Ecophysiological responses of plants to global environmental change since the Last Glacial Maximum

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Abstract
Ecophysiological information on the responses of plants to past global environmental changes may be obtained from Quaternary fossil leaves by measurements of (i) stomatal density, (ii) stomatal dimensions and (iii) 13C discrimination (Δ13C). The stomatal density and stomatal dimensions of leaves can be used to calculate stomatal conductance, while leaf Δ13C values provide independent information on stomatal conductance and plant water use efficiency. In this paper, stomatal conductance is calculated for a sequence of radiocarbon dated fossil leaves of Salix herbacea L. which, together with herbarium and fresh material, represents a time‐series spanning from the Last Glacial Maximum (LGM) (16500 yr BP) to the present day. The calculated values were then tested against leaf Δ13C values previously reported for the same material. Our calculations show that stomatal conductance is negatively correlated with increases in atmospheric CO2 concentration over the last 16500 yr. This represents the first evidence of long‐term response of stomatal conductance to increases in atmospheric CO2 concentration and confirms the response observed in experimental systems exposing plants to lower‐than‐present CO2 concentrations in controlled environments. The calculated decrease in conductance was positively correlated with leaf Δ13C values, supporting this interpretation. The mean leaf Δ13C value for the 18th and 19th centuries was significantly (P≥ 0.05) lower than the mean for the interval LGM‐Holocene (10000 yr BP) implying an increase in plant water‐use‐efficiency over this time. These two lines of evidence, together with the stomatal density record from a glacial cycle, and experimental studies growing C3 plants in glacial‐to‐present CO2 concentrations, strongly imply that the water use efficiency of vegetation during the LGM was lower than at present and that it has increased since that time. Further evidence in support of this conclusion comes from the pattern of world vegetation types present during the LGM previously reconstructed using palaeoecological data. This evidence demonstrates that the distribution of vegetation types during the LGM was significantly different from that of the present day and showed a contraction in the area of rain forest and a major expansion of desert areas.