Wolf M. Mooij

The impact of climate change on lakes in the Netherlands: a review

Climate change will alter freshwater ecosystems but specific effects will vary among regions and the type of water body. Here, we give an integrative review of the observed and predicted impacts of climate change on shallow lakes in the Netherlands and put these impacts in an international perspective. Most of these lakes are man-made and have preset water levels and poorly developed littoral zones. Relevant climatic factors for these ecosystems are temperature, ice-cover and wind. Secondary factors affected by climate include nutrient loading, residence time and water levels. We reviewed the relevant literature in order to assess the impact of climate change on these lakes. We focussed on six management objectives as bioindicators for the functioning of these ecosystems: target species, nuisance species, invading species, transparency, carrying capacity and biodiversity. We conclude that climate change will likely (i) reduce the numbers of several target species of birds; (ii) favour and stabilize cyanobacterial dominance in phytoplankton communities; (iii) cause more serious incidents of botulism among waterfowl and enhance the spreading of mosquito borne diseases; (iv) benefit invaders originating from the Ponto-Caspian region; (v) stabilize turbid, phytoplankton-dominated systems, thus counteracting restoration measures; (vi) destabilize macrophyte-dominated clear-water lakes; (vii) increase the carrying capacity of primary producers, especially phytoplankton, thus mimicking eutrophication; (viii) affect higher trophic levels as a result of enhanced primary production; (ix) have a negative impact on biodiversity which is linked to the clear water state; (x) affect biodiversity by changing the disturbance regime. Water managers can counteract these developments by reduction of nutrient loading, development of the littoral zone, compartmentalization of lakes and fisheries management.

Experimental Evidence for Spatial Self-Organization and Its Emergent Effects in Mussel Bed Ecosystems

Spatial self-organization is the main theoretical explanation for the global occurrence of regular or otherwise coherent spatial patterns in ecosystems. Using mussel beds as a model ecosystem, we provide an experimental demonstration of spatial self-organization. Under homogeneous laboratory conditions, mussels developed regular patterns, similar to those in the field. An individual-based model derived from our experiments showed that interactions between individuals explained the observed patterns. Furthermore, a field study showed that pattern formation affected ecosystem-level processes in terms of improved growth and resistance to wave action. Our results imply that spatial self-organization is an important determinant of the structure and functioning of ecosystems, and it needs to be considered in their conservation.

Challenges and Opportunities for Integrating Lake Ecosystem Modelling Approaches

A large number and wide variety of lake ecosystem models have been developed and published during the past four decades. We identify two challenges for making further progress in this field. One such challenge is to avoid developing more models largely following the concept of others (‘reinventing the wheel’). The other challenge is to avoid focusing on only one type of model, while ignoring new and diverse approaches that have become available (‘having tunnel vision’). In this paper, we aim at improving the awareness of existing models and knowledge of concurrent approaches in lake ecosystem modelling, without covering all possible model tools and avenues. First, we present a broad variety of modelling approaches. To illustrate these approaches, we give brief descriptions of rather arbitrarily selected sets of specific models. We deal with static models (steady state and regression models), complex dynamic models (CAEDYM, CE-QUAL-W2, Delft 3D-ECO, LakeMab, LakeWeb, MyLake, PCLake, PROTECH, SALMO), structurally dynamic models and minimal dynamic models. We also discuss a group of approaches that could all be classified as individual based: super-individual models (Piscator, Charisma), physiologically structured models, stage-structured models and trait-based models. We briefly mention genetic algorithms, neural networks, Kalman filters and fuzzy logic. Thereafter, we zoom in, as an in-depth example, on the multi-decadal development and application of the lake ecosystem model PCLake and related models (PCLake Metamodel, Lake Shira Model, IPH-TRIM3D-PCLake). In the discussion, we argue that while the historical development of each approach and model is understandable given its ‘leading principle’, there are many opportunities for combining approaches. We take the point of view that a single ‘right’ approach does not exist and should not be strived for. Instead, multiple modelling approaches, applied concurrently to a given problem, can help develop an integrative view on the functioning of lake ecosystems. We end with a set of specific recommendations that may be of help in the further development of lake ecosystem models.

INDUCIBLE DEFENSES AND TROPHIC STRUCTURE

Resource edibility is a crucial factor in ecological theory on the relative importance of bottom-up and top-down control. Current theory explains trophic structure in terms of the relative abundance and succession of edible and inedible species across gradients of primary productivity. We argue that this explanation is incomplete owing to its focus on inedibility and the assumption that plants and herbivores have fixed defense levels. Consumer-induced defenses are an important source of variation in the vulnerability of prey and are prevalent in natural communities. Such induced defenses decrease per capita consumption rates of consumers but hardly ever result in complete inedibility. When defenses are inducible a prey population may consist of both undefended and defended individuals. Here we use food chain models with realistic parameter values to show that variation in consumption rates on different prey types causes a gradual instead of stepwise increase in the biomass of all trophic levels in response to enrichment. Such all-level responses have been observed in both aquatic and terrestrial ecosystems and in microbial food chains in the laboratory. We stress that, in addition to the known food web effects of interspecific variation in edibility, intraspecific variation in edibility is another form of within-trophic-level heterogeneity that also has such effects. We conclude that inducible defenses increase the relative importance of bottom-up control.

Seasonal patterns in the mortality of Daphnia species in a shallow lake

To assess the impact of predation by young-of-the-year (0+) fish on the population dynamics of Daphniaspecies, we made independent estimations of the mortality of Daphniaspecies during the year, and of the predation pressure exerted by the juvenile fish. Mortality of daphnids was computed using a model that allowed us to differentiate between different size-classes, while total fish consumption was estimated from the temperature-dependent daily weight increase and the population development of the 0+ fish. The predation pressure on the different size-classes of Daphniaspecies was estimated by combining the total fish consumption with estimates of the selective feeding behaviour of the fish. To make the estimates of fish consumption independent of our current (1989–1991) zooplankton data set, we estimated fish species-specific and fish length-dependent selectivity indices on zooplankton using a different data set (1976–1977).Daphniapopulation densities usually increased in spring and decreased rapidly in early summer. Predation by 0+ fish was not severe enough to explain the large mortality that caused the summer decline; later in the year most of the mortality in the larger size-classes of the daphnids (>1.0 mm) could be explained by fish predation. Resumé: Pour mesurer l’impact de la prédation par les jeunes poissons de l’année (0+) sur la dynamique des populations des espèces de Daphnia, nous avons fait des estimations indépendantes de la mortalité des espèces de Daphniapendant l’année, et de la pression de prédation exercée par les jeunes poissons. Nous avons calculé la mortalité des daphnies à l’aide d’un modèle qui nous permettait de différencier les classes de taille, tandis que la consommation totale des poissons était estimée à partir de l’augmentation de poids quotidienne dépendant de la température et du développement de la population de poissons 0+. Nous avons estimé la pression de prédation sur les différentes classes de taille de Daphniaen combinant la consommation totale des poissons à des estimations du comportement d’alimentation sélective des poissons. Pour rendre les estimations de la consommation des poissons indépendantes de notre série présente (1989–1991) de données sur le zooplancton, nous avons calculé les indices de sélectivité à l’égard du zooplancton qui sont propres à l’espèce de poisson et dépendants de la longueur des poissons, en nous servant d’une série différente de données (1976–1977). Les densités de la population de Daphnia augmentaient généralement au printemps et baissaient rapidement au début de l’été. La prédation par les poissons 0+ n’était pas assez forte pour expliquer l’importante mortalité qui causait le déclin estival; pendant le reste de l’année, la majeure partie de la mortalité chez les plus grandes classes de taille des daphnies (>1,0 mm) pouvait s’expliquer par la prédation des poissons. [Traduit par la Rédaction]

Climate-induced shifts in an experimental phytoplankton community: a mechanistic approach

Climate change is likely to have far-reaching effects on biotic interactions in aquatic ecosystems. We investigated the effect of different spring warming scenarios on the succession of three algal groups (cyanobacteria, diatoms and green algae) in 10-l microcosms. We fitted these microcosm data to a simple mechanistic model to estimate the effect of different climate warming scenarios on the population dynamics of these algal functional groups. Experimental and model results indicate that the different algal functional groups respond differently to climate warming under phosphorus-limited conditions. Whereas the successional sequence, from diatoms to green algae to cyanobacteria, was not affected by the different climate warming scenarios, cyanobacteria showed a stronger response to the different climate warming scenarios than diatoms or green algae. Both the growth rates and peak abundances of cyanobacteria were significantly higher in the average and warm spring scenarios than in the cold spring scenario. Our findings illustrate that integration of models and microcosm experiments are a useful approach in predicting the impacts of rising temperatures on the dynamics of phytoplankton communities.

Allelopathic growth inhibition and colony formation of the green alga Scenedesmus obliquus by the aquatic macrophyte Stratiotes aloides

Laboratory experiments were conducted to elucidate the allelopathic effects of exudates from the aquatic macrophyte Stratiotes aloides on the growth and morphology of the green alga Scenedesmus obliquus. Both water originating from a S. aloides culture and water that had not been in contact with S. aloides was simultaneously inoculated with S. obliquus cells. In all experiments we observed a highly significant inhibitory effect of Stratiotes water on the green alga. The duration of the lag phase as well as the initial biovolume doubling time of S. obliquus were extended. The growth rate of S. obliquus was only significantly inhibited by the younger Stratiotes plants. Furthermore, the mean particle volume (MPV) of Scenedesmus increased significantly in the presence of Stratiotes water in all three experiments. Microscopic analyses confirmed that S. obliquus forms more colonies in the presence of water from a S. aloides culture. Colonies of phytoplankton have higher sinking rates than single phytoplankton cells of the same species and disappear faster from the upper water layers, reducing the competition for light between algae and macrophytes.

A community-based framework for aquatic ecosystem models

Here, we communicate a point of departure in the development of aquatic ecosystem models, namely a new community-based framework, which supports an enhanced and transparent union between the collective expertise that exists in the communities of traditional ecologists and model developers. Through a literature survey, we document the growing importance of numerical aquatic ecosystem models while also noting the difficulties, up until now, of the aquatic scientific community to make significant advances in these models during the past two decades. Through a common forum for aquatic ecosystem modellers we aim to (i) advance collaboration within the aquatic ecosystem modelling community, (ii) enable increased use of models for research, policy and ecosystem-based management, (iii) facilitate a collective framework using common (standardised) code to ensure that model development is incremental, (iv) increase the transparency of model structure, assumptions and techniques, (v) achieve a greater understanding of aquatic ecosystem functioning, (vi) increase the reliability of predictions by aquatic ecosystem models, (vii) stimulate model inter-comparisons including differing model approaches, and (viii) avoid ‘re-inventing the wheel’, thus accelerating improvements to aquatic ecosystem models. We intend to achieve this as a community that fosters interactions amongst ecologists and model developers. Further, we outline scientific topics recently articulated by the scientific community, which lend themselves well to being addressed by integrative modelling approaches and serve to motivate the progress and implementation of an open source model framework.

FUZZY MODELING OF CYANOBACTERIAL SURFACE WATERBLOOMS: VALIDATION WITH NOAA‐AVHRR SATELLITE IMAGES

Surface waterblooms of toxic cyanobacteria (scums) interfere with the use of lakes, for instance in the production of drinking water or for recreation. Routine monitoring data are not sufficient for early warning due to the large temporal and spatial variability in the occurrence of surface waterblooms, and the time lag between the formation of the scum and the availability of relevant information for risk management. We combined a “traditional” dynamic simulation model based upon differential equations with fuzzy logic to describe the three main conditions governing surface waterbloom formation: (1) a preexisting population of cyanobacteria, (2) buoyancy of the cells, and (3) stability of the water column. The attributes and membership functions of the fuzzy model were based on earlier field studies of diel changes in buoyancy and vertical distribution of cyanobacteria. The model was applied without further calibration to the large lake IJsselmeer (1200 km2) in the Netherlands, and we validated the model...

Trade-offs in Daphnia habitat selection

Diel vertical migration (DVM) is a dynamic behavioral pattern found ex- tensively in the worlds oceans and lakes, yet the role of food and temperature distribution on DVM is still unclear. While DVM has been mostly studied in systems with surface food maxima, deep-water food maxima are quite common in lakes and oceans. In such ecosys- tems, optimal conditions of temperature and food are uncoupled. In a Swiss high-mountain lake (Oberer Arosasee) with a deep-water food maximum, we found that Daphnia galeata adults and juveniles exhibit DVM behavior almost throughout the year and migrated up- wards, out of food-rich environments, at night. In a large indoor mesocosm experiment, we were able to show that Daphnia respond to fish-mediated cues by migrating into deeper water layers. In the presence of fish and using natural vertical food and temperature dis- tributions in the mesocosms, we could reproduce the vertical distribution of adult Daphnia observed in the field. The indoor experiments show that food and temperature modulate the actual depth at which the animals stop migrating, whereas fish and ultraviolet radiation likely determine the timing of migration (its synchronization with dawn and dusk). Overall, our results show that the nighttime movement into the surface waters contributes to the fitness ofDaphnia, given the costs and benefits associated with the trade-off between food and temperature.