A proposal of an indicator for quantifying model robustness based on the relationship between variability of errors and of explored conditions

The evaluation of biophysical models is usually carried out by estimating the agreement between measured and simulated data and, more rarely, by using indices for other aspects, like model complexity and overparameterization. In spite of the importance of model robustness, especially for large area applications, no proposals for its quantification are available. In this paper, we would like to open a discussion on this issue, proposing a first approach for a quantification of robustness based on the variability of model error to variability of explored conditions ratio. We used modelling efficiency (EF) for quantifying error in model predictions and a normalized agrometeorological index (SAM) based on cumulated rainfall and reference evapotranspiration to characterize the conditions of application. Population standard deviations of EF and SAM were used to quantify their variability. The indicator was tested for models estimating meteorological variables and crop state variables. The values provided by the robustness indicator (I_R) were discussed according to the models’ features and to the typology and number of processes simulated. I_R increased with the number of processes simulated and, within the same typology of model, with the degree of overparameterization. No correlation were found between I_R and two of the most used indices of model error (RRMSE, EF). This supports its inclusion in integrated systems for model evaluation.     

Directional change in a suite of foraging behaviors in tropical and temperate evolved honey bees (<Emphasis Type="Italic">Apis mellifera</Emphasis> L.)

The concept of a suite of foraging behaviors was introduced as a set of traits showing associative directional change as a characterization of adaptive evolution. I report how naturally selected differential sucrose response thresholds directionally affected a suite of honey bee foraging behaviors. Africanized and European honey bees were tested for their proboscis extension response thresholds to ascending sucrose concentrations, reared in common European colonies and, captured returning from their earliest observed foraging flight. Race constrained sucrose response threshold such that Africanized bees had significantly lower sucrose response thresholds. A Cox proportional hazards regression model of honey bee race and sucrose response threshold indicated that Africanized bees were 29% (P<0.01) more at risk to forage over the 30-day experimental period. Sucrose response threshold organized age of first foraging such that each unit decrease in sucrose response threshold increased risk to forage by 14.3% (P<0.0001). Africanized bees were more likely to return as pollen and water foragers than European foragers. Africanized foragers returned with nectar that was significantly less concentrated than European foragers. A comparative analysis of artificial and naturally selected populations with differential sucrose response thresholds and the common suite of directional change in foraging behaviors is discussed. A suite of foraging behaviors changed with a change in sucrose response threshold that appeared as a product of functional ecological adaptation.Communicated by R.F.A. Moritz     

Beurteilung von Bodenkontaminationen mit Radioaktivität im Gebiet Hannover-List nach Maßstäben und Ansätzen der BBodSchV

Background, aim, and scope In Part 1 of this paper test thresholds for radionuclides in soil have been derived on the basis of methods and scales of the German Ordinance on Soil Protection (BBodSchV). In this paper, these test thresholds are checked with regard to their plausibility. Materials, methods, results are described in Part 1 of this paper. Discussion This contribution demonstrates that the test thresholds derived in Part 1 are plausible and feasible according to the requirements of the established methods. The epidemiological basis for the treatment of ionizing radiation is well founded. Risks can be quantified via the knowledge of cancer risks due to radiation. As a reference value the overall fatal risk coefficient of 5?% per Sv recommended by ICRP is used. The test thresholds obtained are sufficiently far from background levels in soils of northern Germany. Consequently, the test thresholds are applicable without any background corrections. Exposures by external radiation outdoors and exposures by inhalation of radon in cellar rooms result in calculated test thresholds in the order of background levels. Nevertheless, feasible results are obtained because in the case of toxic substances without threshold levels the additional exposures are considered for determination of test thresholds. Conclusions The approach of test thresholds for radioactive soil contamination presented in Part 1 of this paper complies with the criteria of the German methods for calculation of test thresholds in soil protection regarding plausibility and feasibility. It enables the soil protection authorities to assess radioactive contaminations of the ground according to the established rules in the field of soil protection and to make decisions according to Articles 9 and 10 of the German Soil Protection Act. Part 3 of this paper discusses the application of the test thresholds and the possibilities of determining the need for remediation actions.     

Comparison of six correlative models in predictive vegetation mapping on a local scale

A variety of statistical techniques has been used in predictive vegetation modelling (PVM) that attempt to predict occurrence of a given community or species in respect to environmental conditions. We compared the performance of three profile models, BIOCLIM, GARP and MAXENT with three nonparametric models of group discrimination techniques, MARS, NPMR and LRT. The two latter models are relatively new statistical techniques that have just entered the field of PVM. We ran all models on a local scale for a given grassland community (Teucrio-Seslerietum) using the same input data to examine their performance. Model accuracy was evaluated both by Cohen’s kappa statistics (κ) and by area under receiver operating characteristics curve based both on resubstitution of training data and on an independent test data set. MAXENT of profile models and MARS of group discrimination techniques achieved the best prediction.     

Brood pheromone modulates honeybee (Apis mellifera L.) sucrose response thresholds

Foraging behavior and the mechanisms that regulate foraging activity are important components of social organization. Here we test the hypothesis that brood pheromone modulates the sucrose response threshold of bees. Recently the honeybee proboscis extension response to sucrose has been identified as a ”window” into a bee’s perception of sugar. The sucrose response threshold measured in the first week of adult life, prior to foraging age, predicts forage choice. Bees with low response thresholds are more likely to be pollen foragers and bees with high response thresholds are more likely to forage for nectar. There is an associated genetic component to sucrose response thresholds and forage choice such that bees selected to hoard high quantities of pollen have low response thresholds and bees selected to hoard low quantities of pollen have higher response thresholds. The number of larvae in colonies affects the number of bees foraging for pollen. Hexane-extractable compounds from the surface of larvae (brood pheromone) significantly increase the number of pollen foragers. We tested the hypothesis that brood pheromone decreases the sucrose response threshold of bees, to suggest a pheromone- modulated sensory-physiological mechanism for regulating foraging division of labor. Brood pheromone significantly decreased response thresholds as measured in the proboscis extension response assay, a response associated with pollen foraging. A synthetic blend of honeybee brood pheromone stimulated and released pollen foraging in foraging bioassays. Synthetic brood pheromone had dose-dependent effects on the modulation of sucrose response thresholds. We discuss how brood pheromone may act as a releaser of pollen foraging in older bees and a primer pheromone on the development of response thresholds and foraging ontogeny of young bees. Received: 24 May 2000 / Revised: 26 September 2000 / Accepted: 15 October 2000     

Beurteilung von Bodenkontaminationen mit Radioaktivität im Gebiet Hannover-List nach Maßstäben und Ansätzen der BBodSchV

Background, aim, and scope In the district List of Hanover (Lower Saxony) radioactive contaminations of the ground were detected at a site of a former chemical plant. Due to the lack of an ordinance regarding intervention regulations in the case of radioactive contaminations in Germany this situation had to be assessed on the basis of scales and methods of the German regulations concerning soil protection and contaminated sites. In particular it was necessary to develop methods and levels for the assessment of radioactive contaminations. Materials and methods Because radioactivity can be considered as a carcinogenic substance the methodical approaches of the BBodSchV for this group of substances were used in order to derive test thresholds for radioactive contaminations at children’s play areas, residential areas as well as parks and recreation facilities. Results For the assessment of radioactive soil contaminations with naturally occurring radionuclides at children’s play areas and residential areas the ingestion of soil is the decisive pathway of exposure. For children’s play areas a threshold level of 0.2?Bq/g for the sum U-238,max?+?Th-232,max was obtained. At areas with only impacts of ambient radiation from the contaminated ground test thresholds of 0.5?µSv/h are recommended. A special pathway is the migration of radon from the contaminated soil into basement floors of buildings. Taking into account the natural background levels of radon a concentration of 260?Bq/m³ is suggested as a test threshold in the framework of soil protection benchmarks. Discussion and conclusions will be described in Part 2 of the paper.     

Species-specific dinoflagellate vertical distribution in temperature-stratified waters

Thermal stratification is increasing in strength as a result of higher surface water temperature. This could influence the vertical distribution of vertically migrating dinoflagellates. We studied the diel vertical distribution of the dinoflagellates Heterocapsa triquetra and Prorocentrum minimum using stratified laboratory columns with two thermoclines of different strength (ΔT° = 10 or 17 °C), with below cline temperature of 8 °C. Above the thermocline, nutrient depletion simulated the natural summer conditions in the Baltic Sea. Our study shows that H. triquetra and P. minimum can behave differently in terms of their vertical occurrence, both in space and in time when subjected to thermoclines of different strength. Also, both dinoflagellate species showed species-specific distribution patterns. In the ΔT° = 10 °C treatment, H. triquetra cells performed a diel vertical migration (DVM) behavior just above the thermocline, but not in the ΔT° = 17 °C. In the ΔT° = 17 °C, the cells did not migrate and cell densities in the water column decreased over time. Opposing results were observed for P. minimum, where a DVM pattern was found exclusively below the thermocline of ΔT° = 17 °C, while in the ΔT° = 10 °C treatment, no clear DVM pattern was observed, and the highest number of cells were found in the cold bottom water. These results indicate that an increase in thermal stratification can influence species-specific dinoflagellate distribution, behavior, and survival.     

A two-dimensional stochastic model of downy mildew of radish

A two-dimensional stochastic model that simulates the spread of disease over space and time was recently proposed by Xu and Ridout [Xu, X.M., Ridout, M.S., 1998. Effects of initial epidemic conditions, sporulation rate, and spore dispersal gradient on the spatio-temporal dynamics of plant disease epidemics. Phytopathology 88, 1000–1012]. In a theoretical study, the authors showed the ability of their model to generate a broad range of disease patterns and disease progress rates. The objective of our study was to test if this theoretical approach was able to describe disease progress and the disease pattern of a specific disease, downy mildew (Peronospora parasitica) of radish (Raphanus sativus L.). Two field experiments with artificial inoculation were carried out and disease incidence and spatial pattern were assessed twice a week until disease incidence was greater than 0.25. Four model parameters were estimated by an algorithm that uses a least square regression together with an evolutionary optimisation strategy. Moran's I indices of spatial autocorrelation calculated both for measured und simulated data were significantly correlated (α = 0.05, r = 0.61). Also observed variances in measurements and in simulations were closely and significantly correlated (α = 0.05, r = 0.95). Thus, disease pattern (as assessed in terms of variance inflation and spatial autocorrelation) was well described by the model. The model accounted for 94% of the variation in the disease incidence data. It has, therefore, the potential to be developed into a forecast model for risk analysis and for decision support in plant protection. However, in the specific case of downy mildew on radish more experimental data are required for model validation and to parameterise the effects of environment on infection, sporulation and spore dispersal.     

Experimental and environmental factors affect spurious detection of ecological thresholds

Threshold detection methods are increasingly popular for assessing nonlinear responses to environmental change, but their statistical performance remains poorly understood. We simulated linear change in stream benthic macroinvertebrate communities and evaluated the performance of commonly used threshold detection methods based on model fitting (piecewise quantile regression [PQR]), data partitioning (nonparametric change point analysis [NCPA]), and a hybrid approach (significant zero crossings [SiZer]). We demonstrated that false detection of ecological thresholds (type I errors) and inferences on threshold locations are influenced by sample size, rate of linear change, and frequency of observations across the environmental gradient (i.e., sample-environment distribution, SED). However, the relative importance of these factors varied among statistical methods and between inference types. False detection rates were influenced primarily by user-selected parameters for PQR (tau) and SiZer (bandwidth) and secondarily by sample size (for PQR) and SED (for SiZer). In contrast, the location of reported thresholds was influenced primarily by SED. Bootstrapped confidence intervals for NCPA threshold locations revealed strong correspondence to SED. We conclude that the choice of statistical methods for threshold detection should be matched to experimental and environmental constraints to minimize false detection rates and avoid spurious inferences regarding threshold location.     

Parameter estimation for distributed delay based population models from laboratory data: egg hatching of Oulema duftschmidi Redthenbacher (Coleoptera,Chrysomelidae) as an example

The cohort development of poikilotherms under favorable temperature conditions may be described by the time-invariant distributed delay models that require three parameters, i.e. the thermal threshold T₀, the thermal constant W and the variability parameter H representing distributed maturation times, also known as time distributed or stochastic development. Here, a parsimonious method is developed that uses stage-frequency matrices obtained under controlled conditions. The analysis of these matrices permits the estimation of the three parameters, while the incorporation of both developmental threshold and thermal constant into the time-invariant distributed delay model permits the representation of stochastic cohort development under constant temperatures. The evaluated parameters can also be used in time-varying distributed delay models that are particularly useful for a wide range of fluctuating temperature conditions.We consider stage-frequency matrices obtained from cohorts of eggs and larvae of Oulema duftschmidi Redthenbacher at different constant temperatures. A visual inspection of the matrices clearly shows the temperature dependent as well as the time distributed passage from the egg to the larval stage. Under the current range of temperatures, a linear model for developmental rate satisfactorily describes egg development, and estimates the thermal threshold (T₀=11.2 °C), the thermal constant (W=81.3 day-degrees) and the delay order parameter (H=70). The resulting model can be used to represent the development of cohorts of O. duftschmidi eggs in a favorable temperature range.     

Evaluation of information indices as indicators of environmental stress in terrestrial soils

Information indices from Ecosystem Network Analysis (ENA) can be used to quantify the development of an ecosystem in terms of its size and organization. There are two types of indices, i.e. absolute indices that describe both the size and organization of ecosystem (Total System Throughput (TST)—system size, Ascendancy (A)—size of organized flows and Development Capacity (C)—upper limit for A, Overhead (L)—size of unorganized flows) and relative indices that describe only the organization (Average Mutual Information (AMI = A:TST), Flow Diversity (H = C:TST), Relative Overhead (RL = L:TST)).It is theorized that environmental stress impair the ecosystem development and that the effect of stress can be quantified with the ENA information indices. Here we applied ENA on a case of environmental stress in a terrestrial ecosystem, i.e. soils that have endured long-term exposure to elevated copper concentration and altered pH.The absolute indices showed an unexpected pattern of response to pollution, suggesting that ecosystems in polluted soils are more active and better organized than these in unpolluted soils. The relative indices, alternatively, responded to pollution as predicted by theory, i.e. with decrease of stress (pollution level) the level of specialization increased (increase of AMI) and losses of energy, e.g. due to respiration, decreased (decrease of Overhead). The diversity and evenness of flows showed hump-backed relationship with stress. Less polluted soils appeared to be less vulnerable to external disturbances and more efficient in processing energy (higher Relative Ascendancy (RA = A:C)) than polluted soils. The relative information indices were rigid to changes in values of assumed parameters. The relative indices, opposite to absolute indices, appeared to be useful as indicators of environmental stress on the ecosystem level.     

Aggressive thresholds in Dendropsophus ebraccatus: habituation and sensitization to different call types

Males in many chorusing anuran species use aggressive calls during defense of calling spaces from other males. The minimal intensity of another male’s vocalizations that elicits an aggressive call response has been termed the aggressive threshold. Previous studies of aggressive thresholds have shown that they are plastic: males habituated (increased their aggressive thresholds) in response to repeated presentation of stimuli above initial threshold levels. Habituation likely contributes to the stable chorus structure of these species, in which aggressive calling is rare compared to advertisement calls. I have observed high levels of aggressive calling in the treefrog Dendropsophus ebraccatus, suggesting that males of this species do not habituate. In this study, I investigated the plasticity of aggressive thresholds in D. ebraccatus. I measured the aggressive thresholds of males before and after suprathreshold stimulation by both advertisement and aggressive calls. I found that the different call types had different effects: males habituated to advertisement calls but lowered their aggressive thresholds in response to aggressive calls. I consider the latter response to be an example of sensitization, a behavior that has been documented infrequently in vocalizing anurans. Sensitization is a plausible mechanism responsible for the high levels of aggressive calling observed in this species. Given the high costs of aggressive calling, however, it is unclear why a mechanism that increases aggressive call output would be maintained.     

A simulation model for the management of vicuña populations

The population dynamics of the vicuña was modelled based on field data from the Central Galeras Sector, Perú. Rains were simulated in order to obtain net primary productivity and grass availability which act upon density-dependent fecundity and mortality. The model produces a population growth curve that tends to stabilize at densities around 100 vicuñas per km². Harvest and shearing processes were simulated as part of the vicuña population management model. Harvest was based upon a fixed threshold density D_e below which no harvest occurs; for densities above D_e harvesting is applied at a rate proportional to the difference between D_e and the current population density (fixed escapement or ‘bang-bang’ harvest rule). Management optimization was analyzed by determining the optimal escapement density D_e, which maximized either net profits or the number of animals harvested. Analyzing the results as cumulative totals over 20 years of simulation, an optimum harvest production was obtained for a D_e of about 40 vicuñas per km², and maximum profit was obtained for a D_e around 70 vicuñas per km². Analyzing the results of 20 years of simulation as annual averages with an original population density of 40 vicuñas per km², the harvest production and profits were maximized for a D_e of 40 and 60 vicuñas per km², respectively.The model was validated using data from the Reserva Nacional San Guillermo, Argentina, where vicuña and guanaco populations coexist (although their interaction was not modelled). The sensitivity analysis was performed with three different techniques: (a) stepwise multiple linear regression, (b) visual graphic analysis based on a polar coordinates system, and (c) direct evaluation of the effect on management decisions. The curve shape parameters of the fecundity and mortality functions proved to be the most important ones in determining the outcome of the model.     

Mineral licks: motivational factors for visitation and accompanying disease risk at communal use sites of elk and deer

Free-ranging cervids acquire most of their essential minerals through forage consumption, though occasionally seek other sources to account for seasonal mineral deficiencies. Mineral sources occur as natural geological deposits (i.e., licks) or as anthropogenic mineral supplements. In both scenarios, these sources commonly serve as focal sites for visitation. We monitored 11 licks in Rocky Mountain National Park, north-central Colorado, using trail cameras to quantify daily visitation indices (DVI) and soil consumption indices (SCI) for Rocky Mountain elk (Cervus elaphus) and mule deer (Odocoileus hemionus) during summer 2006 and documented elk, mule deer, and moose (Alces alces) visiting licks. Additionally, soil samples were collected, and mineral concentrations were compared to discern levels that explain rates of visitation. Relationships between response variables; DVI and SCI, and explanatory variables; elevation class, moisture class, period of study, and concentrations of minerals were examined. We found that DVI and SCI were greatest at two wet, low-elevation licks exhibiting relatively high concentrations of manganese and sodium. Because cervids are known to seek Na from soils, we suggest our observed association of Mn with DVI and SCI was a likely consequence of deer and elk seeking supplemental dietary Na. Additionally, highly utilized licks such as these provide an area of concentrated cervid occupation and interaction, thus increasing risk for environmental transmission of infectious pathogens such as chronic wasting disease, which has been shown to be shed in the saliva, urine, and feces of infected cervids.     

A peaked function for modeling temperature dependence of plant productivity

We suggests that temperature response of plant productivity can be modeled by the Arrhenius function modified to describe the effect of temperature on enzyme activity: G_A(T) = 2f(T)/(1 + f²(T)), where f(T) = exp(E_a/RT_opt − E_a/RT), R the universal gas constant, E_a the activation energy and T_opt is the optimal temperature. In common with other functions used for modeling the temperature response of plant productivity, the curve of function G is almost symmetrical and bell-shaped. The special convenience of G_A is that it relates the width of the “bell” to thermodynamic concepts, such as activation energy of chemical reactions converting carbon dioxide and water to carbohydrates.     

Surface modelling of global terrestrial ecosystems under three climate change scenarios

A high accuracy and speed method (HASM) of surface modelling is developed to find a solution for error problem and to improve computation speed. A digital elevation model (DEM) is established on spatial resolution of 13.5 km × 13.5 km. Regression formulations among temperature, elevation and latitude are simulated in terms of data from 2766 weather observation stations scattered over the world by using the 13.5 km × 13.5 km DEM as auxiliary data. Three climate scenarios of HadCM3 are refined from spatial resolution of 405 km × 270 km to 13.5 km × 13.5 km in terms of the regression formulations. HASM is employed to simulate surfaces of mean annual bio-temperature, mean annual precipitation and potential evapotranspiration ratio during the periods from 1961 to 1990 (T₁), from 2010 to 2039 (T₂), from 2040 to 2069 (T₃), and from 2070 to 2099 (T₄) on spatial resolution of 13.5 km × 13.5 km. Three scenarios of terrestrial ecosystems on global level are finally developed on the basis of the simulated climate surfaces. The scenarios show that all polar/nival, subpolar/alpine and cold ecosystem types would continuously shrink and all tropical types, except tropical rain forest in scenario A1Fi, would expand because of the climate warming. Especially at least 80% of moist tundra and 22% of nival area might disappear in period T₄ comparing with the ones in the period T₁. Tropical thorn woodland might increase by more than 97%. Subpolar/alpine moist tundra would be the most sensitive ecosystem type because its area would have the rapidest decreasing rate and its mean center would shift the longest distance towards west. Subpolar/alpine moist tundra might be able to serve as an indicator of climatic change. In general, climate change would lead to a continuous reduction of ecological diversity.     

Development of clade-specific Symbiodinium primers for quantitative PCR (qPCR) and their application to detecting clade D symbionts in Caribbean corals

We developed quantitative PCR (qPCR) assays to distinguish each of the four clades (A–D) of dinoflagellate endosymbionts (genus Symbiodinium) commonly found in Caribbean corals. We applied these primer sets, which target portions of the multi-copy ribosomal DNA (rDNA) gene family, to assess the presence/absence of symbionts in clade D (as indicated by the detection of clade D DNA). We detected these symbionts in five of six Caribbean host species/genera (21% of samples analyzed, N = 10 of 47 colonies), from which clade D had rarely or never been observed. This suggests that Symbiodinium in clade D are present in a higher diversity of coral species than previously thought. This qPCR-based approach can improve our understanding of the total microbial diversity associated with corals, particularly in hosts thought to be relatively specific, and has many other potential applications for studies of coral reef ecology and conservation.     

State subsidies induce gray jays to accept greater danger: an ecologically rational response?

Models of strictly rational choice assume that decision-makers evaluate options on relevant dimensions, assign fixed values to options, and then make consistent choices based on these values. If so, recent experience would have no impact on preference. But, recent events change an animal’s state, and preference may change accordingly. We explore how state affects willingness to accept greater danger to obtain larger food rewards. We tested how a supplement in state (hoard size) impacts this willingness in gray jays (Perisoreus canadensis). When subsidized, most of the subjects increased their willingness to trade danger for food. Why would they become less cautious when their hoard was increased? Superficially, it might seem prudent to play it safer in response to a subsidy. But imagining fitness as a sigmoid function of state (hoard size) provides a tentative explanation for our counterintuitive finding. Above a threshold hoard size, a subsidy should weaken the willingness to accept extra danger. Incremental increases in state in the deceleratory phase yield smaller fitness gains, so it would pay to increase emphasis on safety after receiving a subsidy. But below this threshold, incremental increases in state in the acceleratory phase yield bigger fitness gains, and so it would pay to decrease emphasis on safety after receiving a subsidy. Most of our subjects’ choice behavior was, thus, plausibly consistent with the possibility that effective hoard size is considerably smaller than the total number of items stored. We speculate that this response may reflect an ecologically rational compensation for the inevitable loss of hoards via theft and rot.     

Release thresholds strongly determine the range of seed dispersal by wind

The effect of the seed abscission process on the dispersal distance of seeds has never been studied explicitly and is often ignored in studies that aim to estimate the seed shadows of species. To examine the importance of the abscission process for the seed shadow we used a seed trajectory model that keeps track of the release threshold dynamics of the individual seeds on mother plant. We defined the release threshold as the critical wind speed that induces a mechanical force that is just large enough to release a seed from its mother plant. The model used real wind speed sequences and seed appearance over time on the mother plant.Several calculations were performed to investigate the effect of release thresholds dynamics on seed shadow of two herbaceous species with contrasting terminal velocity values (V_t): Centaurea jacea (V_t = 4.1 m s⁻¹) and Hypochaeris radicata (V_t = 0.49 m s⁻¹).Release thresholds were responsible for a two-fold increase of median dispersal distances in both species. Tails of the seed shadows, the fraction of seeds that travel furthest, were even more sensitive and increased with a factor 4.5 for Centaurea and 7.0 for Hypochaeris. Our work indicates that the abscission process appears to be very important and suggests that dispersal distance of plants is currently severely underestimated, which, in turn, has major consequences for our current understanding of the distribution, metapopulation dynamics and survival of plant species.