The Cretaceous Terrestrial Revolution (abbreviated KTR), also known as the Angiosperm Terrestrial Revolution (ATR) by authors who consider it to have lasted into the Palaeogene, [1] describes the intense floral diversification of flowering plants (angiosperms) and the coevolution of pollinating insects, as well as the subsequent faunal radiation of frugivorous, nectarivorous and insectivorous avians, mammals, lissamphibians, squamate reptiles and web-spinning spiders during the Middle to Late Cretaceous, from around 125 Mya to 80 Mya. [2] Alternatively, according to Michael Benton, the ATR is proposed to have lasted from 100 Ma, when the first highly diverse angiosperm leaf floras are known, to 50 Ma, during the Early Eocene Climatic Optimum, by which point most crown lineages of angiosperms had evolved. [1]
Molecular clock analyses of angiosperm evolution suggest that crown group angiosperms may have diverged up to 100 million years before the start of the KTR, although this is possibly due to artefacts of the inabilities of molecular clock estimates to account for explosive accelerations in evolution that may have caused the extremely fast diversification of angiosperms shortly after their first appearance in the fossil record. [3]
The KTR was enabled by the dispersed positions of the continents and the formation of new oceans during the Cretaceous in the aftermath of Pangaea's breakup in the preceding Jurassic period, which enhanced the hydrological cycle and promoted the expansion of temperate climatic zones, fuelling radiations of angiosperms. [4] Among mammals, enhanced tectonic activity generated diversity increases by increasing montane habitats, which promote increased diversity in hot climates. [5]
Another cause of the explosive angiosperm diversification was the evolution of leaf vein densities greater than 2.5–5 mm/mm2, when the leaf interior transport path length of water became shorter than the leaf interior transport path length of CO2. This enabled greater utilisation of CO2 and gave an evolutionary advantage to flowering plants over conifers because they could sequester more CO2 for the same amount of water. [6] The much greater capacity of angiosperms for assimilating CO2 sharply increased global bioproductivity. [7]
The drying of many terrestrial ecosystems during the Middle Cretaceous Hothouse (MKH) benefitted angiosperms, which were able to survive hot and dry environments, and the increased fire activity helped to enhance diversification of angiosperms. [8] Angiosperms enabled more frequent fires than gymnosperms, and they also recovered more quickly from fires than gymnosperms did. This created a feedback loop that advantaged angiosperms over gymnosperms during the Cretaceous. [9]
Although angiosperm diversity drastically grew over the Cretaceous, this did not necessarily translate to ecological dominance, which they only achieved in the Early Cenozoic. [10]
Angiosperms responded to increasing coevolution with frugivores by enlarging the sizes of their fruits, which peaked during the Early Eocene. [11]
Before Lloyd et al.'s 2008 paper described the KTR, it had been widely accepted in paleontology that new families of dinosaurs evolved during the Middle to Late Cretaceous, including the euhadrosaurs, neoceratopsians, ankylosaurids, pachycephalosaurs, carcharodontosaurines, troodontids, dromaeosaurs and ornithomimosaurs. However, the authors of the paper have suggested that the apparent "new diversification" of dinosaurs during this time is due to sampling biases in the fossil record, and better preserved fossils in Cretaceous age sediments than in earlier Triassic or Jurassic sediments. [2] However, later studies still suggest the possibility that the KTR caused a rise in dinosaur diversity. [12] Dinosaurs contributed little to angiosperm diversification, which was instead mainly driven by coevolution with other animals, such as insects and herbivorous mammals. [13] It has been suggested that some pterosaurs may have been seed dispersers symbiotically linked to angiosperms. [14] A comprehensive molecular study of evolution of mammals at the taxonomic level of family also showed important diversification during the KTR. [15] Mammals have been found to have decreased in disparity during the KTR. [16]
Insect diversity overall appears to have been minimally affected by the KTR, as molecular evidence shows that the increase in diversity of pollinating insects was asynchronous with the KTR. [17] However, Early Cretaceous angiosperms were short in stature and would have been heavily reliant on insect pollination, [10] and fossil remains of early angiosperms suggest such a dependence on zoophilous pollination. [18] Genetic evidence indicates a major radiation of phasmatodeans occurred during the KTR, likely in response to a coeval radiation of enantiornitheans and other visual predators. [19] Ants likewise underwent massive increase in diversity as part of the KTR. [20] Similarly, bee pollinator diversification strongly correlates with angiosperm flower appearance and specialization during the same era. [21] Flies, already successful pollinators before the rise of angiosperms, [22] quickly adapted to the new hosts. [23] Beetles became pollinators of angiosperms by the earliest part of the Late Cretaceous. [24] [25] Lepidopterans radiated during the KTR, though the angiosperm radiation is insufficient in and of itself to completely account for their diversification. [26] Among one lineage of sawflies, there was a change in preferred host plants amidst the biotic reorganisation of the KTR. [27] Not all insects were advantaged by this diversification and rearrangement of ecosystems; long-proboscid insects that were mainstays of gymnosperm-dominated ecosystems earlier in the Mesozoic underwent a major decline. [28] Late-surviving eoblattodeans evolved long, slim bodies with long external ovipositors in response to the angiosperm radiation, but this proved to be an evolutionary dead end in the long run and the group went extinct. [29] The so-called "golden age" of neuropterans during the Middle Mesozoic, when gymnosperms dominated the flora, ended with the KTR and its reshaping of the terrestrial environment. [28]
The KTR may have supercharged the contemporary Mesozoic Marine Revolution (MMR) by enhancing weathering and erosion, accelerating the flow of limiting nutrients into the world’s oceans. [30]
For nearly the entirety of Earth's history, including most of the Phanerozoic eon, marine species diversity exceeded terrestrial species diversity, a pattern which was reversed during the Middle Cretaceous as a result of the KTR in what has been termed a biological "great divergence", named after the historical Great Divergence. [31]
The Cretaceous is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the ninth and longest geological period of the entire Phanerozoic. The name is derived from the Latin creta, 'chalk', which is abundant in the latter half of the period. It is usually abbreviated K, for its German translation Kreide.
The Cenozoic is Earth's current geological era, representing the last 66 million years of Earth's history. It is characterized by the dominance of mammals, birds, conifers, and angiosperms. It is the latest of three geological eras of the Phanerozoic Eon, preceded by the Mesozoic and Paleozoic. The Cenozoic started with the Cretaceous–Paleogene extinction event, when many species, including the non-avian dinosaurs, became extinct in an event attributed by most experts to the impact of a large asteroid or other celestial body, the Chicxulub impactor.
The Jurassic is a geologic period and stratigraphic system that spanned from the end of the Triassic Period 201.4 million years ago (Mya) to the beginning of the Cretaceous Period, approximately 145 Mya. The Jurassic constitutes the middle period of the Mesozoic Era as well as the eighth period of the Phanerozoic Eon and is named after the Jura Mountains, where limestone strata from the period were first identified.
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae. The term 'angiosperm' is derived from the Greek words ἀγγεῖον / angeion and σπέρμα / sperma ('seed'), meaning that the seeds are enclosed within a fruit. The group was formerly called Magnoliophyta.
The Mesozoic Era is the era of Earth's geological history, lasting from about 252 to 66 million years ago, comprising the Triassic, Jurassic and Cretaceous Periods. It is characterized by the dominance of gymnosperms such as cycads, ginkgoaceae and araucarian conifers, and of archosaurian reptiles such as the dinosaurs; a hot greenhouse climate; and the tectonic break-up of Pangaea. The Mesozoic is the middle of the three eras since complex life evolved: the Paleozoic, the Mesozoic, and the Cenozoic.
In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well as gene-culture coevolution.
Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds. Pollinating agents can be animals such as insects, for example beetles or butterflies; birds, and bats; water; wind; and even plants themselves. Pollinating animals travel from plant to plant carrying pollen on their bodies in a vital interaction that allows the transfer of genetic material critical to the reproductive system of most flowering plants. When self-pollination occurs within a closed flower. Pollination often occurs within a species. When pollination occurs between species, it can produce hybrid offspring in nature and in plant breeding work.
Gnetophyta is a division of plants, grouped within the gymnosperms, that consists of some 70 species across the three relict genera: Gnetum, Welwitschia, and Ephedra. The earliest unambiguous records of the group date to the Jurassic, and they achieved their highest diversity during the Early Cretaceous. The primary difference between gnetophytes and other gymnosperms is the presence of vessel elements, a system of small tubes (xylem) that transport water within the plant, similar to those found in flowering plants. Because of this, gnetophytes were once thought to be the closest gymnosperm relatives to flowering plants, but more recent molecular studies have brought this hypothesis into question, with many recent phylogenies finding them to be nested within the conifers.
The gymnosperms are a group of seed-producing plants that include conifers, cycads, Ginkgo, and gnetophytes, forming the clade Gymnospermae. The term gymnosperm comes from the composite word in Greek: γυμνόσπερμος, and literally means 'naked seeds'. The name is based on the unenclosed condition of their seeds. The non-encased condition of their seeds contrasts with the seeds and ovules of flowering plants (angiosperms), which are enclosed within an ovary. Gymnosperm seeds develop either on the surface of scales or leaves, which are often modified to form cones, or on their own as in yew, Torreya, and Ginkgo. The life cycle of a gymnosperm involves alternation of generations, with a dominant diploid sporophyte phase, and a reduced haploid gametophyte phase, which is dependent on the sporophytic phase. The term "gymnosperm" is often used in paleobotany to refer to all non-angiosperm seed plants. In that case, to specify the modern monophyletic group of gymnosperms, the term Acrogymnospermae is sometimes used.
An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. A significantly large and diverse radiation within a relatively short geologic time scale is often referred to as an explosion. Radiations may affect one clade or many, and be rapid or gradual; where they are rapid, and driven by a single lineage's adaptation to their environment, they are termed adaptive radiations.
The Yixian Formation is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the Barremian stage of the Early Cretaceous. It is known for its exquisitely preserved fossils, and is mainly composed of basalts interspersed with siliciclastic sediments.
Entomophily or insect pollination is a form of pollination whereby pollen of plants, especially but not only of flowering plants, is distributed by insects. Flowers pollinated by insects typically advertise themselves with bright colours, sometimes with conspicuous patterns leading to rewards of pollen and nectar; they may also have an attractive scent which in some cases mimics insect pheromones. Insect pollinators such as bees have adaptations for their role, such as lapping or sucking mouthparts to take in nectar, and in some species also pollen baskets on their hind legs. This required the coevolution of insects and flowering plants in the development of pollination behaviour by the insects and pollination mechanisms by the flowers, benefiting both groups. Both the size and the density of a population are known to affect pollination and subsequent reproductive performance.
Iguanodontidae is a family of iguanodontians belonging to Styracosterna, a derived clade within Ankylopollexia. The clade is formally defined in the PhyloCode by Daniel Madzia and colleagues in 2021 as "the largest clade containing Iguanodon bernissartensis, but not Hadrosaurus foulkii".
The evolution of plants has resulted in a wide range of complexity, from the earliest algal mats of unicellular archaeplastids evolved through endosymbiosis, through multicellular marine and freshwater green algae, to spore-bearing terrestrial bryophytes, lycopods and ferns, and eventually to the complex seed-bearing 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; for example, the ascendance of flowering plants over gymnosperms in terrestrial environments.
This article attempts to place key plant innovations in a geological context. It concerns itself only with novel adaptations and events that had a major ecological significance, not those that are of solely anthropological interest. The timeline displays a graphical representation of the adaptations; the text attempts to explain the nature and robustness of the evidence.
The Paleocene, or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek παλαιός palaiós meaning "old" and the Eocene Epoch, translating to "the old part of the Eocene".
The Mesozoic–Cenozoic Radiation is the third major extended increase of biodiversity in the Phanerozoic, after the Cambrian Explosion and the Great Ordovician Biodiversification Event, which appeared to exceeded the equilibrium reached after the Ordovician radiation. Made known by its identification in marine invertebrates, this evolutionary radiation began in the Mesozoic, after the Permian extinctions, and continues to this date. This spectacular radiation affected both terrestrial and marine flora and fauna, during which the "modern" fauna came to replace much of the Paleozoic fauna. Notably, this radiation event was marked by the rise of angiosperms during the mid-Cretaceous, and the K-Pg extinction, which initiated the rapid increase in mammalian biodiversity.
The Cretaceous–Paleogene (K–Pg) extinction event, also known as the K–T extinction, was the mass extinction of three-quarters of the plant and animal species on Earth approximately 66 million years ago. The event caused the extinction of all non-avian dinosaurs. Most other tetrapods weighing more than 25 kg (55 lb) also became extinct, with the exception of some ectothermic species such as sea turtles and crocodilians. It marked the end of the Cretaceous period, and with it the Mesozoic era, while heralding the beginning of the current era, the Cenozoic. In the geologic record, the K–Pg event is marked by a thin layer of sediment called the K–Pg boundary, Fatkito boundary or K–T boundary, which can be found throughout the world in marine and terrestrial rocks. The boundary clay shows unusually high levels of the metal iridium, which is more common in asteroids than in the Earth's crust.
A nectar spur is a hollow extension of a part of a flower. The spur may arise from various parts of the flower: the sepals, petals, or hypanthium, and often contain tissues that secrete nectar (nectaries). Nectar spurs are present in many clades across the angiosperms, and are often cited as an example of convergent evolution.
The fossil history of flowering plants records the development of flowers and other distinctive structures of the angiosperms, now the dominant group of plants on land. The history is controversial as flowering plants appear in great diversity in the Cretaceous, with scanty and debatable records before that, creating a puzzle for evolutionary biologists that Charles Darwin named an "abominable mystery". Nonetheless, in April 2024, scientists reported an overview of the origin and development of flowering plants over the years based on extensive genetic studies.