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Late Pleistocene extinctions

February 16, 2024

For extinctions that occurred during the Holocene, see Holocene extinction.

The Late Pleistocene to the beginning of the Holocene saw numerous extinctions of predominantly megafaunal (typically defined as having body masses over 44 kilograms (97 lb)[1]) animal species (the Pleistocene megafauna), which resulted in a collapse in faunal density and diversity across the globe.[2] The extinctions during the Late Pleistocene are differentiated from previous extinctions by the widespread absence of ecological succession to replace these extinct megafaunal species,[3] and the regime shift of previously established faunal relationships and habitats as a consequence. The timing and severity of the extinctions varied by region and are thought to have been driven by varying combinations of human and climatic factors.[3] Human impact on megafauna populations is thought to have been driven by hunting ("overkill"),[4][5] as well as possibly environmental alteration.[6] The relative importance of human vs climatic factors in the extinctions has been the subject of long-running controversy.[3]

Late Pleistocene landscape of northern Spain, by Mauricio Antón. From left to right: wild horse (Equus ferus), woolly mammoth (Mammuthus primigenius) reindeer (Rangifer tarandus) the cave or steppe lion (Panthera spelaea) and the woolly rhinoceros (Coelodonta antiquitatis)
Mural of La Brea Tar Pits by Charles R. Knight, including sabertooth cats (Smilodon fatalis, left) ground sloths (Paramylodon harlani, right) and Columbian mammoths (Mammuthus columbi, background)

Major extinctions occurred in Australia-New Guinea (Sahul) beginning approximately 50,000 years ago and in the Americas about 13,000 years ago, coinciding in time with the early human migrations into these regions.[7] Extinctions in northern Eurasia were staggered over tens of thousands of years between 50,000 and 10,000 years ago,[2] while extinctions in the Americas were virtually simultaneous, spanning only 3000 years at most.[4][8] Overall, during Late Pleistocene about 65% of all megafaunal species worldwide became extinct,[9] rising to 72% in North America, 83% in South America and 88% in Australia,[10] with all mammals over 1,000 kilograms (2,200 lb) becoming extinct in Australia and the Americas.[1] Africa, South Asia and Southeast Asia experienced much lower extinctions than other regions.[10]

Extinctions by biogeographic realm edit

Summary edit

Extinctions range of the continental large and medium-sized mammals from 40,000 to 4,000 years BP in different biogeographic realms[11]
Biogeographic realm Giants
(over 1,000 kg) 		Very large
(400–1,000 kg) 		Large
(150–400 kg) 		Moderately large
(50–150 kg) 		Medium
(10–50 kg) 		Total Regions included
Start Loss % Start Loss % Start Loss % Start Loss % Start Loss % Start Loss %
Afrotropic 6 −1 16.6% 4 −1 25% 25 −3 12% 32 0 0% 69 −2 2.9% 136 -7 5.1% Trans-Saharan Africa and Arabia
Indomalaya 5 −2 40% 6 −1 16.7% 10 −1 10% 20 −3 15% 56 −1 1.8% 97 -8 8.2% Indian subcontinent, Southeast Asia, and southern China
Palearctic 8 −8 100% 10 −5 50% 14 −5 35.7% 23 −3 15% 41 −1 2.4% 96 -22 22.9% Eurasia and North Africa
Nearctic 5 −5 100% 10 −8 80% 26 −22 84.6% 20 −13 65% 25 −9 36% 86 -57 66% North America
Neotropic 9 −9 100% 12 −12 100% 17 −14 82% 20 −11 55% 35 −5 14.3% 93 -51 54% South America, Central America, South Florida, and the Caribbean
Australasia 4 −4 100% 5 −5 100% 6 −6 100% 16 −13 81.2% 25 −10 40% 56 -38 67% Australia, New Guinea, New Zealand, and neighbouring islands.
Global 33 −26 78.8% 46 −31 67.4% 86 −47 54.7% 113 −41 36.3% 215 −23 10.1% 493 -168 34%

Introduction edit

 
The proportion of extinct large mammal species (more than or equal to 10 kg (22 lb)) in each country during the last 132,000 years, only counting extinctions earlier than 1000 years BP

The Late Pleistocene saw the extinction of many mammals weighing more than 40 kilograms (88 lb). The proportion of megafauna extinctions is progressively larger the further the human migratory distance from Africa, with the highest extinction rates in Australia, and North and South America.

The increased extent of extinction mirrors the migration pattern of modern humans: the further away from Africa, the more recently humans inhabited the area, the less time those environments (including its megafauna) had to become accustomed to humans (and vice versa).

There are two main hypotheses to explain this extinction:

  • Climate change associated with the advance and retreat of major ice caps or ice sheets causing reduction in favorable habitat.
  • Human hunting causing attrition of megafauna populations, commonly known as "overkill".[12]

There are some inconsistencies between the current available data and the prehistoric overkill hypothesis. For instance, there are ambiguities around the timing of Australian megafauna extinctions.[12] Evidence supporting the prehistoric overkill hypothesis includes the persistence of megafauna on some islands for millennia past the disappearance of their continental cousins. For instance, ground sloths survived on the Antilles long after North and South American ground sloths were extinct, woolly mammoths died out on remote Wrangel Island 6,000 years after their extinction on the mainland, while Steller's sea cows persisted off the isolated and uninhabited Commander Islands for thousands of years after they had vanished from the continental shores of the north Pacific.[13] The later disappearance of these island species correlates with the later colonization of these islands by humans.

The original debates as to whether human arrival times or climate change constituted the primary cause of megafaunal extinctions necessarily were based on paleontological evidence coupled with geological dating techniques. Recently, genetic analyses of surviving megafaunal populations have contributed new evidence, leading to the conclusion: "The inability of climate to predict the observed population decline of megafauna, especially during the past 75,000 years, implies that human impact became the main driver of megafauna dynamics around this date."[14]

An alternative hypothesis to the theory of human responsibility is climate change associated with the last glacial period. Discredited explanations include the Younger Dryas impact hypothesis[15] and Tollmann's hypothesis that extinctions resulted from bolide impacts.

Recent research indicates that each species responded differently to environmental changes, and no one factor by itself explains the large variety of extinctions. The causes may involve the interplay of climate change, competition between species, unstable population dynamics, and human predation.[16]

Africa edit

Although Africa was one of the least affected regions, the region still suffered extinctions, particularly around the Late Pleistocene-Holocene transition. These extinctions were likely predominantly climatically driven by changes to grassland habitats.[17]

South Asia, Southeast Asia and East Asia edit

 
Giant tapir (Tapirus augustus) restoration
 
Life-sized models of Stegodon
 
Palaeoloxodon namadicus fossil at Indian Museum, Kolkata, India
 
Fossil jaw (Xiahe mandible) of a denisovan

The timing of extinctions on the Indian subcontinent is uncertain due to a lack of reliable dating.[19] Similar issues have been reported for Chinese sites, though there is no evidence for any of the megafaunal taxa having survived into the Holocene in that region.[20] Extinctions in Southeast Asia and South China have been proposed to be the result of environmental shift from open to closed forested habitats.[21]

Europe and northern Asia edit

 
Saiga antelope (Saiga spp.) inhabited a range from England and France to Yukon in the Late Pleistocene, diversifying into two species. S. borealis is now extinct and the critically endangered S. tatarica is now limited to the steppe in Kazakhstan and Mongolia
 
Hippopotamuses (Hippopotamus spp.) inhabited Great Britain until 80,000 BCE, whence due to glacial shifts, hippopotamuses were restricted to southeastern Europe, Mediterranean islands and finally western Asia until 1,000 BCE
 
Reconstruction of the five phenotypes of Pleistocene wild horse. The coat colours and dimensions are based on genetic evidence and historic descriptions
 
Elasmotherium sibiricum reconstruction
 
Cave paintings of the woolly rhinoceros (Coelodonta antiquitatis) in Chauvet-Pont-d'Arc Cave, France
 
The 'Gallery of Lions', representations of the Eurasian cave lion in Chauvet-Pont-d'Arc Cave, France
 
The leopard (Panthera pardus) inhabited the entire expanse of Afro-Eurasia below the 54th parallel north, from modern day Spain and the UK in the west, to South Africa in the south, and Siberia, Japan and Sundaland in the east during the Late Pleistocene
 
Cave bear (Ursus spelaeus) reconstruction
 
The woolly mammoth became extinct around 10,000 BCE – except for diminutive relict populations on St. Paul Island and Wrangel Island, which humans did not colonise until 3,600 BCE and 2,000 BCE respectively
 
Model of the European straight-tusked elephant (Paleoloxodon antiquus)

The Palearctic realm spans the entirety of the European continent and stretches into northern Asia, through the Caucasus and central Asia to northern China, Siberia and Beringia. Extinctions were more severe in Northern Eurasia than in Africa or South and Southeast Asia. These extinctions were staggered over tens of thousands of years, spanning from around 50,000 years Before Present (BP) to around 10,000 years BP, with temperate adapted species like the straight-tusked elephant and the narrow-nosed rhinoceros generally going extinct earlier than cold adapted species like the woolly mammoth and woolly rhinoceros. Climate change has been considered a probable major factor in the extinctions, possibly in combination with human hunting.[2]

See also: List of extinct animals of Europe

North America edit

See also: List of North American animals extinct in the Holocene

Extinctions in North America were concentrated at the end of the Late Pleistocene, around 13,800–11,400 years Before Present, which were coincident with the onset of the Younger Dryas cooling period, as well as the emergence of the hunter-gatherer Clovis culture. The relative importance of human and climactic factors in the North American extinctions has been the subject of significant controversy. Extinctions totalled around 35 genera.[4] The radiocarbon record for North America south of the Alaska-Yukon region has been described as "inadequate" to construct a reliable chronology.[51]

 
Long-horned/Giant bison (Bos latifrons), fossil bison skeleton (public display, Cincinnati Museum of Natural History & Science, Cincinnati, Ohio, United States)
 
Mounted skeleton of a shrub-ox (Euceratherium collinum)
 
Life restoration of Cervalces scotti
 
Tetrameryx shuleri restoration
 
A Chacoan peccary (Catagonus wagneri), believed to be the closest surviving relative of the extinct Platygonus
 
Western camel (Camelops hesternus) reconstruction
 
Life restoration of the Yukon horse (Equus lambei)
 
Reconstruction of Mixotoxodon larenis, a toxodontid notoungulate
 
Saber-toothed cat (Smilodon fatalis) reconstruction
 
Scimitar cat (Homotherium serum) reconstruction
 
American lion (Panthera atrox) reconstruction
 
The dhole (Cuon alpinus), now restricted to the southern portions of Asia, was present from Iberia to Mexico during the Late Pleistocene
 
Giant short-faced bear (Arctodus simus) reconstruction
 
American mastodon (Mammut americanum) reconstruction
 
Columbian mammoth (Mammuthus columbi) reconstruction
 
Giant beaver (Castoroides ohioensis) skeleton displayed at the Field Museum of Natural History, Chicago, Illinois, United States
 
Skull of Paralouatta marianae, one of the two Cuban members of the extinct Antilles monkeys (Xenotrichini)
 
Eremotherium laurillardi skeleton displayed at the Houston Museum of Natural Science
 
Life restoration of Nothrotheriops texanus
 
Glyptotherium reconstruction
 
Californian turkey (Meleagris californica) and megafaunal Californian condor (Gymnogyps amplus) fossil displays at La Brea Tar Pits
 
Teratornis merriami skeleton from the La Brea Tar Pits in flight pose
 
Reconstruction of the Cuban giant owl (Ornimegalonyx oteroi), of Pleistocene Cuba, with the carcass of a large solenodon

North American extinctions (noted as herbivores (H) or carnivores (C)) included:

The survivors are in some ways as significant as the losses: bison (H), grey wolf (C), lynx (C), grizzly bear (C), American black bear (C), deer (e.g. caribou, moose, wapiti (elk), Odocoileus spp.) (H), pronghorn (H), white-lipped peccary (H), muskox (H), bighorn sheep (H), and mountain goat (H); the list of survivors also include species which were extirpated during the Quaternary extinction event, but recolonised at least part of their ranges during the mid-Holocene from South American relict populations, such as the cougar (C), jaguar (C), giant anteater (C), collared peccary (H), ocelot (C) and jaguarundi (C). All save the pronghorns and giant anteaters were descended from Asian ancestors that had evolved with human predators.[77] Pronghorns are the second-fastest land mammal (after the cheetah), which may have helped them elude hunters. More difficult to explain in the context of overkill is the survival of bison, since these animals first appeared in North America less than 240,000 years ago and so were geographically removed from human predators for a sizeable period of time.[78][79][80] Because ancient bison evolved into living bison,[81][82] there was no continent-wide extinction of bison at the end of the Pleistocene (although the genus was regionally extirpated in many areas). The survival of bison into the Holocene and recent times is therefore inconsistent with the overkill scenario. By the end of the Pleistocene, when humans first entered North America, these large animals had been geographically separated from intensive human hunting for more than 200,000 years. Given this enormous span of geologic time, bison would almost certainly have been very nearly as naive as native North American large mammals.

The culture that has been connected with the wave of extinctions in North America is the paleo-American culture associated with the Clovis people (q.v.), who were thought to use spear throwers to kill large animals. The chief criticism of the "prehistoric overkill hypothesis" has been that the human population at the time was too small and/or not sufficiently widespread geographically to have been capable of such ecologically significant impacts. This criticism does not mean that climate change scenarios explaining the extinction are automatically to be preferred by default, however, any more than weaknesses in climate change arguments can be taken as supporting overkill. Some form of a combination of both factors could be plausible, and overkill would be a lot easier to achieve large-scale extinction with an already stressed population due to climate change.

South America edit

 
Fossil of Hippidion, a genus of horse native to South America.
 
Reconstruction of the mother and calf of Macrauchenia, a member of the extinct order Litopterna
 
Skeleton of Toxodon , a member of the extinct order Notoungulata
 
Reconstruction of the Dire wolf (Aenocyon dirus)
 
Life restoration of Arctotherium bonariense
 
Reconstruction of the gomphothere Cuvieronius
 
Skeleton of the giant ground sloth Megatherium
 
Reconstruction of the glytodont Doedicurus clavicaudatus , distributed in the temperate savannah and woodland of South America
 
Fossil reconstruction of Panochthus frenzelianus with metal model
 
Fossil of Smilodon populator
See also: List of South American animals extinct in the Holocene

South America suffered among the worst losses of the continents, with around 83% of its megafauna going extinct.[10] Extinctions are thought to have occurred in the interval 13,000–10,000 years Before Present, coincident with the end of the Antarctic Cold Reversal (a cooling period earlier and less severe than the Northern Hemisphere Younger Dryas) and the emergence of Fishtail projectile points, which became widespread across South America. Fishtail projectile points are thought to have been used in big game hunting, though direct evidence of exploitation of extinct megafauna by humans is rare. Fishtail points rapidly disappeared after the extinction of the megafauna, and were replaced by other styles more suited to hunting smaller prey. Humans have traditionally been less cited as a causal factor in the extinctions than in North America, though some recent scholarship is beginning to challenge this.[83] Rock art shows that humans coexisted with South America's megafauna,[84] a finding also confirmed by anthropogenic interaction with megafaunal remains.[85]

Sahul (Australia-New Guinea) and the Pacific edit

See also: List of Australia-New Guinea species extinct in the Holocene, List of New Zealand species extinct in the Holocene, and List of Oceanian animals extinct in the Holocene
 
Reconstruction of a hippopotamus-sized Diprotodon
 
Some of the marsupial lions were the largest mammalian predators in Australia of their time
 
Reconstruction of Zygomaturus

A scarcity of reliably dated megafaunal bone deposits has made it difficult to construct timelines for megafaunal extinctions in certain areas, leading to a divide among researches about when and how megafaunal species went extinct.[117][118]

There are at least three hypotheses regarding the extinction of the Australian megafauna:

  1. that they went extinct with the arrival of the Aboriginal Australians on the continent,
  2. that they went extinct due to natural climate change.

This theory is based on evidence of megafauna surviving until 40,000 years ago, a full 30,000 years after homo sapiens first landed in Australia, and thus that the two groups coexisted for a long time. Evidence of these animals existing at that time come from fossil records and ocean sediment. To begin with, sediment core drilled in the Indian Ocean off the SW coast of Australia indicate the existence of a fungus called Sporormiella, which survived off the dung of plant-eating mammals. The abundance of these spores in the sediment prior to 45,000 years ago indicates that many large mammals existed in the southwest Australian landscape until that point. The sediment data also indicates that the megafauna population collapsed within a few thousand years, around the 45,000 years ago, suggesting a rapid extinction event.[119] In addition, fossils found at South Walker Creek, which is the youngest megafauna site in northern Australia, indicate that at least 16 species of megafauna survived there until 40,000 years ago. Furthermore, there is no firm evidence of homo sapiens living at South Walker Creek 40,000 years ago, therefore no human cause can be attributed to the extinction of these megafauna. However, there is evidence of major environmental deterioration of South Water Creek 40,000 years ago, which may have caused the extinct event. These changes include increased fire, reduction in grasslands, and the loss of fresh water.[120] The same environmental deterioration is seen across Australia at the time, further strengthening the climate change argument. Australia's climate at the time could best be described as an overall drying of the landscape due to lower precipitation, resulting in less fresh water availability and more drought conditions. Overall, this led to changes in vegetation, increased fires, overall reduction in grasslands, and a greater competition for already scarce fresh water.[121] These environmental changes proved to be too much for the Australian megafauna to cope with, causing the extinction of 90% of megafauna species.

  1. The third hypothesis shared by some scientists is that human impacts and natural climate changes led to the extinction of Australian megafauna. About 75% of Australia is semi-arid or arid, so it makes sense that megafauna species used the same fresh water resources as humans. This competition could have led to more hunting of megafauna.[122] Furthermore, Homo sapiens used fire agriculture[clarification needed] to burn impassable[clarification needed] land. This further diminished the already disappearing grassland which contained plants that were a key dietary component of herbivorous megafauna. While there is no scientific consensus on this, it is plausible that homo sapiens and natural climate change had a combined impact. Overall, there is a great deal of evidence for humans being the culprit, but by ruling out climate change completely as a cause of the Australian megafauna extinction we are not getting the whole picture. The climate change in Australia 45,000 years ago destabilized the ecosystem, making it particularly vulnerable to hunting and fire agriculture by humans; this is probably what led to the extinction of the Australian megafauna.

Several studies provide evidence that climate change caused megafaunal extinction during the Pleistocene in Australia. One group of researchers analyzed fossilized teeth found at Cuddie Springs in southeastern Australia. By analyzing oxygen isotopes, they measured aridity, and by analyzing carbon isotopes and dental microwear texture analysis, they assessed megafaunal diets and vegetation. During the middle Pleistocene, southeastern Australia was dominated by browsers, including fauna that consumed C4 plants. By the late Pleistocene, the C4 plant dietary component had decreased considerably. This shift may have been caused by increasingly arid conditions, which may have caused dietary restrictions. Other isotopic analyses of eggshells and wombat teeth also point to a decline of C4 vegetation after 45 Ka. This decline in C4 vegetation is coincident with increasing aridity. Increasingly arid conditions in southeastern Australia during the late Pleistocene may have stressed megafauna, and contributed to their decline.[123]

 
Procoptodon goliath reconstruction
 
The American flamingo (Phoenicopterus ruber) was one of four species of flamingo present in Australia in the Quaternary, all of which are now either extinct or extirpated. Australia is now the only inhabited continent in the world without flamingoes.
 
Megalania skeleton, Melbourne Museum

In Sahul (a former continent composed of Australia and New Guinea), the sudden and extensive spate of extinctions occurred earlier than in the rest of the world.[124][125][126][127] Most evidence points to a 20,000 year period after human arrival circa 63,000 BCE,[128] but scientific argument continues as to the exact date range.[129] In the rest of the Pacific (other Australasian islands such as New Caledonia, and Oceania) although in some respects far later, endemic fauna also usually perished quickly upon the arrival of humans in the late Pleistocene and early Holocene.

Relationship to later extinctions edit

Main article: Holocene extinction
See also: Timeline of extinctions in the Holocene

There is no general agreement on where the Quaternary extinction event ends, and the Holocene, or anthropogenic, extinction begins, or if they should be considered separate events at all.[136][137] Some have suggested that anthropogenic extinctions may have begun as early as when the first modern humans spread out of Africa between 100,000 and 200,000 years ago, which is supported by rapid megafaunal extinction following recent human colonisation in Australia, New Zealand and Madagascar,[138] in a similar way that any large, adaptable predator moving into a new ecosystem would. In many cases, it is suggested even minimal hunting pressure was enough to wipe out large fauna, particularly on geographically isolated islands.[139][140] Only during the most recent parts of the extinction have plants also suffered large losses.[141]

Overall, the Holocene extinction can be characterised by the human impact on the environment. The Holocene extinction continues into the 21st century, with overfishing, ocean acidification and the amphibian crisis being a few broader examples of an almost universal, cosmopolitan decline of biodiversity.

Causes edit

Human Activity - Hunting edit

The hunting hypothesis suggests that humans hunted megaherbivores to extinction, which in turn caused the extinction of carnivores and scavengers which had preyed upon those animals.[142][143][144] This hypothesis holds Pleistocene humans responsible for the megafaunal extinction. One variant, known as blitzkrieg, portrays this process as relatively quick. Some of the direct evidence for this includes: fossils of some megafauna found in conjunction with human remains, embedded arrows and tool cut marks found in megafaunal bones, and European cave paintings that depict such hunting. Biogeographical evidence is also suggestive: the areas of the world where humans evolved currently have more of their Pleistocene megafaunal diversity (the elephants and rhinos of Asia and Africa) compared to other areas such as Australia, the Americas, Madagascar and New Zealand without the earliest humans. The overkill hypothesis, a variant of the hunting hypothesis, was proposed in 1966 by Paul S. Martin,[145] Professor of Geosciences Emeritus at the Desert Laboratory of the University of Arizona.[146]

 
Despeciation within the genus Homo.
 
Known H. sapiens migration routes in the Pleistocene.

Circumstantially, the close correlation in time between the appearance of humans in an area and extinction there provides weight for this scenario.[147][9][3] Radiocarbon dating has supported the plausibility of this correlation being reflective of causation.[148] The megafaunal extinctions covered a vast period of time and highly variable climatic situations. The earliest extinctions in Australia were complete approximately 50,000 BP, well before the Last Glacial Maximum and before rises in temperature. The most recent extinction in New Zealand was complete no earlier than 500 BP and during a period of cooling. In between these extremes megafaunal extinctions have occurred progressively in such places as North America, South America and Madagascar with no climatic commonality. The only common factor that can be ascertained is the arrival of humans.[149][150] This phenomenon appears even within regions. The mammal extinction wave in Australia about 50,000 years ago coincides not with known climatic changes, but with the arrival of humans. In addition, large mammal species like the giant kangaroo Protemnodon appear to have succumbed sooner on the Australian mainland than on Tasmania, which was colonised by humans a few thousand years later.[151][152] A study published in 2015 supported the hypothesis further by running several thousand scenarios that correlated the time windows in which each species is known to have become extinct with the arrival of humans on different continents or islands. This was compared against climate reconstructions for the last 90,000 years. The researchers found correlations of human spread and species extinction indicating that the human impact was the main cause of the extinction, while climate change exacerbated the frequency of extinctions. The study, however, found an apparently low extinction rate in the fossil record of mainland Asia.[153][154] A 2020 study published in Science Advances found that human population size and/or specific human activities, not climate change, caused rapidly rising global mammal extinction rates during the past 126,000 years. Around 96% of all mammalian extinctions over this time period are attributable to human impacts. According to Tobias Andermann, lead author of the study, "these extinctions did not happen continuously and at constant pace. Instead, bursts of extinctions are detected across different continents at times when humans first reached them. More recently, the magnitude of human driven extinctions has picked up the pace again, this time on a global scale."[155][156] On a related note, the population declines of still extant megafauna during the Pleistocene have also been shown to correlate with human expansion rather than climate change.[14]

The extinction's extreme bias towards larger animals further supports a relationship with human activity rather than climate change.[157] There is evidence that the average size of mammalian fauna declined over the course of the Quaternary,[158] a phenomenon that was likely linked to disproportionate hunting of large animals by humans.[5]

Extinction through human hunting has been supported by archaeological finds of mammoths with projectile points embedded in their skeletons, by observations of modern naive animals allowing hunters to approach easily[159][160][161] and by computer models by Mosimann and Martin,[162] and Whittington and Dyke,[163] and most recently by Alroy.[164]

 
The timing of extinctions follows the "March of Man"

Major objections have been raised regarding the hunting hypothesis. Notable among them is the sparsity of evidence of human hunting of megafauna.[165][166][167] There is no archeological evidence that in North America megafauna other than mammoths, mastodons, gomphotheres and bison were hunted, despite the fact that, for example, camels and horses are very frequently reported in fossil history.[168] Overkill proponents, however, say this is due to the fast extinction process in North America and the low probability of animals with signs of butchery to be preserved.[169] The majority of North American taxa have too sparse a fossil record to accurately assess the frequency of human hunting of them.[10] A study by Surovell and Grund concluded "archaeological sites dating to the time of the coexistence of humans and extinct fauna are rare. Those that preserve bone are considerably more rare, and of those, only a very few show unambiguous evidence of human hunting of any type of prey whatsoever."[170] Eugene S. Hunn points out that the birthrate in hunter-gatherer societies is generally too low, that too much effort is involved in the bringing down of a large animal by a hunting party, and that in order for hunter-gatherers to have brought about the extinction of megafauna simply by hunting them to death, an extraordinary amount of meat would have had to have been wasted.[171]

Second-order predation edit

 
Combination Hypotheses: Climate Change, Overkill + Climate Change, Second-Order Predation + Climate Change
 
Overkill Hypothesis and Second-Order Predation

The Second-Order Predation Hypothesis says that as humans entered the New World they continued their policy of killing predators, which had been successful in the Old World but because they were more efficient and because the fauna, both herbivores and carnivores, were more naive, they killed off enough carnivores to upset the ecological balance of the continent, causing overpopulation, environmental exhaustion, and environmental collapse. The hypothesis accounts for changes in animal, plant, and human populations.

The scenario is as follows:

  • After the arrival of H. sapiens in the New World, existing predators must share the prey populations with this new predator. Because of this competition, populations of original, or first-order, predators cannot find enough food; they are in direct competition with humans.
  • Second-order predation begins as humans begin to kill predators.
  • Prey populations are no longer well controlled by predation. Killing of nonhuman predators by H. sapiens reduces their numbers to a point where these predators no longer regulate the size of the prey populations.
  • Lack of regulation by first-order predators triggers boom-and-bust cycles in prey populations. Prey populations expand and consequently overgraze and over-browse the land. Soon the environment is no longer able to support them. As a result, many herbivores starve. Species that rely on the slowest recruiting food become extinct, followed by species that cannot extract the maximum benefit from every bit of their food.
  • Boom-bust cycles in herbivore populations change the nature of the vegetative environment, with consequent climatic impacts on relative humidity and continentality. Through overgrazing and overbrowsing, mixed parkland becomes grassland, and climatic continentality increases.

The second-order predation hypothesis has been supported by a computer model, the Pleistocene extinction model (PEM), which, using the same assumptions and values for all variables (herbivore population, herbivore recruitment rates, food needed per human, herbivore hunting rates, etc.) other than those for hunting of predators. It compares the overkill hypothesis (predator hunting = 0) with second-order predation (predator hunting varied between 0.01 and 0.05 for different runs). The findings are that second-order predation is more consistent with extinction than is overkill[172][173] (results graph at left). The Pleistocene extinction model is the only test of multiple hypotheses and is the only model to specifically test combination hypotheses by artificially introducing sufficient climate change to cause extinction. When overkill and climate change are combined they balance each other out. Climate change reduces the number of plants, overkill removes animals, therefore fewer plants are eaten. Second-order predation combined with climate change exacerbates the effect of climate change.[174] (results graph at right). The second-order predation hypothesis is further supported by the observation above that there was a massive increase in bison populations.[175]

However, this hypothesis has been criticised on the grounds that the multispecies model produces a mass extinction through indirect competition between herbivore species: small species with high reproductive rates subsidize predation on large species with low reproductive rates.[164] All prey species are lumped in the Pleistocene extinction model. Also, the control of population sizes by predators is not fully supported by observations of modern ecosystems.[176] The hypothesis further assumes decreases in vegetation due to climate change, but deglaciation doubled the habitable area of North America. Any vegetational changes that did occur failed to cause almost any extinctions of small vertebrates, and they are more narrowly distributed on average, which detractors cite as evidence against the hypothesis.

Competition for water edit

In southeastern Australia, the scarcity of water during the interval in which humans arrived in Australia suggests that human competition with megafauna for precious water sources may have played a role in the extinction of the latter.[122]

Landscape alteration edit

One consequence of the colonisation by humans of lands previously uninhabited by them may have been the introduction of new fire regimes because of extensive fire use by humans.[7] There is evidence that anthropogenic fire use had major impacts on the local environments in both Australia[6] and North America.[177]

Climate change edit

At the end of the 19th and beginning of the 20th centuries, when scientists first realized that there had been glacial and interglacial ages, and that they were somehow associated with the prevalence or disappearance of certain animals, they surmised that the termination of the Pleistocene ice age might be an explanation for the extinctions.

The most obvious change associated with the termination of an ice age is the increase in temperature. Between 15,000 BP and 10,000 BP, a 6 °C increase in global mean annual temperatures occurred. This was generally thought to be the cause of the extinctions. According to this hypothesis, a temperature increase sufficient to melt the Wisconsin ice sheet could have placed enough thermal stress on cold-adapted mammals to cause them to die. Their heavy fur, which helps conserve body heat in the glacial cold, might have prevented the dumping of excess heat, causing the mammals to die of heat exhaustion. Large mammals, with their reduced surface area-to-volume ratio, would have fared worse than small mammals. A study covering the past 56,000 years indicates that rapid warming events with temperature changes of up to 16 °C (29 °F) had an important impact on the extinction of megafauna. Ancient DNA and radiocarbon data indicates that local genetic populations were replaced by others within the same species or by others within the same genus. Survival of populations was dependent on the existence of refugia and long distance dispersals, which may have been disrupted by human hunters.[178]

Other scientists have proposed that increasingly extreme weather—hotter summers and colder winters—referred to as "continentality", or related changes in rainfall caused the extinctions. It has been shown that vegetation changed from mixed woodland-parkland to separate prairie and woodland.[179][180][181] This may have affected the kinds of food available. Shorter growing seasons may have caused the extinction of large herbivores and the dwarfing of many others. In this case, as observed, bison and other large ruminants would have fared better than horses, elephants and other monogastrics, because ruminants are able to extract more nutrition from limited quantities of high-fiber food and better able to deal with anti-herbivory toxins.[182][183][184] So, in general, when vegetation becomes more specialized, herbivores with less diet flexibility may be less able to find the mix of vegetation they need to sustain life and reproduce, within a given area. Increased continentality resulted in reduced and less predictable rainfall limiting the availability of plants necessary for energy and nutrition.[185][186][187] It has been suggested that this change in rainfall restricted the amount of time favorable for reproduction.[188][189] This could disproportionately harm large animals, since they have longer, more inflexible mating periods, and so may have produced young at unfavorable seasons (i.e., when sufficient food, water, or shelter was unavailable because of shifts in the growing season). In contrast, small mammals, with their shorter life cycles, shorter reproductive cycles, and shorter gestation periods, could have adjusted to the increased unpredictability of the climate, both as individuals and as species which allowed them to synchronize their reproductive efforts with conditions favorable for offspring survival. If so, smaller mammals would have lost fewer offspring and would have been better able to repeat the reproductive effort when circumstances once more favored offspring survival.[190] A study looking at the environmental conditions across Europe, Siberia and the Americas from 25,000 to 10,000 YBP found that prolonged warming events leading to deglaciation and maximum rainfall occurred just prior to the transformation of the rangelands that supported megaherbivores into widespread wetlands that supported herbivore-resistant plants. The study proposes that moisture-driven environmental change led to the megafaunal extinctions and that Africa's trans-equatorial position allowed rangeland to continue to exist between the deserts and the central forests, therefore fewer megafauna species became extinct there.[178]

Evidence in Southeast Asia, in contrast to Europe, Australia, and the Americas, suggests that climate change and an increasing sea level were significant factors in the extinction of several herbivorous species. Alterations in vegetation growth and new access routes for early humans and mammals to previously isolated, localized ecosystems were detrimental to select groups of fauna.[191]

Some evidence from Europe also suggests climatic changes were responsible for extinctions there, as the individuals extinctions tended to occur during times of environmental change and did not correlate particularly well with human migrations.[2]

In Australia, some studies have suggested that extinctions of megafauna began before the peopling of the continent, favouring climate change as the driver.[192]

In Beringia, megafauna may have gone extinct because of particularly intense paludification and because the land connection between Eurasia and North America flooded before the Cordilleran Ice Sheet retreated far enough to reopen the corridor between Beringia and the remainder of North America.[193] Woolly mammoths became extirpated from Beringia because of climatic factors, although human activity also played a synergistic role in their decline.[194] In North America, a Radiocarbon-dated Event-Count (REC) modelling study found that megafaunal declines in North America correlated with climatic changes instead of human population expansion.[195]

In the North American Great Lakes region, the population declines of mastodons and mammoths have been found to correlate with climatic fluctuations during the Younger Dryas rather than human activity.[196]

In the Argentine Pampas, the flooding of vast swathes of the once much larger Pampas grasslands may have played a role in the extinctions of its megafaunal assemblages.[8]

Critics object that since there were multiple glacial advances and withdrawals in the evolutionary history of many of the megafauna, it is rather implausible that only after the last glacial maximum would there be such extinctions. Proponents of climate change as the extinction event's cause like David J. Meltzer suggest that the last deglaciation may have been markedly different from previous ones.[197] Also, one study suggests that the Pleistocene megafaunal composition may have differed markedly from that of earlier interglacials, making the Pleistocene populations particularly vulnerable to changes in their environment.[198]

Studies propose that the annual mean temperature of the current interglacial that we have seen for the last 10,000 years is no higher than that of previous interglacials, yet most of the same large mammals survived similar temperature increases.[199][200][201] In addition, numerous species such as mammoths on Wrangel Island and St. Paul Island survived in human-free refugia despite changes in climate.[202] This would not be expected if climate change were responsible (unless their maritime climates offered some protection against climate change not afforded to coastal populations on the mainland). Under normal ecological assumptions island populations should be more vulnerable to extinction due to climate change because of small populations and an inability to migrate to more favorable climes.[citation needed]

Critics have also identified a number of problems with the continentality hypotheses. Megaherbivores have prospered at other times of continental climate. For example, megaherbivores thrived in Pleistocene Siberia, which had and has a more continental climate than Pleistocene or modern (post-Pleistocene, interglacial) North America.[203][204][205] The animals that became extinct actually should have prospered during the shift from mixed woodland-parkland to prairie, because their primary food source, grass, was increasing rather than decreasing.[206][205][207] Although the vegetation did become more spatially specialized, the amount of prairie and grass available increased, which would have been good for horses and for mammoths, and yet they became extinct. This criticism ignores the increased abundance and broad geographic extent of Pleistocene bison at the end of the Pleistocene, which would have increased competition for these resources in a manner not seen in any earlier interglacials.[198] Although horses became extinct in the New World, they were successfully reintroduced by the Spanish in the 16th century—into a modern post-Pleistocene, interglacial climate. Today there are feral horses still living in those same environments. They find a sufficient mix of food to avoid toxins, they extract enough nutrition from forage to reproduce effectively and the timing of their gestation is not an issue. Of course, this criticism ignores the obvious fact that present-day horses are not competing for resources with ground sloths, mammoths, mastodons, camels, llamas, and bison. Similarly, mammoths survived the Pleistocene Holocene transition on isolated, uninhabited islands in the Mediterranean Sea until 4,000 to 7,000 years ago,[208] as well as on Wrangel Island in the Siberian Arctic.[209] Additionally, large mammals should have been able to migrate, permanently or seasonally, if they found the temperature too extreme, the breeding season too short, or the rainfall too sparse or unpredictable.[210] Seasons vary geographically. By migrating away from the equator, herbivores could have found areas with growing seasons more favorable for finding food and breeding successfully. Modern-day African elephants migrate during periods of drought to places where there is apt to be water.[211] Large animals also store more fat in their bodies than do medium-sized animals and this should have allowed them to compensate for extreme seasonal fluctuations in food availability.[212]

Some evidence weighs against climate change as a valid hypothesis as applied to Australia. It has been shown that the prevailing climate at the time of extinction (40,000–50,000 BP) was similar to that of today, and that the extinct animals were strongly adapted to an arid climate. The evidence indicates that all of the extinctions took place in the same short time period, which was the time when humans entered the landscape. The main mechanism for extinction was probably fire (started by humans) in a then much less fire-adapted landscape. Isotopic evidence shows sudden changes in the diet of surviving species, which could correspond to the stress they experienced before extinction.[213][214][215]

Some evidence obtained from analysis of the tusks of mastodons from the American Great Lakes region appears inconsistent with the climate change hypothesis. Over a span of several thousand years prior to their extinction in the area, the mastodons show a trend of declining age at maturation. This is the opposite of what one would expect if they were experiencing stresses from deteriorating environmental conditions, but is consistent with a reduction in intraspecific competition that would result from a population being reduced by human hunting.[216]

It may be observed that neither the overkill nor the climate change hypotheses can fully explain events: browsers, mixed feeders and non-ruminant grazer species suffered most, while relatively more ruminant grazers survived.[217] However, a broader variation of the overkill hypothesis may predict this, because changes in vegetation wrought by either Second Order Predation (see below)[174][218] or anthropogenic fire preferentially selects against browse species.[citation needed]

Disease edit

The hyperdisease hypothesis, as advanced by Ross D. E. MacFee and Preston A. Marx, attributes the extinction of large mammals during the late Pleistocene to indirect effects of the newly arrived aboriginal humans.[219][220][221] In more recent times, disease has driven many vulnerable species to extinction; the introduction of avian malaria and avipoxvirus, for example, has greatly decreased the populations of the endemic birds of Hawaii, with some going extinct.[222] The hyperdisease hypothesis proposes that humans or animals traveling with them (e.g., chickens or domestic dogs) introduced one or more highly virulent diseases into vulnerable populations of native mammals, eventually causing extinctions. The extinction was biased toward larger-sized species because smaller species have greater resilience because of their life history traits (e.g., shorter gestation time, greater population sizes, etc.). Humans are thought to be the cause because other earlier immigrations of mammals into North America from Eurasia did not cause extinctions.[219] A similar suggestion is that pathogens were transmitted by the expanding humans via the domesticated dogs they brought with them.[223] A related theory proposes that a highly contagious prion disease similar to chronic wasting disease or scrapie that was capable of infecting a large number of species was the culprit. Animals weakened by this "superprion" would also have easily become reservoirs of viral and bacterial diseases as they succumbed to neurological degeneration from the prion, causing a cascade of different diseases to spread among various mammal species. This theory could potentially explain the prevalence of heterozygosity at codon 129 of the prion protein gene in humans, which has been speculated to be the result of natural selection against homozygous genotypes that were more susceptible to prion disease and thus potentially a tell-tale of a major prion pandemic that affected humans of or younger than reproductive age far in the past and disproportionately killed before they could reproduce those with homozygous genotypes at codon 129.[224]

If a disease was indeed responsible for the end-Pleistocene extinctions, then there are several criteria it must satisfy (see Table 7.3 in MacPhee & Marx 1997). First, the pathogen must have a stable carrier state in a reservoir species. That is, it must be able to sustain itself in the environment when there are no susceptible hosts available to infect. Second, the pathogen must have a high infection rate, such that it is able to infect virtually all individuals of all ages and sexes encountered. Third, it must be extremely lethal, with a mortality rate of c. 50–75%. Finally, it must have the ability to infect multiple host species without posing a serious threat to humans. Humans may be infected, but the disease must not be highly lethal or able to cause an epidemic.[citation needed]

As with other hypotheses, a number of counterarguments to the hyperdisease hypothesis have been put forth. Generally speaking, disease has to be very virulent to kill off all the individuals in a genus or species. Even such a virulent disease as West Nile fever is unlikely to have caused extinction.[225] The disease would need to be implausibly selective while being simultaneously implausibly broad. Such a disease needs to be capable of killing off wolves such as Canis dirus or goats such as Oreamnos harringtoni while leaving other very similar species (Canis lupus and Oreamnos americanus, respectively) unaffected. It would need to be capable of killing off flightless birds while leaving closely related flighted species unaffected. Yet while remaining sufficiently selective to afflict only individual species within genera it must be capable of fatally infecting across such clades as birds, marsupials, placentals, testudines, and crocodilians. No disease with such a broad scope of fatal infectivity is known, much less one that remains simultaneously incapable of infecting numerous closely related species within those disparate clades. On the other hand, this objection does not account for the possibility of a variety of different diseases being introduced around the same era.[citation needed] Numerous species including wolves, mammoths, camelids, and horses had emigrated continually between Asia and North America over the past 100,000 years. For the disease hypothesis to be applicable there it would require that the population remain immunologically naive despite this constant transmission of genetic and pathogenic material.[citation needed] The dog-specific hypothesis in particular cannot account for several major extinction events, notably the Americas (for reasons already covered) and Australia. Dogs did not arrive in Australia until approximately 35,000 years after the first humans arrived there, and approximately 30,000 years after the Australian megafaunal extinction was complete.[citation needed]

Extraterrestrial impact edit

An extraterrestrial impact, which has occasionally been proposed as a cause of the Younger Dryas,[226] is sometimes suggested as a potential cause of the extinction of North America's megafauna due to the temporal proximity between a proposed date for such an impact and the following megafaunal extinctions.[227][4] However, the Younger Dryas impact hypothesis is widely considered a fringe hypothesis that is unsupported by evidence.[228][229]

Geomagnetic field weakening edit

A weakening of the Earth's magnetic field may have caused increased flux of UV-B radiation and has been suggested as a cause of megafaunal extinctions in the Late Quaternary.[230]

Effects edit

The extinction of the megafauna could have caused the disappearance of the mammoth steppe rather than the other way around. Alaska now has low nutrient soil unable to support bison, mammoths, and horses. R. Dale Guthrie has claimed this as a cause of the extinction of the megafauna there; however, he may be interpreting it backwards. The loss of large herbivores to break up the permafrost allows the cold soils that are unable to support large herbivores today. Today, in the arctic, where trucks have broken the permafrost grasses and diverse flora and fauna can be supported.[231][232] In addition, Chapin (Chapin 1980) showed that simply adding fertilizer to the soil in Alaska could make grasses grow again like they did in the era of the mammoth steppe. Possibly, the extinction of the megafauna and the corresponding loss of dung is what led to low nutrient levels in modern-day soil and therefore is why the landscape can no longer support megafauna.

Megafauna play a significant role in the lateral transport of mineral nutrients in an ecosystem, tending to translocate them from areas of high to those of lower abundance. They do so by their movement between the time they consume the nutrient and the time they release it through elimination (or, to a much lesser extent, through decomposition after death).[233] 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.[234][235] 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.[235] The extinction of the mammoths allowed grasslands they had maintained through grazing habits to become birch forests.[236] The new forest and the resulting forest fires may have induced climate change.[236] Such disappearances might be the result of the proliferation of modern humans.[237][238]

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,[239] contributing to the warmer climate of the time (up to 10 °C warmer than at present).[239][240] 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.[239]

Recent studies have indicated that the extinction of megafaunal herbivores may have caused a reduction in atmospheric methane. 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.[241] 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 2–4 times more rapid than any other decrease in the last half million years, suggesting that an unusual mechanism was at work.[242]

The extermination of megafauna left many niches vacant, which has been cited as an explanation for the vulnerability and fragility of many ecosystems to destruction in the later Holocene extinction. The comparative lack of megafauna in modern ecosystems has reduced high-order interactions among surviving species, reducing ecological complexity.[243] This depauperate, post-megafaunal ecological state has been associated with diminished ecological resilience to stressors.[244] Many extant species of plants have adaptations that were advantageous in the presence of megafauna but are now useless in their absence.[245] The demise of megafaunal ecosystem engineers in the Arctic that maintained open grassland environments has been highly detrimental to shorebirds of the genus Numenius.[246]

Human perception edit

Analysis of Maori oral traditions has found that the extinction of New Zealand's megafauna during the Holocene extinction had significant social and linguistic impacts on their culture, suggesting that similar phenomena may have occurred in other societies that lived through megafaunal extinctions.[247]

See also edit

References edit

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February 16, 2024
late, pleistocene, extinctions, extinctions, that, occurred, during, holocene, holocene, extinction, this, article, multiple, issues, please, help, improve, discuss, these, issues, talk, page, learn, when, remove, these, template, messages, this, page, unbalan. For extinctions that occurred during the Holocene see Holocene extinction This article has multiple issues Please help improve it or discuss these issues on the talk page Learn how and when to remove these template messages This page may be unbalanced towards certain viewpoints Please improve the article by adding information on neglected viewpoints or discuss the issue on the talk page January 2023 This article may contain excessive or irrelevant examples Please help improve the article by adding descriptive text and removing less pertinent examples March 2023 This article needs attention from an expert in Palaeontology See the talk page for details WikiProject Palaeontology may be able to help recruit an expert March 2023 Learn how and when to remove this template message The Late Pleistocene to the beginning of the Holocene saw numerous extinctions of predominantly megafaunal typically defined as having body masses over 44 kilograms 97 lb 1 animal species the Pleistocene megafauna which resulted in a collapse in faunal density and diversity across the globe 2 The extinctions during the Late Pleistocene are differentiated from previous extinctions by the widespread absence of ecological succession to replace these extinct megafaunal species 3 and the regime shift of previously established faunal relationships and habitats as a consequence The timing and severity of the extinctions varied by region and are thought to have been driven by varying combinations of human and climatic factors 3 Human impact on megafauna populations is thought to have been driven by hunting overkill 4 5 as well as possibly environmental alteration 6 The relative importance of human vs climatic factors in the extinctions has been the subject of long running controversy 3 Late Pleistocene landscape of northern Spain by Mauricio Anton From left to right wild horse Equus ferus woolly mammoth Mammuthus primigenius reindeer Rangifer tarandus the cave or steppe lion Panthera spelaea and the woolly rhinoceros Coelodonta antiquitatis Mural of La Brea Tar Pits by Charles R Knight including sabertooth cats Smilodon fatalis left ground sloths Paramylodon harlani right and Columbian mammoths Mammuthus columbi background Major extinctions occurred in Australia New Guinea Sahul beginning approximately 50 000 years ago and in the Americas about 13 000 years ago coinciding in time with the early human migrations into these regions 7 Extinctions in northern Eurasia were staggered over tens of thousands of years between 50 000 and 10 000 years ago 2 while extinctions in the Americas were virtually simultaneous spanning only 3000 years at most 4 8 Overall during Late Pleistocene about 65 of all megafaunal species worldwide became extinct 9 rising to 72 in North America 83 in South America and 88 in Australia 10 with all mammals over 1 000 kilograms 2 200 lb becoming extinct in Australia and the Americas 1 Africa South Asia and Southeast Asia experienced much lower extinctions than other regions 10 Contents 1 Extinctions by biogeographic realm 1 1 Summary 1 2 Introduction 1 3 Africa 1 4 South Asia Southeast Asia and East Asia 1 5 Europe and northern Asia 1 6 North America 1 7 South America 1 8 Sahul Australia New Guinea and the Pacific 2 Relationship to later extinctions 3 Causes 3 1 Human Activity Hunting 3 2 Second order predation 3 3 Competition for water 3 4 Landscape alteration 3 5 Climate change 3 6 Disease 3 7 Extraterrestrial impact 3 8 Geomagnetic field weakening 4 Effects 5 Human perception 6 See also 7 References 8 External links 8 1 Hyperdisease hypothesis 8 2 Second order predation 8 3 Other linksExtinctions by biogeographic realm editSummary edit Extinctions range of the continental large and medium sized mammals from 40 000 to 4 000 years BP in different biogeographic realms 11 Biogeographic realm Giants over 1 000 kg Very large 400 1 000 kg Large 150 400 kg Moderately large 50 150 kg Medium 10 50 kg Total Regions includedStart Loss Start Loss Start Loss Start Loss Start Loss Start Loss Afrotropic 6 1 16 6 4 1 25 25 3 12 32 0 0 69 2 2 9 136 7 5 1 Trans Saharan Africa and ArabiaIndomalaya 5 2 40 6 1 16 7 10 1 10 20 3 15 56 1 1 8 97 8 8 2 Indian subcontinent Southeast Asia and southern ChinaPalearctic 8 8 100 10 5 50 14 5 35 7 23 3 15 41 1 2 4 96 22 22 9 Eurasia and North AfricaNearctic 5 5 100 10 8 80 26 22 84 6 20 13 65 25 9 36 86 57 66 North AmericaNeotropic 9 9 100 12 12 100 17 14 82 20 11 55 35 5 14 3 93 51 54 South America Central America South Florida and the CaribbeanAustralasia 4 4 100 5 5 100 6 6 100 16 13 81 2 25 10 40 56 38 67 Australia New Guinea New Zealand and neighbouring islands Global 33 26 78 8 46 31 67 4 86 47 54 7 113 41 36 3 215 23 10 1 493 168 34 Introduction edit nbsp The proportion of extinct large mammal species more than or equal to 10 kg 22 lb in each country during the last 132 000 years only counting extinctions earlier than 1000 years BPThe Late Pleistocene saw the extinction of many mammals weighing more than 40 kilograms 88 lb The proportion of megafauna extinctions is progressively larger the further the human migratory distance from Africa with the highest extinction rates in Australia and North and South America The increased extent of extinction mirrors the migration pattern of modern humans the further away from Africa the more recently humans inhabited the area the less time those environments including its megafauna had to become accustomed to humans and vice versa There are two main hypotheses to explain this extinction Climate change associated with the advance and retreat of major ice caps or ice sheets causing reduction in favorable habitat Human hunting causing attrition of megafauna populations commonly known as overkill 12 There are some inconsistencies between the current available data and the prehistoric overkill hypothesis For instance there are ambiguities around the timing of Australian megafauna extinctions 12 Evidence supporting the prehistoric overkill hypothesis includes the persistence of megafauna on some islands for millennia past the disappearance of their continental cousins For instance ground sloths survived on the Antilles long after North and South American ground sloths were extinct woolly mammoths died out on remote Wrangel Island 6 000 years after their extinction on the mainland while Steller s sea cows persisted off the isolated and uninhabited Commander Islands for thousands of years after they had vanished from the continental shores of the north Pacific 13 The later disappearance of these island species correlates with the later colonization of these islands by humans The original debates as to whether human arrival times or climate change constituted the primary cause of megafaunal extinctions necessarily were based on paleontological evidence coupled with geological dating techniques Recently genetic analyses of surviving megafaunal populations have contributed new evidence leading to the conclusion The inability of climate to predict the observed population decline of megafauna especially during the past 75 000 years implies that human impact became the main driver of megafauna dynamics around this date 14 An alternative hypothesis to the theory of human responsibility is climate change associated with the last glacial period Discredited explanations include the Younger Dryas impact hypothesis 15 and Tollmann s hypothesis that extinctions resulted from bolide impacts Recent research indicates that each species responded differently to environmental changes and no one factor by itself explains the large variety of extinctions The causes may involve the interplay of climate change competition between species unstable population dynamics and human predation 16 Africa edit Although Africa was one of the least affected regions the region still suffered extinctions particularly around the Late Pleistocene Holocene transition These extinctions were likely predominantly climatically driven by changes to grassland habitats 17 Ungulates Even Toed Ungulates Suidae swine Metridiochoerus Kolpochoerus Bovidae bovines antelope Giant buffalo Syncerus antiquus Megalotragus Rusingoryx Antidorcas australis Antidorcas bondi Damaliscus hypsodon Damaliscus niro Gazella atlantica Gazella tingitana Caprinae Makapania Odd toed Ungulates Rhinoceros Rhinocerotidae Stephanorhinus hemitoechus North Africa Ceratotherium mauritanicum Wild Equus spp Caballine horses Equus algericus North Africa Subgenus Asinus asses Equus melkiensis North Africa Zebras Equus capensis Saharan zebra 18 Equus mauritanicus Proboscidea Elephantidae elephants Palaeoloxodon iolensis other authors suggest that this taxon went extinct at the end of the Middle Pleistocene Rodentia Paraethomys filfilae South Asia Southeast Asia and East Asia edit nbsp Giant tapir Tapirus augustus restoration nbsp Life sized models of Stegodon nbsp Palaeoloxodon namadicus fossil at Indian Museum Kolkata India nbsp Fossil jaw Xiahe mandible of a denisovanThe timing of extinctions on the Indian subcontinent is uncertain due to a lack of reliable dating 19 Similar issues have been reported for Chinese sites though there is no evidence for any of the megafaunal taxa having survived into the Holocene in that region 20 Extinctions in Southeast Asia and South China have been proposed to be the result of environmental shift from open to closed forested habitats 21 Ungulates Even Toed Ungulates Several Bovidae spp Bos palaesondaicus ancestor to the banteng Bison hanaizumiensis 22 23 24 Cebu tamaraw Bubalus cebuensis Bubalus grovesi 25 Bubalus wansijocki Short horned water buffalo Bubalus mephistopheles Bubalus palaeokerabau Cervidae Sinomegaceros spp including Sinomegaceros yabei in Japan and Sinomegaceros ordosianus and possibly Sinomegaceros pachyosteus in China 26 Hippopotamidae Hexaprotodon Indian subcontinent and Southeast Asia 27 Odd toed Ungulates Equus spp Equus namadicus Indian subcontinent Yunnan horse Equus yunanensis Giant tapir Tapirus augustus Southeast Asia and Southern China Merck s rhinoceros Stephanorhinus kirchbergensis Eastern Asia Pholidota Giant Asian pangolin Manis palaeojavanica Carnivora Caniformia Arctoidea Bears Ailuropoda baconi ancestor to the giant panda Afrotheria Afroinsectiphilia Orycteropodidae Tubulidentata Aardvark Orycteropus afer extirpated in South Asia circa 13 000 BCE 28 29 Paenungulata Tethytheria Proboscideans Stegodontidae Stegodon spp including Stegodon florensis on Flores Stegodon orientalis in East and Southeast Asia and Stegodon sp in the Indian subcontinent Elephantidae Palaeoloxodon spp Palaeoloxodon namadicus Indian subcontinent possibly also Southeast Asia Palaeoloxodon naumanni Japan Palaeoloxodon huaihoensis China Birds Japanese flightless duck Shiriyanetta hasegawai 30 Leptoptilos robustus Reptiles Crocodilia Alligator munensis Primates Several simian Simiiformes spp Pongo orangutans Pongo weidenreichi South China Various Homo spp archaic humans Homo erectus soloensis Java Homo floresiensis Flores Homo luzonensis Luzon Philippines Denisovans Homo sp Europe and northern Asia edit nbsp Saiga antelope Saiga spp inhabited a range from England and France to Yukon in the Late Pleistocene diversifying into two species S borealis is now extinct and the critically endangered S tatarica is now limited to the steppe in Kazakhstan and Mongolia nbsp Hippopotamuses Hippopotamus spp inhabited Great Britain until 80 000 BCE whence due to glacial shifts hippopotamuses were restricted to southeastern Europe Mediterranean islands and finally western Asia until 1 000 BCE nbsp Reconstruction of the five phenotypes of Pleistocene wild horse The coat colours and dimensions are based on genetic evidence and historic descriptions nbsp Elasmotherium sibiricum reconstruction nbsp Cave paintings of the woolly rhinoceros Coelodonta antiquitatis in Chauvet Pont d Arc Cave France nbsp The Gallery of Lions representations of the Eurasian cave lion in Chauvet Pont d Arc Cave France nbsp The leopard Panthera pardus inhabited the entire expanse of Afro Eurasia below the 54th parallel north from modern day Spain and the UK in the west to South Africa in the south and Siberia Japan and Sundaland in the east during the Late Pleistocene nbsp Cave bear Ursus spelaeus reconstruction nbsp The woolly mammoth became extinct around 10 000 BCE except for diminutive relict populations on St Paul Island and Wrangel Island which humans did not colonise until 3 600 BCE and 2 000 BCE respectively nbsp Model of the European straight tusked elephant Paleoloxodon antiquus The Palearctic realm spans the entirety of the European continent and stretches into northern Asia through the Caucasus and central Asia to northern China Siberia and Beringia Extinctions were more severe in Northern Eurasia than in Africa or South and Southeast Asia These extinctions were staggered over tens of thousands of years spanning from around 50 000 years Before Present BP to around 10 000 years BP with temperate adapted species like the straight tusked elephant and the narrow nosed rhinoceros generally going extinct earlier than cold adapted species like the woolly mammoth and woolly rhinoceros Climate change has been considered a probable major factor in the extinctions possibly in combination with human hunting 2 See also List of extinct animals of Europe Ungulates Even Toed Hoofed Mammals Various Bovidae spp Steppe bison Bison priscus Baikal yak Bos baikalensis 31 European water buffalo Bubalus murrensis European tahr Hemitragus cedrensis 32 33 Giant muskox Praeovibos priscus 34 Northern saiga antelope Saiga borealis 35 Twisted horned antelope Spirocerus kiakhtensis 36 37 Goat horned antelope Parabubalis capricornis 36 37 Various deer Cervidae spp Broad fronted moose Cervalces latifrons Giant deer Megaloceros giganteus Praemegaceros savini Cretan deer Candiacervus Haploidoceros mediterraneus 38 39 All native Hippopotamus spp 40 Hippopotamus amphibius European range Maltese dwarf hippopotamus Hippopotamus melitensis Cyprus dwarf hippopotamus Hippopotamus minor Sicilian dwarf hippopotamus Hippopotamus pentlandi Camelus knoblochi 41 and other Camelus spp Odd Toed Hoofed Mammals Various Equus spp e g Wild horse Equus ferus ssp Equus cf gallicus 42 43 European wild ass Equus hydruntinus Equus cf latipes 37 42 44 Equus lenensis 37 45 Equus cf uralensis 42 All native Rhinoceros Rhinocerotidae spp Elasmotherium Woolly rhinoceros Coelodonta antiquitatis Stephanorhinus spp Merck s rhinoceros Stephanorhinus kirchbergensis Narrow nosed rhinoceros Stephanorhinus hemiotoechus Carnivora Caniformia Canidae Caninae Wolves Cave wolf Canis lupus spelaeus Dire wolf Aenocyon dirus 46 Dholes European dhole Cuon alpinus europaeus Sardinian dhole Cynotherium sardous Arctoidea Various Ursus spp Steppe brown bear Ursus arctos priscus 47 Gamssulzen cave bear Ursus ingressus 48 Pleistocene small cave bear Ursus rossicus Cave bear Ursus spelaeus Giant polar bear Ursus maritimus tyrannus Musteloidea Mustelidae Several otter Lutrinae spp Robust Pleistocene European otter Cyrnaonyx Algarolutra Sardinian giant otter Megalenhydris barbaricina Sardinian dwarf otter Sardolutra Cretan otter Lutrogale cretensis Feliformia Various Felidae spp Eurasian scimitar cat Homotherium latidens Cave lynx Lynx pardinus spelaeus 49 Issoire lynx Lynx issiodorensis Panthera spp Cave lion Panthera spelaea European ice age leopard Panthera pardus spelaea Herpestoidea Cave hyena Crocuta crocuta spelaea hyaena prisca All native Elephant Elephantidae spp Mammoths Woolly mammoth Mammuthus primigenius Dwarf Sardinian mammoth Mammuthus lamarmorai Straight tusked elephant Palaeoloxodon antiquus Europe Dwarf elephant Palaeoloxodon creutzburgi Crete Cyprus dwarf elephant Palaeoloxodon cypriotes Palaeoloxodon mnaidriensis Sicily Rodents Allocricetus bursae Cricetus major alternatively Cricetus cricetus major Dicrostonyx gulielmi ancestor to the Arctic lemming Giant Eurasian porcupine Hystrix refossa Leithia spp Maltese and Sicilian giant dormouse 50 Marmota paleocaucasica Microtus grafi Mimomys spp M pyrenaicus M chandolensis Pliomys lenki Spermophilus citelloides Spermophilus severskensis Spermophilus superciliosus Trogontherium cuiveri Lagomorpha Lepus tanaiticus alternatively Lepus timidus tanaiticus Pika Ochotona spp e g Giant pika Ochotona whartoni Tonomochota spp T khasanensis T sikhotana T major Birds Asian ostrich Struthio asiaticus Yakutian goose Anser djuktaiensis Various European crane spp Genus Grus Grus primigenia Grus melitensis Cretan owl Athene cretensis Primates Homo Denisovans Homo sp Neanderthals Homo sapiens neanderthalensis survived until about 40 000 years ago on the Iberian peninsula Reptiles Solitudo sicula survived in Sicily until about 12 500 years ago Lacerta siculimelitensis from Malta North America edit See also List of North American animals extinct in the HoloceneExtinctions in North America were concentrated at the end of the Late Pleistocene around 13 800 11 400 years Before Present which were coincident with the onset of the Younger Dryas cooling period as well as the emergence of the hunter gatherer Clovis culture The relative importance of human and climactic factors in the North American extinctions has been the subject of significant controversy Extinctions totalled around 35 genera 4 The radiocarbon record for North America south of the Alaska Yukon region has been described as inadequate to construct a reliable chronology 51 nbsp Long horned Giant bison Bos latifrons fossil bison skeleton public display Cincinnati Museum of Natural History amp Science Cincinnati Ohio United States nbsp Mounted skeleton of a shrub ox Euceratherium collinum nbsp Life restoration of Cervalces scotti nbsp Tetrameryx shuleri restoration nbsp A Chacoan peccary Catagonus wagneri believed to be the closest surviving relative of the extinct Platygonus nbsp Western camel Camelops hesternus reconstruction nbsp Life restoration of the Yukon horse Equus lambei nbsp Reconstruction of Mixotoxodon larenis a toxodontid notoungulate nbsp Saber toothed cat Smilodon fatalis reconstruction nbsp Scimitar cat Homotherium serum reconstruction nbsp American lion Panthera atrox reconstruction nbsp The dhole Cuon alpinus now restricted to the southern portions of Asia was present from Iberia to Mexico during the Late Pleistocene nbsp Giant short faced bear Arctodus simus reconstruction nbsp American mastodon Mammut americanum reconstruction nbsp Columbian mammoth Mammuthus columbi reconstruction nbsp Giant beaver Castoroides ohioensis skeleton displayed at the Field Museum of Natural History Chicago Illinois United States nbsp Skull of Paralouatta marianae one of the two Cuban members of the extinct Antilles monkeys Xenotrichini nbsp Eremotherium laurillardi skeleton displayed at the Houston Museum of Natural Science nbsp Life restoration of Nothrotheriops texanus nbsp Glyptotherium reconstruction nbsp Californian turkey Meleagris californica and megafaunal Californian condor Gymnogyps amplus fossil displays at La Brea Tar Pits nbsp Teratornis merriami skeleton from the La Brea Tar Pits in flight pose nbsp Reconstruction of the Cuban giant owl Ornimegalonyx oteroi of Pleistocene Cuba with the carcass of a large solenodonNorth American extinctions noted as herbivores H or carnivores C included Ungulates Even Toed Hoofed Mammals Various Bovidae spp Most forms of Pleistocene bison only Bison bison in North America and Bison bonasus in Eurasia survived Ancient bison Bison antiquus H Long horned Giant bison Bison latifrons H Steppe bison Bison priscus H Bison occidentalis H Several members of Caprinae the muskox survived Giant muskox Praeovibos priscus H Shrub ox Euceratherium collinum H Harlan s muskox Bootherium bombifrons H Soergel s ox Soergelia mayfieldi H Harrington s mountain goat Oreamnos harringtoni smaller and more southern distribution than its surviving relative H Saiga antelope Saiga tatarica extirpated H Deer Stag moose Cervalces scotti H American mountain deer Odocoileus lucasi H Torontoceros hypnogeos H Various Antilocapridae genera pronghorns survived Capromeryx H Stockoceros H Tetrameryx H Pacific pronghorn Antilocapra pacifica H Several peccary Tayassuidae spp Flat headed peccary Platygonus H Long nosed peccary Mylohyus H Collared peccary Dicotyles tajacu extirpated range semi recolonised H Muknalia minimus is a junior synonym Various members of Camelidae Western camel Camelops hesternus H Stilt legged llamas Hemiauchenia ssp H Stout legged llamas Palaeolama ssp H Odd Toed Hoofed Mammals All native forms of Equidae Caballine true horses Equus cf ferus from the Late Pleistocene of North America have historically been assigned to many different species including Equus fraternus Equus scotti and Equus lambei but the taxonomy of these horses is unclear and many of these species may be synonymous with each other perhaps only representing a single species 52 53 54 Stilt legged horse Haringtonhippus francisci Equus francisci H Tapirs Tapirus three species California tapir Tapirus californicus H Merriam s tapir Tapirus merriami H Vero tapir Tapirus veroensis H Order Notoungulata Mixotoxodon 55 56 H Carnivora Feliformia Several Felidae spp Saber Tooths North American saber toothed cat Smilodon fatalis C North American scimitar cat Homotherium serum C American cheetah Miracinonyx not true cheetah Miracinonyx trumani C Cougar Puma concolor megafaunal ecomorph extirpated from North America South American populations recolonised former range C Jaguarundi Herpailurus yagouaroundi extirpated range semi recolonised C Margay Leopardus weidii extirpated C Ocelot Leopardus pardalis extirpated range marginally recolonised C Eurasian lynx Lynx lynx extirpated 57 C Jaguars Pleistocene North American jaguar Panthera onca augusta range semi recolonised by other subspecies C North America Jaguar Panthera balamoides dubious suggested to be a junior synonym of the short faced bear Arctotherium Lions American lion Panthera atrox endemic to North America after 340 000 BP C Cave steppe lion Panthera spelaea present only as far as modern day Yukon C Caniformia Canidae Dire wolf Aenocyon dirus C Pleistocene coyote Canis latrans orcutti C Megafaunal wolf e g Beringian wolf Canis lupus ssp C Dhole Cuon alpinus extirpated C Protocyon troglodytes 58 C Arctoidea Musteloidea Mephitidae Short faced skunk Brachyprotoma obtusata 59 C Mustelidae Steppe polecat Mustela eversmanii extirpated 60 C Various bear Ursidae spp Arctodus simus C Florida spectacled bear Tremarctos floridanus C South American short faced bear Arctotherium wingei 61 58 C Giant polar bear Ursus maritimus tyrannus a possible inhabitant C Afrotheria Afroinsectiphilia Orycteropodidae Tubulidentata Giant anteater Myrmecophaga tridactyla extirpated range partially recolonised 62 63 C Paenungulata Tethytheria All native spp of Proboscidea Mastodons American mastodon Mammut americanum H Pacific mastodon Mammut pacificus H validity uncertain Gomphotheriidae spp Cuvieronius 64 H Mammoth Mammuthus spp Columbian mammoth Mammuthus columbi H Pygmy mammoth Mammuthus exilis H Woolly mammoth Mammuthus primigenius H Sirenia Dugongidae Steller s sea cow Hydrodamalis gigas extirpated from North America survived in Beringia into 18th century H Euarchontoglires Bats Stock s vampire bat Desmodus stocki C Pristine mustached bat Pteronotus Phyllodia pristinus C Rodents Giant beaver Castoroides spp Castoroides ohioensis H Castoroides leiseyorum H Klein s porcupine Erethizon kleini 65 H Giant island deer mouse Peromyscus nesodytes C Neochoerus spp e g Pinckney s capybara Neochoerus pinckneyi H Neochoerus aesopi H Neotoma findleyi Neotoma pygmaea Synaptomys australis All giant hutia Heptaxodontidae spp Blunt toothed giant hutia Amblyrhiza inundata could grow as large as an American black bear H Plate toothed giant hutia Elasmodontomys obliquus H Twisted toothed mouse Quemisia gravis H Osborn s key mouse Clidomys osborn s H Xaymaca fulvopulvis H Lagomorphs Aztlan rabbit Aztlanolagus sp H Giant pika Ochotona whartoni H Eulipotyphla Notiosorex dalquesti Notiosorex harrisi Xenarthrana All remaining ground sloth spp Eremotherium megatheriid giant ground sloth H Nothrotheriops nothrotheriid ground sloth H Megalonychid ground sloth spp Megalonyx H Nohochichak 66 67 H Xibalbaonyx 68 69 H Meizonyx Mylodontid ground sloth spp Paramylodon H All members of Glyptodontidae Glyptotherium 70 H H Beautiful armadillo Dasypus bellus 71 H Pachyarmatherium All Pampatheriidae spp Holmesina H Pampatherium H Birds Water Fowl Ducks Bermuda flightless duck Anas pachyscelus H Californian flightless sea duck Chendytes lawi C Mexican stiff tailed duck Oxyura zapatima 61 H Turkey Meleagris spp Californian turkey Meleagris californica H Meleagris crassipes 61 H Various Gruiformes spp All cave rail Nesotrochis spp e g Antillean cave rail Nesotrochis debooyi C Barbados rail Incertae sedis C Cuban flightless crane Antigone cubensis H La Brea crane Grus pagei H Various flamingo Phoenicopteridae spp Minute flamingo Phoenicopterus minutus 72 C Cope s flamingo Phoenicopterus copei 73 C Dow s puffin Fratercula dowi C Pleistocene Mexican diver spp Plyolimbus baryosteus C Podiceps spp Podiceps parvus 61 C Storks La Brea Asphalt stork Ciconia maltha 61 C Wetmore s stork Mycteria wetmorei 61 C Pleistocene Mexican cormorants spp genus Phalacrocorax 61 Phalacrocorax goletensis C Phalacrocorax chapalensis C All remaining teratorn Teratornithidae spp Aiolornis incredibilis C Cathartornis gracilis C Oscaravis olsoni C Teratornis merriami C Teratornis woodburnensis C Several New World vultures Cathartidae spp Pleistocene black vulture Coragyps occidentalis ssp C Megafaunal Californian condor Gymnogyps amplus C Clark s condor Breagyps clarki C Cuban condor Gymnogyps varonai C Several Accipitridae spp American neophrone vulture Neophrontops americanus 61 74 C Woodward s eagle Amplibuteo woodwardi C Cuban great hawk Buteogallus borrasi C Daggett s eagle Buteogallus daggetti C Fragile eagle Buteogallus fragilis C Cuban giant hawk Gigantohierax suarezi 75 76 C Errant eagle Neogyps errans C Grinnell s crested eagle Spizaetus grinnelli 61 C Willett s hawk eagle Spizaetus willetti 61 C Caribbean titan hawk Titanohierax C Several owl Strigiformes spp Brea miniature owl Asphaltoglaux C Kurochkin s pygmy owl Glaucidium kurochkini C Brea owl Oraristix brea C Cuban giant owl Ornimegalonyx C Bermuda flicker Colaptes oceanicus C Several caracara Caracarinae spp Bahaman terrestrial caracara Caracara sp C Puerto Rican terrestrial caracara Caracara sp C Jamaican caracara Carcara tellustris C Cuban caracara Milvago sp C Hispaniolan caracara Milvago sp C Psittacopasserae Psittaciformes Mexican thick billed parrot Rhynchopsitta phillipsi 61 H Several giant tortoise spp Hesperotestudo H Gopherus spp Gopherus donlaloi H Chelonoidis spp Chelonoidis marcanoi H Chelonoidis alburyorum H The survivors are in some ways as significant as the losses bison H grey wolf C lynx C grizzly bear C American black bear C deer e g caribou moose wapiti elk Odocoileus spp H pronghorn H white lipped peccary H muskox H bighorn sheep H and mountain goat H the list of survivors also include species which were extirpated during the Quaternary extinction event but recolonised at least part of their ranges during the mid Holocene from South American relict populations such as the cougar C jaguar C giant anteater C collared peccary H ocelot C and jaguarundi C All save the pronghorns and giant anteaters were descended from Asian ancestors that had evolved with human predators 77 Pronghorns are the second fastest land mammal after the cheetah which may have helped them elude hunters More difficult to explain in the context of overkill is the survival of bison since these animals first appeared in North America less than 240 000 years ago and so were geographically removed from human predators for a sizeable period of time 78 79 80 Because ancient bison evolved into living bison 81 82 there was no continent wide extinction of bison at the end of the Pleistocene although the genus was regionally extirpated in many areas The survival of bison into the Holocene and recent times is therefore inconsistent with the overkill scenario By the end of the Pleistocene when humans first entered North America these large animals had been geographically separated from intensive human hunting for more than 200 000 years Given this enormous span of geologic time bison would almost certainly have been very nearly as naive as native North American large mammals The culture that has been connected with the wave of extinctions in North America is the paleo American culture associated with the Clovis people q v who were thought to use spear throwers to kill large animals The chief criticism of the prehistoric overkill hypothesis has been that the human population at the time was too small and or not sufficiently widespread geographically to have been capable of such ecologically significant impacts This criticism does not mean that climate change scenarios explaining the extinction are automatically to be preferred by default however any more than weaknesses in climate change arguments can be taken as supporting overkill Some form of a combination of both factors could be plausible and overkill would be a lot easier to achieve large scale extinction with an already stressed population due to climate change South America edit nbsp Fossil of Hippidion a genus of horse native to South America nbsp Reconstruction of the mother and calf of Macrauchenia a member of the extinct order Litopterna nbsp Skeleton of Toxodon a member of the extinct order Notoungulata nbsp Reconstruction of the Dire wolf Aenocyon dirus nbsp Life restoration of Arctotherium bonariense nbsp Reconstruction of the gomphothere Cuvieronius nbsp Skeleton of the giant ground sloth Megatherium nbsp Reconstruction of the glytodont Doedicurus clavicaudatus distributed in the temperate savannah and woodland of South America nbsp Fossil reconstruction of Panochthus frenzelianus with metal model nbsp Fossil of Smilodon populatorSee also List of South American animals extinct in the Holocene South America suffered among the worst losses of the continents with around 83 of its megafauna going extinct 10 Extinctions are thought to have occurred in the interval 13 000 10 000 years Before Present coincident with the end of the Antarctic Cold Reversal a cooling period earlier and less severe than the Northern Hemisphere Younger Dryas and the emergence of Fishtail projectile points which became widespread across South America Fishtail projectile points are thought to have been used in big game hunting though direct evidence of exploitation of extinct megafauna by humans is rare Fishtail points rapidly disappeared after the extinction of the megafauna and were replaced by other styles more suited to hunting smaller prey Humans have traditionally been less cited as a causal factor in the extinctions than in North America though some recent scholarship is beginning to challenge this 83 Rock art shows that humans coexisted with South America s megafauna 84 a finding also confirmed by anthropogenic interaction with megafaunal remains 85 Ungulates Even Toed Hoofed Mammals Several Cervidae spp Morenelaphus Antifer Agalmaceros blicki 86 87 Odocoileus salinae 88 89 Various Camelidae spp Eulamaops Stilt legged llama Hemiauchenia Stout legged llama Palaeolama Odd Toed Hoofed Mammals Several species of tapirs Tapiridae Tapirus rondoniensis Tapirus cristatellus All Pleistocene wild horse genera Equidae Equus neogeus 90 91 36 Hippidion 90 36 92 Hippidion devillei Hippidion principale Hippidion saldiasi All remaining Meridiungulata genera Order Litopterna Macraucheniidae Macrauchenia Macraucheniopsis 93 94 Xenorhinotherium Proterotheriidae Neolicaphrium recens 95 Order Notoungulata Toxodontidae Piauhytherium Some authors regard this taxon as synonym of Trigodonops Mixotoxodon Toxodon Trigodonops Carnivora Feliformia Several Felidae spp Saber toothed cat Smilodon spp 96 North American saber toothed cat Smilodon fatalis South American saber toothed cat Smilodon populator Patagonian jaguar Panthera onca mesembrina some authors have suggested that these remains actually belong to the American lion instead 97 Caniformia Canidae Dire wolf Aenocyon dirus Nehring s wolf Canis nehringi Protocyon spp 98 Protocyon trogolodytes Protocyon tarijense Dusicyon avus Pleistocene bush dog Speothos pacivorus Arctoidea South American short faced bear Arctotherium spp Arctotherium bonairense Arctotherium tarijense Arctotherium wingei Rodents Neochoerus Bats Giant vampire bat Desmodus draculae All remaining Gomphotheridae spp Cuvieronius Notiomastodon Xenarthrans All remaining ground sloth genera Megatheriidae spp Eremotherium Megatherium Nothrotheriidae spp Nothropus Nothrotherium Megalonychidae spp Ahytherium Australonyx Diabolotherium Megistonyx Valgipes 99 Mylodontidae spp Catonyx Glossotherium Lestodon Mylodon Scelidotherium Scelidodon Mylodonopsis Ocnotherium All remaining Glyptodontinae spp Doedicurus 100 101 102 103 Glyptodon Chlamydotherium 104 Heteroglyptodon 105 Hoplophorus Lomaphorus Neosclerocalyptus Neuryurus 104 106 Panochthus Parapanochthus 104 107 Plaxhaplous Sclerocalyptus Several Dasypodidae spp Beautiful armadillo Dasypus bellus Eutatus Pachyarmatherium Propaopus 40 89 All Pampatheriidae spp Holmesina et Chlamytherium occidentale 108 109 Pampatherium 110 Tonnicinctus 110 Birds Psilopterus small terror bird remains dated to the Late Pleistocene 111 112 but these are disputed 113 Various Caracarinae spp Venezuelan caracara Caracara major 114 Seymour s caracara Caracara seymouri 115 Peruvian caracara Milvago brodkorbi 116 Various Cathartidae spp Pampagyps imperator Geronogyps reliquus Wingegyps cartellei Pleistovultur nevesi Crocs amp Gators Caiman venezuelensis Chelonoidis lutzae Argentina Sahul Australia New Guinea and the Pacific edit See also List of Australia New Guinea species extinct in the Holocene List of New Zealand species extinct in the Holocene and List of Oceanian animals extinct in the Holocene nbsp Reconstruction of a hippopotamus sized Diprotodon nbsp Some of the marsupial lions were the largest mammalian predators in Australia of their time nbsp Reconstruction of ZygomaturusA scarcity of reliably dated megafaunal bone deposits has made it difficult to construct timelines for megafaunal extinctions in certain areas leading to a divide among researches about when and how megafaunal species went extinct 117 118 There are at least three hypotheses regarding the extinction of the Australian megafauna that they went extinct with the arrival of the Aboriginal Australians on the continent that they went extinct due to natural climate change This theory is based on evidence of megafauna surviving until 40 000 years ago a full 30 000 years after homo sapiens first landed in Australia and thus that the two groups coexisted for a long time Evidence of these animals existing at that time come from fossil records and ocean sediment To begin with sediment core drilled in the Indian Ocean off the SW coast of Australia indicate the existence of a fungus called Sporormiella which survived off the dung of plant eating mammals The abundance of these spores in the sediment prior to 45 000 years ago indicates that many large mammals existed in the southwest Australian landscape until that point The sediment data also indicates that the megafauna population collapsed within a few thousand years around the 45 000 years ago suggesting a rapid extinction event 119 In addition fossils found at South Walker Creek which is the youngest megafauna site in northern Australia indicate that at least 16 species of megafauna survived there until 40 000 years ago Furthermore there is no firm evidence of homo sapiens living at South Walker Creek 40 000 years ago therefore no human cause can be attributed to the extinction of these megafauna However there is evidence of major environmental deterioration of South Water Creek 40 000 years ago which may have caused the extinct event These changes include increased fire reduction in grasslands and the loss of fresh water 120 The same environmental deterioration is seen across Australia at the time further strengthening the climate change argument Australia s climate at the time could best be described as an overall drying of the landscape due to lower precipitation resulting in less fresh water availability and more drought conditions Overall this led to changes in vegetation increased fires overall reduction in grasslands and a greater competition for already scarce fresh water 121 These environmental changes proved to be too much for the Australian megafauna to cope with causing the extinction of 90 of megafauna species The third hypothesis shared by some scientists is that human impacts and natural climate changes led to the extinction of Australian megafauna About 75 of Australia is semi arid or arid so it makes sense that megafauna species used the same fresh water resources as humans This competition could have led to more hunting of megafauna 122 Furthermore Homo sapiens used fire agriculture clarification needed to burn impassable clarification needed land This further diminished the already disappearing grassland which contained plants that were a key dietary component of herbivorous megafauna While there is no scientific consensus on this it is plausible that homo sapiens and natural climate change had a combined impact Overall there is a great deal of evidence for humans being the culprit but by ruling out climate change completely as a cause of the Australian megafauna extinction we are not getting the whole picture The climate change in Australia 45 000 years ago destabilized the ecosystem making it particularly vulnerable to hunting and fire agriculture by humans this is probably what led to the extinction of the Australian megafauna Several studies provide evidence that climate change caused megafaunal extinction during the Pleistocene in Australia One group of researchers analyzed fossilized teeth found at Cuddie Springs in southeastern Australia By analyzing oxygen isotopes they measured aridity and by analyzing carbon isotopes and dental microwear texture analysis they assessed megafaunal diets and vegetation During the middle Pleistocene southeastern Australia was dominated by browsers including fauna that consumed C4 plants By the late Pleistocene the C4 plant dietary component had decreased considerably This shift may have been caused by increasingly arid conditions which may have caused dietary restrictions Other isotopic analyses of eggshells and wombat teeth also point to a decline of C4 vegetation after 45 Ka This decline in C4 vegetation is coincident with increasing aridity Increasingly arid conditions in southeastern Australia during the late Pleistocene may have stressed megafauna and contributed to their decline 123 nbsp Procoptodon goliath reconstruction nbsp The American flamingo Phoenicopterus ruber was one of four species of flamingo present in Australia in the Quaternary all of which are now either extinct or extirpated Australia is now the only inhabited continent in the world without flamingoes nbsp Megalania skeleton Melbourne MuseumIn Sahul a former continent composed of Australia and New Guinea the sudden and extensive spate of extinctions occurred earlier than in the rest of the world 124 125 126 127 Most evidence points to a 20 000 year period after human arrival circa 63 000 BCE 128 but scientific argument continues as to the exact date range 129 In the rest of the Pacific other Australasian islands such as New Caledonia and Oceania although in some respects far later endemic fauna also usually perished quickly upon the arrival of humans in the late Pleistocene and early Holocene Marsupials Various members of Diprotodontidae Diprotodon Hulitherium tomasetti Maokopia ronaldi Zygomaturus Palorchestes marsupial tapir Various members of Vombatidae Lasiorhinus angustidens giant wombat Phascolonus giant wombat Ramasayia magna giant wombat Vombatus hacketti Hackett s wombat Warendja wakefieldi dwarf wombat Sedophascolomys giant wombat Phascolarctos stirtoni giant koala Marsupial lion Thylacoleo carnifex Various members of Macropodidae Procoptodon short faced kangaroos e g Procoptodon goliah Sthenurus giant kangaroo Simosthenurus giant kangaroo Various Macropus giant kangaroo spp e g Macropus ferragus Macropus titan Macropus pearsoni Protemnodon giant wallaby Troposodon wallaby 124 125 130 131 132 133 Bohra giant tree kangaroo Propleopus oscillans omnivorous giant musky rat kangaroo Nombe Congruus Various forms of Sarcophilus Tasmanian devil Sarcophilus laniarius 25 larger than modern species unclear if it is actually a distinct species from living Tasmanian devil 134 Sarcophilus moornaensis Monotremes egg laying mammals Echidna Murrayglossus hacketti giant echidna Megalibgwilia ramsayi Birds Pygmy Cassowary Casuarius lydekkeri Genyornis a two meter tall 6 6 ft dromornithid Giant malleefowl Progura gallinacea Cryptogyps lacertosus Dynatoaetus gaffae Several Phoenicopteridae spp Xenorhynchopsis spp Australian flamingo 135 Xenorhynchopsis minor Xenorhynchopsis tibialis Reptiles Crocs amp Gators nbsp Quinkana was one of the last surviving land crocodilesIkanogavialis the last fully marine crocodilian Pallimnarchus Australian freshwater mekosuchine crocodiian Quinkana Australian terrestrial mekosuchine crocodilian apex predator Volia a two to three meter long mekosuchine crocodylian apex predator of Pleistocene Fiji Mekosuchus Mekosuchus inexpectatus New Caledonian land crocodile Mekosuchus kalpokasi Vanuatu land crocodile Varanus sp Pleistocene and Holocene New Caledonia Megalania Varanus pricus a giant predatory monitor lizard comparable or larger than the Komodo dragon Snakes Wonambi a five to six metre long Australian constrictor snake Several spp of Meiolaniidae giant armoured turtles Meiolania NinjemysRelationship to later extinctions editMain article Holocene extinction See also Timeline of extinctions in the Holocene There is no general agreement on where the Quaternary extinction event ends and the Holocene or anthropogenic extinction begins or if they should be considered separate events at all 136 137 Some have suggested that anthropogenic extinctions may have begun as early as when the first modern humans spread out of Africa between 100 000 and 200 000 years ago which is supported by rapid megafaunal extinction following recent human colonisation in Australia New Zealand and Madagascar 138 in a similar way that any large adaptable predator moving into a new ecosystem would In many cases it is suggested even minimal hunting pressure was enough to wipe out large fauna particularly on geographically isolated islands 139 140 Only during the most recent parts of the extinction have plants also suffered large losses 141 Overall the Holocene extinction can be characterised by the human impact on the environment The Holocene extinction continues into the 21st century with overfishing ocean acidification and the amphibian crisis being a few broader examples of an almost universal cosmopolitan decline of biodiversity Causes editHuman Activity Hunting edit The hunting hypothesis suggests that humans hunted megaherbivores to extinction which in turn caused the extinction of carnivores and scavengers which had preyed upon those animals 142 143 144 This hypothesis holds Pleistocene humans responsible for the megafaunal extinction One variant known as blitzkrieg portrays this process as relatively quick Some of the direct evidence for this includes fossils of some megafauna found in conjunction with human remains embedded arrows and tool cut marks found in megafaunal bones and European cave paintings that depict such hunting Biogeographical evidence is also suggestive the areas of the world where humans evolved currently have more of their Pleistocene megafaunal diversity the elephants and rhinos of Asia and Africa compared to other areas such as Australia the Americas Madagascar and New Zealand without the earliest humans The overkill hypothesis a variant of the hunting hypothesis was proposed in 1966 by Paul S Martin 145 Professor of Geosciences Emeritus at the Desert Laboratory of the University of Arizona 146 nbsp Despeciation within the genus Homo nbsp Known H sapiens migration routes in the Pleistocene Circumstantially the close correlation in time between the appearance of humans in an area and extinction there provides weight for this scenario 147 9 3 Radiocarbon dating has supported the plausibility of this correlation being reflective of causation 148 The megafaunal extinctions covered a vast period of time and highly variable climatic situations The earliest extinctions in Australia were complete approximately 50 000 BP well before the Last Glacial Maximum and before rises in temperature The most recent extinction in New Zealand was complete no earlier than 500 BP and during a period of cooling In between these extremes megafaunal extinctions have occurred progressively in such places as North America South America and Madagascar with no climatic commonality The only common factor that can be ascertained is the arrival of humans 149 150 This phenomenon appears even within regions The mammal extinction wave in Australia about 50 000 years ago coincides not with known climatic changes but with the arrival of humans In addition large mammal species like the giant kangaroo Protemnodon appear to have succumbed sooner on the Australian mainland than on Tasmania which was colonised by humans a few thousand years later 151 152 A study published in 2015 supported the hypothesis further by running several thousand scenarios that correlated the time windows in which each species is known to have become extinct with the arrival of humans on different continents or islands This was compared against climate reconstructions for the last 90 000 years The researchers found correlations of human spread and species extinction indicating that the human impact was the main cause of the extinction while climate change exacerbated the frequency of extinctions The study however found an apparently low extinction rate in the fossil record of mainland Asia 153 154 A 2020 study published in Science Advances found that human population size and or specific human activities not climate change caused rapidly rising global mammal extinction rates during the past 126 000 years Around 96 of all mammalian extinctions over this time period are attributable to human impacts According to Tobias Andermann lead author of the study these extinctions did not happen continuously and at constant pace Instead bursts of extinctions are detected across different continents at times when humans first reached them More recently the magnitude of human driven extinctions has picked up the pace again this time on a global scale 155 156 On a related note the population declines of still extant megafauna during the Pleistocene have also been shown to correlate with human expansion rather than climate change 14 The extinction s extreme bias towards larger animals further supports a relationship with human activity rather than climate change 157 There is evidence that the average size of mammalian fauna declined over the course of the Quaternary 158 a phenomenon that was likely linked to disproportionate hunting of large animals by humans 5 Extinction through human hunting has been supported by archaeological finds of mammoths with projectile points embedded in their skeletons by observations of modern naive animals allowing hunters to approach easily 159 160 161 and by computer models by Mosimann and Martin 162 and Whittington and Dyke 163 and most recently by Alroy 164 nbsp The timing of extinctions follows the March of Man Major objections have been raised regarding the hunting hypothesis Notable among them is the sparsity of evidence of human hunting of megafauna 165 166 167 There is no archeological evidence that in North America megafauna other than mammoths mastodons gomphotheres and bison were hunted despite the fact that for example camels and horses are very frequently reported in fossil history 168 Overkill proponents however say this is due to the fast extinction process in North America and the low probability of animals with signs of butchery to be preserved 169 The majority of North American taxa have too sparse a fossil record to accurately assess the frequency of human hunting of them 10 A study by Surovell and Grund concluded archaeological sites dating to the time of the coexistence of humans and extinct fauna are rare Those that preserve bone are considerably more rare and of those only a very few show unambiguous evidence of human hunting of any type of prey whatsoever 170 Eugene S Hunn points out that the birthrate in hunter gatherer societies is generally too low that too much effort is involved in the bringing down of a large animal by a hunting party and that in order for hunter gatherers to have brought about the extinction of megafauna simply by hunting them to death an extraordinary amount of meat would have had to have been wasted 171 Second order predation edit nbsp Combination Hypotheses Climate Change Overkill Climate Change Second Order Predation Climate Change nbsp Overkill Hypothesis and Second Order PredationThe Second Order Predation Hypothesis says that as humans entered the New World they continued their policy of killing predators which had been successful in the Old World but because they were more efficient and because the fauna both herbivores and carnivores were more naive they killed off enough carnivores to upset the ecological balance of the continent causing overpopulation environmental exhaustion and environmental collapse The hypothesis accounts for changes in animal plant and human populations The scenario is as follows After the arrival of H sapiens in the New World existing predators must share the prey populations with this new predator Because of this competition populations of original or first order predators cannot find enough food they are in direct competition with humans Second order predation begins as humans begin to kill predators Prey populations are no longer well controlled by predation Killing of nonhuman predators by H sapiens reduces their numbers to a point where these predators no longer regulate the size of the prey populations Lack of regulation by first order predators triggers boom and bust cycles in prey populations Prey populations expand and consequently overgraze and over browse the land Soon the environment is no longer able to support them As a result many herbivores starve Species that rely on the slowest recruiting food become extinct followed by species that cannot extract the maximum benefit from every bit of their food Boom bust cycles in herbivore populations change the nature of the vegetative environment with consequent climatic impacts on relative humidity and continentality Through overgrazing and overbrowsing mixed parkland becomes grassland and climatic continentality increases The second order predation hypothesis has been supported by a computer model the Pleistocene extinction model PEM which using the same assumptions and values for all variables herbivore population herbivore recruitment rates food needed per human herbivore hunting rates etc other than those for hunting of predators It compares the overkill hypothesis predator hunting 0 with second order predation predator hunting varied between 0 01 and 0 05 for different runs The findings are that second order predation is more consistent with extinction than is overkill 172 173 results graph at left The Pleistocene extinction model is the only test of multiple hypotheses and is the only model to specifically test combination hypotheses by artificially introducing sufficient climate change to cause extinction When overkill and climate change are combined they balance each other out Climate change reduces the number of plants overkill removes animals therefore fewer plants are eaten Second order predation combined with climate change exacerbates the effect of climate change 174 results graph at right The second order predation hypothesis is further supported by the observation above that there was a massive increase in bison populations 175 However this hypothesis has been criticised on the grounds that the multispecies model produces a mass extinction through indirect competition between herbivore species small species with high reproductive rates subsidize predation on large species with low reproductive rates 164 All prey species are lumped in the Pleistocene extinction model Also the control of population sizes by predators is not fully supported by observations of modern ecosystems 176 The hypothesis further assumes decreases in vegetation due to climate change but deglaciation doubled the habitable area of North America Any vegetational changes that did occur failed to cause almost any extinctions of small vertebrates and they are more narrowly distributed on average which detractors cite as evidence against the hypothesis Competition for water edit In southeastern Australia the scarcity of water during the interval in which humans arrived in Australia suggests that human competition with megafauna for precious water sources may have played a role in the extinction of the latter 122 Landscape alteration edit One consequence of the colonisation by humans of lands previously uninhabited by them may have been the introduction of new fire regimes because of extensive fire use by humans 7 There is evidence that anthropogenic fire use had major impacts on the local environments in both Australia 6 and North America 177 Climate change edit At the end of the 19th and beginning of the 20th centuries when scientists first realized that there had been glacial and interglacial ages and that they were somehow associated with the prevalence or disappearance of certain animals they surmised that the termination of the Pleistocene ice age might be an explanation for the extinctions The most obvious change associated with the termination of an ice age is the increase in temperature Between 15 000 BP and 10 000 BP a 6 C increase in global mean annual temperatures occurred This was generally thought to be the cause of the extinctions According to this hypothesis a temperature increase sufficient to melt the Wisconsin ice sheet could have placed enough thermal stress on cold adapted mammals to cause them to die Their heavy fur which helps conserve body heat in the glacial cold might have prevented the dumping of excess heat causing the mammals to die of heat exhaustion Large mammals with their reduced surface area to volume ratio would have fared worse than small mammals A study covering the past 56 000 years indicates that rapid warming events with temperature changes of up to 16 C 29 F had an important impact on the extinction of megafauna Ancient DNA and radiocarbon data indicates that local genetic populations were replaced by others within the same species or by others within the same genus Survival of populations was dependent on the existence of refugia and long distance dispersals which may have been disrupted by human hunters 178 Other scientists have proposed that increasingly extreme weather hotter summers and colder winters referred to as continentality or related changes in rainfall caused the extinctions It has been shown that vegetation changed from mixed woodland parkland to separate prairie and woodland 179 180 181 This may have affected the kinds of food available Shorter growing seasons may have caused the extinction of large herbivores and the dwarfing of many others In this case as observed bison and other large ruminants would have fared better than horses elephants and other monogastrics because ruminants are able to extract more nutrition from limited quantities of high fiber food and better able to deal with anti herbivory toxins 182 183 184 So in general when vegetation becomes more specialized herbivores with less diet flexibility may be less able to find the mix of vegetation they need to sustain life and reproduce within a given area Increased continentality resulted in reduced and less predictable rainfall limiting the availability of plants necessary for energy and nutrition 185 186 187 It has been suggested that this change in rainfall restricted the amount of time favorable for reproduction 188 189 This could disproportionately harm large animals since they have longer more inflexible mating periods and so may have produced young at unfavorable seasons i e when sufficient food water or shelter was unavailable because of shifts in the growing season In contrast small mammals with their shorter life cycles shorter reproductive cycles and shorter gestation periods could have adjusted to the increased unpredictability of the climate both as individuals and as species which allowed them to synchronize their reproductive efforts with conditions favorable for offspring survival If so smaller mammals would have lost fewer offspring and would have been better able to repeat the reproductive effort when circumstances once more favored offspring survival 190 A study looking at the environmental conditions across Europe Siberia and the Americas from 25 000 to 10 000 YBP found that prolonged warming events leading to deglaciation and maximum rainfall occurred just prior to the transformation of the rangelands that supported megaherbivores into widespread wetlands that supported herbivore resistant plants The study proposes that moisture driven environmental change led to the megafaunal extinctions and that Africa s trans equatorial position allowed rangeland to continue to exist between the deserts and the central forests therefore fewer megafauna species became extinct there 178 Evidence in Southeast Asia in contrast to Europe Australia and the Americas suggests that climate change and an increasing sea level were significant factors in the extinction of several herbivorous species Alterations in vegetation growth and new access routes for early humans and mammals to previously isolated localized ecosystems were detrimental to select groups of fauna 191 Some evidence from Europe also suggests climatic changes were responsible for extinctions there as the individuals extinctions tended to occur during times of environmental change and did not correlate particularly well with human migrations 2 In Australia some studies have suggested that extinctions of megafauna began before the peopling of the continent favouring climate change as the driver 192 In Beringia megafauna may have gone extinct because of particularly intense paludification and because the land connection between Eurasia and North America flooded before the Cordilleran Ice Sheet retreated far enough to reopen the corridor between Beringia and the remainder of North America 193 Woolly mammoths became extirpated from Beringia because of climatic factors although human activity also played a synergistic role in their decline 194 In North America a Radiocarbon dated Event Count REC modelling study found that megafaunal declines in North America correlated with climatic changes instead of human population expansion 195 In the North American Great Lakes region the population declines of mastodons and mammoths have been found to correlate with climatic fluctuations during the Younger Dryas rather than human activity 196 In the Argentine Pampas the flooding of vast swathes of the once much larger Pampas grasslands may have played a role in the extinctions of its megafaunal assemblages 8 Critics object that since there were multiple glacial advances and withdrawals in the evolutionary history of many of the megafauna it is rather implausible that only after the last glacial maximum would there be such extinctions Proponents of climate change as the extinction event s cause like David J Meltzer suggest that the last deglaciation may have been markedly different from previous ones 197 Also one study suggests that the Pleistocene megafaunal composition may have differed markedly from that of earlier interglacials making the Pleistocene populations particularly vulnerable to changes in their environment 198 Studies propose that the annual mean temperature of the current interglacial that we have seen for the last 10 000 years is no higher than that of previous interglacials yet most of the same large mammals survived similar temperature increases 199 200 201 In addition numerous species such as mammoths on Wrangel Island and St Paul Island survived in human free refugia despite changes in climate 202 This would not be expected if climate change were responsible unless their maritime climates offered some protection against climate change not afforded to coastal populations on the mainland Under normal ecological assumptions island populations should be more vulnerable to extinction due to climate change because of small populations and an inability to migrate to more favorable climes citation needed Critics have also identified a number of problems with the continentality hypotheses Megaherbivores have prospered at other times of continental climate For example megaherbivores thrived in Pleistocene Siberia which had and has a more continental climate than Pleistocene or modern post Pleistocene interglacial North America 203 204 205 The animals that became extinct actually should have prospered during the shift from mixed woodland parkland to prairie because their primary food source grass was increasing rather than decreasing 206 205 207 Although the vegetation did become more spatially specialized the amount of prairie and grass available increased which would have been good for horses and for mammoths and yet they became extinct This criticism ignores the increased abundance and broad geographic extent of Pleistocene bison at the end of the Pleistocene which would have increased competition for these resources in a manner not seen in any earlier interglacials 198 Although horses became extinct in the New World they were successfully reintroduced by the Spanish in the 16th century into a modern post Pleistocene interglacial climate Today there are feral horses still living in those same environments They find a sufficient mix of food to avoid toxins they extract enough nutrition from forage to reproduce effectively and the timing of their gestation is not an issue Of course this criticism ignores the obvious fact that present day horses are not competing for resources with ground sloths mammoths mastodons camels llamas and bison Similarly mammoths survived the Pleistocene Holocene transition on isolated uninhabited islands in the Mediterranean Sea until 4 000 to 7 000 years ago 208 as well as on Wrangel Island in the Siberian Arctic 209 Additionally large mammals should have been able to migrate permanently or seasonally if they found the temperature too extreme the breeding season too short or the rainfall too sparse or unpredictable 210 Seasons vary geographically By migrating away from the equator herbivores could have found areas with growing seasons more favorable for finding food and breeding successfully Modern day African elephants migrate during periods of drought to places where there is apt to be water 211 Large animals also store more fat in their bodies than do medium sized animals and this should have allowed them to compensate for extreme seasonal fluctuations in food availability 212 Some evidence weighs against climate change as a valid hypothesis as applied to Australia It has been shown that the prevailing climate at the time of extinction 40 000 50 000 BP was similar to that of today and that the extinct animals were strongly adapted to an arid climate The evidence indicates that all of the extinctions took place in the same short time period which was the time when humans entered the landscape The main mechanism for extinction was probably fire started by humans in a then much less fire adapted landscape Isotopic evidence shows sudden changes in the diet of surviving species which could correspond to the stress they experienced before extinction 213 214 215 Some evidence obtained from analysis of the tusks of mastodons from the American Great Lakes region appears inconsistent with the climate change hypothesis Over a span of several thousand years prior to their extinction in the area the mastodons show a trend of declining age at maturation This is the opposite of what one would expect if they were experiencing stresses from deteriorating environmental conditions but is consistent with a reduction in intraspecific competition that would result from a population being reduced by human hunting 216 It may be observed that neither the overkill nor the climate change hypotheses can fully explain events browsers mixed feeders and non ruminant grazer species suffered most while relatively more ruminant grazers survived 217 However a broader variation of the overkill hypothesis may predict this because changes in vegetation wrought by either Second Order Predation see below 174 218 or anthropogenic fire preferentially selects against browse species citation needed Disease edit The hyperdisease hypothesis as advanced by Ross D E MacFee and Preston A Marx attributes the extinction of large mammals during the late Pleistocene to indirect effects of the newly arrived aboriginal humans 219 220 221 In more recent times disease has driven many vulnerable species to extinction the introduction of avian malaria and avipoxvirus for example has greatly decreased the populations of the endemic birds of Hawaii with some going extinct 222 The hyperdisease hypothesis proposes that humans or animals traveling with them e g chickens or domestic dogs introduced one or more highly virulent diseases into vulnerable populations of native mammals eventually causing extinctions The extinction was biased toward larger sized species because smaller species have greater resilience because of their life history traits e g shorter gestation time greater population sizes etc Humans are thought to be the cause because other earlier immigrations of mammals into North America from Eurasia did not cause extinctions 219 A similar suggestion is that pathogens were transmitted by the expanding humans via the domesticated dogs they brought with them 223 A related theory proposes that a highly contagious prion disease similar to chronic wasting disease or scrapie that was capable of infecting a large number of species was the culprit Animals weakened by this superprion would also have easily become reservoirs of viral and bacterial diseases as they succumbed to neurological degeneration from the prion causing a cascade of different diseases to spread among various mammal species This theory could potentially explain the prevalence of heterozygosity at codon 129 of the prion protein gene in humans which has been speculated to be the result of natural selection against homozygous genotypes that were more susceptible to prion disease and thus potentially a tell tale of a major prion pandemic that affected humans of or younger than reproductive age far in the past and disproportionately killed before they could reproduce those with homozygous genotypes at codon 129 224 If a disease was indeed responsible for the end Pleistocene extinctions then there are several criteria it must satisfy see Table 7 3 in MacPhee amp Marx 1997 First the pathogen must have a stable carrier state in a reservoir species That is it must be able to sustain itself in the environment when there are no susceptible hosts available to infect Second the pathogen must have a high infection rate such that it is able to infect virtually all individuals of all ages and sexes encountered Third it must be extremely lethal with a mortality rate of c 50 75 Finally it must have the ability to infect multiple host species without posing a serious threat to humans Humans may be infected but the disease must not be highly lethal or able to cause an epidemic citation needed As with other hypotheses a number of counterarguments to the hyperdisease hypothesis have been put forth Generally speaking disease has to be very virulent to kill off all the individuals in a genus or species Even such a virulent disease as West Nile fever is unlikely to have caused extinction 225 The disease would need to be implausibly selective while being simultaneously implausibly broad Such a disease needs to be capable of killing off wolves such as Canis dirus or goats such as Oreamnos harringtoni while leaving other very similar species Canis lupus and Oreamnos americanus respectively unaffected It would need to be capable of killing off flightless birds while leaving closely related flighted species unaffected Yet while remaining sufficiently selective to afflict only individual species within genera it must be capable of fatally infecting across such clades as birds marsupials placentals testudines and crocodilians No disease with such a broad scope of fatal infectivity is known much less one that remains simultaneously incapable of infecting numerous closely related species within those disparate clades On the other hand this objection does not account for the possibility of a variety of different diseases being introduced around the same era citation needed Numerous species including wolves mammoths camelids and horses had emigrated continually between Asia and North America over the past 100 000 years For the disease hypothesis to be applicable there it would require that the population remain immunologically naive despite this constant transmission of genetic and pathogenic material citation needed The dog specific hypothesis in particular cannot account for several major extinction events notably the Americas for reasons already covered and Australia Dogs did not arrive in Australia until approximately 35 000 years after the first humans arrived there and approximately 30 000 years after the Australian megafaunal extinction was complete citation needed Extraterrestrial impact edit An extraterrestrial impact which has occasionally been proposed as a cause of the Younger Dryas 226 is sometimes suggested as a potential cause of the extinction of North America s megafauna due to the temporal proximity between a proposed date for such an impact and the following megafaunal extinctions 227 4 However the Younger Dryas impact hypothesis is widely considered a fringe hypothesis that is unsupported by evidence 228 229 Geomagnetic field weakening edit A weakening of the Earth s magnetic field may have caused increased flux of UV B radiation and has been suggested as a cause of megafaunal extinctions in the Late Quaternary 230 Effects editThe extinction of the megafauna could have caused the disappearance of the mammoth steppe rather than the other way around Alaska now has low nutrient soil unable to support bison mammoths and horses R Dale Guthrie has claimed this as a cause of the extinction of the megafauna there however he may be interpreting it backwards The loss of large herbivores to break up the permafrost allows the cold soils that are unable to support large herbivores today Today in the arctic where trucks have broken the permafrost grasses and diverse flora and fauna can be supported 231 232 In addition Chapin Chapin 1980 showed that simply adding fertilizer to the soil in Alaska could make grasses grow again like they did in the era of the mammoth steppe Possibly the extinction of the megafauna and the corresponding loss of dung is what led to low nutrient levels in modern day soil and therefore is why the landscape can no longer support megafauna Megafauna play a significant role in the lateral transport of mineral nutrients in an ecosystem tending to translocate them from areas of high to those of lower abundance They do so by their movement between the time they consume the nutrient and the time they release it through elimination or to a much lesser extent through decomposition after death 233 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 234 235 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 235 The extinction of the mammoths allowed grasslands they had maintained through grazing habits to become birch forests 236 The new forest and the resulting forest fires may have induced climate change 236 Such disappearances might be the result of the proliferation of modern humans 237 238 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 239 contributing to the warmer climate of the time up to 10 C warmer than at present 239 240 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 239 Recent studies have indicated that the extinction of megafaunal herbivores may have caused a reduction in atmospheric methane 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 241 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 4 times more rapid than any other decrease in the last half million years suggesting that an unusual mechanism was at work 242 The extermination of megafauna left many niches vacant which has been cited as an explanation for the vulnerability and fragility of many ecosystems to destruction in the later Holocene extinction The comparative lack of megafauna in modern ecosystems has reduced high order interactions among surviving species reducing ecological complexity 243 This depauperate post megafaunal ecological state has been associated with diminished ecological resilience to stressors 244 Many extant species of plants have adaptations that were advantageous in the presence of megafauna but are now useless in their absence 245 The demise of megafaunal ecosystem engineers in the Arctic that maintained open grassland environments has been highly detrimental to shorebirds of the genus Numenius 246 Human perception editAnalysis of Maori oral traditions has found that the extinction of New Zealand s megafauna during the Holocene extinction had significant social and linguistic impacts on their culture suggesting that similar phenomena may have occurred in other societies that lived through megafaunal extinctions 247 See also editAustralian megafauna Large animals in Australia past and present era Holocene extinction Ongoing extinction event caused by human activity Late Quaternary prehistoric birds Extinct bird speciesPages displaying short descriptions of redirect targets List of quaternary mammalian fauna of China Megafauna Large animals Pleistocene megafauna Extinction event occurring during the late Quaternary periodPages displaying short descriptions of redirect targets Pleistocene rewilding Ecological practice Toba catastrophe theory Supereruption 74 000 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