Are fern stomatal responses to different stimuli coordinated? Testing responses to light, vapor pressure deficit, and CO2 for diverse species grown under contrasting irradiances.

The stomatal behavior of ferns provides an excellent system for disentangling responses to different environmental signals, which balance carbon gain against water loss. Here, we measured responses of stomatal conductance (gs ) to irradiance, CO2 , and vapor pressure deficit (VPD) for 13 phylogenetically diverse species native to open and shaded habitats, grown under high- and low-irradiance treatments. We tested two main hypotheses: that plants adapted and grown in high-irradiance environments would have greater responsiveness to all stimuli given higher flux rates; and that species' responsiveness to different factors would be correlated because of the relative simplicity of fern stomatal control. We found that species with higher light-saturated gs had larger responses, and that plants grown under high irradiance were more responsive to all stimuli. Open habitat species showed greater responsiveness to irradiance and CO2 , but lower responsiveness to VPD; a case of plasticity and adaptation tending in different directions. Responses of gs to irradiance and VPD were positively correlated across species, but CO2 responses were independent and highly variable. The novel finding of correlations among stomatal responses to different stimuli suggests coordination of hydraulic and photosynthetic signaling networks modulating fern stomatal responses, which show distinct optimization at growth and evolutionary time-scales.