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.

Recruitment dynamics of the grove-dominant tree Microberlinia bisulcata in African rain forest: extending the light response versus adult longevity trade-off concept

In groves of ectomycorrhizal caesalpiniaceous species in the Atlantic coastal forest of Central Africa the dominant tree Microberlinia bisulcata, which is shade-intolerant as a seedling but highly light-responding as a sapling, shows very limited regeneration. M. bisulcata saplings were mapped in an 82.5-ha plot at Korup and found to be located significantly far (>40 m) away from adults, a result confirmed by direct testing in a second 56-ha plot. Sapling growth over 6 years, the distribution of newly emerging seedlings around adults, recruitment of saplings in a large opening and the outward extent of seedlings at the grove edge were also investigated. Two processes appear to have been operating: (1) a very strong and consistent restriction of the very numerous seedlings establishing after masting close to adults, and (2) a strong but highly spatially variable promotion of distant survivors by increased light from the deaths of large trees of species other than M. bisulcata (which itself has very low mortality rate). This leads to an apparent escape-from-adults effect. To maintain saplings in the shade between multiple short periods of release ectomycorrhizal connections to other co-occurring caesalp species may enable a rachet-type mechanism. The recorded sapling dynamics currently contribute an essential part of the long-term cycling of the groves. M. bisulcata is an interesting example of an important group of tropical trees, particularly in Africa, which are both highly light-demanding when young yet capable also of forming very large forest emergents. To more comprehensively explain tropical tree responses, the case is made for adding a new dimension to the trade-off concept of early tree light-response versus adult longevity.

Buttress form of the central African rain forest tree Microberlinia bisulcata, and its possible role in nutrient acquisition

Buttressing is a trait special to tropical trees but explanations for its occurrence remain inconclusive. The two main hypotheses are that they provide structural support and/or promote nutrient acquisition. Studies of the first are common but the second has received much less attention. Architectural measurements were made on adult and juvenile trees of the ectomycorrhizal species Microberlinia bisulcata, in Korup (Cameroon). Buttressing on this species is highly distinctive with strong lateral extension of surface roots of the juveniles leading to a mature buttress system of a shallow spreading form on adults. This contrasts with more vertical buttresses, closer to the stem, found on many other tropical tree species. No clear relationship between main buttress and large branch distribution was found. Whilst this does not argue against the essential structural role of buttresses for these very large tropical trees, the form on M. bisulcata does suggest a likely second role, that of aiding nutrient acquisition. At the Korup site, with its deep sandy soils of very low phosphorus status, and where most nutrient cycling takes place in a thin surface layer of fine roots and mycorrhizas, it appears that buttress form could develop from soil-surface root exploration for nutrients by juvenile trees. It may accordingly allow M. bisulcata to attain the higher greater competitive ability, faster growth rate, and maximum tree size that it does compared with other co-occurring tree species. For sites across the tropics in general, the degree of shallowness and spatial extension of buttresses of the dominant species is hypothesized to increase with decreasing nutrient availability.

Carbon stocks, tree diversity, and the role of organic certification in different cocoa production systems in Alto Beni, Bolivia

This study compares aboveground and belowground carbon stocks and tree diversity in different cocoa cultivation systems in Bolivia: monoculture, simple agroforestry, and successional agroforestry, as well as fallow as a control. Since diversified, agroforestry-based cultivation systems are often considered important for sustainable development, we also evaluated the links between carbon stocks and tree diversity, as well as the role of organic certification in transitioning from monoculture to agroforestry. Biomass, tree diversity, and soil physiochemical parameters were sampled in 15 plots measuring 48 × 48 m. Semi-structured interviews with 52 cocoa farmers were used to evaluate the role of organic certification and farmers’ organizations (e.g., cocoa cooperatives) in promoting tree diversity. Total carbon stocks in simple agroforestry systems (128.4 ± 20 Mg ha−1) were similar to those on fallow plots (125.2 ± 10 Mg ha−1). Successional agroforestry systems had the highest carbon stocks (143.7 ± 5.3 Mg ha−1). Monocultures stored significantly less carbon than all other systems (86.3 ± 4.0 Mg ha−1, posterior probability P(Diff > 0) of 0.000–0.006). Among shade tree species, Schizolobium amazonicum, Centrolobium ochroxylum, and Anadenanthera sp. accumulated the most biomass. High-value timber species (S. amazonicum, C. ochroxylum, Amburana cearensis, and Swietenia macrophylla) accounted for 22.0 % of shade tree biomass. The Shannon index and tree species richness were highest in successional agroforestry systems. Cocoa plots on certified organic farms displayed significantly higher tree species richness than plots on non-certified farms. Thus, expanding the coverage of organic farmers’ organizations may be an effective strategy for fostering transitions from monoculture to agroforestry systems.

Seedling resistance, tolerance and escape from herbivores: insights from co-dominant canopy tree species in a resource-poor African rain forest

* Although plants can reduce the impacts of herbivory in multiple ways, these defensive traits are often studied in isolation and an understanding of the resulting strategies is incomplete. * In the study reported here, empirical evidence was simultaneously evaluated for the three main sets of traits available to plants: (i) resistance through constitutive leaf traits, (ii) tolerance to defoliation and (iii) escape in space, for three caesalpiniaceous tree species Microberlinia bisulcata, Tetraberlinia bifoliolata and T. korupensis, which co-dominate groves within the lowland primary rain forest of Korup National Park (Cameroon). * Mesh cages were placed around individual wild seedlings to exclude insect herbivores at 41 paired canopy gap and understorey locations. After following seedling growth and survival for c. 2 years, caged and control treatments were removed, leaves harvested to determine nutrient and phenolic concentrations, leaf mass per area estimated, and seedling performance in gaps followed for a further c. 2 years to quantify tolerance to the leaf harvesting. * The more nutrient-rich leaves of the weakly shade-tolerant M. bisulcata were damaged much more in gaps than the two strongly shade-tolerant Tetraberlinia species, which had higher leaf mass per area and concentrations of total phenols. Conversely, the faster-growing M. bisulcata was better able to tolerate defoliation in terms of height growth (reflushing capacity), but not at maintaining overall leaf numbers, than the other two species. * Across gaps, insect-mediated Janzen–Connell effects were most pronounced for M. bisulcata, less so for T. korupensis, and not detectable for T. bifoliolata. The three species differed distinctly in their secondary metabolic profiles. * Taken together, the results suggested a conceptual framework linking the three sets of traits, one in which the three co-dominant species adopt different strategies towards herbivore pressure depending on their different responses to light availability. This study is one of the first in a natural forest ecosystem to examine resistance to, tolerance of, and escape from herbivory among a group of co-occurring tropical tree species.

Limitation of seedling growth by potassium and magnesium supply for two ectomycorrhizal tree species of a Central African rain forest and its implication for their recruitment

In the ectomycorrhizal caesalpiniaceous groves of southern Korup National Park, the dominant tree species, Microberlinia bisulcata, displays very poor in situ recruitment compared with its codominant, Tetraberlinia bifoliolata. The reported ex situ experiment tested whether availabilities of soil potassium and magnesium play a role. Seedlings of the two species received applications of K and Mg fertilizer in potted native soil in a local shade house, and their responses in terms of growth and nutrient concentrations were recorded over 2 years. Amended soil concentrations were also determined. Microberlinia responded strongly and positively in its growth to Mg, but less to K; Tetraberlinia responded weakly to both. Added Mg led to strongly increased Mg concentration for Microberlinia while added K changed that concentration only slightly; Tetraberlinia strongly increased its concentration of K with added K, but only somewhat its Mg concentration with added Mg. Additions of Mg and K had small but important antagonistic effects. Microberlinia is Mg-demanding and apparently Mg-limited in Korup soil; Tetraberlinia, whilst K-demanding, appeared not to be K-limited (for growth). Added K enhanced plant P concentrations of both species. Extra applied Mg may also be alleviating soil aluminum toxicity, and hence improving growth indirectly and especially to the benefit of Microberlinia. Mg appears to be essential for Microberlinia seedling growth and its low soil availability in grove soils at Korup may be an important contributing factor to its poor recruitment. Microberlinia is highly shade-intolerant and strongly light-responding, whilst Tetraberlinia is more shade-tolerant and moderately light-responding, which affords an interesting contrast with respect to their differing responses to Mg supply. The study revealed novel aspects of functional traits and likely niche-partitioning among ectomycorrhizal caesalps in African rain forests. Identifying the direct and interacting indirect effects of essential elements on tropical tree seedling growth presents a considerable challenge due the complex nexus of causes involved.

Canopy gaps promote selective stem-cutting by small mammals of two dominant tree species in an African lowland forest: the importance of seedling chemistry

Small mammals can impede tree regeneration by injuring seedlings and saplings in several ways. One fatal way is by severing their stems, but apparently this type of predation is not well-studied in tropical rain forest. Here, we report on the incidence of 'stem-cutting' to new, wild seedlings of two locally dominant, canopy tree species monitored in 40 paired forest understorey and gap-habitat areas in Korup, Cameroon following a 2007 masting event. In gap areas, which are required for the upward growth and sapling recruitment of both species, 137 seedlings of the long-lived, light-demanding, fast-growing large tropical tree (Microberlinia bisulcata) were highly susceptible to stem-cutting (83% of deaths) - it killed 39% of all seedlings over a c. 2-y period. In stark contrast, seedlings of the more shade-tolerant, slower-growing tree species (Tetraberlinia bifoliolata) were hardly attacked (4.3%). In the understorey, however, stem-cutting was virtually absent. Across the gap areas, the incidence of stem-cutting of M. bisulcata seedlings showed significant spatial variation that could not be explained significantly by either canopy openness or Janzen-Connell type effects (proximity and basal area of conspecific adult trees). To examine physical and chemical traits that might explain the species difference to being cut, bark and wood tissues were collected from a separate sample of seedlings in gaps (i.e. not monitored for stem-cutting). These analyses suggested that, compared with T. bifoliolata, the lower stem density, higher Mg and K and fatty acid concentrations in bark, and fewer phenolic and terpene compounds in M. bisulcata seedlings made them more palatable and attractive to small-mammal predators, likely rodents. We conclude that selective stem-cutting is a potent countervailing force to the current local canopy dominance of the grove-forming M. bisulcata by limiting the recruitment and abundance of its saplings. Given the ubiquity of gaps and ground-dwelling rodents in pantropical forests, it would be surprising if this form of lethal browsing was restricted to Korup.

Negative density dependence of seed dispersal and seedling recruitment in a Neotropical palm

Negative density dependence (NDD) of recruitment is pervasive in tropical tree species. We tested the hypotheses that seed dispersal is NDD, due to intraspecific competition for dispersers, and that this contributes to NDD of recruitment. We compared dispersal in the palm Attalea butyracea across a wide range of population density on Barro Colorado Island in Panama and assessed its consequences for seed distributions. We found that frugivore visitation, seed removal and dispersal distance all declined with population density of A. butyracea, demonstrating NDD of seed dispersal due to competition for dispersers. Furthermore, as population density increased, the distances of seeds from the nearest adult decreased, conspecific seed crowding increased and seedling recruitment success decreased, all patterns expected under poorer dispersal. Unexpectedly, however, our analyses showed that NDD of dispersal did not contribute substantially to these changes in the quality of the seed distribution; patterns with population density were dominated by effects due solely to increasing adult and seed density.

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 species and diversity effects on soil water seepage in a tropical plantation

Plant diversity has been shown to influence the water cycle of forest ecosystems by differences in water consumption and the associated effects on groundwater recharge. However, the effects of biodiversity on soil water fluxes remain poorly understood for native tree species plantations in the tropics. Therefore, we estimated soil water fluxes and assessed the effects of tree species and diversity on these fluxes in an experimental native tree species plantation in Sardinilla (Panama). The study was conducted during the wet season 2008 on plots of monocultures and mixtures of three or six tree species. Rainfall and soil water content were measured and evapotranspiration was estimated with the Penman-Monteith equation. Soil water fluxes were estimated using a simple soil water budget model considering water input, output, and soil water and groundwater storage changes and in addition, were simulated using the physically based one-dimensional water flow model Hydrus-1D. In general, the Hydrus simulation did not reflect the observed pressure heads, in that modeled pressure heads were higher compared to measured ones. On the other hand, the results of the water balance equation (WBE) reproduced observed water use patterns well. In monocultures, the downward fluxes through the 200 cm-depth plane were highest below Hura crepitans (6.13 mm day−1) and lowest below Luehea seemannii (5.18 mm day−1). The average seepage rate in monocultures (±SE) was 5.66 ± 0.18 mm day−1, and therefore, significantly higher than below six-species mixtures (5.49 ± 0.04 mm day−1) according to overyielding analyses. The three-species mixtures had an average seepage rate of 5.63 ± 0.12 mm day−1 and their values did not differ significantly from the average values of the corresponding species in monocultures. Seepage rates were driven by the transpiration of the varying biomass among the plots (r = 0.61, p = 0.017). Thus, a mixture of trees with different growth rates resulted in moderate seepage rates compared to monocultures of either fast growing or slow growing tree species. Our results demonstrate that tree-species specific biomass production and tree diversity are important controls of seepage rates in the Sardinilla plantation during the wet season.

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.