As dominance shifted from lycopsids and pteridophytes in the Paleozoic, to gymnosperms in the Mesozoic, to angiosperms in the Tertiary, plant architecture became more sparing in its use of lignin. Lignin-degrading organisms were rare or absent in the Paleozoic, but diverse and abundant in the Tertiary. Thus the terrigenous organic-carbon cycle has quickened over time, the fraction of terrestrial primary production preserved in coals and kerogens has declined, and terrestrial production has been able to increase over time without concomitant rises in atmospheric O2.

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