Under pressure? Epicormic shoots and traumatic growth zones in high-latitude Triassic trees from East Antarctica.

Background and Aims: Investigating the biology of trees that were growing at high latitudes during warmer geological periods is key to understanding the functioning of both past and future forest ecosystems. The aim of this study is to report the first co-occurrence of epicormic shoots and traumatic growth zones in fossil trees from the Triassic of Antarctica and to discuss their biological and environmental implications. Methods: Permineralized woods bearing scars of epicormic shoots were collected from the Triassic Fremouw Formation in Gordon Valley, Central Transantarctic Mountains, Antarctica in 2010. Samples from different portions of three specimens were prepared using standard thin section and hydrofluoric (HF) acid peel techniques, and anatomical details were studied in transmitted light. Key Results: The fossil woods represent the outer part of trunks, with at least 40 growth rings that are 0.2-4.8 mm in width. Anatomical comparisons suggest that they represent a new tree taxon for the Triassic of Antarctica. Numerous small epicormic shoots can be seen crossing the wood almost horizontally and are locally branched. Each specimen also contains several occurrences of traumatic growth zones located in the early wood, in the cells produced either at the very start of the growing season or slightly later. Conclusions: This is the first report of epicormic shoots and traumatic growth zones in the wood of a Triassic tree from Antarctica. Their co-occurrence indicates that these trees from Gordon Valley were subjected to environmental stresses not seen in Triassic trees previously described from this region. This suggests that they had a different biology and/or were growing in a different habitat, which offers a new glimpse into the diversity of high-latitude trees in the Triassic greenhouse climate.

Early Devonian (~410 mya) microfossils resembling Characiopsis (Tribophyceae) and Characium (Chlorophyceae).

Unusual microfossils that occurred associated with fungal spores in the Lower Devonian (~410 mya) Windyfield chert from Scotland were composed of a narrow stipe (2.5-9 µm long) to which was attached an obovoid or elongate drop-shaped cell up to 14 µm long; a basal attachment pad was present in several specimens. The fossils were strikingly similar morphologically to certain present-day unicellular freshwater Tribophyceae and Chlorophyceae, but affinities to the fungal phylum Chytridiomycota also cannot be ruled out. This discovery adds to the inventory of distinctive microbial morphologies in the early non-marine paleoecosystems.

Triassic leech cocoon from Antarctica contains fossil bell animal.

Our understanding of the evolution of life on Earth is limited by the imperfection of the fossil record. One reason for this imperfect record is that organisms without hard parts, such as bones, shells, and wood, have a very low potential to enter the fossil record. Occasionally, however, exceptional fossil deposits that preserve soft-bodied organisms provide a rare glimpse of the true biodiversity during past periods of Earth history. We here present an extraordinary find of a fossil ciliate that is encased inside the wall layer of a more than 200 Ma leech cocoon from Antarctica. The microfossil consists of a helically contractile stalk that attaches to a main body with a peristomial feeding apparatus and a large C-shaped macronucleus. It agrees in every aspect with the living bell animals, such as Vorticella. Vorticellids and similar peritrichs are vital constituents of aquatic ecosystems worldwide, but so far have lacked any fossil record. This discovery offers a glimpse of ancient soft-bodied protozoan biotas, and also highlights the potential of clitellate cocoons as microscopic "conservation traps" comparable to amber.

Morphological and functional stasis in mycorrhizal root nodules as exhibited by a Triassic conifer.

Mycorrhizal root nodules occur in the conifer families Araucariaceae, Podocarpaceae, and Sciadopityaceae. Although the fossil record of these families can be traced back into the early Mesozoic, the oldest fossil evidence of root nodules previously came from the Cretaceous. Here we report on cellularly preserved root nodules of the early conifer Notophytum from Middle Triassic permineralized peat of Antarctica. These fossil root nodules contain fungal arbuscules, hyphal coils, and vesicles in their cortex. Numerous glomoid-type spores are found in the peat matrix surrounding the nodules. This discovery indicates that mutualistic associations between conifer root nodules and arbuscular mycorrhizal fungi date back to at least the early Mesozoic, the period during which most of the modern conifer families first appeared. Notophytum root nodules predate the next known appearance of this association by 100 million years, indicating that this specialized form of mycorrhizal symbiosis has ancient origins.

Paleomycology of the Princeton Chert II. Dark-septate fungi in the aquatic angiosperm Eorhiza arnoldii indicate a diverse assemblage of root-colonizing fungi during the Eocene.

Tissues of the extinct aquatic or emergent angiosperm, Eorhiza arnoldii incertae sedis, were extensively colonized by microfungi, and in this study we report the presence of several types of sterile mycelia. In addition to inter- and intracellular proliferation of regular septate hyphae, the tissues contain monilioid hyphae with intercalary branching. These filamentous mycelia are spatially associated with two distinct morphotypes of intracellular microsclerotia. These quiescent structures are morphologically similar to loose and cerebriform microsclerotia found within the living tissues of some plants, which have been attributed to an informal assemblage of dematiaceous ascomycetes, the dark-septate endophytes. While there are significant challenges to interpreting the ecology of fossilized fungi, these specimens provide evidence for asymptomatic endophytic colonization of the rooting structures of a 48.7 million year old aquatic angiosperm.

Paleomycology of the Princeton Chert I. Fossil hyphomycetes associated with the early Eocene aquatic angiosperm, Eorhiza arnoldii.

The Eocene (~ 48.7 Ma, Ypresian-Lutetian) Princeton Chert of British Columbia, Canada, has long been recognized as a significant paleobotanical locality, and a diverse assemblage of anatomically preserved fossil plants has been extensively documented. Co-occurring fossil fungi also have been observed, but the full scope of their diversity has yet to be comprehensively assessed. Here, we present the first of a series of investigations of fossilized fungi associated with the silicified plants of the Princeton Chert. This report focuses on saprotrophic, facultative-aquatic hyphomycetes observed in cortical aerenchyma tissue of an enigmatic angiosperm, Eorhiza arnoldii. Our use of paleontological thin sections provides the opportunity to observe and infer developmental features, making it possible to more accurately attribute two hyphomycetes that were observed in previous studies. These comprise multiseptate, holothallic, chlamydospore-like phragmoconidia most similar to extant Xylomyces giganteus and basipetal phragmospore-like chains of amerospores like those of extant Thielaviopsis basicola. We also describe a third hyphomycete that previously has not been recognized from this locality; biseptate, chlamydosporic phragmoconidia are distinguished by darkly melanized, inflated apical cells and are morphologically similar to Brachysporiella rhizoidea or Culcitalna achraspora.

Antarctic glossopterid diversity on a local scale: the presence of multiple megasporophyll genera, Upper Permian, Mt. Achernar, Transantarctic Mountains, Antarctica.

PREMISE OF THE STUDY: The glossopterids are a group of plants that thrived during a time of global warming similar to what is happening on the Earth today as well as the transition from archaic plant groups to the ancestors of modern groups. The diversity of the glossopterid clade is based on the megasporangiate structures assigned to the group, because the vegetative and pollen-bearing structures vary little. The presence of numerous reproductive genera from a single Upper Permian locality in the central Transantarctic Mountains provides important data on local glossopterid diversity in Antarctica. METHODS: Impression/compression fossils were imaged with a Leica 5000C digital camera on a dissecting microscope or a Fujifilm FinePix S1pro digital camera. KEY RESULTS: Two megasporangiate taxa are described: Scutum leiophyllum, which represents the first confirmed record of the genus in Antarctica, and Lidgettoniopsis ramulus, a new morphology consisting of a pinnate structure with oppositely attached megasporophylls. Plumsteadia ovata specimens indicate that this genus can be larger than previously recorded and illustrate the vegetative surface with a distinct midrib. CONCLUSIONS: The presence of a laminar, multiovulate structure and a pinnate structure at the same site indicates that local-level glossopterid diversity in Antarctica is greater than previously hypothesized. The discovery of a new megasporophyll morphology in Antarctica (confirming the presence of three distinctive morphologies on the continent) shows that Antarctic glossopterid heterogeneity is on a par with other Gondwanan continents. The diversity of the Antarctic landscape reveals that high polar latitudes can sustain a diverse ecosystem during times of global warming.

Root suckering in a Triassic conifer from Antarctica: paleoecological and evolutionary implications.

PREMISE OF THE STUDY: Although root suckering and other types of sprouting are well studied in extant woody plants, little is known about the distribution of these traits at a macroevolutionary scale. Anatomically preserved fossil plants represent an excellent but understudied source of information of the distribution of sprouting behavior through time and across taxa. METHODS: A block of silicified peat collected in the Middle Triassic Fremouw Formation at the Fremouw Peak locality, Central Transantarctic Mountains, Antarctica, contains a group of anatomically preserved roots of the fossil conifer Notophytum krauselii that bear young shoots. The specimen was prepared using the standard acetate peel technique and studied in reflected and transmitted light. KEY RESULTS: Young sucker shoots bearing well-preserved leaves are produced in groups in some areas of the Notophytum roots. CONCLUSIONS: The production of root suckers in Notophytum indicates that some of the trees growing in polar forests during the Triassic could respond to environmental stresses by regenerating their vegetative structures and had the potential to reproduce vegetatively. The specimens also represent the first anatomical evidence of root suckering in any fossil seed plant, and its occurrence in an early putative podocarp supports the idea that this trait might be ancestral in at least some extant conifer families.

Filamentous cyanobacteria preserved in masses of fungal hyphae from the Triassic of Antarctica.

Permineralized peat from the central Transantarctic Mountains of Antarctica has provided a wealth of information on plant and fungal diversity in Middle Triassic high-latitude forest paleoecosystems; however, there are no reports as yet of algae or cyanobacteria. The first record of a fossil filamentous cyanobacterium in this peat consists of wide, uniseriate trichomes composed of discoid cells up to 25 µm wide, and enveloped in a distinct sheath. Filament morphology, structurally preserved by permineralization and mineral replacement, corresponds to the fossil genus Palaeo-lyngbya, a predominantly Precambrian equivalent of the extant Lyngbya sensu lato (Oscillatoriaceae, Oscillatoriales). Specimens occur exclusively in masses of interwoven hyphae produced by the fungus Endochaetophora antarctica, suggesting that a special micro-environmental setting was required to preserve the filaments. Whether some form of symbiotic relationship existed between the fungus and cyanobacterium remains unknown.

Fungi and fungal interactions in the Rhynie chert: a review of the evidence, with the description of Perexiflasca tayloriana gen. et sp. nov.†.

The Lower Devonian Rhynie chert is one of the most important rock deposits yielding comprehensive information on early continental plant, animal and microbial life. Fungi are especially abundant among the microbial remains, and include representatives of all major fungal lineages except Basidiomycota. This paper surveys the evidence assembled to date of fungal hyphae, mycelial cords and reproductive units (e.g. spores, sporangia, sporocarps), and presents examples of fungal associations and interactions with land plants, other fungi, algae, cyanobacteria and animals from the Rhynie chert. Moreover, a small, chytrid-like organism that occurs singly, in chain-like, linear arrangements, planar assemblages and three-dimensional aggregates of less than 10 to [Formula: see text] individuals in degrading land plant tissue in the Rhynie chert is formally described, and the name Perexiflasca tayloriana proposed for the organism. Perexiflasca tayloriana probably colonized senescent or atrophied plant parts and participated in the process of biological degradation. The fungal fossils described to date from the Rhynie chert constitute the largest body of structurally preserved evidence of fungi and fungal interactions from any rock deposit, and strongly suggest that fungi played important roles in the functioning of the Early Devonian Rhynie ecosystem.This article is part of a discussion meeting issue 'The Rhynie cherts: our earliest terrestrial ecosystem revisited'.

Protoascon missouriensis, a complex fossil microfungus revisited.

The Carboniferous microfungus Protoascon missouriensis has been interpreted variously as an ascomycete, chytridiomycete, zygomycete and oomycete. We offer a more complete interpretation based on a re-examination of the type material that suggests the fossil represents an (a)zygosporangium-suspensor complex of a zygomycete comparable to some modern members of the Mucorales.

Organization, anatomy, and fungal endophytes of a Triassic conifer embryo.

PREMISE OF THE STUDY: Much is known about the Paleozoic conifers and the conifers assignable to extant families that appear in the Jurassic; however, relatively little is known about the transitional conifers of the early Mesozoic, especially the Voltziales. To better understand the evolution of this group, we aim to increase knowledge of voltzialean anatomy and morphology. A fossil embryo of the group is described in this study. • METHODS: Several permineralized seeds, one containing a well-preserved embryo, were collected from the early Middle Triassic Fremouw Peak locality in the Central Transantarctic Mountains. Samples were prepared using the standard acetate peel technique and studied in transmitted light. • KEY RESULTS: The embryo belongs to Parasciadopitys aequata, a member of the Voltziales. The embryo and megagametophyte tissues are exquisitely preserved. The embryo is colonized by two distinct fungi. Sporocarps of the fungi are found in the megagametophyte and in the space between the megagametophyte and nucellus. The additions of embryo and megagametophyte characters to the description of P. aequata have made it one of the most completely known fossil taxa reproductively. • CONCLUSIONS: Preservation of fossil embryos, although rare, can expand the array of characters available in tracing the evolutionary history of plants. The embryo of P. aequata shares similarities with embryos of other extinct and extant conifer families. The association of the embryo with Combresomyces cornifer and Mycocarpon asterineum increases our understanding of the roles of microorganisms in Triassic ecosystems.

Seed ferns from the late Paleozoic and Mesozoic: Any angiosperm ancestors lurking there?

Five orders of late Paleozoic-Mesozoic seed ferns have, at one time or another, figured in discussions on the origin of angiosperms, even before the application of phylogenetic systematics. These are the Glossopteridales, Peltaspermales, Corystospermales, Caytoniales, and Petriellales. Although vegetative features have been used to suggest homologies, most discussion has focused on ovulate structures, which are generally interpreted as megasporophylls bearing seeds, with the seeds partially to almost completely enclosed by the megasporophyll (or cupule). Here we discuss current information about the reproductive parts of these plants. Since most specimens are impression-compression remains, homologizing the ovulate organs, deriving angiospermous homologues, and defining synapomorphies remain somewhat speculative. Although new specimens have increased the known diversity in these groups, a reconstruction of an entire plant is available only for the corystosperms, and thus hypotheses about phylogenetic position are of limited value. We conclude that, in the case of these seed plants, phylogenetic analysis techniques have surpassed the hard data needed to formulate meaningful phylogenetic hypotheses. Speculation on angiosperm origins and transitional stages in these fossils provides for interesting discussion, but currently it is still speculation, as the role of these groups in the origin of angiospermy continues to be cloaked in Darwin's mystery.

Development and ecological implications of dormant buds in the high-Paleolaltitude Triassic sphenophyte Spaciinodum (Equisetaceae).

Spaciinodum collinsonii, a Triassic sphenophyte from the central Transantarctic Mountains, Antarctica, is reinterpreted based on new material in order to clarify discrepancies from previous work and to detail the development and ecology of the Spaciinodum plant. Vegetative stems have alternating nodes and internodes, nodes distinguished by a solid diaphragm of tissue, internodes by the presence of vallecular (cortical) and carinal canals, and a hollow pith. Whorls of branches arise immediately above the nodes, alternating with the leaves of the subjacent nodes. Branches develop in the cortex and are anatomically similar to the stems. While Spaciinodum is similar to extant Equisetum, it is distinctive in that its large vallecular canals form a complete ring within the cortex and are separated only by thin fimbrils of tissue. The majority of specimens of Spaciinodum are now believed to be dormant buds with condensed nodes and internodes, with progressively longer internodal regions more basally. More apical portions of buds have cellular internodes because the areas where the canals will form have not yet ruptured from elongation. The abundance of buds and the absence of elongated stems in the permineralized peat deposit suggest that Spaciinodum underwent dormancy during the dark Antarctic winters.

Secondary phloem anatomy of Cycadeoidea (Bennettitales).

Secondary phloem anatomy of several species of Cycadeoidea is described from trunks in the Wieland Collection, Peabody Museum of Natural History. The trunks were collected from the Lakota Formation, Lower Cretaceous, Black Hills of South Dakota. Secondary phloem is extensively developed and consists of alternating, tangential bands of fibers and sieve elements, with rare phloem parenchyma. Uniseriate rays, 2-22 cells high, occur between every one to three files of the axial system. Fibers are long, more than 1200 µm, approximately 26.6-34.2 µm in diameter, and have slit-like apertures on the lateral walls. Sieve elements range from 16-25 µm in diameter and are up to 500 µm long. Elliptical sieve areas appear on both end and radial walls and measure 10 µm across; minute spots, which may represent sieve pores, are present within the sieve areas. Secondary phloem of North American Cycadeoidea is similar in organization (alternating tangential bands) and cell types (sieve cells, fibers, axial parenchyma) to that known in other extant and fossil cycadophytes and some seed ferns. The unusual pattern of cell types and thickness of secondary phloem is discussed in the context of plant habit, phloem efficiency, and potential phylogenetic importance.

Cataphylls of the Middle Triassic cycad Antarcticycas schopfii and new insights into cycad evolution.

Cataphylls associated with the Middle Triassic stem genus Antarcticycas are described, and their impact on understanding cycad evolution is discussed. The cataphylls of Antarcticycas are triangular in outline and flattened adaxially with lateral flanges. The outer surfaces are covered with a ramentum of filamentous hairs, the epidermis is a single cell layer thick, and the ground tissue is parenchymatous with mucilage canals and sclereids. Vascular bundles form a distinct inverted omega-shaped pattern characteristic of the Cycadales observed in petioles of extant species. The structures in Antarcticycas are interpreted as cataphylls based on overall morphology, presence of straight vascular strands in the cortex of the associated stem, and lack of fascicular cambia in the vascular bundles. Because much of the overall diversity of Cycadales is represented by fossils, integrating fossil taxa into explicit phylogenetic hypotheses is important for understanding cycad evolution. Therefore, character and minimum age mapping were performed on a phylogeny of extant and fossil taxa including Antarcticycas. The results suggest that major extant lineages of Cycadales had diverged by the Permian to Triassic and that certain synapomorphies for Cycadales had evolved by the Permian. Evidence of insect feeding on Antarcticycas suggests that associations between cycads and insects are ancient.

Anatomy of umkomasia (corystospermales) from the triassic of antarctica.

The permineralized, corystosperm, cupulate, ovule-bearing organ Umkomasia resinosa is described from the early Middle Triassic of Antarctica. This is the first description of anatomically preserved Umkomasia, which consists of a determinate cupulate branch with helically arranged, recurved, pedicellate megasporophylls, each of which bears one or two abaxially attached unitegmic ovules. Cupules are ovoid, bilobed with elongate ventral and dorsal openings or unlobed with a single ventral opening, and have a two-zoned parenchymatous cortex and abundant secretory cavities. Ovules are small, orthotropous, and possess a thin integument that contains numerous secretory cavities. The ovules are broadly attached at the base, with a bifid integumentary apex that extends past the cupule lobes. The cupulate branch displays stem-like anatomy, producing paired traces into each cupule stalk. These structurally preserved ovulate organs can be related to other corystosperm organs from Antarctica, particularly the pollen-organ Pteruchus fremouwensis. Both anatomical and morphological features support interpretation of corystosperm reproductive structures as branching systems rather than as compound sporophylls. As a result of an increased understanding of the organization of Umkomasia, it appears doubtful that any direct relationship exists between the corystosperm and angiosperm lineages.

Ashicaulis woolfei n. sp.: additional evidence for the antiquity of osmundaceous ferns from the Triassic of Antarctica.

Numerous small fern trunks and dispersed osmundaceous frond fragments occur within a Middle Triassic silicified peat near Fremouw Peak in the Transantarctic Mountains of Antarctica. These specimens form the basis of a new species of osmundaceous ferns that further helps to characterize the early Mesozoic vegetation of high latitude Gondwana. Ashicaulis woolfei n. sp. consists of small, upright trunks with a persistent armor of frond bases, adventitious roots, and vegetative frond parts. In cross section the trunks are ∼2.5 cm in diameter and include up to 45 frond bases. Stems range from 5 to 8 mm in diameter with a xylem cylinder of 8-9 xylem segments separated by leaf gaps. Phyllotaxy is variable, approaching 2/5 or 3/8, with 10-12 frond traces in the cortex. Stipes have parenchymatous, stipular wings that are usually devoid of sclerenchyma; fronds are pinnate with alternate-subopposite pinnatifid pinnules. Although the absence of fertile pinnules and sporangia precludes assigning the fossils to a living genus, this species demonstrates that ferns with stelar architecture and histology similar to Osmunda subgenus Osmundastrum (Osmundaceae) were present in the Southern Hemisphere by the mid-Triassic.