Declining CO 2 over the Cretaceous has been suggested as an evolutionary driver of the high leaf vein densities (7-28 mm mm −2 ) that are unique to the angiosperms throughout all of Earth history. Photosynthetic modeling indicated the link between high vein density and productivity documented in the modern low-CO 2 regime would be lost as CO 2 concentrations increased but also implied that plants with very low vein densities (less than 3 mm mm −2 ) should experience substantial disadvantages with high CO 2 . Thus, the hypothesized relationship between CO 2 and plant evolution can be tested through analysis of the concurrent histories of alternative lineages, because an extrinsic driver like atmospheric CO 2 should affect all plants and not just the flowering plants. No such relationship is seen. Regardless of CO 2 concentrations, low vein densities are equally common among nonangiosperms throughout history and common enough to include forest canopies and not just obligate shade species that will always be of limited productivity. Modeling results can be reconciled with the fossil record if maximum assimilation rates of nonflowering plants are capped well below those of flowering plants, capturing biochemical and physiological differences that would be consistent with extant plants but previously unrecognized in the fossil record. Although previous photosynthetic modeling suggested that productivity would double or triple with each Phanerozoic transition from low to high CO 2 , productivity changes are likely to have been limited before a substantial increase accompanying the evolution of flowering plants.
Role of CO 2 in Plant Evolution and Productivity over Geological Time ScalesA tmospheric CO 2 concentrations have fluctuated greatly over the past 400 million years: CO 2 levels are thought to have decreased by a factor of 10 or more as a result of the Devonian evolution of deep rooting plants and, subsequently, to have varied between levels somewhat less than and fivefold greater than preindustrial levels (1, 2). Just as land plant evolution has been a primary driver of these changes, in turn, CO 2 has often been assigned a dominant role in plant evolution. CO 2 changes have been implicated for the radiation of vascular plants, seed plants, and flowering plants; for the spread of C 4 photosynthesis and grasslands; and for the evolution of arborescence and both laminate leaves in general and the high vein density leaves of angiosperms in particular (3-12). Furthermore, models of plant function have repeatedly predicted that large swings in terrestrial productivity of 200-300% accompany these changes in atmospheric CO 2 (13-16).Plants require CO 2 , and productivity can be adversely affected by the CO 2 minima of the recent geological past that have approached the CO 2 compensation point for plant growth (17, 18), but did the positive relationship with CO 2 carry to a doubling or tripling of current productivity during the Mesozoic highs in CO 2 concentration? This suggestion raises a number of complications. Fo...