Adding energy gradients and long‐distance dispersal to a neutral model improves predictions of Madagascan bird diversity

Macroecological patterns are likely the result of both stochastically neutral mechanisms and deterministic differences between species. In Madagascar, the simplest stochastically neutral hypothesis - the mid-domain effects (MDE) hypothesis - has already been rejected. However, rejecting the MDE hypothesis does not necessarily refute the existence of all other neutral mechanisms. Here, we test whether adding complexity to a basic neutral model improves predictions of biodiversity patterns. The simplest MDE model assumes that: (1) species' ranges are continuous and unfragmented, (2) are randomly located throughout the landscape, and (3) can be stacked independently and indefinitely. We designed a simulation based on neutral theory that allowed us to weaken each of these assumptions incrementally by adjusting the habitat capacity as well as the likelihood of short- and long-distance dispersal. Simulated outputs were compared to four empirical patterns of bird diversity: the frequency distributions of species richness and range size, the within-island latitudinal diversity gradient, and the distance-decay of species compositional similarity. Neutral models emulated empirical diversity patterns for Madagascan birds accurately. The frequency distribution of range size, latitudinal diversity gradient, and the distance-decay of species compositional similarity could be attributed to stochastic long-distance migration events and zero-sum population dynamics. However, heterogenous environmental gradients improved predictions of the frequency distribution of species richness. Patterns of bird diversity in Madagascar can broadly be attributed to stochastic long-distance migration events and zero-sum population dynamics. This implies that rejecting simple hypotheses, such as MDE, does not serve as evidence against stochastic processes in general. However, environmental gradients were necessary to explain patterns of species richness and deterministic differences between species are probably important for explaining the distributions of narrow-range and endemic species.