A palaeobotanical perspective on the Permian extinction.


Leaf bank of Glossopteris leaves (Adapted from Mcloughlin, 2012)

Leaf bank of Glossopteris leaves (Adapted from Mcloughlin, 2012)

The fossil record indicates that more than 95% of all species that ever lived are now extinct. Occasionally, extinction events reach a global scale with many species of all ecological types dying out in a near geological instant. These are mass extinctions. They were originally identified in the marine fossil record and have been interpreted as a result of catastrophic events or major environmental changes that occurred too rapidly for organisms to adapt. Mass extinctions are probably due to a set of different possible causes like basaltic super-eruptions, impacts of asteroids, global climate changes, or continental drift. A central question in the understanding of mass extinctions is whether the extinction was a sudden or gradual event. This question may be addressed by examining the pattern of last occurrences of fossil species in a stratigraphic section.

Jack Sepkoski and David M. Raup identified five major extinction events in Earth’s history: at the end of the Ordovician period, Late Devonian, End Permian, End Triassic and the End Cretaceous. The most recently identified mass extinction occurred during the Middle Permian, about  262 million years ago, and it was first recognised in the marine realm as a turnover among foraminifera, with fusulinaceans among the principal casualties.

Sin título

Total diversity patterns of continental diversity (solid line) and marine diversity (dotted line) at the family level. Arrows indicate the mass extinction events. (From Cascales-Miñana and Cleal 2015)

Extinction dynamics in the marine and terrestrial biotas followed different trajectories, and only the Permo-Triassic event coincided with a clear and abrupt diminution of both realms. Moreover, analysis of the paleobotanical record has suggested that plants may have suffered an additional extinction event, that is not reflected significantly in the marine realm, at the Carboniferous–Permian boundary. Evidence also suggests that  terrestrial environments suffered a single global pulse of extinction in the latest Permian, affecting both the fauna and flora (Cascales-Miñana and Cleal 2015).

During the end-Permian Event, the woody gymnosperm vegetation (cordaitaleans and glossopterids) were replaced by spore-producing plants (mainly lycophytes) before the typical Mesozoic woody vegetation evolved. The palynological record suggests that wooded terrestrial ecosystems took four to five million years to reform stable ecosystems, while spore-producing lycopsids had an important ecological role in the post-extinction interval. A key factor for plant resilience is the time-scale: if the duration of the ecological disruption did not exceed that of the viability of seeds and spores, those plant taxa have the potential to recover (Traverse, 1988).



Borja Cascales-Miñana, José B. Diez, Philippe Gerrienne & Christopher J.Cleal (2015): A palaeobotanical perspective on the great end-Permian biotic crisis, HistoricalBiology, DOI: 10.1080/08912963.2015.1103237

Aberhan M. 2014. Mass extinctions: ecological diversity maintained. NatGeosci. 7:171–172.

Cascales-Miñana B, Cleal CJ. 2014. The plant fossil record reflects just two great extinction events. Terra Nova. 26(3):195–200.

Darwin and the flowering plant evolution in South America.


Retimonocolpites sp. (Adapted from Llorens and Loinaze, 2015)

Charles Darwin’s fascination and frustration with the evolutionary events associated with the origin and early radiation of flowering plants are legendary. In a letter to Oswald Heer, a famous Swiss botanist, and paleontologist, Darwin wrote: “the sudden appearance of so many Dicotyledons in the Upper Chalk appears to me a most perplexing phenomenon to all who believe in any form of evolution, especially to those who believe in extremely gradual evolution, to which view I know that you are strongly opposed”. Heer discussed about the early angiosperm fossil record with Darwin, in a letter dated 1 March 1875: “if we say that the Dicotyledons begin with the Upper Cretaceous, we must still concede that this section of the vegetable kingdom, which forms the bulk of modern vegetation, appears relatively late and that, in geological terms, it underwent a substantial transformation within a brief period of time.” 

Darwin’s defense of a gradualist perspective led him to suggest that prior to the Cretaceous record of flowering plants, angiosperms had slowly evolved and diversified on a remote landmass. On 22 July 1879, in a letter to Joseph Dalton Hooker, Darwin refers to the early evolution of flowering plants as an “abominable mystery”. Nearing the end of his life, he wrote to Hooker another letter about a lost fossil record in the earliest phases of angiosperm diversification:  “Nothing is more extraordinary in the history of the Vegetable Kingdom, as it seems to me, than the apparently very sudden or abrupt development of the higher plants. I have sometimes speculated whether there did not exist somewhere during long ages an extremely isolated continent, perhaps near the South Pole.”

Letter from Charles Darwin to Joseph Dalton Hooker, written 22 July 1879 (provenance: Cambridge University Library DAR 95: 485–488)

Letter from Charles Darwin to Joseph Dalton Hooker, written 22 July 1879 (provenance: Cambridge University Library DAR 95: 485–488)

While the oldest records of the different groups of angiosperms are still in discussion, the outcrops of the Baquero Group, located in Argentinean Patagonia, contain one of the richest and most diverse Early Cretaceous floras in the Southern Hemisphere. The unit comprises three formations: Anfiteatro de Tico, Bajo Tigre and Punta del Barco. The first reports of angiosperm remains for the Anfiteatro de Tico Formation were made in 1967. The dominant types are Clavatipollenites, and Retimonocolpites.

Pollen grains  could enter into the fossil record by falling directly into swamps or lakes, or being washed into them or into the rivers and seas. The ones which are not buried in reducing sediments will tend to become oxidized and be destroyed. They reflects the ecology of their parent plants and their habitats and provide a continuous record of their evolutionary history. Gymnosperms pollen often is saccate (grains with two or three air sacs attached to the central body), while Angiosperm pollen shows more variation and covers a multitude of combinations of features: they could be  in groups of four (tetrads),  in pairs (dyads),  or single (monads). The individual grains can be inaperturate, or have one or more pores, or slit-like apertures or colpi (monocolpate, tricolpate).


Clavatipollenites sp. SEM (Adapted from Archangelsky 2013)

Clavatipollenites pollen grains are interpreted as related to the modern family Chloranthaceae. The genus was established by Couper for dispersed monosulcate pollen grains recovered from the Early Cretaceous of Britain. Currently, the genus has a very broad definition. The genus Retimonocolpites include elongated to subcircular semitectate, columellate and microreticulate pollen grains with well defined monocolpate aperture (Llorens and Loinaze, 2015). The new species Jusinghipollisticoensis sp. nov. represents one of the oldest records of trichotomosulcate, and extends the geographical distribution of Early Cretaceous trichotomosulcate pollen grains to southern South America.

The data also indicates strong similarities between the Baquero Group assemblages and other coeval units from Argentina, Australia and United States.


M. Llorens, V.S. Perez Loinaze, Late Aptian angiosperm pollen grains from Patagonia: Earliest steps in flowering plant evolution at middle latitudes in southern South America, Cretaceous Research 57 (2016) 66-78

Archangelsky, S.,et al. (2009). Early angiosperm diversification: evidence from southern South America. Cretaceous Research, 30, 1073-1082.

Doyle, J. A., & Endress, P. K. (2014). Integrating Early Cretaceous fossils into the phylogeny of living Angiosperms: ANITA. Lines and relatives of Chloranthaceae. International Journal of Plant Sciences, 175, 555-560.