Competition for microsites during recruitment in semiarid annual plant communities

The concept of microsites for recruitment is central to plant ecology, but it is unclear whether these sites are abstract constructs or real entities. I hypothesize that, in generally microsite-limited communities, microsites comprise a limiting physical resource for which different species compete. I tested this hypothesis on winter-annual communities on biocrust in the semiarid Northern Negev of Israel, in which most species are microsite-limited, while the dominant grass (Stipa capensis) has overcome this limitation by efficient microsite acquisition and a lack of secondary seed dormancy. I tested whether the dominant suppresses the subordinate species, collectively, during recruitment, rather than during growth. To this end, I conducted a field experiment with three blocks of six plots (6 m × 6 m) with two treatments - mowing in spring 2006 (intershrub, intershrub + shrub patches, and none) and shrub-patch removal (0% or 50% of the patches). I collected data from four seed traps per plot before spring 2007 and from five plant samples per plot at the end of spring. Mowing significantly reduced both seed and plant density of the dominant species, reflecting seed-limited recruitment, and increased subordinate plant density by competitive release. Multiple regressions of per-plant and per-gram effects and responses showed that competition was a direct effect of the dominant's density. Total and per-group biomass was proportional to density, implying density-independent per capita growth. Subordinate species number also increased with their density, due to the sample-size effect. These findings indicate that the seed-limited dominant diffusely suppresses the subordinates during recruitment, supporting the microsite competition hypothesis. The shift from growth resources to microsites extends the role of inter-specific competition along productivity and disturbance gradients, and highlights the asymmetric relationship between the two kinds of competition, as microsite competition is only observable if initial abundances are not overshadowed by density-dependent growth and mortality. The findings also demonstrate that (1) lacking secondary seed dormancy is an evolutionarily stable strategy in dryland annuals, alongside seed dormancy in microsite-limited species, and (2) biomass removal (e.g., by herbivory) increases small-scale biodiversity, enhancing the sustainability of dryland grazing, but without compensatory growth.