South America has some of the most diverse floras and insect faunas that are known, but its Cenozoic fossil record of insects and insect herbivory is sparse. We quantified insect feeding on 3,599 leaves from the speciose Laguna del Hunco flora (Chubut, Argentina), which dates to the early Eocene climatic optimum (52 million years ago) and compared the results with three well preserved, rich, and identically analyzed early-and middle-Eocene floras from the following sites in North America: Republic, WA; Green River, UT; and Sourdough, WY. We found significantly more damage diversity at Laguna del Hunco than in the North American floras, whether measured on bulk collections or on individual plant species, for both damage morphotypes and feeding groups. An ancient history of rich, specialized plant-insect associations on diverse plant lineages in warm climates may be a major factor contributing to the current biodiversity of South America.paleobotany ͉ Argentina ͉ herbivory ͉ Laguna del Hunco ͉ paleoecology S outh America is well known for its highly diverse** floras (1-3) and associated insect faunas (4-7). Observations in South America and elsewhere show strong positive linkage between plant and insect-herbivore diversity (8)(9)(10)(11)(12)(13)(14). Proposed mechanisms include dependency of insect diversity on plant diversity (8, 9, 13), coevolution of plants and insects (15-21), herbivore selection against host density (22-25), and herbivore intensification of abiotic factors that select for habitat specialization (26,27). Paleontological data can be used to test whether plant and herbivore diversity were correlated in the past (28-30), although data bearing on the primary mechanisms are not typically available from fossils. However, throughout South America, insect body fossils have a limited Cenozoic record (31), and insect damage on fossil plants has rarely been reported (32,33). Here, we seek to determine whether a diverse fossil flora from South America has an elevated level of plant-insect associations. We quantify insect-feeding damage on one of the most speciose known assemblages of fossil plants, from earlyEocene Patagonia and, for comparison, on three well preserved, diverse early-and middle-Eocene floras from comparable absolute latitudes of North America.
Study Areas, Specimens, and MethodsThe Patagonian flora comes from tuffaceous lake beds exposed at Laguna del Hunco (LH; S42.5°, W70°; paleolatitude, Ϸ47°S) in Chubut, Argentina (34-36). These sediments were deposited 52 million years ago (35,36), during the early Eocene global climatic optimum (37). At this time, thermophilic organisms reached middle and high latitudes of both hemispheres (38-42), and the LH flora contains a large proportion of tropical as well as temperate lineages (34)(35)(36)39). Our unbiased macrofloral sample, in which all material was collected or tallied, totals 6,521 specimens from 25 quarries, of which 4,303 specimens (66%) are identifiable to 186 species of plant organs. These include 152 leaf species, of which 132 are ma...