{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T00:50:35Z","timestamp":1787014235659,"version":"build-2736575974"},"reference-count":83,"publisher":"Wiley","issue":"3","license":[{"start":{"date-parts":[[2021,2,10]],"date-time":"2021-02-10T00:00:00Z","timestamp":1612915200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Geobiology"],"published-print":{"date-parts":[[2021,5]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Modern cryptogamic ground covers (CGCs), comprising assemblages of bryophytes (hornworts, liverworts, mosses), fungi, bacteria, lichens and algae, are thought to resemble early divergent terrestrial communities. However, limited in situ plant and other fossils in the rock record, and a lack of CGC\u2010like soils reported in the pre\u2010Silurian sedimentological record, have hindered understanding of the structure, composition and interactions within the earliest CGCs. A key question is how the earliest CGC\u2010like organisms drove weathering on primordial terrestrial surfaces (regolith), leading to the early stages of soil development as proto\u2010soils, and subsequently contributing to large\u2010scale biogeochemical shifts in the Earth System. Here, we employed a novel qualitative, quantitative and multi\u2010dimensional imaging approach through X\u2010ray micro\u2010computed tomography, scanning electron, and optical microscopy to investigate whether different combinations of modern CGC organisms from primordial\u2010like settings in Iceland develop organism\u2010specific soil forming features at the macro\u2010 and micro\u2010scales. Additionally, we analysed CGCs growing on hard rocky substrates to investigate the initiation of weathering processes non\u2010destructively in 3D. We show that thalloid CGC organisms (liverworts, hornworts) develop thin organic layers at the surface (&lt;1\u00a0cm) with limited subsurface structural development, whereas leafy mosses and communities of mixed organisms form profiles that are thicker (up to\u00a0~\u00a07\u00a0cm), structurally more complex, and more organic\u2010rich. We term these thin layers and profiles proto\u2010soils. Component analyses from X\u2010ray micro\u2010computed tomography data show that thickness and structure of these proto\u2010soils are determined by the type of colonising organism(s), suggesting that the evolution of more complex soils through the Palaeozoic may have been driven by a shift in body plan of CGC\u2010like organisms from flattened and appressed to upright and leafy. Our results provide a framework for identifying CGC\u2010like proto\u2010soils in the rock record and a new proxy for understanding organism\u2013soil interactions in ancient terrestrial biospheres and their contribution to the early stages of soil formation.<\/jats:p>","DOI":"10.1111\/gbi.12431","type":"journal-article","created":{"date-parts":[[2021,2,11]],"date-time":"2021-02-11T01:53:42Z","timestamp":1613008422000},"page":"292-306","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":35,"title":["Cryptogamic ground covers as analogues for early terrestrial biospheres: Initiation and evolution of biologically mediated proto\u2010soils"],"prefix":"10.1111","volume":"19","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6328-3998","authenticated-orcid":false,"given":"Ria L.","family":"Mitchell","sequence":"first","affiliation":[{"name":"Earth Sciences Department The Natural History Museum London UK"},{"name":"Sheffield Tomography Centre (STC) Kroto Research Institute The University of Sheffield Sheffield UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christine","family":"Strullu\u2010Derrien","sequence":"additional","affiliation":[{"name":"Earth Sciences Department The Natural History Museum London UK"},{"name":"Institut de Syst\u00e9matique, Evolution, Biodiversit\u00e9 (ISYEB) UMR7205 Mus\u00e9um National d\u2019Histoire naturelle Sorbonne Universit\u00e9 CNRS Paris France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dan","family":"Sykes","sequence":"additional","affiliation":[{"name":"Imaging and Analysis Centre (IAC) The Natural History Museum London UK"},{"name":"Henry Moseley X\u2010ray Imaging Facility School of Materials The Royce Institute The University of Manchester Manchester UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Silvia","family":"Pressel","sequence":"additional","affiliation":[{"name":"Life Sciences Department The Natural History Museum London UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jeffrey G.","family":"Duckett","sequence":"additional","affiliation":[{"name":"Life Sciences Department The Natural History Museum London UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Paul","family":"Kenrick","sequence":"additional","affiliation":[{"name":"Earth Sciences Department The Natural History Museum London UK"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2021,2,10]]},"reference":[{"key":"e_1_2_7_2_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/ern005"},{"key":"e_1_2_7_3_1","first-page":"307","article-title":"Estimating the Phanerozoic history of the Ascomycota lineages: Combining fossil and molecular data","volume":"77","author":"Beimforde C.","year":"2014","journal-title":"Molecular Phylogenetics and Evolution"},{"key":"e_1_2_7_4_1","doi-asserted-by":"publisher","DOI":"10.1890\/1540-9295(2003)001[0181:TWAYFD]2.0.CO;2"},{"key":"e_1_2_7_5_1","volume-title":"Biological Soil Crusts: Structure, Function, and Management","author":"Belnap J.","year":"2001"},{"key":"e_1_2_7_6_1","first-page":"1534","article-title":"Genomic and fossil windows into the secret lives of the most ancient fungi","volume":"1740","author":"Berbee M. 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