Accurate reconstruction of the biomass, structure, and productivity of ancient forests from their fossilized remnants remains an interesting challenge in paleoecology. In well-preserved Tertiary fossil Metasequoia forests of Canada's Arctic, in situ stumps and fragments of stems, treetops, and branches contain substantial information about tree dimensions that can be used to determine tree height, stand biomass, and other characteristics such as canopy depth and structure, and the history of stand development. To validate a method for reconstructing the biomass of the Eocene floodplain Metasequoia forests of Axel Heiberg Island, we measured stump diameters and spacing, and stem, branch, and treetop characteristics in living Metasequoia glyptostroboides and Chamaecyparis thyoides stands in ways that simulate the limited measurements that can be made in well-preserved fossil forests in Canada and probably elsewhere. We used those limited measurements to estimate tree height and volume, branch and foliar dry weights, and tree biomass. The estimates derived from the limited data set are usually within 15% of the estimates derived from the methods currently used in forest ecology for determining those metrics in modern forests. Under appropriate conditions, the biomass of ancient forests can be estimated with reasonable confidence.

Abundant fossil plant remains are preserved in the high-latitude middle Eocene deposits of the Buchanan Lake Formation on Axel Heiberg Island, Nunavut Territory, Canada. Intact leaf litter, logs, and stumps preserved in situ as mummified remains offer an opportunity to determine the structure, biomass, and productivity of two Taxodiaceae-dominated forests that grew north of the Arctic Circle (paleolatitude 75–80°N). We excavated fossil tree trunks and treetops to develop equations that describe the height, structure, and mass of the aboveground components of Eocene-age Metasequoia trees. We combined those data with surveys of in situ stumps to determine the structure, biomass, and productivity of two fossil forests, “N” and “HR.” We calculated a canopy height of 40 ± 3 m for the N forest and 25 m ± 2 m for the HR forest. Buried knots in dissected logs and abundant branch-free bole wood indicate that these were tall, closed-canopy forests. Stem tapers indicate that the overstory was of uniform height. Stem volume equaled 2095 m3 ha−1 and stem biomass was 628 Mg ha−1 in the N forest. Volume and biomass in the HR forest were much smaller, 211 m3 ha−1 and 63.3 Mg ha−1, respectively. We estimated understory tree biomass to be 40 Mg ha−1 in the N forest and 3.5 Mg ha−1 in the HR forest. Recovery of seven fossil treetops with exposed branch stubs enabled us to make estimates of branchwood and foliar biomass using allometric equations derived from modern, plantation-grown Metasequoia glyptostroboides. Estimated stand-level branch biomass was 13 and 6.7 Mg ha−1 in the N and HR forests, respectively. Standing foliar biomass was estimated to be 3.2 and 2.1 Mg ha−1 in the N and HR forests, respectively. Using annual ring widths, the reconstructed parabolic stems, and wood density of modern Metasequoia, we calculated annual wood production to be 2.3 Mg ha−1 yr−1 for the N forest and 0.55 Mg ha−1 yr−1 for the HR forest Assuming that the ancient Metasequoia were deciduous like their living relatives, annual aboveground net primary productivity was 5.5 Mg ha−1 yr−1 for the N forest and 2.8 Mg ha−1 yr−1 for the HR forest. Our estimated biomass and productivity values for N are similar to those of modern old-growth forests of the Pacific Northwest of the United States and old-growth coastal Cordillera forests of southern Chile.

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.