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Allometric trophic network (ATN) models offer high flexibility and scalability while minimizing the number of parameters and have been successfully applied to investigate complex food web dynamics and their influence on food web diversity and stability. However, the realism of ATN model energetics has never been assessed in detail, despite their critical influence on dynamic biomass and production patterns. Here, we compare the energetics of the currently established original ATN model, considering only biomass-dependent basal respiration, to an extended ATN model version, considering both basal and assimilation-dependent activity respiration. The latter is crucial in particular for unicellular and invertebrate organisms which dominate the metabolism of pelagic and soil food webs. Based on metabolic scaling laws, we show that the extended ATN version reflects the energy transfer through a chain of four trophic levels of unicellular and invertebrate organisms more realistically than the original ATN version. Depending on the strength of top-down control, the original ATN model yields trophic transfer efficiencies up to 71% at either the third or the fourth trophic level, which considerably exceeds any realistic values. In contrast, the extended ATN version yields realistic trophic transfer efficiencies 30% at all trophic levels, in accordance with both physiological considerations and empirical evidence from pelagic systems. Our results imply that accounting for activity respiration is essential for consistently implementing the metabolic theory of ecology in ATN models and for improving their quantitative predictions, which makes them more powerful tools for investigating the dynamics of complex natural communities.
It is of major interest to estimate the feedback of arctic ecosystems to the global warming we expect in upcoming decades. The speed of this response is driven by the potential of species to migrate, tracking their climate optimum. For this, sessile plants have to produce and disperse seeds to newly available habitats, and pollination of ovules is needed for the seeds to be viable. These two processes are also the vectors that pass genetic information through a population. A restricted exchange among subpopulations might lead to a maladapted population due to diversity losses. Hence, a realistic implementation of these dispersal processes into a simulation model would allow an assessment of the importance of diversity for the migration of plant species in various environments worldwide. To date, dynamic global vegetation models have been optimized for a global application and overestimate the migration of biome shifts in currently warming temperatures. We hypothesize that this is caused by neglecting important fine-scale processes, which are necessary to estimate realistic vegetation trajectories. Recently, we built and parameterized a simulation model LAVESI for larches that dominate the latitudinal treelines in the northernmost areas of Siberia. In this study, we updated the vegetation model by including seed and pollen dispersal driven by wind speed and direction. The seed dispersal is modelled as a ballistic flight, and for the pollination of ovules of seeds produced, we implemented a wind-determined and distance-dependent probability distribution function using a von Mises distribution to select the pollen donor. A local sensitivity analysis of both processes supported the robustness of the model's results to the parameterization, although it highlighted the importance of recruitment and seed dispersal traits for migration rates. This individual-based and spatially explicit implementation of both dispersal processes makes it easily feasible to inherit plant traits and genetic information to assess the impact of migration processes on the genetics. Finally, we suggest how the final model can be applied to substantially help in unveiling the important drivers of migration dynamics and, with this, guide the improvement of recent global vegetation models.
Changes in species’ distributions are classically projected based on their climate envelopes. For Siberian forests, which have a tremendous significance for vegetation-climate feedbacks, this implies future shifts of each of the forest-forming larch (Larix) species to the north-east. However, in addition to abiotic factors, reliable projections must assess the role of historical biogeography and biotic interactions. Here, we use sedimentary ancient DNA and individual-based modelling to investigate the distribution of larch species and mitochondrial haplotypes through space and time across the treeline ecotone on the southern Taymyr peninsula, which at the same time presents a boundary area of two larch species. We find spatial and temporal patterns, which suggest that forest density is the most influential driver determining the precise distribution of species and mitochondrial haplotypes. This suggests a strong influence of competition on the species’ range shifts. These findings imply possible climate change outcomes that are directly opposed to projections based purely on climate envelopes. Investigations of such fine-scale processes of biodiversity change through time are possible using paleoenvironmental DNA, which is available much more readily than visible fossils and can provide information at a level of resolution that is not reached in classical palaeoecology.
It is of major interest to estimate the feedback of arctic ecosystems to the global warming we expect in upcoming decades. The speed of this response is driven by the potential of species to migrate, tracking their climate optimum. For this, sessile plants have to produce and disperse seeds to newly available habitats, and pollination of ovules is needed for the seeds to be viable. These two processes are also the vectors that pass genetic information through a population. A restricted exchange among subpopulations might lead to a maladapted population due to diversity losses. Hence, a realistic implementation of these dispersal processes into a simulation model would allow an assessment of the importance of diversity for the migration of plant species in various environments worldwide. To date, dynamic global vegetation models have been optimized for a global application and overestimate the migration of biome shifts in currently warming temperatures. We hypothesize that this is caused by neglecting important fine-scale processes, which are necessary to estimate realistic vegetation trajectories. Recently, we built and parameterized a simulation model LAVESI for larches that dominate the latitudinal treelines in the northernmost areas of Siberia. In this study, we updated the vegetation model by including seed and pollen dispersal driven by wind speed and direction. The seed dispersal is modelled as a ballistic flight, and for the pollination of ovules of seeds produced, we implemented a wind-determined and distance-dependent probability distribution function using a von Mises distribution to select the pollen donor. A local sensitivity analysis of both processes supported the robustness of the model's results to the parameterization, although it highlighted the importance of recruitment and seed dispersal traits for migration rates. This individual-based and spatially explicit implementation of both dispersal processes makes it easily feasible to inherit plant traits and genetic information to assess the impact of migration processes on the genetics. Finally, we suggest how the final model can be applied to substantially help in unveiling the important drivers of migration dynamics and, with this, guide the improvement of recent global vegetation models.
Changes in species' distributions are classically projected based on their climate envelopes. For Siberian forests, which have a tremendous significance for vegetation-climate feedbacks, this implies future shifts of each of the forest-forming larch (Larix) species to the north-east. However, in addition to abiotic factors, reliable projections must assess the role of historical biogeography and biotic interactions. Here, we use sedimentary ancient DNA and individual-based modelling to investigate the distribution of larch species and mitochondrial haplotypes through space and time across the treeline ecotone on the southern Taymyr peninsula, which at the same time presents a boundary area of two larch species. We find spatial and temporal patterns, which suggest that forest density is the most influential driver determining the precise distribution of species and mitochondrial haplotypes. This suggests a strong influence of competition on the species' range shifts. These findings imply possible climate change outcomes that are directly opposed to projections based purely on climate envelopes. Investigations of such fine-scale processes of biodiversity change through time are possible using paleoenvironmental DNA, which is available much more readily than visible fossils and can provide information at a level of resolution that is not reached in classical palaeoecology.