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Summary Leaf nitrogen concentration often is higher in leguminous plants, which associate with dinitrogen‐fixing bacteria, compared with nonlegume plants. However, the range of nitrogen concentrations in legumes is wide, likely related to the range of nitrogen fixation strategies. We evaluated how carbon and nitrogen allocation to roots, stems and leaves is influenced by the type of strategy of nitrogen fixation regulation. We grew herbaceous annual legumes (Medicago truncatula, Hymenocarpos circinnatus and Vicia palaestina) under two nitrogen availability treatments (none/sufficient), with and without bacterial inoculation. We found facultative downregulation of the rate of nitrogen fixation when nitrogen was available in H. circinnatus, and an obligate similar fixation rate in both nitrogen treatments in M. truncatula and V. palaestina. Uninoculated plants invested more biomass in roots and contained lower nitrogen concentrations. However, nitrogen concentration in the entire plant and in the leaves was lower and more plastic in the species with a facultative fixation strategy, whereas species with an obligate fixation strategy also maintained high nitrogen concentrations. Our results suggest a suite of functional traits associated with the strategies of allocation and symbiotic nitrogen fixation. This suite of traits probably shapes successional and functional niches of different leguminous species in specious plant communities.

Summary Plants, especially perennials, growing in drylands and seasonally dry ecosystems are uniquely adapted to dry conditions. Legume shrubs and trees, capable of symbiotic dinitrogen (N2) fixation, often dominate in drylands. However, the strategies that allow symbiotic fixation in these ecosystems, and their influence on the nitrogen cycle, are largely unresolved. We evaluated the climatic, biogeochemical and ontogenetic factors influencing nitrogen fixation in an abundant Mediterranean legume shrub, Calicotome villosa. We measured nodulation, fixation rate, nitrogen allocation and soil biogeochemistry in three field sites over a full year. A controlled experiment evaluated differences in plant regulation of fixation as a function of soil nutrient availability and seedling and adult developmental stages. We found a strong seasonal pattern, shifting between high fixation rates during the rainy season at flowering and seed‐set times to almost none in the rainless season. Under controlled conditions, plants downregulated fixation in response to soil nitrogen availability, but this response was stronger in seedlings than in adult shrubs. Finally, we did not find elevated soil nitrogen under N2‐fixing shrubs. We conclude that seasonal nitrogen fixation, regulation of fixation, and nitrogen conservation are key adaptations influencing the dominance of dryland legumes in the community, with broader consequences on the ecosystem nitrogen cycle.

The importance of symbiotic dinitrogen (N2) fixation in shaping the coupled nitrogen–carbon cycle is now known for most humid terrestrial ecosystems. However, whether N2 fixation can play a key role in the nitrogen and carbon budget of water‐limited and seasonally dry ecosystems remains a mystery. The maintenance of metabolically and physiologically costly symbiotic fixation in water‐limited environments is highly complex. These costs are particularly high during the first developmental season, when allocation to deep rooting and drought resistance mechanisms is essential for seedling survival of prolonged seasonal drought. We, therefore, evaluated how drought‐adapted legume species change their allocation to symbiotic nitrogen fixation as a function of soil nitrogen availability. We tested this on seedlings of a suite of four common Mediterranean legume shrubs with a strong seasonal behaviour, which we grew under controlled nitrogen and phosphorus availabilities. We asked: (1) Do species differ in their investment and regulation of nitrogen fixation? (2) Is fixation regulated via plant allocation to nodules, fixation rate or both? and (3) Does phosphorus availability limit symbiotic nitrogen fixation? All Mediterranean perennial legumes in the experiment established and grew, nodulated, and fixed nitrogen, even under severe nitrogen limitation. The four species reacted similarly to nitrogen supply, by strongly downregulating fixation through both decreased nodulation and lower rate of fixation. However, we found a significant interspecific difference in fixation (both nodulation and rate), biomass production and growth rate. Our experimental species presented a range of fixation investment strategies corresponding to life‐history and resource partitioning patterns. Phosphorus limitation had a minor influence on both fixation and plant growth. Synthesis. The high physiological cost of symbiotic fixation imposes the need to tightly regulate fixation in perennial legumes coping with severe water stress. Control of fixation allows legume species to colonize recently disturbed nitrogen‐deficient habitats, cope with grazing, survive long seasonal droughts and recover nitrogen fixation later in the wet season, and survive over time by reducing nitrogen inputs to the ecosystem.