page_title: Evolutionary history of plants
section_title: Flowers
hierarchical_section_title: Evolutionary history of plants / Evolution of plant morphology / Flowers
caption_attribution_description: Bennettiales from the Sedgwick Museum's collection. Note the "cone" in the top-right of the image, and leaf traces. Width is ~15cm.
context_page_description: The evolution of plants has resulted in a wide range of complexity, from the earliest algal mats, through multicellular marine and freshwater green algae, terrestrial bryophytes, lycopods and ferns, to the complex gymnosperms and angiosperms of today. While many of the earliest groups continue to thrive, as exemplified by red and green algae in marine environments, more recently derived groups have displaced previously ecologically dominant ones, e.g. the ascendance of flowering plants over gymnosperms in terrestrial environments.
  There is evidence that cyanobacteria and multicellular photosynthetic eukaryotes lived in freshwater communities on land as early as 1 billion years ago, and that communities of complex, multicellular photosynthesizing organisms existed on land in the late Precambrian, around 850 million years ago.
  Evidence of the emergence of embryophyte land plants first occurs in the mid-Ordovician, and by the middle of the Devonian, many of the features recognised in land plants today were present, including roots and leaves.
context_section_description: Flowers are modified leaves possessed only by the angiosperms, which are relatively late to appear in the fossil record. The group originated and diversified during the Early Cretaceous and became ecologically significant thereafter. Flower-like structures first appear in the fossil records some ~130 mya, in the Cretaceous. However, in 2018, scientists reported the finding of a fossil flower from about 180 million years ago, 50 million years earlier than thought earlier. The interpretation has been however highly disputed. 
  Colorful and/or pungent structures surround the cones of plants such as cycads and Gnetales, making a strict definition of the term "flower" elusive.
  The main function of a flower is reproduction, which, before the evolution of the flower and angiosperms, was the job of microsporophylls and megasporophylls. A flower can be considered a powerful evolutionary innovation, because its presence allowed the plant world to access new means and mechanisms for reproduction.
  The flowering plants have long been assumed to have evolved from within the gymnosperms; according to the traditional morphological view, they are closely allied to the Gnetales. However, as noted above, recent molecular evidence is at odds with this hypothesis, and further suggests that Gnetales are more closely related to some gymnosperm groups than angiosperms, and that extant gymnosperms form a distinct clade to the angiosperms, the two clades diverging some 300 million years ago.
  The relationship of stem groups to the angiosperms is important in determining the evolution of flowers. Stem groups provide an insight into the state of earlier "forks" on the path to the current state. Convergence increases the risk of misidentifying stem groups. Since the protection of the megagametophyte is evolutionarily desirable, probably many separate groups evolved protective encasements independently. In flowers, this protection takes the form of a carpel, evolved from a leaf and recruited into a protective role, shielding the ovules. These ovules are further protected by a double-walled integument.
  Penetration of these protective layers needs something more than a free-floating microgametophyte. Angiosperms have pollen grains comprising just three cells. One cell is responsible for drilling down through the integuments, and creating a conduit for the two sperm cells to flow down. The megagametophyte has just seven cells; of these, one fuses with a sperm cell, forming the nucleus of the egg itself, and another joins with the other sperm, and dedicates itself to forming a nutrient-rich endosperm. The other cells take auxiliary roles. This process of "double fertilisation" is unique and common to all angiosperms.
  In the fossil record, there are three intriguing groups which bore flower-like structures. The first is the Permian pteridosperm Glossopteris, which already bore recurved leaves resembling carpels. The Mesozoic Caytonia is more flower-like still, with enclosed ovules – but only a single integument. Further, details of their pollen and stamens set them apart from true flowering plants.
  The Bennettitales bore remarkably flower-like organs, protected by whorls of bracts which may have played a similar role to the petals and sepals of true flowers; however, these flower-like structures evolved independently, as the Bennettitales are more closely related to cycads and ginkgos than to the angiosperms.
  However, no true flowers are found in any groups save those extant today. Most morphological and molecular analyses place Amborella, the nymphaeales and Austrobaileyaceae in a basal clade called "ANA". This clade appear to have diverged in the early Cretaceous, around 130 million years ago – around the same time as the earliest fossil angiosperm, and just after the first angiosperm-like pollen, 136 million years ago.
  The magnoliids diverged soon after, and a rapid radiation had produced eudicots and monocots by 125 million years ago. By the end of the Cretaceous 66 million years ago, over 50% of today's angiosperm orders had evolved, and the clade acco
caption: The inflorescences of the Bennettitales are strikingly similar to flowers
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