Influences of canopy structure and physiological traits on flux partitioning between understory and overstory in an eastern Siberian boreal larch forest

Boreal forests play an important role in the global balance of energy and CO₂. Our previous study of elaborate eddy covariance observations in a Siberian boreal larch forest, conducted both above the forest canopy and at the forest floor, revealed a significant contribution of latent heat flux (LE) from the cowberry understory to the whole ecosystem LE. Thus, in the present study, we examined what factors control the partitioning of whole ecosystem LE and CO₂ flux into the understory and overstory vegetation, using detailed leaf-level physiology (for both understory and overstory vegetation) and soil respiration property measurements as well as a multilayer soil-vegetation-atmosphere transfer (SVAT) model. The modeling results showed that the larch overstory's leaf area index (LAI) and vertical profile of leaf photosynthetic capacity were major factors determining the flux partitioning in this boreal forest ecosystem. This is unlike other forest ecosystems that tend to have dense LAI. We concluded that control of the larch overstory's LAI had a relationship with both the coexistence of the larch with the cowberry understory and with the water resources available to the total forest ecosystem.     

Application of a temperature-dependent von Bertalanffy growth model to bullhead (Cottus gobio)

The most studied and commonly applied model of fish growth is the von Bertalanffy model. However, this model does not take water temperature into account, which is one of the most important environmental factors affecting the life cycle of fish, as many physiological processes that determine growth, e.g. metabolic rate and oxygen supply, are directly influenced by temperature. In the present study we propose a version of the von Bertalanffy growth model that includes mean annual water temperatures by correlating the growth coefficient, k, explicitly and the asymptotic length, L_∞, implicitly to water temperature. All relationships include parameters with an obvious biological relevance that makes them easier to identify. The model is used to fit growth data of bullhead (Cottus gobio) at different locations in the Bez River network (Drme, France). We show that temperature explains much of the growth variability at the different sampling sites of the network.     

Climate, canopy conductance and leaf area development controls on evapotranspiration in a boreal coniferous forest over a 10-year period: A united model assessment

The aim of this work was to test a process-based model (hydrological model combined with forest growth model) on the simulation of seasonal variability of evapotranspiration (ET) in an even-aged boreal Scots pine (Pinus sylvestris L.) stand over a 10 year period (1999-2008). The water flux components (including canopy transpiration (E_t) and evaporation from canopy (E_c) and ground surface (E_g) were estimated in order to output the long-term stand water budget considering the interaction between climate variations and stand development. For validation, half-hourly data on eddy water vapor fluxes were measured during the 10 growing seasons (May-September). The model predicted well the seasonal course of ET compared to the measured values, but slightly underestimated the water fluxes both in non-drought and drought (2000, 2003 and 2006) years. The prediction accuracy was, on average, higher in drought years. The simulated ET over the 10 years explained, on average, 58% of the daily variations and 84% of the monthly amount of ET. Water amount from E_t contributed most to the ET, with the fractions of E_t, E_c and E_g being, on average, 67, 11 and 23% over the 10-year period, respectively. Regardless of weather conditions, the daily ET was strongly dependent on air temperature (T_a) and vapor pressure deficit (D_a), but less dependent on soil moisture (W_s). On cloudy and rainy days, there was a non-linear relationship between the ET and solar radiation (R_o). During drought years, the model predicted lower daily canopy stomatal conductance (g_cs) compared with non-drought years, leading to a lower level of E_t. The modeled daily g_cs responded well to D_a and W_s. In the model simulation, the annual LAI increased by 35% between 1999 and 2008. The ratio of E_c: ET correlated strongly with LAI. Furthermore, LAI reduced the proportion of E_g as a result of the increased share of E_c and E_t and radiation interception. Although the increase of LAI affected positively E_t, the contribution of E_t in ET was not significantly correlated with LAI. To conclude, although the model predicted reasonably well the seasonal course of ET, the calculation time steps of different processes in the model should be homogenized in the future to increase the prediction accuracy.     

A comparison of four process-based models and a statistical regression model to predict growth of Eucalyptus globulus plantations

In forest management and ecological research, consideration of the impacts and risks of climate change or management optimisation is complex. Computer models have long been applied as tools for these tasks. Process-based forest growth models claim to overcome the limitations of empirical statistical models, but the capacity of different process-based models and modelling approaches have rarely been compared directly. This study evaluates stepwise multiple regression models in comparison to four process-based modelling approaches (3-PG, 3-PG+, CABALA and Forest-DNDC) for greenfield predictions of Eucalyptus globulus plantation growth from 2 to 8 years after planting throughout southern Australia.     

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.     

Zooplankton population model coupled to a biogeochemical model of the North Western Mediterranean Sea ecosystem

A stage structured population (SSP) model based on Fennel's [Fennel, W., 2001. Modelling copepods with links to circulation models. Journal of Plankton Research, 23, 1217–1232] equations is applied to Centropages typicus (Kröyer), a dominant copepod species of the North Western Mediterranean Sea (NWMS) and a prey of small pelagic fish. The model considers five groups of stages and development rates are represented by a mechanistic formulation depending on individual specific growth in each stage. Individual growth is calculated from the individual energy budget depending on food availability and temperature.     

Sanitary felling of Norway spruce due to spruce bark beetles in Slovenia: A model and projections for various climate change scenarios

A model is presented to predict sanitary felling of Norway spruce (Picea abies) due to spruce bark beetles (Ips typographus, Pityogenes chalcographus) in Slovenia according to different climate change scenarios. The model incorporates 21 variables that are directly or indirectly related to the dependent variable, and that can be arranged into five groups: climate, forest, landscape, topography, and soil. The soil properties are represented by 8 variables, 4 variables define the topography, 4 describe the climate, 4 define the landscape, and one additional variable provides the quantity of Norway spruce present in the model cell. The model was developed using the M5′ model tree. The basic spatial unit of the model is 1 km², and the time resolution is 1 year. The model evaluation was performed by three different measures: (1) the correlation coefficient (51.9%), (2) the Theil's inequality coefficient (0.49) and (3) the modelling efficiency (0.32). Validation of the model was carried out by 10-fold cross-validation. The model tree consists of 28 linear models, and model was calculated for three different climate change scenarios extending over a period until 2100, in 10-year intervals. The model is valid for the entire area of Slovenia; however, climate change projections were made only for the Maribor region (596 km²). The model assumes that relationships among the incorporated factors will remain unchanged under climate change, and the influence of humans was not taken into account. The structure of the model reveals the great importance of landscape variables, which proved to be positively correlated with the dependent variable. Variables that describe the water regime in the model cell were also highly correlated with the dependent variable, with evapotranspiration and parent material being of particular importance. The results of the model support the hypothesis that bark beetles do greater damage to Norway spruce artificially planted out of its native range in Slovenia, i.e., lowlands and soils rich in N, P, and K. The model calculation for climate change scenarios in the Maribor region shows an increase in sanitary felling of Norway spruce due to spruce bark beetles, for all scenarios. The model provides a path towards better understanding of the complex ecological interactions involved in bark beetle outbreaks. Potential application of the results in forest management and planning is discussed.     

Simulation modeling to understand how selective foraging by beaver can drive the structure and function of a willow community

Beaver–willow (Castor-Salix) communities are a unique and vital component of healthy wetlands throughout the Holarctic region. Beaver selectively forage willow to provide fresh food, stored winter food, and construction material. The effects of this complex foraging behavior on the structure and function of willow communities is poorly understood. Simulation modeling may help ecologists understand these complex interactions. In this study, a modified version of the SAVANNA ecosystem model was developed to better understand how beaver foraging affects the structure and function of a willow community in a simulated riparian ecosystem in Rocky Mountain National Park, Colorado (RMNP). The model represents willow in terms of plant and stem dynamics and beaver foraging in terms of the quantity and quality of stems cut to meet the energetic and life history requirements of beaver. Given a site where all stems were equally available, the model suggested a simulated beaver family of 2 adults, 2 yearlings, and 2 kits required a minimum of 4 ha of willow (containing about10 stems m⁻²) to persist in a steady-state condition. Beaver created a willow community where the annual net primary productivity (ANPP) was 2 times higher and plant architecture was more diverse than the willow community without beaver. Beaver foraging created a plant architecture dominated by medium size willow plants, which likely explains how beaver can increase ANPP. Long-term simulations suggested that woody biomass stabilized at similar values even though availability differed greatly at initial condition. Simulations also suggested that willow ANPP increased across a range of beaver densities until beaver became food limited. Thus, selective foraging by beaver increased productivity, decreased biomass, and increased structural heterogeneity in a simulated willow community.     

On the use of growth rate parameters for projecting population sizes: Application to aphids

This paper extends the application of the cumulative size based mechanistic model, which has previously been shown to describe diverse aphid population size data well. The mechanistic model is reviewed with a focus on the explanatory role of the birth and death rate formulation. An analysis of two data sets, one on the mustard aphid and the other on the pecan aphid, indicates that multiple linear regression equations based on the estimated birth and death rate parameters alone account for nearly all (R² > 0.95) of the variability in two key population attributes, namely the peak count and the cumulative density. This indicates that population size variables may be projected directly from the growth rate parameters using linear equations. Such linear relationships based on the birth and death rate parameters are shown to hold also for certain generalized mechanistic models for which the analytical solution is not available. The birth and death rate coefficients, therefore, constitute a new succinct set of variables that could be included in the predictive modeling of aphid populations, as well as other insect and animal populations with local collapse which follow similar growth dynamics.     

On the sudden disappearance of Egeria densa from a Ramsar wetland site of Southern Chile: A climatic event trigger model

Contemporary shallow lakes theory proposes that these ecosystems may experience abrupt regime shifts due to small changes in controlling variables or triggers. So far, these triggers have been related mostly to nutrients as the immediate driver. During May 2004 the río Cruces wetland, a Ramsar site located in Southern Chile, underwent a major regime shift, from a clear water state, vastly dominated by the invasive macrophyte Egeria densa, to a turbid water state. In this article we show, through the analysis of long-term meteorological data that late fall 2004 was anomalous due to the presence of a high-pressure cell that persisted most of the month of May over Southern Chile. This climatic event caused an almost complete absence of precipitations and lower temperatures during this period, including several freezing nights. Eco-physiological experiments showed that 6 h exposure to desiccation kill the macrophyte. We developed a simple-biology dynamic model, under Stella Research 9.1, to show that the climatic anomaly of May 2004, plus the increased sedimentation of the wetland's floodplains, and the associated response of E. densa, explains its sudden disappearance from río Cruces wetland.     

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.     

Dynamic model of Lake Chozas (León, NW Spain)—Decrease in eco-exergy from clear to turbid phase due to introduction of exotic crayfish

We developed a dynamic model of the phosphorus cycle in Lake Chozas, a small shallow water body in León (NW Spain). The calibrated model simulated seasonal dynamics of phosphorus concentrations in major components of the lake's ecological network before and after 1997, the year when an invasive allochthonous crustacean, the Louisiana red swamp crayfish (Procambarus clarkii), was introduced into the lake. The shift from clean to turbid phase, due to grazing by crayfish on submerged vegetation, caused a gradual decrease in eco-exergy, reflecting an increase in entropy, related to breakdown of ecosystem internal equilibria. This case study verifies the hypothesis of Marchi et al. (2010) that, after an initial relatively stable state, the allochthonous species may cause an increase in entropy indicating perturbation of the ecosystem.     

An individual-based model to study the reproduction of egg bearing copepods: Application to Eurytemora affinis (Copepoda Calanoida) from the Seine estuary,France

Limited empirical studies have elucidated the daily egg production and associated reproductive processes of egg bearing copepod. Herein, we present an individual-based model which constitutes a realistic representation of the reproduction in egg bearing copepods. The model has been parameterized using an extensive set of experimental data obtained from the literature and from the laboratory and field experiments on the estuarine copepod Eurytemora affinis. The proposed model takes into account the adult female longevity, the clutch size and interclutch duration, which is a function of egg maturation time and latency time required by the female after egg hatching to produce a new clutch. The embryonic development time and hatching success are also taken into account. The effect of temperature on the means and variances of above-mentioned reproductive parameters has been also incorporated. A multi agent system based generic platform “Mobidyc” has been used to generate and calibrate the model. The model demonstrates the reproductive parameters of females of E. affinis which is validated through individual based experiments. Temperature specific simulations provide a dynamical explanation of temperature effect on the cumulative egg production. The daily survival principally affects the number of clutches produced per female during its life span. The results obtained in the present study by combining temperature and survival effects reveal the relatively greater importance of the first factor on the daily egg production of egg-carrying copepods. The present model is generic and hence easily applicable to other animals with comparable reproductive strategy.     

Modelling the responses of wildlife to human disturbance: An evaluation of alternative management scenarios for black-crowned night-herons

The impact of anthropogenic disturbance on wildlife is increasing becoming a source of concern as the popularity of outdoor recreation rises. There is now more pressure on site managers to simultaneously ensure the continued persistence of wildlife and provide recreational opportunities. Using ‘Simulation of Disturbance Activities’, a model designed to investigate the impact of recreational disturbance on wildlife, we demonstrate how a simulation modelling approach can effectively inform such management decisions. As an example, we explored the implications of various design and management options for a proposed recreational area containing a historic breeding bird colony. By manipulating the proximity, orientation and intensity of recreation, we were able to evaluate the impact of recreational activities on the behaviour of black-crowned night-heron nestlings (Nycticorax nycticorax). Using a classification and regression tree (CART) procedure to analyse simulation output, we explored the dynamics of multiple strategies in concert. Our analysis revealed that there are inherent advantages in implementing multiple strategies as opposed to any single strategy. Nestlings were not disturbed by recreation when bird-watching facility placement (proximity and orientation) and type were considered in combination. In comparison, proximity alone only led to a <10% reduction in disturbance. Thus we demonstrate how simulation models based on customised empirical data can bridge the gap between field studies and active management, enabling users to test novel management scenarios that are otherwise logistically difficult. Furthermore, such models potentially have broad application in understanding human-wildlife interactions (e.g. exploring the implications of roads on wildlife, probability of bird strikes around airports, etc.). They therefore represent a valuable decision-making tool in the ecological design of urban infrastructures.     

COSMOS,a spatially explicit model to simulate the epidemiology of Cosmopolites sordidus in banana fields

A stochastic individual-based model called COSMOS was developed to simulate the epidemiology of banana weevil Cosmopolites sordidus, a major pest of banana fields. The model is based on simple rules of local movement of adults, egg laying of females, development and mortality, and infestation of larvae inside the banana plants. The biological parameters were estimated from the literature, and the model was validated at the small-plot scale. Simulated and observed distributions of attacks were similar except for five plots out of 18, using a Kolmogorov–Smirnov test. These exceptions may be explained by variation in predation of eggs and measurement error. An exhaustive sensitivity analysis using the Morris method showed that predation rate of eggs, demographic parameters of adults and mortality rate of larvae were the most influential parameters. COSMOS was therefore used to test different spatial arrangements of banana plants on the epidemiology of C. sordidus. Planting bananas in groups increased the time required to colonise plots but also the percentage of banana plants with severe attacks. Spatial heterogeneity of banana stages had no effect on time required to colonise plots but increased the mean level of attacks. Our model helps explain key factors of population dynamics and the epidemiology of this tropical pest.     

Assessing environmental factors in red spruce (Picea rubens Sarg.) growth in the Great Smoky Mountains National Park, USA: From conceptual model, envirogram, to simulation model

This study provides a method for assessing a multiplicity of environmental factors in red spruce growth in the Great Smoky Mountains National Park (GSMNP) of Southeastern USA. Direct and indirect factors in the annual growth increment are first organized into a schematic input-output envirogram (ARIRS), and this information is then used to construct a simulation model (ARIM). The envirogram represents a structured conceptualization of most environmental factors involved in growth, as developed from relevant literature. This interdisciplinary synthesis distinguishes direct vs. indirect factors in growth and takes account of the systems ecology concept that indirect factors may be as important as or more important than direct ones in regulating growth. The ARIRS envirogram summarizes hierarchically organized, within- and cross-scale, local-to-global interactions, and its construction makes it obvious that growth is influenced by many cross-scale spatiotemporal interactions. More research on genecology is still needed to clarify the role of phenotypic plasticity and adaptive capacity in nutrient cycling, global change, and human disturbance.     

Simulation of NO and N2O emissions from a spruce forest during a freeze/thaw event using an N-flux submodel from the PnET-N-DNDC model integrated to CoupModel

The amount of nitrogen gases (N₂O, NO and N₂) emitted from forest soils depends on interactions between soil properties, climatic factors and soil management. To increase the understanding of nitrogen processes in soil ecosystems, two dynamic models, CoupModel (coupled heat and mass transfer model for soil–plant–atmosphere systems) and the denitrification–decomposition (DNDC) model were selected. Both are dynamic models with different submodels for soil, vegetation, hydrology and climate system. CoupModel has a higher degree of detail on soil physical and abiotic components, whereas the DNDC model contains details of microbiological processes involved in production of nitrogen gases. To improve the previous simple submodel of nitrogen emission in CoupModel, we included a submodel corresponding to the forest version of DNDC containing photosynthesis/evapotranspiration-nitrogen (PnET-N-DNDC model).     

Waterfowls habitat modeling: Simulation of nest site selection for the migratory Little Tern (Sterna albifrons) in the Nakdong estuary

This paper aims to find patterns in nest site selection by Little Terns Sterna albifrons, in the Nakdong estuary in South Korea. This estuary is important waterfowl stopover and breeding habitat, located in the middle of the East Asia-Australasian Flyway. The Little Tern is a common species easily observed near the seashore but their number is gradually declining around the world. We investigated their nests and eggs on a barrier islet in the Nakdong estuary during the breeding season (May to June, 2007), and a pattern for the nest site selection was identified using genetic programming (GP). The GP generated a predictive rule-set model for the number of Little Tern nests (training: R² = 0.48 and test: 0.46). The physical features of average elevation, variation of elevation, plant coverage, and average plant height were estimated to determine the influence on nest numbers for Little Tern. A series of sensitivity analyses stressed that mean elevation and vegetation played an important role in nest distribution for Little Tern. The influence of these two variables could be maximized when elevation changed moderately within the sampled quadrats. The study results are regarded as a good example of applying GP to vertebrate distribution patterning and prediction with several important advantages compared to conventional modeling techniques, and can help establish a management or restoration strategy for the species.     

Modelling effects of chemical exposure on birds wintering in agricultural landscapes: The western burrowing owl (Athene cunicularia hypugaea) as a case study

We describe an ecotoxicological model that simulates the sublethal and lethal effects of chronic, low-level, chemical exposure on birds wintering in agricultural landscapes. Previous models estimating the impact on wildlife of chemicals used in agro-ecosystems typically have not included the variety of pathways, including both dermal and oral, by which individuals are exposed. The present model contains four submodels simulating (1) foraging behavior of individual birds, (2) chemical applications to crops, (3) transfers of chemicals among soil, insects, and small mammals, and (4) transfers of chemicals to birds via ingestion and dermal exposure. We demonstrate use of the model by simulating the impacts of a variety of commonly used herbicides, insecticides, growth regulators, and defoliants on western burrowing owls (Athene cunicularia hypugaea) that winter in agricultural landscapes in southern Texas, United States. The model generated reasonable movement patterns for each chemical through soil, water, insects, and rodents, as well as into the owl via consumption and dermal absorption. Sensitivity analysis suggested model predictions were sensitive to uncertainty associated with estimates of chemical half-lives in birds, soil, and prey, sensitive to parameters associated with estimating dermal exposure, and relatively insensitive to uncertainty associated with details of chemical application procedures (timing of application, amount of drift). Nonetheless, the general trends in chemical accumulations and the relative impacts of the various chemicals were robust to these parameter changes. Simulation results suggested that insecticides posed a greater potential risk to owls of both sublethal and lethal effects than do herbicides, defoliants, and growth regulators under crop scenarios typical of southern Texas, and that use of multiple indicators, or endpoints provided a more accurate assessment of risk due to agricultural chemical exposure. The model should prove useful in helping prioritize the chemicals and transfer pathways targeted in future studies and also, as these new data become available, in assessing the relative danger to other birds of exposure to different types of agricultural chemicals.     

Allowing macroalgae growth forms to emerge: Use of an agent-based model to understand the growth and spread of macroalgae in Florida coral reefs,with emphasis on Halimeda tuna

The growth patterns of macroalgae in three-dimensional space can provide important information regarding the environments in which they live, and insights into changes that may occur when those environments change due to anthropogenic and/or natural causes. To decipher these patterns and their attendant mechanisms and influencing factors, a spatially explicit model has been developed. The model SPREAD (SPatially-explicit Reef Algae Dynamics), which incorporates the key morphogenetic characteristics of clonality and morphological plasticity, is used to investigate the influences of light, temperature, nutrients and disturbance on the growth and spatial occupancy of dominant macroalgae in the Florida Reef Tract. The model species, Halimeda and Dictyota spp., are modular organisms, with an “individual” being made up of repeating structures. These species can also propagate asexually through clonal fragmentation. These traits lead to potentially indefinite growth and plastic morphology that can respond to environmental conditions in various ways. The growth of an individual is modeled as the iteration of discrete macroalgal modules whose dynamics are affected by the light, temperature, and nutrient regimes. Fragmentation is included as a source of asexual reproduction and/or mortality. Model outputs are the same metrics that are obtained in the field, thus allowing for easy comparison. The performance of SPREAD was tested through sensitivity analysis and comparison with independent field data from four study sites in the Florida Reef Tract. Halimeda tuna was selected for initial model comparisons because the relatively untangled growth form permits detailed characterization in the field. Differences in the growth patterns of H. tuna were observed among these reefs. SPREAD was able to closely reproduce these variations, and indicate the potential importance of light and nutrient variations in producing these patterns.