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Major Events in the Evolution of Plants


This page provides a high level review of some of the main threads of the evolution of embryophytes (“plants”), from their first appearance in the fossil record to the present day.

Keywords: evolution, plants


“Plants – with three phyla of bryophytes (mosses, liverworts, and hornworts) and the seven living, or extant, phyla of vascular plants – constitute a kingdom of photosynthetic organisms adapted for life on land…. All plants are multicellular and are composed of eukaryotic cells that contain vacuoles and are surrounded by cell walls that contain cellulose. Their principle mode of nutrition is photosynthesis, although a few plants have become heterotrophic. … Reproduction in plants is primarily sexual, with cycles of alternating haploid and diploid generations. … The unifying character of the Plantae is the presence of an embryo during the sporophytic [diploid] phase of the lifecycle” (Raven et al. 2005, p. 233, 235).

This definition excludes acritarchs (by definition, if something is a plant it cannot be an acritarch) and other “algae”. Discussion of these groups is often included in general paleobotanical texts (such as the excellent Taylor et al. 2008) but it is important to understand that these organisms are no more “plants” than an elephant is.

“Tracing early plant evolution is often focused on individual taxa, their taxonomy, appearances and stratigraphic ranges” Kraft et al. 2019, p. 144).


“Although the oldest macrofossil evidence of land plants is of middle Silurian (Wenlock) age, palynology indicates that they probably first appeared in late Ordovician times” (Cleal & Thomas 2009, p. 203).

“The search for the oldest land plant (embryophyte) reflects an effort to prove a substantial step of terrestrialization. It is of importance for molecular biologists and (neo)botanists to calibrate the molecular clock and establish a starting point for phylogenetic studies of early vascular plants (e.g. Clarke et al. 2011; Kenrick et al. 2012; Gerrienne et al. 2016). The first appearance of such plants, based on macroscopic evidence, can be provisionally considered to have occurred in the Late Ordovician in Poland (Salamon et al. 2018). However, the earliest unequivocal record based on Cooksonia (the oldest accepted macrofossil land plant; Fig. 1, left) from almost coeval sites in Ireland (Edwards et al. 1983) and Bohemia (Libertin et al. 2003, 2018a), points to the appearance of land plants in the Wenlock (middle Silurian). On the other hand, fossil spores and isolated sporangia indicate a much older appearance of embryophytes, probably as early as in the Middle Ordovician (Clarke et al. 2011; Gerrienne et al. 2016; Vavrdova 1984), even if a coeval macrofossil record is missing” (Kraft et al. 2019, p. 144).


The first major diversification of land plant life, such as rhyniophytes, zosterophylls, drepanophycaleans, lycophytes, trimerophytes, and (in the southern hemisphere) Baragwanathia (Fig. 1, right), began as early as Late Ludlow (Late Silurian) in Australia through Gedinnian (Early Devonian) on Laurentia.

“Throughout Silurian times, only small rhyniophytes (or rhyniophytoids) and lycophytes are known…. Remarkably, even in these early phases of their evolution, plants had an almost worldwide distribution with records from North and South America, Europe, Africa, central Asia, China and Australia, presumably reflecting the wide dispersal potential of their spores and the lack of any competition. The global composition of these floras also appears to be fairly uniform, although this may in part be due to the problems of differentiating biological species within the plexus of these morphologically very simple plants” (Cleal & Thomas 2009, p. 203).

Fig. 1: Cooksonia cf. hemisphaerica, left, and Baragwanathia brevifolia (Kraft et al. 2019, fig. 4).


Cooksonia, the early or even primordial land plant, apparently played a key role: it has been described from a number of regions, ranges widely from the Silurian to the early Devonian, and is represented by several species such as C. pertoni, C. paranensis, C. banksii, and doubtfully C. cambrensis, C. hemisphaerica (Gonez and Gerrienne 2010a) with many other specimens having been described in open nomenclature. Cooksonia also occupies a special position in the colonization of terrestrial habitats, which is a major aspect of early plant research based on broad studies of associations, their successions and distributions (e.g. Edwards & Wellman 2001; Kenrick et al. 2012). This complex paleoecological approach has attracted considerable attention over the past two decades” Kraft et al. 2019, p. 144).


“From its primitive rhyniophyte and lycophyte precursors, land vegetation rapidly diversified during Devonian times. By the end of the period, all of the major divisions of vascular plants except the flowering plants had appeared, with the first record of ferns (or at least their pre-fern ancestors) being in Middle Devonian times, and of sphenophytes and gymnosperms in Late Devonian times…” (Cleal & Thomas 2009, p. 203).

“The Devonian was pivotal in the evolutionary history of land plants: a diverse range of plant architectures evolved and plants radiated into a remarkable array of ecological niches (Gensel & Andrews 1984, Gerrienne et al. 2004, Stein et al. 2012)” (Xu et al. 2017, p. 524).

“The Devonian Period also saw the development of several structures that made the vascular plants better adapted to life on land…. Secondary wood is the most widely adopted means by which plants can significantly increase their stature and it is first seen in the fossil record in the Middle Devonian Series. The resulting trees (and therefore presumably forests) must have had a dramatic impact on the Devonian landscape. Photosynthetic efficiency was also enhanced by the development of planated leaves in Devonian times” (Cleal & Thomas 2009, p. 203).

Numerous plant taxa have been reported from Devonian deposits around the world and phytogeographical provincialism began to develop in the global flora (see Raymond et al. 2006). However, plants with simple form and diminutive size, i.e., with axis widths less than 2 mm, from the Early–Middle Devonian, have been paid little attention (see Cai & Wang 1995, Wang & Berry 2003, Wang et al. 2004, 2007). Those basal euphyllophytes have quite distinct morphological characters in their branching patterns and sporangial attachment, and played key roles in the evolution of early land plants (Stewart & Rothwell 1993, Kenrick & Crane 1997)” (Xu et al. 2017, p. 524; Fig. 2).

The lycopsids apparently arose in the Silurian (Kotyk et al. 2002) but their fossil record is better known from the Devonian onwards.

The sphenophytes appear to have arisen in the Devonian.


Fig. 2: The Early Devonian plant, Pauthecophyton; composite from Xu et al. 2019, fig. 3A and D.


“In Pennsylvanian [= Late Carboniferous] times, there was a dramatic change in vegetation. In low paleolatitudes, the very first tropical rain forests appeared: the so-called ‘Coal Forests’ of europe, North America and China (known as the Amerosinian Palaeokingdom...), so named because of the vast reserves of coal-forming peat that were laid down by them…. These forests were dominated by the giant lycophytes…, which were perfectly adapted to the wetland habitats that existed over much of the then tropics” (Cleal & Thomas 2009, p. 205-206).


Early Cretaceous vegetation was broadly similar to that of Late Jurassic times, both in distribution and general composition. Low paleolatitudes at that time were arid, having desert and sub-desert conditions, and here the floras were dominated by cheirolepidiacean conifers and matoniacean ferns. Northern mid paleolatitude floras were more diverse, including ferns, bennettitaleans, cycads, conifers and some ginkgos. At higher latitudes, diversity declined again, the floras dominated by leptostrobaleans and ginkgos. Southern mid paleolatitudes were dominated by bennettitaleans and cheirolepidiacean conifers. (Adapted from Cleal & Thomas 2009.)

The most striking event in the evolution of plants during the Cretaceous was certainly the enormous radiation of angiosperms.

The angiosperms (flowering plants) are the most diverse group of land plants living today, comprising some 270,000 described species, placed in about 380 families and 83 orders (Mayr 2001, p. 64).

The group may have evolved from either the Gnetales or possibly the Bennettitales (Willis & McElwain 2002, p. 184).

The enormous radiation of this group has largely occurred since the mid-Cretaceous, coevolving with a similar radiation of insects. However, the angiosperms must have arisen earlier: perhaps as early as the Triassic or even the late Carboniferous (Qui et al. 1999). Evidence supporting the earliest date estimates is mainly provided by calibrated genetic divergence studies, though fossil angiosperm-like pollen and leaves have been found dating back to the late Triassic. (Several form-species of Crinopolles-type pollen possessing a tectate wall have been described, dating to perhaps 220 Ma. The oldest leaves are somewhat younger, perhaps 210 Ma, and include the problematic taxa Furcula and Sanmiguelia.)

That being said, however, current orthodoxy is that the first true angiosperms evolved in the Early Cretaceous, probably in the Valanginian (~140 to ~133 Ma) or Hauterivian (~133 to ~129 Ma) ages. The oldest unequivocal angiosperm pollen grains first appear in the fossil record “during the Valanginian-Hauterivian; they spread out of the tropics in the Aptian and Albian [~125.0 to 100.5 Ma], and radiated in the Late Cretaceous” (Harris & Arens 2016, p. 640).

Some of the earliest “body” fossils are “small plants, possibly rooted aquatics or wetland herbs” (Wing 2004, p. 90).

Genetic evidence (Zanis et al. 2002) strongly suggests that the most ‘primitive’ (basal) living angiosperm is a little known shrub called Amborella trichopoda; a small shrub with tiny greenish-yellow flowers and red fruit, native to the South Pacific island of New Caledonia. The Nymphaeales (waterlilies and their relatives) are also contenders for the distinction of being the most basal living angiosperms.

Perhaps the most basal fossil group yet to be well-delineated within the angiosperm clade is the Archaefructaceae, a family of herbaceous aquatic plants recovered from the Lower Cretaceous or possibly uppermost Jurassic Yixian Formation of western Liaoning, China. These plants had reproductive axes that lacked petals and sepals, and bore stamens in pairs below conduplicate (sharply folded together lengthwise) carpels. One combined morphological and molecular “total evidence” analysis places the Archaefructaceae as a sister group to all extant angiosperms, including Amborella and the Nymphaeales (Sun et al. 2002, p. 900).


Fig. 3: Amborella trichopoda. [Photograph by Tim Stephens, courtesy of the Arboretum of the University of California, Santa Cruz.]


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Cleal, C.J.; Thomas, B.A. 2009: An Introduction to Plant Fossils. Cambridge University Press: 1-248.

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Harris, E.B.; Arens, N.C. 2016: A mid-Cretaceous angiosperm-dominated macroflora from the Cedar Mountain Formation of Utah, USA. Journal of Paleontology 90 (4): 640-662.

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Kenrick, P.; Wellman, C.H.; Schneider, H.; Edgecombe, G.D. 2012: A timeline for terrestrialization: consequences for the carbon cycle in the Palaeozoic. Philos. Trans. R. Soc. Lond. B 367 (1588): 519-536.

Kotyk, M.E.; Basinger, J.F.; Gensel, P.G.; de Freitas, T.A. 2002: Morphologically complex plant macrofossils from the late Silurian of Arctic Canada. American Journal of Botany 89 (6): 1004-1013.

Kraft, P.; Psenicka, J.; Sakala, J.; Frýda, J. 2019: Initial plant diversification and dispersal event in upper Silurian of the Prague Basin. Palaeogeography, Palaeoclimatology, Palaeoecology 514 (2019) 144-155.

Libertín, M.; Kvacek, J.; Bek, J.; Žárský, V.; Štorch, P. 2018a: Sporophytes of polysporangiate land plants from the early Silurian period may have been photosynthetically autonomous. Nat. Plants 4: 269-271.

Libertín, M.; Labuta, R.; Dašková, J. 2003: The oldest vascular plants from the Bohemian Massif. Zprávy o geologických výzkumech v roce 2002. (In Czech, English summary.) 127.

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Qui, Y.-L.; Lee, J.; Bernasconi-Quadroni, F.; Soltis, D.E.; Soltis, P.; Zanis, M.; Zimmer, E.A.; Chen, Z.; Savolainen, V.; Chase, M.W. 1999: The Earliest Angiosperms. Nature 402: 404-407.

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Salamon, M.; Gerrienne, P.; Steemans, P.; Gorzelak, P.; Filipiak, P.; Le Hérissé, A.; Paris, F.; Cascales-Miñana, B.; Brachaniec, T.; Misz-Kennan, M.; Niedzwiedzki, R.; Trela, W. 2018: Putative Late Ordovician land plants. New Phytol. 218: 1305-1309.

Stein, W.E.; Berry, C.M.; Hernick, L.V.A.; Mannolini, F. 2012: Surprisingly complex community discovered in the mid-Devonian fossil forest at Gilboa. Nature 483: 78-81.

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Sun, G.; Ji, Q.; Dilcher, D.L.; Zheng, S.; Nixon, K.C.; Wang, X. 2002: Archaefructaceae, a new basal angiosperm family. Science 296: 899-904.

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Wang, Y., Xu, H., Fu, Q.; Tang, P. 2004: A new diminutive plant from the Hujiersite Formation (late Middle Devonian) of North Xinjiang, China. Acta Palaeontologica Sinica 43: 461-471.

Wang, Y.; Berry, C.M. 2003: A reconsideration of Dimeripteris cornuta Schweitzer and Cai, a diminutive fossil plant from the Middle Devonian of Yunnan, China. Geobios 36: 437-446.

Wang, Y.; Berry, C.M.; Hao, S.; Xu, H.; Fu, Q. 2007: The Xichong flora of Yunnan, China: diversity in late Mid Devonian plant assemblages. Geological Journal 42: 339-350.

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Xu, H.; Wang, Y.; Tang, P.; Wang, Y. 2017: A new diminutive euphyllophyte from the Middle Devonian of West Junggar, Xinjiang, China and its evolutionary implications. Alcerhinga 41 (4): 524-531.

Zanis, M.J.; Soltis, D.E.; Soltis, P.S.; Mathews, S.; Donoghue, M.J. 2002: The root of the angiosperms revisited. Proceedings of the National Academy of Sciences of the USA 99: 6848-6853.

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