Carnegiea gigantea (saguaro, Cactaceae) is a protected and keystone species in the Sonoran Desert upon which nearly all of the ecosystem’s fauna rely. Little is known about its branching habits, which are particularly important as branching is directly related to seed production. This study assessed the ages and heights of plants when branching commenced under four different environmental regimes. Significant variations were observed in the onset of branching across the populations by both age (78–139 years) and height (4.4–5.5 m), likely related to the interplay between summer precipitation (and thus growth rate) and winter precipitation, which increases branching in any given individual. In all four populations, branches developed from areoles that are themselves reproductive (past or present); i.e., the lower areoles that pre-date the start of reproduction do not appear to contribute to branch production. Further, unlike the wide range of ages at the start of branching, the delay between the start of fruiting and the production of branches appears to be relatively consistent across populations, hovering around a 30-year time lag. Of greatest concern is the advanced age (around 140 years) at which branching begins at the most marginal, arid site, which is around the same time that old-age mortality may set in for the species (generally), dramatically diminishing reproductive output in an already marginal population. If this keystone species declines in this already arid to hyper-arid environment, it would have dramatic and irreversible consequences for the fauna and the ecosystem in this region.