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.     

Development and validation of an individual based Daphnia magna population model: The influence of crowding on population dynamics

An individual-based model was developed to predict the population dynamics of Daphnia magna at laboratory conditions from individual life-history traits observed in experiments with different feeding conditions. Within the model, each daphnid passes its individual life cycle including feeding on algae, aging, growing, developing and – when maturity is reached – reproducing. The modelled life cycle is driven by the amount of ingested algae and the density of the Daphnia population. At low algae densities the population dynamics is mainly driven by food supply, when the densities of algae are high, the limiting factor is “crowding” (a density-dependent mechanism due to chemical substances released by the organisms or physical contact, but independent of food competition).     

Data-directed modelling of Daphnia dynamics in a long-term micro-ecosystem experiment

The micro-ecosystem under consideration consists of three compartments forming a closed chain in which water circulates. Three trophic levels are represented in different compartments: autotrophs (algae, mainly Chlorella vulgaris), herbivores (Daphnia magna) and microbial decomposers. From a 20 years experiment with this system, data has been selected for this study. The dynamics of algae and Daphnia magna in only one of the compartments were modeled by different systems of differential and difference equations. We describe the successive steps in the process of model development, and the fitting of parameters using a Nelder-Mead simplex calibration method. Identification problems were overcome by taking values for physiological parameters in agreement with the literature. It turned out that a logistic type of model gives the best result for the structured Daphnia population because of the set up of the experiment: algae grow and reproduce in the upstream compartment. For this reason well-known plant–herbivore models did not comply with the data. The results of the parameter estimation procedure are discussed. The estimated grazing rate by Daphnia was smaller than expected. Possibly the Daphnia fed also on detritus and decomposing algae which were not measured.     

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.     

Effects of resource availability,predators, conspecifics and heterospecifics on decorating behaviour by the majid crab <Emphasis Type="Italic">Tiarinia cornigera</Emphasis>

Utilization of algae for decorating by the intertidal majid crab Tiarinia cornigera was examined by laboratory experiments in relation to availability of algae, presence of a predator, and the presence of conspecific and heterospecific crabs. Different availabilities of decorating materials had a positive correlation on decorating amount by juveniles, while the correlation was not so clear in subadults. The amount of decorating decreased with increasing density of conspecific crabs in the presence of a predator, but there was so clear relationship in the absence of a predator. The decrease in decorating under high density could be due to intraspecific aggression, because the superior crab, in fighting, was found to be decorated with more algae. Tiarinia cornigera was found to be superior to the co-occurring different majid species Micippa platipes in fighting. When T. cornigera and M. platipes were housed together, the former was decorated with more algae than the latter in the presence of a predator, but in the absence of a predator, the amount of algae was not different. Thus, presence of a predator may motivate intraspecific competition in T. cornigera as well as interspecific competition between T. cornigera and M. platipes for decorating materials.     

Patterns of co-occurrence and interactions between age classes of the common triplefin,Forsterygion lapillum

Many marine organisms have pelagic larval stages that settle into benthic habitats occupied by older individuals; however, a mechanistic understanding of intercohort interactions remains elusive for most species. Patterns of spatial covariation in the densities of juvenile and adult age classes of a small temperate reef fish, the common triplefin (Forsterygion lapillum), were evaluated during the recruitment season (Feb–Mar, 2011) in Wellington, New Zealand (41°17′S, 174°46′E). The relationship between juvenile and adult density among sites was best approximated by a dome-shaped curve, with a negative correlation between densities of juveniles and adults at higher adult densities. The curve shape was temporally variable, but was unaffected by settlement habitat type (algal species). A laboratory experiment using a “multiple-predator effects” design tested the hypothesis that increased settler mortality in the presence of adults (via enhanced predation risk or cannibalism) contributed to the observed negative relationship between juveniles and adults. Settler mortality did not differ between controls and treatments that contained either one (p = 0.08) or two (p = 0.09) adults. However, post hoc analyses revealed a significant positive correlation between the mean length of juveniles used in experimental trials and survival of juveniles in these treatments, suggesting that smaller juveniles may be vulnerable to cannibalism. There was no evidence for risk enhancement or predator interference when adults were present alongside a heterospecific predator (F. varium). These results highlight the complex nature of intercohort relationships in shaping recruitment patterns and add to the growing body of literature recognizing the importance of age class interactions.     

Modeling individual and population dynamics in a consumer–resource system: Behavior under food limitation and crowding and the effect on population cycling in Daphnia

In population modeling, a considerable level of complexity is often required to provide trustworthy results, comparable with field observations. By assuring sufficient detail at the individual level while preserving the potential to explore the consequences at higher levels, individual-based modeling may thus provide a useful tool to investigate dynamics at different levels of organization. Still, population dynamics resulting from such models are often at odds with observations from the field. This may be partly caused by a lack of focus on the individual dynamics under conditions of food stress and starvation. I developed a physiologically structured, individual-based simulation model to investigate life history of Daphnia and its effect on population dynamics in response to the productivity of the system. In verifying model behavior with available literature data on life history and physiology, I paid special attention to the dynamics of food intake and the verification of individual level results under conditions of food limitation and starvation. I show that the maximum filtering rates under low food levels used in the current model are much closer to measured filtering rates than the ones used in other models. Being consistent with results on physiology and life history from experiments at a wide range of food availability (including starvation), the model generates low amplitude or high amplitude population density cycles depending on the productivity of the system, as observed in field and experimental populations of Daphnia and with the minimum population densities being one to two orders of magnitude lower in the high amplitude than in the low amplitude cycles. To generate results which are not only qualitatively but also quantitatively comparable to experimental and field observations, however, a crowding effect on the filtering response has to be incorporated in the model.     

Elevated CO2 affects the behavior of an ecologically and economically important coral reef fish

We tested the effect of near-future CO₂ levels (≈490, 570, 700, and 960 μatm CO₂) on the olfactory responses and activity levels of juvenile coral trout, Plectropomus leopardus, a piscivorous reef fish that is also one of the most important fisheries species on the Great Barrier Reef, Australia. Juvenile coral trout reared for 4 weeks at 570 μatm CO₂ exhibited similar sensory responses and behaviors to juveniles reared at 490 μatm CO₂ (control). In contrast, juveniles reared at 700 and 960 μatm CO₂ exhibited dramatically altered sensory function and behaviors. At these higher CO₂ concentrations, juveniles became attracted to the odor of potential predators, as has been observed in other reef fishes. They were more active, spent less time in shelter, ventured further from shelter, and were bolder than fish reared at 490 or 570 μatm CO₂. These results demonstrate that behavioral impairment of coral trout is unlikely if pCO₂ remains below 600 μatm; however, at higher levels, there are significant impacts on juvenile performance that are likely to affect survival and energy budgets, with consequences for predator–prey interactions and commercial fisheries.     

Temperature-dependent ranges of coexistence in a model of a two-prey-one-predator microbial food web

The objective of our study was to analyze the effects of temperature on the population dynamics of a three-species food web consisting of two prey bacteria (Pedobacter sp. and Acinetobacter johnsonii) and a protozoan predator (Tetrahymena pyriformis) as model organisms. We assessed the effects of temperature on the growth rates of all three species with the objective of developing a model with four differential equations based on the experimental data. The following hypotheses were tested at a theoretical level: Firstly, temperature changes can affect the dynamic behavior of a system by temperature-dependent parameters and interactions and secondly, food web response to temperature cannot be derived from the single species temperature response. The main outcome of the study is that temperature changes affect the parameter range where coexistence is possible within all three species. This has significant consequences on our ideas regarding the evaluation of effects of global warming.     

Effects of cadmium and chlordane on cellular apoptosis in ecotoxicological test species

Cadmium and chlordane are compounds chosen for a preliminary study on the impact of environmental contaminants on apoptosis in different representative species. These two potentially toxic substances decrease in vitro apoptosis of Daphnia magna cells. This appears to be related to the chemical concentrations and the time exposure. Cadmium was also directly introduced to the cell culture medium of Syrian hamster embryos (SHE) or indirectly via daphnids, which had firstly consumed algae contaminated with cadmium. A very sharp transfer of cadmium toxicity from algae to daphnids is noted and an inhibition of the apoptosis in SHE cells can be observed.     

Langmuir circulations and plankton patchiness

A computer simulation model of oceanic Langmuir circulations illustrates the role of these vortical water movements as physical forcing mechanisms affecting the horizontal and vertical distributions of planktonic organisms. The diel migration of omnivorous and herbivorous zooplankton (approximately by literature values for the swimming speeds of Meganyctiphanes norvegica and Calanus finmarchicus, respectively) and the sinking of phytoplankton interact with the Langmuir cells, resulting in a spatially heterogeneous community. The patchy distributions, in turn, affect the prey—predator interactions in this simple food web. Situations involving threshold and non-threshold feeding strategies as well as constant and changing circulations are investigated. The simulations are restricted to twilight and nocturnal events.     

Influence of food quantity on the kinetics of cadmium uptake and loss via food and seawater in Mytilus edulis

The difference between the cadmium uptake via food and seawater in Mytilus edulis has been studied. This was done by labelling algae with Cd-109 and seawater with Cd-115m. Mussels were fed on six different quantities of Isochrysis galbana. Cadmium uptake via algae was more efficient at low food levels, while accumulation from seawater was linearly correlated with food quantities. Cadmium from food contributed only little to the body burden (0.2–0.5%). Half-lives for the elimination of cadmium ranged from 96–190 d and increased with decreasing availability of algae, presumably due to slowed down metabolism. Differences in elimination patterns suggest a release of both isotopes from different storage depots. A computer model shows that the food pathway can only play a significant role if algae are highly contaminated. It also demonstrates the paradox that in long-term studies the highest contribution of food-derived cadmium to the body burden must be expected near maintenance food concentrations.     

Modelling predation by transient leopard seals for an ecosystem-based management of Southern Ocean fisheries

Correctly quantifying the impacts of rare apex marine predators is essential to ecosystem-based approaches to fisheries management, where harvesting must be sustainable for targeted species and their dependent predators. This requires modelling the uncertainty in such processes as predator life history, seasonal abundance and movement, size-based predation, energetic requirements, and prey vulnerability. We combined these uncertainties to evaluate the predatory impact of transient leopard seals on a community of mesopredators (seals and penguins) and their prey at South Georgia, and assess the implications for an ecosystem-based management. The mesopredators are highly dependent on Antarctic krill and icefish, which are targeted by regional fisheries. We used a state-space formulation to combine (1) a mark-recapture open-population model and individual identification data to assess seasonally variable leopard seal arrival and departure dates, numbers, and residency times; (2) a size-based bioenergetic model; and (3) a size-based prey choice model from a diet analysis. Our models indicated that prey choice and consumption reflected seasonal changes in leopard seal population size and structure, size-selective predation and prey vulnerability. A population of 104 (90–125) leopard seals, of which 64% were juveniles, consumed less than 2% of the Antarctic fur seal pup production of the area (50% of total ingested energy, IE), but ca. 12–16% of the local gentoo penguin population (20% IE). Antarctic krill (28% IE) were the only observed food of leopard seal pups and supplemented the diet of older individuals. Direct impacts on krill and fish were negligible, but the “escapement” due to leopard seal predation on fur seal pups and penguins could be significant for the mackerel icefish fishery at South Georgia. These results suggest that: (1) rare apex predators like leopard seals may control, and may depend on, populations of mesopredators dependent on prey species targeted by fisheries; and (2) predatory impacts and community control may vary throughout the predator's geographic range, and differ across ecosystems and management areas, depending on the seasonal abundance of the prey and the predator's dispersal movements. This understanding is important to integrate the predator needs as natural mortality of its prey in models to set prey catch limits for fisheries. Reliable estimates of the variability of these needs are essential for a precautionary interpretation in the context of an ecosystem-based management.     

The concepts of emergent and collective properties in individual-based models—Summary and outlook of the Bornhöved case studies

Ecology requires the conceptual and technical ability to analyse complex and dynamic systems consisting of a high and variable number of components and relations. These components are part of a variable interaction structure in a spatially heterogeneous context. The components of ecological interaction networks can give rise to self-organised, and scale-dependent interaction patterns and processes, which are the underlying causes of the overall ecological systems states.The individual-based modelling approach provides a widely applicable simulation framework based on a ‘hierarchy theory’ view of ecological systems.Here, we summarise and generalise the theoretical implications of the modelling studies presented in this volume in the field of terrestrial and aquatic, animal and plant ecology. The case studies cover a representative profile of processes related to ecological applications, such as food web interactions, population dynamics, dispersal, energy physiology, nutrient allocation and mutual impact of morphological and physiological development. The generic approach applied in this context allows a hierarchical representation of ecological systems and their components. Model results are obtained as self-organised structural relation networks and as aggregated quantitative states. In order to address different model characteristics we distinguish collective and emergent properties. Collective properties are those that are attributed equally to different organisation levels of the system. Emergent properties result from the activities of lower level entities on a higher organisation level, while not being present on the lower level. They can be subdivided into aggregational and connective properties. Emergent properties that are aggregational are those which emerge as a result of an aggregation procedure by an observer on the higher level which does not make sense or is not applicable on lower levels. Emergent properties that are connective, however, are based on an interaction network of lower level entities, which brings about the specific system characteristic.This classification of model results will allow to generalise the achievements and potential of the individual-based modelling approach in ecology.     

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.     

Incorporating consumer–resource spatial interactions in reserve design

The persistence of species in reserves depends in large part on the persistence of functional ecological interactions. Despite their importance, however, ecological interactions have not yet been explicitly incorporated into conservation prioritization methods. We develop here a general method for incorporating consumer–resource interactions into spatial reserve design. This method protects spatial consumer–resource interactions by protecting areas that maintain the connectivity between the distribution of consumers and resources. We illustrate our method with a conservation planning case study of a mammalian predator, American marten (Martes americana), and its two primary prey species, Red-backed vole (Clethrionomys rutilus) and Deer mouse (Peromyscus maniculatus). The conservation goal was to identify a reserve for marten that comprised 12% of a forest management unit in the boreal forest in Québec, Canada. We compared reserves developed using analysis variants that utilized different levels of information about predator and prey habitat distributions, species-specific connectivity requirements, and interaction connectivity requirements. The inclusion of consumer–resource interactions in reserve-selection resulted in spatially aggregated reserves that maintained local habitat quality for the species. This spatial aggregation was not induced by applying a qualitative penalty for the boundary length of the reserve, but rather was a direct consequence of modelling the spatial needs of the interacting consumer and resources. Our method for maintaining connectivity between consumers and their resources within reserves can be applied even under the most extreme cases of either complete spatial overlap or complete spatial segregation of consumer–resource distributions. The method has been made available via public software.     

In situ beziehungen zwischen nahrungsangebot und aufgenommener Nahrung bei 5 Blennius-Arten (Pisces) des Mittelmeeres

Eight epiphytal samples were taken from the upper littoral at Banyuls-sur-Mer (French Mediterranean coast) and the fauna living on the thalli of algae or between bivalves was examined. Intestinal food contents of 67 specimens of 5 Blennius species (B. trigloides, B. canevae, B. sphinx, B. incognitus, and B. dalmatinus) occurring in the same biotope were also investigated. The epiphytal samples consisted of a rich supply of amphipods, bivalves, and algae. Amphipods are the preferred food of the Blennius species examined. These fishes also consume large amounts of copepods and algae, and small quantities of halacarids, bivalves, and ostracods. In addition to serving as a substrate for the main food-animals of the fishes, the algae themselves constitute an important food source. Presumably, for these reasons, the fishes remain close to the area inhabited by the algae on rocky substrates.     

Eutrophication alters social preferences in three-spined sticklebacks (Gasterosteus aculeatus)

Algae blooms, which can be caused by eutrophication, drastically influence the ecology and behaviour of aquatic organisms. Such impact is often demonstrated in the context of mate choice and predator–prey interactions. In contrast, the influence of increased turbidity on social behaviour is less well understood, although it may have strong influence, at both the level of the individual and the population. We aimed to address this gap in our knowledge by using the well-described preference of three-spined sticklebacks (Gasterosteus aculeatus) to shoal with the larger of two shoals as model behaviour. In our experiments focal fish had the choice between two shoals of different sizes, either in clear or in turbid water containing green algae. Fish in clear water spent significantly more time near the larger shoal, while fish in algae water showed no significant preferences. Furthermore, fish tested in clear water changed more often between the shoals than fish tested in algae water. These results indicate that eutrophication-induced algae blooms have the potential to alter social decisions of sticklebacks. Such changes of social decisions do not only influence the behaviour of individuals, furthermore it might influence entire populations. This might eventually lead to changes of the structure of the social system.     

Modelling llama population development under environmental and market constraints in the Bolivian highlands

Changes in the size of animal populations over time are mainly determined by demographic and environmental factors. Livestock population dynamics are additionally influenced by harvesting decisions taken by herders. In Bolivia, not much is known about current llama husbandry and the main influencing factors determining population sizes. We collected data on demography, environmental factors and market values affecting the current and future llama population in three different regions in Bolivia. We generated a population model and assessed the future development of the llama population including environmental factors (rangeland carrying capacity, disturbance phenomena), herd structure and dynamics, and economic market demands. We calibrated and validated the llama model on the basis of 20-year data sets of the regions of Oruro, Potosi and La Paz, respectively. Model calibration by means of the Gauss-Marquardt-Levenberg algorithm yielded a model efficiency of 0.94. For model validation, however, the simulation slightly overestimated the observed llama population yielding model efficiencies of 0.91 and 0.87 for Potosi and La Paz, respectively. Model outcomes were most sensitive to death and birth rates of juveniles and death rate of females compared to environmental or other demographic factors. Population trajectories approached an overall carrying capacity for Oruro, Potosi and La Paz of 8.8 × 10⁵, 9.1 × 10⁵, and 9.0 × 10⁵ llama individuals after 100 years of simulation. Hence, detailed monitoring of demographic, environmental, and economic factors can improve predictions of llama population development over time. Further management should focus on improving birth rates and lowering female mortality through providing supplemental food and shelters against the harsh environmental conditions of the Andean highlands.     

Toxicity of the nerve agent tabun to Daphnia magna,a new experimental species in military toxicology

Daphnia magna, a freshwater microcrustacean, is currently tested as an alternative experimental species in research dealing with nerve agents poisonings treatment. Because of this, the toxicity of the nerve agent tabun (a cyanide-group containing organophosphate) to Daphnia had to be examined by estimating the EC₅₀ values. The immobilization of daphnids was chosen as the end-point. It was found that D. magna is sensitive to small amounts of tabun, even after 15 min exposure, and tabun toxicity increases with time. The estimated EC₅₀ values for 15, 30, 45, and 60 min exposure were as follows: 67.39, 38.10, 26.95, and 21.9 μg l^?1. In addition, the toxicity of media to which tabun was added 24 h before the start of experiments was examined. The results obtained indicate that daphnids can be used in experiments with nerve-agent intoxication treatment.