Abstract
The storage effect is a general explanation for coexistence in a
variable environment. The generality of the storage effect is both a
strength — it can be quantified in many systems — and a challenge
— there is not a clear relationship between the abstract conditions
for storage effect and species’ life-history traits (e.g., dormancy,
stage-structure, non-overlapping generations), thus precluding a simple
ecological interpretation of the storage effect. Our goal here is to
provide a clearer understanding of the conditions for the storage effect
as a step towards a better general explanation for coexistence in a
variable environment. Our approach focuses on dividing one of the key
conditions for the storage effect, covariance between environment and
competition, into two pieces, namely that there must be a causal
relationship between environment and competition, and that the effects
of the environment do not change too quickly. This finer-grained
definition can explain a number of previous results, including 1) that
the storage effect promotes annual plant coexistence when the
germination rate fluctuates, but not when the seed yield fluctuates, 2)
that the storage effect is more likely to be induced by resource
competition than apparent competition, and 3) that the spatial storage
effect is more probable than the temporal storage effect. Additionally,
our expanded definition suggests two novel mechanisms by which the
temporal storage effect can arise: transgenerational plasticity, and
causal chains of environmental variables. These mechanisms produce
coexistence via the storage effect without any need for stage structure
or a temporally autocorrelated environment.