved the models fitted. The lognormal also described well the seedling recruitment distribution of this species in 121 ground plots. By contrast, the lower intensity of masting and more limited dispersal of the heavier-seeded T. bifoliolata prevented reliable IM. For this species, seed density as function of distance to traps suggested a maximum dispersal distance of 40–50 m, and a correspondingly more aggregated seedling recruitment pattern ensued than for M. bisulcata. From this integrated field study, we conclude that the reproductive traits of M. bisulcata give it a considerable advantage over T. bifoliolata by better dispersing more seeds per capita to reach more suitable establishment sites, and combined with other key traits they explain its local dominance in the forest. Understanding the linkages between size at onset of maturity, individual fecundity, and dispersal capability can better inform the life-history strategies, and hence management, of co-occurring tree species in tropical forests.

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.

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.