Sharply increased insect herbivory during the Paleocene–Eocene Thermal Maximum

Abstract
The Paleocene–Eocene Thermal Maximum (PETM, 55.8 Ma), an abrupt global warming event linked to a transient increase in p CO 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 p CO 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 p CO 2 increase and warming temperatures.