Links between environment and stomatal size through evolutionary time in Proteaceae

The size of plant stomata (adjustable pores that determine uptake of CO2 and loss of water from leaves) is considered to be evolutionarily important.  This study uses fossils from the major southern hemisphere family Proteaceae to test whether stomatal cell size responded to Cenozoic climate change.  We measured the length and abundance of guard cells (the cells forming stomata), the area of epidermal pavement cells, stomatal index and maximum stomatal conductance from a comprehensive sample of fossil cuticles of Proteaceae, and extracted published estimates of past temperature and atmospheric CO2.  We developed a novel test based on stochastic modelling of trait evolution to test correlations among traits.  Guard cell length increased, and stomatal density decreased significantly with decreasing palaeotemperature.  However, contrary to expectations, stomata tended to be smaller and more densely packed at higher atmospheric CO2.  Thus, associations between stomatal traits and paleoclimate over the last 70 MY in Proteaceae suggest that stomatal size is significantly affected by environmental factors other than atmospheric CO2.  Guard cell length, pavement cell area, stomatal density, stomatal index covaried in ways consistent with coordinated development of leaf tissues.

Methods
The data set includes stomatal and epidermal cell dimensions from leaves of fossil and extant species of Proteaceae. The extant species represent a phylogenetically and ecologically diverse sample of the family. The fossils are all suitably preserved fossils currently known of the family.