Insect-damaged fossil leaves record food web response to ancient climate change and extinction.

Plants and herbivorous insects have dominated terrestrial ecosystems for over 300 million years. Uniquely in the fossil record, foliage with well-preserved insect damage offers abundant and diverse information both about producers and about ecological and sometimes taxonomic groups of consumers. These data are ideally suited to investigate food web response to environmental perturbations, and they represent an invaluable deep-time complement to neoecological studies of global change. Correlations between feeding diversity and temperature, between herbivory and leaf traits that are modulated by climate, and between insect diversity and plant diversity can all be investigated in deep time. To illustrate, I emphasize recent work on the time interval from the latest Cretaceous through the middle Eocene (67-47 million years ago (Ma)), including two significant events that affected life: the end-Cretaceous mass extinction (65.5 Ma) and its ensuing recovery; and globally warming temperatures across the Paleocene-Eocene boundary (55.8 Ma). Climatic effects predicted from neoecology generally hold true in these deep-time settings. Rising temperature is associated with increased herbivory in multiple studies, a result with major predictive importance for current global warming. Diverse floras are usually associated with diverse insect damage; however, recovery from the end-Cretaceous extinction reveals uncorrelated plant and insect diversity as food webs rebuilt chaotically from a drastically simplified state. Calibration studies from living forests are needed to improve interpretation of the fossil data.

Habitat-related error in estimating temperatures from leaf margins in a humid tropical forest.

Leaf margin characters are strong predictors of mean annual temperature (MAT) in modern plant communities and widely used tools for reconstructing paleoclimates from fossil floras. However, the frequency of nonentire-margined species may vary dramatically between different habitats of the same forest. In this paper we explore the potential for this habitat variation to introduce error into temperature reconstructions, based on field data from a modern lowland forest in Amazonian Ecuador.The data show that the provenance of leaves can influence temperature estimates to an important degree and in a consistent direction. Woody plants growing along lakes and rivers underestimated MAT by 2.5°-5°C, while those in closed-canopy forest provided very accurate predictions. The high proportion of liana species with toothed leaves in lakeside and riverside samples appears to be responsible for a large part of the bias. Samples from closed-canopy forest that included both lianas and trees, however, were more accurate than tree-only or liana-only samples.We conclude that paleotemperature reconstructions based on leaf margin characters will be misleading to the extent that fossilization provides a better record of certain habitats than others. The preponderance of lake and river deposits in the angiosperm fossil record suggests that underestimation of mean annual paleotemperature may be common.

Insect herbivory, plant defense, and early Cenozoic climate change.

Insect damage on fossil leaves from the Central Rocky Mountains, United States, documents the response of herbivores to changing regional climates and vegetation during the late Paleocene (humid, warm temperate to subtropical, predominantly deciduous), early Eocene (humid subtropical, mixed deciduous and evergreen), and middle Eocene (seasonally dry, subtropical, mixed deciduous and thick-leaved evergreen). During all three time periods, greater herbivory occurred on taxa considered to have short rather than long leaf life spans, consistent with studies in living forests that demonstrate the insect resistance of long-lived, thick leaves. Variance in herbivory frequency and diversity was highest during the middle Eocene, indicating the increased representation of two distinct herbivory syndromes: one for taxa with deciduous, palatable foliage, and the other for hosts with evergreen, thick-textured, small leaves characterized by elevated insect resistance. Leaf galling, which is negatively correlated with moisture today, apparently increased during the middle Eocene, whereas leaf mining decreased.

Timing the radiations of leaf beetles: hispines on gingers from latest cretaceous to recent.

Stereotyped feeding damage attributable solely to rolled-leaf hispine beetles is documented on latest Cretaceous and early Eocene ginger leaves from North Dakota and Wyoming. Hispine beetles (6000 extant species) therefore evolved at least 20 million years earlier than suggested by insect body fossils, and their specialized associations with gingers and ginger relatives are ancient and phylogenetically conservative. The latest Cretaceous presence of these relatively derived members of the hyperdiverse leaf-beetle clade (Chrysomelidae, more than 38,000 species) implies that many of the adaptive radiations that account for the present diversity of leaf beetles occurred during the Late Cretaceous, contemporaneously with the ongoing rapid evolution of their angiosperm hosts.

Response of plant-insect associations to paleocene-eocene warming

The diversity of modern herbivorous insects and their pressure on plant hosts generally increase with decreasing latitude. These observations imply that the diversity and intensity of herbivory should increase with rising temperatures at constant latitude. Insect damage on fossil leaves found in southwestern Wyoming, from the late Paleocene-early Eocene global warming interval, demonstrates this prediction. Early Eocene plants had more types of insect damage per host species and higher attack frequencies than late Paleocene plants. Herbivory was most elevated on the most abundant group, the birch family (Betulaceae). Change in the composition of the herbivore fauna during the Paleocene-Eocene interval is also indicated.