One of the central tenets of ecological stoichiometry is that consumer growth rate is strongly determined by food phosphorus (P) content. In planktonic organisms population growth rates of zooplankton have repeatedly been shown to be reduced when fed with P-limited algal food sources. However, P-limitation may also affect other quality-related aspects of algae, such as biochemical composition or palatability. We studied the population growth, detailed life history and body elemental composition of the herbivorous rotifer, Brachionus calyciflorus, in response to three different food quality treatments: algae cultured in high phosphorus conditions (average algal molar C:P ≈ 112, ‘HP’), algae cultured in low P conditions (molar C:P ≈ 631, ‘LP’) and low-P cultured algae spiked with P just before feeding (molar C:P ≈ 113, ‘LP+P’). LP+P algae thus combined high P content with a history of growth under P-limited conditions. Total P content and the C:P ratio of rotifers in the LP+P treatment equaled those of rotifers in the HP treatment. Rotifer population growth rates were higher in HP than in LP and intermediate in the LP+P treatment. Similarly, many life history traits observed for animals in the LP+P treatment, such as somatic growth rate, age at maturity, and egg production rate were also intermediate to those observed in the LP and HP treatments. However, there were important deviations from this pattern: size at first reproduction and egg mortality in the LP+P treatment equaled the HP treatment, whereas size and development time of the first eggs equaled those of the LP treatment. Our results indicate that elemental limitation cannot fully explain reduced performance of consumers fed with P-limited algae and strongly suggest that indirect, non-stoichiometric effects of P-limitation, e.g., via changes in biochemical composition or morphology of the algae also play a major role. Furthermore, our study highlights that such indirect effects have a differential impact on major fitness components and may as such also determine the population dynamics and demographic structure of consumer populations.
Anthropogenic activities have reshaped the relative supply rates of essential elements to organisms. Recent studies suggested that consumer performance is strongly reduced by food that is either very high or very low in relative phosphorus content. However, the generality of such ‘stoichiometric knife‐edge’ and its underlying mechanisms are poorly understood. We studied the response of a planktonic rotifer to a 10‐fold food carbon : phosphorus (C : P) gradient and confirmed the existence of the stoichiometric knife‐edge. Interestingly, we observed a complete homeostatic breakdown associated with strong growth reductions at high food C : P. In contrast, at low food C : P, animals maintained homeostasis despite pronounced performance reductions. Our results suggest that the mechanisms underlying adverse effects of stoichiometric imbalance are determined by both the identity of elements that are limiting and those that are present in excess. Negative effects of excess P reveal an additional way of how eutrophication may affect consumers.
Brachionus calyciflorus is arguably the most studied freshwater monogonont rotifer. Although it has been recognised as a cryptic species complex for more than a decade, a formal (re‐)description of the four species known so far (B. calyciflorus, Brachionus dorcas, Brachionus elevatus, and Brachionus fernandoi) has only recently been made. Information on the ecology of these species is very scant and fragmented. The aim of this study was to test for ecological divergence between these four species, specifically their life history strategy and population demography. We conducted a life history experiment using 12–16 genotypes per species. For each species, genotypes were extracted from at least three different natural populations. In addition, we performed population‐level culture experiments with the aim to compare population growth rates and demographic structure of experimental populations among species. Finally, we searched the literature for life history studies with molecular data allowing retrospective species identification. We found pronounced differences in life history traits between B. fernandoi and the other three species. B. fernandoi had higher egg and juvenile development times and a lower egg production rate and mictic ratio. We detected no significant life history differences among B. calyciflorus, B. elevatus, and B. dorcas. Population growth rates of B. fernandoi and B. calyciflorus were higher than those of B. elevatus and B. dorcas. Life history divergence resulted in marked differences in the demographic structure of populations. Populations of B. fernandoi contained larger fractions of pre‐reproductive females and lower fractions of adult females with sexual eggs than populations of B. calyciflorus, B. elevatus, and B. dorcas. Mortality was found to be highest in B. elevatus and lowest in B. calyciflorus populations. Our results show that a reverse taxonomy approach is powerful in revealing sources of variation in ecologically relevant traits of cryptic species, such as life history and demographic structure. Explicit consideration of this variation is crucial for future studies of their dynamics in natural communities.
Nutrient limitation of primary producers has repeatedly been shown to negatively affect consumers, directly through stoichiometric mismatch and indirectly via alterations in the producer's biochemical quality or palatability. In this study, we assessed whether direct and indirect impacts of phosphorus‐limitation on a planktonic consumer are transferred to the next generation via maternal effects and whether these effects reflect an anticipatory adaptive strategy. For this, we subjected cultures of the algivorous monogonont rotifer Brachionus calyciflorus to three food quality treatments, i.e. P‐limited (LP), P‐replete (HP) and P‐enriched LP algae (i.e. algae with an LP‐growth history but with molar C:P ratios equal to those of HP‐algae). After two generations, we subjected offspring of these cultures to each of the three food quality treatments and monitored life history traits. In addition, we tested starvation resistance. Our results showed very strong negative maternal effects of low P food on offspring performance. These negative effects prevailed irrespective of contemporary diets, suggesting transmissive and selfish maternal effects rather than anticipatory adaptive effects. The relative strength of direct and indirect maternal P‐limitation effects varied among different traits. Adult body size was predominantly determined by direct effects of P‐shortage in maternal as well as contemporary food (LP < LP + P and LP + P = HP). In contrast, whereas egg size was negatively affected by direct effects of P‐limitation in the maternal diet, a contemporary diet of LP and LP + P algae resulted in larger eggs than HP algae. Animals born from such larger eggs showed no higher growth rates, but they were more resistant to starvation, likely as the result of higher maternal allocation of energy rich molecules to the eggs. The present study shows that maternal food conditions represent an important factor that should be taken into account in studies of stoichiometric mismatch between producers and consumers.
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