Mammuthus Africanavus Plio Tunisia
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A mammoth is any species of the
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
elephantid genus ''Mammuthus'', one of the many genera that make up the
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
of trunked
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s called
proboscidea The Proboscidea (; , ) are a taxonomic order of afrotherian mammals containing one living family (Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. From ...
ns. The various species of mammoth were commonly equipped with long, curved tusks and, in northern species, a covering of long
hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and f ...
. They lived from the Pliocene epoch (from around 5 million years ago) into the Holocene at about 4,000 years ago, and various species existed in Africa, Europe, Asia, and North America. They were members of the family Elephantidae, which also contains the two genera of modern elephants and their ancestors. Mammoths are more closely related to living Asian elephants than African elephants. The oldest representative of ''Mammuthus'', the South African mammoth (''M. subplanifrons''), appeared around 5 million years ago during the early Pliocene in what is now southern and eastern Africa. Descendant species of these mammoths moved north and continued to propagate into numerous subsequent species, eventually covering most of Eurasia before migrating into North America around 1.5-1.3 million years ago, becoming ancestral to the Columbian mammoth (''M. columbi''). The last species to emerge, the woolly mammoth (''M. primigenius''), developed about 400,000 years ago in East Asia, with some surviving on Russia's Wrangel Island in the Arctic Ocean, as well as possibly the Taymyr Peninsula on mainland Siberia, until as recently as roughly 3,700 to 4,000 years ago, still extant during the construction of the
Great Pyramid The Great Pyramid of Giza is the biggest Egyptian pyramid and the tomb of Fourth Dynasty pharaoh Khufu. Built in the early 26th century BC during a period of around 27 years, the pyramid is the oldest of the Seven Wonders of the Ancient World, ...
of ancient Egypt.


Evolution

The earliest known proboscideans, the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
that contains the elephants, existed about 55 million years ago around the Tethys Sea area. The closest relatives of the Proboscidea are the sirenians and the
hyrax Hyraxes (), also called dassies, are small, thickset, herbivorous mammals in the order Hyracoidea. Hyraxes are well-furred, rotund animals with short tails. Typically, they measure between long and weigh between . They are superficially simil ...
es. The family Elephantidae is known to have existed six million years ago in Africa, and includes the living elephants and the mammoths. Among many now extinct clades, the mastodon is only a distant relative of the mammoths, and part of the separate Mammutidae family, which diverged 25 million years before the mammoths evolved. The following cladogram shows the placement of the genus ''Mammuthus'' among other proboscideans, based on hyoid characteristics: Since many remains of each species of mammoth are known from several localities, it is possible to reconstruct the evolutionary history of the genus through morphological studies. Mammoth species can be identified from the number of enamel ridges on their molars; the primitive species had few ridges, and the amount increased gradually as new species evolved and replaced the former ones. At the same time, the crowns of the teeth became longer, and the skulls become higher from top to bottom and shorter from the back to the front over time to accommodate this. The first known members of the genus ''Mammuthus'' are the African species ''
Mammuthus subplanifrons ''Mammuthus subplanifrons'', the South African Mammoth, is the oldest representative of the genus ''Mammuthus'', appearing around 5 million years ago during the early Pliocene in what is today South Africa and countries of East Africa East ...
'' from the Pliocene and ''
Mammuthus africanavus The African mammoth, ''Mammuthus africanavus'' (literally, "African ancestor mammoth"), is the second oldest mammoth species, having first appeared around 3 million years ago during the late Pliocene, with a last appearance around 1.65 million ...
'' from the Pleistocene. The former is thought to be the ancestor of later forms. Mammoths entered Europe around 3 million years ago; the earliest known type has been named ''M. rumanus'', which spread across Europe and China. Only its molars are known, which show it had 8–10 enamel ridges. A population evolved 12–14 ridges and split off from and replaced the earlier type, becoming ''
M. meridionalis ''Mammuthus meridionalis'', or the southern mammoth, is an extinct species of mammoth native to Europe and Central Asia from the Gelasian stage of the Early Pleistocene, living from 2.5–0.8 mya. Taxonomy The taxonomy of extinct elephant ...
''. In turn, this species was replaced by the steppe mammoth, ''
M. trogontherii The steppe mammoth (''Mammuthus trogontherii'', sometimes ''Mammuthus armeniacus'') is an extinct species of Elephantidae that ranged over most of northern Eurasia during the late Early and Middle Pleistocene, approximately 1.8 million-200,000 y ...
'', with 18–20 ridges, which evolved in East Asia ca. 1.8 million years ago. Steppe mammoth populations replaced ''
M. meridionalis ''Mammuthus meridionalis'', or the southern mammoth, is an extinct species of mammoth native to Europe and Central Asia from the Gelasian stage of the Early Pleistocene, living from 2.5–0.8 mya. Taxonomy The taxonomy of extinct elephant ...
'' in Europe between 1 and 0.7 million years ago. The Columbian mammoth, ''
M. columbi The Columbian mammoth (''Mammuthus columbi'') is an extinct species of mammoth that inhabited the Americas as far north as the Northern United States and as far south as Costa Rica during the Pleistocene epoch. It was one of the last in a line ...
'', evolved from a population of ''M. trogontherii'' that had entered North America over 1 million years ago. Mammoths derived from ''M. trogontherii'' evolved molars with 26 ridges between 800,000 and 400,000 years ago in Siberia, becoming the woolly mammoth, ''M. primigenius''. The woolly mammoth would replace the steppe mammoth in Europe during the late Middle Pleistocene around 200,000 years ago. A 2011 genetic study showed that two examined specimens of the Columbian mammoth were grouped within a subclade of woolly mammoths. This suggests that the two populations interbred and produced fertile offspring. It also suggested that a North American form known as "''M. jeffersonii''" may be a hybrid between the two species. By the late Pleistocene, mammoths in continental Eurasia had undergone a major transformation, including a shortening and heightening of the cranium and mandible, increase in molar hypsodonty index, increase in plate number, and thinning of dental enamel. Due to this change in physical appearance, it became customary to group European mammoths separately into distinguishable clusters: # Early Pleistocene – ''Mammuthus meridionalis'' # Middle Pleistocene – ''Mammuthus trogontherii'' # Late Pleistocene – ''Mammuthus primigenius'' There is speculation as to what caused this variation within the three chronospecies. Variations in environment, climate change, and migration surely played roles in the evolutionary process of the mammoths. Take ''M. primigenius'' for example: Woolly mammoths lived in opened grassland biomes. The cool steppe-tundra of the Northern Hemisphere was the ideal place for mammoths to thrive because of the resources it supplied. With occasional warmings during the ice age, climate would change the landscape, and resources available to the mammoths altered accordingly. In February 2021, scientists reported that DNA from million-year old mammoth
remains Remains or The Remains may refer to: Music *The Remains (band), a 1960s American rock band *The Ramainz, originally The Remains, a Ramones tribute band Albums * ''Remains'' (Alkaline Trio album), 2007 * ''Remains'' (Annihilator album), 1997 * ' ...
had become the oldest ever sequenced.


Etymology and early observations

The word ''mammoth'' was first used in Europe during the early 17th century, when referring to ''maimanto'' tusks discovered in Siberia. John Bell, who was on the Ob River in 1722, said that mammoth tusks were well known in the area. They were called "mammon's horn" and were often found in washed-out river banks. Some local people claimed to have seen a living mammoth, but they came out only at night and always disappeared under water when detected. He bought one and presented it to
Hans Sloan Sir Hans Sloane, 1st Baronet (16 April 1660 – 11 January 1753), was an Irish physician, naturalist, and collector, with a collection of 71,000 items which he bequeathed to the British nation, thus providing the foundation of the British Mu ...
who pronounced it an elephant's tooth. The folklore of some native peoples of Siberia, who would routinely find mammoth bones, and sometimes frozen mammoth bodies, in eroding river banks, had various interesting explanations for these finds. Among the
Khanty people The Khanty (Khanty: ханти, ''hanti''), also known in older literature as Ostyaks (russian: остяки) are a Ugric indigenous people, living in Khanty–Mansi Autonomous Okrug, a region historically known as "Yugra" in Russia, together ...
of the
Irtysh River The Irtysh ( otk, 𐰼𐱅𐰾:𐰇𐰏𐰕𐰏, Ertis ügüzüg, mn, Эрчис мөрөн, ''Erchis mörön'', "erchleh", "twirl"; russian: Иртыш; kk, Ертіс, Ertis, ; Chinese: 额尔齐斯河, pinyin: ''É'ěrqísī hé'', Xiao'erj ...
basin, a belief existed that the mammoth was some kind of a water spirit. According to other Khanty, the mammoth was a creature that lived underground, burrowing its tunnels as it went, and would die if it accidentally came to the surface. The concept of the mammoth as an underground creature was known to the Chinese, who received some mammoth ivory from the Siberian natives; accordingly, the creature was known in China as ''yǐn shǔ'' 隐鼠, "the hidden rodent". Thomas Jefferson, who famously had a keen interest in paleontology, is partially responsible for transforming the word ''mammoth'' from a noun describing the prehistoric elephant to an adjective describing anything of surprisingly large size. The first recorded use of the word as an adjective was in a description of a large wheel of cheese (the "
Cheshire Mammoth Cheese The Cheshire Mammoth Cheese was a gift from the town of Cheshire, Massachusetts to President of the United States of America, President Thomas Jefferson in 1802. The cheese was created by combining the milk from every cow in the town, and made in ...
") given to Jefferson in 1802.Simpson, J. (2009).
Word Stories: Mammoth
" ''Oxford English Dictionary Online'', Oxford University Press. Accessed 05-JUN-2009.


Description

Like their modern relatives, mammoths were quite large. The largest known species reached heights in the region of at the shoulder and weights of up to , while exceptionally large males may have exceeded . However, most species of mammoth were only about as large as a modern Asian elephant (which are about 2.5 m to 3 m high at the shoulder, and rarely exceeding 5 tonnes). Both sexes bore tusks. A first, small set appeared at about the age of six months, and these were replaced at about 18 months by the permanent set. Growth of the permanent set was at a rate of about per year. Based on studies of their close relatives, the modern elephants, mammoths probably had a
gestation Gestation is the period of development during the carrying of an embryo, and later fetus, inside viviparous animals (the embryo develops within the parent). It is typical for mammals, but also occurs for some non-mammals. Mammals during pregna ...
period of 22 months, resulting in a single calf being born. Their social structure was probably the same as that of
African African or Africans may refer to: * Anything from or pertaining to the continent of Africa: ** People who are native to Africa, descendants of natives of Africa, or individuals who trace their ancestry to indigenous inhabitants of Africa *** Ethn ...
and Asian elephants, with females living in herds headed by a matriarch, whilst bulls lived solitary lives or formed loose groups after sexual maturity. Scientists discovered and studied the remains of a mammoth calf, and found that fat greatly influenced its form, and enabled it to store large amounts of nutrients necessary for survival in temperatures as low as . The fat also allowed the mammoths to increase their muscle mass, allowing the mammoths to fight against enemies and live longer. Woolly mammoths evolved a suite of adaptations for arctic life, including morphological traits such as small ears and tails to minimize heat loss, a thick layer of subcutaneous fat, long thick fur, and numerous sebaceous glands for insulation , as well as a large brown-fat hump like deposit behind the neck that may have functioned as a heat source and fat reservoir during winter.


Diet

Depending on the species or race of mammoth, the diet differed somewhat depending on location, although all mammoths ate similar things. For the Columbian mammoth, ''
M. columbi The Columbian mammoth (''Mammuthus columbi'') is an extinct species of mammoth that inhabited the Americas as far north as the Northern United States and as far south as Costa Rica during the Pleistocene epoch. It was one of the last in a line ...
'', the diet was mainly grazing. American Columbian mammoths fed primarily on cactus leaves, trees, and shrubs. These assumptions were based on mammoth feces and mammoth teeth. Mammoths, like modern day elephants, have hypsodont molars. These features also allowed mammoths to live an expansive life because of the availability of grasses and trees. For the Mongochen mammoth, its diet consisted of herbs, grasses, larch, and shrubs, and possibly alder. These inferences were made through the observation of mammoth feces, which scientists observed contained non-arboreal pollen and moss spores. European mammoths had a major diet of C3 carbon fixation plants. This was determined by examining the isotopic data from the European mammoth teeth. The arctic tundra and steppe where the mammoths lived appears to have been dominated by forbs, not grass. These were richer in protein and easier to digest than grasses and wooden plants, which came to dominate the areas when the climate became wetter and warmer. This could have been a major contributor to why the arctic megafauna went extinct. The Yamal baby mammoth Lyuba, found in 2007 in the Yamal Peninsula in Western Siberia, suggests that baby mammoths, as do modern baby elephants, ate the dung of adult animals. The evidence to show this is that the dentition (teeth) of the baby mammoth had not yet fully developed to chew grass. Furthermore, there was an abundance of ascospores of
coprophilous fungi Coprophilous fungi (''dung-loving'' fungi) are a type of saprobic fungi that grow on animal dung. The hardy spores of coprophilous species are unwittingly consumed by herbivores from vegetation, and are excreted along with the plant matter. The f ...
from the pollen spectrum of the baby's mother. Coprophilous fungi are fungi that grow on animal dung and disperse spores in nearby vegetation, which the baby mammoth would then consume. Spores might have gotten into its stomach while grazing for the first few times. Coprophagy may be an
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the po ...
, serving to populate the infant's gut with the needed microbiome for digestion. Mammoths alive in the Arctic during the
Last Glacial Maximum The Last Glacial Maximum (LGM), also referred to as the Late Glacial Maximum, was the most recent time during the Last Glacial Period that ice sheets were at their greatest extent. Ice sheets covered much of Northern North America, Northern Eur ...
consumed mainly forbs, such as ''
Artemisia Artemisia may refer to: People * Artemisia I of Caria (fl. 480 BC), queen of Halicarnassus under the First Persian Empire, naval commander during the second Persian invasion of Greece * Artemisia II of Caria (died 350 BC), queen of Caria under th ...
'';
graminoid In botany and ecology, graminoid refers to a herbaceous plant with a grass-like morphology, i.e. elongated culms with long, blade-like leaves. They are contrasted to forbs, herbaceous plants without grass-like features. The plants most ofte ...
s were only a minor part of their diet.


Extinction

The woolly mammoth (''M. primigenius'') was the last species of the genus. Most populations of the woolly mammoth in North America and Eurasia, as well as all the Columbian mammoths (''M. columbi'') in North America, died out around the time of the last glacial retreat, as part of a mass extinction of
megafauna In terrestrial zoology, the megafauna (from Greek μέγας ''megas'' "large" and New Latin ''fauna'' "animal life") comprises the large or giant animals of an area, habitat, or geological period, extinct and/or extant. The most common threshold ...
in northern Eurasia and the Americas. Until recently, the last woolly mammoths were generally assumed to have vanished from Europe and southern Siberia about 12,000 years ago, but new findings show some were still present there about 10,000 years ago. Slightly later, the woolly mammoths also disappeared from continental northern Siberia. A small population survived on
St. Paul Island, Alaska Saint Paul Island (russian: Остров Святого Павла, Ostrov Svyatogo Pavla) is the largest of the Pribilof Islands, a group of four Alaskan volcanic islands located in the Bering Sea between the United States and Russia. The city o ...
, up until 3750 BC, and the small mammoths of Wrangel Island survived until about 2000 BC Recent eDNA research of sediments indicates mammoths survived in north central Siberia at least as late as 2000 BC, in continental northeast Siberia until at least 5300 BC, and until at least 6600 BC in North America. The warming trend (Holocene) that occurred 12,000 years ago, accompanied by a glacial retreat and rising sea levels, has been suggested as a contributing factor. Forests replaced open woodlands and grasslands across the continent. The available habitat would have been reduced for some megafaunal species, such as the mammoth. However, such climate changes were nothing new; numerous very similar warming episodes had occurred previously within the ice age of the last several million years without producing comparable megafaunal extinctions, so climate alone is unlikely to have played a decisive role. The spread of advanced human hunters through northern Eurasia and the
Americas The Americas, which are sometimes collectively called America, are a landmass comprising the totality of North and South America. The Americas make up most of the land in Earth's Western Hemisphere and comprise the New World. Along with th ...
around the time of the extinctions, however, was a new development, and thus might have contributed significantly. Whether the general mammoth population died out for climatic reasons or due to overhunting by humans is controversial. During the transition from the Late Pleistocene epoch to the Holocene epoch, there was shrinkage of the distribution of the mammoth because progressive warming at the end of the Pleistocene epoch changed the mammoth's environment. The mammoth steppe was a periglacial landscape with rich herb and grass vegetation that disappeared along with the mammoth because of environmental changes in the climate. Mammoths had moved to isolated spots in Eurasia, where they disappeared completely. Also, it is thought that Late Paleolithic and Mesolithic human hunters might have affected the size of the last mammoth populations in Europe. There is evidence to suggest that humans did cause the mammoth extinction, although there is no definitive proof. It was found that humans living south of a mammoth steppe learned to adapt themselves to the harsher climates north of the steppe, where mammoths resided. It was concluded that if humans could survive the harsh north climate of that particular mammoth steppe then it was possible humans could hunt (and eventually extinguish) mammoths everywhere. Another hypothesis suggests mammoths fell victim to an infectious disease. A combination of climate change and hunting by humans may be a possible explanation for their extinction. ''
Homo erectus ''Homo erectus'' (; meaning "upright man") is an extinct species of archaic human from the Pleistocene, with its earliest occurrence about 2 million years ago. Several human species, such as '' H. heidelbergensis'' and '' H. antecessor' ...
'' is known to have consumed mammoth meat as early as 1.8 million years ago, though this may mean only successful scavenging, rather than actual hunting. Later humans show greater evidence for hunting mammoths; mammoth bones at a 50,000-year-old site in South Britain suggest that
Neanderthals Neanderthals (, also ''Homo neanderthalensis'' and erroneously ''Homo sapiens neanderthalensis''), also written as Neandertals, are an Extinction, extinct species or subspecies of archaic humans who lived in Eurasia until about 40,000 years ag ...
butchered the animals, while various sites in Eastern Europe dating from 15,000 to 44,000 years old suggest humans (probably '' Homo sapiens'') built dwellings using mammoth bones (the age of some of the earlier structures suggests that Neanderthals began the practice). However, the American Institute of Biological Sciences notes that bones of dead elephants, left on the ground and subsequently trampled by other elephants, tend to bear marks resembling butchery marks, which have allegedly been misinterpreted as such by
archaeologists Archaeology or archeology is the scientific study of human activity through the recovery and analysis of material culture. The archaeological record consists of artifacts, architecture, biofacts or ecofacts, sites, and cultural landscap ...
. In 2021, a study using ancient environmental DNA concluded that the extinction of the mammoth was primarily caused by dramatic vegetation changes at the end of the
Last Glacial Maximum The Last Glacial Maximum (LGM), also referred to as the Late Glacial Maximum, was the most recent time during the Last Glacial Period that ice sheets were at their greatest extent. Ice sheets covered much of Northern North America, Northern Eur ...
, due to a changed climate and precipitation regime. During the Last Glacial Maximum, the Arctic would have had a homogeneous flora consisting of mammoth steppe-associated plants, supporting a unique community of grazing mammals, including the mammoth. Increased precipitation following the end of the LGM caused caused a significant
paludification Paludification is the most common process by which peatlands in the boreal zone are formed. Formation The process is characterized by peat initialization on previously drier and vegetated habitats over inorganic soils, with no fully aquatic ...
of the steppe, which has been linked to a general decrease in species diversity of the region's former inhabitants, and not just mammoths. In addition, the study found mammoths to have persisted on the mainland for far longer than previously thought, with their shrinking distribution roughly tracking the shrinkage of the mammoth steppe; their final mainland refugium, the Taymyr Peninsula, was roughly concurrent with the Wrangel population. The close relationship between decreasing mammoth populations and steppe, as well as the lack of a sudden decline that would be associated with the human overkill hypothesis, indicates that humans and mammoths may have coexisted for millennia, and thus humans may have only played a minor role in the species' extinction. Many hypotheses also seek to explain the regional extinction of mammoths in specific areas. Scientists have speculated that the mammoths of Saint Paul Island (Alaska), an isolated enclave where mammoths survived until about 8,000 years ago, died out as the island shrank by 80–90% when sea levels rose, eventually making it too small to support a viable population. Similarly, genome sequences of the Wrangel Island mammoths indicate a sharp decline in genetic diversity, though the extent to which this played a role in their extinction is still unclear. Another hypothesis, said to be the cause of mammoth extinction in Siberia, comes from the idea that many may have drowned. While traveling to the Northern River, many of these mammoths broke through the ice and drowned. This also explains bones remains in the Arctic Coast and some of the New Siberian Islands. Dwarfing occurred with the
pygmy mammoth The pygmy mammoth or Channel Islands mammoth (''Mammuthus exilis'') is an extinct species of dwarf elephant descended from the Columbian mammoth (''M. columbi'') of mainland North America. This species became extinct during the Quaternary extin ...
on the outer Channel Islands of California, but at an earlier period. Those animals were very likely killed by early Paleo-Native Americans, and habitat loss caused by a rising sea level that split Santa Rosae into the outer Channel Islands.


Mammoth-elephant hybrid

One proposed scientific use of this preserved genetic material is to recreate living mammoths. This has long been discussed theoretically but has only recently become the subject of formal effort due to advances in molecular biology techniques and cloning of mammals.The Long Now Fouldation - Revive and Restore
According to one research team, a mammoth cannot be recreated, but they will try to eventually grow in an "artificial womb" a hybrid elephant with some woolly mammoth traits.The Mammoth Genome Project
Pennsylvania State University The Pennsylvania State University (Penn State or PSU) is a Public university, public Commonwealth System of Higher Education, state-related Land-grant university, land-grant research university with campuses and facilities throughout Pennsylvan ...
. Accessed: October 2018.
The outcome would be an elephant-mammoth hybrid with no more than 1% mammoth genes. Other projects are working on gradually adding mammoth genes to elephant cells ''in vitro''."Mammoth: Back from the Dead"
. ''National Geographic Channel''.


See also

* ''
Genesis 2.0 ''Genesis 2.0'' is a documentary film made by Swiss director and producer Christian Frei and Russian filmmaker Maxim Abugaev. The feature-length film was released in January 2018 in the World Cinema Documentary section at the Sundance Film Festiv ...
'', a documentary *
Ivory trade The ivory trade is the commercial, often illegal trade in the ivory tusks of the hippopotamus, walrus, narwhal, mammoth, and most commonly, African and Asian elephants. Ivory has been traded for hundreds of years by people in Africa and Asia, ...
*
La Brea tar pits La Brea Tar Pits is an active paleontological research site in urban Los Angeles. Hancock Park was formed around a group of tar pits where natural asphalt (also called asphaltum, bitumen, or pitch; ''brea'' in Spanish) has seeped up from the gro ...
*
List of mammoths A ''list'' is any set of items in a row. List or lists may also refer to: People * List (surname) Organizations * List College, an undergraduate division of the Jewish Theological Seminary of America * SC Germania List, German rugby union ...
* The Mammoth Site in Hot Springs, South Dakota *
Niederweningen Mammoth Museum The Mammutmuseum Niederweningen (literally: Mammoth Museum Niederweningen) is a paleontological and geological museum in the municipality of Niederweningen in the Wehntal valley, Canton of Zürich, Switzerland, and one of the few mammoth museums ...
* Pleistocene Park * Waco Mammoth National Monument


References


Further reading

* * * * * * * * * * * * {{Authority control Cenozoic mammals of Africa Cenozoic mammals of Asia Cenozoic mammals of Europe Cenozoic mammals of North America Holocene extinctions Pleistocene proboscideans Prehistoric elephants Taxa named by Joshua Brookes Zanclean first appearances