The Paleocene-Eocene Thermal Maximum (PETM, 55.8 Ma), an abrupt global warming event linked to a transient increase in pCO 2, was comparable in rate and magnitude to modern anthropogenic climate change. Here we use plant fossils from the Bighorn Basin of Wyoming to document the combined effects of temperature and pCO 2 on insect herbivory. We examined 5,062 fossil leaves from five sites positioned before, during, and after the PETM (59 -55.2 Ma). The amount and diversity of insect damage on angiosperm leaves, as well as the relative abundance of specialized damage, correlate with rising and falling temperature. All reach distinct maxima during the PETM, and every PETM plant species is extensively damaged and colonized by specialized herbivores. Our study suggests that increased insect herbivory is likely to be a net long-term effect of anthropogenic pCO 2 increase and warming temperatures.Bighorn Basin ͉ paleobotany ͉ plant-insect interactions ͉ rapid climate change D uring the 21st century, global surface temperature is expected to increase 1.8-4.0°C as higher atmospheric concentrations of greenhouse gases (especially CO 2 ) are generated by human activities (1). Food webs incorporating plants and phytophagous insects account for up to 75% of modern global biodiversity (2), so their response to this anthropogenic change will have a profound effect on the biosphere. Experiments show that plants grown in elevated CO 2 tend to accumulate more carbon and have a higher carbon:nitrogen ratio; they are, therefore, nutritionally poorer (3-5), leading to an average compensatory increase in insect consumption rates (6) as nitrogen becomes limiting. Modern insect herbivory and herbivore diversity are greatest overall in the tropics (7-10), implying a broad correlation between temperature and herbivory, and Pliocene-Pleistocene fossils show rapid shifts in the geographic ranges of insects in response to climate change (11). These existing data provide limited insight into future changes, however. The complexity of plant-insect food webs makes it difficult to generalize from experiments to the response of natural ecosystems over long time scales (12). Modern and PliocenePleistocene insect biogeographic patterns have not been directly linked to pCO 2 and do not document the response of plant-insect food webs to rapid increases in temperature and pCO 2 . Well preserved Paleocene-Eocene fossil angiosperm leaves show insect feeding damage and, therefore, can be used to investigate the net effects of increasing temperature and pCO 2 on full plant-insect food webs over long time scales.Beginning in the late Paleocene, global temperatures gradually warmed to the sustained Cenozoic maximum at Ϸ53 Ma (13). The Paleocene-Eocene Thermal Maximum (PETM) is a transient spike of high temperature and pCO 2 representing Ϸ100 thousand years (ky), superimposed on a longer interval of gradual warming (14, 15); it is one of the best deep-time analogues for the modern time scale of global warming. The PETM is marked by a negative carbon isotope ...