Living with the invader: two native deer, two outcomes along a gradient of axis deer invasion

Abstract

Exotic species can reshape native communities, in the extreme, to local extirpation. Before that, natives may adjust where and when they use the landscape, in ways that allow coexistence and may be the earliest sign of trouble. Distinguishing coexistence built on such adjustment from one built on pre-existing niche differences requires comparing a native where the invader is absent with the same native where it is present, across sites at different invasion stages. Using eight years (2018-2026) of camera-trap data from 103 stations across Uruguay (154,590 camera-nights), we asked how the invasive axis deer (Axis axis) affects the space and time use of two native cervids: the forest-dwelling brown brocket (Subulo gouazoubira) and the grassland-restricted pampas deer (Ozotoceros bezoarticus). Activity-overlap, directional-avoidance, hierarchical activity, and two-species occupancy analyses showed sharply divergent dyads. A. axis and S. gouazoubira shared forest yet segregated in space and along the understory stratum the brocket requires and the invader avoids, and shifted their diel activity by context; understory cover fell with A. axis abundance, consistent with habitat modification. For A. axis and O. bezoarticus, segregation was intrinsic, opposite diel niches and habitats, with no support for a detectable shift in pampas deer behavior. The only behavioral signal was asymmetric: the invader, rather than the native, appeared to adjust, though few shared sites warrant caution. The same invader thus drives different mechanisms, but coexistence is provisional: A. axis is still expanding, far from equilibrium, so these shifts read best as early warning signals.