History of scientific understanding
For much of history, the modern understanding of extinction as the end of a species was incompatible with the prevailing worldview. Prior to the 19th century, much of Western society adhered to the belief that the world was created by God and as such was complete and perfect. This concept reached its heyday in the 1700s with the peak popularity of a theological concept called the great chain of being, in which all life on earth, from the tiniest microorganism to God, is linked in a continuous chain. The extinction of a species was impossible under this model, as it would create gaps or missing links in the chain and destroy the natural order. Thomas Jefferson was a firm supporter of the great chain of being and an opponent of extinction, famously denying the extinction of the woolly mammoth on the grounds that nature never allows a race of animals to become extinct.
A series of fossils were discovered in the late 17th century that appeared unlike any living species. As a result, the scientific community embarked on a voyage of creative rationalization, seeking to understand what had happened to these species within a framework that did not account for total extinction. In October 1686, Robert Hooke presented an impression of a nautilus to the Royal Society that was more than two feet in diameter, and morphologically distinct from any known living species. Hooke theorized that this was simply because the species lived in the deep ocean and no one had discovered them yet. While he contended that it was possible a species could be "lost", he thought this highly unlikely. Similarly, in 1695, Sir Thomas Molyneux published an account of enormous antlers found in Ireland that did not belong to any extant taxa in that area. Molyneux reasoned that they came from the North American moose and that the animal had once been common on the British Isles. Rather than suggest that this indicated the possibility of species going extinct, he argued that although organisms could become locally extinct, they could never be entirely lost and would continue to exist in some unknown region of the globe. The antlers were later confirmed to be from the extinct deer Megaloceros. Hooke and Molyneux's line of thinking was difficult to disprove. When parts of the world had not been thoroughly examined and charted, scientists could not rule out that animals found only in the fossil record were not simply "hiding" in unexplored regions of the Earth.
Georges Cuvier is credited with establishing the modern conception of catastrophic extinction in a 1796 lecture to the French Institute, though he would spend most of his career trying to convince the wider scientific community of his theory. Cuvier was a well-regarded paleontologist, lauded for his ability to reconstruct the anatomy of an unknown species from a few fragments of bone. His primary evidence for extinction came from mammoth skulls found near Paris. Cuvier recognized them as distinct from any known living species of elephant, and argued that it was highly unlikely such an enormous animal would go undiscovered. In 1798, he studied a fossil from the Paris Basin that was first observed by Robert de Lamanon in 1782, first hypothesizing that it belonged to a canine but then deciding that it instead belonged to an animal that was unlike living ones. His study paved the way to his naming of the extinct mammal genus Palaeotherium in 1804 based on the skull and additional fossil material along with another extinct contemporary mammal genus Anoplotherium. In both genera, he noticed that their fossils shared some similarities with other mammals like ruminants and rhinoceroses but still had distinct differences. In 1812, Cuvier, along with Alexandre Brongniart and Geoffroy Saint-Hilaire, mapped the strata of the Paris basin. They saw alternating saltwater and freshwater deposits, as well as patterns of the appearance and disappearance of fossils throughout the record. From these patterns, Cuvier inferred historic cycles of catastrophic flooding, extinction, and repopulation of the earth with new species.
Cuvier's fossil evidence showed that very different life forms had existed in the past to those that exist today, a fact that was accepted by most scientists. The primary debate focused on whether this turnover caused by extinction was gradual or abrupt in nature. Cuvier understood extinction to be the result of cataclysmic events that wipe out huge numbers of species, as opposed to the gradual decline of a species over time. In his Essay on the Theory of the Earth (1813), Cuvier proposed that now-extinct species had been wiped out by periodic catastrophic flooding events. His catastrophic view of the nature of extinction garnered him many opponents in the newly emerging school of uniformitarianism.
Jean-Baptiste Lamarck, a gradualist and colleague of Cuvier, saw the fossils of different life forms as evidence of the mutable character of species. While Lamarck did not deny the possibility of extinction, he believed that it was exceptional and rare and that most of the change in species over time was due to gradual change. Unlike Cuvier, Lamarck was skeptical that catastrophic events of a scale large enough to cause total extinction were possible. In his geological history of the earth titled Hydrogeologie, Lamarck instead argued that the surface of the earth was shaped by gradual erosion and deposition by water, and that species changed over time in response to the changing environment.
Charles Lyell, a noted geologist and founder of uniformitarianism, believed that past processes should be understood using present day processes. Like Lamarck, Lyell acknowledged that extinction could occur, noting the total extinction of the dodo and the extirpation of indigenous horses to the British Isles. He similarly argued against mass extinctions, believing that any extinction must be a gradual process. Lyell also showed that Cuvier's original interpretation of the Parisian strata was incorrect. Instead of the catastrophic floods inferred by Cuvier, Lyell demonstrated that patterns of saltwater and freshwater deposits, like those seen in the Paris basin, could be formed by a slow rise and fall of sea levels.
The concept of extinction was integral to Charles Darwin's On the Origin of Species, with less fit lineages disappearing over time. For Darwin, extinction was a constant side effect of competition. Because of the wide reach of On the Origin of Species, it was widely accepted that extinction occurred gradually and evenly (a concept now referred to as background extinction). It was not until 1982, when David Raup and Jack Sepkoski published their seminal paper on mass extinctions, that Cuvier was vindicated and catastrophic extinction was accepted as an important mechanism. The current understanding of extinction is a synthesis of the cataclysmic extinction events proposed by Cuvier, and the background extinction events proposed by Lyell and Darwin.
