Leaf mass per area (MA) is a central ecological trait that is intercorrelated with leaf life span, photosynthetic rate, nutrient concentration, and palatability to herbivores. These coordinated variables form a globally convergent leaf economics spectrum, which represents a general continuum running from rapid resource acquisition to maximized resource retention. Leaf economics are little studied in ancient ecosystems because they cannot be directly measured from leaf fossils. Here we use a large extant data set (65 sites; 667 species-site pairs) to develop a new, easily measured scaling relationship between petiole width and leaf mass, normalized for leaf area; this enables MA estimation for fossil leaves from petiole width and leaf area, two variables that are commonly measurable in leaf compression floras. The calibration data are restricted to woody angiosperms exclusive of monocots, but a preliminary data set (25 species) suggests that broad-leaved gymnosperms exhibit a similar scaling. Application to two well-studied, classic Eocene floras demonstrates that MA can be quantified in fossil assemblages. First, our results are consistent with predictions from paleobotanical and paleoclimatic studies of these floras. We found exclusively low-MA species from Republic (Washington, U.S.A., 49 Ma), a humid, warm-temperate flora with a strong deciduous component among the angiosperms, and a wide MA range in a seasonally dry, warm-temperate flora from the Green River Formation at Bonanza (Utah, U.S.A., 47 Ma), presumed to comprise a mix of short and long leaf life spans. Second, reconstructed MA in the fossil species is negatively correlated with levels of insect herbivory, whether measured as the proportion of leaves with insect damage, the proportion of leaf area removed by herbivores, or the diversity of insect-damage morphotypes. These correlations are consistent with herbivory observations in extant floras and they reflect fundamental trade-offs in plant-herbivore associations. Our results indicate that several key aspects of plant and plant-animal ecology can now be quantified in the fossil record and demonstrate that herbivory has helped shape the evolution of leaf structure for millions of years.

We document evidence of endophytic oviposition on fossil compression/impression leaves from the early Eocene Laguna del Hunco and middle Eocene Rio Pichileufu floras of Patagonia, Argentina. Based on distinctive morphologies and damage patterns of elongate, ovoid, lens-, or teardrop-shaped scars in the leaves, we assign this insect damage to the ichnogenus Paleoovoidus, consisting of an existing ichnospecies, P. rectus, and two new ichnospecies, P. arcuatum and P. bifurcatus. In P. rectus, the scars are characteristically arranged in linear rows along the midvein; in P. bifurcatus, scars are distributed in double rows along the midvein and parallel to secondary veins; and in P. arcuatum, scars are deployed in rectilinear and arcuate rows. In some cases, the narrow, angulate end of individual scars bear a darkened region encompassing a circular hole or similar feature indicating ovipositor tissue penetration. A comparison to the structure and surface pattern of modern ovipositional damage on dicotyledonous leaves suggests considerable similarity to certain zygopteran Odonata. Specifically, members of the Lestidae probably produced P. rectus and P. bifurcatus, whereas species of Coenagrionidae were responsible for P. arcuatum. Both Patagonian localities represent an elevated diversity of potential fern, gymnosperm, and especially angiosperm hosts, the targets of all observed oviposition. However, we did not detect targeting of particular plant families. Our results indicate behavioral stasis for the three ovipositional patterns for at least 50 million years. Nevertheless, synonymy of these oviposition patterns with mid-Mesozoic ichnospecies indicates older origins for these distinctive modes of oviposition.