The Santa Monica Mountains in southern California represent a mediterranean-type ecosystem, dominated by chaparral shrub communities adapted to protracted summer drought. However, ferns also thrive in the chaparral understory. While most fern species require abundant water, previous research has revealed several strategies that chaparral ferns use to survive protracted drought, including several desiccation-tolerant (DT) “resurrection” ferns. DT species can survive near-complete desiccation of the entire plant and subsequently resurrect when re-watered. However, the large sporophyte stage is just one part of the fern life cycle. Fern sporophytes only develop following the survival and reproduction of independent fern gametophytes, small plants lacking structures to limit water loss such as a cuticle and stomata. Many fern gametophytes are ephemeral, living just a few months in humid habitats before fertilization and emergence of the sporophyte. To our knowledge, no one has previously examined fern gametophytes in situ in the chaparral understory. Previous studies have shown that the physiological adaptations of tropical gametophytes parallel the ecological strategies of the sporophyte, but does this pattern hold in the chaparral understory? We hypothesized that 1) chaparral fern gametophytes will reflect the diversity of sporophyte survival strategies, and 2) chaparral fern gametophytes have adapted to survive seasonal drought. We examined physiological, ecological, and developmental aspects of four fern species. Woodwardia fimbriata (evergreen) grows in riparian streams, while Dryopteris arguta (evergreen), Adiantum jordanii (deciduous), and Pentagramma triangularis (resurrection) grow in the dry chaparral understory. All four species are terrestrial. Field studies revealed that the gametophyte microclimate differs from the sporophyte microclimate. Gametophyte thalli experienced wetter, cooler, and shadier conditions than sporophyte leaves (p < 0.05). This disequilibrium between the near-soil and free-air environments may allow gametophytes to remain hydrated and metabolically active long past the last spring rain event. Lab-grown gametophytes from at least two species (D. arguta and A. jordanii) were DT, as shown by partial recovery of dark-adapted chlorophyll fluorescence (Fv/Fm) following desiccation (Fv/Fm = 0). However, recovery from desiccation varied with the substrate on which the gametophytes were dried. D. arguta gametophytes showed some recovery following growth and desiccation on soil, but gametophytes grown on agar and dried on filter paper did not recover from desiccation. These differences may be due to the slower drying speed of the soil substrate or to morphological differences in the gametophytes. Developmental characteristics of lab-grown gametophytes appear to match the ecology of the sporophytes. The species with the largest sporophyte (W. fimbriata) also has the fastest growing gametophyte (shown by average area at the end of 20 days) compared to the other three species. Taken together, our results suggest that the ecological and physiological diversity of chaparral fern sporophytes is also reflected in the gametophyte stage. As we consider how climate change will impact already water-limited ecosystems, it is critical that we understand survival limits of fern gametophytes.