A Thorough Review on Ethnomedicinal Value of Bioactive Compounds of Pteridophytes against Cancer: Clinical Applications and Future Prospects.

Most cancers have become immune to normal cancer therapy, like chemotherapy and radiation. Therefore, exploring more effective and economical treatment options is important. Plants and herbs contain substances called phytochemicals, which have biological effects. Many phytochemicals having antioxidant and anticancer properties have been studied previously. There is increasing evidence that phytochemicals' anti-carcinogenic benefits originate from their ability to inhibit oxidation, inflammation, cell proliferation, and angiogenesis. These phytochemicals inhibit the spread of cancer by controlling the cell cycle and other molecular processes, such as metastasis. Along with therapeutic potential, other advantages, like their abundance, greater tolerability, and economic use, increase their utility in cancer therapeutics. In recent years, a number of scientists have examined lycophytes and ferns for their potential medicinal and phytochemical properties. This analysis emphasizes the significance of chemicals obtained from ferns and their derivatives in therapeutics. The authors discuss the pteridophyte's anti-cancer properties and other medical uses in this article. This information may help researchers in further research related to the most promising anticancer phytochemicals and their possibility as alternative drugs against cancer.

Insights into Chemical Diversity and Potential Health-Promoting Effects of Ferns.

The scientific community is focusing on how to enhance human health and immunity through functional foods, and dietary supplements are proven to have a positive as well as a protective effect against infectious and chronic diseases. Ferns act as a taxonomical linkage between higher and lower plants and are endowed with a wide chemical diversity not subjected to sufficient scrutinization before. Even though a wealth of traditional medicinal fern uses were recorded in Chinese medicine, robust phytochemical and biological investigations of these plants are lacking. Herein, an extensive search was conducted using the keywords ferns and compounds, ferns and NMR, ferns and toxicity, and the terms ferns and chemistry, lignans, Polypodiaceae, NMR, isolation, bioactive compounds, terpenes, phenolics, phloroglucinols, monoterpenes, alkaloids, phenolics, and fatty acids were utilized with the Boolean operators AND, OR, and NOT. Databases such as PubMed, Web of Science, Science Direct, Scopus, Google Scholar, and Reaxys were utilized to reveal a wealth of information regarding fern chemistry and their health-promoting effects. Terpenes followed by phenolics represented the largest number of isolated active compounds. Regarding the neuroprotective effects, Psilotium, Polypodium, and Dryopteris species possessed as their major phenolics component unique chemical moieties including catechins, procyanidins, and bioflavonoids. In this updated chemical review, the pharmacological and chemical aspects of ferns are compiled manifesting their chemical diversity in the last seven years (2017-2024) together with a special focus on their nutritive and potential health-promoting effects.

Lineage diversification and rampant hybridization among subspecies explain taxonomic confusion in the endemic Hawaiian fern Polypodium pellucidum.

PREMISE: Polypodium pellucidum, a fern endemic to the Hawaiian Islands, encompasses five ecologically and morphologically variable subspecies, suggesting a complex history involving both rapid divergence and rampant hybridization. METHODS: We employed a large target-capture data set to investigate the evolution of genetic, morphological, and ecological variation in P. pellucidum. With a broad sampling across five Hawaiian Islands, we deciphered the evolutionary history of P. pellucidum, identified nonhybrid lineages and intraspecific hybrids, and inferred the relative influence of geography and ecology on their distributions. RESULTS: Polypodium pellucidum is monophyletic, dispersing to the Hawaiian archipelago 11.53-7.77 Ma and diversifying into extant clades between 5.66 and 4.73 Ma. We identified four nonhybrid clades with unique morphologies, ecological niches, and distributions. Additionally, we elucidated several intraspecific hybrid combinations and evidence for undiscovered or extinct "ghost" lineages contributing to extant hybrid populations. CONCLUSIONS: We provide a foundation for revising the taxonomy of P. pellucidum to account for cryptic lineages and intraspecific hybrids. Geologic succession of the Hawaiian Islands through cycles of volcanism, vegetative succession, and erosion has determined the available habitats and distribution of ecologically specific, divergent clades within P. pellucidum. Intraspecific hybrids have likely arisen due to ecological and or geological transitions, often persisting after the local extinction of their progenitors. This research contributes to our understanding of the evolution of Hawai'i's diverse fern flora and illuminated cryptic taxa to allow better-informed conservation efforts.

Mycorrhizal association and its relation with pteridophytes.

Mycorrhizal association is one of the earliest and diversely distributed symbiotic associations on the Earth. This association helped early terrestrial plants to colonize the land by improved supply of nutrients like phosphate, nitrogen and zinc. It also helped plants to tolerate unfavorable soil conditions with increased water retention capacity, resistance to drought and pathogens. In return, fungi benefitted with carbon as their food source from the plants. More than 80% of terrestrial plants including pteridophytes, gymnosperms and angiosperms are reported to form arbuscular mycorrhizal (AM) association. Plants with root systems appeared on land during the Devonian period and many of them like pteridophytes still exist today. Various molecular and fossil studies confirm that the plants belonging to Ordovician-Devonian are associated with fungi, which are very similar to genus Glomus. AM association is very common in pteridophytes and the growth of its sporophyte and gametophyte is directly affected in the presence of AM association. Pteridophytes as early land plants with root systems have a very significant place in the plant kingdom. They have evolved and adapted to fill various habitats and facilitated early terrestrialization of other land plants by providing suitable niche with the help of AM fungi. In spite of pteridophytes being a very important plant group in the land system, very few reports are available on fungal-pteridophyte association. The present review is an effort to gather information about AM association in pteridophytes that might help in unraveling the evolution and significance of plant and fungi association.

An adventurous journey toward and away from fern apomixis: Insights from genome size and spore abortion patterns.

PREMISE: Apomixis in ferns is relatively common and obligatory. Sterile hybrids may restore fertility via apomixis at a cost of long-term genetic stagnation. In this study, we outlined apomixis as a possible temporary phase leading to sexuality and analyzed factors relating to transitioning to and away from apomixis, such as unreduced and reduced spore formation in apomict and apo-sex hybrid ferns. METHODS: We analyzed the genome size of 15 fern species or hybrids ("taxa") via flow cytometry. The number of reduced and unreduced gametophytes was established as a proxy for viable spore formation of either type. We also calculated the spore abortion ratio (sign of reduced spores) in several taxa, including the apo-sex hybrid Dryopteris × critica and its 16 apomictically formed offspring. RESULTS: Four of 15 sampled taxa yielded offspring variable in genome size. Specifically, each variable taxon formed one viable reduced plant among 12-451 sampled gametophytes per taxon. Thus, haploid spore formation in the studied apomicts was very rare but possible. Spore abortion analyses indicated gradually decreasing abortion (haploid spore formation) over time. In Dryopteris × critica, abortion decreased from 93.8% to mean 89.5% in one generation. CONCLUSIONS: Our results support apomixis as a transitionary phase toward sexuality. Newly formed apomicts hybridize with sexual relatives and continue to form haploid spores early on. Thus, they may get the genomic content necessary for regular meiosis and restore sexuality. If the missing relative goes extinct, the lineage gets locked into apomixis as may be the case with the Dryopteris affinis complex.

Chloroplast genome structure analysis of Equisetum unveils phylogenetic relationships to ferns and mutational hotspot region.

Equisetum is one of the oldest extant group vascular plants and is considered to be the key to understanding vascular plant evolution. Equisetum is distributed almost all over the world and has a high degree of adaptability to different environments. Despite the fossil record of horsetails (Equisetum, Equisetaceae) dating back to the Carboniferous, the phylogenetic relationship of this genus is not well, and the chloroplast evolution in Equisetum remains poorly understood. In order to fill this gap, we sequenced, assembled, and annotated the chloroplast genomes of 12 species of Equisetum, and compared them to 13 previously published vascular plants chloroplast genomes to deeply examine the plastome evolutionary dynamics of Equisetum. The chloroplast genomes have a highly conserved quadripartite structure across the genus, but these chloroplast genomes have a lower GC content than other ferns. The size of Equisetum plastomes ranges from 130,773 bp to 133,684 bp and they encode 130 genes. Contraction/expansion of IR regions and the number of simple sequences repeat regions underlie large genomic variations in size among them. Comparative analysis revealed we also identified 13 divergence hotspot regions. Additionally, the genes accD and ycf1 can be used as potential DNA barcodes for the identification and phylogeny of the genus Equisetum. Twelve photosynthesis-related genes were specifically selected in Equisetum. Comparative genomic analyses implied divergent evolutionary patterns between Equisetum and other ferns. Phylogenomic analyses and molecular dating revealed a relatively distant phylogenetic relationship between Equisetum and other ferns, supporting the division of pteridophyte into Lycophytes, Equisetaceae and ferns. The results show that the chloroplast genome can be used to solve phylogenetic problems within or between Equisetum species, and also provide genomic resources for the study of Equisetum systematics and evolution.

The traditional utilization, biological activity and chemical composition of edible fern species.

ETHNOPHARMACOLOGICAL RELEVANCE: Ferns form an important part of the human diet. Young fern fiddleheads are mostly consumed as vegetables, while the rhizomes are often extracted for starch. These edible ferns are also often employed in traditional medicine, where all parts of the plant are used, mostly to prepare extracts. These extracts are applied either externally as lotions and baths or internally as potions, decoctions and teas. Ailments traditionally treated with ferns include coughs, colds, fevers, pain, burns and wounds, asthma, rheumatism, diarrhoea, or skin diseases (eczema, rashes, itching, leprosy). AIM OF THE REVIEW: This review aims to compile the worldwide knowledge on the traditional medicinal uses of edible fern species correlating to reported biological activities and isolated bioactive compounds. MATERIALS AND METHODS: The articles and books published on edible fern species were searched through the online databases Web of Science, Pubmed and Google Scholar, with critical evaluation of the hits. The time period up to the end of 2022 was included. RESULTS: First, the edible fern species were identified based on the literature data. A total of 90 fern species were identified that are eaten around the world and are also used in traditional medicine. Ailments treated are often associated with inflammation or bacterial infection. However, only the most common and well-known fern species, were investigated for their biological activity. The most studied species are Blechnum orientale L., Cibotium barometz (L.) J. Sm., Diplazium esculentum (Retz.) Sw., Marsilea minuta L., Osmunda japonica Thunb., Polypodium vulgare L., and Stenochlaena palustris (Burm.) Bedd. Most of the fern extracts have been studied for their antioxidant, anti-inflammatory and antimicrobial activities. Not surprisingly, antioxidant capacity has been the most studied, with results reported for 28 edible fern species. Ferns have been found to be very rich sources of flavonoids, polyphenols, polyunsaturated fatty acids, carotenoids, terpenoids and steroids and most of these compounds are remarkable free radical scavengers responsible for the outstanding antioxidant capacity of fern extracts. As far as clinical trials are concerned, extracts from only three edible fern species have been evaluated. CONCLUSIONS: The extracts of edible fern species exert antioxidant anti-inflammatory and related biological activities, which is consistent with their traditional medicinal use in the treatment of wounds, burns, colds, coughs, skin diseases and intestinal diseases. However, studies to prove pharmacological activities are scarce, and require chemical-biological standardization. Furthermore, correct botanical classification needs to be included in publications to simplify data acquisition. Finally, more in-depth phytochemical studies, allowing the linking of traditional use to pharmacological relevance are needed to be done in a standardized way.

Genus Equisetum L: Taxonomy, toxicology, phytochemistry and pharmacology.

INTRODUCTION: The genus Equisetum (Equisetaceae) is cosmopolitan in distribution, with 41 recognized species. Several species of Equisetum are widely used in treating genitourinary and related diseases, inflammatory and rheumatic problems, hypertension, and wound healing in traditional medicine practices worldwide. This review intends to present information on the traditional uses, phytochemical components, pharmacological activities, and toxicity of Equisetum spp. and to analyze the new insights for further study. METHODS: Relevant literature has been scanned and collected via various electronic repositories, including PubMed, Science Direct, Google Scholar, Springer Connect, and Science Online, from 1960 to 2022. RESULTS: Sixteen Equisetum spp. were documented as widely used in traditional medicine practices by different ethnic groups throughout the world. A total of 229 chemical compounds were identified from Equisetum spp. with the major group of constituents being flavonol glycosides and flavonoids. The crude extracts and phytochemicals of Equisetum spp. exhibited significant antioxidant, antimicrobial, anti-inflammatory, antiulcerogenic, antidiabetic, hepatoprotective, and diuretic properties. A wide range of studies have also demonstrated the safety of Equisetum spp. CONCLUSION: The reported pharmacological properties of Equisetum spp. support its use in traditional medicine, though there are gaps in understanding the traditional usage of these plants for clinical experiments. The documented information revealed that the genus is not only a great herbal remedy but also has several bioactives with the potential to be discovered as novel drugs. Detailed scientific investigation is still needed to fully understand the efficacy of this genus; hence, very few Equisetum spp. were studied in detail for phytochemical and pharmacological investigation. Moreover, its bioactives, structure-activity connection, in vivo activity, and associated mechanism of action ought to be explored further.

Giant Fern Genomes Show Complex Evolution Patterns: A Comparative Analysis in Two Species of Tmesipteris (Psilotaceae).

Giant genomes are rare across the plant kingdom and their study has focused almost exclusively on angiosperms and gymnosperms. The scarce genetic data that are available for ferns, however, indicate differences in their genome organization and a lower dynamism compared to other plant groups. Tmesipteris is a small genus of mainly epiphytic ferns that occur in Oceania and several Pacific Islands. So far, only two species with giant genomes have been reported in the genus, T. tannensis (1C = 73.19 Gbp) and T. obliqua (1C = 147.29 Gbp). Low-coverage genome skimming sequence data were generated in these two species and analyzed using the RepeatExplorer2 pipeline to identify and quantify the repetitive DNA fraction of these genomes. We found that both species share a similar genomic composition, with high repeat diversity compared to taxa with small (1C < 10 Gbp) genomes. We also found that, in general, characterized repetitive elements have relatively high heterogeneity scores, indicating ancient diverging evolutionary trajectories. Our results suggest that a whole genome multiplication event, accumulation of repetitive elements, and recent activation of those repeats have all played a role in shaping these genomes. It will be informative to compare these data in the future with data from the giant genome of the angiosperm Paris japonica, to determine if the structures observed here are an emergent property of massive genomic inflation or derived from lineage specific processes.

Evolution of genome space occupation in ferns: linking genome diversity and species richness.

BACKGROUND AND AIMS: The dynamics of genome evolution caused by whole genome duplications and other processes are hypothesized to shape the diversification of plants and thus contribute to the astonishing variation in species richness among the main lineages of land plants. Ferns, the second most species-rich lineage of land plants, are highly suitable to test this hypothesis because of several unique features that distinguish fern genomes from those of seed plants. In this study, we tested the hypothesis that genome diversity and disparity shape fern species diversity by recording several parameters related to genome size and chromosome number. METHODS: We conducted de novo measurement of DNA C-values across the fern phylogeny to reconstruct the phylogenetic history of the genome space occupation in ferns by integrating genomic parameters such as genome size, chromosome number and average DNA amount per chromosome into a time-scaled phylogenetic framework. Using phylogenetic generalized least square methods, we determined correlations between chromosome number and genome size, species diversity and evolutionary rates of their transformation. KEY RESULTS: The measurements of DNA C-values for 233 species more than doubled the taxon coverage from ~2.2 % in previous studies to 5.3 % of extant diversity. The dataset not only documented substantial differences in the accumulation of genomic diversity and disparity among the major lineages of ferns but also supported the predicted correlation between species diversity and the dynamics of genome evolution. CONCLUSIONS: Our results demonstrated substantial genome disparity among different groups of ferns and supported the prediction that alterations of reproductive modes alter trends of genome evolution. Finally, we recovered evidence for a close link between the dynamics of genome evolution and species diversity in ferns for the first time.

A comprehensive review on Nepalese wild vegetable food ferns.

Ferns are used as traditional and fascinating foods in many countries. They are also considered to possess important ethnomedicinal values; however, ferns are one of the underutilized plant resources by both scientific and local communities. Pharmagonostical studies reveal that ferns and fern-allies possess several biological activities including antibacterial, antiviral, antifungal, antimalarial, antidiarrheal, anthelmintic, analgesic, anti-inflammatory, antidiabetic, anticancer, neuroprotective, nephroprotective, hepatoprotective, antifertility, etc. Flavonoids and terpenoids are major secondary metabolites present in ferns. Ugonins, particularly isolated from Helminthostachys zeylanica, have found diverse bioactivities. Ptaquiloside, a norsesquiterpene glucoside, found in Pteridium revolutum, Dryopteris cochleata and Polystichum squarrosum, is one of the hazardous metabolites of ferns which is responsible for the toxic effect. Alkaloids are reported to be present in the ferns; however, the qualitative data are uncertain. Some fern metabolites, such as cyanogenic glycosides and terpenoids, are considered to possess defensive activity against animal attacks. Some ferns are also used for manuring as biological alternative to pesticides. Nepalese have consumed at least 33 species of ferns and fern-allies belonging to 13 families, 20 genera as cooked vegetable foods. The aim of this review is compilation of information available on their distribution, ethnomecinal values, pharmcognosy, pharmacology and phytochemistry.

(Fern flora of Viçosa, Minas Gerais State, Brazil: Dennstaedtiaceae, Lindsaeaceae and Saccolomataceae / Flora de samambaias de Viçosa, Estado de Minas Gerais, Brasil: Dennstaedtiaceae, Lindsaeaceae and Saccolomataceae

As part of an ongoing project treating the ferns and lycophytes from the region of Viçosa, Minas Gerais State, Brazil, we here present the taxonomic treatment of the early divergent lineages of the leptosporangiate ferns: the families Dennstaedtiaceae, Lindsaeaceae and Saccolomataceae. We have been sampling the remnant forest patches since 2012; we also fully reviewed the collection of herbarium VIC and other online collections: F, IAN, NY, PH, RB, U, UC, UPCB, US, and WTU. In the region of Viçosa, six taxa belonging to those families occur Dennstaedtia cicutaria and Pteridium esculentum subsp. arachnoideum (Dennstaedtiaceae); Lindsaea lancea var. lancea, L. quadrangularis subsp. quadrangularis, and L. stricta var. stricta (Lindsaeaceae); and Saccoloma elegans (Saccolomataceae). Among these taxa, only L. quadrangularis subsp. quadrangularis and S. elegans are endemic to the Brazilian Atlantic Rainforest; the remaining are widespread in the Neotropics. We present keys, descriptions, illustrations, examined specimens, and comments.

Como parte de um projeto em andamento sobre samambaias e licófitas da região de Viçosa, Estado de Minas Gerais, Brasil, o tratamento taxonômico das primeiras linhagens divergentes de samambaias leptosporangiadas: as famílias Dennstaedtiaceae, Lindsaeaceae e Saccolomataceae são apresentadas. Expedições de campo foram realizadas desde 2012 em remanescentes florestais da região. Também foi realizada uma revisão completa da coleção do herbário VIC, cuja data de início é 1930, bem como de outras coleções online de herbários: F, IAN, NY, PH, RB, U, UC, UPCB, US, and WTU. Na região de Viçosa, existem seis táxons pertencentes a essas famílias: Dennstaedtia cicutaria e Pteridium esculentum subsp. arachnoideum (Dennstaedtiaceae); Lindsaea lancea var. lancea, L. quadrangularis subsp. quadrangularis e L. stricta var. stricta (Lindsaeaceae); e Saccoloma elegans (Saccolomataceae). Dentre esses táxons, apenas L. quadrangularis subsp. quadrangularis e S. elegans são endêmicos da Mata Atlântica brasileira; as demais estão amplamente distribuídas na região Neotropical. São apresentadas chaves, descrições, ilustrações, materiais examinados e comentários.

The relationship between chlorophyllous spores and mycorrhizal associations in ferns: evidence from an evolutionary approach.

PREMISE: Approximately 14% of all fern species have physiologically active chlorophyllous spores that are much more short-lived than the more common and dormant achlorophyllous spores. Most chlorophyllous-spored species (70%) are epiphytes and account for almost 37% of all epiphytic ferns. Chlorophyllous-spored ferns are also overrepresented among fern species in habitats with waterlogged soils, of which nearly 60% have chlorophyllous spores. Ferns in these disparate habitat types also have a low incidence of mycorrhizal associations. We therefore hypothesized that autotrophic chlorophyllous spores represent an adaptation of ferns to habitats with scarce mycorrhizal associations. METHODS: We evaluated the coevolution of chlorophyllous spores and mycorrhizal associations in ferns and their relation to habitat type using phylogenetic comparative methods. RESULTS: Although we did not find support for the coevolution of spore type and mycorrhizal associations, we did find that chlorophyllous spores and the absence of mycorrhizal associations have coevolved with epiphytic and waterlogged habitats. Transition rates to epiphytic and waterlogged habitats were significantly higher in species with chlorophyllous spores compared to achlorophyllous lineages. CONCLUSIONS: Spore type and mycorrhizal associations appear to play important roles in the radiation of ferns into different habitat types. Future work should focus on clarifying the functional significance of these associations.

Origins of the seed: The "golden-trio hypothesis".

The seed is an evolutionary innovation in the plant kingdom. While human civilization depends heavily on seed production, how the seed trait emerged remains elusive. In this opinion article, a "golden-trio hypothesis" is proposed based on our investigations of LEC1 gene functions in Adiantum capillus-veneris. This hypothesis posits that a "seed program" arose from spatiotemporal integration of three key components: assimilate flow, ABA-mediated stress responses, and stress-induced LEC1 expression. Thus, the evolutionary innovation of seeds should be considered not a simple event resulting from new genes; rather, it represents the outcome of a series of physiological and morphological innovations that emerged prior to and regardless of the origin of the seed program. This new perspective could help us tackle some long-standing questions around the puzzling origin of seeds.

Deep vicariance and frequent transoceanic dispersal shape the evolutionary history of a globally distributed fern family.

PREMISE: The historical biogeography of ferns is typically expected to be dominated by long-distance dispersal due to their minuscule spores. However, few studies have inferred the historical biogeography of a large and widely distributed group of ferns to test this hypothesis. Our aims were to determine the extent to which long-distance dispersal vs. vicariance have shaped the history of the fern family Blechnaceae, to explore ecological correlates of dispersal and diversification, and to determine whether these patterns differ between the northern and southern hemispheres. METHODS: We used sequence data for three chloroplast loci to infer a time-calibrated phylogeny for 154 of 265 species of Blechnaceae, including representatives of all genera in the family. This tree was used to conduct ancestral range reconstruction and stochastic character mapping, estimate diversification rates, and identify ecological correlates of diversification. RESULTS: Blechnaceae originated in Eurasia and began diversifying in the late Cretaceous. A lineage comprising most extant diversity diversified principally in the austral Pacific region around the Paleocene-Eocene Thermal Maximum. Land connections that existed near the poles during periods of warm climates likely facilitated migration of several lineages, with subsequent climate-mediated vicariance shaping current distributions. Long-distance dispersal is frequent and asymmetrical, with New Zealand/Pacific Islands, Australia, and tropical America being major source areas. CONCLUSIONS: Ancient vicariance and extensive long-distance dispersal have shaped the history of Blechnaceae in both the northern and southern hemispheres. The exceptional diversity in austral regions appears to reflect rapid speciation in these areas; mechanisms underlying this evolutionary success remain uncertain.

Exploring the High Variability of Vegetative Desiccation Tolerance in Pteridophytes.

In the context of plant evolution, pteridophytes, which is comprised of lycophytes and ferns, occupy an intermediate position between bryophytes and seed plants, sharing characteristics with both groups. Pteridophytes is a highly diverse group of plant species that occupy a wide range of habitats including ecosystems with extreme climatic conditions. There is a significant number of pteridophytes that can tolerate desiccation by temporarily arresting their metabolism in the dry state and reactivating it upon rehydration. Desiccation-tolerant pteridophytes exhibit a strategy that appears to be intermediate between the constitutive and inducible desiccation tolerance (DT) mechanisms observed in bryophytes and angiosperms, respectively. In this review, we first describe the incidence and anatomical diversity of desiccation-tolerant pteridophytes and discuss recent advances on the origin of DT in vascular plants. Then, we summarize the highly diverse adaptations and mechanisms exhibited by this group and describe how some of these plants could exhibit tolerance to multiple types of abiotic stress. Research on the evolution and regulation of DT in different lineages is crucial to understand how plants have adapted to extreme environments. Thus, in the current scenario of climate change, the knowledge of the whole landscape of DT strategies is of vital importance as a potential basis to improve plant abiotic stress tolerance.

Taxonomic importance of spore morphology of selected taxa of Asplenium (Aspleniaceae) from Pakistan.

Taxonomic importance of spore morphology of seven taxa of genus Asplenium (Aspleniaceae) from Pakistan is illustrated with SEM. Spore morphology of the taxa were monolete with bilateral symmetry, distal and proximal view were ellipsoidal, spheroidal, equatorial view were hemicircular; perispore surface were alate, costate, costate-alate, with reticulate and tuberculate ornamentation. Taxonomic importance of spore morphology of seven taxa of Asplenium (Aspleniaceae, polypodiales) from Pakistan. Basionym, Synonyms with updated nomenclature. Spores are monolete, distal and proximal view ellipsoidal, spheroidal, equatorial view hemicircular, perispore with alate, costate, reticulate, and tuberculate surface. Habitat ecology with specimens examined from different areas of Pakistan and their worldwide general distribution.

Ferns and lycophytes in coal mining waste and tailing landfills.

Mineral coal extraction in Santa Catarina State (Brazil) Carboniferous Basin has degraded the local ecosystem, restricting the use of its areas. One of the biggest environmental impacts in the mining areas is the uncontrolled disposal of waste and sterile mining with high concentrations of pyrite, which in the presence of air and water is oxidized promoting the formation of acid mine drainage (AMD). These contaminants can be leached into water resources, restrict the use of water and soil, and cause threats to fauna and flora. This study aimed to characterize these areas as to the content of Cd, Pb, Ni and Zn metals in the tailings and waste resulting from coal mining and to survey the species of ferns and lycophytes present. Wastes and tailing samples and specimens of ferns and lycophytes were collected in 23 landfills in six municipalities in the region and in four underlying areas used as controls. Chemical and physical analyses (pH in water and pH in KCl, Ca, Mg, P, K, Na, Mn, Fe, Al, clay and OM contents) were carried out and the total contents of heavy metals Cd, Pb, Ni and Zn were determined. Sampling of ferns and lycophytes was carried out by walking. The levels of heavy metals, Cd, Ni and Zn, were below the prevention concentrations established by CONAMA Resolution 420/2009. Pb levels were above prevention values in four landfills. Sixteen species of ferns and one lycophyte were found, with hemicryptophytes the most frequent and helophytes the most adapted to the environment. Of the species found, Pteridium esculentum (G. Forst.) Cockayne, Pityrogramma calomelanos (L.) Link and Telmatoblechnum serrulatum (Rich.) Perrie, DJ Ohlsen & Brownsey demonstrated resistance to degraded and contaminated environments with Pb, which may constitute an alternative for project monitoring and environmental recovery.

Apomictic fern fathers: an experimental approach to the reproductive characteristics of sexual, apomict, and hybrid fern gametophytes.

PREMISE: Apomixis and hybridization are two essential and complementary factors in the evolution of plants, including ferns. Hybridization combines characteristics from different species, while apomixis conserves features within a lineage. When combined, these two processes result in apo-sex hybrids. The conditions leading to the formation of these hybrids are poorly understood in ferns. METHODS: We cultivated spores from 66 fern samples (43 apomicts, 7 apo-sex hybrids, and 16 sexuals), and measured their development in vitro over 16 weeks. We evaluated germination, lateral meristem formation rates, sexual expression, and production of sporophytes and then compared ontogenetic patterns among the three groups. RESULTS: The three examined groups formed antheridia (male gametangia) but differed in overall gametophyte development. Sexual species created archegonia (female, 86% of viable samples), but no sporophytes. Apomicts rarely created nonfunctional archegonia (8%) but usually produced apogamous sporophytes (75%). Surprisingly, apomictic and sexual species showed similar development speed. The sexually reproducing parents of viable studied hybrids formed about twice as many meristic gametophytes as the apomictic parents (39% vs. 20%, respectively). CONCLUSIONS: We present the most thorough comparison of gametangial development of sexual and apomictic ferns, to date. Despite expectations, apomictic reproduction might not lead to earlier sporophyte formation. Apomicts produce functional sperm and thus can contribute this type of gamete to their hybrids. The development patterns found in the parents of hybrids indicate a possible increase of hybridization rates by antheridiogens. The apo-sex hybrids always inherit the apomictic reproductive strategy and are thus capable of self-perpetuation.

Ants associated with fronds of the tropical bracken fern Pteridium esculentum subsp. arachnoideum / Formigas associadas às frondes da samambaia Pteridium esculentum subsp. arachnoideum

Abstract: Pteridium is a cosmopolitan genus of ferns that possess nectaries on its fronds (fern leaves), thereby attracting ants. Foliar (or extrafloral) nectaries are nectar-producing glands that are not related to pollination, but rather attract ants and other arthropods. Foliar nectaries are found in 101 fern species, belonging to 11 genera and six families. The aim of the study is to characterize the community of ants that visit the fronds of Pteridium esculentum subsp. arachnoideum, as well as daily and seasonal ant abundance in different frond development stages. The study was conducted in the Atlantic Forest of Rio de Janeiro state, Brazil. Bimonthly collections were established, where 30 expanding fronds and 30 fully expanded fronds were randomly marked. In each 1-hour shift starting at 8:30 am and ending at 5:30 pm, the fronds were observed for the presence of ants. Thirty three ant species were recorded on the Pteridium esculentum subsp. arachnoideum fronds, distributed into six subfamilies and 13 genera. The most abundant species were Solenopsis sp.1 and Ectatomma tuberculatum. Eight ant species were observed foraging the nectaries of tropical bracken fern fronds. Ectatomma tuberculatum has been observed feeding on the nectaries and patrolling the fronds. Ant activity peak was on mid-day during the rainy season. The tropical bracken fern Pteridium esculentum subsp. arachnoideum has a rich (the highest recorded until now on Pteridium species) and diverse ant community on its fronds, mainly on the expanding fronds. The presence of generalist predatory ants (Ectatomma tuberculatum and Solenopsis sp.1) during the entire study period suggests a positive interaction between ants and Pteridium esculentum subsp. arachnoideum.

Resumo: Pteridium é um gênero cosmopolita de samambaias que possui nectários em suas frondes (folhas de samambaias), atraindo formigas. Nectários foliares (ou extraflorais) são glândulas produtoras de néctar que não estão relacionadas com a polinização, mas podem atrair formigas e outros artrópodes. Nectários foliares já foram registrados em 101 espécies de samambaias, pertencentes a 11 gêneros e seis famílias. O objetivo do estudo é caracterizar a comunidade de formigas que visitam as frondes de Pteridium esculentum subsp. arachnoideum, bem como a abundância diária e sazonal das formigas em diferentes estágios foliares. O estudo foi realizado na Mata Atlântica do estado do Rio de Janeiro, Brasil. Foram estabelecidas coletas bimestrais, onde foram marcadas aleatoriamente 30 frondes em expansão e 30 frondes totalmente expandidas. Em cada turno de 1 hora com início às 8h30 e término às 17h30, as frondes marcadas foram observadas quanto à presença de formigas. Trinta e três espécies de formigas foram registradas nas frondes do Pteridium esculentum subsp. arachnoideum, distribuídas em seis subfamílias e 13 gêneros. As espécies mais abundantes foram Solenopsis sp.1 e Ectatomma tuberculatum. Foram registradas oito espécies de formigas forrageando os nectários foliares da samambaia. Ectatomma tuberculatum foi observada se alimentando nos nectários e patrulhando as frondes. As formigas tiveram o pico de atividade ao meio-dia e na estação chuvosa. A samambaia tropical Pteridium esculentum subsp. arachnoideum tem uma elevada riqueza (a maior já registrada até o momento para espécies de Pteridium) e diversidade de formigas em suas frondes, com maior frequência nas frondes não totalmente expandidas. A presença de formigas predadoras generalistas (Ectatomma tuberculatum e Solenopsis sp.1) durante todo o período de estudo sugere uma interação positiva entre as formigas e Pteridium esculentum subsp. arachnoideum.