Extremely thin but very robust: Surprising cryptogam trait combinations at the end of the leaf economics spectrum.

Leaf economics spectrum (LES) describes the fundamental trade-offs between leaf structural, chemical, and physiological investments. Generally, structurally robust thick leaves with high leaf dry mass per unit area (LMA) exhibit lower photosynthetic capacity per dry mass (A _mass). Paradoxically, "soft and thin-leaved" mosses and spikemosses have very low A _mass, but due to minute-size foliage elements, their LMA and its components, leaf thickness (LT) and density (LD), have not been systematically estimated. Here, we characterized LES and associated traits in cryptogams in unprecedented details, covering five evolutionarily different lineages. We found that mosses and spikemosses had the lowest LMA and LT values ever measured for terrestrial plants. Across a broad range of species from different lineages, A _mass and LD were negatively correlated. In contrast, A _mass was only related to LMA when LMA was greater than 14 g cm^{- 2}. In fact, low A _mass reflected high LD and cell wall thickness in the studied cryptogams. We conclude that evolutionarily old plant lineages attained poorly differentiated, ultrathin mesophyll by increasing LD. Across plant lineages, LD, not LMA, is the trait that represents the trade-off between leaf robustness and physiology in the LES.