Hostname: page-component-cd9895bd7-q99xh Total loading time: 0 Render date: 2024-12-24T16:01:26.618Z Has data issue: false hasContentIssue false
  • English
  • Français

An introduction to the Rhynie chert

Published online by Cambridge University Press:  18 July 2019

Russell J Garwood Open the ORCID record for Russell J Garwood [Opens in a new window] ,
Heather Oliver  and
Alan R T Spencer Open the ORCID record for Alan R T Spencer [Opens in a new window]
Russell J Garwood*
Affiliation:
School of Earth and Environmental Science, University of Manchester, Manchester, UK Earth Sciences Department, Natural History Museum, London, UK
Heather Oliver
Affiliation:
Sharples School, Hill Cot Road, Astley Bridge, Bolton, UK
Alan R T Spencer
Affiliation:
Earth Sciences Department, Natural History Museum, London, UK Department of Earth Sciences and Engineering, Imperial College London, London, UK
*
Author for correspondence: Russell J Garwood, Email: [email protected]
Get access Rights & Permissions [Opens in a new window]

Abstract

The terrestrialization of life has profoundly affected the biosphere, geosphere and atmosphere, and the Geological Magazine has published key works charting the development of our understanding of this process. Integral to this understanding – and featuring in one of the Geological Magazine publications – is the Devonian Rhynie chert Konservat-Lagerstätte located in Aberdeenshire, Scotland. Here we provide a review of the work on this important early terrestrial deposit to date. We begin by highlighting contributions of note in the Geological Magazine improving understanding of terrestrialization and Palaeozoic terrestrial ecosystems. We then introduce the Rhynie chert. The review highlights its geological setting: the Caledonian context of the Rhynie Basin and its nature at the time of deposition of the cherts which host its famous fossils. There follows an introduction to the development of the half-graben in which the cherts and host sediments were deposited, the palaeoenvironment this represented and the taphonomy of the fossils themselves. We subsequently provide an overview of the mineralization and geochemistry of the deposit, and then the fossils found within the Rhynie chert. These include: six plant genera, which continue to provide significant insights into the evolution of life on land; a range of different fungi, with recent work starting to probe plant–fungus interactions; lichens, amoebae and a range of unicellular eukaryotes and prokaryotes (algae and cyanobacteria); and finally a range of both aquatic and terrestrial arthropods. Through continued study coupled with methodological advances, Rhynie fossils will continue to provide unique insights into early life on land.

Keywords

chertterrestrializationDevonianLagerstätte
Type
Original Article
Information
Geological Magazine , Volume 157 , Special Issue 1: SPECIAL ISSUE: Commemorating the 155th anniversary of Geological Magazine , January 2020 , pp. 47 - 64
Copyright
© Cambridge University Press 2019 

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Abbott, GD, Fletcher, IW, Tardio, S and Hack, E (2018) Exploring the geochemical distribution of organic carbon in early land plants: a novel approach. Philosophical Transactions of the Royal Society B: Biological Sciences 373, 20160499. doi: 10.1098/rstb.2016.0499.CrossRefGoogle ScholarPubMed
Anderson, L, Crighton, B and Hass, H (2003) A new univalve crustacean from the Early Devonian Rhynie chert hot-spring complex. Transactions of the Royal Society of Edinburgh: Earth Sciences 94, 355–69.CrossRefGoogle Scholar
Anderson, LI, Dunlop, JA, Eagar, RMC, Horrocks, CA and Wilson, HM (1999) Soft-bodied fossils from the roof shales of the Wigan Four Foot coal seam, Westhoughton, Lancashire, UK. Geological Magazine 135, 321–9.10.1017/S0016756899002575CrossRefGoogle Scholar
Anderson, LI and Trewin, NH (2003) An Early Devonian arthropod fauna from the Windyfield Cherts, Aberdeenshire, Scotland. Palaeontology 46, 467–509.CrossRefGoogle Scholar
Arber, EAN (1907) II. – A note on fossil plants from the Carboniferous limestone of Chepstow. Geological Magazine 4, 4–5.CrossRefGoogle Scholar
Arber, EAN (1912) I. – A note on some fossil plants from the Kent Coal-field. Geological Magazine 9, 97–9.CrossRefGoogle Scholar
Arber, EAN (1913) V. – On the discovery of fossil plants in the Old Hill Marls of the South Staffordshire Coal-field. Geological Magazine 10, 215–16.CrossRefGoogle Scholar
Baldwin, W (1911) VI.–Fossil myriopods from the Middle Coal-Measures of Sparth Bottoms, Rochdale, Lancashire. Geological Magazine 8, 74–80.CrossRefGoogle Scholar
Barghoorn, ES and Darrah, WC (1938) Horneophyton, a necessary change of name for Horneabrade. Botanical Museum Leaflets, Harvard University 6, 142–4.Google Scholar
Baron, M, Hillier, S, Rice, CM, Czapnik, K and Parnell, J (2003) Fluids and hydrothermal alteration assemblages in a Devonian gold-bearing hot-spring system, Rhynie, Scotland. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 94, 309–24.CrossRefGoogle Scholar
Berry, CM, Morel, E, Mojica, J and Villarroel, C (2000) Devonian plants from Colombia, with discussion of their geological and palaeogeographical context. Geological Magazine 137, 257–68.CrossRefGoogle Scholar
Bolton, H (1905) I – Notes on the geological horizon and palæontology of the ‘Soapstone Bed’, in the Lower Coal-Measures, near Colne, Lancashire. Geological Magazine 2, 433–7.CrossRefGoogle Scholar
Boyce, CK, Cody, GD, Feser, M, Jacobsen, C, Knoll, AH and Wirick, S (2002) Organic chemical differentiation within fossil plant cell walls detected with X-ray spectromicroscopy. Geology 30, 1039–42.2.0.CO;2>CrossRefGoogle Scholar
Boyce, CK, Hazen, RM and Knoll, AH (2001) Nondestructive, in situ, cellular-scale mapping of elemental abundances including organic carbon in permineralized fossils. Proceedings of the National Academy of Sciences 98, 5970–4.CrossRefGoogle ScholarPubMed
Brade, SB-G (1928) An important specimen of Euphoberia ferox from the Middle Coal Measures of Crawcrook. Geological Magazine 65, 400–6.CrossRefGoogle Scholar
Brongniart, C (1885) I. – The fossil insects of the primary rocks: a rapid survey of the entomological fauna of the Palæozoic Period 1. Geological Magazine 2, 481–91.CrossRefGoogle Scholar
Busrewil, MT, Pankhurst, RJ and Wadsworth, WJ (1973) The igneous rocks of the Boganclogh area, N.E. Scotland. Scottish Journal of Geology 9, 165–76.CrossRefGoogle Scholar
Channing, A (2018) A review of active hot-spring analogues of Rhynie: environments, habitats and ecosystems. Philosophical Transactions of the Royal Society B: Biological Sciences 373, 20160490. doi: 10.1098/rstb.2016.0490.CrossRefGoogle ScholarPubMed
Channing, A and Edwards, D (2009a) Silicification of higher plants in geothermally influenced wetlands: Yellowstone as a Lower Devonian Rhynie analog. PALAIOS 24, 505–21.CrossRefGoogle Scholar
Channing, A and Edwards, D (2009b) Yellowstone hot spring environments and the palaeo-ecophysiology of Rhynie chert plants: towards a synthesis. Plant Ecology & Diversity 2, 111–43.CrossRefGoogle Scholar
Cingolani, CA, Berry, CM, Morel, E and Tomezzoli, R (2002) Middle Devonian lycopsids from high southern palaeolatitudes of Gondwana (Argentina). Geological Magazine 139, 641–9.CrossRefGoogle Scholar
Claridge, MF and Lyon, AG (1961) Lung-books in the Devonian Palaeocharinidae (Arachnida). Nature 191, 1190–1.CrossRefGoogle Scholar
Clarke, PD and Wadsworth, WJ (1970) The Insch layered intrusion. Scottish Journal of Geology 6, 7–25.CrossRefGoogle Scholar
Cleal, CJ (1986) Fossil plants of the Severn Coalfield and their biostratigraphical significance. Geological Magazine 123, 553–68.CrossRefGoogle Scholar
Cocks, LRM, Fortey, RA and Rushton, AWA (2010) Correlation for the Lower Palaeozoic. Geological Magazine 147, 171–80.CrossRefGoogle Scholar
Croft, WN and George, EA (1959) Blue-green algae from the Middle Devonian of Rhynie, Aberdeenshire. Bulletin of the British Museum (Natural History) Geology 3, 341–53.Google Scholar
Davies, NS and Gibling, MR (2010) Cambrian to Devonian evolution of alluvial systems: the sedimentological impact of the earliest land plants. Earth-Science Reviews 98, 171–200.10.1016/j.earscirev.2009.11.002CrossRefGoogle Scholar
Deline, B, Greenwood, JM, Clark, JW, Puttick, MN, Peterson, KJ and Donoghue, PCJ (2018) Evolution of metazoan morphological disparity. Proceedings of the National Academy of Sciences 115, E8909–18.CrossRefGoogle ScholarPubMed
Dewey, JF and Shackleton, RM (1984) A model for the evolution of the Grampian tract in the early Caledonides and Appalachians. Nature 312, 115–21.CrossRefGoogle Scholar
Dotzler, N, Taylor, TN and Krings, M (2007) A prasinophycean alga of the genus Cymatiosphaera in the Early Devonian Rhynie chert. Review of Palaeobotany and Palynology 147, 106–11.CrossRefGoogle Scholar
Draut, AE, Clift, PD, Amato, JM, Blusztajn, J and Schouten, H (2009) Arc–continent collision and the formation of continental crust: a new geochemical and isotopic record from the Ordovician Tyrone Igneous Complex, Ireland. Journal of the Geological Society 166, 485–500.CrossRefGoogle Scholar
Droop, GTR and Charnley, NR (1985) Comparative geobarometry of pelitic hornfelses associated with the Newer Gabbros: a preliminary study. Journal of the Geological Society 142, 53–62.CrossRefGoogle Scholar
Dubinin, VB (1962) Class Acaromorpha: mites or gnathosomic chelicerate arthropods. In Fundamentals of Palaeontology (ed. Rodendorf, B.), pp. 447–73. Moscow: Academy of Sciences of the USSR. Google Scholar
Dunlop, JA (1998) A redescription of the trigonotarbid arachnid Pseudokreischeria pococki (Gill, 1924). Bulletin of the British Arachnological Society 11, 49–53.Google Scholar
Dunlop, JA, Anderson, LI, Kerp, H and Hass, H (2003) Preserved organs of Devonian harvestmen. Nature 425, 916.CrossRefGoogle ScholarPubMed
Dunlop, JA, Anderson, LI, Kerp, H and Hass, H (2004) A harvestman (Arachnida: Opiliones) from the Early Devonian Rhynie cherts, Aberdeenshire, Scotland. Transactions of the Royal Society of Edinburgh: Earth Sciences 94, 341–54.CrossRefGoogle Scholar
Dunlop, J and Garwood, R (2014) Tomographic reconstruction of the exceptionally preserved trigonotarbid arachnid Eophrynus prestvicii . Acta Palaeontologica Polonica 59, 443–54.Google Scholar
Dunlop, JA and Garwood, RJ (2018) Terrestrial invertebrates in the Rhynie chert ecosystem. Philosophical Transactions of the Royal Society B: Biological Sciences 373, 20160493. doi: 10.1098/rstb.2016.0493.CrossRefGoogle ScholarPubMed
Dunlop, JA and Horrocks, CA (1997) Phalangiotarbid arachnids from the Coal Measures of Lancashire, UK. Geological Magazine 134, 369–81.CrossRefGoogle Scholar
Dunlop, JA, Scholtz, G. and Selden, PA (2013) Water-to-land transitions. In Arthropod Biology and Evolution (eds Minelli, A, Boxshall, G and Fusco, G), pp. 417–39. Berlin and Heidelberg: Springer.CrossRefGoogle Scholar
Edwards, DS (1980) Evidence for the sporophytic status of the lower Devonian plant Rhynia gwynne-vaughanii Kidston and Lang. Review of Palaeobotany and Palynology 29, 177–88.CrossRefGoogle Scholar
Edwards, DS (1986) Aglaophyton major, a non-vascular land-plant from the Devonian Rhynie Chert. Botanical Journal of the Linnean Society 93, 173–204.CrossRefGoogle Scholar
Edwards, D, Kenrick, P and Dolan, L (2018) History and contemporary significance of the Rhynie cherts – our earliest preserved terrestrial ecosystem. Philosophical Transactions of the Royal Society B: Biological Sciences 373, 20160489. doi: 10.1098/rstb.2016.0489.CrossRefGoogle ScholarPubMed
Edwards, DS and Lyon, AG (1983) Algae from the Rhynie Chert. Botanical Journal of the Linnean Society 86, 37–55.CrossRefGoogle Scholar
Edwards, D and Rogerson, ECW (1979) New records of fertile Rhyniophytina from the late Silurian of Wales. Geological Magazine 116, 93–8.CrossRefGoogle Scholar
Eggert, DA (1974) The Sporangium of Horneophyton lignieri (Rhyniophytina). American Journal of Botany 61, 405–13.CrossRefGoogle Scholar
El-Saadawy, WE-S and Lacey, WS (1979) Observations on Nothia aphylla Lyon ex Høeg. Review of Palaeobotany and Palynology 27, 119–47.CrossRefGoogle Scholar
Engel, MS and Grimaldi, DA (2004) New light shed on the oldest insect. Nature 427, 627–30.CrossRefGoogle ScholarPubMed
Ewers, GR (1991) Oxygen isotopes and the recognition of siliceous sinters in epithermal ore deposits. Economic Geology 86, 173–8.CrossRefGoogle Scholar
Fayers, SR, Dunlop, JA and Trewin, NH (2005) A new Early Devonian trigonotarbid arachnid from the Windyfield Chert, Rhynie, Scotland. Journal of Systematic Palaeontology 2, 269–84.CrossRefGoogle Scholar
Fayers, S and Trewin, N (2003) A new crustacean from the Early Devonian Rhynie chert, Aberdeenshire, Scotland. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 93, 355–82.CrossRefGoogle Scholar
Fayers, SR and Trewin, NH (2004) A review of the palaeoenvironments and biota of the Windyfield chert. Transactions of the Royal Society of Edinburgh: Earth Sciences 94, 325–39.CrossRefGoogle Scholar
Garwood, RJ and Dunlop, JA (2010) Fossils explained 58 – Trigonotarbids. Geology Today 26, 34–7.CrossRefGoogle Scholar
Garwood, RJ, Dunlop, JA, Knecht, BJ and Hegna, TA (2017) The phylogeny of fossil whip spiders. BMC Evolutionary Biology 17, 105. doi: 10.1186/s12862-017-0931-1.CrossRefGoogle ScholarPubMed
Garwood, RJ and Dunlop, JA (2014) The walking dead: Blender as a tool for paleontologists with a case study on extinct arachnids. Journal of Paleontology 88, 735–46.CrossRefGoogle Scholar
Garwood, RJ, Dunlop, JA and Sutton, MD (2009) High-fidelity X-ray micro-tomography reconstruction of siderite-hosted Carboniferous arachnids. Biology Letters 5, 841–4.CrossRefGoogle ScholarPubMed
Garwood, RJ and Edgecombe, GD (2011) Early terrestrial animals, evolution, and uncertainty. Evolution: Education and Outreach 4, 489–501.Google Scholar
Garwood, RJ, Sharma, PP, Dunlop, JA and Giribet, G (2014) A Paleozoic stem group to mite harvestmen revealed through integration of phylogenetics and development. Current Biology 24, 1017–23.CrossRefGoogle ScholarPubMed
Garwood, RJ and Sutton, MD (2012) The enigmatic arthropod Camptophyllia . Palaeontologia Electronica 15, 12 pp. doi: 10.1111/1475-4983.00174.Google Scholar
The Geological Magazine (1921) Obituary Henry Woodward. Geological Magazine 58, 481–4.CrossRefGoogle Scholar
Gill, EL (1911) II.–A Carboniferous arachnid from Lancashire. Geological Magazine 8, 395–8.CrossRefGoogle Scholar
Gill, EL (1924) Fossil arthropods from the Tyne Coalfield. Geological Magazine 61, 455–71.CrossRefGoogle Scholar
Gray, J, Boucot, AJ, Grahn, Y and Himes, G (1992) A new record of early Silurian land plant spores from the Paraná Basin, Paraguay (Malvinokaffric Realm). Geological Magazine 129, 741–52.CrossRefGoogle Scholar
Greenslade, PJ and Whalley, P (1986) The systematic position of Rhyniella praecursor Hirst and Maulik (Collembola), the earliest known hexapod. In Second International Symposium on Apterygota, Siena, Italy (ed. Dallai, R), pp. 319–23.Google Scholar
Grimaldi, D and Engel, MS (2005) Evolution of the Insects. Cambridge University Press.Google Scholar
Habgood, K, Hass, H and Kerp, H (2003) Evidence for an early terrestrial food web: coprolites from the Early Devonian Rhynie chert. Transactions of the Royal Society of Edinburgh: Earth Sciences 94, 371–89.CrossRefGoogle Scholar
Hass, H, Taylor, TN and Remy, W (1994) Fungi from the Lower Devonian Rhynie chert: mycoparasitism. American Journal of Botany 81, 29–37.CrossRefGoogle Scholar
Haug, C (2018) Feeding strategies in arthropods from the Rhynie and Windyfield cherts: ecological diversification in an early non-marine biota. Philosophical Transactions of the Royal Society B: Biological Sciences 373, 20160492. doi: 10.1098/rstb.2016.0492.CrossRefGoogle Scholar
Haug, C and Haug, JT (2017) The presumed oldest flying insect: more likely a myriapod? PeerJ 5, e3402. doi: 10.7717/peerj.3402.CrossRefGoogle ScholarPubMed
Hetherington, AJ and Dolan, L (2018a) Bilaterally symmetric axes with rhizoids composed the rooting structure of the common ancestor of vascular plants. Philosophical Transactions of the Royal Society B: Biological Sciences 373, 20170042. doi: 10.1098/rstb.2016.0042.CrossRefGoogle ScholarPubMed
Hetherington, AJ and Dolan, L (2018b) Stepwise and independent origins of roots among land plants. Nature 561, 235–8.CrossRefGoogle ScholarPubMed
Hirst, S (1923) On some arachnid remains from the Old Red Sandstone. Annals and Magazine of Natural History 12, 455–74.CrossRefGoogle Scholar
Hirst, S and Maulik, S (1926) On some arthropod remains from the Rhynie chert (Old Red Sandstone). Geological Magazine 63, 69–71.CrossRefGoogle Scholar
Jackson, JW, Brade-Birke, HK and Brade-Birke, SG (1919) III. – Notes on Myriopoda. XIX. A revision of some fossil material from Sparth Bottoms, Lancs. Geological Magazine 6, 406–11.CrossRefGoogle Scholar
Jarzembowski, EA and Schneider, JW (2007) The stratigraphical potential of blattodean insects from the late Carboniferous of southern Britain. Geological Magazine 144, 449–56.CrossRefGoogle Scholar
Johnson, EW, Briggs, DEG, Suthren, RJ, Wright, JL and Tunnicliff, SP (1994) Non-marine arthropod traces from the subaerial Ordovician Borrowdale volcanic group, English Lake District. Geological Magazine 131, 395–406.CrossRefGoogle Scholar
Jones, FM, Dunlop, JA, Friedman, M and Garwood, RJ (2014) Trigonotarbus johnsoni Pocock, 1911, revealed by X-ray computed tomography, with a cladistic analysis of the extinct trigonotarbid arachnids. Zoological Journal of the Linnean Society 172, 49–70.CrossRefGoogle Scholar
Jones, T and Woodward, H (1899) II. – Contributions to Fossil Crustacea. Geological Magazine 6, 388–95.CrossRefGoogle Scholar
Kamenz, C, Dunlop, JA, Scholtz, G, Kerp, H and Hass, H (2008) Microanatomy of Early Devonian book lungs. Biology Letters 4, 212–15.CrossRefGoogle ScholarPubMed
Karatygin, IV, Snigirevskaya, NS and Demchenko, KN (2006) Species of the genus Glomites kring as plant mycobionts in Early Devonian ecosystems. Paleontological Journal 40, 572–9.CrossRefGoogle Scholar
Karatygin, IV, Snigirevskaya, NS and Vikulin, S (2009) The most ancient terrestrial lichen Winfrenatia reticulata: a new find and new interpretation. Palaeontological Journal 43, 107–14.CrossRefGoogle Scholar
Kelman, R, Feist, M, Trewin, NH and Hass, H (2003) Charophyte algae from the Rhynie chert. Transactions of the Royal Society of Edinburgh: Earth Sciences 94, 445–55.CrossRefGoogle Scholar
Kenrick, P (2018) Changing expressions: a hypothesis for the origin of the vascular plant life cycle. Philosophical Transactions of the Royal Society B: Biological Sciences 373, 20170149. doi: 10.1098/rstb.2016.0149.CrossRefGoogle ScholarPubMed
Kenrick, P and Strullu-Derrien, C (2014) The origin and early evolution of roots. Plant Physiology 166, 570–80.CrossRefGoogle ScholarPubMed
Kenrick, P, Wellman, CH, Schneider, H and Edgecombe, GD (2012) A timeline for terrestrialization: consequences for the carbon cycle in the Palaeozoic. Philosophical Transactions of the Royal Society B: Biological Sciences 367, 519–36.CrossRefGoogle ScholarPubMed
Kerp, H (2018) Organs and tissues of Rhynie chert plants. Philosophical Transactions of the Royal Society B: Biological Sciences 373, 20160495. doi: 10.1098/rstb.2016.0495.CrossRefGoogle ScholarPubMed
Kerp, H, Hass, H and Mosbrugger, V (2001) New data on Nothia aphylla Lyon 1964 ex El-Saadawy et Lacey 1979, a poorly known plant from the Lower Devonian Rhynie Chert. In Plants Invade the Land: Evolutionary and Environmental Perspectives (eds. Gensel, PG & Edwards, D), pp. 52–82. New York, USA: Columbia University Press.Google Scholar
Kerp, H, Trewin, NH and Hass, H (2003) New gametophytes from the Early Devonian Rhynie chert. Transactions of the Royal Society of Edinburgh: Earth Sciences 94, 411–28.CrossRefGoogle Scholar
Kevan, PG, Chaloner, WG and Saville, DBO (1975) Interrelationships of early terrestrial arthropods and plants. Palaeontology 18, 391–417.Google Scholar
Kidston, R and Lang, WH (1917) XXIV. – On Old Red Sandstone Plants showing structure, from the Rhynie Chert Bed, Aberdeenshire. Part I. Rhynia Gwynne-Vaughani, Kidston and Lang. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 87, 427–50.CrossRefGoogle Scholar
Kidston, R and Lang, WH (1920a) XXIV. – On Old Red Sandstone Plants showing structure, from the Rhynie Chert Bed, Aberdeenshire. Part II. Additional notes on Rhynia Gwynne-Vaughani, Kidston and Lang; with descriptions of Rhynia major, n.sp., and Hornea Lignieri, n.g., n.sp. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 52, 603–27.CrossRefGoogle Scholar
Kidston, R and Lang, WH (1920b) XXVI. – On Old Red Sandstone plants showing structure, from the Rhynie Chert Bed, Aberdeenshire. Part III. Asteroxylon Mackiei, Kidston and Lang. Transactions of the Royal Society of Edinburgh 52, 643–80.CrossRefGoogle Scholar
Kidston, R and Lang, W (1921a) XXXIII. – On Old Red Sandstone plants showing structure, from the Rhynie Chert Bed, Aberdeenshire. Part V. The Thallophyta occurring in the peat-bed; the succession of the plants throughout a vertical section of the bed, and the conditions of accumulation and preservation of the deposit. Transactions of the Royal Society of Edinburgh 52, 855–902.CrossRefGoogle Scholar
Kidston, R and Lang, WH (1921b) XXXII.–On Old Red Sandstone plants showing structure, from the Rhynie Chert Bed, Aberdeenshire. Part IV. Restorations of the vascular cryptogams, and discussion of their bearing on the general morphology of the Pteridophyta and the origin of the organisation of land-plants. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 52, 831–54.CrossRefGoogle Scholar
Kindle, EM (1913) I.–Note on a process of fossilization in the Palæozoic Lycopods. Geological Magazine 10, 336–40.CrossRefGoogle Scholar
Kinny, PD, Strachan, RA, Friend, CRL, Kocks, H, Rogers, G and Paterson, BA (2003) U–Pb geochronology of deformed metagranites in central Sutherland, Scotland: evidence for widespread late Silurian metamorphism and ductile deformation of the Moine Supergroup during the Caledonian orogeny. Journal of the Geological Society 160, 259–69.CrossRefGoogle Scholar
Kofuji, R, Yagita, Y, Murata, T and Hasebe, M (2018) Antheridial development in the moss Physcomitrella patens: implications for understanding stem cells in mosses. Philosophical Transactions of the Royal Society B: Biological Sciences 373, 20160494. doi: 10.1098/rstb.2016.0494.CrossRefGoogle ScholarPubMed
Krings, M and Harper, CJ (2019) A microfossil resembling Merismopedia (Cyanobacteria) from the 410-million-yr-old Rhynie and Windyfield cherts – Rhyniococcus uniformis revisited. Nova Hedwigia 108, 17–35.CrossRefGoogle Scholar
Krings, M, Harper, CJ and Taylor, EL (2018) Fungi and fungal interactions in the Rhynie chert: a review of the evidence, with the description of Perexiflasca tayloriana gen. et sp. nov.†. Philosophical Transactions of the Royal Society B: Biological Sciences 373, 20160500. doi: 10.1098/rstb.2016.0500.CrossRefGoogle ScholarPubMed
Krings, M, Kerp, H, Hass, H, Taylor, TN and Dotzler, N (2007a) A filamentous cyanobacterium showing structured colonial growth from the Early Devonian Rhynie chert. Review of Palaeobotany and Palynology 146, 265–76.CrossRefGoogle Scholar
Krings, M and Sergeev, VN (2019) A coccoid, colony-forming cyanobacterium from the Lower Devonian Rhynie chert that resembles Eucapsis (Synechococcales) and Entophysalis (Chroococcales). Review of Palaeobotany and Palynology 268, 65–71. doi: 10.1016/j.revpalbo.2019.06.002.CrossRefGoogle Scholar
Krings, M, Taylor, TN, Hass, H, Kerp, H, Dotzler, N and Hermsen, EJ (2007b) An alternative mode of early land plant colonization by putative endomycorrhizal fungi. Plant Signaling & Behavior 2, 125–6.CrossRefGoogle ScholarPubMed
Krings, M, Taylor, TN, Hass, H, Kerp, H, Dotzler, N and Hermsen, EJ (2007c) Fungal endophytes in a 400-million-yr-old land plant: infection pathways, spatial distribution, and host responses. The New Phytologist 174, 648–57.CrossRefGoogle Scholar
Kustatscher, E, Dotzler, N, Taylor, TN and Krings, M (2014a) Microalgae from the Lower Devonian Rhynie chert: a new Cymatiosphaera. Zitteliana 54, 165–9.Google Scholar
Kustatscher, E, Dotzler, N, Taylor, TN and Krings, M (2014b) Microfossils with suggested affinities to the Pyramimonadales (Pyramimonadophyceae, Chlorophyta) from the Lower Devonian Rhynie Chert. Acta Palaeobotanica 54, 163–71.CrossRefGoogle Scholar
Labandeira, CC (2005) Invasion of the continents: cyanobacterial crusts to tree-inhabiting arthropods. Trends in Ecology & Evolution 20, 253–62.CrossRefGoogle ScholarPubMed
Lambert, RSJ and McKerrow, WS (1976) The Grampian Orogeny. Scottish Journal of Geology 12, 271–92.CrossRefGoogle Scholar
Levitan, MA, Dontsova, EI, Lisitsyn, AP and Bogdanov, YA (1975) The origin of chert in the sediments of the Pacific Ocean from data of oxygen isotopic analysis and a study of the distribution of chert. Geochemistry International 12, 95–104.Google Scholar
Lozano-Fernandez, J, Carton, R, Tanner, AR, Puttick, MN, Blaxter, M, Vinther, J, Olesen, J, Giribet, G, Edgecombe, GD and Pisani, D (2016) A molecular palaeobiological exploration of arthropod terrestrialization. Philosophical Transactions of the Royal Society B: Biological Sciences 371, 20150133. doi: 10.1098/rstb.2016.0133.CrossRefGoogle ScholarPubMed
Lyon, AG (1957) Germinating spores in the Rhynie Chert. Nature 180, 1219–21.CrossRefGoogle Scholar
Lyon, AG and Edwards, D (1991) The first zosterophyll from the Lower Devonian Rhynie Chert, Aberdeenshire. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 82, 324–32.CrossRefGoogle Scholar
Mark, DF, Rice, CM, Fallick, AE, Trewin, NH, Lee, MR, Boyce, A and Lee, JKW (2011) 40Ar/39Ar dating of hydrothermal activity, biota and gold mineralization in the Rhynie hot-spring system, Aberdeenshire, Scotland. Geochimica et Cosmochimica Acta 75, 555–69.CrossRefGoogle Scholar
Mark, DF, Rice, CM and Trewin, NH (2013) Discussion on ‘A high-precision U–Pb age constraint on the Rhynie Chert Konservat-Lagerstätte: time scale and other implications’: Journal, Vol. 168, 863–872. Journal of the Geological Society 170, 701–3.CrossRefGoogle Scholar
Marshall, JEA (1991) Palynology of the Stonehaven Group, Scotland: evidence for a Mid Silurian age and its geological implications. Geological Magazine 128, 283–6.CrossRefGoogle Scholar
Mills, BJW, Batterman, SA and Field, KJ (2018) Nutrient acquisition by symbiotic fungi governs Palaeozoic climate transition. Philosophical Transactions of the Royal Society B: Biological Sciences 373, 20160503. doi: 10.1098/rstb.2016.0503.CrossRefGoogle ScholarPubMed
Oehler, D (1977) Pyrenoid-like structures in late Precambrian algae from the Bitter Springs Formation of Australia. Journal of Paleontology 51, 885–901.Google Scholar
Parry, SF, Noble, SR, Crowley, QG and Wellman, CH (2011) A high-precision U–Pb age constraint on the Rhynie Chert Konservat-Lagerstätte: time scale and other implications. Journal of the Geological Society, London 168, 863–72.CrossRefGoogle Scholar
Parry, SF, Noble, SR, Crowley, QG and Wellman, CH (2013) Reply to discussion on ‘A high-precision U–Pb age constraint on the Rhynie Chert Konservat-Lagerstätte: time scale and other implications’: Journal, 168, 863–872. Journal of the Geological Society 170, 703–6.CrossRefGoogle Scholar
Pocock, RI (1902) II. – Eophrynus and allied Carboniferous Arachnida. Geological Magazine 9, 439–48.CrossRefGoogle Scholar
Pocock, RI (1903a) II.–A new Carboniferous arachnid. Geological Magazine 4, 247–51.CrossRefGoogle Scholar
Pocock, RI (1903b) V.–Details of Further remarks upon the Carboniferous Arachnid Anthracosiro with the description of a second species of the genus. Geological Magazine 4, 405–8.CrossRefGoogle Scholar
Poinar, G (2015) The geological record of parasitic nematode evolution. In Fossil Parasites (eds Littlewood, T and De Baets, K), pp. 53–92. Advances in Parasitology 90. Amsterdam: Elsevier.CrossRefGoogle Scholar
Poinar, G, Kerp, H and Hass, H (2008) Palaeonema phyticum gen. n., sp. n. (Nematoda: Palaeonematidae fam. n.), a Devonian nematode associated with early land plants. Nematology 10, 9–14.Google Scholar
Powell, CL, Trewin, NH and Edwards, D (2000) Palaeoecology and plant succession in a borehole through the Rhynie cherts, Lower Old Red Sandstone, Scotland. In New Perspectives on the Old Red Sandstone (eds Friend, PF and Williams, BPJ), pp. 439–57. Geological Society of London, Special Publication no. 180.Google Scholar
Preston, LJ and Genge, MJ (2010) The Rhynie Chert, Scotland, and the search for life on Mars. Astrobiology 10, 549–60.CrossRefGoogle ScholarPubMed
Raven, JA (2018) Evolution and palaeophysiology of the vascular system and other means of long-distance transport. Philosophical Transactions of the Royal Society B: Biological Sciences 373, 20160497. doi: 10.1098/rstb.2016.0497.CrossRefGoogle ScholarPubMed
Remy, W, Gensel, PG and Hass, H (1993) The gametophyte generation of some Early Devonian land plants. International Journal of Plant Sciences 154, 35–58.CrossRefGoogle Scholar
Remy, W and Remy, R (1980a) Devonian gametophytes with anatomically preserved gametangia. Science 208, 295–6.CrossRefGoogle ScholarPubMed
Remy, W and Remy, R (1980b) Lyonophyton rhyniensis n. gen. et nov. spec., ein Gametophyt aus dem Chert von Rhynie (Unterdevon, Schottland). Argumenta Palaeobotanica 6, 37–72.Google Scholar
Rice, CM and Ashcroft, WA (2003) The geology of the northern half of the Rhynie Basin, Aberdeenshire, Scotland. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 94, 299–308.CrossRefGoogle Scholar
Rice, CM, Ashcroft, WA, Batten, DJ, Boyce, AJ, Caulfield, JBD, Fallick, AE, Hole, MJ, Jones, E, Pearson, MJ, Rogers, G, Saxton, JM, Stuart, FM, Trewin, NH and Turner, G (1995) A Devonian auriferous hot spring system, Rhynie, Scotland. Journal of the Geological Society 152, 229–50.CrossRefGoogle Scholar
Rice, CM and Trewin, NH (1988) A Lower Devonian gold-bearing hot spring system, Rhynie, Scotland. Transactions of the Institute of Mining and Metallurgy 97, B141–4.Google Scholar
Rice, CM, Trewin, NH and Anderson, LI (2002) Geological setting of the Early Devonian Rhynie cherts, Aberdeenshire, Scotland: an early terrestrial hot spring system. Journal of the Geological Society 159, 203–14.CrossRefGoogle Scholar
Richardson, JB (1967) Some British Lower Devonian spore assemblages and their stratigraphic significance. Review of Palaeobotany and Palynology 1, 111–29.CrossRefGoogle Scholar
Rickards, RB (2000) The age of the earliest club mosses: the Silurian Baragwanathia flora in Victoria, Australia. Geological Magazine 137, 207–9.CrossRefGoogle Scholar
Rolfe, WDI (1967) Rochdalia, a Carboniferous insect nymph. Palaeontology 10, 307–13.Google Scholar
Rota-Stabelli, O, Daley, AC and Pisani, D (2013) Molecular timetrees reveal a Cambrian colonization of land and a new scenario for ecdysozoan evolution. Current Biology 23, 392–8.CrossRefGoogle Scholar
Scott, DH (1923) Studies in Fossil Botany. Volume II: Spermophyta, 3rd ed. London: A & C Black Ltd, 446 pp.Google Scholar
Scott, DH (1924) Extinct Plants and Problems of Evolution. London: Macmillan & Co., 240 pp.Google Scholar
Scourfield, DJ (1926) On a new type of crustacean from the Old Red Sandstone (Rhynie Chert Bed, Aberdeenshire) – Lepidocaris rhyniensis, gen. et sp. nov. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 214, 153–87.CrossRefGoogle Scholar
Scourfield, DJ (1940) The oldest known fossil insect (Rhyniella praecursor Hirst & Maulik) – further details from additional specimens. Proceedings of the Linnean Society of London 152, 113–31.CrossRefGoogle Scholar
Scudder, SH (1876) Fossil Palæozoic insects. Geological Magazine 3, 519–20.CrossRefGoogle Scholar
Selden, P (2012) Terrestrialisation (Precambrian-Devonian). In Encyclopedia of Life Sciences. Chichester: John Wiley & Sons, Ltd. doi: 10.1002/9780470015902. a0001641.pub3.Google Scholar
Selden, PA, Shear, WA and Bonamo, PMP (1991) A spider and other arachnids from the Devonian of New York, and reinterpretations of Devonian Araneae. Palaeontology 34, 241–81.Google Scholar
Smith, A, Braddy, SJ, Marriott, SB and Briggs, DEG (2003) Arthropod trackways from the Early Devonian of South Wales: a functional analysis of producers and their behaviour. Geological Magazine 140, 63–72.CrossRefGoogle Scholar
Stephenson, D (ed.) (2000) Caledonian Igneous rocks of Great Britain. Peterborough: Joint Nature Conservation Committee, 648 pp.Google Scholar
Strachan, RA (2012) The Grampian Orogeny: mid-Ordovician arc-continent collision along the Laurentian margin of Iapetus. In Geological History of Britain and Ireland (eds Woodcock, N & Strachan, R), pp. 91–109. Chichester: John Wiley & Sons, Ltd.CrossRefGoogle Scholar
Strachan, RA, Smith, M, Harris, AL, Fettes, DJ and Trewin, NH (2003) The Northern Highland and Grampian terranes. In The Geology of Scotland (ed. Trewin, NH), pp. 81–147. London and Bath: Geological Society of London.CrossRefGoogle Scholar
Strullu-Derrien, C, Goral, T, Longcore, JE, Olesen, J, Kenrick, P and Edgecombe, GD (2016) A new Chytridiomycete fungus intermixed with crustacean resting eggs in a 407-million-year-old continental freshwater environment. PLOS ONE 11, e0167301. doi: 10.1371/journalpone.0167301.CrossRefGoogle Scholar
Strullu-Derrien, C, Kenrick, P, Goral, T and Knoll, AH (2019) Testate amoebae in the 407-million-year-old Rhynie Chert. Current Biology 29, R93–R95.CrossRefGoogle ScholarPubMed
Strullu-Derrien, C, Kenrick, P, Pressel, S, Duckett, JG, Rioult, J-P and Strullu, D-G (2014) Fungal associations in Horneophyton ligneri from the Rhynie Chert (c. 407 million year old) closely resemble those in extant lower land plants: novel insights into ancestral plant-fungus symbioses. The New Phytologist 203, 964–79.CrossRefGoogle ScholarPubMed
Strullu-Derrien, C, Spencer, ART, Goral, T, Dee, J, Honegger, R, Kenrick, P, Longcore, JE and Berbee, ML (2018) New insights into the evolutionary history of Fungi from a 407 Ma Blastocladiomycota fossil showing a complex hyphal thallus. Philosophical Transactions of the Royal Society B: Biological Sciences 373, 20160502. doi: 10.1098/rstb.2016.0502.CrossRefGoogle ScholarPubMed
Strullu-Derrien, C, Wawrzyniak, Z, Goral, T and Kenrick, P (2015) Fungal colonization of the rooting system of the early land plant Asteroxylon mackiei from the 407-Myr-old Rhynie Chert (Scotland, UK). Botanical Journal of the Linnean Society 179, 201–3.CrossRefGoogle Scholar
Taylor, TN, Hass, H and Kerp, H (1997) A cyanolichen from the Lower Devonian Rhynie chert. American Journal of Botany 84, 992–1004.CrossRefGoogle ScholarPubMed
Taylor, TN, Klavins, SD, Krings, M, Taylor, EL, Kerp, H and Hass, H (2003) Fungi from the Rhynie chert: a view from the dark side. Transactions of the Royal Society of Edinburgh: Earth Sciences 94, 457–73.CrossRefGoogle Scholar
Taylor, TN and Krings, M (2015) A colony-forming microorganism with probable affinities to the Chroococcales (cyanobacteria) from the Lower Devonian Rhynie chert. Review of Palaeobotany and Palynology 219, 147–56.CrossRefGoogle Scholar
Taylor, TN, Remy, W and Hass, H (1992) Parasitism in a 400-million-year-old green alga. Nature 357, 493–4.CrossRefGoogle Scholar
Taylor, TN, Remy, W, Hass, H and Kerp, H (1995a) Fossil arbuscular Mycorrhizae from the Early Devonian. Mycologia 87, 560–73.CrossRefGoogle Scholar
Taylor, TN, Remy, W, Hass, H and Kerp, H (1995b) The oldest fossil lichen. Nature 378, 244.CrossRefGoogle Scholar
Tillyard, RJ (1928) Some remarks on the Devonian fossil insects from the Rhynie chert beds, Old Red Sandstone. Transactions of the Royal Entomological Society of London 76, 65–71.CrossRefGoogle Scholar
Tosdal, R and Richards, J (2001) Magmatic and structural controls on the development of porphyry Cu ± Mo ± Au deposits. Reviews in Economic Geology 14, 157–81.Google Scholar
Trewin, NH (1994) Depositional environment and preservation of biota in the Lower Devonian hot-springs of Rhynie, Aberdeenshire, Scotland. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 84, 433–42.CrossRefGoogle Scholar
Trewin, NH (1996) The Rhynie cherts: an early Devonian ecosystem preserved by hydrothermal activity. Ciba Foundation Symposium 202, 131–45; discussion 145–9.Google ScholarPubMed
Trewin, NH, Fayers, SR and Kelman, R (2003) Subaqueous silicification of the contents of small ponds in an Early Devonian hot-spring complex, Rhynie, Scotland. Canadian Journal of Earth Sciences 40, 1697–1712.CrossRefGoogle Scholar
Trewin, NH and Rice, CM (1992) Stratigraphy and sedimentology of the Devonian Rhynie chert locality. Scottish Journal of Geology 28, 37–47.CrossRefGoogle Scholar
Trewin, NH and Rice, CM (2004) The Rhynie Hot-Spring System: Geology, Biota and Mineralisation. Edinburgh: Royal Society of Edinburgh, 246 pp.Google Scholar
Trewin, NH and Wilson, E (2004) Correlation of the Early Devonian Rhynie chert beds between three boreholes at Rhynie, Aberdeenshire. Scottish Journal of Geology 40, 73–81.CrossRefGoogle Scholar
Vecoli, M, Clément, G and Meyer-Berthaud, B (eds) (2010) The Terrestrialization Process: Modelling Complex Interactions at the Biosphere–Geosphere Interface. Geological Society of London, Special Publication no. 339.Google Scholar
Wacey, D, Battison, L, Garwood, RJ, Hickman-Lewis, K and Brasier, MD (2017) Advanced analytical techniques for studying the morphology and chemistry of Proterozoic microfossils. In Earth System Evolution and Early Life: A Celebration of the Work of Martin Brasier (eds Brasier, AT, McIlroy, D and McLoughlin, N), pp. 81–104. Geological Society of London, Special Publication no. 448.Google Scholar
Wang, B, Dunlop, JA, Selden, PA, Garwood, RJ, Shear, WA, Müller, P and Lei, X (2018) Cretaceous arachnid Chimerarachne yingi gen. et sp. nov. illuminates spider origins. Nature Ecology & Evolution 2, 614–22.CrossRefGoogle ScholarPubMed
Wellman, CH (2004) Palaeoecology and palaeophytogeography of the Rhynie Chert plants: evidence from integrated analysis of in situ and dispersed spores. Proceedings of the Royal Society B: Biological Sciences 271, 985–92.CrossRefGoogle ScholarPubMed
Wellman, CH (2006) Spore assemblages from the Lower Devonian ‘Lower Old Red Sandstone’ deposits of the Rhynie Outlier, Scotland. Transactions of the Royal Society of Edinburgh: Earth Sciences 97, 167–211.CrossRefGoogle Scholar
Wellman, CH (2018) Palaeoecology and palaeophytogeography of the Rhynie chert plants: further evidence from integrated analysis of in situ and dispersed spores. Philosophical Transactions of the Royal Society B: Biological Sciences 373, 20160491. doi: 10.1098/rstb.2016.0491.CrossRefGoogle ScholarPubMed
Wellman, CH, Thomas, RG, Edwards, D and Kenrick, P (1998) The Cosheston Group (Lower Old Red Sandstone) in southwest Wales: age, correlation and palaeobotanical significance. Geological Magazine 135, 397–412.CrossRefGoogle Scholar
Whalley, P. and Jarzembowski, EA (1981) A new assessment of Rhyniella, the earliest known insect, from the Devonian of Rhynie, Scotland. Nature 291, 317.CrossRefGoogle Scholar
Woodward, H (1868) III. – Contributions to British Fossil Crustacea. Geological Magazine 5, 258–61.CrossRefGoogle Scholar
Woodward, H (1870) I. – Contributions to British Fossil Crustacea. Geological Magazine 7, 493–7.CrossRefGoogle Scholar
Woodward, H (1871a) II. – On Euphoberia Brownii, H. Woodw., a new species of myriapod from the Coal-measures of the West of Scotland. Geological Magazine 8, 102–4.CrossRefGoogle Scholar
Woodward, H (1871b) I. – On the discovery of a new and very perfect arachnide from the ironstone of the Dudley Coal-field. Geological Magazine 8, 383–7.CrossRefGoogle Scholar
Woodward, H (1871c) I. – Relics of the Carboniferous and other old land-subfaces. Geological Magazine 8, 492–500.CrossRefGoogle Scholar
Woodward, H (1872) I. – On a new Arachnide from the Coal-measures of Lancashire. Geological Magazine 9, 385–7.CrossRefGoogle Scholar
Woodward, H (1873) V. – On some supposed fossil remains of Arachnida (?) and Myriopoda from the English Coal-Measures. Geological Magazine 10, 104–12.CrossRefGoogle Scholar
Woodward, H (1874) I. – On the dawn and development of life on the Earth. Geological Magazine 1, 289–300.CrossRefGoogle Scholar
Woodward, H (1885) II. – Iguanodon Mantelli, Meyer. Geological Magazine 2, 10–15.CrossRefGoogle Scholar
Woodward, H (1887a) I. – On some spined myriapods from the Carboniferous series of England. Geological Magazine 4, 1–10.CrossRefGoogle Scholar
Woodward, H (1887b) I. – On the discovery of the larval stage of a cockroach, Etoblattina Peachii, H. Woodw., from the Coal-Measures of Kilmaurs, Ayrshire. Geological Magazine 4, 433–5.CrossRefGoogle Scholar
Woodward, H (1887c) I. – Some new British Carboniferous cockroaches. Geological Magazine 24, 49–58.CrossRefGoogle Scholar
Woodward, H (1906) IV. – A fossil insect from the Coal-measures of Longton, North Staffordshire. Geological Magazine 3, 25–9.CrossRefGoogle Scholar
Woodward, H (1913a) III. – The position of the Merostomata. Geological Magazine 10, 293–300.CrossRefGoogle Scholar
Woodward, H (1913b) IV. – Rochdalia Parkeri, a new branchiopod crustacean from the Middle Coal-Measures of Sparth, Rochdale. Geological Magazine 10, 352–6.CrossRefGoogle Scholar
Xu, H-H and Wang, Y (2008) The palaeogeographical significance of specimens attributed to Protolepidodendron scharyanum Krejči (Lycopsida) from the Middle Devonian of North Xinjiang, China. Geological Magazine 145, 295–9.CrossRefGoogle Scholar