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. Rising atmospheric CO2 is expected to increase globaltemperatures, plant water-use efficiency, and carbon storage in theterrestrial biosphere. A CO2 fertilization effect on terrestrialvegetation is predicted to cause global greening as the potential ecospacefor forests expands. However, leaf-level fertilization effects, such asincreased productivity and water-use efficiency, have not been documentedfrom fossil leaves in periods of heightened atmospheric CO2. Here, weuse leaf gas-exchange modeling on a well-preserved fossil flora from earlyMiocene New Zealand, as well as two previously published tropical florasfrom the same time period, to reconstruct atmospheric CO2, leaf-levelproductivity, and intrinsic water-use efficiency. Leaf gas-exchange ratesreconstructed from early Miocene fossils, which grew at southern temperateand tropical latitudes when global average temperatures were5–6 ∘C higher than today, reveal that atmospheric CO2 was∼450–550 ppm. Early Miocene CO2 was similar toprojected values for 2040 CE and is consistent with an Earth system sensitivityof 3–7 ∘C to a doubling of CO2. The Southern Hemispheretemperate leaves had higher reconstructed productivity than modern analogs,likely due to a longer growing season. This higher productivity waspresumably mirrored at northern temperate latitudes as well, where a greateravailability of landmass would have led to increased carbon storage inforest biomass relative to today. Intrinsic water-use efficiency of bothtemperate and tropical forest trees was high, toward the upper limit of therange for modern trees, which likely expanded the habitable range in regionsthat could not support forests with high moisture demands under loweratmospheric CO2. Overall, early Miocene elevated atmospheric CO2sustained globally higher temperatures, and our results provide the firstempirical evidence of concomitant enhanced intrinsic water-use efficiency,indicating a forest fertilization effect.

Elevated CO&lt;sub&gt;2&lt;/sub&gt;, increased leaf-level productivity, and water-use efficiency during the early Miocene

Elevated CO<sub>2</sub>, increased leaf-level productivity, and water-use efficiency during the early Miocene