@article{MehnerLischkeScharnweberetal.2018, author = {Mehner, Thomas and Lischke, Betty and Scharnweber, Inga Kristin and Attermeyer, Katrin and Brothers, Soren and Gaedke, Ursula and Hilt, Sabine and Brucet, Sandra}, title = {Empirical correspondence between trophic transfer efficiency in freshwater food webs and the slope of their size spectra}, series = {Ecology : a publication of the Ecological Society of America}, volume = {99}, journal = {Ecology : a publication of the Ecological Society of America}, number = {6}, publisher = {Wiley}, address = {Hoboken}, issn = {0012-9658}, doi = {10.1002/ecy.2347}, pages = {1463 -- 1472}, year = {2018}, abstract = {The density of organisms declines with size, because larger organisms need more energy than smaller ones and energetic losses occur when larger organisms feed on smaller ones. A potential expression of density-size distributions are Normalized Biomass Size Spectra (NBSS), which plot the logarithm of biomass independent of taxonomy within bins of logarithmic organismal size, divided by the bin width. Theoretically, the NBSS slope of multi-trophic communities is exactly - 1.0 if the trophic transfer efficiency (TTE, ratio of production rates between adjacent trophic levels) is 10\% and the predator-prey mass ratio (PPMR) is fixed at 10(4). Here we provide evidence from four multi-trophic lake food webs that empirically estimated TTEs correspond to empirically estimated slopes of the respective community NBSS. Each of the NBSS considered pelagic and benthic organisms spanning size ranges from bacteria to fish, all sampled over three seasons in 1 yr. The four NBSS slopes were significantly steeper than -1.0 (range -1.14 to -1.19, with 95\% CIs excluding -1). The corresponding average TTEs were substantially lower than 10\% in each of the four food webs (range 1.0\% to 3.6\%, mean 1.85\%). The overall slope merging all biomass-size data pairs from the four systems (-1.17) was almost identical to the slope predicted from the arithmetic mean TTE of the four food webs (-1.18) assuming a constant PPMR of 10(4). Accordingly, our empirical data confirm the theoretically predicted quantitative relationship between TTE and the slope of the biomass-size distribution. Furthermore, we show that benthic and pelagic organisms can be merged into a community NBSS, but future studies have yet to explore potential differences in habitat-specific TTEs and PPMRs. We suggest that community NBSS may provide valuable information on the structure of food webs and their energetic pathways, and can result in improved accuracy of TTE-estimates.}, language = {en} } @article{LischkeMehnerHiltetal.2017, author = {Lischke, Betty and Mehner, Thomas and Hilt, Sabine and Attermeyer, Katrin and Brauns, Mario and Brothers, Soren M. and Grossart, Hans-Peter and Koehler, Jan and Scharnweber, Inga Kristin and Gaedke, Ursula}, title = {Benthic carbon is inefficiently transferred in the food webs of two eutrophic shallow lakes}, series = {Freshwater biology}, volume = {62}, journal = {Freshwater biology}, publisher = {Wiley}, address = {Hoboken}, issn = {0046-5070}, doi = {10.1111/fwb.12979}, pages = {1693 -- 1706}, year = {2017}, abstract = {The sum of benthic autotrophic and bacterial production often exceeds the sum of pelagic autotrophic and bacterial production, and hence may contribute substantially to whole-lake carbon fluxes, especially in shallow lakes. Furthermore, both benthic and pelagic autotrophic and bacterial production are highly edible and of sufficient nutritional quality for animal consumers. We thus hypothesised that pelagic and benthic transfer efficiencies (ratios of production at adjacent trophic levels) in shallow lakes should be similar. We performed whole ecosystem studies in two shallow lakes (3.5ha, mean depth 2m), one with and one without submerged macrophytes, and quantified pelagic and benthic biomass, production and transfer efficiencies for bacteria, phytoplankton, epipelon, epiphyton, macrophytes, zooplankton, macrozoobenthos and fish. We expected higher transfer efficiencies in the lake with macrophytes, because these provide shelter and food for macrozoobenthos and may thus enable a more efficient conversion of basal production to consumer production. In both lakes, the majority of the whole-lake autotrophic and bacterial production was provided by benthic organisms, but whole-lake primary consumer production mostly relied on pelagic autotrophic and bacterial production. Consequently, transfer efficiency of benthic autotrophic and bacterial production to macrozoobenthos production was an order of magnitude lower than the transfer efficiency of pelagic autotrophic and bacterial production to rotifer and crustacean production. Between-lake differences in transfer efficiencies were minor. We discuss several aspects potentially causing the unexpectedly low benthic transfer efficiencies, such as the food quality of producers, pelagic-benthic links, oxygen concentrations in the deeper lake areas and additional unaccounted consumer production by pelagic and benthic protozoa and meiobenthos at intermediate or top trophic levels. None of these processes convincingly explain the large differences between benthic and pelagic transfer efficiencies. Our data indicate that shallow eutrophic lakes, even with a major share of autotrophic and bacterial production in the benthic zone, can function as pelagic systems with respect to primary consumer production. We suggest that the benthic autotrophic production was mostly transferred to benthic bacterial production, which remained in the sediments, potentially cycling internally in a similar way to what has previously been described for the microbial loop in pelagic habitats. Understanding the energetics of whole-lake food webs, including the fate of the substantial benthic bacterial production, which is either mineralised at the sediment surface or permanently buried, has important implications for regional and global carbon cycling.}, language = {en} } @article{LischkeWeithoffWickhametal.2016, author = {Lischke, Betty and Weithoff, Guntram and Wickham, Stephen A. and Attermeyer, Katrin and Großart, Hans-Peter and Scharnweber, Inga Kristin and Hilt, Sabine and Gaedke, Ursula}, title = {Large biomass of small feeders: ciliates may dominate herbivory in eutrophic lakes}, series = {Journal of plankton research}, volume = {38}, journal = {Journal of plankton research}, publisher = {Oxford Univ. Press}, address = {Oxford}, issn = {0142-7873}, doi = {10.1093/plankt/fbv102}, pages = {2 -- 15}, year = {2016}, abstract = {The importance of ciliates as herbivores and in biogeochemical cycles is increasingly recognized. An opportunity to observe the potential consequences of zooplankton dominated by ciliates arose when winter fish kills resulted in strong suppression of crustaceans by young planktivorous fish in two shallow lakes. On an annual average, ciliates made up 38-76\% of the total zooplankton biomass in both lakes during two subsequent years. Consequently, ciliate biomass and their estimated grazing potential were extremely high compared with other lakes of various trophic states and depths. Grazing estimates based on abundance and size suggest that ciliates should have cleared the water column of small (<5 mu m) and intermediate (5-50 mu m) sized phytoplankton more than once a day. Especially, small feeders within the ciliates were important, likely exerting a strong top-down control on small phytoplankton. Particle-attached bacteria were presumably strongly suppressed by intermediate-sized ciliate feeders. In contrast to other lakes, large phytoplankton was proportionately very abundant. The phytoplankton community had a high evenness, which may be attributed to the feeding by numerous fast growing and selective ciliate species. Our study highlights ciliates as an important trophic link and adds to the growing awareness of the role of winter processes for plankton dynamics.}, language = {en} } @article{MehnerAttermeyerBraunsetal.2016, author = {Mehner, T. and Attermeyer, Katrin and Brauns, Mario and Brothers, Soren M. and Diekmann, J. and Gaedke, Ursula and Grossart, Hans-Peter and Koehler, J. and Lischke, Betty and Meyer, N. and Scharnweber, Inga Kristin and Syvaranta, J. and Vanni, M. J. and Hilt, S.}, title = {Weak Response of Animal Allochthony and Production to Enhanced Supply of Terrestrial Leaf Litter in Nutrient-Rich Lakes}, series = {Ecosystems}, volume = {19}, journal = {Ecosystems}, publisher = {Springer}, address = {New York}, issn = {1432-9840}, doi = {10.1007/s10021-015-9933-2}, pages = {311 -- 325}, year = {2016}, abstract = {Ecosystems are generally linked via fluxes of nutrients and energy across their boundaries. For example, freshwater ecosystems in temperate regions may receive significant inputs of terrestrially derived carbon via autumnal leaf litter. This terrestrial particulate organic carbon (POC) is hypothesized to subsidize animal production in lakes, but direct evidence is still lacking. We divided two small eutrophic lakes each into two sections and added isotopically distinct maize litter to the treatment sections to simulate increased terrestrial POC inputs via leaf litter in autumn. We quantified the reliance of aquatic consumers on terrestrial resources (allochthony) in the year subsequent to POC additions by applying mixing models of stable isotopes. We also estimated lake-wide carbon (C) balances to calculate the C flow to the production of the major aquatic consumer groups: benthic macroinvertebrates, crustacean zooplankton, and fish. The sum of secondary production of crustaceans and benthic macroinvertebrates supported by terrestrial POC was higher in the treatment sections of both lakes. In contrast, total secondary and tertiary production (supported by both autochthonous and allochthonous C) was higher in the reference than in the treatment sections of both lakes. Average aquatic consumer allochthony per lake section was 27-40\%, although terrestrial POC contributed less than about 10\% to total organic C supply to the lakes. The production of aquatic consumers incorporated less than 5\% of the total organic C supply in both lakes, indicating a low ecological efficiency. We suggest that the consumption of terrestrial POC by aquatic consumers facilitates a strong coupling with the terrestrial environment. However, the high autochthonous production and the large pool of autochthonous detritus in these nutrient-rich lakes make terrestrial POC quantitatively unimportant for the C flows within food webs.}, language = {en} } @article{vanGervenBredervelddeKleinetal.2015, author = {van Gerven, Luuk P. A. and Brederveld, Robert J. and de Klein, Jeroen J. M. and DeAngelis, Don L. and Downing, Andrea S. and Faber, Michiel and Gerla, Daan J. and Janse, Jan H. and Janssen, Annette B. G. and Jeuken, Michel and Kooi, Bob W. and Kuiper, Jan J. and Lischke, Betty and Liu, Sien and Petzoldt, Thomas and Schep, Sebastiaan A. and Teurlincx, Sven and Thiange, Christophe and Trolle, Dennis and van Nes, Egbert H. and Mooij, Wolf M.}, title = {Advantages of concurrent use of multiple software frameworks in water quality modelling using a database approach}, series = {Fundamental and applied limnology : official journal of the International Association of Theoretical and Applied Limnology}, volume = {186}, journal = {Fundamental and applied limnology : official journal of the International Association of Theoretical and Applied Limnology}, number = {1-2}, publisher = {Schweizerbart}, address = {Stuttgart}, issn = {1863-9135}, doi = {10.1127/fal/2015/0631}, pages = {5 -- 20}, year = {2015}, abstract = {Water quality modelling deals with multidisciplinary questions ranging from fundamental to applied. Addressing this broad range of questions requires multiple analysis techniques and therefore multiple frameworks. Through the recently developed database approach to modelling (DATM), it has become possible to run a model in multiple software frameworks without much overhead. Here we apply DATM to the ecosystem model for ditches PCDitch and its twin model for shallow lakes PCLake. Using DATM, we run these models in six frameworks (ACSL, DELWAQ, DUFLOW, GRIND for MATLAB, OSIRIS and R), and report on the possible model analyses with tools provided by each framework. We conclude that the dynamic link between frameworks and models resulting from DATM has the following main advantages: it allows one to use the framework one is familiar with for most model analyses and eases switching between frameworks for complementary model analyses, including the switch between a 0-D and 1-D to 3-D setting. Moreover, the strength of each framework - including runtime performance - can now be easily exploited. We envision that a community-based further development of the concept can contribute to the future development of water quality modelling, not only by addressing multidisciplinary questions but also by facilitating the exchange of models and process formulations within the community of water quality modellers.}, language = {en} } @article{HiltWankeScharnweberetal.2015, author = {Hilt, Sabine and Wanke, Thomas and Scharnweber, Inga Kristin and Brauns, Mario and Syvaranta, Jari and Brothers, Soren M. and Gaedke, Ursula and K{\"o}hler, Jan and Lischke, Betty and Mehner, Thomas}, title = {Contrasting response of two shallow eutrophic cold temperate lakes to a partial winterkill of fish}, series = {Hydrobiologia : acta hydrobiologica, hydrographica, limnologica et protistologica}, volume = {749}, journal = {Hydrobiologia : acta hydrobiologica, hydrographica, limnologica et protistologica}, number = {1}, publisher = {Springer}, address = {Dordrecht}, issn = {0018-8158}, doi = {10.1007/s10750-014-2143-7}, pages = {31 -- 42}, year = {2015}, abstract = {Food-web effects of winterkill are difficult to predict as the enhanced mortality of planktivorous fish may be counterbalanced by an even higher mortality of piscivores. We hypothesised that a winterkill in a clear and a turbid shallow lake would equalise their fish community composition, but seasonal plankton successions would differ between lakes. After a partial winterkill, we observed a reduction of fish biomass by 16 and 43\% in a clear-water and a turbid small temperate lake, respectively. Fish biomass and piscivore shares (5\% of fish biomass) were similar in both lakes after this winterkill, but young-of-the-year (YOY) abundances were higher in the turbid lake. Top-down control by crustaceans was only partly responsible for low phytoplankton biomass at the end of May following the winterkill in both lakes. Summer phytoplankton biomass remained low in the clear-water lake despite high abundances of YOY fish (mainly roach). In contrast, the crustacean biomass of the turbid lake was reduced in summer by a high YOY abundance (sunbleak and roach), leading to a strong increase in phytoplankton biomass. The YOY abundance of fish in shallow eutrophic lakes may thus be more important for their summer phytoplankton development after winterkill than the relative abundance of piscivores.}, language = {en} } @article{LischkeHiltJanseetal.2014, author = {Lischke, Betty and Hilt, Sabine and Janse, Jan H. and Kuiper, Jan J. and Mehner, Thomas and Mooij, Wolf M. and Gaedke, Ursula}, title = {Enhanced input of terrestrial particulate organic matter reduces the resilience of the clear-water state of shallow lakes: A model study}, series = {Ecosystems}, volume = {17}, journal = {Ecosystems}, number = {4}, publisher = {Springer}, address = {New York}, issn = {1432-9840}, doi = {10.1007/s10021-014-9747-7}, pages = {616 -- 626}, year = {2014}, abstract = {The amount of terrestrial particulate organic matter (t-POM) entering lakes is predicted to increase as a result of climate change. This may especially alter the structure and functioning of ecosystems in small, shallow lakes which can rapidly shift from a clear-water, macrophyte-dominated into a turbid, phytoplankton-dominated state. We used the integrative ecosystem model PCLake to predict how rising t-POM inputs affect the resilience of the clear-water state. PCLake links a pelagic and benthic food chain with abiotic components by a number of direct and indirect effects. We focused on three pathways (zoobenthos, zooplankton, light availability) by which elevated t-POM inputs (with and without additional nutrients) may modify the critical nutrient loading thresholds at which a clear-water lake becomes turbid and vice versa. Our model results show that (1) increased zoobenthos biomass due to the enhanced food availability results in more benthivorous fish which reduce light availability due to bioturbation, (2) zooplankton biomass does not change, but suspended t-POM reduces the consumption of autochthonous particulate organic matter which increases the turbidity, and (3) the suspended t-POM reduces the light availability for submerged macrophytes. Therefore, light availability is the key process that is indirectly or directly changed by t-POM input. This strikingly resembles the deteriorating effect of terrestrial dissolved organic matter on the light climate of lakes. In all scenarios, the resilience of the clear-water state is reduced thus making the turbid state more likely at a given nutrient loading. Therefore, our study suggests that rising t-POM input can add to the effects of climate warming making reductions in nutrient loadings even more urgent.}, language = {en} } @article{MooijBredervelddeKleinetal.2014, author = {Mooij, Wolf M. and Brederveld, Robert J. and de Klein, Jeroen J. M. and DeAngelis, Don L. and Downing, Andrea S. and Faber, Michiel and Gerla, Daan J. and Hipsey, Matthew R. and Janse, Jan H. and Janssen, Annette B. G. and Jeuken, Michel and Kooi, Bob W. and Lischke, Betty and Petzoldt, Thomas and Postma, Leo and Schep, Sebastiaan A. and Scholten, Huub and Teurlincx, Sven and Thiange, Christophe and Trolle, Dennis and van Dam, Anne A. and van Gerven, Luuk P. A. and van Nes, Egbert H. and Kuiper, Jan J.}, title = {Serving many at once: How a database approach can create unity in dynamical ecosystem modelling}, series = {Environmental modelling \& software with environment data news}, volume = {61}, journal = {Environmental modelling \& software with environment data news}, publisher = {Elsevier}, address = {Oxford}, issn = {1364-8152}, doi = {10.1016/j.envsoft.2014.04.004}, pages = {266 -- 273}, year = {2014}, abstract = {Simulation modelling in ecology is a field that is becoming increasingly compartmentalized. Here we propose a Database Approach To Modelling (DATM) to create unity in dynamical ecosystem modelling with differential equations. In this approach the storage of ecological knowledge is independent of the language and platform in which the model will be run. To create an instance of the model, the information in the database is translated and augmented with the language and platform specifics. This process is automated so that a new instance can be created each time the database is updated. We describe the approach using the simple Lotka-Volterra model and the complex ecosystem model for shallow lakes PCLake, which we automatically implement in the frameworks OSIRIS, GRIND for MATLAB, ACSL, R, DUFLOW and DELWAQ. A clear advantage of working in a database is the overview it provides. The simplicity of the approach only adds to its elegance. (C) 2014 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY-NC-SA license (http://creativecommons.org/licenses/by-nc-sa/3.0/).}, language = {en} } @article{BrothersHiltAttermeyeretal.2013, author = {Brothers, Soren M. and Hilt, Sabine and Attermeyer, Katrin and Grossart, Hans-Peter and Kosten, Sarian and Lischke, Betty and Mehner, Thomas and Meyer, Nils and Scharnweber, Inga Kristin and K{\"o}hler, Jan}, title = {A regime shift from macrophyte to phytoplankton dominance enhances carbon burial in a shallow, eutrophic lake}, series = {Ecosphere : the magazine of the International Ecology University}, volume = {4}, journal = {Ecosphere : the magazine of the International Ecology University}, number = {11}, publisher = {Wiley}, address = {Washington}, issn = {2150-8925}, doi = {10.1890/ES13-00247.1}, pages = {17}, year = {2013}, abstract = {Ecological regime shifts and carbon cycling in aquatic systems have both been subject to increasing attention in recent years, yet the direct connection between these topics has remained poorly understood. A four-fold increase in sedimentation rates was observed within the past 50 years in a shallow eutrophic lake with no surface in-or outflows. This change coincided with an ecological regime shift involving the complete loss of submerged macrophytes, leading to a more turbid, phytoplankton-dominated state. To determine whether the increase in carbon (C) burial resulted from a comprehensive transformation of C cycling pathways in parallel to this regime shift, we compared the annual C balances (mass balance and ecosystem budget) of this turbid lake to a similar nearby lake with submerged macrophytes, a higher transparency, and similar nutrient concentrations. C balances indicated that roughly 80\% of the C input was permanently buried in the turbid lake sediments, compared to 40\% in the clearer macrophyte-dominated lake. This was due to a higher measured C burial efficiency in the turbid lake, which could be explained by lower benthic C mineralization rates. These lower mineralization rates were associated with a decrease in benthic oxygen availability coinciding with the loss of submerged macrophytes. In contrast to previous assumptions that a regime shift to phytoplankton dominance decreases lake heterotrophy by boosting whole-lake primary production, our results suggest that an equivalent net metabolic shift may also result from lower C mineralization rates in a shallow, turbid lake. The widespread occurrence of such shifts may thus fundamentally alter the role of shallow lakes in the global C cycle, away from channeling terrestrial C to the atmosphere and towards burying an increasing amount of C.}, language = {en} } @phdthesis{Lischke2015, author = {Lischke, Betty}, title = {Food web regulation under different forcing regimes in shallow lakes}, url = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus4-89149}, school = {Universit{\"a}t Potsdam}, pages = {131}, year = {2015}, abstract = {The standing stock and production of organismal biomass depends strongly on the organisms' biotic environment, which arises from trophic and non-trophic interactions among them. The trophic interactions between the different groups of organisms form the food web of an ecosystem, with the autotrophic and bacterial production at the basis and potentially several levels of consumers on top of the producers. Feeding interactions can regulate communities either by severe grazing pressure or by shortage of resources or prey production, termed top-down and bottom-up control, respectively. The limitations of all communities conglomerate in the food web regulation, which is subject to abiotic and biotic forcing regimes arising from external and internal constraints. This dissertation presents the effects of alterations in two abiotic, external forcing regimes, terrestrial matter input and long-lasting low temperatures in winter. Diverse methodological approaches, a complex ecosystem model study and the analysis of two whole-lake measurements, were performed to investigate effects for the food web regulation and the resulting consequences at the species, community and ecosystem scale. Thus, all types of organisms, autotrophs and heterotrophs, at all trophic levels were investigated to gain a comprehensive overview of the effects of the two mentioned altered forcing regimes. In addition, an extensive evaluation of the trophic interactions and resulting carbon fluxes along the pelagic and benthic food web was performed to display the efficiencies of the trophic energy transfer within the food webs. All studies were conducted in shallow lakes, which is worldwide the most abundant type of lakes. The specific morphology of shallow lakes allows that the benthic production contributes substantially to the whole-lake production. Further, as shallow lakes are often small they are especially sensitive to both, changes in the input of terrestrial organic matter and the atmospheric temperature. Another characteristic of shallow lakes is their appearance in alternative stable states. They are either in a clear-water or turbid state, where macrophytes and phytoplankton dominate, respectively. Both states can stabilize themselves through various mechanisms. These two alternative states and stabilizing mechanisms are integrated in the complex ecosystem model PCLake, which was used to investigate the effects of the enhanced terrestrial particulate organic matter (t-POM) input to lakes. The food web regulation was altered by three distinct pathways: (1) Zoobenthos received more food, increased in biomass which favored benthivorous fish and those reduced the available light due to bioturbation. (2) Zooplankton substituted autochthonous organic matter in their diet by suspended t-POM, thus the autochthonous organic matter remaining in the water reduced its transparency. (3) T-POM suspended into the water and reduced directly the available light. As macrophytes are more light-sensitive than phytoplankton they suffered the most from the lower transparency. Consequently, the resilience of the clear-water state was reduced by enhanced t-POM inputs, which makes the turbid state more likely at a given nutrient concentration. In two subsequent winters long-lasting low temperatures and a concurrent long duration of ice coverage was observed which resulted in low overall adult fish biomasses in the two study lakes - Schulzensee and Gollinsee, characterized by having and not having submerged macrophytes, respectively. Before the partial winterkill of fish Schulzensee allowed for a higher proportion of piscivorous fish than Gollinsee. However, the partial winterkill of fish aligned both communities as piscivorous fish are more sensitive to low oxygen concentrations. Young of the year fish benefitted extremely from the absence of adult fish due to lower predation pressure. Therefore, they could exert a strong top-down control on crustaceans, which restructured the entire zooplankton community leading to low crustacean biomasses and a community composition characterized by copepodites and nauplii. As a result, ciliates were released from top-down control, increased to high biomasses compared to lakes of various trophic states and depths and dominated the zooplankton community. While being very abundant in the study lakes and having the highest weight specific grazing rates among the zooplankton, ciliates exerted potentially a strong top-down control on small phytoplankton and particle-attached bacteria. This resulted in a higher proportion of large phytoplankton compared to other lakes. Additionally, the phytoplankton community was evenly distributed presumably due to the numerous fast growing and highly specific ciliate grazers. Although, the pelagic food web was completely restructured after the subsequent partial winterkills of fish, both lakes were resistant to effects of this forcing regime at the ecosystem scale. The consistently high predation pressure on phytoplankton prevented that Schulzensee switched from the clear-water to the turbid state. Further mechanisms, which potentially stabilized the clear-water state, were allelopathic effects by macrophytes and nutrient limitation in summer. The pelagic autotrophic and bacterial production was an order of magnitude more efficient transferred to animal consumers than the respective benthic production, despite the alterations of the food web structure after the partial winterkill of fish. Thus, the compiled mass-balanced whole-lake food webs suggested that the benthic bacterial and autotrophic production, which exceeded those of the pelagic habitat, was not used by animal consumers. This holds even true if the food quality, additional consumers such as ciliates, benthic protozoa and meiobenthos, the pelagic-benthic link and the potential oxygen limitation of macrobenthos were considered. Therefore, low benthic efficiencies suggest that lakes are primarily pelagic systems at least at the animal consumer level. Overall, this dissertation gives insights into the regulation of organism groups in the pelagic and benthic habitat at each trophic level under two different forcing regimes and displays the efficiency of the carbon transfer in both habitats. The results underline that the alterations of external forcing regimes affect all hierarchical level including the ecosystem.}, language = {en} }