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.

Untangling a Holocene pollen record with forest model simulations and independent climate data

Adaptation potential of forests to rapid climatic changes can be assessed from vegetation dynamics during past climatic changes as preserved in fossil pollen data. However, pollen data reflect the integrated effects of climate and biotic processes, such as establishment, survival, competition, and migration. To disentangle these processes, we compared an annually laminated late Würm and Holocene pollen record from the Central Swiss Plateau with simulations of a dynamic forest patch model. All input data used in the simulations were largely independent from pollen data; i.e. the presented analysis is non-circular. Temperature and precipitation scenarios were based on reconstructions from pollen-independent sources. The earliest arrival times of the species at the study site after the last glacial were inferred from pollen maps. We ran a series of simulations under different combinations of climate and immigration scenarios. In addition, the sensitivity of the simulated presence/absence of four major species to changes in the climate scenario was examined. The pattern of the pollen record could partly be explained by the used climate scenario, mostly by temperature. However, some features, in particular the absence of most species during the late Würm could only be simulated if the winter temperature anomalies of the used scenario were decreased considerably. Consequently, we had to assume in the simulations, that most species immigrated during or after the Younger Dryas (12 000 years BP), Abies and Fagus even later. Given the timing of tree species immigration, the vegetation was in equilibrium with climate during long periods, but responded with lags at the time-scale of centuries to millennia caused by a secondary succession after rapid climatic changes such as at the end of Younger Dryas, or immigration of dominant taxa. Climate influenced the tree taxa both directly and indirectly by changing inter-specific competition. We concluded, that also during the present fast climatic change, species migration might be an important process, particularly if geographic barriers, such as the Alps are in the migrational path.

Effects of climate, fire, and humans on forest dynamics: forest simulations compared to the palaeological record

In order to find out which factors influenced the forest dynamics in northern Italy during the Holocene, a palaeoecological approach involving pollen analysis was combined with ecosystem modelling. The dynamic and distribution based forest model DisCForm was run with different input scenarios for climate, species immigration, fire, and human impact and the similarity of the simulations with the original pollen record was assessed. From the comparisons of the model output and the pollen core, it appears that immigration was most important in the first part of the Holocene, and that fire and human activity had a major influence in the second half. Species not well represented in the simulation outputs are species with a higher abundance in the past than today (Corylus), with their habitat in riparian forests (Alnus) or with a strong response to human impact (Castanea).

Tracing of 15N in a mountain spruce forest: comparison of experimental data with th TRACE model

Despite numerous studies about nitrogen-cycling in forest ecosystems, many uncertainties remain, especially regarding the longer-term nitrogen accumulation. To contribute to filling this gap, the dynamic process-based model TRACE, with the ability to simulate 15N tracer redistribution in forest ecosystems was used to study N cycling processes in a mountain spruce forest of the northern edge of the Alps in Switzerland (Alptal, SZ). Most modeling analyses of N-cycling and C-N interactions have very limited ability to determine whether the process interactions are captured correctly. Because the interactions in such a system are complex, it is possible to get the whole-system C and N cycling right in a model without really knowing if the way the model combines fine-scale interactions to derive whole-system cycling is correct. With the possibility to simulate 15N tracer redistribution in ecosystem compartments, TRACE features a very powerful tool for the validation of fine-scale processes captured by the model. We first adapted the model to the new site (Alptal, Switzerland; long-term low-dose N-amendment experiment) by including a new algorithm for preferential water flow and by parameterizing of differences in drivers such as climate, N deposition and initial site conditions. After the calibration of key rates such as NPP and SOM turnover, we simulated patterns of 15N redistribution to compare against 15N field observations from a large-scale labeling experiment. The comparison of 15N field data with the modeled redistribution of the tracer in the soil horizons and vegetation compartments shows that the majority of fine-scale processes are captured satisfactorily. Particularly, the model is able to reproduce the fact that the largest part of the N deposition is immobilized in the soil. The discrepancies of 15N recovery in the LF and M soil horizon can be explained by the application method of the tracer and by the retention of the applied tracer by the well developed moss layer, which is not considered in the model. Discrepancies in the dynamics of foliage and litterfall 15N recovery were also observed and are related to the longevity of the needles in our mountain forest. As a next step, we will use the final Alptal version of the model to calculate the effects of climate change (temperature, CO2) and N deposition on ecosystem C sequestration in this regionally representative Norway spruce (Picea abies) stand.

A model-data comparison of Holocene timberline changes in the Swiss Alps reveals past and future drivers of mountain forest dynamics

Mountain vegetation is strongly affected by temperature and is expected to shift upwards with climate change. Dynamic vegetation models are often used to assess the impact of climate on vegetation and model output can be compared with paleobotanical data as a reality check. Recent paleoecological studies have revealed regional variation in the upward shift of timberlines in the Northern and Central European Alps in response to rapid warming at the Younger Dryas/Preboreal transition ca. 11700years ago, probably caused by a climatic gradient across the Alps. This contrasts with previous studies that successfully simulated the early Holocene afforestation in the (warmer) Central Alps with a chironomid-inferred temperature reconstruction from the (colder) Northern Alps. We use LandClim, a dynamic landscape vegetation model to simulate mountain forests under different temperature, soil and precipitation scenarios around Iffigsee (2065m a.s.l.) a lake in the Northwestern Swiss Alps, and compare the model output with the paleobotanical records. The model clearly overestimates the upward shift of timberline in a climate scenario that applies chironomid-inferred July-temperature anomalies to all months. However, forest establishment at 9800 cal. BP at Iffigsee is successfully simulated with lower moisture availability and monthly temperatures corrected for stronger seasonality during the early Holocene. The model-data comparison reveals a contraction in the realized niche of Abies alba due to the prominent role of anthropogenic disturbance after ca. 5000 cal. BP, which has important implications for species distribution models (SDMs) that rely on equilibrium with climate and niche stability. Under future climate projections, LandClim indicates a rapid upward shift of mountain vegetation belts by ca. 500m and treeline positions of ca. 2500m a.s.l. by the end of this century. Resulting biodiversity losses in the alpine vegetation belt might be mitigated with low-impact pastoralism to preserve species-rich alpine meadows.

Deforestation and forest degradation monitoring and assessment of biomass and carbon stock of lowland rainforest in the Analanjirofo region, Madagascar

Madagascar is currently developing a policy and strategies to enhance the sustainable management of its natural resources, encouraged by United Nations Framework Convention on Climate Change (UNFCCC) and REDD. To set up a sustainable financing scheme methodologies have to be provided that estimate, prevent and mitigate leakage, develop national and regional baselines, and estimate carbon benefits. With this research study this challenge was tried to be addressed by analysing a lowland rainforest in the Analanjirofo region in the district of Soanierana Ivongo, North East of Madagascar. For two distinguished forest degradation stages: “low degraded forest” and “degraded forest” aboveground biomass and carbon stock was assessed. The corresponding rates of carbon within those two classes were calculated and linked to a multi-temporal set of SPOT satellite data acquired in 1991, 2004 and 2009. Deforestation and particularly degradation and the related carbon stock developments were analysed. With the assessed data for the 3 years 1991, 2004 and 2009 it was possible to model a baseline and to develop a forest prediction for 2020 for Analanjirofo region in the district of Soanierana Ivongo. These results, developed applying robust methods, may provide important spatial information regarding the priorities in planning and implementation of future REDD+ activities in the area.

Transient dominance in a central African rain forest

The large-crowned emergent tree Microberlinia bisulcata dominates rain forest groves at Korup National Park, Cameroon, along with two codominants, Tetraberlinia bifoliolata and T. korupensis. M. bisulcata has a pronounced modal size frequency distribution around 110 cm stem diameter: its recruitment potential is very poor. It is a long-lived light-demanding species, one of many found in African forests. Tetraberlinia species lack modality, are more shade tolerant, and recruit better. All three species are ectomycorrhizal. M. bisulcata dominates grove basal area, even though it has similar numbers of trees (≥50 cm stem diameter) as each of the other two species. This situation presented a conundrum that prompted a long-term study of grove dynamics. Enumerations of two plots (82.5 and 56.25 ha) between 1990 and 2010 showed mortality and recruitment of M. bisulcata to be very low (both rates 0.2% per year) compared with Tetraberlinia (2.4% and 0.8% per year), and M. bisulcata grows twice as fast as the Tetraberlinia. Ordinations indicated that these three species determined community structure by their strong negative associations while other species showed almost none. Ranked species abundance curves fitted the Zipf-Mandelbrot model well and allowed “overdominance” of M. bisulcata to be estimated. Spatial analysis indicated strong repulsion by clusters of large (50 to <100 cm) and very large (≥100 cm) M. bisulcata of their own medium-sized (10 to <50 cm) trees and all sizes of Tetraberlinia. This was interpreted as competition by M. bisulcata increasing its dominance, but also inhibition of its own replacement potential. Stem coring showed a modal age of 200 years for M. bisulcata, but with large size variation (50–150 cm). Fifty-year model projections suggested little change in medium, decreases in large, and increases in very large trees of M. bisulcata, accompanied by overall decreases in medium and large trees of Tetraberlinia species. Realistically increasing very-large-tree mortality led to grove collapse without short-term replacement. M. bisulcata most likely depends on climatic events to rebuild its stands: the ratio of disturbance interval to median species' longevity is important. A new theory of transient dominance explains how M. bisulcata may be cycling in abundance over time and displaying nonequilibrium dynamics.

The importance of glacier and forest change in hydrological climate-impact studies

Changes in land cover alter the water balance components of a catchment, due to strong interactions between soils, vegetation and the atmosphere. Therefore, hydrological climate impact studies should also integrate scenarios of associated land cover change. To reflect two severe climate-induced changes in land cover, we applied scenarios of glacier retreat and forest cover increase that were derived from the temperature signals of the climate scenarios used in this study. The climate scenarios were derived from ten regional climate models from the ENSEMBLES project. Their respective temperature and precipitation changes between the scenario period (2074–2095) and the control period (1984–2005) were used to run a hydrological model. The relative importance of each of the three types of scenarios (climate, glacier, forest) was assessed through an analysis of variance (ANOVA). Altogether, 15 mountainous catchments in Switzerland were analysed, exhibiting different degrees of glaciation during the control period (0–51%) and different degrees of forest cover increase under scenarios of change (12–55% of the catchment area). The results show that even an extreme change in forest cover is negligible with respect to changes in runoff, but it is crucial as soon as changes in evaporation or soil moisture are concerned. For the latter two variables, the relative impact of forest change is proportional to the magnitude of its change. For changes that concern 35% of the catchment area or more, the effect of forest change on summer evapotranspiration is equally or even more important than the climate signal. For catchments with a glaciation of 10% or more in the control period, the glacier retreat significantly determines summer and annual runoff. The most important source of uncertainty in this study, though, is the climate scenario and it is highly recommended to apply an ensemble of climate scenarios in the impact studies. The results presented here are valid for the climatic region they were tested for, i.e., a humid, mid-latitude mountainous environment. They might be different for regions where the evaporation is a major component of the water balance, for example. Nevertheless, a hydrological climate-impact study that assesses the additional impacts of forest and glacier change is new so far and provides insight into the question whether or not it is necessary to account for land cover changes as part of climate change impacts on hydrological systems.

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.

Robust Proximal Femur Segmentation in Conventional X-Ray Images via Random Forest Regression on Multi-resolution Gradient Features

In this paper, we propose a fully automatic, robust approach for segmenting proximal femur in conventional X-ray images. Our method is based on hierarchical landmark detection by random forest regression, where the detection results of 22 global landmarks are used to do the spatial normalization, and the detection results of the 59 local landmarks serve as the image cue for instantiation of a statistical shape model of the proximal femur. To detect landmarks in both levels, we use multi-resolution HoG (Histogram of Oriented Gradients) as features which can achieve better accuracy and robustness. The efficacy of the present method is demonstrated by experiments conducted on 150 clinical x-ray images. It was found that the present method could achieve an average point-to-curve error of 2.0 mm and that the present method was robust to low image contrast, noise and occlusions caused by implants.

Historical and idealized climate model experiments: an intercomparison of Earth system models of intermediate complexity

Both historical and idealized climate model experiments are performed with a variety of Earth system models of intermediate complexity (EMICs) as part of a community contribution to the Intergovernmental Panel on Climate Change Fifth Assessment Report. Historical simulations start at 850 CE and continue through to 2005. The standard simulations include changes in forcing from solar luminosity, Earth's orbital configuration, CO2, additional greenhouse gases, land use, and sulphate and volcanic aerosols. In spite of very different modelled pre-industrial global surface air temperatures, overall 20th century trends in surface air temperature and carbon uptake are reasonably well simulated when compared to observed trends. Land carbon fluxes show much more variation between models than ocean carbon fluxes, and recent land fluxes appear to be slightly underestimated. It is possible that recent modelled climate trends or climate–carbon feedbacks are overestimated resulting in too much land carbon loss or that carbon uptake due to CO2 and/or nitrogen fertilization is underestimated. Several one thousand year long, idealized, 2 × and 4 × CO2 experiments are used to quantify standard model characteristics, including transient and equilibrium climate sensitivities, and climate–carbon feedbacks. The values from EMICs generally fall within the range given by general circulation models. Seven additional historical simulations, each including a single specified forcing, are used to assess the contributions of different climate forcings to the overall climate and carbon cycle response. The response of surface air temperature is the linear sum of the individual forcings, while the carbon cycle response shows a non-linear interaction between land-use change and CO2 forcings for some models. Finally, the preindustrial portions of the last millennium simulations are used to assess historical model carbon-climate feedbacks. Given the specified forcing, there is a tendency for the EMICs to underestimate the drop in surface air temperature and CO2 between the Medieval Climate Anomaly and the Little Ice Age estimated from palaeoclimate reconstructions. This in turn could be a result of unforced variability within the climate system, uncertainty in the reconstructions of temperature and CO2, errors in the reconstructions of forcing used to drive the models, or the incomplete representation of certain processes within the models. Given the forcing datasets used in this study, the models calculate significant land-use emissions over the pre-industrial period. This implies that land-use emissions might need to be taken into account, when making estimates of climate–carbon feedbacks from palaeoclimate reconstructions.

Recent trends in Inner Asian forest dynamics to temperature and precipitation indicate high sensitivity to climate change

Semi-arid ecosystems play an important role in regulating global climate with the fate of these ecosystems in the Anthropocene depending upon interactions among temperature, precipitation, and CO2. However, in cool-arid environments, precipitation is not the only limitation to forest productivity. Interactions between changes in precipitation and air temperature may enhance soil moisture stress while simultaneously extending growing season length, with unclear consequences for net carbon uptake. This study evaluates recent trends in productivity and phenology of Inner Asian forests (in Mongolia and Northern China) using satellite remote sensing, dendrochronology, and dynamic global vegetation model (DGVM) simulations to quantify the sensitivity of forest dynamics to decadal climate variability and trends. Trends in photosynthetically active radiation fraction (FPAR) between 1982 and 2010 show a greening of about 7% of the region in spring (March, April, May), and 3% of the area ‘browning’ during summertime (June, July, August). These satellite observations of FPAR are corroborated by trends in NPP simulated by the LPJ DGVM. Spring greening trends in FPAR are mainly explained by long-term trends in precipitation whereas summer browning trends are correlated with decreasing precipitation. Tree ring data from 25 sites confirm annual growth increments are mainly limited by summer precipitation (June, July, August) in Mongolia, and spring precipitation in northern China (March, April, May), with relatively weak prior-year lag effects. An ensemble of climate projections from the IPCC CMIP3 models indicates that warming temperatures (spring, summer) are expected to be associated with higher summer precipitation, which combined with CO2 causes large increases in NPP and possibly even greater forest cover in the Mongolian steppe. In the absence of a strong direct CO2 fertilization effect on plant growth (e.g., due to nutrient limitation), water stress or decreased carbon gain from higher autotrophic respiration results in decreased productivity and loss of forest cover. The fate of these semi-arid ecosystems thus appears to hinge upon the magnitude and subtleties of CO2 fertilization effects, for which experimental observations in arid systems are needed to test and refine vegetation models.

Site- and species-specific responses of forest growth to climate across the European continent

Aim To evaluate the climate sensitivity of model-based forest productivity estimates using a continental-scale tree-ring network. Location Europe and North Africa (30–70° N, 10° W–40° E). Methods We compiled close to 1000 annually resolved records of radial tree growth for all major European tree species and quantified changes in growth as a function of historical climatic variation. Sites were grouped using a neural network clustering technique to isolate spatiotemporal and species-specific climate response patterns. The resulting empirical climate sensitivities were compared with the sensitivities of net primary production (NPP) estimates derived from the ORCHIDEE-FM and LPJ-wsl dynamic global vegetation models (DGVMs). Results We found coherent biogeographic patterns in climate response that depend upon (1) phylogenetic controls and (2) ambient environmental conditions delineated by latitudinal/elevational location. Temperature controls dominate forest productivity in high-elevation and high-latitude areas whereas moisture sensitive sites are widespread at low elevation in central and southern Europe. DGVM simulations broadly reproduce the empirical patterns, but show less temperature sensitivity in the boreal zone and stronger precipitation sensitivity towards the mid-latitudes. Main conclusions Large-scale forest productivity is driven by monthly to seasonal climate controls, but our results emphasize species-specific growth patterns under comparable environmental conditions. Furthermore, we demonstrate that carry-over effects from the previous growing season can significantly influence tree growth, particularly in areas with harsh climatic conditions – an element not considered in most current-state DGVMs. Model–data discrepancies suggest that the simulated climate sensitivity of NPP will need refinement before carbon-cycle climate feedbacks can be accurately quantified.