The preservation and palaeoecology of the Lower Carboniferoussilicified plant deposits at Esnost,near Autun,France

The mineralogy and preservation history of the anatomically preserved plants (permineralizations) of Lower Carboniferous age (Upper Visean) from Esnost, near Autun, France is described. The plants are all preserved in silica and are associated with acid volcanism. The permineralized plants occur within a range of different sediments each of which is characterized by a specific floral association and has a distinct mineralization. The structure of the mineralization is dependent on whether fossilization occurred within a site of growth of a plant or at a site of deposition of plant fragments. The timing of mineralization, whether early or late, also influenced the form of the crystals. The most common plant-bearing lithology at Esnost is the Archaeocalamites «peat. This lithology contains fine roots which are interpreted as the terminal roots of Archaeocalamites. These were important in providing channels through the semi-compacted peat for the permineralizing siliceous fluids. The Botryopteris cherts, dominated by one species of coenopterid fern, were fossilized at a much earlier stage giving exceptionally good preservation of the anatomical detail. The third most important plant-bearing sediment is the Diplolabis chert. This represents mineralization at the site of deposition of fragments of this fern. Within this site, mineralization was slow and episodic involving the considerable loss of organic material from the plant organs, resulting in some cases in a mineralization approaching petrifaction. Fusain was rare in the deposits but occurred within the Diplolabis cherts and the fusain-bearing chert where it is associated with a series of microorganisms of unknown affinities. The relationship of the sediments, plants and mineralization offers a basis for reconstructing the original ecology of the plants within the volcanic terrain. These plant deposits are compared to those of the same age, associated with basaltic volcanism, at Pettycur, Fife, Scotland.     

A new fossil record of Thyrsopteridaceae (Cyatheales) from the mid-Cretaceous amber of Myanmar

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.     

阔鳞瘤蕨颈卵器形成与卵发生的初步研究

运用光学显微镜与透射电镜对阔鳞瘤蕨（Phymatosorus hainanensis(Noot.) S.G.Lu）颈卵器形成和卵发生进行了研究。阔鳞瘤蕨颈卵器产生于雌配子体生长点下方分枝毛状体内侧。切片观察表明颈卵器起源于配子体表面的原始细胞,该细胞经两次不等分裂形成3个细胞,上下两个细胞分别发育为颈卵器的颈部与底部壁细胞,中间的细胞为初生细胞,含有较丰富的细胞器。初生细胞进行两次不等分裂产生颈沟细胞、腹沟细胞与卵细胞。成熟颈卵器内颈沟细胞和腹沟细胞退化,卵细胞上表面产生受精孔。本研究阐述了阔鳞瘤蕨颈卵器形成和卵发生的细胞学过程,对阐明蕨类植物雌性生殖器官的发育特征有一定的科学意义。     

Lithotrophic iron-oxidizing bacteria produce organic stalks to control mineral growth: implications for biosignature formation

Neutrophilic Fe-oxidizing bacteria (FeOB) are often identified by their distinctive morphologies, such as the extracellular twisted ribbon-like stalks formed by Gallionella ferruginea or Mariprofundus ferrooxydans. Similar filaments preserved in silica are often identified as FeOB fossils in rocks. Although it is assumed that twisted iron stalks are indicative of FeOB, the stalk''s metabolic role has not been established. To this end, we studied the marine FeOB M. ferrooxydans by light, X-ray and electron microscopy. Using time-lapse light microscopy, we observed cells excreting stalks during growth (averaging 2.2 μm h⁻¹). Scanning transmission X-ray microscopy and near-edge X-ray absorption fine structure (NEXAFS) spectroscopy show that stalks are Fe(III)-rich, whereas cells are low in Fe. Transmission electron microscopy reveals that stalks are composed of several fibrils, which contain few-nanometer-sized iron oxyhydroxide crystals. Lepidocrocite crystals that nucleated on the fibril surface are much larger (∼100 nm), suggesting that mineral growth within fibrils is retarded, relative to sites surrounding fibrils. C and N 1s NEXAFS spectroscopy and fluorescence probing show that stalks primarily contain carboxyl-rich polysaccharides. On the basis of these results, we suggest a physiological model for Fe oxidation in which cells excrete oxidized Fe bound to organic polymers. These organic molecules retard mineral growth, preventing cell encrustation. This model describes an essential role for stalk formation in FeOB growth. We suggest that stalk-like morphologies observed in modern and ancient samples may be correlated confidently with the Fe-oxidizing metabolism as a robust biosignature.     

Morphologies and elemental compositions of calcium crystals in phyllodes and branchlets of Acacia robeorum (Leguminosae: Mimosoideae)

Background and Aims

Formation of calcium oxalate crystals is common in the plant kingdom, but biogenic formation of calcium sulfate crystals in plants is rare. We investigated the morphologies and elemental compositions of crystals found in phyllodes and branchlets of Acacia robeorum, a desert shrub of north-western Australia.

Methods

Morphologies of crystals in phyllodes and branchlets of A. robeorum were studied using scanning electron microscopy (SEM), and elemental compositions of the crystals were identified by energy-dispersive X-ray spectroscopy. Distributional patterns of the crystals were studied using optical microscopy together with SEM.

Key Results

According to the elemental compositions, the crystals were classified into three groups: (1) calcium oxalate; (2) calcium sulfate, which is a possible mixture of calcium sulfate and calcium oxalate with calcium sulfate being the major component; and (3) calcium sulfate · magnesium oxalate, presumably mixtures of calcium sulfate, calcium oxalate, magnesium oxalate and silica. The crystals were of various morphologies, including prisms, raphides, styloids, druses, crystal sand, spheres and clusters. Both calcium oxalate and calcium sulfate crystals were observed in almost all tissues, including mesophyll, parenchyma, sclerenchyma (fibre cells), pith, pith ray and cortex; calcium sulfate · magnesium oxalate crystals were only found in mesophyll and parenchyma cells in phyllodes.

Conclusions

The formation of most crystals was biologically induced, as confirmed by studying the crystals formed in the phyllodes from seedlings grown in a glasshouse. The crystals may have functions in removing excess calcium, magnesium and sulfur, protecting the plants against herbivory, and detoxifying aluminium and heavy metals.     

Filicales y Bennettitales del Cretácico Inferior de Carazuelo, provincia de Soria, España

Silicified woods of the fern Tempskya riojana Barale and Viera and the Bennettitales Cycadeoidea barbarae Crisafulli nov. sp., found together with undetermined conifer woods, from the NW of Carazuelo, province Soria, are described. This site is located south of the so-called San Saturio fault which runs parallel to Road N-234. The fossil trunks are included in the “Conglomerados poligénicos, areniscas y arcillas” fluviatile sediments, dated as Paleogene. The original source of these fossils could be Tera and/or Oncala Groups, both Early Cretaceous in age and outcropping close by at Fuentetecha, Fuensaúco and North of Soria. Similar plant fossil findings from this region are reported and some mistakes in the geological mapping are analyzed. The Tempskya specimens were histologically poorly preserved, but had enough anatomical characters to be undoubtedly assigned to Tempskya riojana. The new species Cycadeoidea barbarae is described, with fairly well preserved cellular structure, viz., pith, vascular cylinder, cortex, foliar/branch traces, as well as strobili surrounded by foliar scales and apical seminal cones. An important and new aspect of the new species is that it includes morphologically and anatomically preserved both fructifications and vegetative parts, whereas these are usually find separated by other species of Bennettitales.     

Heterogeneity of silica and glycan-epitope distribution in epidermal idioblast cell walls in Adiantum raddianum laminae

Laminae of Adiantum raddianum Presl., a fern belonging to the family Pteridaceae, are characterised by the presence of epidermal fibre-like cells under the vascular bundles. These cells were thought to contain silica bodies, but their thickened walls leave no space for intracellular silica suggesting it may actually be deposited within their walls. Using advanced electron microscopy in conjunction with energy dispersive X-ray microanalysis we showed the presence of silica in the cell walls of the fibre-like idioblasts. However, it was specifically localised to the outer layers of the periclinal wall facing the leaf surface, with the thick secondary wall being devoid of silica. Immunocytochemical experiments were performed to ascertain the respective localisation of silica deposition and glycan polymers. Epitopes characteristic for pectic homogalacturonan and the hemicelluloses xyloglucan and mannan were detected in most epidermal walls, including the silica-rich cell wall layers. The monoclonal antibody, LM6, raised against pectic arabinan, labelled the silica-rich primary wall of the epidermal fibre-like cells and the guard cell walls, which were also shown to contain silica. We hypothesise that the silicified outer wall layers of the epidermal fibre-like cells support the lamina during cell expansion prior to secondary wall formation. This implies that silicification does not impede cell elongation. Although our results suggest that pectic arabinan may be implicated in silica deposition, further detailed analyses are needed to confirm this. The combinatorial approach presented here, which allows correlative screening and in situ localisation of silicon and cell wall polysaccharide distribution, shows great potential for future studies.     

The Range of Bioinclusions and Pseudoinclusions Preserved in a New Turonian (~90 Ma) Amber Occurrence from Southern Australia

A new Turonian amber occurrence, representing the oldest in situ amber locality in Australia and the southern-most locality in Gondwana, has recently been discovered in the Otway Basin of Victoria. The amber was collected from petroleum cores and many pieces contain a range of inclusions that can provide information on the depositional history of the resin. To date, one species of fern spore (Cyathidites minor) and one species of lycophyte spore (Kraeuselisporites sp?) have been conclusively identified in the amber, along with filamentous microorganisms and degraded plant matter. Several samples are also rife with pseudoinclusions as reported recently in other ambers. The abundance of preserved particulate debris and wind dispersed spores suggest that the Otway amber formed subaerially. Furthermore, based on the range of bioinclusions and forms of pseudoinclusions preserved within a single piece of amber, the locus of hardening for individual samples is variably interpreted as occurring in the tree tops, on the tree trunk or on the ground surface. Notably, specific inclusion assemblages are associated with certain colours of amber. By extension, and in accordance with recent studies, amber colour may be indicative of depositional environment. Variation in the environment of solidification may, therefore, be sufficient to account for the broad range of morphological characteristics preserved in a single amber deposit.     

The distribution of TPX2 in dividing leaf cells of the fern Asplenium nidus

Plant cell division requires the dynamic organisation of several microtubule arrays. The mechanisms of regulation of the above arrays are under rigorous research. Among several factors that are involved in plant microtubule dynamics, the Targeting Protein for Xklp2 (TPX2) has been found to play a role in spindle organisation, in combination with Aurora kinases, in dividing cells of angiosperms. Microtubule organisation in dividing cells of ferns exhibits certain peculiarities. Accordingly, the presence and distribution of a TPX2 homologue might be helpful in understanding the patterns and regulatory mechanisms of microtubule arrays in this plant group. In this study, a putative TPX2 homologue was identified using Western blotting in the fern Asplenium nidus. It was found, using immunostaining and CLSM, that it is co‐localised with perinuclear preprophase microtubules and the prophase spindle, and follows the microtubule pattern during metaphase/anaphase and telophase. During cytokinesis, while in angiosperms TPX2 is degraded, in A. nidus the TPX2 signal persists, co‐localising with the phragmoplast. In early post‐cytokinetic cells, a TPX2 signal is present on the nuclear surface facing the daughter cell wall and, thereafter it is co‐localised with the fern‐specific microtubule aggregation that lines the new wall, which is possibly involved in cortical microtubule assembly.     

Fossil evidence of eupolypod ferns in the mid-Cretaceous of Myanmar

Divergence time estimates based on DNA sequence variation of extant species suggest that eupolypod ferns were diverse already in the Cretaceous; however, fossil evidence to support this assumption remains exceedingly rare. Holttumopteris burmensis gen. et sp. nov., a fertile fern foliage fragment preserved in a piece of Albian–Cenomanian Burmese amber from Myanmar, is characterized by divided fertile leaves with catadromous, free lateral veins. Sporangia possess a vertical annulus interrupted by the sporangium stalk and contain monolete spores with a lophate perine. The sporangia occur clustered in discrete sori overarched by a laterally attached, reniform indusium. Reconstruction of ancestral character states suggests affinities of Holttumopteris to the Thelypteridaceae; however, several taxonomically relevant characters are not preserved. This discovery is nevertheless important because H. burmensis represents the first unequivocal fossil of a representative of the eupolypods from the middle Cretaceous.     

HALLETHECA RETICULATUS GEN. ET SP. N.: A SYNANGIATE PENNSYLVANIAN PTERIDOSPERM POLLEN ORGAN

A new structurally preserved synangiate pollen organ is described from the upper Pennsylvanian (Mattoon Formation) of southeastern Illinois. The specimen of Halletheca reticulatus gen. et sp. nov. measures approximately 1.5 cm long by approximately 5.0 mm in diam, and consists of five elongate sporangial tubes equidistantly arranged around a solid central column, and embedded in a thin-walled ground tissue. The vascular system consists of five terete bundles characterized by scalariform tracheids. Sporangia are thick-walled and contain pollen of the Monoletes-type. Ultrastructural studies of the pollen grain wall show it to consist of a uniform network of muri which branch and re-unite to form small lumina. A comparison of the ultrastructure of the pollen grain wall of the new fructification is made with the wall organization found in similar pollen of Dolerotheca. Relationships between the new taxon and other presumed seed fern pollen organs differing in preservation mode are discussed.     

Heinrichsia cheilanthoides gen. et sp. nov., a fossil fern in the family Pteridaceae (Polypodiales) from the Cretaceous amber forests of Myanmar

Divergence time estimates suggest that most clades constituting the fern family Pteridaceae (Polypodiales) were in existence by the Early Cretaceous. However, fossil evidence to corroborate this remains exceedingly rare. Burmese amber is an important source of new information on the radiation of derived fern lineages during the Cretaceous Terrestrial Revolution. This study describes Heinrichsia cheilanthoides gen. et sp. nov., a fern with suggested affinities to Pteridaceae, based on fertile foliage portions preserved in Early Cretaceous (~100 Ma) amber from Myanmar. Heinrichsia cheilanthoides is characterized by a pinnate‐pinnatifid frond that bears apical, marginal sori protected by a pseudoindusium. Sporangia are of the polypod type and contain tetrahedral‐globose, trilete spores with a striate perine. This discovery provides a new calibration point to test and refine molecular clock‐based concepts of the evolutionary history of the Pteridaceae. Heinrichsia cheilanthoides further substantiates the suggestion that the Cretaceous forests of Myanmar were home to a rich fern flora.     

Occurrence of an in situ fern grove in the Aptian Douiret Formation, Tataouine area, South-Tunisia

A level with fern stumps was discovered in the Aptian Douiret Formation, South-Tunisia. These stumps are preserved as external moulds or casts, without any preservation of anatomical structures. These stumps are considered to be affiliated with the numerous fossil plants assigned to the fern genera Alstaettia and Piazopteris that are widely distributed in coeval strata from the same region, either as leaf imprints or as permineralized remains. The record of in situ fossil forests for the Southern Hemisphere reveals that their systematic components are different, i.e. mainly corystosperms and/or conifers, and rarely under tidal influence. The way this fern grove settled in a margino-littoral environment is discussed.     

Polen y esporas de la Formación La Irene (Maastrichtiano), sudoeste de la provincia de Santa Cruz, Argentina: primeros resultados

Palynological assemblages are reported for the first time for the La Irene Formation, southwestern Santa Cruz Province, Argentina. This unit represents the sedimentation during a regressive period from transitional or delta-plain to fluvial environments. Palynological assemblages are scarce and moderately well preserved. They mainly consist of continental elements (wood remains, cuticles, spores and pollen grains) with scarse marine palynomorphs (dinoflagellate cysts). The spore-pollen assemblages are dominated by fern spores, followed by angiosperm and gymnosperm pollen grains. Bryophyte spores and fungal remains are also present. Among ferns, Cyatheaceae and spores of Laevigatosporites, of uncertain affinity, are dominant. Of the angiosperm pollen, those of Chloranthaceae (Clavatipollenites sp.) and Arecaceae (Arecipites spp., Longapertites sp., Spinizonocolpites hialinus Archangelsky and Zamaloa) are the most abundant. Pollen of Liliaceae (Liliacidites spp.), Proteaceae (Proteacidites sp., Peninsulapollis gillii (Cookson) Dettmann and Jarzen, Retidiporites camachoii Archangelsky) and Ericaceae (Ericipites scabratus Harris) are also present. Gymnosperm pollen is represented by Podocarpaceae (Podocarpidites spp.) and Ephedraceae (Equisetosporites sp.). These palynological suites would represent a fern-angiosperm dominated coastal vegetation, developed under warm and at least locally humid climatic conditions. La Irene Formation is considered Maastrichtian in age based on stratigraphic evidence, which is, in turn consistent with the ages suggested by the species ranges and the similarities observed with others previously described assemblages. This is the southernmost record of Spinizonocolpites, similar to the extant mangrove palm Nypa.     

Early devonian flora of the trout valley formation of northern maine

The data on the plant fossils of the Trout Valley Formation of north-central Maine, U.S.A., are summarized. Evidence from lithological correlation, from plant megafossils and paleopalynology suggest a late Early Devonian (Emsian) age for the Formation. Plant megafossils present are several species of Psilophyton, Pertica quadrifaria, Kaulangiophyton akantha, Prototaxites sp., Drepanophycus sp., Taeniocrada sp. and Thursophyton sp. The microfossil assemblage, through poorly preserved, contains Deltoidospora sp., Apiculiretusispora sp., Emphanisporites rotatus, E. annulatus, Clivosisporites cf. C. verrucata, Grandispora sp. and a chitinozoan, Sphaerochitina sp. The ancient landscape is suggested as having been an area a few square miles in extent with modest relief and comparable to a modern brackish or fresh-water marsh. Three other floras, one in Maine, a second in New Brunswick, Canada, and a third along the coast of Gaspé Bay, Quebec, are thought to have been at least partially contemporaneous with the Trout Valley flora.     

Effects of Lead on Ultrastructure of Isoetes sinensis Palmer (Isoetaceae), a Critically Endangered Species in China

Isoetes sinensis Palmer (Isoetaceae) is a critically endangered fern that is a marsh plant (that is an aquatic or amphibious plant) in China. To evaluate damage or influence of lead (Pb) on cell ultrastructure in I. sinensis, we used 2000mg·L^-1 Pb(NO₃)₂ solution to treat I. sinensis for 35d, and used transmission electron microscope (TEM) to observe the cell ultrastructure of leaf blades and roots of the plant. Our results indicated that Pb induced distinct changes of the organelles including chloroplast, mitochondria, nucleolus and vacuole. The level of damage organ was lower leaf > upper leaf > root The typical performance of the damages caused by lead shown that part of the nucleolus cracked; the cristae dilated, matrix vacuolized and membrane structure blurred in mitochondria; the vacuole cracked; grana lamella decreased, stroma lamella loosed, starch grains decreased, and membrane structure was disrupted in chloroplasts; Pb deposits were present on cell wall. The damages to chloroplasts and mitochondria were relatively severe, while damage to the nucleus was relatively lighter. The damage to the cell ultrastructure of leaf blades with direct contact with Pb was more severe than that without direct contact with Pb.     

Plant cytoplasm preservation in a baked root of Abies

Plant cytoplasm was thought impossible to be preserved in fossils. This stereotype is now undermined by increasing reports of plant cytoplasm in fossils. Previous simulations of high temperature preservation for plant cytoplasm were performed at short time scales, leaving the effectiveness and durability of such preservation an open question. Here we attempt to investigate the long time effect of high temperature on plant cytoplasm preservation in a root of Abies concolor that was baked in the wild at least 8 years ago. Light microscopy (LM), scanning electron microscopy (SEM), and transmission electron microscopy (TEM) indicate that cytoplasm and possible organelles such as mitochondria can be preserved in the baked tissue. The high frequency of wildfire in nature suggests that the potential for plant cytoplasm preservation in fossils is greater than commonly assumed. The good preservation of plant cytoplasm in the studied material after 8 years indicates that plant cytoplasm indeed can be preserved longer than previously proven.     

Mineralized soft-tissue structure and chemistry in a mummified hadrosaur from the Hell Creek Formation,North Dakota (USA)

An extremely well-preserved dinosaur (Cf. Edmontosaurus sp.) found in the Hell Creek Formation (Upper Cretaceous, North Dakota) retains soft-tissue replacement structures and associated organic compounds. Mineral cements precipitated in the skin apparently follow original cell boundaries, partially preserving epidermis microstructure. Infrared and electron microprobe images of ossified tendon clearly show preserved mineral zonation, with silica and trapped carbon dioxide forming thin linings on Haversian canals within apatite. Furthermore, Fourier transform infrared spectroscopy (FTIR) of materials recovered from the skin and terminal ungual phalanx suggests the presence of compounds containing amide groups. Amino acid composition analyses of the mineralized skin envelope clearly differ from the surrounding matrix; however, intact proteins could not be obtained using protein mass spectrometry. The presence of endogenously derived organics from the skin was further demonstrated by pyrolysis gas chromatography mass spectrometry (Py-GCMS), indicating survival and presence of macromolecules that were in part aliphatic (see the electronic supplementary material).     

Plant Parasitic Nematodes Associated with Leatherleaf Fern

Seven species of plant parasitic nematodes were found to be associated with leatherleaf fern (Rumohra adiantiformis) in central Florida. Of these, Pratylenchus penetrans, Tylenchorhynchus claytoni, and Criconemoides curvatum were commonly encountered. Nematode communities generally included two or three species of plant parasitic nematodes, with greatest diversity in nematode species occurring in ferneries shaded by oak trees. Species diversity was not correlated with fernery age. Leatherleaf fern was tolerant of P. penetrans and T. claytoni in microplot tests.     

Holmesopteris faironiae,fougère nouvelle du Carbonifère supérieur des Monts Cantabriques (NW de l'Espagne) et de Belgique

Isolated fern rachises have been discovered in twoWestphalian A coal balls of Belgium and Spain. They represent a new taxon, Holmesopteris faironiae n.gen., n.sp., characterized by the special location of the protoxylem poles. The morphology, the anatomy and the foliar branching of this new taxon suggest that it may belong to the family Botryopteridaceae; Stephanian species of the genus Botryopteris seem closely related to the new fern.