title: Energy storage and fecundity explain deviations from ecological stoichiometry predictions under global warming and size-selective predation.
abstract: A key challenge for ecologists is to predict how single and joint effects of global warming and predation risk translate from the individual level up to ecosystem functions. Recently, stoichiometric theory linked these levels through changes in body stoichiometry, predicting that both higher temperatures and predation risk induce shifts in energy storage (increases in C-rich carbohydrates and reductions in N-rich proteins) and body stoichiometry (increases in C : N and C : P). This promising theory, however, is rarely tested and assumes that prey will divert energy away from reproduction under predation risk, while under size-selective predation, prey instead increase fecundity. We exposed the water flea Daphnia magna to 4 °C warming and fish predation risk to test whether C-rich carbohydrates increase and N-rich proteins decrease, and as a result, C : N and C : P increase under warming and predation risk. Unexpectedly, warming decreased body C : N, which was driven by reductions in C-rich fat and sugar contents while the protein content did not change. This reflected a trade-off where the accelerated intrinsic growth rate under warming occurred at the cost of a reduced energy storage. Warming reduced C : N less and only increased C : P and N : P in the fish-period Daphnia. These evolved stoichiometric responses to warming were largely driven by stronger warming-induced reductions in P than in C and N and could be explained by the better ability to deal with warming in the fish-period Daphnia. In contrast to theory predictions, body C : N decreased under predation risk due to a strong increase in the N-rich protein content that offsets the increase in C-rich fat content. The higher investment in fecundity (more N-rich eggs) under predation risk contributed to this stronger increase in protein content. Similarly, the lower body C : N of pre-fish Daphnia also matched their higher fecundity. Warming and predation risk independently shaped body stoichiometry, largely by changing levels of energy storage molecules. Our results highlight that two widespread patterns, the trade-off between rapid development and energy storage and the increased investment in reproduction under size-selective predation, cause predictable deviations from current ecological stoichiometry theory.
authors.affiliation: Laboratory of Aquatic Ecology, Evolution and Conservation, KU Leuven, Deberiotstraat 32, B-3000, Leuven, Belgium. chao.zhang@bio.kuleuven.be.
authors.forename: Chao
authors.initials: C
authors.lastname: Zhang
authors.affiliation: Laboratory of Aquatic Ecology, Evolution and Conservation, KU Leuven, Deberiotstraat 32, B-3000, Leuven, Belgium.
authors.forename: Mieke
authors.initials: M
authors.lastname: Jansen
authors.affiliation: Laboratory of Aquatic Ecology, Evolution and Conservation, KU Leuven, Deberiotstraat 32, B-3000, Leuven, Belgium.
authors.forename: Luc
authors.initials: L
authors.lastname: De Meester
authors.affiliation: Laboratory of Aquatic Ecology, Evolution and Conservation, KU Leuven, Deberiotstraat 32, B-3000, Leuven, Belgium.
authors.forename: Robby
authors.initials: R
authors.lastname: Stoks
mesh_terms: D000818:Animals; D060829:Arthropod Proteins / Q000378:metabolism; D049628:Body Size; D050260:Carbohydrate Metabolism; D003621:Daphnia / Q000502:physiology*; D004734:Energy Metabolism; D005298:Fertility; D005399:Fishes / Q000502:physiology*; D020387:Food Chain; D057232:Global Warming; D011235:Predatory Behavior
chemical_list: D060829:Arthropod Proteins
keywords: 
  Daphnia magna
  ; RNA : DNA; general stress paradigm; global climate change; multiple stressors; non-consumptive predator effects; rapid evolution; stress ecology