783:- 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.
31:
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727:, 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
381:, 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.
310:", 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.
925:
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.
282:- 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.
2303:
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".
2128:
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
682:, 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.
254:, 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
1990:
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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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
801:
Fungal interactions are known to promote speciation in modern plants, and presumably also affected
Devonian diversity by providing a selection pressure.
433:). 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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1425:
Wellman, Charles H. (2006). "Spore assemblages from the Lower
Devonian 'Lower Old Red Sandstone' deposits of the Rhynie outlier, Scotland".
895:
2368:
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".
1047:
Trewin, N.H.; Wilson, E. (2004). "Correlation of the Early
Devonian Rhynie chert beds between three boreholes at Rhynie, Aberdeenshire".
242:
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
217:. 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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1368:"Geological setting of the Early Devonian Rhynie cherts, Aberdeenshire, Scotland: an early terrestrial hot spring system"
1138:"A new Middle – Late Jurassic flora and hot spring chert deposit from the Deseado Massif, Santa Cruz province, Argentina"
48:
2090:
Berbee, Mary L.; Taylor, JW (2007). "Rhynie chert: a window into a lost world of complex plant?fungus interactions".
2044:"Fungal endophytes in a 400-million-yr-old land plant: infection pathways, spatial distribution, and host responses"
1253:"Subaqueous silicification of the contents of small ponds in an Early Devonian hot-spring complex, Rhynie, Scotland"
2611:
1101:
Trewin (2003). "History of research on the geology and palaeontology of the Rhynie area, Aberdeenshire, Scotland".
989:"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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399:, 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".
243:
545:, which inhabited the alkaline freshwater pools towards the end of the sinter apron, has been characterized.
239:
461:. 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
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1327:
630:, was found in the Rhynie chert, pushing dates for the origination of hexapods (a group that includes the
1955:
Taylor, T.N.; Remy, W.; Hass, H. (1992). "Fungi from the Lower Devonian Rhynie chert: Chytridiomycetes".
1662:"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".
595:
1203:
Trewin, NH (1996). "The Rhynie cherts: an early Devonian ecosystem preserved by hydrothermal activity".
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1803:"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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2216:"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.
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250:, 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.
27:
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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650:, oomycota (Peronosporomycetes) and glomeromycetes; indeed the only fungal groups
388:, is not observed elsewhere in the fossil record until the advent of amber in the
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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).
2003:
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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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2335:
Earth and Environmental Science Transactions of the Royal Society of Edinburgh
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1807:
Earth and Environmental Science Transactions of the Royal Society of Edinburgh
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1594:"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 (
213:, 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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158:) 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
2012:
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902:. Department of Earth Sciences, University of Bristol. Archived from
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1846:, the earliest known insect, from the Devonian of Rhynie, Scotland"
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427:), and which were likely to have colonised waterlogged surfaces (
1207:. Novartis Foundation Symposia. Vol. 202. pp. 131–45.
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133:
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Garwood, Russell J; Oliver, Heather; Spencer, Alan R T (2019).
660:
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
1727:
Transactions of the Royal Society of Edinburgh: Earth Sciences
1715:, The Biota of Early Terrestrial Ecosystems: The Rhynie Chert.
1559:
Transactions of the Royal Society of Edinburgh: Earth Sciences
1520:
Transactions of the Royal Society of Edinburgh: Earth Sciences
1513:"Embryophytic sporophytes in the Rhynie and Windyfield cherts"
1462:
Transactions of the Royal Society of Edinburgh: Earth Sciences
1427:
Transactions of the Royal Society of Edinburgh: Earth Sciences
1103:
Transactions of the Royal Society of Edinburgh: Earth Sciences
2472:"4 hundred million year old vesicular-arbuscular mycorrhizae"
392:. This allows the study of structures such as the air spaces
987:
Parry, S.F.; Noble S.R.; Crowley Q.G.; Wellman C.H. (2011).
523:, remains enigmatic, but may represent aquatic land plants.
928:"A Devonian auriferous hot spring system, Rhynie, Scotland"
746:
The Rhynie chert, by preserving a snapshot of an ecosystem
2413:"Terrestrial invertebrates in the Rhynie chert ecosystem"
1801:
Dunlop, J.A.; Anderson, L.I.; Kerp, H.; Hass, H. (2007).
298:
Sedimentary textures which appear to have formed in the
1762:"Early Terrestrial Animals, Evolution, and Uncertainty"
1760:Garwood, Russell J.; Edgecombe, Gregory D. (2011).
1136:Channing; Zamuner, Alba B.; ZÚÑiga, Adolfo (2007).
842:
840:
838:
836:
1312:Geological Society of London, Special Publications
1366:Rice, C.M.; Trewin, N.H.; Anderson, L.I. (2002).
2250:The Origin And Early Evolution Of Plants On Land
1557:(Kidston and Lang) Barghoorn and Darrah, 1938".
1599:Proceedings of the National Academy of Sciences
1251:Trewin, N.H.; Fayers, S.R.; Kelman, R. (2003).
2411:Dunlop, Jason A.; Garwood, Russell J. (2017).
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642:Fungi known from the Rhynie chert include the
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972:: CS1 maint: multiple names: authors list (
1840:Whalley, Paul; Jarzembowski, E. A. (1981).
1361:
1359:
2636:
2619:
2605:
2597:
2470:Remy W, Taylor TN, Hass H, Kerp H (1994).
1592:Taylor, T. N.; Kerp, H.; Hass, H. (2005).
162:are famous for their exceptional state of
2505:
2495:
2446:
2428:
2231:
2059:
2011:
1869:
1777:
1689:
1629:
1619:
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932:Journal of the Geological Society, London
186:) can be seen in cross-sections. Fungal
29:
832:
436:The preservation of spores attached to
209:, in a small field near the village of
2570:
2560:
2285:
2275:
965:
2130:Review of Palaeobotany and Palynology
849:"An introduction to the Rhynie chert"
403:to firmly deduce that plants such as
205:The bed is under at least 1 metre of
97:). It is exposed near the village of
34:Hand sample of the Rhynie chert from
7:
807:are also found in the Rhynie chert.
109:, is located some 700 m away.
2532:Taylor, T.N.; Taylor, E.L. (2000).
215:Site of Special Scientific Interest
1670:Proceedings of the Royal Society B
1257:Canadian Journal of Earth Sciences
25:
2247:Kenrick, P.; Crane, P.R. (2000).
1766:Evolution: Education and Outreach
1375:Journal of the Geological Society
993:Journal of the Geological Society
120:fossil bed consists of primitive
2735:Protected areas of Aberdeenshire
2590:"Rhynie Chert Learning Resource"
2233:10.1111/j.1744-7917.2007.00152.x
2104:10.1111/j.1469-8137.2007.02080.x
2061:10.1111/j.1469-8137.2007.02008.x
539:). A well-preserved charophyte,
331:stems (axes). Scale bar is 1 cm.
302:themselves are preserved with a
2256:. University of Chicago Press.
634:) back to the Silurian period.
2150:10.1016/j.revpalbo.2007.05.002
817:Evolutionary history of plants
626:), which resembles the modern
1:
1342:10.1144/GSL.SP.2000.180.01.23
2760:Environment of Aberdeenshire
2539:(Free access @ Google books)
2476:Proc. Natl. Acad. Sci. U.S.A
1660:Wellman, Charles H. (2004).
413:aquatic, as once believed.
140:, lichens, algae and fungi.
2214:Labandeira, CONRAD (2007).
1049:Scottish Journal of Geology
690:In the rare instances that
2786:
2317:10.1163/156854108783360159
2004:10.3852/mycologia.97.1.269
1957:American Journal of Botany
93:detail or completeness (a
2740:Devonian System of Europe
2634:
2592:. University of Aberdeen.
2347:10.1017/S0263593300000754
1819:10.1017/S0263593300000730
1779:10.1007/s12052-011-0357-y
1739:10.1017/S0263593300000808
1571:10.1017/S0263593300000791
1532:10.1017/S0263593300000778
1474:10.1017/S0263593300000729
1439:10.1017/S0263593300001449
1213:10.1002/9780470514986.ch8
1165:10.1017/S0016756807003263
1115:10.1017/S0263593300000699
1013:10.1144/0016-76492010-043
865:10.1017/S0016756819000670
89:exhibiting extraordinary
2765:Paleontology in Scotland
2497:10.1073/pnas.91.25.11841
1511:Edwards, Dianne (2003).
952:10.1144/gsjgs.152.2.0229
705:section on the basis of
221:and 90 m down-dip.
2370:Journal of Paleontology
1621:10.1073/pnas.0501985102
1395:10.1144/0016-764900-181
721:A new genus of lichen,
261:Conditions of formation
240:Alexander Geoffrey Lyon
156: million years ago
2430:10.1098/rstb.2016.0493
1713:University of Aberdeen
1682:10.1098/rspb.2004.2686
462:
343:
332:
269:the early terrestrial
170:have been counted and
126:water-conducting cells
43:
2750:Lower Devonian Series
1842:"A new assessment of
1666:and dispersed spores"
1555:Horneophyton lignieri
456:
338:
326:
244:University of Münster
105:; a second unit, the
99:Rhynie, Aberdeenshire
33:
2725:1910 in paleontology
2641:Lower/Early Devonian
623:Rhyniella praecursor
64:57.33667°N 2.84139°W
2648:Upper/Late Devonian
2488:1994PNAS...9111841R
2482:(25): 11841–11843.
2142:2007RPaPa.146..265K
1915:10.1038/nature02291
1907:2004Natur.427..627E
1862:1981Natur.291..317W
1612:2005PNAS..102.5892T
1387:2002JGSoc.159..203R
1324:2000GSLSP.180..439P
1269:2003CaJES..40.1697T
1157:2007GeoM..144..401C
1145:Geological Magazine
1069:10.1144/sjg40010073
1061:2004ScJG...40...73T
1005:2011JGSoc.168..863P
944:1995JGSoc.152..229R
900:Fossil Lagerstätten
853:Geological Magazine
822:Geology of Scotland
596:Eophalangium sheari
248:Aberdeen University
246:, and from 1987 by
225:History of research
87:sedimentary deposit
60: /
2423:(1739): 20160493.
906:on 1 December 2017
896:"The Rhynie Chert"
463:
354:still attached to
344:
333:
300:hydrothermal vents
235:William Henry Lang
116:. The bulk of the
69:57.33667; -2.84139
44:
2712:
2711:
2707:
2706:
2548:978-0-8247-8831-5
2263:978-0-226-28497-2
1963:(11): 1233–1241.
1263:(11): 1697–1712.
894:Nunn, Elizabeth.
736:Chroococcidiopsis
616:The oldest known
506:Trichopherophyton
348:permineralisation
16:(Redirected from
2777:
2730:1910 in Scotland
2637:
2621:
2614:
2607:
2598:
2593:
2578:
2572:
2568:
2566:
2558:
2556:
2555:
2540:
2520:
2519:
2509:
2499:
2467:
2461:
2460:
2450:
2432:
2408:
2402:
2401:
2365:
2359:
2358:
2330:
2321:
2320:
2300:
2294:
2293:
2287:
2283:
2281:
2273:
2271:
2270:
2255:
2244:
2238:
2237:
2235:
2211:
2205:
2204:
2160:
2154:
2153:
2136:(1–4): 265–276.
2125:
2116:
2115:
2087:
2074:
2073:
2063:
2039:
2026:
2025:
2015:
1987:
1981:
1980:
1952:
1935:
1934:
1901:(6975): 627–30.
1890:
1884:
1883:
1873:
1871:10.1038/291317a0
1837:
1831:
1830:
1798:
1792:
1791:
1781:
1757:
1751:
1750:
1722:
1716:
1710:
1704:
1703:
1693:
1676:(1542): 985–92.
1657:
1644:
1643:
1633:
1623:
1589:
1583:
1582:
1550:
1544:
1543:
1517:
1508:
1491:
1485:
1457:
1451:
1450:
1422:
1413:
1412:
1410:
1409:
1372:
1363:
1354:
1353:
1335:
1333:10.1.1.1029.3013
1307:
1292:
1291:
1289:
1288:
1279:. Archived from
1248:
1235:
1234:
1200:
1177:
1176:
1142:
1133:
1127:
1126:
1098:
1081:
1080:
1044:
1025:
1024:
984:
978:
977:
971:
963:
922:
916:
915:
913:
911:
891:
885:
884:
844:
644:chytridiomycetes
157:
114:volcanic deposit
107:Windyfield chert
75:
74:
72:
71:
70:
65:
61:
58:
57:
56:
53:
21:
2785:
2784:
2780:
2779:
2778:
2776:
2775:
2774:
2715:
2714:
2713:
2708:
2703:
2687:
2671:
2645:Middle Devonian
2630:
2628:Devonian Period
2625:
2588:
2585:
2569:
2559:
2553:
2551:
2549:
2538:
2531:
2528:
2526:Further reading
2523:
2469:
2468:
2464:
2410:
2409:
2405:
2367:
2366:
2362:
2332:
2331:
2324:
2302:
2301:
2297:
2284:
2274:
2268:
2266:
2264:
2253:
2246:
2245:
2241:
2213:
2212:
2208:
2177:10.2307/2446290
2171:(7): 992–1004.
2162:
2161:
2157:
2127:
2126:
2119:
2092:New Phytologist
2089:
2088:
2077:
2048:New Phytologist
2041:
2040:
2029:
1989:
1988:
1984:
1969:10.2307/2445050
1954:
1953:
1938:
1892:
1891:
1887:
1839:
1838:
1834:
1800:
1799:
1795:
1759:
1758:
1754:
1724:
1723:
1719:
1711:
1707:
1659:
1658:
1647:
1591:
1590:
1586:
1552:
1551:
1547:
1515:
1510:
1509:
1494:
1459:
1458:
1454:
1424:
1423:
1416:
1407:
1405:
1370:
1365:
1364:
1357:
1309:
1308:
1295:
1286:
1284:
1277:10.1139/e03-065
1250:
1249:
1238:
1223:
1202:
1201:
1180:
1140:
1135:
1134:
1130:
1100:
1099:
1084:
1046:
1045:
1028:
986:
985:
981:
964:
924:
923:
919:
909:
907:
893:
892:
888:
846:
845:
834:
830:
813:
744:
719:
703:Oscillatoriales
688:
640:
581:, the possible
551:
529:
519:Another group,
321:
316:
263:
227:
203:
164:ultrastructural
152:
151:, formed about
68:
66:
62:
59:
54:
51:
49:
47:
46:
28:
23:
22:
15:
12:
11:
5:
2783:
2781:
2773:
2772:
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2762:
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2732:
2727:
2717:
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2710:
2709:
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2702:
2701:
2696:
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2659:
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2649:
2646:
2643:
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2609:
2601:
2595:
2594:
2584:
2583:External links
2581:
2580:
2579:
2571:|journal=
2547:
2527:
2524:
2522:
2521:
2462:
2403:
2382:10.1666/13-088
2376:(4): 735–746.
2360:
2341:(4): 371–389.
2322:
2295:
2286:|journal=
2262:
2254:(Google books)
2239:
2226:(4): 259–275.
2220:Insect Science
2206:
2155:
2117:
2075:
2027:
1982:
1936:
1885:
1832:
1813:(4): 341–354.
1793:
1772:(3): 489–501.
1752:
1733:(4): 445–455.
1717:
1705:
1645:
1606:(16): 5892–7.
1584:
1565:(4): 429–443.
1545:
1526:(4): 397–410.
1492:
1490:
1489:
1468:(4): 325–339.
1452:
1433:(2): 167–211.
1414:
1381:(2): 203–214.
1355:
1318:(1): 439–457.
1293:
1236:
1221:
1178:
1128:
1109:(4): 285–297.
1082:
1026:
999:(4): 863–872.
979:
938:(2): 229–250.
917:
886:
831:
829:
826:
825:
824:
819:
812:
809:
756:, provoking a
743:
740:
718:
715:
711:microbial mats
687:
684:
639:
636:
610:Palaeocharinus
605:trigonotarbids
565:euthycarcinoid
550:
547:
528:
525:
517:
516:
509:
502:
495:
488:
481:
474:
401:paleobotanists
362:is preserved.
320:
317:
315:
312:
262:
259:
231:Robert Kidston
226:
223:
202:
199:
178:—of the class
176:trigonotarbids
149:Early Devonian
136:), along with
132:, but no true
84:Lower Devonian
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
2782:
2771:
2770:Pragian Stage
2768:
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2541:. CRC Press.
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2113:
2109:
2105:
2101:
2097:
2093:
2086:
2084:
2082:
2080:
2076:
2071:
2067:
2062:
2057:
2054:(3): 648–57.
2053:
2049:
2045:
2038:
2036:
2034:
2032:
2028:
2023:
2019:
2014:
2009:
2005:
2001:
1998:(1): 269–85.
1997:
1993:
1986:
1983:
1978:
1974:
1970:
1966:
1962:
1958:
1951:
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1856:(5813): 317.
1855:
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1564:
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1321:
1317:
1313:
1306:
1304:
1302:
1300:
1298:
1294:
1283:on 2012-12-16
1282:
1278:
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1262:
1258:
1254:
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1241:
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1222:9780470514986
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841:
839:
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833:
827:
823:
820:
818:
815:
814:
810:
808:
806:
802:
799:
796:
795:
789:
788:environment.
786:
785:trigonotarbid
782:
778:
776:
772:
771:
766:
763:
759:
755:
754:
753:Palaeonitella
749:
741:
739:
737:
733:
730:
726:
725:
716:
714:
712:
708:
704:
700:
695:
693:
692:cyanobacteria
686:Cyanobacteria
685:
683:
681:
680:
674:
671:
667:
663:
658:
655:
653:
649:
645:
637:
635:
633:
629:
625:
624:
619:
614:
612:
611:
607:in the genus
606:
603:(mites), and
602:
598:
597:
593:
589:
588:
584:
580:
579:
575:
571:
570:
566:
562:
561:
557:
548:
546:
544:
543:
542:Palaeonitella
538:
534:
526:
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127:
123:
119:
115:
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108:
104:
100:
96:
92:
88:
85:
81:
76:
73:
41:
37:
32:
19:
2755:Lagerstätten
2552:. Retrieved
2534:
2479:
2475:
2465:
2420:
2416:
2406:
2373:
2369:
2363:
2338:
2334:
2308:
2304:
2298:
2267:. Retrieved
2249:
2242:
2223:
2219:
2209:
2168:
2164:
2158:
2133:
2129:
2098:(3): 475–9.
2095:
2091:
2051:
2047:
1995:
1991:
1985:
1960:
1956:
1898:
1894:
1888:
1853:
1849:
1843:
1835:
1810:
1806:
1796:
1769:
1765:
1755:
1730:
1726:
1720:
1708:
1673:
1669:
1663:
1603:
1597:
1587:
1562:
1558:
1554:
1548:
1523:
1519:
1465:
1461:
1455:
1430:
1426:
1406:. Retrieved
1378:
1374:
1315:
1311:
1285:. Retrieved
1281:the original
1260:
1256:
1204:
1148:
1144:
1131:
1106:
1102:
1055:(1): 73–81.
1052:
1048:
996:
992:
982:
968:cite journal
935:
931:
920:
908:. Retrieved
904:the original
899:
889:
859:(1): 47–64.
856:
852:
803:
800:
792:
790:
779:
774:
773:parasitised
768:
758:hypertrophic
751:
747:
745:
742:Interactions
722:
720:
698:
696:
689:
679:Prototaxites
677:
675:
659:
656:
651:
641:
621:
615:
608:
594:
585:
576:
569:Heterocrania
567:
558:
552:
540:
536:
532:
530:
521:Nematophytes
518:
511:
504:
497:
490:
485:Horneophyton
483:
476:
469:
464:
459:Horneophyton
458:
448:
441:
435:
428:
424:Horneophyton
422:
417:
415:
410:
404:
393:
383:
374:Horneophyton
372:
366:
364:
360:plant litter
345:
340:
328:
314:Preservation
297:
288:
274:
264:
256:
228:
204:
142:
111:
106:
80:Rhynie chert
79:
77:
45:
18:Rhynie Chert
2311:(1): 9–14.
910:23 November
805:Mycorrhizae
775:Aglaophyton
724:Winfrenatia
670:Saprotrophy
648:ascomycetes
628:springtails
587:Leverhulmia
560:Lepidocaris
537:Rhynchertia
478:Asteroxylon
471:Aglaophyton
443:Aglaophyton
430:Asteroxylon
406:Aglaophyton
358:; even the
292:Yellowstone
192:mycorrhizal
182:—(known as
124:(which had
95:Lagerstätte
67: /
55:002°50′29″W
2719:Categories
2657:Lochkovian
2554:2008-05-16
2305:Nematology
2269:2008-05-16
2013:1808/16786
1408:2008-05-15
1371:(abstract)
1287:2008-05-15
1151:(2): 401.
828:References
781:Coprolites
770:Palaeonema
732:Gloeocapsa
666:holocarpic
592:harvestman
574:springtail
556:crustacean
549:Arthropods
386:parenchyma
306:texture; "
304:brecciated
207:overburden
184:book lungs
138:arthropods
52:57°20′12″N
2699:Famennian
2573:ignored (
2563:cite book
2439:0962-8436
2398:131202472
2390:0022-3360
2355:129545961
2288:ignored (
2278:cite book
1992:Mycologia
1844:Rhyniella
1827:128563568
1788:1936-6426
1747:128869547
1579:128501945
1482:129845220
1447:128754463
1350:129847888
1328:CiteSeerX
1123:128424299
1077:128937466
1021:128679831
960:128977213
881:182210855
873:0016-7568
707:biomarker
699:bona fide
697:However,
578:Rhyniella
533:Mackiella
513:Ventarura
438:sporangia
308:geyserite
271:ecosystem
267:petrified
195:symbionts
180:Arachnida
130:sporangia
2694:Frasnian
2683:Givetian
2678:Eifelian
2516:11607500
2457:29254958
2201:25684294
2193:21708654
2165:Am J Bot
2112:17447903
2070:17447919
2022:16389979
1923:14961119
1700:15255055
1640:15809414
1540:52103830
1403:55042118
1173:53975045
811:See also
765:nematode
662:Eucarpic
390:Triassic
377:grew on
356:rhizomes
352:rhizoids
201:Location
118:Devonian
103:Scotland
40:Scotland
2662:Pragian
2484:Bibcode
2448:5745329
2185:2446290
2138:Bibcode
1977:2445050
1931:4431205
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1858:Bibcode
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1265:Bibcode
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1153:Bibcode
1057:Bibcode
1001:Bibcode
940:Bibcode
762:enoplid
748:in situ
729:coccoid
717:Lichens
632:insects
618:hexapod
397:stomata
275:in situ
168:Stomata
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2667:Emsian
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590:, the
583:insect
572:, the
563:, the
499:Rhynia
492:Nothia
394:behind
379:sinter
368:Rhynia
341:Rhynia
329:Rhynia
319:Plants
284:graben
219:strike
211:Rhynie
188:hyphae
172:lignin
160:cherts
134:leaves
122:plants
91:fossil
36:Rhynie
2745:Chert
2507:45331
2394:S2CID
2351:S2CID
2197:S2CID
2181:JSTOR
1973:JSTOR
1927:S2CID
1876:S2CID
1823:S2CID
1743:S2CID
1575:S2CID
1536:S2CID
1516:(PDF)
1478:S2CID
1443:S2CID
1399:S2CID
1346:S2CID
1169:S2CID
1141:(PDF)
1119:S2CID
1073:S2CID
1017:S2CID
956:S2CID
877:S2CID
638:Fungi
601:Acari
527:Algae
280:tuffs
252:Cores
82:is a
2575:help
2543:ISBN
2512:PMID
2453:PMID
2435:ISSN
2386:ISSN
2290:help
2258:ISBN
2189:PMID
2108:PMID
2066:PMID
2018:PMID
1919:PMID
1784:ISSN
1696:PMID
1636:PMID
1227:PMID
1217:ISBN
974:link
912:2017
869:ISSN
734:and
664:and
535:and
409:were
233:and
128:and
78:The
2502:PMC
2492:doi
2443:PMC
2425:doi
2421:373
2378:doi
2343:doi
2313:doi
2228:doi
2173:doi
2146:doi
2134:146
2100:doi
2096:174
2056:doi
2052:174
2008:hdl
2000:doi
1965:doi
1911:doi
1899:427
1866:doi
1854:291
1815:doi
1774:doi
1735:doi
1686:PMC
1678:doi
1674:271
1626:PMC
1616:doi
1604:102
1567:doi
1528:doi
1470:doi
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1391:doi
1379:159
1338:doi
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1273:doi
1209:doi
1161:doi
1149:144
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