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Paleopalynology evidence and megafossil records indicated that tree fern order Cyatheales, played an important role in Mesozoic terrestrial ecosystem. Few fossil records of the family Thyrsopteridaceae have been reported so far. In the present study, we describe a distinctive fertile pinnule segment of a fern plant preserved in a mid-Cretaceous amber from Myanmar. The well-preserved fertile pinnule is identified as a new species of the extant tree fern genus Thyrsopteris (Thyrsopteridaceae): Thyrsopteris cyathindusia n. sp. It represents the second fossil species of Thyrsopteridaceae found in the mid-Cretaceous amber from Myanmar. The new species shows distinctive characteristics of the sporophyll that are considered paleoecologically/paleobiogeographically significant for evolution of the Thyrsopteridaceae.

Premise of research. Excavations at the Clarkia fossil beds in northern Idaho have recovered numerous exceptionally well-preserved plant fossils over the past 45 years. We report on a compression f...

New remains corresponding to Cyathealean tree ferns are described for the Aptian Cerro Negro Formation at Byers Peninsula, Livingston Island (South Shetland Islands, Antarctica). A new genus and species, Rafaherbstia nishidai, is proposed for several permineralized dictyostelic stems characterized by a heterogeneous pith with a central star-shaped sclerenchyma mass, U-shaped meristeles developing small hooks at the margins of the leaf gap, sclerenchyma sheaths surrounding the vascular strands, and leaf traces omega-shaped and multi parted. The new taxon shows a combination of features that preclude its placement in one of the eight recognized families of Cyatheales, supporting that previous diversity of tree ferns was higher than in modern times. Fragmentary isolated sori with preserved sporangia containing trilete spores, recovered from the sedimentary matrix surrounding the stems, are also described. These new tree ferns increase the diversity of Cyatheales in the Cerro Negro Formation, and support warm climatic conditions in this region during the Aptian. Fil: Vera, Ezequiel Ignacio. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Departamento de Geología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”; Argentina Fil: Cesari, Silvia Nelida. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”; Argentina

Variation in rates of molecular evolution (heterotachy) is a common phenomenon among plants. Although multiple theoretical models have been proposed, fundamental questions remain regarding the combined effects of ecological and morphological traits on rate heterogeneity. Here, we used tree ferns to explore the correlation between rates of molecular evolution in chloroplast DNA sequences and several morphological and environmental factors within a Bayesian framework. We revealed direct and indirect effects of body size, biological productivity, and temperature on substitution rates, where smaller tree ferns living in warmer and less productive environments tend to have faster rates of molecular evolution. In addition, we found that variation in the ratio of nonsynonymous to synonymous substitution rates (dN/dS) in the chloroplast rbcL gene was significantly correlated with ecological and morphological variables. Heterotachy in tree ferns may be influenced by effective population size associated with variation in body size and productivity. Macroevolutionary hypotheses should go beyond explaining heterotachy in terms of mutation rates and instead, should integrate population-level factors to better understand the processes affecting the tempo of evolution at the molecular level.

A new species of hairy tree fern (Dicksoniaceae - Cyatheales) is described and illustrated: Dicksonia utteridgei from stunted ridge forest in central New Guinea. The species is the only one in the genus with a combination of fully pinnate to bipinnate-pinnatifid pinnules and bristly spreading petiole hairs.

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.

An unusual tree fern trapped in mid-Cretaceous Myanmar amber is described as Thyrsopteris cretacea sp. nov. (Thyrsopteridaceae). The fossil most resembles T. elegans, a present-day species endemic to the Juan Fernandez Islands, based on the character combination of pinnate fertile lamina segments with short lobes bearing terminal sori, cyathiform indusia, clavate receptacles, and oblique annuli. Thyrsopteris cretacea is the first described mid-Cretaceous tree fern preserved in amber. It adds to the diversity previously ascribed to the Thyrsopteridaceae, which has been based on Eocene fossils, and it extends the fossil record of the family further back to the mid-Cretaceous. Most previous fossils of Thyrsopteridaceae have been from the Southern Hemisphere and are therefore considered Gondwanan. Thyrsopteris cretacea represents one of the few occurrences of the family in Laurasia.

To determine the mycorrhizal status of pteridophyte gametophytes in diverse taxa, the mycorrhizal colonization of wild gametophytes was investigated in terrestrial cordate gametophytes of pre-polypod leptosporangiate ferns, i.e., one species of Osmundaceae (Osmunda banksiifolia), two species of Gleicheniaceae (Diplopterygium glaucum, Dicranopteris linearis), and four species of Cyatheales including tree ferns (Plagiogyriaceae: Plagiogyria japonica, Plagiogyria euphlebia; Cyatheaceae: Cyathea podophylla, Cyathea lepifera). Microscopic observations revealed that 58 to 97% of gametophytes in all species were colonized with arbuscular mycorrhizal (AM) fungi. Fungal colonization was limited to the multilayered midrib (cushion) tissue in all gametophytes examined. Molecular identification using fungal SSU rDNA sequences indicated that the AM fungi in gametophytes primarily belonged to the Glomeraceae, but also included the Claroideoglomeraceae, Gigasporaceae, Acaulosporaceae, and Archaeosporales. This study provides the first evidence for AM fungal colonization of wild gametophytes in the Plagiogyriaceae and Cyatheaceae. Taxonomically divergent photosynthetic gametophytes are similarly colonized by AM fungi, suggesting that mycorrhizal associations with AM fungi could widely occur in terrestrial pteridophyte gametophytes.

The patterns of geographic differentiation in fern species have been linked to climatic differences across regions and the distribution of available habitat. In this paper, the association between some climatic features and patterns of geographic differentiation in American tree ferns was evaluated. For this, the occurrence ranges of 190 species were plotted and then analysed using track analysis. Here we identify six generalised tracks that span the main highland regions of tropical America: the Andes, the Guyana Highlands, the Brazilian Atlantic coast, lower Central America, the Greater Antilles, and upper Central America-Mexico. We did not find an association between cloud forest habitat and the differentiation pattern revealed by generalised tracks in Central America. Instead, these patterns are congruent with well-documented geological boundaries in the region. Climatic variables associated with cloud forest habitat were extracted from each generalised track and subjected to ANOVA, showing t...

Marattiaceous ferns are conspicuous fossil element in upper Paleozoic and lower Mesozoic deposits, but become rarer to almost absent in Cretaceous to Holocene deposits. In this contribution, fragmentary synangia recovered from the Aptian Cerro Negro Formation (Livingston Island, Antarctica) are described. Synangia are bivalvate, containing at least four lachrymiform sporangia per valve. Valves are fused at the base, and connect to the leaf blade by a short peduncle. Sporangia contain more than 400 oval to reniform (probably monolete), smooth to microgranulate spores. The reproductive structures have morphological similarities with extant Marattia and Ptisana, and the apparent absence of labia suggests a closer relationship with the latter genus. The presence of Marattiaceae, along with previously described ferns (e.g. Cyatheales) supports warm climatic conditions during the Aptian in this region of Antarctica. Fil: Vera, Ezequiel Ignacio. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Departamento de Geología; Argentina Fil: Cesari, Silvia Nelida. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina

We present a taxonomic revision of Cyathea, clade Hymenophyllopsis, a monophyletic group restricted to the tepuis of the Guiana Shield, a region in northern South America. We recognize fifteen species, including four newly described here: C. indefinita, C. minima, C. pseudoctenitoides, and C. tatei. The main diagnostic characters for the species are: rhizome (symmetry and scales), lamina division (pinnatisect to 5-pinnate), pinnae (congested or sparse, recurverd to ascending), scales and/or hairs of the rachis (shape, size, margins), as well as the shape and margin of the indusium (bivalvate or cyatheoid, entire to deeply lacerate). An identification key, descriptions, taxonomic comments, illustrations, and geographical distribution maps for all recognized taxa are provided.

A new genus and species of cyathealean fern, Yavanna chimaerica gen. et sp. nov., is erected for several permineralized stems recovered from the Early Cretaceous (Aptian) Cerro Negro Formation, which crops out at the South Shetland Islands, Antarctica. The new species is characterized by solenostelic erect stems with medullary bundles traversing the pith, and surrounded by persistent petiole bases and adventitious roots. Proximal petiole bases present a one-parted modified omega-shaped trace, which becomes three-parted distally. The anatomy of the new fern show similarities with both Thyrsopteris elegans Kunze and with the Cyatheaceae s.s., suggesting that it is a representative of an extinct lineage among the Cyatheales. Fil: Vera, Ezequiel Ignacio. Universidad de Buenos Aires. Facultad de Cs.exactas y Naturales. Departamento de Geologia. Cat.de Paleontologia; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales; Argentina

Analyses of richness and endemism of Cyatheales (tree ferns) in tropical America were performed and evidence of a diversity gradient is presented. For this, the occurrence ranges of 239 species were plotted into a 5° × 5° grid-cell map and then analyzed using species richness and endemism indices. Here we show that species richness and endemism are not distributed randomly over the landscape, but do aggregate into defined regions of high diversity in tropical America: the northern Andes, lower Central America, upper Central America and Mexico, the Guyana Highlands, southeastern Brazil, and the Antilles. These distributional patterns are congruent with the geographical distribution of cloud forest, which in turn is determined by topography, high humidity, and persistent cloud immersion. The mountain regions of tropical America, especially the cloud forests, harbour most of the species of American Cyatheales and have high levels of habitat loss and climatic fragility. Conservation policies for Cyatheales are centred on the local use and trade of many tree fern species, but none such policies focus on cloud forest habitat loss. This makes tree ferns a critically endangered group of plants. In the face of the current environmental crisis and global climate change, the presence of Cyatheales in these regions sounds the alarm on their conservation priorities.

The study of abundant permineralized fern remains collected in the Oligocene Rancahue Formation (Neuquen Province, Argentina) allowed the identification of two different taxa. The name Cuyenopteris patagoniensis nov. gen., nov. sp. is proposed for several stems and fronds characterized by the presence of caulinar dictyosteles with root gaps, producing C-shaped leaf traces composed by two hippocampiform bundles flanked by eight to ten small round bundles, half of them diverging from cauline meristeles and the rest from the hippocampiform bundles. The presence of these features, along with frond bases with adaxial grooves and a pair of pneumatophores, suggest affinities with the Blechnaceae or Dryopteridaceae. The second identified taxon is Alsophilocaulis calveloi Menendez. Based on features observed in these new materials, including characteristics of the mantle of adventitious roots and anatomy of the petiole bases, an emended diagnosis for this latter species is proposed.

Three new species of Cyathea , two from Venezuela and one from Venezuela and the border region between Venezuela and Brazil, are here described: Cyathea alsophiloides , C. cylindrica , and C. lellingeriana . These species are endemic to the tepuis of the Guayana Shield and occur mainly at high elevations (1850–2555 m, although C. lellingeriana also grows down to 770 m). Descriptions, illustrations and mapped distributions are provided for each species, and their similarities to and differences from other species of the Hymenophyllopsis clade in Cyathea are discussed.

Two new specimens of a cyathealean fern stems collected from the Lower Cretaceous (Aptian) Cerro Negro Formation, which crops out at the South Shetland Islands, Antarctica, are described in detail. The specimens are dictyostelic stems with U-shaped meristeles without sclerenchyma sheaths and petiole bases with numerous, small, vascular strands, and a mantle of adventitious roots. The anatomy of the stems show similarities with Cibotiaceae, but given they are not well preserved, some critical features that would allow a conclusive referral to this family are lacking. Preserved in the surrounding matrix, fragmentary pinnules were found, showing sori with bivalvate indusia, and sporangia with more than 20 spores, attached to an elongate receptacle. The features observed in these fertile remains suggest cyathealean affinities, but are difficult to refer to a particular family. These findings increase the diversity of cyathealean tree ferns in the Cerro Negro Formation, supporting previously suggested warm climatic conditions for this region of Antarctica during the Aptian. Fil: Vera, Ezequiel Ignacio. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Departamento de Geología. Cátedra de Paleontología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”; Argentina

Two new genera and species of permineralized tree ferns, Natalipteris wildei and Kwazulupteris schaarschmidtii, are defined and described in detail. Natalipteris wildei is a solenostelic stem without well-developed sclerenchyma sheaths in their vascular strands, and has a single vascular strand in the petiole bases, which are somewhat similar to the one present in Cibotium and Nishidacaulis. On the other hand, K. schaarschmidtii is a dictyostelic fern, with petiole bases with fused adaxial arcs and a single isolated meristele in the petiolar pith. Both taxa present features that preclude their placement in the recognized families of Cyatheales, but may be referred to the "core tree ferns" clade. Fossil specimens were found in the Senckenberg Forschungsinstitut Palaeobotanical Collection and, although they lack precise stratigraphic provenance, it is suggested that they were collected from the Mzinene Formation (Albian-Turonian). Fil: Vera, Ezequiel Ignacio. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”; Argentina. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Departamento de Geología. Cátedra de Paleontología; Argentina Fil: Herbst, Rafael. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Tucumán. Instituto Superior de Correlación Geológica. Universidad Nacional de Tucumán. Facultad de Ciencias Naturales e Instituto Miguel Lillo. Departamento de Geología. Cátedra Geología Estructural. Instituto Superior de Correlación Geológica; Argentina

A new genus and species of Mesozoic fern, Alienopteris livingstonensis gen. et sp. nov., is defined, based on permineralized stems recovered from the Aptian Cerro Negro Formation (Antarctica). This new taxon is characterized by the presence of Y-shaped meristeles in the stem, surrounding an heterogeneous pith, and the presence of complex petiole bases, consisting in abaxial and adaxial archs, similar to the ones present in the Cyatheaceae s.l.. Since no living nor fossil filicalean fern possesses such mixture of features, and taking into account that no leaves and reproductive structures were found, phylogenetic relationships are not clear. Two independent cladistic analysis show clear placement of Alienopteris livingstonensis among the ‘‘core tree ferns.’’ Fil: Vera, Ezequiel Ignacio. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina

Tree ferns are a well-established clade within leptosporangiate ferns. Most of the 600 species (in seven families and 13 genera) are arborescent, but considerable morphological variability exists, spanning the giant scaly tree ferns (Cyatheaceae), the low, erect plants (Plagiogyriaceae), and the diminutive endemics of the Guayana Highlands (Hymenophyllopsidaceae). In this study, we investigate phylogenetic relationships within tree ferns based on analyses of four protein-coding, plastid loci (atpA, atpB, rbcL, and rps4). Our results reveal four well-supported clades, with genera of Dicksoniaceae (sensu Kubitzki, 1990) interspersed among them: (A) (Loxomataceae, (Culcita, Plagiogyriaceae)), (B) (Calochlaena, (Dicksonia, Lophosoriaceae)), (C) Cibotium, and (D) Cyatheaceae, with Hymenophyllopsidaceae nested within. How these four groups are related to one other, to Thyrsopteris, or to Metaxyaceae is weakly supported. Our results show that Dicksoniaceae and Cyatheaceae, as currently recognised, are not monophyletic and new circumscriptions for these families are needed.