A model approach to evaluate the effect of temperature and food concentration on individual life-history and population dynamics of Daphnia

We present a model framework for the simulation of growth and reproduction of Daphnia at varying conditions of food concentration and temperature. The core of our framework consists of an individual level model that simulates allocation of assimilated carbon into somatic growth, maintenance costs, and reproduction on the basis of a closed carbon budget. A fixed percentage of assimilated carbon is allocated into somatic growth and maintenance costs. Special physiological adaptations in energy acquisition and usage allow realistic model performance even at very low food concentrations close to minimal food requirements. All model parameters are based on physiological measures taken from the literature. Model outputs were thoroughly validated on data from a life-table experiment with Daphnia galeata. For the first time, a successful model validation was performed at such low food concentrations. The escalator boxcar train (EBT) was used to integrate this individual level model into a stage-structured population model. In advance to previous applications of the EBT to Daphnia we included an additional clutch compartment into the model structure that accounts for the characteristic time delay between egg deposition and hatching in cladocerans. By linking two levels of biological organisation, this model approach represents a comprehensive framework for studying Daphnia both at laboratory conditions and in the field. We compared outputs of our stage-structured model with predictions by two other models having analogous parameterisation: (i) another individual level Daphnia model (Kooijman–Metz model) and (ii) a classical unstructured population model. In contrast to our Daphnia model, the Kooijman–Metz model lacks the structure to account for the optimisation of energy acquisition and maintenance requirements by individual daphnids. The unstructured population model showed different patterns of population dynamics that were not in concordance with typical patterns observed in the field. Thus, we conclude our model provides a comprehensive tool for the simulation of growth and reproduction of Daphnia and corresponding population dynamics.     

Is natural hatching asynchrony optimal? An experimental investigation of sibling competition patterns in a facultatively siblicidal seabird

In unpredictable environments, any tactic that enables avian parents to adjust brood size and, thus, energy expenditure to environmental conditions should be favoured. Hatching asynchrony (HA), which occurs whenever incubation commences before clutch completion, may comprise such a tactic. For instance, the sibling rivalry hypothesis states that the hierarchy among chicks, concomitant to HA, should both facilitate the adjustment of brood size to environmental conditions and reduce several components of sibling competition as compared to synchronous hatching, at both brood and individual levels. We thus predicted that brood aggression, begging and feeding rates should decrease and that older chick superiority should increase with HA increasing, leading to higher growth and survival rates. Accordingly, we investigated the effects of an experimental upward and downward manipulation of HA magnitude on behaviour, growth and survival of black-legged kittiwake (Rissa tridactyla) chicks. In line with the sibling rivalry hypothesis, synchronous hatching increased aggression and tended to increase feeding rates by parents at the brood level. Begging rates, however, increased with HA contrary to our expectations. At the individual level, as HA magnitude increased, the younger chick was attacked and begged proportionally more often, experienced a slower growth and a higher mortality than its sibling. Overall, the occurrence of energetic costs triggered by synchronous hatching both for parents and chicks, together with the lower growth rate and increased mortality of the younger chick in highly asynchronous broods suggest that natural HA magnitude may be optimal.     

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.     

Modeling effects of toxin exposure in fish on long-term population size, with an application to selenium toxicity in bluegill (Lepomis macrochirus)

A primary goal in ecotoxicology is the prediction of population-level effects of contaminant exposure based on individual-level response. Assessment of toxicity at the population level has predominately focused on the population growth rate (PGR), but the PGR may not be a relevant toxicological endpoint for populations at equilibrium. Equilibrium population size may be a more meaningful endpoint than the PGR because a population with smaller equilibrium size is more susceptible to the negative effects of environmental variability. We address the individual-to-population extrapolation problem with modeling utilizing classical mathematical theory. We developed and analyzed a general model applicable to many freshwater fish species, that includes density-dependent juvenile survival and additional juvenile mortality due to toxicity exposure, and we quantified effect on equilibrium population size as a means of assessing toxicity. Individual-level effects are typically greater than population-level effects until the individual effect is large, due to compensatory density-dependent relationships. These effects are sensitive to the recruitment potential of a population, in particular the low-density first-year survival rate S_b. Assuming high S_b could result in underestimating effects of population-level toxicity. The equilibrium size depends directly on S_b, the reproductive potential, the toxin concentration at which mean mortality is 50% (LC₅₀), and the rate at which individual mortality increases with increasing toxin concentration. More experimental data are needed to decrease the uncertainty in estimating these parameters. We then used existing data for selenium toxicity in bluegill sunfish to parameterize a simulation version of the model as an example to assess the effects of environmental stochasticity on toxicity response. Effects of environmental variability resulted in simulated extinctions at much lower toxin concentrations than predicted deterministically.     

Ecotoxicology and population dynamics: Using DEBtox models in a Leslie modeling approach

Although the ecological risks of toxic chemicals are usually assessed on the basis of individual responses, such as survival, reproduction or growth, ecotoxicologists are now attempting to assess the impact of environmental pollution on the dynamics of naturally exposed populations. The main issue is how to infer the likely impact on the population of the toxic effects observed at the individual level. Dynamic energy budget in toxicology (DEBtox) is the most user-friendly software currently available to analyze the experimental data obtained in toxicity tests performed on individuals. Because toxic effects are diverse and because the sensitivity of individuals varies considerably depending on life-cycle stage, Leslie models offer a convenient way of predicting toxicant effects on population dynamics.In the present study, we first show how parameter inputs, estimated from individual data using DEBtox, can be coupled using a Leslie matrix population model. Then, using experimental data obtained with Chironomus riparius, we show how the effects of a pesticide (methiocarb) on the population growth rate of a laboratory population can be estimated. Lastly, we perform a complex sensitivity analysis to pinpoint critical age classes within the population for the purposes of the field management of populations.     

Performance level and efficiency of two differing predator-avoidance strategies depend on nutritional state of the prey fish

Animal prey has developed a variety of behavioural strategies to avoid predation. Many fish species form shoals in the open water or seek refuge in structurally complex habitats. Since anti-predator strategies bear costs and are energy-demanding, we hypothesised that the nutritional state of prey should modify the performance level and efficiency of such strategies. In aquaria either containing or lacking a structured refuge habitat, well-fed or food-deprived juvenile roach (Rutilus rutilus) were exposed to an open-water predator (pikeperch, Sander lucioperca). Controls were run without predators. In the presence of the predator, roach enhanced the performance of the anti-predator strategy and increased the use of the refuge habitat whereby food-deprived roach were encountered more often in the structure than well-fed roach. Nonetheless more starved than well-fed roach were fed upon by the predator. In the treatments offering only open-water areas, roach always formed dense shoals in the presence of the predator. The shoal density, however, was lower in starved roach. Starving fish in shoals experienced the highest predation mortality across all experimental treatments. The experiment confirmed the plasticity of the anti-predator behaviour in roach and demonstrated that food deprivation diminished the efficiency of shoaling more strongly than the efficiency of hiding. The findings may be relevant to spatial distribution of prey and predator–prey interactions under natural conditions because when prey are confronted with phases of reduced resource availability, flexible anti-predator strategies may lead to dynamic habitat use patterns.     

Modelling the spatial population dynamics of the green oak leaf roller (Tortrix viridana) using density dependent competitive interactions: Effects of herbivore mortality and varying host-plant quality

Cyclic population dynamics of forest insects with periods of more than two generations have been discussed in relation to a variety of extrinsic and intrinsic forces. In the present study, we employed the selection pressure of density dependent competitive interactions according to Witting's equations (Witting, 2000) as driver for a discrete spatiotemporal model of the green oak leaf roller (Tortrix viridana). The model was successfully parameterised to rebuild the cyclic population dynamics of an empirical data set of a 30-year leaf roller monitoring in Russia. Our analysis focussed on the role of herbivore mortality and host plant food quality, which have a significant effect on T. viridana population dynamics. An additional egg or larvae mortality lowers population density and can lead to selection pressures that favour individuals with higher growth rate. This increased population growth rate can not only compensate the additional mortality, but also can lead to higher average moth abundances in subsequent generations. Furthermore, we analysed the effect of inter- and intraspecific variation in host plant quality on herbivore population dynamics and the spatial distribution of abundance and defoliation patterns. We found significant effects of the qualitative composition of a trees neighbourhood on the herbivore population of the respective tree. Also, the patchy damage patterns observable in reality have been reproduced by the present model. The applicability of the model approach and the putative genetic processes underlying Witting's model are discussed.     

Behavioral response to predators reverses the outcome of competition between prey species

Summary In a laboratory experiment it was shown that piscivorous predators reversed the outcome of competitive interactions between two fish prey species, juveniles of roach (Rutilus rutilus) and perch (Perca fluviatilis), by behaviorally affecting their use of two available habitats, an open water habitat and a structurally complex refuge. The shift in the competitive relationship was the result of predators forcing the juvenile fishes into a prey refuge with high structural complexity. While roach was competitively superior in the unstructured habitat, perch was superior in the structurally complex prey refuge. The reversal in competitive relationship was demonstrated both with respect to foraging rate and growth rate and resulted from the high structural complexity in the prey refuge interfering with the roach's swimming performance. Because survival and growth patterns through the juvenile stages have profound effects on the population/community dynamics of size-structured populations such as those of fish, behaviorally induced changes in competitive ability should have significant implications also at the population and community levels.     

Finite population properties of individual predictors based on spatial patterns

In this study a conceptual framework for assessing the statistical properties of a non-stochastic spatial interpolator is developed through the use of design-based finite population inference tools. By considering the observed locations as the result of a probabilistic sampling design, we propose a standardized weighted predictor for spatial data starting from a deterministic interpolator that usually does not provide uncertainty measures. The information regarding the coordinates of the spatial locations is known at the population level and is directly used in constructing the weighting system. Our procedure captures the spatial pattern by means of the Euclidean distances between locations, which are fixed and do not require any further assessment after the sample has been drawn. The predictor for any individual value turns in a ratio of design-based random quantities. We illustrate the predictor design-based statistical properties, i.e. asymptotically p-unbiasedness and p-consistency, for simple random sampling without replacement. An application to a couple of environmental datasets is presented, for assessing predictor performances in correspondence of different population characteristics. A comparison with the equivalent non-spatial predictor is presented.     

Invasion dynamics of the varnish clam (Nuttallia obscurata): a matrix demographic modeling approach

Marine invaders have become a significant threat to native biodiversity and ecosystem function. In this study, the invasion of the varnish clam (Nuttallia obscurata) in British Columbia, Canada, is investigated using a matrix modeling approach to identify the life history characteristics most crucial for population growth and to investigate population differences. Mark-recapture analyses and field collections from 2003 to 2004 were used to determine individual growth, survival rates, and fecundity for two sites. A multi-state matrix model was used to determine population growth rates and to conduct sensitivity and elasticity analyses. A life table response experiment was also used to determine what life history stage contributed most to observed differences in population growth rates. Population survey data were used in conjunction with the matrix model to determine plausible recruitment levels and to investigate recruitment scenarios. Both populations are currently declining but are likely sustainable because of the pulsed nature of large recruitment events. Survival of larger clams (>40 mm) is the most important for population growth based on elasticity and sensitivity analyses. Adult survival also had the largest influence on observed differences between site-specific population growth rates. The two populations studied differed in recruitment dynamics; one experiencing annual recruitment with higher post-settlement mortality and the other, episodic recruitment and lower post-settlement mortality. The most influential factor for the successful invasion of the varnish clam appears to be survival of the larger size classes. Therefore, any process that decreases adult survival (e.g., predation, commercial harvest) will have the greatest impact on population growth.     

Aphid population response to agricultural landscape change: A spatially explicit,individual-based model

A spatially explicit individual-based simulation model has been developed to represent aphid population dynamics in agricultural landscapes. The application of the model to Rhopalosiphum padi (L.) population dynamics is detailed, including an outline of the construction of the model, its parameterisation and validation. Over time, the aphids interact with the landscape and with one another. The landscape is modified by varying a simple pesticide regime, and the multi-scale spatial and temporal implications for a population of aphids is analysed. The results show that a spatial modelling approach that considers the effects on the individual of landscape properties and factors such as wind speed and wind direction provides novel insight into aphid population dynamics both spatially and temporally. This forms the basis for the development of further simulation models that can be used to analyse how changes in landscape structure impact upon important species distributions and population dynamics.     

Growth and production of the inshore marine copepod Pseudodiaptomus marinus in the central part of the Inland Sea of Japan

The growth and production of the inshore marine copepod Pseudodiaptomus marinus was studied in the central part of the Inland Sea of Japan. The stage-specific growth rate was determined under controlled laboratory conditions by examining the length-weight relationship and development rates at various temperatures. The stage duration was short and constant from NII to CII, beyond which development was retarded. Males developed faster than females in CIV and CV. The specific growth rate was highest in copepodite stages followed by the nauplii and adult females (=egg production rate). The daily production of P. marinus was estimated from the stage-specific growth rate and stage-specific abundance in nature as the sum of the individual stages. The production changed seasonally with water temperature and population biomass. Daily production and biomass (P/B) ratios increased linearly with temperature. Total annual production was 20.7 mg C m^-3 yr^-1.     

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.     

Individual-based modeling of flooding and salinity effects on a coastal swamp forest

Coastal swamps are among the rapidly vanishing wetland habitats in Louisiana. Increased flooding, nutrient and sediment deprivation, and salt-water intrusion have been implicated as probable causes of the decline of coastal swamps. We developed a two-species individual-based forest succession model to compare the growth and composition of a cypress-tupelo swamp under various combinations of flooding intensity and salinity levels, using historical time-series of stage and salinity data as inputs. Our model simulates forest succession over 500 years by representing the growth, mortality, and reproduction of individual Taxodium distichum (baldcypress) and Nyssa aquatica (water tupelo) trees in a 1-km² spatial grid of 10 m × 10 m cells that vary in water levels and salinity through differences in elevation. We independently adjusted the elevations of each cell to obtain different grid-wide mean elevations and standard deviations of elevation; this affected the temporal and spatial pattern of flooding. We calibrated the model by adjusting selected parameters until averaged basal area, stem density and wood production rates under two different mean elevations (partially versus highly flooded) were qualitatively similar to comparable values reported for swamps in the literature. Corroboration involved comparing model predictions to four well-monitored contrasting habitat sites within the Maurepas Basin, Louisiana, USA. Model predictions of both species combined showed the same patterns among sites as the data, but the model overestimated wood production and the dominance of T. distichum. Exploratory simulations predicted that increased flooding leads to swamps with reduced basal areas and stem densities, while increased salinity resulted in lower basal areas at low salinity concentration (∼1-3 psu) and complete tree mortality at higher salinity concentrations (∼2-6 psu). Our model can provide insight into the succession dynamics of coastal swamps and information for the effective design of restoration actions.     

Influence of low and decreasing food levels on Daphnia-algal interactions: Numerical experiments with a new dynamic energy budget model

Based on numerical experiments with a new physiologically structured population model we demonstrate that predator physiology under low food and under starving conditions can have substantial implications for population dynamics in predator-prey interactions. We focused on Daphnia-algae interactions as model system and developed a new dynamic energy budget (DEB) model for individual daphnids. This model integrates the κ-rule approach common to net assimilation models into a net-production model, but uses a fixed allocation of net-productive energy in juveniles. The new DEB-model agrees well with the results of life history experiments with Daphnia. Compared to a pure κ-rule model the new allocation scheme leads to significant earlier maturation at low food levels and thus is in better agreement with the data. Incorporation of the new DEB-model into a physiologically structured population model using a box-car elevator technique revealed that the dynamics of Daphnia-algae interactions are highly sensitive to the assumptions on the energy allocation of juveniles under low food conditions. Additionally we show that also other energy allocation rules of our DEB-model concerning decreasing food levels and starving conditions at the individual level have strong implications for Daphnia-algae interactions at the population level. With increasing carrying capacity of algae a stable equilibrium with coexistence of Daphnia occurs and algae shifts to limit cycles. The amplitudes of the limit cycles increase with increasing percentage of sustainable weight loss. If a κ-rule energy allocation is applied to juveniles, the stable equilibrium occurs for a much narrower range of algal carrying capacities, the algal concentration at equilibrium is about 2 times larger, and the range of algae carrying capacities at which daphnids become extinct extends to higher carrying capacities than in the new DEB-model. Because predator-prey dynamics are very sensitive to predator physiology under low food and starving conditions, empirical constraints of predator physiology under these conditions are essential when comparing model results with observations in laboratory experiments or in the field.     

Roach (<Emphasis Type="Italic">Rutilus rutilus</Emphasis>) as an important predator on blue mussel (<Emphasis Type="Italic">Mytilus edulis</Emphasis>) populations in a brackish water environment,the northern Baltic Sea

Suspension-feeding bivalves are organisms of major functional importance in several aquatic environments around the world. They are also important food items for many fish and benthivorous seabirds. It has commonly been thought that predation pressure on blue mussel (Mytilus edulis) populations is negligible in the Baltic Sea, owing to the scarcity of major invertebrate predators such as starfish and crabs. It has recently been shown, however, that the blue mussel is the main food item for roach (Rutilus rutilus) in the archipelago areas of the western Gulf of Finland, where this freshwater fish species has become increasingly abundant, mainly due to increased eutrophication. To quantify the influence of roach predation on blue mussel populations we measured the standing biomass and size structure of the local blue mussel population and used a bioenergetic model to estimate mussel consumption by individual roach during two consecutive summers, 1997 and 1998. The results of the model were combined with existing data on roach abundance, giving annual consumption estimates of 75–105 kg blue mussel dry weight ha^–1 in the study area, approximately two-thirds of these consumed mussels being >10 mm. This corresponds to approximately one-third of the standing population of mussels >10 mm in the area. Our results suggest that the predation effects of vertebrates on Baltic blue mussel populations are not insignificant, as previously believed. Predation by roach and other predators may have an important structuring effect on unstable blue mussel communities within the Gulf of Finland, where the species lives at the edge of its range.Communicated by M. Kühl, Helsingør