Demographic mechanisms in the coexistence of two closely related perennials in a fluctuating environment.

The demographic variability and life history differentiation of two closely related shrubs (Atriplex canescens and A. acanthocarpa) were investigated in the Chihuahuan Desert, and the results were interpreted in the context of theories of coexistence in fluctuating environments. Demographic information was recorded during three annual intervals and analyzed employing matrix projection models. A. canescens had lower lambda (finite rate of population increase), higher longevity and generation time and slower convergence to a stable population structure than A. acanthocarpa. In favorable years for recruitment (those when, for both species, lambda > 1), the lambda of A. acanthocarpa was higher than that of A. canescens; in unfavorable years (when lambda < 1), the reverse was true. Regardless of conditions (year), A. acanthocarpa had a type 2 survivorship curve (constant rate of mortality with age), while A. canescens had a type 3 survivorship curve (declining mortality with age). Elasticity analyses highlighted the larger influence that fecundity and growth would have in modifying the lambda of A. acanthocarpa in comparison to that of A. canescens. In contrast, survival would have a larger influence on the lambda of A. canescens. Atriplex acanthocarpa behaved as an opportunistic species that benefitted from sporadic favorable conditions and declined rapidly when conditions deteriorated. In contrast, A. canescens behaved as a tolerant species able to withstand years when conditions were poor, but which could not gain any advantage over A. acanthocarpa when conditions improved. By each having a relative advantage over the other on opposite ends of the contrasting climatic conditions experienced in the Chihuahuan Desert, they are able to coexist. Their contrasting life histories agreed with the theoretical predictions for the operation of the two mechanisms of species coexistence in fluctuating environments: the storage effect and the relative non-linearity of competition. Based on these results, we conclude by speculating on the nature of succession in arid communities.