Did Angiosperms Appear in the Early Permian, 165 Million Years Before the Cretaceous? Paleobotanist Xin Wang Thinks They Did

     I will first note that researcher Xin Wang’s work has been a bit controversial. There is certainly no widespread evidence that angiosperms, plants with seeds enclosed within an ovary, which develop into a fruit, existed before the Cretaceous period. Wang has done other research suggesting that angiosperms existed in the Jurassic period, but extending this to the Early Permian is indeed a big stretch. The Wikipedia entry on ‘flowering plants’ notes that Wang’s work has not been widely accepted:

“…angiosperms appeared suddenly and rapidly diversified during the Early Cretaceous (beginning ~130 mya), much later than other major plant groups. Claimed records of flowering plants prior to this are not widely accepted, as all supposed pre-Cretaceous "flowers" can be explained through being misidentifications of other seed plants. Furthermore, almost all of these controversial fossils are described in papers co-authored by the researcher Xin Wang, such as the particularly debated Nanjinganthus.”

     The authors, including Wang, of a new paper in the journal Plant Biosystems suggest that there is indeed evidence of angiosperms dating back to the Early Permian and cite some examples. They think that it is simply that the structures of angiosperms were not recognized as such:

“The early age and the lack of conduplicate carpels in these fossils deter many from recognizing them as angiosperms. Such a hesitation is also rooted in the long held “no-angiosperms-until-Cretaceous” belief.”

“Arguably, Permian angiosperms cannot win full acceptance until the origin of apocarpous gynoecium composed of multiple carpels is critically re-analysed in the light of fossil evidence. Therefore, carpel-like fossils related angiosperms are badly needed in palaeobotany. Here, we report the first carpel-like structures, Permocarpelloides gen. nov., from the Lower Permian (Asselian Shanxi Formation) of Shanxi, China, to complement the former evidence of Permian angiosperms. The Permian age of this fossil pins down a 295-Ma-long history for angiosperms. The unique morphology of Permocarpelloides supports the existence of an apocarpous gynoecium in the Permian, and its differences from typical magnolialean carpels shed new light upon carpel origin and evolution.”

     The specimens showing the ovules were found in the Shanxi region of China, in the Shanxi Formation, which consists of coal beds, shale, siltstone, and limestone. The specimens were found in a siltstone, aged to the Asselian stage of the earliest Permian, dated approximately 295–298.9 million years ago. Incidentally, some of the rocks around where I live, likely including those on my own property, are of this age. The specimens are of a plant they named Permocarpelloides shuozhouensis.  

    It is thought that the early gymnosperms: conifers and seeded cycads, emerged 300 million years ago when the climate became much drier, and seasonality became more pronounced. That is also when insects began interacting with them, although there is no direct evidence in the fossil record for gymnosperms being pollinated by insects until the Jurassic. It is thought that when angiosperms developed in the Cretaceous, they were pollinated by insects such as beetles. If Wang is correct, then those insects would be needed for pollination. While there was a significant diversification of insects beginning in the Early Permian, there is no evidence of pollination of gymnosperms, nor of angiosperms if they even existed. There were beetles around then, and some flying insects as well, but modern pollinators like bees did not emerge until the Early Cretaceous.   

     The paragraph below explains the reasoning behind seeing the specimen as an angiosperm and why it differs from the seeds of conifers and seed ferns present at the time.   

“A caveat for the above conclusion is that many Conifers may enclose their seeds after pollination (Tomlinson and Takaso 2002) and the same is also known for some fossil taxa, for example, Caytonia Thomas (Harris 1933). However, in our opinion these exceptions do not undermine our interpretation of Permocarpelloides as angiosperm carpels/fruits. In Conifers, almost all the enclosed seeds occur on the adaxial side of scales and bracts, in contrast to the seeds/ovules positioned between two lines along the carpel margins in Permocarpelloides. In the case of Caytonia, the ovules/seeds are on the adaxial side of the cupule wall and the general morphology of cupule and basal opening of the cupule have no equivalents in Permocarpelloides. Another ghost alternative is seed ferns, which originally designated plants bearing seeds and fern-like foliage. We currently have no clue about the foliage of Permocarpelloides thus comparison based on this trait is impossible. Yet, there are no known seed ferns bearing reproductive organs anyhow similar to Permocarpelloides, in spite of the great diversity documented in seed ferns (Taylor and Taylor 2009; Taylor et al. 1994, 2006, 2007; Doyle 2006). These differences are, in our opinion, deep enough to outweigh doubt over interpretation of Permocarpelloides as an early angiosperm.”

     They also write about other possible interpretations of the new species, such as designating it to a new group of gymnosperms. Below, they explain that and suggest that it could just be new evidence for the suspected blurring of the difference between gymnosperms and early angiosperms in the fossil record.

“Even if Permocarpelloides were finally placed in a new group of gymnosperms, this treatment would not reduce the evolutionary significance: Permocarpelloides would then be the first gymnosperm displaying angio-ovuly and angiospermy, suggestive of the blurry boundary between gymnosperms and angiosperms that has long be theoretically assumed but never confirmed by fossil evidence so far. Independent of being an angiosperm or a gymnosperm, Permocarpelloides is a peerless plant that sheds novel light on the evolution of seed plants.”

     Below, they suggest other possible evidence for Early Permian angiosperms:

“The possible ecological relationship between angiosperms and Permian insects (Khramov et al. 2023; Khramov et al. 2020; Peña-Kairath et al. 2023) deserves further investigations. Furthermore, gene and phylogenomic studies have converged to a pre-Cretaceous origin of angiosperms (Martin et al. 1989; Wolfe et al. 1989; Becker et al. 2000; Shi and de Peer 2026; Li et al. 2019; Zuntini et al. 2024; Ma et al. 2025; Ramshaw et al. 1972). Agreement among these independent lines of evidence, including fossil evidence, insects, and molecular dating, unveil a previously hidden fact: angiosperms have a long history dated back to the Early Permian.”

     Since I have rocks of this exact age quite nearby, I plan to investigate further. There are many different plant and marine fossils nearby, some in the slightly older Monongahela Group. That group contains world-class petrified wood and coal balls, which have well-preserved plant fossils. Coal balls are concretions that form in coal beds. The permineralization is not quartz or opal as in the case of petrified wood, but calcium and magnesium minerals. Back in the late 80s when I was an undergrad, I worked for a paleobotanist. One thing I did was make acetate peels of these coal balls in order to get full 3D representations of plant fossils. I have identified a limestone down in the creek below my house that appears to be fossiliferous, and I plan to collect more specimens when access is better in the late fall and winter. Here on the ridgetop is likely where the earliest Permian outcrops are, and most rocks preserved on the ridge are sandstones and siltstones. There are similarities and differences between the site here and the site in China. Both were positioned near the equator at the time of deposition. However, the U.S. area was part of the mass of continents known as Pangaea, and the Shanxi region of China was a small isolated island. As my historical geology teacher taught me: "isolation yields speciation." Thus, the species found there may never have occurred on Pangaea. The beginning of the Permian marks when ice sheets formed at the poles, especially the Southern polar area, resulting in a global drop in sea level. This followed the Alleghenian uplift in Appalachia. It represents the last time this area was offshore or nearshore, and the Appalachian Mountains have been eroding away ever since. 

     Below is the paleogeography at the Permian-Pennsylvanian boundary. Both the Euroamerican and Cathaysian plates are along the equator, but the Cathaysian plate is isolated. Thus, I would say it is a long shot to find similar species here. 

     However, the ice sheets led to more land being exposed, so land corridors from the Cathaysian plate to the Euramerican plate occurred soon after the advent of the Permian, as shown below.

References:

 

Permian carpel-like organs and their implications on origin of angiosperms. Xin Wang, Qiang Fu, Weijia Huang & Jie Sun. Plant Biosystems. Volume 160, article number 257 (2026). August 25, 2026. Permian carpel-like organs and their implications on origin of angiosperms | Plant Biosystems | Springer Nature Link

Flowering plant. Wikipedia. Flowering plant - Wikipedia

Permian. Wikipedia. Permian - Wikipedia