Paleobotany is the study of plant or plant-like fossils.

Lepidodendron leaf fossils are in or on a piece of bituminous coal. Credit: Martin.{{Free media}}

The Lepidodendrales, quillwort-like large tree-like plants from the Carboniferous also left fossils in bituminous coal as on the left.

Def. the "branch of paleontology that deals with the study of plant fossils"[1] is called paleobotany.

"The groups could then be arranged from the dominant group (first rank) to the least common group (fifth rank)."[2]

"The dominant group in the Palaeozoic microplankton is the Acritarcha."[3]

"Pteridophyta is the second dominant group."[4]

"The angiosperms are both the dominant group of land plants and by far the most important plants for human use."[5]

"Statistical analyses indicate that the Phanerozoic history of vascular land plants (tracheophytes) may be interpreted in terms of the successive radiations of four major plant groups, each characterized by a common morphological and/or reproductive grade. Following initial invasion of the land, the diversification of each group coincides with a decline in species numbers of the previously dominant group."[6]


Illustration of Asteroxylon mackiei is from the Rhynie Chert. Credit: Kidston, R., & Lang, W. H.{{free media}}

Asteroxylon ("star-shaped xylem") is an extinct genus of vascular plants of the Division Lycopodiophyta known from anatomically preserved specimens described from the famous Early Devonian Rhynie chert and Windyfield chert in Aberdeenshire, Scotland.[7][8] Asteroxylon is considered the most basal member of the Lycopsida.[9]

This plant consisted of aerial, isotomously and anisotomously branching stems that reached 12 mm in diameter and 40 cm in length.[10] The possibly procumbent aerial stems arose from a leaf-less rhizome which bore smaller-diameter, positively geotropic root-like branches.[10] The rhizomes, which represent an independent origin of roots,[11] reached a depth of up to 20 cm below the surface.[12] The xylem or conducting tissue at the center of the aerial stems is distinctly star-shaped in cross-section and has been considered an early actinostele or an "Asteroxylon-type" protostele.[13] The tracheids are of the primitive annular or helical type (so-called G-type).[14] "Leaves" – not true leaves, but protrusions – were of the form of unbranched strap-shaped enations up to 5 mm long; a single vascular trace branched from the main bundle in the centre of the stem to terminate at the base of each enation.[9][13] Enations and axes bore stomata, indicating that their tissues were capable of photosynthesis.[15]

Asteroxylon differs from other similar Early Devonian lycopsids such as Drepanophycus and Baragwanathia in that the singular vascular leaf trace in these latter plants extends into the leaf.[9] The leaves of Drepanophycus and Baragwanathia are therefore considered to be true microphylls or, alternatively, small leaves.[16]

The type species is Asteroxylon mackiei.


A fossil leaf from the extinct Betula leopoldae, 48.5 million years old; Klondike Mountain Formation, Republic, Ferry County, Washington, USA. Credit: Kevmin.{{free media}}

Betula leopoldae fossils have been identified from a number of locations in Western North America, the 49 mya Klondike Mountain Formation near Republic, Washington[17] along with the Allenby Formation near Princeton, British Columbia,[18] the Falkland fossil site[19] near Falkland, British Columbia, and McAbee Fossil Beds both of the Kamloops Groups Tranquille Formation,[20][21] and the Quilchena fossil site[22] near Quilchena, British Columbia.[23]

In general, the geologic ages for the Okanagan Highland locations are of Early Eocene, with the sites that have current uranium-lead dating or argon–argon dating radiometric dates indicating Ypresian ages, while the undated sites or those given older dates being possibly slightly younger and Lutetian in age.[24]

Betula leopoldae was described from a series of type specimens collected in the Republic, Washington area during the early 1980s. The paratype leaf, UW 31256 plus the holotype leaf UW 39722, are in the paleobotanical collections of Burke Museum, while the counterpart for the holotype, UCMP 9286 is in the University of California Museum of Paleontology in California. Working from these two specimens, the species was studied by Jack A. Wolfe of the University of California and Wesley C. Wehr of the Burke Museum.[17] They published their 1987 type description in a United States Geological Survey monograph on the North Eastern Washington dicot fossils. The specific epithet leopoldae is a matronym recognizing paleobotanist and conservationist Estella Leopold, though this was not noted in the type description.[25] In a paper which appeared that same year, Peter Crane and Ruth Stockey described a series of B. leopoldae leaves along with catkins, flowering bodies, and pollen from the Allenby Formation. Crane and Stockey noted B. leopoldae to be the oldest reproductive plus vegetative record for a Betula species at that time.[18] A B. leopoldae leaf from the Klondike mountain formation was figured by Conrad Labandeira in 2002 which displayed distinct interior foliage feeding damage from insect feeding, in which a series of four leaf blade sections had been removed between successive secondary veins.[26]


Fossil of Calamites, an extinct plant, photographed at Museo di Storia Naturale di Verona. Credit: Ghedoghedo{{free media}}

"Specimens of Calamites cistii (Sphenophyta; Pennsylvanian, France) are described showing endophytic cavities, located in the outer cortex of the stem, a tissue that is rarely preserved. This new record shifts the appearance of this behavior back 60 Ma."[27] Two "specimens of the arborescent Calamites cistii (Sphenophyta) [were] collected from the Pennsylvanian basin of Graissessac (Hérault, France)".[27] "The specimens belong to the species Calamites cistii Brongniart, 1828 (Sphenophyta). They are housed in the Collections de Paléobotanique, Service général des Collections, University Montpellier 2 (LPM)."[27]


This image is of Ginkgoites huttoni from Scalby Ness, Scarborough, England. Credit: Ghedoghedo.{{Free media}}
Fossil Ginkgo huttonii leaves are from the Jurassic of England. Credit: Dlloyd.{{free media}}

The image at the right shows fronds impressed onto shale in a specimen on display at the Paläontologische Museum München. The fossil is from Scalby Ness, Scarborough, England.

Ginkgo huttonii is an extinct Ginkgo species from the Jurassic of England, also known by the name, Ginkgoites huttonii, the genus, Ginkgoites, referring to a group of extinct members of the Ginkgoaceae, where G. huttonii was a broad-leaved, deciduous gymnosperm[28] bearing resemblance to the only living member of the Ginkgoaceae, Ginkgo biloba.[29]

Ginkgo huttonii is known largely by compression fossils of its leaves similar to other members of the Ginkgoites, the fossil leaves are simple, four-lobed, and have dense, radially disposed venation.[30][29] G. huttonii fossil seeds are frequently found as well as at least a few fossilized male catkins.[31] G. huttonii wood has yet to be described but it is likely the plant was similar to the extant, G. biloba, with wood akin to that of modern day conifers.[32]

G. huttonii is heavily represented in the Jurassic flora of Yorkshire, England - a flora which has been studied in depth since the 1800s.[32][33] The order Ginkgoales had a wide distribution throughout the northern hemisphere from the Lower Jurassic through the Cretaceous.[34]


Neuropteris is a common fossil in bituminous coal. Credit: Jstuby.{{Free media}}
Neuropteris is a fairly common fossil in bituminous coal. Credit: Gunnar Ries.{{free media}}

Neuropteris, a fern, leaf impressions and fossils occur in bituminous coal such as in the image on the right. These coal seams and strata are dated to the Carboniferous period.

Neuropteris is an extinct Pteridospermatophyta (seed fern) that existed in the Carboniferous period,[35] known only from fossils.

Major species include Neuropteris loschi.



The earliest fossil record of Camellia are the leaves of †C. abensis from the upper Eocene of Japan, †C. abchasica from the lower Oligocene of Bulgaria and †C. multiforma from the lower Oligocene of Washington, United States.[36]


  1. Plant classification before genomics may not agree with classification after application of genomics.

See also



  1. SemperBlotto (17 February 2007). "paleobotany". San Francisco, California: Wikimedia Foundation, Inc. Retrieved 2013-10-25. {{cite web}}: |author= has generic name (help)
  2. Hermann W. Pfefferkorn; Margaret C. Thomson (December 1982). "Changes in dominance patterns in Upper Carboniferous plant-fossil assemblages". Geology 10 (12): 641-4. doi:10.1130/0091-7613(1982)10<641:CIDPIU>2.0.CO;2. Retrieved 2012-08-08. 
  3. C. Downie (March 1967). "The geological history of the microplankton". Review of Palaeobotany and Palynology 1 (1-4): 269-81. doi:10.1016/0034-6667(67)90128-5. Retrieved 2012-08-08. 
  4. Duan Shuying (April 1987). "A comparison between the Upper Triassic floras of China and the Rhaeto‐Liassic floras of Europe and East Greenland". Lethaia 20 (2): 177-84. doi:10.1111/j.1502-3931.1987.tb02035.x. Retrieved 2012-08-08. 
  5. CS Gasser; K robinson-Beers (October 1993). "Pistil development". The Plant Cell 5 (10): 1231-9. PMID 12271024. Retrieved 2012-08-08. 
  6. Karl J. Niklas; Brace H. Tiffney; Andrew H. Knoll (16 June 1983). "Patterns in vascular land plant diversification". Nature 303 (5918): 614-6. doi:10.1038/303614a0. Retrieved 2016-02-07. 
  7. Kidston, R.; Lang, W. H. (1920). "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 (3): 643–680. doi:10.1017/S0080456800004506. 
  8. Rice, C. M.; Ashcroft, W. A.; Batten, D. J.; Boyce, A. J.; Caulfield, J. B. D.; Fallick, A. E.; Hole, M. J.; Jones, E. et al. (1995). "A Devonian auriferous hot spring system, Rhynie, Scotland". Journal of the Geological Society, London 152 (2): 229–250. doi:10.1144/gsjgs.152.2.0229. 
  9. 9.0 9.1 9.2 Hao, Shougang; Xue, Jinzhuang (2013). The Early Devonian Posongchong Flora of Yunnan - A Contribution to an Understanding of the Evolution and Early Diversification of Vascular Plants. Beijing: Science Press. pp. 244–245. ISBN 978-7-03-036616-0. 
  10. 10.0 10.1 Strullu‐Derrien, Christine; Wawrzyniak, Zuzanna; Goral, Tomasz; Kenrick, Paul (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 (1): 201–213. doi:10.1111/boj.12307. 
  11. Hetherington, Alexander J.; Dolan, Liam (2018). "Stepwise and independent origins of roots among land plants". Nature 561 (7722): 235–238. doi:10.1038/s41586-018-0445-z. PMID 30135586. 
  12. Smoot, E.L.; Jansen, R.K.; Taylor, T.N. (1981). "A Phylogenetic Analysis of the Land Plants: A Botanical Commentary". Taxon 30 (1): 65–67. doi:10.2307/1219392. 
  13. 13.0 13.1 Kerp, Hans; Wellman, Charles H.; Krings, Michael; Kearney, Patricia; Hass, Hagen (2013). "Reproductive organs and in situ spores of Asteroxylon mackiei Kidston & Lang, the most complex plant from the lower Devonian Rhynie chert". International Journal of Plant Sciences 174 (3): 293–308. doi:10.1086/668613. 
  14. Kenrick, Paul; Crane, Peter R. (1997). The Origin and Early Diversification of Land Plants: A Cladistic Study. Washington, D.C.: Smithsonian Institution Press. ISBN 978-1-56098-730-7. 
  15. Wilson, Jonathon P.; Fischer, Woodward W. (2011). "Hydraulics of Asteroxylon mackei, an early Devonian vascular plant, and the early evolution of water transport tissue in terrestrial plants". Geobiology 9 (2): 121–130. doi:10.1111/j.1472-4669.2010.00269.x. PMID 21244621. 
  16. Tomescu, Alexandru M.F. (2009). "Megaphylls, microphylls and the evolution of leaf development". Trends in Plant Science 14 (1): 5–12. doi:10.1016/j.tplants.2008.10.008. PMID 19070531. 
  17. 17.0 17.1 Wolfe, J.A.; Wehr, W.C. (1987). "Middle Eocene dicotyledonous plants from Republic, northeastern Washington". United States Geological Survey Bulletin 1597: 1–25. 
  18. 18.0 18.1 Crane, P. R.; Stockey, R. A. (1987). "Betula leaves and reproductive structures from the Middle Eocene of British Columbia, Canada". Canadian Journal of Botany 65 (12): 2490–2500. doi:10.1139/b87-338. 
  19. Smith, R.Y.; Basinger, J.F.; Greenwood, D.R. (2012). "Early Eocene plant diversity and dynamics in the Falkland flora, Okanagan Highlands, British Columbia, Canada". Palaeobiodiversity and Palaeoenvironments 92 (3): 309–328. doi:10.1007/s12549-011-0061-5. 
  20. Dillhoff, R.M.; Leopold, E.B.; Manchester, S.R. (2005). "The McAbee flora of British Columbia and its relations to the Early-Middle Eocene Okanagan Highlands flora of the Pacific Northwest". Canadian Journal of Earth Sciences 42 (2): 151–166. doi:10.1139/e04-084. 
  21. Lowe, A. J.; Greenwood, D. R.; West, C. K.; Galloway, J. M.; Sudermann, M.; Reichgelt, T. (2018). "Plant community ecology and climate on an upland volcanic landscape during the Early Eocene Climatic Optimum: McAbee Fossil Beds, British Columbia, Canada". Palaeogeography, Palaeoclimatology, Palaeoecology 511: 433-448. 
  22. Mathewes, R. W.; Greenwood, D. R.; Archibald, S. B. (2016). "Paleoenvironment of the Quilchena flora, British Columbia, during the Early Eocene Climatic Optimum". Canadian Journal of Earth Sciences 53 (6): 574–590. doi:10.1139/cjes-2015-0163. 
  23. Ludvigsen, Rolf (2011). Life in Stone: A Natural History of British Columbia's Fossils. UBC Press. p. 243. ISBN 978-0774841511. 
  24. Greenwood, D. R.; Archibald, S. B.; Mathewes, R. W.; Moss, P. T. (2005). "Fossil biotas from the Okanagan Highlands, southern British Columbia and northeastern Washington State: climates and ecosystems across an Eocene landscape". Canadian Journal of Earth Sciences 42 (2): 167–185. doi:10.1139/e04-100. 
  25. Pigg, K. B.; DeVore, M. L. (2007). "East meets West: the contrasting contributions of David L. Dilcher and Jack A. Wolfe to Eocene systematic paleobotany in North America". Courier Forschungsinstitut Senckenberg 258: 89. 
  26. Labandeira, C. C. (2002). "Paleobiology of middle Eocene plant-insect associations from the Pacific Northwest: a preliminary report". Rocky Mountain Geology 37 (1): 31–59. doi:10.2113/gsrocky.37.1.31. 
  27. 27.0 27.1 27.2 Olivier Béthoux, Jean Galtier, and André Nel (1 August 2004). "Earliest Evidence of Insect Endophytic Oviposition". Palaios 19 (4): 408-413. doi:10.1669/0883-1351(2004)019<0408:EEOIEO>2.0.CO;2. Retrieved 12 October 2021. 
  28. Ernest M. Gifford (1998). "Ginkgophyte". Britannica. Encyclopaedia Britannica, Inc.
  29. 29.0 29.1 Villar de Seoane, Liliana 1997. Comparative study between Ginkgoites tigrensis Archangelsky and Ginkgo biloba Linn, leaves. Palaeobotanist 46(3):1-12.
  30. Nosova, Natalya; Zhang, Jian-Wei (August 2011). "Revision of Ginkgoites obrustschewii (Steward) Seward (Ginkgoales) and the new material from the Jurassic of Northwestern China". Review of Palaeobotany and Palynology 166 (3–4): 286–294. doi:10.1016/j.revpalbo.2011.06.002. 
  31. Steur, Hans (March 1, 2020). "Ginkgo-like plants from Yorkshire". The Jurassic Flora of North Yorkshire. Retrieved June 10, 2020.
  32. 32.0 32.1 H.A. van Konijnenburg-van Cittert, Johanna (2008). "The Jurassic fossil plant record of the UK area". Proceedings of the Geologists' Association. 
  33. M. Slater, Sam; H. Wellman, CHarles; Romano, Michael; Vajda, Vivi (2018). "Dinosaur-plant interactions within a Middle Jurassic ecosystem-palynology of the Burniston Bay dinosaur footpring locality, Yorkshire, UK". Palaeobiodiversity and Palaeoenvironments 98: 139–151. doi:10.1007/s12549-017-0309-9. 
  34. "Ginkgoales: Fossil Record". Ginkgoales. UC Berkeley. 1997.
  35. "Neuropteris". Retrieved 2016-12-01.
  36. Journal of Plant Research, September 2016, Volume 129, Issue 5, pp 823–831, Camellia nanningensis sp. nov.: the earliest fossil wood record of the genus Camellia (Theaceae) from East Asia by Lu-Liang Huang, Jian-Hua Jin, Cheng Quan and Alexei A.

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