The importance of being atomic: Ecological invasions as random walks instead of waves

Invasions are one of the most easily identified spatial phenomena in ecology, and have inspired a rich variety of theories for ecologists' and naturalists' consideration. However, a number of arguments over the sensitivities of invasion rates to stochasticity, density-dependence, dimension, and discreteness persist in the literature. The standard mathematical approach to invasions is based on Fisher's analysis of traveling waves solutions for the spread of an advantageous allele. In this paper, we exploit an alternative theory based on Ellner's premise that species invasions are best interpreted not as waves, but as random walks, and that the discreteness of living organisms is fundamentally important. Using a density-dependent invasion model in a stationary environment with indivisible (atomic) individuals where reproduction and dispersal are stochastic and independent, we show 4 key properties of Ellner's invasions previously suggested by simulation analysis: (1) greater spatial dispersal stochasticity quickens invasions, (2) greater demographic stochasticity slows invasions, (3) negative density-dependence slows invasions, and (4) greater temporal dispersal stochasticity quickens invasions. We prove the first three results by using generating functions and stochastic-dominance methods to rank furthest-forward dispersal distributions. The fourth result is proven in the special case of atomless theory, but remains an open conjecture in atomic theory. In addition, we explain why, unlike atomless invasions, an infinitely wide atomic invasion in two-dimensions can travel faster than a finite-width invasion and a one-dimensional invasion. The paper concludes with a classification of invasion dynamics based on dispersal kernel tails.