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Thermal variation over space and time shapes global patterns of biodiversity and responses of species to climate change. Vertical microclimate gradients from the understory to the canopy mediate biogeographic patterns of arboreal species across larger spatial scales. Yet, the magnitude to which microhabitats in the canopy, such as large epiphytes, alter the exposure of arboreal species to hot and variable canopy conditions across biogeographic gradients is not well known. We first examine the extent to which maximum temperatures and temperature variation in epiphytic bird's nest ferns (BNFs) and the understory are lower than those in the canopy across an elevation gradient in the Philippines and the environmental factors influencing these differences. We find that differences in maximum temperatures (i.e., offset) and temperature variation (i.e., buffering) between the canopy and BNFs are greater than differences between the canopy and understory. Offset and buffering are largest when canopy temperatures are high and are positively impacted by precipitation. Furthermore, we address the theory that exposure to higher thermal variation in the canopy may increase the range sizes of arboreal species. We find that lower thermal variation over time in BNFs reduces the thermal overlap between low and high elevations relative to canopy temperatures. Arboreal species that shelter in BNFs during the day may therefore have narrow physiological tolerances and consequently smaller elevational ranges. By providing thermal regimes more similar to the understory than the canopy, BNFs may therefore effectively function as "arboreal soil", mediating impacts of arboreality on patterns of biodiversity and trait variation. [ABSTRACT FROM AUTHOR]

Aspleniaceae is a diverse fern family with around 730 species distributed across various climates. Historically, its genera were differentiated by leaf morphology, but molecular studies revealed inconsistencies with this method, leading to a revised classification into two genera. Anatomical studies, particularly of petiole color and vascular bundles, offer insight into phylogenetic relationships. Our research examines petiole anatomy to understand its link to classification and phylogeny, and to investigate the functional structures of Aspleniaceae species, thus contributing to a deeper understanding of their diversity and evolution. The present study analyzed 49 Aspleniaceae species, previously categorized into 9 genera, collected from the field and botanic gardens. Sections were stained with safranin O-fast green and examined with various microscopes to assess anatomical structures, contributing to the understanding of Aspleniaceae phylogeny. We identified three petiole types based on vascular and cellular structure. Hymenasplenium species shared common traits such as cell wall thickness in the epidermis, while Asplenium groups exhibited variation in the epidermal and hypodermal cell wall composition affecting petiole color. This research contributes to understanding phylogenetic relationships within Aspleniaceae, providing a detailed anatomical basis for classification. [ABSTRACT FROM AUTHOR]

Phenolic compounds possess the potential to function as antioxidants, hence the growing interest in ethnomedicine to use them for treatment or prevention of diseases associated with oxidative stress. This study measured the total phenolics and flavonoid contents (TPC and TFC) and in vitro antioxidant activity of Asplenium platyneuron extract and its fractions. The methanol extract was partitioned successively in n-hexane, ethyl acetate (EtOAc) and n-butanol. Thereafter, the EtOAc soluble was fractionated using vacuum liquid chromatography (VLC) in gradients of n-hexane/EtOAc and dichloromethane/methanol. The free radical scavenging activity was determined by using diphenyl-picrylhydrazyl (DPPH), and the reducing power by ferric reducing antioxidant power (FRAP) and the total antioxidant capacity assays. The TPC of EtOAc fraction was 9.31 ± 0.76 mg GAE/g extract compared with 1.33 ± 0.61 mg GAE/g extract of n-hexane fraction. The TFC for EtOAc was 139.73 ± 8.03 mg QE/g extract higher than 35.73 ± 15.01 mg QE/g extract of n-butanol fraction. The ethylacetate fraction exhibited the highest DPPH free radical scavenging capacity of 80.50%, and FRAP of 413.00 ± 1.28 mg GAE/g extract with IC50 of 1.31 ± 0.68mg/mL. There was a strong correlation between the antioxidant activity and the TFC/TPC. The VLC fractions, AP2 and AP7 (IC50 1.43 ± 1.34 and 1.41± 1.11 mg/mL respectively) elicited antioxidant activity higher than other fractions. Dereplication analysis identified three phenolics; apigenin 6-C-hexoside-8-C-pentoside, kaempferol 3-O-caffeoylsophoroside and schaftoside from A. platyneuron. The high TPC/TFC and its strong antioxidant properties makes A. platyneuron a promising source of novel lead antioxidants. [ABSTRACT FROM AUTHOR]

Abstract A new species of Asplenium L. (Aspleniaceae) is described and illustrated. Asplenium sylvaticum is endemic of the Yungas and Paraná biogeographic provinces in the Southern Cone. The main diagnostic characters for this taxon are: apical pinnae with one to four caudate lobes at the base, similar in size to lateral pinnae, smooth pinnate blades with 18-36 pairs of caudate pinnae and spores with reticulate perispore, winged folds with scarce equinulaes on the margins and the laesurae wide with smooth margin. The new species belongs to the A. serra species complex group, characterised by xeromorphic habit, creeping rhizome and coriaceous fronds with branched scales on both surfaces, and resembles the other species inhabiting in Argentina, A. achalense and A. serra, in overall leaf architecture and shape but differs of the aforementioned species (and the other species of the A. serra complex) by the unique combination of characters of its rhizomatic scales, fronds and spores. A key with the morphological and palynological characters that differentiate A. sylvaticum from the other taxa belonging to the A. serra complex in the South American Cone and Bolivia is given.

Abstract We present a floristic-taxonomic survey of the species of Aspleniaceae in Mato Grosso as part of a series on the flora of lycophytes and ferns in this state. The study is based on an analysis of national and foreign herbaria specimens. In the state of Mato Grosso, there are two genera and 22 species of Aspleniaceae: Asplenium abscissum, A. angustum, A. auritum, A. cirrhatum, A. claussenii, A. cristatum, A. dimidiatum, A. flabellulatum, A. formosum, A. hallii, A. inaequilaterale, A. juglandifolium, A. otites, A. poloense, A. praemorsum, A. pumilum, A. resiliens, A. salicifolium, A. serra, A. serratum, A. stuebelianum, and Hymenasplenium delitescens. This corresponds to 27,5% of the Aspleniaceae species reported for Brazil. The species predominantly occur in forest environments. Dichotomous keys to the genera and species, descriptions, diagnostic illustrations, and notes on the geographic distribution, habitat, and morphological affinities between species are provided.

Background and Aims Substrate preferences are often treated as species traits and are used to distinguish different habits, i.e. an epiphytic, lithophytic or terrestrial habit. Such a categorization, however, ignores substantial intraspecific variation. An approach that takes biological variability within a species into account is needed. Methods We focused on four large genera of ferns and lycophytes and found relevant information in >500 sources, such as online databases, checklists, floras and species descriptions. Translating textual information into a quantitative index, we quantified the propensity to grow on either substrate as a continuous trait for 1475 species. Key Results Only a minority of species exhibited strict substrate fidelity, but a majority of them showed clear habitat preferences. The relative frequencies of intermediates between strict lithophytes, epiphytes and terrestrials does not support the frequent notion of ecological similarity of the lithophytic and epiphytic habitat. Conclusions The compiled data are useful immediately for ecological and evolutionary studies with the focal taxa. More importantly, we propose the replacement of the concept of distinct habits with one of gradual differences. This should have a profound impact on any such study with plants in general. [ABSTRACT FROM AUTHOR]

The biodiversity hotspot of central Chile is home to a high proportion of endemic species, but some of these species are inconspicuous and not easily observed. During a botanical exploration in the Los Queules National Reserve (Chile), a population of Asplenium trilobum Cav. was identified. The plants were found growing on the bark of a Myrceugenia parvifolia (DC.) Kausel tree in a small swamp next to specimens of Drimys winteri J.R.Forst. & G.Forst. (35°59'11.84"S; 72°41'11.53"W). Several previously unrecorded species were found, including Carex cf. excelsa Poepp. ex Kunth, Chusquea cf. quila Kunth, Ercilla cf. spicata (Bertero) Moq., and Boquila trifoliolata (DC.) Decne., highlighting the importance of exploring and documenting this biodiversity hotspot. The discovery in this wilderness area extends the distribution 86 km north on the continent, which was previously limited to the east of the municipality of Penco in the Biobío region (36°44'9.26"S; 72°57'42.5"W). This paper presents an observed specimen, its locality, and associated species. [ABSTRACT FROM AUTHOR]

We present a floristic-taxonomic survey of the species of Aspleniaceae in Mato Grosso as part of a series on the flora of lycophytes and ferns in this state. The study is based on an analysis of national and foreign herbaria specimens. In the state of Mato Grosso, there are two genera and 22 species of Aspleniaceae: Asplenium abscissum, A. angustum, A. auritum, A. cirrhatum, A. claussenii, A. cristatum, A. dimidiatum, A. flabellulatum, A. formosum, A. hallii, A. inaequilaterale, A. juglandifolium, A. otites, A. poloense, A. praemorsum, A. pumilum, A. resiliens, A. salicifolium, A. serra, A. serratum, A. stuebelianum, and Hymenasplenium delitescens. This corresponds to 27,5% of the Aspleniaceae species reported for Brazil. The species predominantly occur in forest environments. Dichotomous keys to the genera and species, descriptions, diagnostic illustrations, and notes on the geographic distribution, habitat, and morphological affinities between species are provided. [ABSTRACT FROM AUTHOR]

Asplenium (Aspleniaceae) is taxonomically one of the most complex fern genera with about 700 species, nearly cosmopolitan in distribution. In Argentina, Asplenium constitutes the most speciesrich fern genus, with 38 taxa. Among these species, the delimitation of Asplenium achalense remain poorly understood. Some authors consider A. serra as a variant of A. achalense, or even the last species has been considered as a complex of species. The finding of a population of Asplenium achalense in Yungas after three decades without observations in the wild and neither registering more recently herbarium specimens, made it possible to analyse living material and observe new diagnostic characters present in rachises and laminae scales, glandular hairs in petioles and rachises along with spore wall features, that are critical to delimitate A. achalense as a well-defined taxon, endemics of north-west to central Argentina. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Asplenium platyneuron growing at two locations (outer sunlit location and inner shaded location) in a low brick and masonry wall in upper Manhattan (New York City) was studied to assess the effects of the two different microenvironments on the ecophysiology of the two fern populations. The intensity of illumination at the inner location was ca. 40% of the intensity at the outer location, where the photosynthetic active radiation (PAR) at mid-day was ca. 1,484 µmol m–2 s–1. Generally, root space temperatures tracked air temperature, varying from 26 to 31 °C for the outer location, and 25 to 28.7 °C for the inner location. Based on prior published literature with other fern species, we hypothesized that samples of A. platyneuron growing on the outer surface exposed to maximum illumination would exhibit higher maximum photosynthesis rates, higher respiration in the light, and smaller leaves compared to the less illuminated samples growing on the inner surface. The results supported our hypothesized relationships. Mean leaf length (cm) was longer for leaves of plants at the shaded inner location (30 ± 1.1) compared to those of plants at the more illuminated outer location (9 ± 1.2). The maximum photosynthetic rate (Amax) for the outside sample of ferns (6.3 ± 0.25) was statistically significantly different from the Amax for the inside sample (4.3 ± 0.72) as hypothesized. The mean respiration in the light was significantly larger for the ferns growing outside (-0.52 ± 0.03) compared to those inside (-0.36 ± 0.04). Additionally, light response (A–Q) curves and relevant other physiological evidence are presented for ferns at the two sampling sites. [ABSTRACT FROM AUTHOR]

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Background and Objectives: Asplenium incisum, a natural plant, is known to possess numerous pharmacological and biochemical properties. However, the inhibitory effect of A. incisum against Porphyromonas gingivalis and other factors related to periodontal disease have not yet been demonstrated. This study aimed to investigate the potential of A. incisum extract as a phytotherapeutic candidate for improving periodontal diseases by assessing its antibacterial, anti-inflammatory, and anti-osteoclastogenic activities. Materials and Methods: The inhibition of proliferation of P. gingivalis by A. incisum and the sustainability of its antibacterial activity were evaluated in this study. The production of inflammatory cytokines (tumor necrosis factor-&@945; (TNF-&@945;) and interleukin-6 (IL-6)) and nitric oxide (NO) from lipopolysaccharide-stimulated RAW264.7 cells was assessed using an enzyme-linked immunosorbent assay. To identify the anti-osteoclastogenic activity, tartrate-resistant acid phosphatase (TRAP) staining and TRAP activity analyses were performed on bone marrow macrophages. Results: The proliferation of P. gingivalis was significantly inhibited by A. incisum (p < 0.001), and the antibacterial activity was sustained for up to 3 days. A. incisum showed antiinflammatory activities by significantly decreasing the release of TNF-&@945;, IL-6 (p < 0.05), and NO (p < 0.01). In addition, A. incisum significantly suppressed TRAP-positive cells and TRAP activity (at 30 μg/mL, p < 0.01) without causing any cytotoxicity (p > 0.05). Conclusions: A. incisum showed antibacterial, anti-inflammatory, and anti-osteoclastogenic activities, suggesting it has strong therapeutic potential against periodontal diseases. [ABSTRACT FROM AUTHOR]

The complete plastome sequence of Asplenium × bimixtum (Aspleniaceae) was determined. The whole genome is 153,098 bp in length, consisting of large (86,394 bp) and small (21,508 bp) single-copy regions, and a pair of identical inverted repeats (22,598 bp). The genome contains 88 protein-coding genes, 33 tRNA genes, 8 rRNA genes, and 1 pseudogene (infA). The phylogenetic analysis based on 84 protein-coding genes showed that Asplenium is monophyletic with high bootstrap value.

While most ferns avoid freezing as they have a tropical distribution or shed their fronds, wintergreen species in temperate and boreoalpine ecosystems have to deal with sub-zero temperatures. Increasing evidence has revealed overlapping mechanisms of desiccation and freezing tolerance in angiosperms, but the physiological mechanisms behind freezing tolerance in ferns are far from clear. We evaluated photochemical and hydraulic parameters in five wintergreen fern species differing in their ability to tolerate desiccation. We assessed frond freezing tolerance, ice nucleation temperature and propagation pattern, and xylem anatomical traits. Dynamics of photochemical performance and xanthophyll cycle were evaluated during freeze–thaw events under controlled conditions and, in selected species, in the field. Only desiccation-tolerant species, which possessed a greater fraction of narrow tracheids (<18 μm) than sensitive species, tolerated freezing. Frond freezing occurred in the field at –3.4 ± 0.9 °C (SD) irrespective of freezing tolerance, freezable water content, or tracheid properties. Even in complete darkness, maximal photochemical efficiency of photosystem II was down-regulated concomitantly with zeaxanthin accumulation in response to freezing. This was reversible upon re-warming only in tolerant species. Our results suggest that adaptation for freezing tolerance is associated with desiccation tolerance through complementary xylem properties (which may prevent risk of irreversible cavitation) and effective photoprotection mechanisms. The latter includes de-epoxidation of xanthophylls in darkness, a process evidenced for the first time directly in the field. [ABSTRACT FROM AUTHOR]

The Asplenium pekinense complex mainly comprises one diploid, A. sarelii Hook. (rare), one autotetraploid, A. pekinense Hance (best known and very common), and shares two allotetraploids, A. anogrammoides Christ (common but often misidentified) and A. altajense (Komarov) Grubov (rare and endemic) with the A. varians complex. The latter is further constituted by two diploids, A. tenuicaule Hayata (widespread) and A. semivarians Viane & Reichstein (rare), as well as other three tetraploids, A. kansuense Ching (barely known), A. varians Wallich ex Hooker & Greville (well‐known, relatively common, and morphologically variable), and A. kukkonenii Viane & Reichstein (rare and often misidentified). These two species complexes are notorious for their taxonomic difficulty based on general morphology, which is mainly caused by their history of reticulate evolution. Here, we collected most species within the two complexes, and obtained ploidy information by spore size measurement and flow cytometry investigation. Phylogenetic analyses using DNA markers representing maternally inherited chloroplast and biparentally inherited nuclear genomes helped to reconstruct the reticulate evolution history. The present results support previous hypotheses that A. sarelii is the ancestor of both A. pekinense and A. anogrammoides, as well as that A. tenuicaule is the common progenitor of A. anogrammoides, A. varians, and A. kukkonenii. We also unraveled the autotetraploid origin of A. kansuense from A. tenuicaule for the first time, and found that A. altajense shares essentially identical genomes with A. anogrammoides. [ABSTRACT FROM AUTHOR]

Several studies have demonstrated the usefulness of leaf epidermal, and spore morphological characters in the taxonomy of genus Asplenium. However, works on the Pakistani species of Asplenium are not existent. With the objective to verify the efficacy of leaf epidermal and spore morphological traits, the leaf epidermis and spore morphology in nine Asplenium species occurring in Malakand division was studied. The spores were studied under light microscope (LM) and scanning electron microscope (SEM), whereas for leaf epidermal anatomy, the LM was used. The spores are monolete, ellipsoidal in shape, the equatorial and polar diameter ranges between 28.3 and 50.2 × 27.6 and 45.8 μm. The exospore thickness varied from 0.5 to 3.8 μm. The perispore is 0.8–3.5 μm thick, ornamented, and morphologically variable. The spores characters specifically the perispore ornamentation are useful in distinguishing species within the genus. The irregular spore shape and unusual development of perispore wall surface characterize aborted spores in the species of Asplenium × alternifolium. The most informative quantitative characters appeared to be length and width of epidermal cells, and length and width of stomata were useful to distinguish species. The most significant qualitative characters to distinguish species were the anticlinal wall pattern. Our study has shown that considerable variations exist in the leaf epidermis of Asplenium species, at least some of which has taxonomic significance. We confirmed the prevalent taxonomic value of stomatal and epidermal cells traits. The leaf epidermal anatomy and spore morphological features showed to be a good source of information for taxonomy of the genus Asplenium. [ABSTRACT FROM AUTHOR]

Extracts of the aerial (fronds) and underground parts (rhizome and roots) of Asplenium adiantum-nigrum L. and Asplenium ruta-muraria L. were analyzed for their phenolic composition and antioxidant properties (scavenging capacities against DPPH• and ABTS•+), and were comparatively assayed for antimicrobial and antibiofilm activity. Total of the 17 phenolic compounds, belonging to the classes of phenolic acids (hydroxybenzoic and hydroxycinnamic acids), flavonoids (flavan-3-ols, flavonols), and xanthones were identified and quantified using UHPLC/DAD/(–)HESI-MS/MS analysis. The present study shows that two analyzed ferns are significantly diverse in terms of their phenolic profile. Flavonoids considerably contribute to the total phenolic content of A. ruta-muraria , ranging from 47% in roots to around 75% in rhizome and fronds, while A. adiantum-nigrum preferably accumulates xanthones. Antioxidant activity of fern methanol extracts was ascribed to their phenolics content. Extracts of rhizomes of both Asplenium ferns exhibited the highest antimicrobial activity, while frond extracts were more efficient in inhibition of Candida albicans biofilm formation. The present study provides experimental evidence that methanol extracts from two analyzed Asplenium fern species exhibit strong antibacterial effect against Gram-negative bacteria which candidate them as a potentially useful natural antimicrobial agents. [ABSTRACT FROM AUTHOR]

Hybridization is a ubiquitous force in plant evolution. In ferns, hybrids are often easily recognized by their intermediate morphology and abortive spores and thus provide a good model for studying reproductive isolating barriers between species. Asplenium ×aran-tohanum is a rare fern hybrid, despite wide coexistence of its parental species (A. billotii and A. trichomanes subsp. quadrivalens) in Western Europe. We made a complete characterization of its three known individuals, including macro- and micromorphology, sporogenesis, gametophyte reproduction, and chloroplast DNA inheritance and evolution. The hybrid expressed morphological characters that were mostly intermediate between those of the parents, but some characters were more similar to one parent or the other. Sizes of both guard cells and spores indicate that the hybrid is tetraploid, as are both parents, and one parent (A. billotii) consistently acted as the female. A very small fraction of spores (~ 7%) were viable and the resulting gametophytes could not form sporophytes, either sexually or apogamously, suggesting that effective postzygotic barriers exist between the parents. The lineages of these taxa diverged about 35 million years ago, which may explain the strong reproductive isolation and rarity of the hybrid. Asplenium ×aran-tohanum appears to be an evolutionary dead end, probably formed recurrently at the places where it grows, but incapable of completing its life cycle or producing viable offspring. [ABSTRACT FROM AUTHOR]

Asplenium Genus is present inside Alexandru Beldie Herbarium from „Marin Drăcea" National Institute for Research and Development in Forestry. This fact is consolidated by the presence of 92 vouchers containing plants of this genus, as well as information about them. The data refers to their harvesting location which covers almost the entire country and important names of the specialists who contributed to the development of the collection by gathering or identifying Asplenium plants. The present paper organizes and presents the Asplenium species present in the herbarium, reaching a total number of 16 species and 88 samples based on the year and place of their harvest, as well as the specialist who collected the plant. Additional criteria are also followed, such as: drawer number, voucher number, botanical collection, species name, date of collection, place of collection, the specialist who collected and / or determined the species and the degree of conservation. This last criterion was placed on a scale from 1 to 4 with 1 meaning a very good state of preservation while 4 represents a very poor state of preservation. In addition, the most important Asplenium species from the herbarium are also described from the point of view of their abundance in the collection. The paper starts with a short description of the herbarium, followed by the presentation of the studied material (92 vouchers that contain 16 Asplenium species). This stage details also the materials and methods used for elaborating the paper, organizing the plants and describing the most important ones. The herbarium hosts a sample of a rare species mentioned in the Red Book of superior Romanian plants (Asplenium lepidum Presl.). Alternatively, the collection can take pride in old Asplenium plants with an historical value that were collected 180 years ago (Asplenium trichomanes L, 1842). In addition, the paper graphically represents the periods in which the plants were harvested, as well as a map with the harvesting places on the Romanian territory. The conclusions present important aspects regarding the species and samples of Asplenium present in this herbarium. [ABSTRACT FROM AUTHOR]

The genus Asplenium L. comprises about 700 species of terrestrial, epiphytic, and saxicolous habits distributed in temperate and tropical regions around the world, exhibiting a high chemical richness with variable biological activity. In this review, compounds with antioxidant activity that constitute a pharmacological potential in diseases of the central nervous system are detailed. Asplenium nidus L. species presents a high concentration of phenols and flavonoids evidenced by antioxidant activity assays, such as DPPH, FRAP, total phenols, total flavonoids, and ORAC, and represent compounds with bioactive potential, including neuroprotection. The species Asplenium adiantum-nigrum L. presents a high antioxidant potential of its rhizome extracts exhibited in DPPH and ABTS assays, attributed to the high concentration of mangiferin. the xanthone mangiferin is a compound also present in other species of genera, such as Asplenium ceterach L. and Asplenium montanum Willd., in significant amounts. This xanthone has studies on its neuroprotective effect through different targets, some of them being the acetylcholinesterase enzymes, the 5-lipooxygenase enzyme, and the antioxidant activity itself. All these mechanisms of action of mangiferin constitute an object of study for its effect on memory loss, which can be relayed in Alzheimer’s disease.

Introducción y objetivos: Las características externas e internas de los pecíolos brindan información útil para los estudios taxonómicos en helechos. El objetivo de este trabajo fue analizar los caracteres diagnósticos del pecíolo en las 37 especies de Asplenium que crecen en Argentina, para contribuir a su identificación e indagar sobre su significado adaptativo. M&M: Se emplearon especímenes de herbario y material fresco. Los materiales se analizaron con microscopios estereoscópico y óptico; en el último caso las muestras fueron procesadas siguiendo las técnicas histológicas convencionales. Los caracteres analizados fueron: color, longitud, presencia de alas, tejido mecánico sub-epidérmico, líneas de aireación, banda circumendodermal (BC), forma y número de los haces vasculares y contorno del xilema. Resultados: Algunas características de los pecíolos permiten el agrupamiento de especies. Los caracteres diagnósticos son: color, presencia/ausencia de alas, número de haces vasculares, contorno de los haces, contorno del xilema y presencia/ausencia de BC. Conclusiones: Los caracteres del pecíolo por si solos no permiten la identificación de los taxa a nivel específico, si bien junto con otros contribuyen a la sistemática del género. La anatomía del pecíolo no evidencia en general relación con el hábito de estas plantas. El tipo y la disposición del tejido mecánico sub-epidérmico y la presencia de BC se asocian a especies que crecen en ambientes expuestos y secos, pero no son exclusivos de los taxa que crecen bajo estas condiciones."

The ferns Asplenium ceterach L., Asplenium scolopendrium L. and Asplenium trichomanes L. have wide application in traditional medicine worldwide. However, the scientific research on their anticancer and antibacterial properties is insufficient. The present article aims to provide more information on this topic. Extracts derived from the aerial parts of A. ceterach, A. scolopendrium and A. trichomanes were examined using a panel of in vitro assays with different bacterial and mammalian cells. The cytotoxicity and anticancer activity of the samples were analyzed by 3-(4,5-dimethylthiazol-2-yl)-2,5-diphenyltetrazolium bromide (MTT) and Trypan blue assays with three human (A549, FL, HeLa) and three murine (3T3, TIB-71, LS48) cell lines. Inhibitory effects on the growth of Gram-positive (Bacillus cereus) and Gram-negative (Pseudomonas aeruginosa) bacteria were determined by the agar diffusion assay. Apoptosis-inducing properties of the extracts were analyzed by flow cytometry. Superoxide dismutase (SOD) activity in extract-treated cells was investigated by ELISA. The obtained results demonstrate selective anticancer activity of all three Asplenium species. The extract from A. ceterach displayed the strongest inhibitory properties against human cervical cancer cells and bacterial cells. It induced a lower level of cytotoxicity against mouse cell lines, indicating a species-specific effect. The extract from A. trichomanes demonstrated better anticancer and antibacterial properties than the sample from A. scolopendrium. Further experiments linked the mechanism of action of A. ceterach extract with oxidative stress-inducing potential and strong proapoptotic potential against the cervical cancer cell line HeLa. A. trichomanes and A. scolopendrium extracts appeared to be potent inducers of necrotic cell death.

Background and aims - Relatively few studies have addressed the sun-shade response of fern species. Moreover, there is no information on species-specific plasticity patterns of such response, their relationship with species ecological requirements and the costs of such plasticity. The present study aims at filling these gaps by analysing the sun-shade plastic response of two Asplenium species that differ in their ecological requirements. Methods - We measured 27 leaf morphological, anatomical and physiological parameters using standard methods for A. ceterach and A. trichomanes in the field. The parameters were combined through Principal Component Analysis in order to highlight an integrated sun-shade response across species. Linear regression analysis was carried out to highlight the relationship between the calculated species plasticity patterns and the structural control on photosynthetic process. Key results - A significant degree of phenotypic plasticity was found for both species. Moreover, sun and shade leaves shared a common slope for the morpho-functional relationships reflecting no additional costs in terms of carbon assimilation. Even if the plastic responses of the two species scaled positively (R² = 0.68, P = 4.667e07), A. trichomanes was characterized by a slightly higher anatomical plasticity (plasticity index = 0.19), while A. ceterach showed a higher physiological plasticity (0.60). Conclusion - A remarkable acclimation capacity for the two Asplenium species in response to different light conditions was highlighted. Nevertheless, A. ceterach seems to be more suited to cope with full sunlight conditions as compared to A. trichomanes, according to species ecological requirements. [ABSTRACT FROM AUTHOR]

Background and aims: The morphology and the anatomy of the petioles provide useful information for taxonomic issues in ferns. The aim of this study was to analyze the diagnostic characters of petioles in 37 species of Asplenium that grow in Argentina, to contribute to their identification and to ascertain on their adaptive significance. M&M: Herbarium specimens and fresh material were employed. Materials were analyzed under stereoscopic and light microscopes; in the latter case, the samples were processed following the conventional histological techniques. The characters analyzed were: colour, length, presence of wings, sub-epidermic mechanical tissue, aeration lines, circumendodermal band (CB), shape and number of vascular bundles and xylem contour. Results: Some characteristics of the petioles allow the grouping of species. The diagnostic features are: colour, presence/absence of wings, number of vascular bundles, bundles contour, xylem contour and presence/absence of CB. Conclusions: Petiole characters alone do not allow the identification of the taxa at specific level, although along with others they contribute to the systematics of the genus. The anatomy of the petiole does not show in general terms relationship with the habit of these plants. The type and distribution of subepidermic mechanical tissue and the presence of CB are associated with species that inhabit exposed and dry environments, but these features are not exclusive of the taxa that grow under these conditions. [ABSTRACT FROM AUTHOR]

Abstract Enzyme-assisted extraction (EAE) possesses the advantages of being environmentally friendly and easily operated owing to relatively mild reaction conditions. Thus, mucilage from Asplenium australasicum was extracted with xylanase, glucanase and simultaneous application of both enzymatic preparations then investigated for their physicochemical properties. It was observed that compared to the control, the yield of mucilage increased significantly by EAE (from 3.64% to 6.04–6.86%), while the average molecular weight also increased slightly. SEM results showed that enzymes can erode the raw material which contributed to the higher extraction efficiency of polysaccharide. Furthermore, EAE modified the monosaccharide composition profile of A. australasicum mucilage considerably, especially there was a significant increase in glucose and galacturonic acid content. Although the FT-IR fingerprint did not change pronouncedly due to the action of the enzymes, but the degree of polysaccharide branching seemed to be more complicated, thus significantly reducing the intrinsic viscosity and water holding capacity of mucilage. Graphical abstract Unlabelled Image Highlights • Mucilage from the fronds of A. australasicum (J. Sm.) Hook was extracted by endo-1,4-β-xylanase and/or β-glucanase. • Enzyme treatment could effectively erode the plant cell wall and improve the extraction yield of A. australasicum mucilage. • EAE further modified the monosaccharide compositions and physical properties of A. australasicum mucilage. [ABSTRACT FROM AUTHOR]

Dental calculus, or calcified plaque, is a potential source of archaeological information relating to the lives of past societies. The recovery and identification of organic microremains entrapped within dental calculus provide new data regarding diet, health, and lifestyle. This paper presents, for the first time, microscopic evidence of the sporangium annulus of a fern within the dental calculus of an adult male from the medieval necropolis of Can Reiners (Balearic Islands, Spain). The sporangium evidence was compared with modern sporangia collected in northern Mallorca and the Iberian Peninsula. The results revealed that the features of the Can Reiners sporangium sample were compatible with the species Asplenium trichomanes. Through a historical and modern botanical literature review, it is known that this species has been widely regarded as a cure for kidney stones and alopecia. Moreover, this species was used as an expectorant, diuretic, and emmenagogue (herb that provokes menstruation or abortion). Considering that sporangia are attached to leaves and when they are mature only spores are released in the air, the presence of the annulus within dental calculus provides direct evidence of the consumption of ferns by the population living in the Balearic Islands during the Middle Ages. Although the use of these plants as constituents of diet is not attested, we may suggest that they could be employed as medicinal plants. [ABSTRACT FROM AUTHOR]

In this work we include interesting chorological news regarding to three ferns which are unknown or scarce in the Iberian Peninsula. Firstly, it is documented for the first time the presence of Polystichum braunii in the Iberian Peninsula. On the other hand, we add the third known locality for the hybrid Asplenium × aran-tohanum. Finally, we extend notably the distribution of Struthiopteris spicant var. homophyllum towards the interior of the Iberian Peninsula. [ABSTRACT FROM AUTHOR]

The complete plastome sequence of Asplenium × bimixtum (Aspleniaceae) was determined. The whole genome is 153,098 bp in length, consisting of large (86,394 bp) and small (21,508 bp) single-copy regions, and a pair of identical inverted repeats (22,598 bp). The genome contains 88 protein-coding genes, 33 tRNA genes, 8 rRNA genes, and 1 pseudogene (infA). The phylogenetic analysis based on 84 protein-coding genes showed that Asplenium is monophyletic with high bootstrap value. [ABSTRACT FROM AUTHOR]

Asplenium nidus L. is one of the most important terrestrial fern in Andaman and Nicobar Islands with great ornamental value. Inter simple sequence repeat (ISSR) analysis was used to study the genetic diversity of the species A. nidus L. collected from different geographical locations of the Island. Among 15 ISSR primers screened, 9 primers produced amplification and produced a total of 222 reproducible amplified bands. The number of amplified bands varied from 10 in ISSR 11 to 44 in UBC 842 with an average of 24.7 bands per primer. The range of polymorphic bands and percent polymorphism observed were 1 to 5 and 33.3 to 83.3, respectively. The polymorphic information content value of ISSR marker ranged from 0.365 to 0.485. Cluster analysis grouped the twelve accessions into three clusters (Cluster I, II and III) at Jaccard's similarity coefficient value of 0.69 on basis of geographical locations of the accessions. The cluster I was further divided into two sub-clusters Ia and Ib at the coefficient value of 0.77 and it includes the accessions belonging to South Andaman, one each accession collected from two different locations, Little Andaman and Rangat (Middle Andaman) were grouped separately as clusters II and III, respectively. The genetic variability studies with ISSR marker grouped the accessions of A. nidus L.on basis of their geographical locations and it would help in rapid identification of polymorphism and assist in germplasm collection, conservation and domestication programmes. [ABSTRACT FROM AUTHOR]

We describe Pleopeltis constricta Sprunt & Hickey as a new species from El Salvador and Mexico and additionally provide eight new fern records for the State of Veracruz. Furthermore, we propose one new combination each in Pecluma (P. liebmannii (C. Chr.) A. R. Sm. & Carv.-Hern.) and Pleopeltis (P. acicularis (Weath.) A. R. Sm. & T. Krömer). These novelties bring the new total for Veracruz to 574 species. [ABSTRACT FROM AUTHOR]

Dispersal is a key process in plant population dynamics. In ferns, two successive vectors are needed: the sporangium catapulting mechanism, and wind or gravity. However, some rock ferns have a growth habit that suggests a kind of autochory by placing spores on the rock surface. Moreover, some ferns show modifications of the sporangial dehiscence. To determine the role of growth habit in spore dispersal, we checked the sporangial opening mechanism and explored the spatial distribution of plants on the walls. The presence of spores of Asplenium celtibericum, a rupicolous fern, in the rock surface was checked. In addition, its sporangial dehiscence, plant size and position in the wall were analysed. Spores and indehiscent sporangia were present on walls at each sampling moment. Their highest number was found close to the plants. There was a positive correlation between crack width and plant size. However, most plants occupy the upper half of the cliffs. The growth habit of A. celtibericum is instrumental to deposit the spores over the neighbouring rock surface, thus enhancing the probability of spores to find suitable crevices for germination. Furthermore, dispersal of indehiscent sporangia might promote intergametophytic mating, and the modified sporangial opening mechanism extends the dispersive period. [ABSTRACT FROM AUTHOR]

The fern genus Hymenasplenium (Aspleniaceae) is one of the two genera in the family. It is generally recognized among modern pteridologists. However, its infrageneric relationships and species diversity have been unclear and controversial. The molecular studies so far have had small taxon and character sampling. In the present study, DNA sequences of six plastid markers of 158 accessions representing ca. 40 out of ca. 50 known species of Hymenasplenium , and 16 species of Asplenium were used to infer a phylogeny with maximum likelihood, Bayesian inference, and maximum parsimony approaches. Our major results include: (1) Hymenasplenium as currently defined is strongly supported as monophyletic; (2) three major clades representing early splits in Hymenasplenium are identified, with the Old World species being strongly supported as monophyletic; it is ambiguous if the New World species are monophyletic; (3) extensive cryptic speciation in the Old World is discovered demonstrating the complexity of evolution of the genus; and (4) six strongly or moderately supported subclades in the Old World clade are revealed, differing from one another in molecular, morphological, and geographical features. [ABSTRACT FROM AUTHOR]

The utility of three plastid DNA regions to identify fern species was explored with focus on the European representatives of the Asplenium trichomanes aggregate. The sampling included representatives of the three diploid and the four tetraploid taxa recognized in the European flora plus Macaronesia. Besides European samples, the compiled data set comprised specimens of a putative Hawaiian endemic and one species occurring in Southeast Asia. By combining the sequences of three non-coding plastid regions, 13 haplotypes were recovered of which four were found in more than one taxon. Evidences for four distinct diploid lineages were found that correspond to Asplenium anceps, A. inexpectans, A. trichomanes s.s., and A. tripteropus. The four tetraploids occurring in Europe shared haplotypes with A. inexpectans. Thus, DNA barcoding can successfully identify the diploids, but fail to separate the tetraploids from their diploid ancestors. As a consequence, barcoding analyses of ferns need to take into account the differences of ploidy level measured by evidence independent from the DNA barcode. Evidence for uneven accumulation of intra-species DNA variation was recovered by comparing all species. Furthermore, the study provided evidence that the current taxonomy of these ferns requires to be revised. The two European diploids form well-separated clades and need to be recognized as A. inexpectans and A. trichomanes s.s. To keep name consistency for all European tetraploids, a new name Asplenium jessenii is introduced to replace A. trichomanes subsp. hastatum. [ABSTRACT FROM AUTHOR]

The Asplenium normale D. Don complex comprises several taxa that are either diploid or tetraploid. The tetraploids are assumed to have originated from diploid ancestors by relatively recent autopolyploidization or allopolyploidization. Some of the diploids are readily recognized morphologically but most of the taxa have until now been placed into a single species. However, phylogenetic studies have challenged this treatment and emphasized the notion that the taxonomic treatment of this complex needs to be revised. An integrative taxonomic approach was employed to delimit species in the complex using cytological, morphological, and DNA sequence data. Initially, we employed a diploid first approach to establish a robust taxonomic framework. Special efforts were made to collect and identify the diploid progenitors of each polyploid lineage identified in the plastid DNA based phylogenetic hypothesis. A total of six distinct diploid species were identified. The distinctive nature of the six diploids is strongly supported by sequence differences in plastid DNA and nuclear loci, as well as by the results of morphometric analysis. Diagnostic morphological characters were identified to distinguish the six diploid species, resulting in their revised taxonomy, which includes two novel species, namely, Asplenium normaloides and A. guangdongense. Further studies to strengthen the taxonomic classification of all of the tetraploid taxa are warranted. [ABSTRACT FROM AUTHOR]

Abstract Asplenium L. is estimated to be the largest fern genus and is very prone to polyploidy and hybridization. We investigated the phylogenetic relationships of two geographically restricted complexes in the Mediterranean Basin, and searched for evidence of multiple and/or reciprocal origins of the allopolyploids. The A. obovatum Viv. and A. adiantum-nigrum L. complexes include a total of five diploid and four polyploid taxa and are linked by the tetraploid A. balearicum Shivas. We collected 36 new samples representing all nine taxa, from which we amplified five chloroplast regions. We also included sequences from GenBank and performed maximum likelihood (ML) and Bayesian inference (BI) analyses to identify multiple and reciprocal origins. We conducted a larger molecular dating analysis to assess relationships with the rest of Asplenium and estimate divergence times. This matrix included 137 species of Asplenium and three outgroups, including sequences previously published. All sequences of allotetraploids A. adiantum-nigrum and A. balearicum grouped with A. onopteris L., with no sign of reciprocal parentage, but possible evidence of multiple origins. The tetraploids A. foreziense Le Grand and A. billotii F. W. Schultz grouped with both respective parents, indicating probable multiple and reciprocal origins. The best supported case of reciprocal parentage is A. foreziense. In the larger analysis, the species studied resolve in two different clades that correspond to the two complexes, and the groups are not closely related, with a conservative estimate of 45.43 millions of years ago (Ma) for their divergence. There is evidence of multiple origins of all four allopolyploids, though only with strong support for one, and of reciprocal origins for two of the four allopolyploids, though again only one is well supported. Asplenium balearicum represents a hybridization event between members of clades whose ancestors diverged roughly 45 Ma, making this divergence one of the oldest to have descendants later produce hybrid offspring. [ABSTRACT FROM AUTHOR]

The paper aims to determine the lead-induced physiological, biochemical and enzymatic changes in Asplenium scolopendrium, which could represent biomarkers used in environmental assessment. Of all the physiological processes, photosynthesis and respiration were analyzed and the enzymatic and biochemical determinations focused on catalase activity, assimilatory pigment concentration, polyphenol content and lead presence in tissues.The stress induced by the exposure to Pb of the species Asplenium scolopendrium determined an increase in the carotenoid content, the catalase activity, the total polyphenol content and also enhanced the respiration potential. No significant changes were recorded regarding the chlorophyll content and the photosynthetic activity. The recorded changes may be used as non-specific markers in the assessment of the impact of Pb on plants (Asplenium scolopendrium). [ABSTRACT FROM AUTHOR]

* Background and Aims A key structural adaptation of vascular plants was the evolution of specialized vascular and mechanical tissues, innovations likely to have generated novel cell wall architectures. While collenchyma is a strengthening tissue typically found in growing organs of angiosperms, a similar tissue occurs in the petiole of the fern Asplenium rutifolium. * Methods The in situ cell wall (ultra)structure and composition of this tissue was investigated and characterized mechanically as well as structurally through nano-indentation and wide-angle X-ray diffraction, respectively. * Key Results Structurally the mechanical tissue resembles sclerenchyma, while its biomechanical properties and molecular composition both share more characteristics with angiosperm collenchyma. Cell wall thickening only occurs late during cell expansion or after cell expansion has ceased. * Conclusions If the term collenchyma is reserved for walls that thicken during expansive growth, the mechanical tissue in A. rutifolium represents sclerenchyma that mimics the properties of collenchyma and has the ability to modify its mechanical properties through sclerification. These results support the view that collenchyma does not occur in ferns and most probably evolved in angiosperms. [ABSTRACT FROM AUTHOR]

Asplenium is one genus ferns aspleniaceae tribal members. Identification of the classification of the species can be determined by one of the morphological characteristics of the spores. Spore morphological characteristics include the shape, size, and type aperture ornamentation. The purpose of this study is to describe and analyze the species based on morphological characters spores. The method used is descriptive explorative to explore ferns Asplenium. The research found that five types of Asplenium is Asplenium nidus, Asplenium robustum, Asplenium salignum, Asplenium tenerum, and Asplenium unilaterale. The difference of 5 species Asplenium lies in the type of ornamentation characteristic morphology and size of the spores, whereas the equation is the shape, size and type categories aperture spores. Asplenium merupakan salah satu marga tumbuhan paku anggota suku Aspleniaceae. Identifikasi penggolongan spesies dapat ditentukan melalui salah satu karakteristik morfologi spora. Karakteristik morfologi spora meliputi bentuk, ukuran, apertura dan tipe ornamentasi. Tujuan penelitian ini adalah mendeskripsikan dan menganalisis karakter spesies berdasarkan morfologi spora. Metode penelitian yang digunakan ialah deskriptif eksploratif untuk mengeksplorasi tumbuhan paku Asplenium. Hasil penelitian ditemukan 5 jenis dari Asplenium yaitu Asplenium nidus, Asplenium robustum, Asplenium salignum, Asplenium tenerum, dan Asplenium unilaterale. Perbedaan dari 5 jenis spesies Asplenium terletak pada ciri morfologi tipe ornamentasi dan ukuran spora, sedangkan persamaan terletak pada bentuk, kategori ukuran dan tipe apertura spora.