Costs of acquiring phosphorus by vascular land plants: patterns and implications for plant coexistence

A survey of 659 papers mostly published since 1987 was conducted to compile a checklist of mycorrhizal occurrence among 3,617 species (263 families) of land plants. A plant phylogeny was then used to map the mycorrhizal information to examine evolutionary patterns. Several findings from this survey enhance our understanding of the roles of mycorrhizas in the origin and subsequent diversification of land plants. First, 80 and 92% of surveyed land plant species and families are mycorrhizal. Second, arbuscular mycorrhiza (AM) is the predominant and ancestral type of mycorrhiza in land plants. Its occurrence in a vast majority of land plants and early-diverging lineages of liverworts suggests that the origin of AM probably coincided with the origin of land plants. Third, ectomycorrhiza (ECM) and its derived types independently evolved from AM many times through parallel evolution. Coevolution between plant and fungal partners in ECM and its derived types has probably contributed to diversification of both plant hosts and fungal symbionts. Fourth, mycoheterotrophy and loss of the mycorrhizal condition also evolved many times independently in land plants through parallel evolution.

Nitrogen (N) tends to limit plant productivity on young soils; phosphorus (P) becomes increasingly limiting in ancient soils because it gradually disappears through leaching and erosion. Plant traits that are regarded as adaptations to N- and P-limited conditions include mycorrhizas and cluster roots. Mycorrhizas 'scavenge' P from solution or 'mine' insoluble organic N. Cluster roots function in severely P-impoverished landscapes, 'mining' P fixed as insoluble inorganic phosphates. The 'scavenging' and 'mining' strategies of mycorrhizal species without and non-mycorrhizal species with cluster roots, respectively, allow functioning on soils that differ markedly in P availability. Based on recent advances in our understanding of these contrasting strategies of nutrient acquisition, we provide an explanation for the distribution of mycorrhizal species on less P-impoverished soils, and for why, globally, cluster-bearing species dominate on severely P-impoverished, ancient soils, where P sensitivity is relatively common.