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Megafauna

In zoology, megafauna are large animals. The precise definition of the term varies widely, though a common threshold is approximately 45 kilograms (99 lb), this lower end being centered on humans, with other thresholds being more relative to the sizes of animals in an ecosystem, the spectrum of lower-end thresholds ranging from 10 kilograms (22 lb) to 1,000 kilograms (2,200 lb), the latter largely restricted to megaherbivores. Large body size is generally associated with other traits, such as having a slow rate of reproduction and, in large herbivores, reduced or negligible adult mortality from predation.

History
One of the earliest occurrences of the term "megafauna" is Alfred Russel Wallace's 1876 work The geographical distribution of animals. He described the animals as "the hugest, and fiercest, and strangest forms". In the 20th and 21st centuries, the term usually refers to large animals. There are variations in thresholds used to define megafauna as a whole or certain groups of megafauna. Many scientific literature adopt Paul S. Martin's proposed threshold of to classify animals as megafauna. However, for freshwater species, is the preferred threshold. Some scientists define herbivorous terrestrial megafauna as having a weight exceeding , and terrestrial carnivorous megafauna as more than . Additionally, Owen-Smith coined the term megaherbivore to describe herbivores that weighed over , which has seen some use by other researchers. Among living animals, the term megafauna is most commonly used for the largest extant terrestrial mammals, which include (but are not limited to) elephants, giraffes, hippopotamuses, rhinoceroses, and larger bovines. Of these five categories of large herbivores, only bovines are presently found outside of Africa and Asia, but all the others were formerly more wide-ranging, with their ranges and populations continually shrinking and decreasing over time. Wild equines are another example of megafauna, but their current ranges are largely restricted to the Old World, specifically in Africa and Asia. Megafaunal species may be categorized according to their dietary type: megaherbivores (e.g., elephants), megacarnivores (e.g., lions), and megaomnivores (e.g., bears). == Definition ==
Definition
Since the term "megafauna" was first proposed, the scientific community has not settled on a single, succinct definition. However, recent scholarship has been in support of ecological definitions. A 2020 study in Proceedings of the Royal Society B: Biological Sciences analyzed 276 pieces of literature to categorize the different uses of the term. The study determined two typical uses: "keystone megafauna" and "functional megafauna," with a subcategory of "apex megafauna." Their function is further illustrated by the effects of their absence. Losing these species can cause natural ecological processes to unravel, leading to further environmental decline. Keystone megafauna's larger mass and range is significant to such roles they play in their ecosystems. Distinguishing a keystone megafauna from the other categories depends on the individual species' size and their implications for the function of the habitat. Although small species can also function as keystones, they do not meet the sizing criteria of megafauna. The actual mass minimum with this definition will vary between the individuals SPECIFY classification, like how herbivores typically reach around 1000kg to be considered whereas carnivores need only be around 13-16kg. Apex megafauna is an additional term to be used for these species within this category. The difference for these megafauna species, humans for example, lies in their unlikeliness to become prey which increases significantly into adulthood, whether it is because of predation status or their size. Large animals that are not predators can fit into this apex ideation because, similarly, their range is more influenced by the capacity of the habitat rather than the typical phenomenon of being hunted for population control. ==Ecological strategy==
Ecological strategy
Megafauna tend to display the ecological role of K-strategists, with high longevity, slow population growth rates, low mortality rates, and (at least for the largest) few or no natural predators capable of killing adults. Megafauna are considerable contributors to the environments they populate in. The lasting effects of megafauna have been studied, such as Enquist's paper that considered the results from notable megafauna extinction events. During the Anthropocene, many larger species declined in populations (flora and fauna both), which was found to have profound downward effects on the total biosphere activity. This can be attributed to losing these species that assist in many natural processes, like creating new soil, cycling carbon, and population control of the other species. According to simulations and studies done based on current global conditions, removing megafauna from a habitat would leave niches to be filled by smaller plants and animals, upsetting the stable chain of systems that is currently functioning. ==Evolution of large body size==
Evolution of large body size
One observation that has been made about the evolution of larger body size is that rapid rates of increase that are often seen over relatively short time intervals are not sustainable over much longer time periods. In an examination of mammal body mass changes over time, the maximum increase possible in a given time interval was found to scale with the interval length raised to the 0.25 power. However, when considered from the standpoint of rate of size increase per generation, the exponential increase is found to have continued until the appearance of Indricotherium 30 Ma ago. (Since generation time scales with body mass0.259, increasing generation times with increasing size cause the log mass vs. time plot to curve downward from a linear fit.) Megaherbivores eventually attained a body mass of over . The largest of these, indricotheres and proboscids, have been hindgut fermenters, which are believed to have an advantage over foregut fermenters in terms of being able to accelerate gastrointestinal transit in order to accommodate very large food intakes. A similar trend emerges when rates of increase of maximum body mass per generation for different mammalian clades are compared (using rates averaged over macroevolutionary time scales). Among terrestrial mammals, the fastest rates of increase of body mass0.259 vs. time (in Ma) occurred in perissodactyls (a slope of 2.1), followed by rodents (1.2) and proboscids (1.1), all of which are hindgut fermenters. The rate of increase for artiodactyls (0.74) was about a third of the perissodactyls. The rate for carnivorans (0.65) was slightly lower yet, while primates, perhaps constrained by their arboreal habits, had the lowest rate (0.39) among the mammalian groups studied. It has also been suggested that maximum size for mammalian carnivores is constrained by the stress the humerus can withstand at top running speed. Analysis of the variation of maximum body size over the last 40 Ma suggests that decreasing temperature and increasing continental land area are associated with increasing maximum body size. The former correlation would be consistent with Bergmann's rule, and might be related to the thermoregulatory advantage of large body mass in cool climates, The ancestors of cetaceans are believed to have been the semiaquatic pakicetids, no larger than dogs, of about 53 million years (Ma) ago. By 40 Ma ago, cetaceans had attained a length of or more in Basilosaurus, an elongated, serpentine whale that differed from modern whales in many respects and was not ancestral to them. Following this, the evolution of large body size in cetaceans appears to have come to a temporary halt and then to have backtracked, although the available fossil records are limited. However, in the period from 31 Ma ago (in the Oligocene) to the present, cetaceans underwent a significantly more rapid sustained increase in body mass (a rate of increase in body mass0.259 of a factor of 3.2 per million years) than achieved by any group of terrestrial mammals. The cooling trend in Earth's recent history may have generated more localities of high plankton abundance via wind-driven upwellings, facilitating the evolution of gigantic whales. The largest mammal carnivorans of all time are marine pinnipeds, the largest of which is the southern elephant seal, which can reach more than in length and weigh up to . Other large pinnipeds include the northern elephant seal at , walrus at , and Steller sea lion at . The sirenians are another group of marine mammals which adapted to fully aquatic life around the same time as the cetaceans did. Sirenians are closely related to elephants. The largest sirenian was the Steller's sea cow, which reached up to in length and weighed , and was hunted to extinction in the 18th century. In flightless birds species: 1. Dinornis novaezealandiae 2. Emeus crassus 3. Anomalopteryx didiformis 4. Dinornis robustus Because of the small initial size of all mammals following the extinction of the non-avian dinosaurs, nonmammalian vertebrates had a roughly ten-million-year-long window of opportunity (during the Paleocene) for evolution of gigantism without much competition. During this interval, apex predator niches were often occupied by reptiles, such as terrestrial crocodilians (e.g. Pristichampsus), large snakes (e.g. Titanoboa) or varanid lizards, or by flightless birds Flightless paleognaths, termed ratites, have traditionally been viewed as representing a lineage separate from that of their small flighted relatives, the Neotropic tinamous. However, recent genetic studies have found that tinamous nest well within the ratite tree, and are the sister group of the extinct moa of New Zealand. Similarly, the small kiwi of New Zealand have been found to be the sister group of the extinct elephant birds of Madagascar. or herbivorous Aepyornis, ever grew to masses much above ; thus, they never attained the size of the largest mammalian carnivores, let alone that of the largest mammalian herbivores. It has been suggested that the increasing thickness of avian eggshells in proportion to egg mass with increasing egg size places an upper limit on the size of birds. The largest species of Dromornis, D. stirtoni, may have gone extinct after it attained the maximum avian body mass and was then outcompeted by marsupial diprotodonts that evolved to sizes several times larger. In giant turtles Giant tortoises were important components of late Cenozoic megafaunas, being present in every nonpolar continent until the arrival of homininans. The largest known terrestrial tortoise was Megalochelys atlas, an animal that probably weighed about . Some earlier aquatic Testudines, e.g. the marine Archelon of the Cretaceous and freshwater Stupendemys of the Miocene, were considerably larger, weighing more than . ==Megafaunal mass extinctions==
Megafaunal mass extinctions
Timing and possible causes over the last 450,000 years (based on Antarctic temperatures and global ice volume), showing that there were no unique climatic events that would account for any of the megafaunal extinction pulses Numerous extinctions occurred during the latter half of the Last Glacial Period when most large mammals went extinct in the Americas, Australia-New Guinea, and Eurasia, including over 80% of all terrestrial animals with a body mass greater than . Small animals and other organisms like plants were generally unaffected by the extinctions, which is unprecented in previous extinctions during the last 30 million years. Various theories have attributed the wave of extinctions to human hunting, climate change, disease, extraterrestrial impact, competition from other animals or other causes. However, this extinction near the end of the Pleistocene was just one of a series of megafaunal extinction pulses that have occurred during the last 50,000 years over much of the Earth's surface, with Africa and Asia (where the local megafauna had a chance to evolve alongside modern humans) being comparatively less affected. The latter areas did suffer gradual attrition of megafauna, particularly of the slower-moving species (a class of vulnerable megafauna epitomized by giant tortoises), over the last several million years. Outside the mainland of Afro-Eurasia, these megafaunal extinctions followed a highly distinctive landmass-by-landmass pattern that closely parallels the spread of humans into previously uninhabited regions of the world, and which shows no overall correlation with climatic history (which can be visualized with plots over recent geological time periods of climate markers such as marine oxygen isotopes or atmospheric carbon dioxide levels). Australia and nearby islands (e.g., Flores) were struck first around 46,000 years ago, followed by Tasmania about 41,000 years ago (after formation of a land bridge to Australia about 43,000 years ago). The role of humans in the extinction of Australia and New Guinea's megafauna has been disputed, with multiple studies showing a decline in the number of species prior to the arrival of humans on the continent and the absence of any evidence of human predation; the impact of climate change has instead been cited for their decline. North America 13,000 years ago and South America about 500 years later, Cyprus 10,000 years ago, the Antilles 6,000 years ago, New Caledonia and nearby islands 3,000 years ago, Madagascar 2,000 years ago, New Zealand 700 years ago, the Mascarenes 400 years ago, and the Commander Islands 250 years ago. Nearly all of the world's isolated islands could furnish similar examples of extinctions occurring shortly after the arrival of humans, though most of these islands, such as the Hawaiian Islands, never had terrestrial megafauna, so their extinct fauna were smaller, but still displayed island gigantism. In at least some areas, interstadials were periods of expanding human populations. An analysis of Sporormiella fungal spores (which derive mainly from the dung of megaherbivores) in swamp sediment cores spanning the last 130,000 years from Lynch's Crater in Queensland, Australia, showed that the megafauna of that region virtually disappeared about 41,000 years ago, at a time when climate changes were minimal; the change was accompanied by an increase in charcoal, and was followed by a transition from rainforest to fire-tolerant sclerophyll vegetation. The high-resolution chronology of the changes supports the hypothesis that human hunting alone eliminated the megafauna, and that the subsequent change in flora was most likely a consequence of the elimination of browsers and an increase in fire. from analysis of Sporormiella fungal spores from a lake in eastern North America and from study of deposits of Shasta ground sloth dung left in over half a dozen caves in the American Southwest. Continuing human hunting and environmental disturbance has led to additional megafaunal extinctions in the recent past, and has created a serious danger of further extinctions in the near future (see examples below). Direct killing by humans, primarily for meat or other body parts, is the most significant factor in contemporary megafaunal decline. A number of other mass extinctions occurred earlier in Earth's geologic history, in which some or all of the megafauna of the time also died out. Famously, in the Cretaceous–Paleogene extinction event, the non-avian dinosaurs and most other giant reptiles were eliminated. However, the earlier mass extinctions were more global and not so selective for megafauna; i.e., many species of other types, including plants, marine invertebrates and plankton, went extinct as well. Thus, the earlier events must have been caused by more generalized types of disturbances to the biosphere. Consequences of depletion of megafauna Depletion of herbivorous megafauna results in increased growth of woody vegetation, and a consequent increase in wildfire frequency. Megafauna may help to suppress the growth of invasive plants. Large herbivores and carnivores can suppress the abundance of smaller animals, resulting in their population increase when megafauna are removed. In South America's Amazon Basin, it is estimated that such lateral diffusion was reduced over 98% following the megafaunal extinctions that occurred roughly 12,500 years ago. Given that phosphorus availability is thought to limit productivity in much of the region, the decrease in its transport from the western part of the basin and from floodplains (both of which derive their supply from the uplift of the Andes) to other areas is thought to have significantly impacted the region's ecology, and the effects may not yet have reached their limits. Effect on methane emissions Large populations of megaherbivores have the potential to contribute greatly to the atmospheric concentration of methane, which is an important greenhouse gas. Modern ruminant herbivores produce methane as a byproduct of foregut fermentation in digestion and release it through belching or flatulence. Today, around 20% of annual methane emissions come from livestock methane release. In the Mesozoic, it has been estimated that sauropods could have emitted 520 million tons of methane to the atmosphere annually, contributing to the warmer climate of the time (up to 10 °C (18 °F) warmer than at present). This large emission follows from the enormous estimated biomass of sauropods, and because methane production of individual herbivores is believed to be almost proportional to their mass. One study examined the methane emissions from the bison that occupied the Great Plains of North America before contact with European settlers. The study estimated that the removal of the bison caused a decrease of as much as 2.2 million tons per year. Another study examined the change in the methane concentration in the atmosphere at the end of the Pleistocene epoch after the extinction of megafauna in the Americas. After early humans migrated to the Americas about 13,000 BP, their hunting and other associated ecological impacts led to the extinction of many megafaunal species there. Calculations suggest that this extinction decreased methane production by about 9.6 million tons per year. This suggests that the absence of megafaunal methane emissions may have contributed to the abrupt climatic cooling at the onset of the Younger Dryas. The decrease in atmospheric methane that occurred at that time, as recorded in ice cores, was 2 to 4 times more rapid than any other decrease in the last half million years, suggesting that an unusual mechanism was at work. == Current studies ==
Current studies
Current studies being completed on megafauna species are reflective, and do not tend to offer a set definition of the subject; additionally, some consider there to be a significant lack in cross-disciplinary studies. Megaflora also contribute a huge amount to their ecosystems, offering shelter for many species and food for even more, including many megafauna. ==Gallery==
Gallery
Pleistocene extinct megafauna File:Dinornis novaezealandiae.png|Moa (Dinornis pictured) File:Diprotodon optatum (2).jpg|Diprotodon optatum File:Varanus priscus Melbourne Museum.jpg|"Megalania" (Varanus priscus) File:Panthera leo atrox Sergiodlarosa.jpg|American lions (Panthera atrox) File:Woolly mammoth (Mammuthus primigenius) - Mauricio Antón.jpg|alt=Woolly mammoths vanished after humans invaded their habitat in Eurasia and N. America.|Woolly mammoth File:Archaeoindris fontoynonti.jpg|The subfossil lemur Archaeoindris File:Giant Haasts eagle.jpg|Haast's eagle File:Macrauchenia patachonica Life Reconstruction.png|Restoration of Macrauchenia, a camel-sized member of the extinct ungulate order Litopterna File:Doedicurus and Glyptodon.jpg|Life restoration of the glyptodonts Doedicurus (front) and Glyptodon Other extinct Cenozoic megafauna File:Dromornis stirtoni.jpg|Dromornis stirtoni File:Indricotherium11.jpg|alt=Asian indricothere rhino Paraceratherium was among the largest land mammals, about twice a bush elephant's mass.|Asian paraceratheriid rhino Paraceratherium was among the largest land mammals File:Megalodon jaws on display at the National Baltimore Aquarium.jpg|Reconstructed jaws of megalodon (Otodus megalodon) File:Deinotherium12.jpg|Deinotherium File:Kelenken.jpg|Kelenken guillermoi File:Gastornis.png|Gastornis gigantea Extant File:Nandu Rhea americana Tierpark Hellabrunn-1.jpg|Greater rhea File:Kbnpsilverbackandchild 0.5.jpg|Eastern gorilla File:Hunting Tiger Ranthambore.jpg|Bengal tiger File:Polar Bear 2004-11-15.jpg|Polar bear File:Ngorongoro Spitzmaulnashorn edit1crop.jpg|Black rhinoceros File:Ovibos moschatus qtl3.jpg|Muskox File:Alaska moose.jpg|Moose File:Nijlpaard.jpg|Hippopotamus File:Mother and baby sperm whale.jpg|Adult sperm whale and calf File:Killerwhales jumping.jpg|Orcas File:Southern Cassowary 7071.jpg|Southern cassowary File:Ostrich Struthio camelus Tanzania 3742 cropped Nevit.jpg|Common ostrich File:SaltwaterCrocodile('Maximo').jpg|Saltwater crocodile File:Komodo dragon Varanus komodoensis Ragunan Zoo 2.JPG|Komodo dragon File:Anaconda Loreto Peru.jpg|Green anaconda File:Mola alexandrini (Bump-head Mola).jpg|Giant sunfish File:Lates niloticus 2.jpg|Nile perch File:White shark.jpg|Great white shark File:Manta alfredi fushivaru thila.jpg|Reef manta ray File:Giant squid Ranheim2.jpg|Carcass of a giant squid ==See also==
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