Research on the ecological impacts of environmental change has primarily focused at the species level, leaving the responses of ecosystem-level properties like energy flow poorly understood. This is especially so over millennial timescales inaccessible to direct observation. Here we examine how energy flow within a Great Basin small mammal community responded to climate-driven environmental change during the past 12,800 y, and use this baseline to evaluate responses observed during the past century. Our analyses reveal marked stability in energy flow during rapid climatic warming at the terminal Pleistocene despite dramatic turnover in the distribution of mammalian body sizes and habitat-associated functional groups. Functional group turnover was strongly correlated with climatedriven changes in regional vegetation, with climate and vegetation change preceding energetic shifts in the small mammal community. In contrast, the past century has witnessed a substantial reduction in energy flow caused by an increase in energetic dominance of smallbodied species with an affinity for closed grass habitats. This suggests that modern changes in land cover caused by anthropogenic activities-particularly the spread of nonnative annual grasslandshas led to a breakdown in the compensatory dynamics of energy flow. Human activities are thus modifying the small mammal community in ways that differ from climate-driven expectations, resulting in an energetically novel ecosystem. Our study illustrates the need to integrate across ecological and temporal scales to provide robust insights for long-term conservation and management.paleoecology | Quaternary | desert rodents | body size | invasive grass T he chronic and cumulative anthropogenic impacts of the past century have spurred considerable shifts in climates and habitats over local to global scales. Efforts to understand ecological responses to these changes have largely been focused at the species level (e.g., range shifts) over annual to decadal timescales. However, climate and habitat change impact biotas across all levels of ecological organization, from individuals to entire ecosystems. Furthermore, many ecological processes unfold over timescales inaccessible to direct observation or experimentation (1). Evaluating the response of higher-level aggregate properties (e.g., energy flow) to environmental change across centennial to millennial timescales can thus provide the temporal context needed to assess whether modern systems now fall within or outside their natural range of variability.Emphasis on species-level responses to environmental change is rooted in the paradigm that populations are highly dynamic while the aggregate properties of a community remain relatively stable and are thus more robust to perturbation. Aggregate properties have been shown to reflect resource availability (2, 3), and their stability has often been attributed to compensatory dynamics among individual species or functional groups (4). However, recent work has shown that marked change in aggrega...