The Lilliput effect is an observed decrease in animal body size in genera that have survived a major extinction. [1] There are several hypotheses as to why these patterns appear in the fossil record, some of which are: [2]
The term was coined in by Urbanek (1993) in a paper concerning the end-Silurian extinction of graptoloids [3] and is derived from an island in Gulliver’s Travels , Lilliput, inhabited by a race of miniature people. The size decrease may just be a temporary phenomenon restricted to the survival period of the extinction event. Atkinson et al. (2019) coined the term Brobdingnag effect [4] to describe a related phenomenon, operating in the opposite direction, whereby new species evolving after the Triassic-Jurassic mass extinction that began the period with small body sizes underwent substantial size increases. [4] The term is also from Gulliver's Travels , where Brobdingnag is a place inhabited by a race of giants.
Trends in body size changes are seen throughout the fossil record in many organisms, and major changes (shrinking and dwarfing) in body size can significantly affect the morphology of the animal itself as well as how it interacts with the environment. [2] Since Urbanek's publication several researchers have described a decrease in body size in fauna post-extinction event, although not all use the term "Lilliput effect" when discussing this trend in body size decrease. [5] [6] [7]
The Lilliput effect has been noted by several authors to have occurred after the Permian-Triassic mass extinction: Early Triassic fauna, both marine and terrestrial, is notably smaller than those preceding and following in the geologic record. [1]
The extinction event may have been more severe for the larger-bodied species, leaving only species of smaller-bodied animals behind. [1] As such, organisms in the smaller species which then make up the recovering ecosystem, will take time to evolve larger bodies to replace the extinct species and re-occupy the vacant ecological niche for a large-bodied animal. [1] Taxa whose animals are larger may be evolutionarily selected against for several reasons, including [1]
Stanley (1973) hypothesized that newly emerged animal taxa tend to develop at an originally small size, hence a sudden proliferation of new species would tend to produce many initially small organisms. [8]
It is possible that the extinction event selectively removed larger individuals within any single lineage, without extinguishing the entire species, but leaving as survivors only the individuals with a naturally smaller body size. The smaller survivors then form the new breeding population, and pass on that trait to their descendents. Because of the selection during the extinction, compared to the previously "normal"-sized members of the species who lived before the extinction event occurred, later members of that species living after the extinction, who are descended only from the smaller survivors, would be reduced in size, constituting a "new-normal". [1]
Approximately 251.9 million years ago, the Permian–Triassicextinction event forms the boundary between the Permian and Triassic geologic periods, and with them the Paleozoic and Mesozoic eras. It is Earth's most severe known extinction event, with the extinction of 57% of biological families, 83% of genera, 81% of marine species and 70% of terrestrial vertebrate species. It is also the greatest known mass extinction of insects. It is the greatest of the "Big Five" mass extinctions of the Phanerozoic. There is evidence for one to three distinct pulses, or phases, of extinction.
The Silurian is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at 443.8 million years ago (Mya), to the beginning of the Devonian Period, 419.2 Mya. The Silurian is the shortest period of the Paleozoic Era. As with other geologic periods, the rock beds that define the period's start and end are well identified, but the exact dates are uncertain by a few million years. The base of the Silurian is set at a series of major Ordovician–Silurian extinction events when up to 60% of marine genera were wiped out.
The Triassic is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago (Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is the first and shortest period of the Mesozoic Era. Both the start and end of the period are marked by major extinction events. The Triassic Period is subdivided into three epochs: Early Triassic, Middle Triassic and Late Triassic.
The Triassic–Jurassic (Tr-J) extinction event (TJME), often called the end-Triassic extinction, was a Mesozoic extinction event that marks the boundary between the Triassic and Jurassic periods, 201.4 million years ago, and is one of the top five major extinction events of the Phanerozoic eon, profoundly affecting life on land and in the oceans. In the seas, the entire class of conodonts and 23–34% of marine genera disappeared. On land, all archosauromorphs other than crocodylomorphs, pterosaurs, and non-avian dinosaurs became extinct; some of the groups which died out were previously abundant, such as aetosaurs, phytosaurs, and rauisuchids. Some remaining non-mammalian therapsids and many of the large temnospondyl amphibians had become extinct prior to the Jurassic as well. However, there is still much uncertainty regarding a connection between the Tr-J boundary and terrestrial vertebrates, due to a lack of terrestrial fossils from the Rhaetian (uppermost) stage of the Triassic. Plants, crocodylomorphs, dinosaurs, pterosaurs and mammals were left largely untouched; this allowed the dinosaurs, pterosaurs, and crocodylomorphs to become the dominant land animals for the next 135 million years.
The Late Ordovician mass extinction (LOME), sometimes known as the end-Ordovician mass extinction or the Ordovician-Silurian extinction, is the first of the "big five" major mass extinction events in Earth's history, occurring roughly 445 million years ago (Ma). It is often considered to be the second-largest known extinction event just behind the end-Permian mass extinction, in terms of the percentage of genera that became extinct. Extinction was global during this interval, eliminating 49–60% of marine genera and nearly 85% of marine species. Under most tabulations, only the Permian-Triassic mass extinction exceeds the Late Ordovician mass extinction in biodiversity loss. The extinction event abruptly affected all major taxonomic groups and caused the disappearance of one third of all brachiopod and bryozoan families, as well as numerous groups of conodonts, trilobites, echinoderms, corals, bivalves, and graptolites. Despite its taxonomic severity, the Late Ordovician mass extinction did not produce major changes to ecosystem structures compared to other mass extinctions, nor did it lead to any particular morphological innovations. Diversity gradually recovered to pre-extinction levels over the first 5 million years of the Silurian period.
The Late Devonian extinction consisted of several extinction events in the Late Devonian Epoch, which collectively represent one of the five largest mass extinction events in the history of life on Earth. The term primarily refers to a major extinction, the Kellwasser event, also known as the Frasnian-Famennian extinction, which occurred around 372 million years ago, at the boundary between the Frasnian age and the Famennian age, the last age in the Devonian Period. Overall, 19% of all families and 50% of all genera became extinct. A second mass extinction called the Hangenberg event, also known as the end-Devonian extinction, occurred 359 million years ago, bringing an end to the Famennian and Devonian, as the world transitioned into the Carboniferous Period.
The Andean-Saharan glaciation, also known as the Early Paleozoic Ice Age (EPIA), the Early Paleozoic Icehouse, the Late Ordovician glaciation, the end-Ordovician glaciation, or the Hirnantian glaciation, occurred during the Paleozoic from approximately 460 Ma to around 420 Ma, during the Late Ordovician and the Silurian period. The major glaciation during this period was formerly thought only to consist of the Hirnantian glaciation itself but has now been recognized as a longer, more gradual event, which began as early as the Darriwilian, and possibly even the Floian. Evidence of this glaciation can be seen in places such as Arabia, North Africa, South Africa, Brazil, Peru, Bolivia, Chile, Argentina, and Wyoming. More evidence derived from isotopic data is that during the Late Ordovician, tropical ocean temperatures were about 5 °C cooler than present day; this would have been a major factor that aided in the glaciation process.
Tabulata, commonly known as tabulate corals, are an order of extinct forms of coral. They are almost always colonial, forming colonies of individual hexagonal cells known as corallites defined by a skeleton of calcite, similar in appearance to a honeycomb. Adjacent cells are joined by small pores. Their distinguishing feature is their well-developed horizontal internal partitions (tabulae) within each cell, but reduced or absent vertical internal partitions. They are usually smaller than rugose corals, but vary considerably in shape, from flat to conical to spherical.
The Early Triassic is the first of three epochs of the Triassic Period of the geologic timescale. It spans the time between 251.9 Ma and 247.2 Ma. Rocks from this epoch are collectively known as the Lower Triassic Series, which is a unit in chronostratigraphy. The Early Triassic is the oldest epoch of the Mesozoic Era. It is preceded by the Lopingian Epoch and followed by the Middle Triassic Epoch. The Early Triassic is divided into the Induan and Olenekian ages. The Induan is subdivided into the Griesbachian and Dienerian subages and the Olenekian is subdivided into the Smithian and Spathian subages.
The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between 237 Ma and 201.4 Ma. It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic Epoch. The corresponding series of rock beds is known as the Upper Triassic. The Late Triassic is divided into the Carnian, Norian and Rhaetian ages.
The Lystrosaurus Assemblage Zone is a tetrapod assemblage zone or biozone which correlates to the upper Adelaide and lower Tarkastad Subgroups of the Beaufort Group, a fossiliferous and geologically important geological Group of the Karoo Supergroup in South Africa. This biozone has outcrops in the south central Eastern Cape and in the southern and northeastern Free State. The Lystrosaurus Assemblage Zone is one of eight biozones found in the Beaufort Group, and is considered to be Early Triassic in age.
The Lau event was the last of three relatively minor mass extinctions during the Silurian period. It had a major effect on the conodont fauna, but barely scathed the graptolites, though they suffered an extinction very shortly thereafter termed the Kozlowskii event that some authors have suggested was coeval with the Lau event and only appears asynchronous due to taphonomic reasons. It coincided with a global low point in sea level caused by glacioeustasy and is closely followed by an excursion in geochemical isotopes in the ensuing late Ludfordian faunal stage and a change in depositional regime.
The Hangenberg event, also known as the Hangenberg crisis or end-Devonian extinction, is a mass extinction that occurred at the end of the Famennian stage, the last stage in the Devonian Period. It is usually considered the second-largest extinction in the Devonian Period, having occurred approximately 13 million years after the Late Devonian mass extinction at the Frasnian-Famennian boundary. The event is named after the Hangenberg Shale, which is part of a sequence that straddles the Devonian-Carboniferous boundary in the Rhenish Massif of Germany.
Moschorhinus is an extinct genus of therocephalian synapsid in the family Akidnognathidae with only one species: M. kitchingi, which has been found in the Late Permian to Early Triassic of the South African Karoo Supergroup. It was a large carnivorous therapsid, reaching 1.1–1.5 metres (3.6–4.9 ft) in total body length with the largest skull comparable to that of a lion in size, and had a broad, blunt snout which bore long, straight canines.
The Ireviken event was the first of three relatively minor extinction events during the Silurian period. It occurred at the Llandovery/Wenlock boundary. The event is best recorded at Ireviken, Gotland, where over 50% of trilobite species became extinct; 80% of the global conodont species also became extinct in this interval.
Gnathorhiza is an extinct genus of prehistoric lobe-finned fish (lungfish) which lived from the Carboniferous period to the Early Triassic epoch. It is the only known lungfish genus to have crossed the Permo-Triassic boundary. Several species have been described, ranging in size from 5 to 50 centimeters.
The Carboniferous rainforest collapse (CRC) was a minor extinction event that occurred around 305 million years ago in the Carboniferous period. The event occurred at the end of the Moscovian and continued into the early Kasimovian stages of the Pennsylvanian.
Productida is an extinct order of brachiopods in the extinct class Strophomenata. Members of Productida first appeared during the Silurian. They represented the most abundant group of brachiopods during the Permian period, accounting for 45-70% of all species. The vast majority of species went extinct during the Permian-Triassic extinction event, though a handful survived into the Early Triassic. Many productids are covered in hollow tubular spines, which are characteristic of the group. A number of functions for the spines have been proposed, including as a defensive mechanism against predators.
Alatoconchidae is an extinct family of prehistoric bivalves that lived in the early to middle Permian period. Genera belonging to Alatoconchidae are characterized by their shell that is strongly compressed in the dorsoventral direction. Some species reached large sizes of as much as 1 metre (3.3 ft) long. It is hypothesized that some species in this family got energy from chemosynthetic bacteria.
The Lundgreni Event, also known as the Mid-Homerian Biotic Crisis, was an extinction event during the middle Homerian age of the Silurian period. Evidence for the event has been observed in Silurian marine deposits in the Iberian Peninsula, Bohemia, and Poland.