Recently, the importance of body mass and allometric scaling for the structure and dynamics of ecological networks has been highlighted in several ground-breaking studies. However, advances in the understanding of generalities across ecosystem types are impeded to a considerable extent by a methodological dichotomy contrasting a considerable portion of marine ecology on the one hand opposite to traditional community ecology on the other hand. Many marine ecologists are bound to the taxonomy-neglecting size spectrum approach when describing and analysing community patterns. In contrast, the mindset of the other school is focused on taxonomies according to the Linnean system at the cost of obscuring information due to applying species or population averages of body masses and other traits. Following other pioneering studies, we addressed this lingering gap, and studied non-linear interaction strengths (i.e. functional responses) between two taxonomically-distinct terrestrial arthropod predators (centipedes and spiders) of varying individual body masses and their prey. We fitted three non-linear functional response models to the data: (1) a taxonomic model not accounting for variance in body masses amongst predator individuals, (2) an allometric model ignoring taxonomic differences between predator individuals, and (3) a combined model including body mass and taxonomic effects. Ranked according to their AICs, the combined model performs better than the allometric model, which provides a superior fit to the data than the taxonomic model. These results strongly indicate that the body masses of predator and prey individuals were responsible for most of the variation in non-linear interaction strengths. Taxonomy explained some specific patterns in allometric exponents between groups and revealed mechanistic insights in predation efficiencies. Reconciling quantitative allometric models as employed by the marine size-spectrum approach with taxonomic information may thus yield quantitative results that are generalized across ecosystem types and taxonomic groups. Using these quantitative models as novel null models should also strengthen subsequent taxonomic analyses.
Macrofauna invertebrates of forest floors provide important functions in the decomposition process of soil organic matter, which is affected by the nutrient stoichiometry of the leaf litter. Climate change effects on forest ecosystems include warming and decreasing litter quality (e.g. higher C : nutrient ratios) induced by higher atmospheric CO 2 concentrations. While litter-bag experiments unravelled separate effects, a mechanistic understanding of how interactions between temperature and litter stoichiometry are driving decomposition rates is lacking. In a laboratory experiment, we filled this void by quantifying decomposer consumption rates analogous to predator-prey functional responses that include the mechanistic parameters handling time and attack rate. Systematically, we varied the body masses of isopods, the environmental temperature and the resource between poor (hornbeam) and good quality (ash). We found that attack rates increased and handling times decreased (i) with body masses and (ii) temperature. Interestingly, these relationships interacted with litter quality: small isopods possibly avoided the poorer resource, whereas large isopods exhibited increased, compensatory feeding of the poorer resource, which may be explained by their higher metabolic demands. The combination of metabolic theory and ecological stoichiometry provided critically important mechanistic insights into how warming and varying litter quality may modify macrofaunal decomposition rates.
Ecological communities consist of small abundant and large non-abundant species. The energetic equivalence rule is an often-observed pattern that could be explained by equal energy usage among abundant small organisms and non-abundant large organisms. To generate this pattern, metabolism (as an indicator of individual energy use) and abundance have to scale inversely with body mass, and cancel each other out. In contrast, the pattern referred to as biomass equivalence states that the biomass of all species in an area should be constant across the body-mass range. In this study, we investigated forest soil communities with respect to metabolism, abundance, population energy use, and biomass. We focused on four land-use types in three different landscape blocks (Biodiversity Exploratories). The soil samples contained 870 species across 12 phylogenetic groups. Our results indicated positive sublinear metabolic scaling and negative sublinear abundance scaling with species body mass. The relationships varied mainly due to differences among phylogenetic groups or feeding types, and only marginally due to land-use type. However, these scaling relationships were not exactly inverse to each other, resulting in increasing population energy use and biomass with increasing body mass for most combinations of phylogenetic group or feeding type with land-use type. Thus, our results are mostly inconsistent with the classic perception of energetic equivalence, and reject the biomass equivalence hypothesis while documenting a specific and nonrandom pattern of how abundance, energy use, and biomass are distributed across size classes. However, these patterns are consistent with two alternative predictions: the resource-thinning hypothesis, which states that abundance decreases with trophic level, and the allometric degree hypothesis, which states that population energy use should increase with population average body mass, due to correlations with the number of links of consumers and resources. Overall, our results suggest that a synthesis of food web structures with metabolic theory may be most promising for predicting natural patterns of abundance, biomass, and energy use.
To maintain constant chemical composition, i.e. elemental homeostasis, organisms have to consume resources of sufficient quality to meet their own specific stoichiometric demand. Therefore, concentrations of elements indicate resource quality, and rare elements in the environment may act as limiting factors for individual organisms scaling up to constrain population densities. We investigated how the biomass densities of invertebrate populations of temperate forest soil communities depend on 1) the stoichiometry of the basal litter according to ecological stoichiometry concepts and 2) the population average body mass as predicted by metabolic theory. We used a large data set on biomass densities of 4959 populations across 48 forests in three regions of Germany. Following various ecological stoichiometry hypotheses, we tested for effects of the carbon‐to‐element ratios of 10 elements. Additionally, we included the abiotic litter characteristics habitat size (represented by litter depth), litter diversity and pH, as well as forest type as an indicator for human management. Across 12 species groups, we found that the biomass densities scaled significantly with population‐averaged body masses thus supporting metabolic theory. Additionally, 10 of these allometric scaling relationships exhibited interactions with stoichiometric and abiotic co‐variables. The four most frequent co‐variables were 1) forest type, 2) the carbon‐to‐phosphorus ratio (C:P), 3) the carbon‐to‐sodium ratio (C:Na), and the carbon‐to‐nitrogen ratio (C:N). Hence, our analyses support the sodium shortage hypothesis for microbi‐detritivores, the structural elements hypothesis for some predator groups (concerning N), and the secondary productivity hypothesis (concerning P) across all trophic groups in our data. In contrast, the ecosystem size hypothesis was only supported for some meso‐ and macrofauna detritivores. Our study is thus providing a comprehensive analysis how the elemental stoichiometry of the litter as the basal resource constrain population densities across multiple trophic levels of soil communities.
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