Aim:To disentangle the influence of environmental factors at different spatial grains (regional and local) on fern and lycophyte species richness and to ask how regional and plot-level richness are related to each other.Location: Global.

Functional traits determine how species interact with their abiotic and biotic environment. In turn, functional diversity describes how assemblages of species as a whole are adapted to their environment, which also determines how they might react to changing conditions. To fully understand functional diversity, it is fundamental to (a) disentangle the influences of environmental filtering and species richness from each other, (b) assess if the trait space saturates at high levels of species richness, and (c) understand how changes in species numbers affect the relative importance of the trait niche expansion and packing. In the present study, we determined functional diversity of fern assemblages by describing morphological traits related to resource acquisition along four tropical elevational transects with different environmental conditions and species richness. We used several functional diversity indices and their standardized effect size to consider different aspects of functional diversity. We contrasted these aspects of functional diversity with climate data and species richness using linear models and linear mixed models. Our results show that functional morphological trait diversity was primarily driven by species richness and only marginally by environmental conditions. Moreover, increasing species richness contributed progressively to packing of the morphological niche space, while at the same time decreasing morphological expansion until a saturation point was reached. Overall, our findings suggest that the density of co-occurring species is the fundamental driving force of morphological niche structure, and environmental conditions have only an indirect influence on fern resource acquisition strategies.

Echinopsis chiloensis is an endemic cactus from Chile, distributed in a temperature and rainfall gradient between 30° and 35° South latitude, with mean temperatures increasing and precipitation decreasing toward the north. It is the main host of the mistletoe Tristerix aphyllus, a holoparasite completely dependent on the cactus for water, carbon, and minerals. In this study, we investigated the consequences of parasitism over the fitness and physiology of this cactus throughout its distribution range and how it is affected by the environment. We measured five functional traits in eight populations latitudinally distributed, the first three only for the host: reproductive fitness, stomatal traits (density and size), and photosynthesis (during winter and summer); and the last two for the host and parasite: stable isotopes (∂13C and ∂15N), and nutrients (carbon and nitrogen content). The results showed a negative effect of parasitism over fitness of infected cacti. However, the higher nitrogen concentrations in cactus tissues toward the south improved overall fitness. Regarding photosynthesis, we only observed a negative effect of parasitism during the dry season (summer), which is also negatively affected by the increase in summer temperatures and decrease in winter rainfall toward the north. There were no differences in nutrient concentration or in the isotopic signature of healthy and infected cacti. Conversely, we observed a higher carbon and lower nitrogen concentration in mistletoes than in cacti regardless of latitude. The loss of temperature seasonality toward the north increases the C:N ratio, and the values between the parasite and its host diverge. ∂15N was similar between parasites and hosts while ∂13C of the parasite was enriched when compared to its host. Overall, the infection by T. aphyllus affects Echinopsis chiloensis fitness but showed no strong effects over the cactus physiology, except for the summer photosynthesis. Therefore, our data revealed that E. chiloensis response to T. aphyllus infection is sensitive to environmental changes in a way that could be strongly impacted by the desertification projected for this area due to climate change.

Circadian regulation plays a vital role in optimizing plant responses to the environment. However, while circadian regulation has been extensively studied in angiosperms, very little is known for lycophytes and ferns, leaving a gap in our understanding of the evolution of circadian rhythms across the plant kingdom. Here, we investigated circadian regulation in gas exchange through stomatal conductance and photosynthetic efficiency in a phylogenetically broad panel of 21 species of lycophytes and ferns over a 46-hour period under constant light and a selected few under more natural conditions with day-night cycles. No rhythm was detected under constant light for either lycophytes or ferns, except for two semi-aquatic species of the family Marsileaceae (Marsilea azorica and Regnellidium diphyllum), which showed rhythms in stomatal conductance. Furthermore, these results indicated the presence of a light-driven stomatal control for ferns and lycophytes, with a possible passive fine-tuning through leaf water status adjustments. These findings support previous evidence for the fundamentally different regulation of gas exchange in lycophytes and ferns compared to angiosperms, and they suggest the presence of alternative stomatal regulations in Marsileaceae, an aquatic family already well known for numerous other distinctive physiological traits. Overall, our study provides evidence for heterogeneous circadian regulation across plant lineages, highlighting the importance of broad taxonomic scope in comparative plant physiology studies.