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772:- fossilised droppings - give a useful insight of what animals ate, even if the animals cannot be identified. Coprolites found in the Rhynie chert are typically between 0.5 and 3 mm in size, and contain a variety of contents. Analysis of coprolites allows the identification of different feeding modes, including detritovory and herbivory; some coprolites are so densely packed with spores that it is possible that these made up a substantial proportion of some organisms' diets. The
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organism to be identified - are identical to those found elsewhere in "normal" environments. There is no clear-cut evidence that the plants of the Rhynie assemblage were specifically adapted to stressed environments, and it is likely that the flora in fact represents those members of the global fauna that happened to be capable of colonising and surviving a hot spring environment by virtue of fortuitous preadaptations.
20:
325:
716:, has been recovered from the Rhynie chert. The lichen comprises a thallus, made of layered, aseptate hyphae; a number of depressions are formed on its top surface. Each depression contains a net of hyphae holding a sheathed cyanobacterium. The fungus appears to be related to the Zygomycetes, and the photobiont resembles the
370:, the sediment formed by the hot springs. These two colonisers were subsequently joined by other genera. The time between sinter deposition events was too short to allow the populations to develop to climax communities, and correspondingly early colonisers appear most frequently, pseudo-randomly, in logged sequences.
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displayed three responses to fungal infestation: the hyphae of some (mutualistic) colonists were encased by plant cell walls; other (parasitic) fungi were met with typical host responses of increased rhizome cell size; while yet other fungi solicited an increase in thickness and pigmentation of cell
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and almost instantaneously, in much the same fashion that organisms are petrified by hot springs today - although the astounding fidelity of preservation has not been found in recent deposits. Hot springs, with temperatures between 90 and 120 °C (194 and 248 °F), were active in a number of
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which are typically alkaline (pH 8.7) and tepid 20 to 28 °C (68 to 82 °F). The springs were periodically active, and flowed into an alluvial plain containing small lakes. By analogy with
Yellowstone, the chert itself probably formed in a marshy area towards the latter end of the extent of
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Analysis of spores shows that the flora was lacking in some elements common elsewhere at this time, likely due to its setting in a mountainous region, rather than in a lowland flood plain like most other fossil deposits. However, the spores, which are distinctive enough to permit their producing
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species found in the deposit were predators: it is possible for many of the arthropods to deduce their likely ecological role, however, it is unclear if this community was representative of a typical terrestrial arthropod community of the time, or rather was specific to the stressed Rhynie
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episodes; the water had probably cooled to under 30 °C (86 °F) before it reached the fossilised organisms. Their activity is preserved in 53 beds, 80 mm (3 in) thick on average, over a 35.41 m (116.17 ft) sequence, interbedded with sands, shales and
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may be present, and parasitism is common; one individual has even been found parasitising a germinating gametophyte. The fungi were aquatic, and grew in both plants and algae; they are also found preserved "loose" in the chert matrix. Their flagellate spores are preserved.
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outwash from the springs. Living vegetation covered around 55% of the land area, with litter covering 30% and the remaining 15% of the ground being bare. A braided river flowing to the north periodically deposited the sandy layers found in cores when it flooded its banks.
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Fossils were formed as silica formed in the hot springs themselves; when silica-rich water flooded the surrounding areas; and when it permeated into the surrounding soil. The texture of the sinter formed resemble those found today in freshwater streams at
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As a result of its exquisite preservation, the Rhynie chert boasts the most diverse non-marine fauna of its time, and is important for our understanding of arthropod terrestrialisation. Typical members of the Rhynie chert arthropod fauna include the
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the branching patterns of the early plants emerged is possible, whereas typical fossils only show that branching was present. The analysis of rhizomes and rhizoids makes it possible to discern which plants had an active water uptake system (e.g.
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299:", a sediment with a botryoidal form reminiscent of modern vent margins, is also found. Spores collected from within surrounding rocks had been heated to different degrees, implying a complex history of local heating by volcanic processes.
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Rice, C. M., Ashcroft, W. A., Batten, D. J., Boyce, A. J., Caulfield, J. B. D., Fallick, A. E., Hole, M. J., Jones, E., Pearson, M. J., Rogers, G., Saxton, J. M., Stuart, F. M., Trewin, N. H. & Turner, G. (1995).
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worked furiously to describe the plant fossils between 1917 and 1921. The arthropods were examined soon afterwards by different workers. Interest in the chert then waned until the field was reinvigorated by
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allows spore genera to be matched with their producers - something that is otherwise very difficult to do. The chert also allows the identification of the gametophyte phases of taxa such as
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Until recently, the Rhynie chert was the only such deposit known from the geological record, although recent work has turned up other localities from different time periods and continents.
271:- which speak of local volcanic activity. Deposition was very rapid. The fluids originated from a shallowly dipping extensional fault system to the west, which bounded an extensional half-
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walls. Once inside a plant cell, fungi produced spores, which are found in decaying plant cells; the cells may have decayed as a defence mechanism to prevent the fungi from spreading.
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are found in the fossil record, their presence is usually the subject of much controversy, for their simple form is difficult to distinguish from inorganic structures such as bubbles.
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yet known from the Rhynie are the
Zygomycota (although they may have formed lichens - see later), and the Basidiomycota, the latter of which may not even have evolved by Rhynie time.
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Poinar Jr, George; Kerp, Hans; Hass, Hagen (2008). "Palaeonema phyticum gen. n., sp. n. (Nematoda: Palaeonematidae fam. n.), a
Devonian nematode associated with early land plants".
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Krings; Kerp, Hans; Hass, Hagen; Taylor, Thomas N.; Dotzler, Nora (2007). "A filamentous cyanobacterium showing structured colonial growth from the Early
Devonian Rhynie chert".
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absence. The fossils are filamentous, around 3 μm in diameter, and grew on plants and the sediment itself. They occasionally form structured colonies which go on to create
671:, growing as a mound a metre or more taller than anything in the community, whose isotopic composition varied like a saprotroph and whose septate pores resemble those of fungi.
243:, allowing an insight into the evolution of the chert over time, were drilled in 1988 and 1997, accompanied by further trenching efforts, which unearthed the Windyfield chert.
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The chert was discovered by
William Mackie while mapping the western margin of the Rhynie basin in 1910–1913. Trenches were cut into the chert at the end of this period, and
1979:
Taylor, T.N.; Hass, H; Kerp, H; Krings, M; Hanlin, RT (2005). "Perithecial ascomycetes from the 400 million year old Rhynie chert: an example of ancestral polymorphism".
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228:
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Powell, C. L.; Trewin, N. H.; Edwards, D. (2000). "Palaeoecology and plant succession in a borehole through the Rhynie cherts, Lower Old Red
Sandstone, Scotland".
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response. Herbivory is also evident, judging by boring and piercing wounds in various states of repair, and the mouthparts of arthropods. In addition, a fossil
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in high fidelity, gives a unique opportunity to observe interactions between species and kingdoms. There is evidence of parasitic behaviour by fungi on algae
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Fungal interactions are known to promote speciation in modern plants, and presumably also affected
Devonian diversity by providing a selection pressure.
422:). In some cases, it is possible to see different mechanisms of repairing wounds, and to deduce that they were caused by fungal or bacterial infection.
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741:
530:
557:
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Wellman, Charles H. (2006). "Spore assemblages from the Lower
Devonian 'Lower Old Red Sandstone' deposits of the Rhynie outlier, Scotland".
884:
2357:
Garwood, Russell; Dunlop, Jason (2015). "The walking dead: Blender as a tool for paleontologists with a case study on extinct arachnids".
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Habgood, K.S.; Hass, H.; Kerp, H. (2003). "Evidence for an early terrestrial food web: coprolites from the Early
Devonian Rhynie chert".
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Trewin, N.H.; Wilson, E. (2004). "Correlation of the Early
Devonian Rhynie chert beds between three boreholes at Rhynie, Aberdeenshire".
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in the late 1950s, and new material was collected by further trenching from 1963 to 1971. Since 1980, the chert has been examined by the
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Plants demonstrate best the great value of the exceptional preservation of the Rhynie chert. The presence of soft tissue, including
206:. A second unit, the Windyfield chert, is some 700 m from the Rhynie. The Rhynie chert extends for at least 80 m along
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The Rhynie chert contains exceptionally preserved plant, fungus, lichen and animal material preserved in place by an overlying
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forms are known - i.e. some forms grew specialised fruiting bodies while others did not show specialisation in this fashion.
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1357:"Geological setting of the Early Devonian Rhynie cherts, Aberdeenshire, Scotland: an early terrestrial hot spring system"
1127:"A new Middle – Late Jurassic flora and hot spring chert deposit from the Deseado Massif, Santa Cruz province, Argentina"
203:
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37:
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2079:
Berbee, Mary L.; Taylor, JW (2007). "Rhynie chert: a window into a lost world of complex plant?fungus interactions".
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2033:"Fungal endophytes in a 400-million-yr-old land plant: infection pathways, spatial distribution, and host responses"
1242:"Subaqueous silicification of the contents of small ponds in an Early Devonian hot-spring complex, Rhynie, Scotland"
1090:
Trewin (2003). "History of research on the geology and palaeontology of the Rhynie area, Aberdeenshire, Scotland".
978:"A high-precision U–Pb age constraint on the Rhynie Chert Konservat-Lagerstätte: time scale and other implications"
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to flattened charcoal films. On occasion, plants may have their vertical axes preserved in growth position, with
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388:, whereas the conventional record at its best allows no more than the counting of stomata. It has also enabled
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Kelman, Ruth; Feist, Monique; Trewin, Nigel H.; Hass, Hagen (2003). "Charophyte algae from the Rhynie chert".
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534:, which inhabited the alkaline freshwater pools towards the end of the sinter apron, has been characterized.
450:. Marked examples: centre – single corm with rhizoids; left – linked corms with rhizoids. Scale bar is 1 cm.
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Thin section of a piece of Rhynie chert viewed by transmitted light showing the cross-section of a stem of
619:, was found in the Rhynie chert, pushing dates for the origination of hexapods (a group that includes the
1944:
Taylor, T.N.; Remy, W.; Hass, H. (1992). "Fungi from the Lower Devonian Rhynie chert: Chytridiomycetes".
1651:"Palaeoecology and palaeophytogeography of the Rhynie chert plants: evidence from integrated analysis of
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Fayers; Trewin, Nigel H. (2003). "A review of the palaeoenvironments and biota of the Windyfield chert".
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Trewin, NH (1996). "The Rhynie cherts: an early Devonian ecosystem preserved by hydrothermal activity".
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1321:
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1792:"A harvestman (Arachnida: Opiliones) from the Early Devonian Rhynie cherts, Aberdeenshire, Scotland"
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Plants responded to fungal colonisation in different ways, depending on the fungus. The rhizoids of
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cyanobacteria are preserved in the Rhynie chert. The aquatic organisms are thought to belong to the
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The Chytridiomycetes, or Chytrids, are a basal group of fungi, closely related to the true fungi.
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2237:
2205:"The origin of herbivory on land: Initial patterns of plant tissue consumption by arthropods"
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Taylor, T.N.; Hass, H; Kerp, H (1997). "A cyanolichen from the Lower Devonian Rhynie chert".
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plants, with eggs, juveniles and adults all recorded from within their stomatal chambers.
239:, whose researchers confirmed that the chert was indeed produced in a hot spring setting.
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Contains useful reconstructions of both the plant associations, and the regional setting.
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Early Devonian sedimentary deposit exhibiting extraordinary fossil detail or completeness
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Ciba Foundation Symposium 202: Evolution of Hydrothermal Ecosystems on Earth (and Mars?)
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Wellman, Charles H.; Kerp, Hans; Hass, Hagen (2003). "Spores of the Rhynie chert plant
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Plants were only found on the land - none lived in the water of lakes or hot springs.
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The chert was formed when silica-rich water from volcanic springs rose rapidly and
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639:, oomycota (Peronosporomycetes) and glomeromycetes; indeed the only fungal groups
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typically grew on sandy surfaces, and is often preserved there in life position;
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Seven land plant taxa have been identified in the Rhynie and Windyfield cherts:
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Surface view of a polished piece of Rhynie chert showing many cross-sections of
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Krings, Michael; Taylor, TN; Hass, H; Kerp, H; Dotzler, N; Hermsen, EJ (2007).
1992:
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Engel, Michael S.; Grimaldi, DA (2004). "New light shed on the oldest insect".
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preservation, with individual cell walls easily visible in polished specimens.
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Earth and Environmental Science Transactions of the Royal Society of Edinburgh
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Earth and Environmental Science Transactions of the Royal Society of Edinburgh
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1583:"Life history biology of early land plants: Deciphering the gametophyte phase"
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Surface view of a polished piece of Rhynie chert showing many corms/tubers of
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Several putative chlorophytes have been discovered in the Rhynie assemblage (
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202:, so is effectively inaccessible to collectors; besides which, the site is a
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The largest organism present in Rhynie was probably a fungus, the enigmatic
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This fossil bed is remarkable for two reasons. First, the age of the site (
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The preservation of plants varies from perfect three-dimensional cellular
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The Rhynie chert ecosystem: a model for understanding fungal interactions
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147:) places it at an early stage in the colonisation of land. Second, these
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remnants detected in the plant material, and the breathing apparatus of
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Further, as plants are preserved in situ, the study of exactly how and
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Philosophical Transactions of the Royal Society B: Biological Sciences
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891:. Department of Earth Sciences, University of Bristol. Archived from
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1835:, the earliest known insect, from the Devonian of Rhynie, Scotland"
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416:), and which were likely to have colonised waterlogged surfaces (
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1196:. Novartis Foundation Symposia. Vol. 202. pp. 131–45.
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Garwood, Russell J; Oliver, Heather; Spencer, Alan R T (2019).
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The chytrids display a range of behaviour in the Rhynie chert.
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can be seen entering plant material, acting as decomposers and
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Transactions of the Royal Society of Edinburgh: Earth Sciences
1704:, The Biota of Early Terrestrial Ecosystems: The Rhynie Chert.
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Transactions of the Royal Society of Edinburgh: Earth Sciences
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Transactions of the Royal Society of Edinburgh: Earth Sciences
1502:"Embryophytic sporophytes in the Rhynie and Windyfield cherts"
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Transactions of the Royal Society of Edinburgh: Earth Sciences
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Transactions of the Royal Society of Edinburgh: Earth Sciences
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Transactions of the Royal Society of Edinburgh: Earth Sciences
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2461:"4 hundred million year old vesicular-arbuscular mycorrhizae"
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381:. This allows the study of structures such as the air spaces
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114:
110:
1943:
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Parry, S.F.; Noble S.R.; Crowley Q.G.; Wellman C.H. (2011).
512:, remains enigmatic, but may represent aquatic land plants.
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917:"A Devonian auriferous hot spring system, Rhynie, Scotland"
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The Rhynie chert, by preserving a snapshot of an ecosystem
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122:
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1978:
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2402:"Terrestrial invertebrates in the Rhynie chert ecosystem"
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Dunlop, J.A.; Anderson, L.I.; Kerp, H.; Hass, H. (2007).
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Sedimentary textures which appear to have formed in the
1751:"Early Terrestrial Animals, Evolution, and Uncertainty"
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1749:Garwood, Russell J.; Edgecombe, Gregory D. (2011).
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1125:Channing; Zamuner, Alba B.; ZÚÑiga, Adolfo (2007).
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1301:Geological Society of London, Special Publications
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1355:Rice, C.M.; Trewin, N.H.; Anderson, L.I. (2002).
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2239:The Origin And Early Evolution Of Plants On Land
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1546:(Kidston and Lang) Barghoorn and Darrah, 1938".
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822:
1588:Proceedings of the National Academy of Sciences
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1240:Trewin, N.H.; Fayers, S.R.; Kelman, R. (2003).
2400:Dunlop, Jason A.; Garwood, Russell J. (2017).
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631:Fungi known from the Rhynie chert include the
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961:: CS1 maint: multiple names: authors list (
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1829:Whalley, Paul; Jarzembowski, E. A. (1981).
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2459:Remy W, Taylor TN, Hass H, Kerp H (1994).
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2017:
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1581:Taylor, T. N.; Kerp, H.; Hass, H. (2005).
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151:are famous for their exceptional state of
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921:Journal of the Geological Society, London
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175:) can be seen in cross-sections. Fungal
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425:The preservation of spores attached to
198:, in a small field near the village of
2706:
1191:
1089:
213:
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2119:Review of Palaeobotany and Palynology
838:"An introduction to the Rhynie chert"
392:to firmly deduce that plants such as
194:The bed is under at least 1 metre of
86:). It is exposed near the village of
23:Hand sample of the Rhynie chert from
969:
796:are also found in the Rhynie chert.
98:, is located some 700 m away.
2615:
2521:Taylor, T.N.; Taylor, E.L. (2000).
204:Site of Special Scientific Interest
13:
2514:
1695:
1659:Proceedings of the Royal Society B
1246:Canadian Journal of Earth Sciences
14:
2770:
2571:
2236:Kenrick, P.; Crane, P.R. (2000).
1755:Evolution: Education and Outreach
1364:Journal of the Geological Society
982:Journal of the Geological Society
109:fossil bed consists of primitive
2724:Protected areas of Aberdeenshire
2579:"Rhynie Chert Learning Resource"
2222:10.1111/j.1744-7917.2007.00152.x
2093:10.1111/j.1469-8137.2007.02080.x
2050:10.1111/j.1469-8137.2007.02008.x
674:
528:). A well-preserved charophyte,
320:stems (axes). Scale bar is 1 cm.
291:themselves are preserved with a
2285:
2245:. University of Chicago Press.
1822:
730:
623:) back to the Silurian period.
302:
2139:10.1016/j.revpalbo.2007.05.002
907:
806:Evolutionary history of plants
615:), which resembles the modern
1:
1331:10.1144/GSL.SP.2000.180.01.23
816:
537:
2749:Environment of Aberdeenshire
2528:(Free access @ Google books)
2465:Proc. Natl. Acad. Sci. U.S.A
1649:Wellman, Charles H. (2004).
402:aquatic, as once believed.
129:, lichens, algae and fungi.
7:
2203:Labandeira, CONRAD (2007).
1038:Scottish Journal of Geology
799:
679:In the rare instances that
189:
10:
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2306:10.1163/156854108783360159
1993:10.3852/mycologia.97.1.269
1946:American Journal of Botany
705:
82:detail or completeness (a
2729:Devonian System of Europe
2623:
2581:. University of Aberdeen.
2336:10.1017/S0263593300000754
1808:10.1017/S0263593300000730
1768:10.1007/s12052-011-0357-y
1728:10.1017/S0263593300000808
1560:10.1017/S0263593300000791
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1428:10.1017/S0263593300001449
1202:10.1002/9780470514986.ch8
1154:10.1017/S0016756807003263
1104:10.1017/S0263593300000699
1002:10.1144/0016-76492010-043
854:10.1017/S0016756819000670
307:
78:exhibiting extraordinary
2754:Paleontology in Scotland
2486:10.1073/pnas.91.25.11841
1500:Edwards, Dianne (2003).
941:10.1144/gsjgs.152.2.0229
694:section on the basis of
626:
515:
210:and 90 m down-dip.
2359:Journal of Paleontology
1610:10.1073/pnas.0501985102
1384:10.1144/0016-764900-181
710:A new genus of lichen,
250:Conditions of formation
229:Alexander Geoffrey Lyon
145: million years ago
2419:10.1098/rstb.2016.0493
1702:University of Aberdeen
1671:10.1098/rspb.2004.2686
451:
332:
321:
258:the early terrestrial
159:have been counted and
115:water-conducting cells
32:
2739:Lower Devonian Series
1831:"A new assessment of
1655:and dispersed spores"
1544:Horneophyton lignieri
445:
327:
315:
233:University of Münster
94:; a second unit, the
88:Rhynie, Aberdeenshire
22:
2714:1910 in paleontology
2630:Lower/Early Devonian
612:Rhyniella praecursor
53:57.33667°N 2.84139°W
2637:Upper/Late Devonian
2477:1994PNAS...9111841R
2471:(25): 11841–11843.
2131:2007RPaPa.146..265K
1904:10.1038/nature02291
1896:2004Natur.427..627E
1851:1981Natur.291..317W
1601:2005PNAS..102.5892T
1376:2002JGSoc.159..203R
1313:2000GSLSP.180..439P
1258:2003CaJES..40.1697T
1146:2007GeoM..144..401C
1134:Geological Magazine
1058:10.1144/sjg40010073
1050:2004ScJG...40...73T
994:2011JGSoc.168..863P
933:1995JGSoc.152..229R
889:Fossil Lagerstätten
842:Geological Magazine
811:Geology of Scotland
585:Eophalangium sheari
237:Aberdeen University
235:, and from 1987 by
214:History of research
76:sedimentary deposit
49: /
2412:(1739): 20160493.
895:on 1 December 2017
885:"The Rhynie Chert"
452:
343:still attached to
333:
322:
289:hydrothermal vents
224:William Henry Lang
105:. The bulk of the
58:57.33667; -2.84139
33:
2701:
2700:
2696:
2695:
2537:978-0-8247-8831-5
2252:978-0-226-28497-2
1952:(11): 1233–1241.
1252:(11): 1697–1712.
883:Nunn, Elizabeth.
725:Chroococcidiopsis
605:The oldest known
495:Trichopherophyton
337:permineralisation
2766:
2719:1910 in Scotland
2626:
2625:
2610:
2603:
2596:
2587:
2586:
2582:
2567:
2561:
2557:
2555:
2547:
2545:
2544:
2529:
2509:
2508:
2498:
2488:
2456:
2450:
2449:
2439:
2421:
2397:
2391:
2390:
2354:
2348:
2347:
2319:
2310:
2309:
2289:
2283:
2282:
2276:
2272:
2270:
2262:
2260:
2259:
2244:
2233:
2227:
2226:
2224:
2200:
2194:
2193:
2149:
2143:
2142:
2125:(1–4): 265–276.
2114:
2105:
2104:
2076:
2063:
2062:
2052:
2028:
2015:
2014:
2004:
1976:
1970:
1969:
1941:
1924:
1923:
1890:(6975): 627–30.
1879:
1873:
1872:
1862:
1860:10.1038/291317a0
1826:
1820:
1819:
1787:
1781:
1780:
1770:
1746:
1740:
1739:
1711:
1705:
1699:
1693:
1692:
1682:
1665:(1542): 985–92.
1646:
1633:
1632:
1622:
1612:
1578:
1572:
1571:
1539:
1533:
1532:
1506:
1497:
1480:
1474:
1446:
1440:
1439:
1411:
1402:
1401:
1399:
1398:
1361:
1352:
1343:
1342:
1324:
1322:10.1.1.1029.3013
1296:
1281:
1280:
1278:
1277:
1268:. Archived from
1237:
1224:
1223:
1189:
1166:
1165:
1131:
1122:
1116:
1115:
1087:
1070:
1069:
1033:
1014:
1013:
973:
967:
966:
960:
952:
911:
905:
904:
902:
900:
880:
874:
873:
833:
633:chytridiomycetes
146:
103:volcanic deposit
96:Windyfield chert
64:
63:
61:
60:
59:
54:
50:
47:
46:
45:
42:
2774:
2773:
2769:
2768:
2767:
2765:
2764:
2763:
2704:
2703:
2702:
2697:
2692:
2676:
2660:
2634:Middle Devonian
2619:
2617:Devonian Period
2614:
2577:
2574:
2559:
2558:
2549:
2548:
2542:
2540:
2538:
2527:
2517:
2515:Further reading
2512:
2457:
2453:
2398:
2394:
2355:
2351:
2320:
2313:
2290:
2286:
2274:
2273:
2264:
2263:
2257:
2255:
2253:
2242:
2234:
2230:
2201:
2197:
2166:10.2307/2446290
2160:(7): 992–1004.
2150:
2146:
2115:
2108:
2081:New Phytologist
2077:
2066:
2037:New Phytologist
2029:
2018:
1977:
1973:
1958:10.2307/2445050
1942:
1927:
1880:
1876:
1827:
1823:
1788:
1784:
1747:
1743:
1712:
1708:
1700:
1696:
1647:
1636:
1579:
1575:
1540:
1536:
1504:
1498:
1483:
1447:
1443:
1412:
1405:
1396:
1394:
1359:
1353:
1346:
1297:
1284:
1275:
1273:
1266:10.1139/e03-065
1238:
1227:
1212:
1190:
1169:
1129:
1123:
1119:
1088:
1073:
1034:
1017:
974:
970:
954:
953:
912:
908:
898:
896:
881:
877:
834:
823:
819:
802:
733:
708:
692:Oscillatoriales
677:
629:
570:, the possible
540:
518:
508:Another group,
310:
305:
252:
216:
192:
153:ultrastructural
141:
140:, formed about
57:
55:
51:
48:
43:
40:
38:
36:
35:
17:
12:
11:
5:
2772:
2762:
2761:
2756:
2751:
2746:
2741:
2736:
2731:
2726:
2721:
2716:
2699:
2698:
2694:
2693:
2691:
2690:
2685:
2679:
2677:
2675:
2674:
2669:
2663:
2661:
2659:
2658:
2653:
2648:
2642:
2639:
2638:
2635:
2632:
2624:
2621:
2620:
2613:
2612:
2605:
2598:
2590:
2584:
2583:
2573:
2572:External links
2570:
2569:
2568:
2560:|journal=
2536:
2516:
2513:
2511:
2510:
2451:
2392:
2371:10.1666/13-088
2365:(4): 735–746.
2349:
2330:(4): 371–389.
2311:
2284:
2275:|journal=
2251:
2243:(Google books)
2228:
2215:(4): 259–275.
2209:Insect Science
2195:
2144:
2106:
2064:
2016:
1971:
1925:
1874:
1821:
1802:(4): 341–354.
1782:
1761:(3): 489–501.
1741:
1722:(4): 445–455.
1706:
1694:
1634:
1595:(16): 5892–7.
1573:
1554:(4): 429–443.
1534:
1515:(4): 397–410.
1481:
1479:
1478:
1457:(4): 325–339.
1441:
1422:(2): 167–211.
1403:
1370:(2): 203–214.
1344:
1307:(1): 439–457.
1282:
1225:
1210:
1167:
1117:
1098:(4): 285–297.
1071:
1015:
988:(4): 863–872.
968:
927:(2): 229–250.
906:
875:
820:
818:
815:
814:
813:
808:
801:
798:
745:, provoking a
732:
729:
707:
704:
700:microbial mats
676:
673:
628:
625:
599:Palaeocharinus
594:trigonotarbids
554:euthycarcinoid
539:
536:
517:
514:
506:
505:
498:
491:
484:
477:
470:
463:
390:paleobotanists
351:is preserved.
309:
306:
304:
301:
251:
248:
220:Robert Kidston
215:
212:
191:
188:
167:—of the class
165:trigonotarbids
138:Early Devonian
125:), along with
121:, but no true
73:Lower Devonian
15:
9:
6:
4:
3:
2:
2771:
2760:
2759:Pragian Stage
2757:
2755:
2752:
2750:
2747:
2745:
2742:
2740:
2737:
2735:
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2641:
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2622:
2618:
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2606:
2604:
2599:
2597:
2592:
2591:
2588:
2580:
2576:
2575:
2565:
2553:
2539:
2533:
2530:. CRC Press.
2526:
2525:
2519:
2518:
2506:
2502:
2497:
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2487:
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2474:
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2240:
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2199:
2191:
2187:
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2179:
2175:
2171:
2167:
2163:
2159:
2155:
2148:
2140:
2136:
2132:
2128:
2124:
2120:
2113:
2111:
2102:
2098:
2094:
2090:
2086:
2082:
2075:
2073:
2071:
2069:
2060:
2056:
2051:
2046:
2043:(3): 648–57.
2042:
2038:
2034:
2027:
2025:
2023:
2021:
2012:
2008:
2003:
1998:
1994:
1990:
1987:(1): 269–85.
1986:
1982:
1975:
1967:
1963:
1959:
1955:
1951:
1947:
1940:
1938:
1936:
1934:
1932:
1930:
1921:
1917:
1913:
1909:
1905:
1901:
1897:
1893:
1889:
1885:
1878:
1870:
1866:
1861:
1856:
1852:
1848:
1845:(5813): 317.
1844:
1840:
1836:
1834:
1825:
1817:
1813:
1809:
1805:
1801:
1797:
1793:
1786:
1778:
1774:
1769:
1764:
1760:
1756:
1752:
1745:
1737:
1733:
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1710:
1703:
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1681:
1676:
1672:
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1664:
1660:
1656:
1654:
1645:
1643:
1641:
1639:
1630:
1626:
1621:
1616:
1611:
1606:
1602:
1598:
1594:
1590:
1589:
1584:
1577:
1569:
1565:
1561:
1557:
1553:
1549:
1545:
1538:
1530:
1526:
1522:
1518:
1514:
1510:
1503:
1496:
1494:
1492:
1490:
1488:
1486:
1476:
1475:
1472:
1468:
1464:
1460:
1456:
1452:
1445:
1437:
1433:
1429:
1425:
1421:
1417:
1410:
1408:
1393:
1389:
1385:
1381:
1377:
1373:
1369:
1365:
1358:
1351:
1349:
1340:
1336:
1332:
1328:
1323:
1318:
1314:
1310:
1306:
1302:
1295:
1293:
1291:
1289:
1287:
1272:on 2012-12-16
1271:
1267:
1263:
1259:
1255:
1251:
1247:
1243:
1236:
1234:
1232:
1230:
1221:
1217:
1213:
1211:9780470514986
1207:
1203:
1199:
1195:
1188:
1186:
1184:
1182:
1180:
1178:
1176:
1174:
1172:
1163:
1159:
1155:
1151:
1147:
1143:
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1135:
1128:
1121:
1113:
1109:
1105:
1101:
1097:
1093:
1086:
1084:
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1080:
1078:
1076:
1067:
1063:
1059:
1055:
1051:
1047:
1043:
1039:
1032:
1030:
1028:
1026:
1024:
1022:
1020:
1011:
1007:
1003:
999:
995:
991:
987:
983:
979:
972:
964:
958:
950:
946:
942:
938:
934:
930:
926:
922:
918:
910:
894:
890:
886:
879:
871:
867:
863:
859:
855:
851:
847:
843:
839:
832:
830:
828:
826:
821:
812:
809:
807:
804:
803:
797:
795:
791:
788:
785:
784:
778:
777:environment.
775:
774:trigonotarbid
771:
767:
765:
761:
760:
755:
752:
748:
744:
743:
742:Palaeonitella
738:
728:
726:
722:
719:
715:
714:
703:
701:
697:
693:
689:
684:
682:
681:cyanobacteria
675:Cyanobacteria
672:
670:
669:
663:
660:
656:
652:
647:
644:
642:
638:
634:
624:
622:
618:
614:
613:
608:
603:
601:
600:
596:in the genus
595:
592:(mites), and
591:
587:
586:
582:
578:
577:
573:
569:
568:
564:
560:
559:
555:
551:
550:
546:
535:
533:
532:
531:Palaeonitella
527:
523:
513:
511:
504:
503:
499:
497:
496:
492:
490:
489:
485:
483:
482:
478:
476:
475:
471:
469:
468:
464:
462:
461:
457:
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455:
449:
444:
440:
436:
434:
433:
428:
423:
421:
420:
415:
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408:
403:
401:
397:
396:
391:
387:
384:
380:
376:
371:
369:
365:
364:
359:
358:
352:
350:
346:
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338:
331:
326:
319:
314:
300:
298:
294:
290:
285:
282:
276:
274:
270:
265:
261:
257:
247:
244:
242:
238:
234:
230:
225:
221:
211:
209:
205:
201:
197:
187:
185:
182:
178:
174:
170:
166:
162:
158:
154:
150:
144:
139:
135:
130:
128:
124:
120:
116:
112:
108:
104:
99:
97:
93:
89:
85:
81:
77:
74:
70:
65:
62:
30:
26:
21:
2744:Lagerstätten
2541:. Retrieved
2523:
2468:
2464:
2454:
2409:
2405:
2395:
2362:
2358:
2352:
2327:
2323:
2297:
2293:
2287:
2256:. Retrieved
2238:
2231:
2212:
2208:
2198:
2157:
2153:
2147:
2122:
2118:
2087:(3): 475–9.
2084:
2080:
2040:
2036:
1984:
1980:
1974:
1949:
1945:
1887:
1883:
1877:
1842:
1838:
1832:
1824:
1799:
1795:
1785:
1758:
1754:
1744:
1719:
1715:
1709:
1697:
1662:
1658:
1652:
1592:
1586:
1576:
1551:
1547:
1543:
1537:
1512:
1508:
1454:
1450:
1444:
1419:
1415:
1395:. Retrieved
1367:
1363:
1304:
1300:
1274:. Retrieved
1270:the original
1249:
1245:
1193:
1137:
1133:
1120:
1095:
1091:
1044:(1): 73–81.
1041:
1037:
985:
981:
971:
957:cite journal
924:
920:
909:
897:. Retrieved
893:the original
888:
878:
848:(1): 47–64.
845:
841:
792:
789:
781:
779:
768:
763:
762:parasitised
757:
747:hypertrophic
740:
736:
734:
731:Interactions
711:
709:
687:
685:
678:
668:Prototaxites
666:
664:
648:
645:
640:
630:
610:
604:
597:
583:
574:
565:
558:Heterocrania
556:
547:
541:
529:
525:
521:
519:
510:Nematophytes
507:
500:
493:
486:
479:
474:Horneophyton
472:
465:
458:
453:
448:Horneophyton
447:
437:
430:
424:
417:
413:Horneophyton
411:
406:
404:
399:
393:
382:
372:
363:Horneophyton
361:
355:
353:
349:plant litter
334:
329:
317:
303:Preservation
286:
277:
263:
253:
245:
217:
193:
131:
100:
95:
69:Rhynie chert
68:
66:
34:
2300:(1): 9–14.
899:23 November
794:Mycorrhizae
764:Aglaophyton
713:Winfrenatia
659:Saprotrophy
637:ascomycetes
617:springtails
576:Leverhulmia
549:Lepidocaris
526:Rhynchertia
467:Asteroxylon
460:Aglaophyton
432:Aglaophyton
419:Asteroxylon
395:Aglaophyton
347:; even the
281:Yellowstone
181:mycorrhizal
171:—(known as
113:(which had
84:Lagerstätte
56: /
44:002°50′29″W
2708:Categories
2646:Lochkovian
2543:2008-05-16
2294:Nematology
2258:2008-05-16
2002:1808/16786
1397:2008-05-15
1360:(abstract)
1276:2008-05-15
1140:(2): 401.
817:References
770:Coprolites
759:Palaeonema
721:Gloeocapsa
655:holocarpic
581:harvestman
563:springtail
545:crustacean
538:Arthropods
375:parenchyma
295:texture; "
293:brecciated
196:overburden
173:book lungs
127:arthropods
41:57°20′12″N
2688:Famennian
2562:ignored (
2552:cite book
2428:0962-8436
2387:131202472
2379:0022-3360
2344:129545961
2277:ignored (
2267:cite book
1981:Mycologia
1833:Rhyniella
1816:128563568
1777:1936-6426
1736:128869547
1568:128501945
1471:129845220
1436:128754463
1339:129847888
1317:CiteSeerX
1112:128424299
1066:128937466
1010:128679831
949:128977213
870:182210855
862:0016-7568
696:biomarker
688:bona fide
686:However,
567:Rhyniella
522:Mackiella
502:Ventarura
427:sporangia
297:geyserite
260:ecosystem
256:petrified
184:symbionts
169:Arachnida
119:sporangia
2683:Frasnian
2672:Givetian
2667:Eifelian
2505:11607500
2446:29254958
2190:25684294
2182:21708654
2154:Am J Bot
2101:17447903
2059:17447919
2011:16389979
1912:14961119
1689:15255055
1629:15809414
1529:52103830
1392:55042118
1162:53975045
800:See also
754:nematode
651:Eucarpic
379:Triassic
366:grew on
345:rhizomes
341:rhizoids
190:Location
107:Devonian
92:Scotland
29:Scotland
2651:Pragian
2473:Bibcode
2437:5745329
2174:2446290
2127:Bibcode
1966:2445050
1920:4431205
1892:Bibcode
1869:4339420
1847:Bibcode
1680:1691674
1653:in situ
1597:Bibcode
1372:Bibcode
1309:Bibcode
1254:Bibcode
1220:9243014
1142:Bibcode
1046:Bibcode
990:Bibcode
929:Bibcode
751:enoplid
737:in situ
718:coccoid
706:Lichens
621:insects
607:hexapod
386:stomata
264:in situ
157:Stomata
134:Pragian
2656:Emsian
2534:
2503:
2493:
2444:
2434:
2426:
2385:
2377:
2342:
2249:
2188:
2180:
2172:
2099:
2057:
2009:
1964:
1918:
1910:
1884:Nature
1867:
1839:Nature
1814:
1775:
1734:
1687:
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1627:
1620:556298
1617:
1566:
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1469:
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1390:
1337:
1319:
1218:
1208:
1160:
1110:
1064:
1008:
947:
868:
860:
783:Nothia
756:named
579:, the
572:insect
561:, the
552:, the
488:Rhynia
481:Nothia
383:behind
368:sinter
357:Rhynia
330:Rhynia
318:Rhynia
308:Plants
273:graben
208:strike
200:Rhynie
177:hyphae
161:lignin
149:cherts
123:leaves
111:plants
80:fossil
25:Rhynie
2734:Chert
2496:45331
2383:S2CID
2340:S2CID
2186:S2CID
2170:JSTOR
1962:JSTOR
1916:S2CID
1865:S2CID
1812:S2CID
1732:S2CID
1564:S2CID
1525:S2CID
1505:(PDF)
1467:S2CID
1432:S2CID
1388:S2CID
1335:S2CID
1158:S2CID
1130:(PDF)
1108:S2CID
1062:S2CID
1006:S2CID
945:S2CID
866:S2CID
627:Fungi
590:Acari
516:Algae
269:tuffs
241:Cores
71:is a
2564:help
2532:ISBN
2501:PMID
2442:PMID
2424:ISSN
2375:ISSN
2279:help
2247:ISBN
2178:PMID
2097:PMID
2055:PMID
2007:PMID
1908:PMID
1773:ISSN
1685:PMID
1625:PMID
1216:PMID
1206:ISBN
963:link
901:2017
858:ISSN
723:and
653:and
524:and
398:were
222:and
117:and
67:The
2491:PMC
2481:doi
2432:PMC
2414:doi
2410:373
2367:doi
2332:doi
2302:doi
2217:doi
2162:doi
2135:doi
2123:146
2089:doi
2085:174
2045:doi
2041:174
1997:hdl
1989:doi
1954:doi
1900:doi
1888:427
1855:doi
1843:291
1804:doi
1763:doi
1724:doi
1675:PMC
1667:doi
1663:271
1615:PMC
1605:doi
1593:102
1556:doi
1517:doi
1459:doi
1424:doi
1380:doi
1368:159
1327:doi
1305:180
1262:doi
1198:doi
1150:doi
1138:144
1100:doi
1054:doi
998:doi
986:168
937:doi
925:152
850:doi
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