The earliest evidence of wildfire is documented from two localities: the early mid-Silurian Pen-y-lan Mudstone, Rumney, Wales (UK), and the late Silurian Winnica Formation, Winnica, Poland. Nematophytes dominate both charcoal assemblages. Reflectance data indicate low-temperature fires with localized intense conditions. Fire temperatures are greater in the older and less evolved assemblage. These charcoal assemblages and others, new and previously documented, from the Silurian and earliest Devonian are compared to box models of atmospheric oxygen concentration (pO2). Based on modern charring experiments, these data indicate pO2 is divergent from the broad trends predicted by the COPSE-revisited and GEOCARBSULFOR models. Sustained burns require a minimum pO2 threshold of 16%, or ~0.75 present atmospheric level. This threshold was first met and, our charcoal data indicate, was exceeded in the mid-Silurian and then, later in the Silurian, attained again repeatedly.

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