Hydathodes in ferns: their phylogenetic distribution, structure and function.

BACKGROUND AND AIMS: Ferns are the second largest group of vascular plants and are distributed nearly worldwide. Although ferns have been integrated into some comparative ecological studies focusing on hydathodes, there is a considerable gap in our understanding of the functional anatomy of these secretory tissues that are found on the vein endings of many fern leaves. In this study, we aimed to investigate the phylogenetic distribution, structure and function of fern hydathodes. METHODS: We performed a global review on fern hydathodes and their phylogenetic distribution, carried out an ancestral character state reconstruction, and studied the structure, guttation and elemental composition of salt residues of eight species, and the diurnal patterns of xylem pressure of two species. KEY RESULTS: Hydathodes are known from 1189 fern species, 92 genera and 19 families of 2 orders, Equisetales and Polypodiales. Stochastic character mapping indicated multiple gains and losses of hydathodes at the genus level, occurring especially during the last 50 million years of fern evolution. Hydathodes were located on the adaxial leaf surface and characterized by a cytoplasm-rich, pore-free epidermis, and became functional for several weeks after nearly complete leaf expansion. In two species, positive xylem pressure built up at night, potentially facilitating guttation. Guttation fluid was rich in Ca and often Si, but also contained P, Mg, Na and Al. CONCLUSIONS: Stochastic character mapping and the structural and functional diversity of hydathodes indicate multiple origins, and their presence/absence in closely related taxa implies secondary losses during fern evolution. Positive xylem pressure and high air humidity play an important role as drivers of guttation. Hydathodes may contribute to the regulation of leaf nutrient stoichiometry by the release of excessive compounds and minerals other than waste products, but the presence of essential chemical elements in salt residues also indicates possible leakage.

Nectaries in ferns: their taxonomic distribution, structure, function, and sugar composition.

PREMISE: Extrafloral nectaries have mainly been studied in angiosperms, but have also been reported in 39 fern species. Here we provide a global review of nectaries in ferns and examined their structure, function, and nectar sugar composition in two genera. METHODS: We searched in the literature and living plant collections of botanical gardens for indications of fern nectaries, observed nectar-feeding animals, studied the morphoanatomy in the two genera Aglaomorpha and Campyloneurum, and analyzed the total sugar concentrations and ratios of 16 species. Diurnal nectar release was observed with time-lapse photography. RESULTS: We found evidence for nectaries in 101 species of ferns from 11 genera and 6 families. Most of the nectary-bearing species were tree ferns (Cyatheaceae) and epiphytic ferns of the family Polypodiaceae. Nectaries consisted of cytoplasm-rich parenchyma with large nuclei and an epidermis with or without stomata, were attached to amphiphloic vascular bundles, and released nectar on the lower leaf surface mainly on expanding leaves during the night. Sugar concentrations varied between species (3.8-15.3%) but not between genera, and were sucrose-dominant (3 spp.), sucrose-rich (7), or hexose-rich (3). In the greenhouse, introduced ants, scale insects, and snails fed on the nectar. CONCLUSIONS: The wide taxonomic distribution, variable morphology, locations, and sugar compositions point to multiple evolutionary origins of fern nectaries. Nectar release in young leaves might attract mutualistic ants to protect leaves against herbivores only during this most vulnerable developmental stage. Even ex-situ, fern nectar is a valuable food source because it attracted several opportunistic animal species.

Can fern spores develop hydration memory in response to priming?

Fern spores and seeds initiate germination with fast water uptake, followed by a stationary phase with no appreciable water uptake and biochemical and metabolic processes that precede germination. After that, seed, germination is avoided by dehydration, as part of the priming treatments. After dehydration, seeds maintain their metabolic advances (hydration memory). As a result, rehydrated seeds germinate rapidly. We hypothesized that, as seeds, fern spores may be capable of developing hydration memory. To assess priming, spores of six fern species were exposed to: four or eight days of hydration in water (hydro-priming) or in a soil matrix (matrix-priming); or 1 month of hydration in the soil of the collection site (natural-priming). At the end of the treatments, the spores were dehydrated in the dark and germinated under laboratory conditions. Germination was evaluated using lag-time, germination rate and germination percentage. Priming treatments shortened lag time and/or increased germination rate or germination percentage in relation to the controls. Matrix-priming (8 days) reduced the spore germination percentage in three species. Our results provide evidence that fern spores possess a hydration memory that probably evolved in the soil bank and suggests that hydration-dehydration cycles within the natural soil might provide advantages for successful germination.

A Review of Galls on Ferns and Lycophytes.

Fern-insect interactions have not received the same attention as angiosperm-insect interactions have. It has even been stated that ferns may have very few interactions with animals because of their lack of flowers, fruits, and seeds. Consequently, for many decades fern-insect interactions have been overlooked and underestimated, especially for highly developed interactions such as those with gall-formers. The present work aims to review the galls of ferns and lycophytes worldwide, to provide an updated checklist including unpublished data and to estimate the global gall diversity of ferns and lycophytes. We recorded 93 host species, belonging to 41 genera. Galls were found in 20 fern families and one lycophyte family (Selaginellaceae). Most galls occur within the more derived ferns of the order Polypodiales, especially the fern families Polypodiaceae (21 host species), Dryopteridaceae (14 host species) and Athyriaceae (11 host species). Thirty-eight of the 133 gall morphotypes were induced by mites and 95 by insects of six orders (Coleoptera, Diptera, Hymenoptera, Lepidoptera, Thysanoptera, and Hemiptera). Among the insects, Cecidomyiidae (Diptera) caused most of the galls (35 morphotypes). So far, most galls have been reported from the Neotropical region (40 spp.) and Oriental region (28 spp.).

The evolution of aluminum accumulation in ferns and lycophytes.

PREMISE OF THE STUDY: This paper investigates the occurrence and evolution of aluminum (Al) accumulation within ferns and lycophytes, which is characterized by Al concentrations above 1000 mg·kg-1 in aboveground plant tissues. We hypothesize that this feature is more common in ferns than in angiosperms, and potentially correlated with growth form and other chemical elements. METHODS: Aluminum concentrations were obtained from novel analyses and literature for a total of 354 specimens and 307 species. Moreover, a semiquantitative aluminon test was applied for a subset of 105 species and validated against exact Al measurements. KEY RESULTS: Molecular phylogenetic analyses showed that the major Al-accumulating groups were primarily found in the Gleicheniales and Cyatheales, and largely absent in the Polypodiales. At the species and generic level, Al accumulation was typically either absent or present, and mixed results within a single species and genus were limited to less than 30% of the species and genera tested. Epiphytic ferns had significantly lower Al levels than terrestrial ferns, although this finding was not significant after phylogenetic correction. In addition, a significant, positive correlation was found between Al and iron, while Al was negatively correlated with phosphorus and potassium concentrations. CONCLUSIONS: Aluminum accumulation is most common outside of the Polypodiales and occurs in 38% of the species studied, indicating that this trait is indeed common within subtropical and tropical ferns, a finding that could be in line with their role as pioneer species on landslides and soils with high levels of soluble Al.