Ecophysiology of four co-occurring lycophyte species: an investigation of functional convergence

Lycophytes are the most early divergent extant lineage of vascular land plants. The group has a broad global distribution ranging from tundra to tropical forests and can make up an important component of temperate northeast US forests. We know very little about the in situ ecophysiology of this group and apparently no study has evaluated if lycophytes conform to functional patterns expected by the leaf economics spectrum hypothesis. To determine factors influencing photosynthetic capacity (Amax), we analysed several physiological traits related to photosynthesis to include stomatal, nutrient, vascular traits, and patterns of biomass distribution in four coexisting temperate lycophyte species: Lycopodium clavatum, Spinulum annotinum, Diphasiastrum digitatum and Dendrolycopodium dendroideum. We found no difference in maximum photosynthetic rates across species, yet wide variation in other traits. We also found that Amax was not related to leaf nitrogen concentration and is more tied to stomatal conductance, suggestive of a fundamentally different sets of constraints on photosynthesis in these lycophyte taxa compared with ferns and seed plants. These findings complement the hydropassive model of stomatal control in lycophytes and may reflect canalization of function in this group. Our data also demonstrate functional ecological similarities: De. dendroideum and D. digitatum are species that have substantial belowground biomass investment and are consistently more similar to each other across multiple traits than either is to the more surficial S. annotinum and L. clavatum. Such differences may partition environments in ways that allow for the close coexistence of these species.