Above- and belowground responses of Arctic tundra ecosystems to altered soil nutrients and mammalian herbivory

Theory and observation indicate that changes in the rate of primary production can alter the balance between the bottom-up influences of plants and resources and the top-down regulation of herbivores and predators on ecosystem structure and function. The exploitation ecosystem hypothesis (EEH) posited that as aboveground net primary productivity (ANPP) increases, the additional biomass should support higher trophic levels. We developed an extension of EEH to include the impacts of increases in ANPP on belowground consumers in a similar manner as aboveground, but indirectly through changes in the allocation of photosynthate to roots. We tested our predictions for plants aboveground and for phytophagous nematodes and their predators belowground in two common arctic tundra plant communities subjected to 11 years of increased soil nutrient availability and/or exclusion of mammalian herbivores. The less productive dry heath (DH) community met the predictions of EEH aboveground, with the greatest ANPP and plant biomass in the fertilized plots protected from herbivory. A palatable grass increased in fertilized plots while dwarf evergreen shrubs and lichens declined. Belowground, phytophagous nematodes also responded as predicted, achieving greater biomass in the higher ANPP plots, whereas predator biomass tended to be lower in those same plots (although not significantly). In the higher productivity moist acidic tussock (MAT) community, aboveground responses were quite different. Herbivores stimulated ANPP and biomass in both ambient and enriched soil nutrient plots; maximum ANPP occurred in fertilized plots exposed to herbivory. Fertilized plots became dominated by dwarf birch (a deciduous shrub) and cloudberry (a perennial forb); under ambient conditions these two species coexist with sedges, evergreen dwarf shrubs, and Sphagnum mosses. Phytophagous nematodes did not respond significantly to changes in ANPP, although predator biomass was greatest in control plots. The contrasting results of these two arctic tundra plant communities suggest that the predictions of EEH may hold for very low ANPP communities, but that other factors, including competition and shifts in vegetation composition toward less palatable species, may confound predicted responses to changes in productivity in higher ANPP communities such as the MAT studied here.     

Modeling the role of macroalgae in a shallow sub-estuary of Narragansett Bay, RI (USA)

Proliferation of macroalgal mats is a frequent consequence of nutrient-driven eutrophication in shallow, photic coastal marine ecosystems. These macroalgae have the potential to significantly modify water quality, plankton productivity, nutrient cycling, and dissolved oxygen dynamics. We developed a model for Ulva lactuca and Gracilaria tikvahiae in Greenwich Bay, RI (USA), a shallow sub-estuary of Narragansett Bay, as part of a larger estuarine ecosystem model. The model predicts the biomass of both species in units of carbon, nitrogen, and phosphorus as a function of primary production, respiration, grazing, decay, and physical exchange, with particular attention to the effects of biomass layering on light attenuation and suppression of metabolic rates. The model successfully reproduced the magnitude and seasonal cycle of area-weighted and peak biomass in Greenwich Bay along with tissue C:N ratios, and highlighted the importance of grazing and inclusion of self-limitation primarily in the form of self-shading to overcome an order of magnitude difference in rates of production and respiration. Inclusion of luxury nutrient uptake demonstrated the importance of internal nutrient storage in fueling production when nutrients are limiting. Macroalgae were predicted to contribute a small fraction of total system primary production and their removal had little effect on predicted water quality. Despite a lack of data for calibration and a fair amount of sensitivity to individual parameter values, which highlights the need for further autecological studies to constrain formulations, the model successfully predicted macroalgal biomass dynamics and their role in ecosystem functioning. Our formulations should be exportable to other temperate systems where macroalgae occur in abundance.     

Evaluating weather effects on interannual variation in net ecosystem productivity of a coastal temperate forest landscape: A model intercomparison

Forest productivity is strongly affected by seasonal weather patterns and by natural or anthropogenic disturbances. However weather effects on forest productivity are not currently represented in inventory-based models such as CBM-CFS3 used in national forest C accounting programs. To evaluate different approaches to modelling these effects, a model intercomparison was conducted among CBM-CFS3 and four process models (ecosys, CN-CLASS, Can-IBIS and 3PG) over a 2500 ha landscape in the Oyster River (OR) area of British Columbia, Canada. The process models used local weather data to simulate net primary productivity (NPP), net ecosystem productivity (NEP) and net biome productivity (NBP) from 1920 to 2005. Other inputs used by the process and inventory models were generated from soil, land cover and disturbance records. During a period of intense disturbance from 1928 to 1943, simulated NBP diverged considerably among the models. This divergence was attributed to differences among models in the sizes of detrital and humus C stocks in different soil layers to which a uniform set of soil C transformation coefficients was applied during disturbances. After the disturbance period, divergence in modelled NBP among models was much smaller, and attributed mainly to differences in simulated NPP caused by different approaches to modelling weather effects on productivity. In spite of these differences, age-detrended variation in annual NPP and NEP of closed canopy forest stands was negatively correlated with mean daily maximum air temperature during July-September (T_amax) in all process models (R² = 0.4-0.6), indicating that these correlations were robust. The negative correlation between T_amax and NEP was attributed to different processes in different models, which were tested by comparing CO₂ fluxes from these models with those measured by eddy covariance (EC) under contrasting air temperatures (T_a). The general agreement in sensitivity of annual NPP to T_amax among the process models led to the development of a generalized algorithm for weather effects on NPP of coastal temperate coniferous forests for use in inventory-based models such as CBM-CFS3: NPP′ = NPP − 57.1 (T_amax − 18.6), where NPP and NPP′ are the current and temperature-adjusted annual NPP estimates from the inventory-based model, 18.6 is the long-term mean daily maximum air temperature during July-September, and T_amax is the mean value for the current year. Our analysis indicated that the sensitivity of NPP to T_amax was nonlinear, so that this algorithm should not be extrapolated beyond the conditions of this study. However the process-based methodology to estimate weather effects on NPP and NEP developed in this study is widely applicable to other forest types and may be adopted for other inventory based forest carbon cycle models.     

Effects of local fisheries and ocean productivity on the northeastern Ionian Sea ecosystem

To better understand the effects of fisheries and ocean productivity on the northeastern Ionian Sea we constructed an Ecopath with Ecosim model with 22 functional groups. Data on biomass, production/biomass, consumption/biomass, and diet for each group were estimated or extrapolated from the literature. Fisheries landings and discards were also included. Temporal trajectories were simulated using Ecosim. The model was fitted with time-series data for the most important groups from 1964 to 2006. Simulations highlighted a decline of top predators and of most of the commercial species since the late 1970s. The model shows that the decline of fish resources was mainly caused by an intensive fishing pressure that occurred in the area until the end of the 1990s and also by changes in primary production that impacted the trajectories of the main functional groups. In particular, simulated changes through time in PP impacted the abundance trends of all the commercial species, showing a cascade-up effect through the ecosystem. The application of Ecopath with Ecosim was a useful tool for understanding the trends of the main functional groups of the northeastern Ionian Sea. The model underlined that management actions are needed to restore and protect target species including marine mammals, pelagic and demersal fishes. In particular, measures to reduce overfishing, illegal fishing activities and to respect existing legislations are in need. Moreover, the adoption of marine protected areas could be an effective management measure to guarantee prey survival and to sustain marine predators.     

Inter-annual variability of ecosystem production in boreal jack pine forests (1975-2004) estimated from tree-ring data using CBM-CFS3

We describe and apply a method of using tree-ring data and an ecosystem model to reconstruct past annual rates of ecosystem production. Annual data on merchantable wood volume increment and mortality obtained by dendrochronological stand reconstruction were used as input to the Carbon Budget Model of the Canadian Forest Sector (CBM-CFS3) to estimate net ecosystem production (NEP), net primary production (NPP), and heterotrophic respiration (Rh) annually from 1975 to 2004 at 10 boreal jack pine (Pinus banksiana Lamb.) stands in Saskatchewan and Manitoba, Canada. From 1975 (when sites aged 41-60 years) to 2004 (when they aged 70-89 years), all sites were moderate C sinks except during some warmer than average years where estimated Rh increased. Across all sites and years, estimated annual NEP averaged 57 g Cm⁻² yr⁻¹ (range −31 to 176 g Cm⁻² yr⁻¹), NPP 244 g Cm⁻² yr⁻¹ (147-376 g Cm⁻² yr⁻¹), and Rh 187 g Cm⁻² yr⁻¹ (124-270 g Cm⁻² yr⁻¹). Across all sites, NPP was related to stand age and density, which are proxies for successional changes in leaf area. Regionally, warm spring temperature increased NPP and defoliation by jack pine budworm 1 year previously reduced NPP. Our estimates of NPP, Rh, and NEP were plausible when compared to regional eddy covariance and carbon stock measurements. Inter-annual variability in ecosystem productivity contributes uncertainty to inventory-based assessments of regional forest C budgets that use yield curves predicting averaged growth over time. Our method could expand the spatial and temporal coverage of annual forest productivity estimates, providing additional data for the development of empirical models accounting for factors not presently considered by these models.     

Woody encroachment decreases diversity across North American grasslands and savannas

Woody encroachment is a widespread and acute phenomenon affecting grasslands and savannas worldwide. We performed a meta-analysis of 29 studies from 13 different grassland/savanna communities in North America to determine the consequences of woody encroachment on plant species richness. In all 13 communities, species richness declined with woody plant encroachment (average decline = 45%). Species richness declined more in communities with higher precipitation (r2 = 0.81) and where encroachment was associated with a greater change in annual net primary productivity (ANPP; r2 = 0.69). Based on the strong positive correlation between precipitation and ANPP following encroachment (r2 = 0.87), we hypothesize that these relationships occur because water-limited woody plants experience a greater physiological and demographic release as precipitation increases. The observed relationship between species richness and ANPP provides support for the theoretical expectation that a trade-off occurs between richness and productivity in herbaceous communities. We conclude that woody plant encroachment leads to significant declines in species richness in North American grassland/savanna communities.     

The use of simulation modeling to evaluate the mechanisms responsible for the nutritional benefits of food-for-protection mutualisms

In some mutualisms, a plant or insect provides a food resource in exchange for protection from herbivores, competitors or predators. This food resource can benefit the consumer, but the relative importance of different mechanisms responsible for this benefit is unclear. We used a colony-level simulation model to test the relative importance of increased larval production, increased worker foraging and increased worker survival for colony growth of fire ants, Solenopsis invicta, that consume plant-based foods. Increased food for larvae had the largest effect on colony growth of S. invicta followed by decreased worker mortality. Increased foraging rate had a small effect in the simulation model but data from a small laboratory experiment and another published study suggest that plant-based foods have little or no effect on foraging rates of S. invicta. Colony growth steadily increased the longer plant-based food was available and colonies were most responsive to plant-based food in the early summer (i.e., June). Our results demonstrate that population level simulation modeling can be a useful tool for examining the ecology of mutualistic interactions and the mechanisms through which species interact.     

Water tables constrain height recovery of willow on Yellowstone's northern range.

Excessive levels of herbivory may disturb ecosystems in ways that persist even when herbivory is moderated. These persistent changes may complicate efforts to restore ecosystems affected by herbivores. Willow (Salix spp.) communities within the northern range in Yellowstone National Park have been eliminated or degraded in many riparian areas by excessive elk (Cervus elaphus L.) browsing. Elk browsing of riparian willows appears to have diminished following the reintroduction of wolves (Canis lupis L.), but it remains uncertain whether reduced herbivory will restore willow communities. The direct effects of elk browsing on willows have been accompanied by indirect effects from the loss of beaver (Castor canadensis Kuhl) activity, including incision of stream channels, erosion of fine sediments, and lower water tables near streams historically dammed by beaver. In areas where these changes have occurred, lowered water tables may suppress willow height even in the absence of elk browsing. We conducted a factorial field experiment to understand willow responses to browsing and to height of water tables. After four years of protection from elk browsing, willows with ambient water tables averaged only 106 cm in height, with negligible height gain in two of three study species during the last year of the experiment. Willows that were protected from browsing and had artificially elevated water tables averaged 147 cm in height and gained 19 cm in the last year of the experiment. In browsed plots, elevated water tables doubled height gain during a period of slightly reduced browsing pressure. We conclude that water availability mediates the rate of willow height gain and may determine whether willows grow tall enough to escape the browse zone of elk and gain resistance to future elk browsing. Consequently, in areas where long-term beaver absence has resulted in incised stream channels and low water tables, a reduction in elk browsing alone may not be sufficient for recovery of tall willow stands. Because tall willow stems are important elements of habitat for beaver, mitigating water table decline may be necessary in these areas to promote recovery of historical willow-beaver mutualisms.     

Predator diversity and identity drive interaction strength and trophic cascades in a food web

Declining predator diversity may drastically affect the biomass and productivity of herbivores and plants. Understanding how changes in predator diversity can propagate through food webs to alter ecosystem function is one of the most challenging ecological research topics today. We studied the effects of predator removal in a simple natural food web in the Sierra Nevada mountains of California (USA). By excluding the predators of the third trophic level of a food web in a full-factorial design, we monitored cascading effects of varying predator diversity and composition on the herbivorous beetle Chrysomela aeneicollis and the willow Salix orestera, which compose the first and second trophic levels of the food web. Decreasing predator diversity increased herbivore biomass and survivorship, and consequently increased the amount of plant biomass consumed via a trophic cascade. Despite this simple linear mean effect of diversity on the strength of the trophic cascade, we found additivity, compensation, and interference in the effects of multiple predators on herbivores and plants. Herbivore survivorship and predator-prey interaction strengths varied with predator diversity, predator identity, and the identity of coexisting predators. Additive effects of predators on herbivores and plants may have been driven by temporal niche separation, whereas compensatory effects and interference occurred among predators with a similar phenology. Together, these results suggest that while the general trends of diversity effects may appear linear and additive, other information about species identity was required to predict the effects of removing individual predators. In a community that is not temporally well-mixed, predator traits such as phenology may help predict impacts of species loss on other species. Information about predator natural history and food web structure may help explain variation in predator diversity effects on trophic cascades and ecosystem function.     

Assessing self-organization of plant communities—A thermodynamic approach

Thermodynamics is a powerful tool for the study of system development and has the potential to be applied to studies of ecological complexity. Here, we develop a set of thermodynamic indicators including energy capture and energy dissipation to quantify plant community self-organization. The study ecosystems included a tropical seasonal rainforest, an artificial tropical rainforest, a rubber plantation, and two Chromolaena odorata (L.) R.M. King & H. Robinson communities aged 13 years and 1 year. The communities represent a complexity transect from primary vegetation, to transitional community, economic plantation, and fallows and are typical for Xishuangbanna, southwestern China. The indicators of ecosystem self-organization are sensitive to plant community type and seasonality, and demonstrate that the tropical seasonal rainforest is highly self-organized and plays an important role in local environmental stability via the land surface thermal regulation. The rubber plantation is at a very low level of self-organization as quantified by the thermodynamic indicators, especially during the dry season. The expansion of the area of rubber plantation and shrinkage of tropical seasonal rainforest would likely induce local surface warming and a larger daily temperature range.     

Grassland establishment under varying resource availability: a test of positive and negative feedback

The traditional logic of carbon (C) and nitrogen (N) interactions in ecosystems predicts further increases or decreases in productivity (positive feedback) in response to high and low fertility in the soil, respectively; but the potential for development of feedback in ecosystems recovering from disturbance is less well understood. Furthermore, this logic has been challenged in grassland ecosystems where frequent fires or grazing may reduce the contribution of aboveground litter inputs to soil organic matter pools and nutrient supply for plant growth, relative to forest ecosystems. Further, if increases in plant productivity increase soil C content more than soil N content, negative feedback may result from increased microbial demand for N making less available for plant growth. We used a field experiment to test for feedback in an establishing grassland by comparing aboveground net primary productivity (ANPP) and belowground pools and fluxes of C and N in soil with enriched, ambient, and reduced N availability. For eight years annual N enrichment increased ANPP, root N, and root tissue quality, but root C:N ratios remained well above the threshold for net mineralization of N. There was no evidence that N enrichment increased root biomass, soil C or N accrual rates, or storage of C in total, microbial, or mineralizable pools within this time frame. However, the net nitrogen mineralization potential (NMP) rate was greater following eight years of N enrichment, and we attributed this to N saturation of the microbial biomass. Grassland developing under experimentally imposed N limitation through C addition to the soil exhibited ANPP, root biomass and quality, and net NMP rate similar to the ambient soil. Similarity in productivity and roots in the reduced and ambient N treatments was attributed to the potentially high nitrogen-use efficiency (NUE) of the dominant C4 grasses, and increasing cover of legumes over time in the C-amended soil. Thus, in a developing ecosystem, positive feedback between soil N supply and plant productivity may promote enhanced long-term N availability and override progressive N limitation as C accrues in plant and soil pools. However, experimentally imposed reduction in N availability did not feed back to reduce ANPP, possibly due to shifts in NUE and functional group composition.     

A parsimonious optimal foraging model explaining mortality patterns in Serengeti wildebeest

Based on data collected over 24 years in the Serengeti in Tanzania, Sinclair and Arcese (1995) indicated that the sensitivity of blue wildebeest Connochaetes taurinus to predation risk by lions Panthera leo may cause them to change habitats between open (low risk) and wooded (risky) habitats. They found that, in poor rainfall years, predators kill wildebeest that are in better condition than those that die of natural causes. In good rainfall years, predators kill wildebeest that are in worse condition than those that die of natural causes. Sinclair and Arcese (1995) proposed the “predation-sensitive food” hypothesis. This hypothesis suggests that, as food becomes limiting, animals take greater risks to obtain more food, and some of these animals are killed. I propose a more parsimonious hypothesis based on the marginal value theorem that is consistent with the observations made by Sinclair and Arcese (1995). Wildebeest follow a single decision rule in good and poor rainfall years, viz. move when foraging elsewhere increases your rate of intake of nutritious food. Similarly, predators follow a single decision rule in good and poor rainfall years, viz. take the prey item that maximizes the intake of energy per unit effort expended. This parsimonious model does not require differences in predator sensitivity as required by Sinclair and Arcese's (1995) model. I indicate ways in which my model can be falsified.     

Modeling seasonal course of carbon fluxes and evapotranspiration in response to low temperature and moisture in a boreal Scots pine ecosystem

Environmental conditions act above and below ground, and regulate carbon fluxes and evapotranspiration. The productivity of boreal forest ecosystems is strongly governed by low temperature and moisture conditions, but the understanding of various feedbacks between vegetation and environmental conditions is still unclear. In order to quantify the seasonal responses of vegetation to environmental factors, the seasonality of carbon and heat fluxes and the corresponding responses for temperature and moisture in air and soil were simulated by merging a process-based model (CoupModel) with detailed measurements representing various components of a forest ecosystem in Hyytiälä, southern Finland. The uncertainties in parameters, model assumptions, and measurements were identified by generalized likelihood uncertainty estimation (GLUE). Seasonal and diurnal courses of sensible and latent heat fluxes and net ecosystem exchange (NEE) of CO₂ were successfully simulated for two contrasting years. Moreover, systematic increases in efficiency of photosynthesis, water uptake, and decomposition occurred from spring to summer, demonstrating the strong coupling between processes. Evapotranspiration and NEE flux both showed a strong response to soil temperature conditions via different direct and indirect ecosystem mechanisms. The rate of photosynthesis was strongly correlated with the corresponding water uptake response and the light use efficiency. With the present data and model assumptions, it was not possible to precisely distinguish the various regulating ecosystem mechanisms. Our approach proved robust for modeling the seasonal course of carbon fluxes and evapotranspiration by combining different independent measurements. It will be highly interesting to continue using long-term series data and to make additional tests of optional stomatal conductance models in order to improve our understanding of the boreal forest ecosystem in response to climate variability and environmental conditions.     

Metabolism and model of an estuarine bay ecosystem affected by a coastal power plant

The effects of a coastal power plant on an outer estuarine bay ecosystem on the west coast of Florida were evaluated with measurements and an ecological model. Field measurements of community metabolism and biomass were taken from the thermally affected bay and from similar control bays. Model simulations were used to help understand these observations in terms of ecosystem structure and functioning.In the outer discharge bay the direct impact of the thermal plume was diluted and spread overlarge areas. The ecosystem developed structure and functions with lower biomass than in the control bays but with slightly faster rates of organic turnover. The productive turnover time of producer biomass during the summer was about 5 days in the discharge bay and about 6 days in the control bays. Power plant influence on total community metabolism was small with less than 10% difference in annual averages between the discharge and control bays (5.22 and 5.58 g O₂/m²/day). The selection for faster metabolic turnover rates in the discharge by was evidenced by a dominance of plankton metabolism over benthic metabolism. The annual average gross planktonic production was around 3 g O₂/m²/day in the discharge bay and around 2 g/m²/day in the control bays.In the model, temperature served as a stimulant to both productivity and respiration. When the isolated effects of increased temperature were simulated the model responded with lower producer biomass and faster rates of organic turnover, as was found in field measurements. These simulations also showed increased nutrient recycling and indicated patterns of temperature-induced migrations. Since power plant operation affected water exchange in the bays, several levels of total water exchange were simulated. These simulations indicated the importance of water exchange as a stabilizing factor, especially for sensitive compartments with rapid turnover rates (i.e. plankton and phosphorus stocks). Simulations of the effects of future power plant units on the bay ecosystem showed no large changes in total metabolism but indicated larger effects of plankton entrainment mortality and temperature-induced migrations of larger organisms.     

Spatially distributed modeling of the long-term carbon balance of a boreal landscape

Spatially and temporally distributed information on the sizes of biomass carbon (C) pools (BCPs) and soil C pools (SCPs) is vital for improving our understanding of biosphere-atmosphere C fluxes. Because the sizes of C pools result from the integrated effects of primary production, age-effects, changes in climate, atmospheric CO₂ concentration, N deposition, and disturbances, a modeling scheme that interactively considers these processes is important. We used the InTEC model, driven by various spatio-temporal datasets to simulate the long-term C-balance in a boreal landscape in eastern Canada. Our results suggested that in this boreal landscape, mature coniferous stands had stabilized their productivity and fluctuated as a weak C-sink or C-source depending on the interannual variations in hydrometeorological factors. Disturbed deciduous stands were larger C-sinks (NEP₂₀₀₄ = 150 gC m⁻² yr⁻¹) than undisturbed coniferous stands (e.g. NEP₂₀₀₄ = 8 gC m⁻² yr⁻¹). Wetlands had lower NPP but showed temporally consistent C accumulation patterns. The simulated spatio-temporal patterns of BCPs and SCPs were unique and reflected the integrated effects of climate, plant growth and atmospheric chemistry besides the inherent properties of the C pool themselves. The simulated BCPs and SCPs generally compared well with the biometric estimates (BCPs: r = 0.86, SCPs: r = 0.84). The largest BCP biases were found in recently disturbed stands and the largest SCP biases were seen in locations where moss necro-masses were abundant. Reconstructing C pools and C fluxes in the ecosystem in such a spatio-temporal manner could help reduce the uncertainties in our understanding of terrestrial C-cycle.     

Interpreting spatial heterogeneity of crop yield with a process model and remote sensing

A process-based crop growth model (Vegetation Interface Processes (VIP) model) is used to estimate crop yield with remote sensing over the North China Plain. Spatial pattern of the key parameter—maximum catalytic capacity of Rubisco (V_cmax) for assimilation is retrieved from Normalized Difference of Vegetation Index (NDVI) from Terra-MODIS and statistical yield records. The regional simulation shows that the agreements between the simulated winter wheat yields and census data at county-level are quite well with R² being 0.41-0.50 during 2001-2005. Spatial variability of photosynthetic capacity and yield in irrigated regions depend greatly on nitrogen input. Due to the heavy soil salinity, the photosynthetic capacity and yield in coastal region is less than 50 μmol C m⁻² s⁻¹ and 3000 kg ha⁻¹, respectively, which are much lower than that in non-salinized region, 84.5 μmol C m⁻² s⁻¹ and 5700 kg ha⁻¹. The predicted yield for irrigated wheat ranges from 4000 to 7800 kg ha⁻¹, which is significantly larger than that of rainfed, 1500-3000 kg ha⁻¹. According to the path coefficient analysis, nitrogen significantly affects yield, by which water exerts noticeably indirect influences on yield. The effect of water on yield is regulated, to a certain extent, by crop photosynthetic capacity and nitrogen application. It is believed that photosynthetic parameters retrieved from remote sensing are reliable for regional production prediction with a process-based model.     

Relationships between simulated water stress and mortality and growth rates in underplanted Toona ciliata Roem. in subtropical Argentinean plantations

Toona ciliata Roem. (Australian red cedar) requires a nurse-tree overstory to prevent damage from drought and irradiation in some regions of north-eastern Argentina. T. ciliata was planted in the understory of Pinus taeda L. (625 stems/ha), Pinus elliottii Engelm. × Pinus caribaea Morelet (625 stems/ha), and Grevillea robusta A. Cunn. (833 stems/ha) nurse trees, which were thinned to 0, 25, 50, 75 and 100% of the initial densities. We measured initial T. ciliata mortality and growth as well as Leaf Area Index (LAI) based on light transmission. T. ciliata soil water availability and its effect on early growth and mortality were examined by modelling drought stress using the two-dimensional forest hydrology model ForWaDy. Simulated patterns in T. ciliata water stress for the different overstory treatments were consistent with observed patterns of mortality. Early mortality was lowest with a G. robusta overstory, with corresponding lowest drought stress values and high modelled soil water contents in the top soil layer in intermediate and high overstory densities. Mortality was highest with a P. elliottii × P. caribaea overstory in treatments with the highest modelled drought stress values in the most open treatments. The model supported our field observations by indicating that water stress was an important limitation to T. ciliata survival and growth on our study sites. The linkage between T. ciliata establishment success, early growth and soil water availability as indicated by ForWaDy, leads us to conclude that the model is a suitable stand management tool for guiding establishment of T. ciliata plantations.     

Analysis of annual fluctuations of C. nodosa in the Venice lagoon: Modeling approach

A simple simulation model was developed to describe the growth trends of Cymodocea nodosa (Ucria) Ascherson based on data sets from the Venice lagoon. The model reproduces the seasonal fluctuations in the above and belowground biomass and in shoot density. The modeling results are in good agreement with data on net production, growth rates and chemical–physical parameters of water. It was assumed that light and temperature are the most important factors controlling C. nodosa development, and that the growth was not limited by nutrient availability. The aim was to simulate biomass production as a function of external forcing variables (light, water temperature) and internal control (plant density). A series of simulation experiments were performed with the basic model showing that among the most important phenomena affecting C. nodosa growth are: (1) inhibition of production and recruitment of new shoots by high temperature and (2) light attenuation due to seasonal fluctuation.     

Estimating California ecosystem carbon change using process model and land cover disturbance data: 1951-2000

Land use change, natural disturbance, and climate change directly alter ecosystem productivity and carbon stock level. The estimation of ecosystem carbon dynamics depends on the quality of land cover change data and the effectiveness of the ecosystem models that represent the vegetation growth processes and disturbance effects. We used the Integrated Biosphere Simulator (IBIS) and a set of 30- to 60-m resolution fire and land cover change data to examine the carbon changes of California's forests, shrublands, and grasslands. Simulation results indicate that during 1951-2000, the net primary productivity (NPP) increased by 7%, from 72.2 to 77.1 Tg C yr⁻¹ (1 teragram = 10¹² g), mainly due to CO₂ fertilization, since the climate hardly changed during this period. Similarly, heterotrophic respiration increased by 5%, from 69.4 to 73.1 Tg C yr⁻¹, mainly due to increased forest soil carbon and temperature. Net ecosystem production (NEP) was highly variable in the 50-year period but on average equalled 3.0 Tg C yr⁻¹ (total of 149 Tg C). As with NEP, the net biome production (NBP) was also highly variable but averaged −0.55 Tg C yr⁻¹ (total of -27.3 Tg C) because NBP in the 1980s was very low (-5.34 Tg C yr⁻¹). During the study period, a total of 126 Tg carbon were removed by logging and land use change, and 50 Tg carbon were directly removed by wildland fires. For carbon pools, the estimated total living upper canopy (tree) biomass decreased from 928 to 834 Tg C, and the understory (including shrub and grass) biomass increased from 59 to 63 Tg C. Soil carbon and dead biomass carbon increased from 1136 to 1197 Tg C.Our analyses suggest that both natural and human processes have significant influence on the carbon change in California. During 1951-2000, climate interannual variability was the key driving force for the large interannual changes of ecosystem carbon source and sink at the state level, while logging and fire were the dominant driving forces for carbon balances in several specific ecoregions. From a long-term perspective, CO₂ fertilization plays a key role in maintaining higher NPP. However, our study shows that the increase in C sequestration by CO₂ fertilization is largely offset by logging/land use change and wildland fires.     

Modeling amphibian energetics, habitat suitability, and movements of western toads, Anaxyrus (=Bufo) boreas, across present and future landscapes

Effective conservation of amphibian populations requires the prediction of how amphibians use and move through a landscape. Amphibians are closely coupled to their physical environment. Thus an approach that uses the physiological attributes of amphibians, together with knowledge of their natural history, should be helpful. We used Niche Mapper™ to model the known movements and habitat use patterns of a population of Western toads (Anaxyrus (=Bufo) boreas) occupying forested habitats in southeastern Idaho. Niche Mapper uses first principles of environmental biophysics to combine features of topography, climate, land cover, and animal features to model microclimates and animal physiology and behavior across landscapes. Niche Mapper reproduced core body temperatures (T_c) and evaporation rates of live toads with average errors of 1.6 ± 0.4 °C and 0.8 ± 0.2 g/h, respectively. For four different habitat types, it reproduced similar mid-summer daily temperature patterns as those measured in the field and calculated evaporation rates (g/h) with an average error rate of 7.2 ± 5.5%. Sensitivity analyses indicate these errors do not significantly affect estimates of food consumption or activity. Using Niche Mapper we predicted the daily habitats used by free-ranging toads; our accuracy for female toads was greater than for male toads (74.2 ± 6.8% and 53.6 ± 15.8%, respectively), reflecting the stronger patterns of habitat selection among females. Using these changing to construct a cost surface, we also reconstructed movement paths that were consistent with field observations. The effect of climate warming on toads depends on the interaction of temperature and atmospheric moisture. If climate change occurs as predicted, results from Niche Mapper suggests that climate warming will increase the physiological cost of landscapes thereby limiting the activity for toads in different habitats.