Successional loss of two key food tree species best explains decline in group size of Panamanian howler monkeys (Alouatta palliata)

Negative impacts of discrete, short-term disturbances to wildlife populations are well-documented. The consequences of more gradual environmental change are less apparent and harder to study because they play out over longer periods and are often indirect in their action. Yet, they can drive the decline of wildlife populations even in seemingly pristine and currently well-protected habitats. One such environmental change is a successional shift in a community's species composition as it regenerates from disturbance caused by past human land use. Early and middle successional tree species often provide key foods to folivores and frugivores, but the abundance of these resources drops as the forest matures, with adverse repercussions for these consumers. Our 44-year record (1974-2018) of howler monkey (Alouatta palliata) group sizes and demographic composition from Barro Colorado Island, Panama, a protected reserve, documents an example of this phenomenon. After 70 years of relative stability, the mean size of howler monkey groups exhibited a marked decline, beginning in 2003. This downward trajectory in group size has continued through the most recent census in 2018. The composition of howler groups also changed significantly during the study period, with the patterns of decline differing among age/sex classes. There is no evidence that these changes were caused by increased rates of emigration, group fission, predation, parasitism, or disease. Rather, they are best explained by an island-wide, succession-driven decline in the densities of two species of free-standing fig trees, Ficus yoponensis and F. insipida, which together were providing ~36% of BCI howlers' annual diet. in Spanish is available with online material.