Not dead yet: the seasonal water relations of two perennial ferns during California's exceptional drought.

The understory of the redwood forests of California's coast harbors perennial ferns, including Polystichum munitum and Dryopteris arguta. Unusual for ferns, these species are adapted to the characteristic Mediterranean-type dry season, but the mechanisms of tolerance have not been studied. The water relations of P. munitum and D. arguta were surveyed for over a year, including measures of water potential (Ψ), stomatal conductance (gs) and frond stipe hydraulic conductivity (K). A dehydration and re-watering experiment on potted P. munitum plants corroborated the field data. The seasonal Ψ varied from 0 to below -3 MPa in both species, with gs and K generally tracking Ψ; the loss of K rarely exceeded 80%. Quantile regression analysis showed that, at the 0.1 quantile, 50% of K was lost at -2.58 and -3.84 MPa in P. munitum and D. arguta, respectively. The hydraulic recovery of re-watered plants was attributed to capillarity. The seasonal water relations of P. munitum and D. arguta are variable, but consistent with laboratory-based estimates of drought tolerance. Hydraulic and Ψ recovery following rain allows perennial ferns to survive severe drought, but prolonged water deficit, coupled with insect damage, may hamper frond survival. The legacy effects of drought on reproductive capacity and community dynamics are unknown.