Disentangling endogenous versus exogenous pattern formation in spatial ecology: a case study of the ant Azteca sericeasur in southern Mexico.

Spatial patterns in ecology can be described as reflective of environmental heterogeneity (exogenous), or emergent from dynamic relationships between interacting species (endogenous), but few empirical studies focus on the combination. The spatial distribution of the nests of Azteca sericeasur, a keystone tropical arboreal ant, is thought to form endogenous spatial patterns among the shade trees of a coffee plantation through self-regulating interactions with controlling agents (i.e. natural enemies). Using inhomogeneous point process models, we found evidence for both types of processes in the spatial distribution of A. sericeasur. Each year's nest distribution was determined mainly by a density-dependent relationship with the previous year's lagged nest density; but using a novel application of a Thomas cluster process to account for the effects of nest clustering, we found that nest distribution also correlated significantly with tree density in the later years of the study. This coincided with the initiation of agricultural intensification and tree felling on the coffee farm. The emergence of this significant exogenous effect, along with the changing character of the density-dependent effect of lagged nest density, provides clues to the mechanism behind a unique phenomenon observed in the plot, that of an increase in nest population despite resource limitation in nest sites. Our results have implications in coffee agroecological management, as this system provides important biocontrol ecosystem services. Further research is needed, however, to understand the effective scales at which these relationships occur.