Charles Darwin famously called it an 'ábominable mystery'. He was referring to the sudden appearance and diversification of flowering plants in the Cretaceous fossil record. He noticed that these early fossils resembled modern flowering plants. 'Primitive' or ancestral stages were missing. Today, biologists categorize these as crown and stem representatives of a group.
The first fossil evidence of flowering plants is from 140-130 million year old sediments. These are early types of pollen grains with one aperture (uniaperturate). Triaperturate pollen is found in slightly younger 125 million year old rocks. Towards the end of the early Cretaceous, by around 100 million years ago, flowers, leaves, and other organs appear from several continents representing all the major groups of angiosperms.
The picture below is of an early Cretaceous (~100 million year old) flowering plant from the lotus family. The location is northeast Brazil. There is a remarkable preservation of the whole plant, with connected roots, rhizome, leaves, and aggregate fruit.
Source: William Vieira Gobo et.al. Nature Scientific Reports 2023- A new remarkable Early Cretaceous nelumbonaceous fossil bridges the gap between herbaceous aquatic and woody protealeans.
Taking a long view of their evolutionary pattern, angiosperm diversification is structured in three phases. The first phase was a steady expansion through early to late Cretaceous. There was more rapid diversification in late Cretaceous by around 70 million years ago. Enumeration of floral species through the Cretaceous indicate that angiosperms made up about 5% of species in early Cretaceous, increasing to 80% by Maastrichtian times (late Cretaceous). Despite this increase in species numbers, in terms of biomass, angiosperms were still a small component of Cretaceous floras. Their domination of floral communities, including the origin of modern wet tropical forests, began in the Paleogene (65-24 million years ago) after the end Cretaceous mass extinction. Michael J. Benton, Peter Wilf, and Herve Sauquet have provided a good overview in New Phytologist of this pivotal phase of ecosystem change.
These evolutionary changes did not occur in isolation. Throughout the Cretaceous, significant changes were occurring to terrestrial ecosystems, with the origination of many plant and animal groups. This extended phase of ecosystem reorganization is known as the Cretaceous Terrestrial Revolution. Angiosperm diversification is thought to have played a key role in this transformation of land biodiversity, so much so, that the phase from about 100 million years to 50 million years ago is known as the Angiosperm Terrestrial Revolution.
The Cretaceous -Paleogene mass extinction hit angiosperms hard, as well as altering the trajectory of their evolution. For example, there was a 40% loss of diversity of flowering plants in Colombia following the mass extinction. But certain attributes of angiosperms, such as their partnerships with other organisms, their ability to efficiently capture energy and enhance photosynthetic rates, and an underlying genetic propensity to speciate, resulted in them expanding rapidly in the post extinction landscape. Angiosperm evolution opened up opportunities for a variety of land creatures including insects, spiders, lizards, birds, and mammals, eventually driving up terrestrial biodiversity to 10 times more as marine biodiversity.
Paleobiologists are interested in understanding the interaction and impact angiosperm diversification could have had on other groups of plants and animals. Of particular interest is the diversification of insects in the Cretaceous and Paleogene.
Modern insect lineages began diversifying by 245 million years ago, long before angiosperms evolved. Gymnosperm and insect communities preserved in amber and sediments show that insects had an intricate relationship with host gymnosperms like cycads, conifers and ginkgoaleans. Insect pollination of gymnosperms predated the origin of angiosperms by at least 100 million years and their fossil record show phases of diversification even when angiosperms were rare.
Did angiosperm evolution also drive a rise in insect diversity?
Pollinator insects particularly would seem to benefit from an abundance
in flowering plants, and if so, what co-evolutionary patterns are
apparent from the fossil record?
David Perise and Fabien Condamine have tackled this question in a new study in Nature Communications. I will share this beautifully compiled infographic from the paper that conveys so clearly the patterns of angiosperm and insect diversification through the Cretaceous and Cenozoic.
Digging into published databases, the researchers compiled data on the origination and extinction times of angiosperm and insect families. They then statistically analyzed whether angiosperm and insect origination and extinction times, and pulses of their diversification coincide. Their analysis showed that angiosperms seemed to have played a dual role in insect evolution. They mitigated insect extinction through the Cretaceous and spurred on the origination of new insect groups in the Cenozoic. Besides a broad analysis of insects, they also found that pollinator insects like bees and long proboscid butterflies show a pronounced diversification alongside angiosperm lineages.
The success of angiosperms in the late Cretaceous and Cenozoic coincided with the decline in gymnosperms. Intrinsic mechanisms of genomic rearrangements in angiosperms resulted in repeated evolution of novel traits and specializations. They competitively displaced gymnosperms. The impact on gymnosperm dependent insects was variable. Generalist insect pollinators such as several beetle lineages transitioned to angiosperms. Much of the co-diversification of angiosperm and insects can be explained by this shift of gymnosperm pollinators to angiosperm hosts. Gymnosperm specialized insect groups did not fare that well. For example, gymnosperms like Cheirolepidiaceae and Bennettitales went extinct by the latest Cretaceous. This was followed by the extinction of insect groups that were dependent on these plants such as some specialized long-proboscid flies, scorpionflies and lacewings.
Insect diversification did not depend only on angiosperms. Analysis also shows that warmer climate phases negatively impacted insect diversity and coincided with higher insect extinction rates. There seems also to be a relationship with other plant types. Spore plant and gymnosperm diversity had a positive impact on origination rates of insects. Ecosystem relationships and dependencies are multifarious and complex as this analysis between angiosperm and insect co-evolution shows.
Darwin's anxiety over flowering plants reflected his insistence that evolution is gradual. Nature does not make leaps, he stressed. He explained abruptness in the fossil record by invoking missing strata due to non deposition and erosion. Regarding flowering plants, he suggested that fossils were perhaps preceded by a period of cryptic evolution of that lineage that took place in a remote area or a lost continent, although he conceded that this was a poor explanation. However, this latter view, that a substantial lag time or a long fuse precedes the bang, continues to resonate among many biologists. Molecular methods that compares accumulated genetic difference to calculate the time of divergence of groups indicate a fairly long gap between the genetic branching of lineages and their first fossil appearance.
Most familiar is the example of the origin of animals. Molecular data indicate that animals originated by 750 million years ago, yet unequivocal animal fossils appear by 570 million years ago, close to 200 million years later. Similarly, some molecular estimates put angiosperm origins to pre-Cretaceous times, stretching back 240-200 million years ago to Triassic-Jurassic, a good 100 million to 60 million years before first appearance of fossils.
This idea of a phylogenetic fuse has been recently criticized. Published in Systematic Biology, Graham E. Budd and Richard P. Mann have undertaken a critical examination of molecular clock methods. Their analysis indicate that popular methods used to assign probabilities to maximum age of lineages are biased against rapid lineage radiations being true evolutionary events. In their view, the mismatch between molecular dates of lineage origin and the timing of the first appearance of their fossils is an artifact. They point out that the coincident appearance of fossils from widespread localities in a particular sequence and across different modes of preservation faithfully records evolution. The time gap between the origin and later diversification of lineages is not that deep.
The 'abominable mystery' of the sudden appearance of fossil groups may in fact be a real biological motif in earth history, signalling the rapid radiation of lineages filling ecologic spaces following an environmental crises and evolutionary innovation.
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