Woolly mammoth

Woolly mammoth
Skeleton reconstruction of a woolly mammoth in the Southeast Bavarian Natural History and Mammoth Museum in Siegsdorf
Systematics
without rank: Paenungulata without rank: Tethytheria Order : Russell animals (Proboscidea) Family : Elephants (Elephantidae) Genre : Mammoths ( Mammuthus ) Type : Woolly mammoth
Scientific name
Mammuthus primigenius
( Blumenbach , 1799)

The woolly mammoth , also woolly or fur mammoth ( Mammuthus primigenius ), is an extinct species from the elephant family . This type of mammoth developed in the transition from the Old - to the Middle Pleistocene around 800,000 to 600,000 years ago in Siberia and inhabited the cold-age steppes in northern Eurasia and North America . It represents the final link of the Plio- and Pleistocene development of the mammoths. At the end of the Pleistocene , the woolly mammoth died out in large parts of its range, but survived regionally in some remaining populations until the middle Holocene (1800 BC).

distribution and habitat

Distribution area

Maximum distribution of the woolly mammoth during the Vistula glacial period

The woolly mammoth was an animal adapted to the cold and preferred boreal to subarctic climates . It originated in the interior of Siberia around 800,000 to 600,000 years ago and spread in Eurasia in the late Middle Pleistocene around 300,000 to 250,000 years ago. In Central Europe it has been found in the Saale Glaciation (300,000 to 126,000 years ago), including from the Geiseltal ( Saxony-Anhalt ). At the end of this freezing period, however, the species quickly withdrew to its original areas.

At the end of the Eem warm period (126,000 to 115,000 years ago) the woolly mammoth began another phase of expansion towards the south, south-west and south-east, but also to the north. Finds from Averley ( Essex , England ), where it has already been demonstrated in late warm-period sediments , show that this second wave of propagation must have taken place very quickly . In the last glacial period that followed (115,000 to about 12,000 years ago) the range of the woolly mammoth reached its maximum extent. The species occurred in western Eurasia across the board from western to central to eastern and the southern part of northern Europe ; Exceptions were the ice-covered areas such as the northern part of Fennoscandinavia . In addition, it penetrated southern Europe , where it reached its southernmost point in Europe on the Iberian Peninsula in Granada ( Spain ) at latitude 37 ° north. But it has also advanced to the Apennines , the Balkan Peninsula and the Black Sea .

In West Asia , the woolly mammoth reached the southern Caucasus region , while in Central Asia it also appeared in Kazakhstan , bypassing the desert regions of the Kyzylkum . Further to the east, it also populated the northern part of Mongolia . In East Asia , it penetrated into northeast China , where its southernmost sites on the Huang He are at about 35 ° to 36 ° north latitude, which is also the world's southernmost limit of distribution of this proboscis species. In the far east of Asia, finds are known from the northern part of the Korean peninsula , from Kamchatka , Sakhalin (both Russia ) and from Hokkaidō ( Japan ).

Furthermore, there was a northern expansion, in the course of which Yakutia and the far north and northeast of Asia were settled, such as the Taimyr Peninsula and Wrangel Island . The Taimyr Peninsula represents the northernmost distribution limit of the woolly mammoth, which is approximately at the 76th parallel. The woolly mammoth must have reached the Chukchi Peninsula around 100,000 years ago. This vast north-east Asian expanse made it possible for the animals to immigrate to North America via the dry Bering Strait .

In North America, the woolly mammoth was found mainly in Alaska and northern Canada , while it reached the Great Lakes in the east . From there it advanced into the prairie areas and the Great Plains . This is also where the southernmost American limits of distribution are located, which are around 36 ° to 38 ° north latitude. The proboscis could not penetrate the western part of North America because the Rocky Mountains were then covered with ice. Remains of the woolly mammoth, originally reported from Mexico , are obviously confused with the prairie mammoth ( Mammuthus columbi ).

During the Vistula Glaciation, the entire area populated by the woolly mammoth spanned an area of ​​33.3 million square kilometers, regardless of individual temporal differences and enclosed areas that could not be inhabited by the animals. At the end of the last icing phase, the woolly mammoth withdrew from its new refuges, but soon largely died out in both Eurasia and North America. Some remaining populations, however, survived far into the Holocene . Among these populations there was one isolated on Wrangel Island, another on Saint Paul Island (Alaska) , which lies in the southern part of the Bering Sea , and two more on the Chukchi Peninsula and the Taimyr Peninsula of the Eurasian mainland.

habitat

The woolly mammoth preferred open landscapes and was adapted to the hard grass and shrub vegetation of the cold- age loess steppe and steppentundra. For this reason, it predominantly settled in flat regions and lowlands, occurred under favorable climatic conditions also in high altitudes and mountainous terrain, but largely avoided desert areas. It was particularly common in water-rich areas such as river valleys or lake banks. Due to the high distribution during the last glacial period, it sometimes lived in its southern distribution areas in a more forested or park-like, but still clearly steppe-like biotope. During the warmer climatic phases of the last glaciation period, it also inhabited a forest-steppe-like habitat in western Russia.

The woolly mammoth was not only namesake, but also an important inhabitant of the mammoth steppe , which occupied large parts of Eurasia and North America during the cold ages. This mammoth steppe , a hybrid of steppe and tundra vegetation , was characterized by a high level of solar radiation and a long-lasting high pressure situation, caused by its close proximity to the glacier edges . This resulted in the formation of a nutrient-rich plant cover. In this landscape, which no longer exists under today's climatic conditions, the s that occurred in the cold ages of the late Middle and Young Pleistocene flourished. G. Mammuthus-Coelodonta fauna complex , whose character animal was the woolly rhinoceros as well as the woolly mammoth . Other accompanying elements of this faunal complex were the bison , the reindeer , the saiga antelope and the musk ox .

Due to the long time frame and the large area of ​​distribution, the woolly mammoth occasionally coexisted with other mammoth and proboscis species. Especially at the beginning of the Saale glaciation in western Eurasia it still appeared together with the steppe mammoth ( Mammuthus trogontherii ), which however soon died out here. Since this older mammoth form seems to have persisted in eastern Eurasia, especially in China, until the late Young Pleistocene - here it is occasionally but incorrectly called Mammuthus sungari in its late form - both woolly and steppe mammoths keep coming back to individual Sites together. In North America, overlaps with the prairie mammoth occurred predominantly at the southern distribution limits of the woolly mammoth, which, however, generally had a more western and southern habitat. In addition, there was also a common habitat use here with the American mastodon ( Mammut americanum ), which also existed up to the Young Pleistocene , but which is not a representative of the elephants, but belongs to the much more primitive mammoth form of the mammoths (Mammutidae).

It is unclear whether the woolly mammoth also appeared together with the European forest elephant ( Palaeoloxodon antiquus ), which was adapted to warmer climates . In Central Europe this could have been the case at least in the transition phases between warm and cold periods. The rapid repopulation of western Eurasia by the woolly mammoth in the late Eem warm period makes it probable that the extreme ranges of the two proboscis were not geographically very far apart or even overlapped. It is also possible that the woolly mammoth, as it spread to the south, again moved very close to the last refuge areas of the European forest elephant in southern Europe during the subsequent glacial period.

Finds of the woolly mammoth

Locations in general

Skeletal reconstruction of a woolly mammoth in the Museum of Geology in Warsaw

There are many sites of discovery of the woolly mammoth due to its large distribution area. Usually only the most resilient skeletal elements such as the teeth are found, less often it is skeletal parts or even complete animals. There are many finds in gravel, sand and clay pits, the deposits of which show former rivers or lake basins. Occasionally they can also be found in caves or sinkholes or submarine on the floor of the North Sea and Baltic Sea or the coastal Arctic Ocean , whose shelf areas had dried out during the last cold period due to the lower sea level.

The mammoth cemetery in a loop of the Bjorjoljoch in Siberia, which was discovered in 1970 and dated to an age of around 12,000 years, is a very important site that is excellent for mammoth research . The entire 'cemetery' contained more than 8,800 mammoth bones from a total of 156 individuals. Outstanding is a 175 cm long rear leg of a woolly mammoth that was still completely covered with skin and hair. Why so many animals died here is debatable, but since there are more female animals, it is assumed that several herds or a large group of herds drowned while crossing the river. In the absence of complete skeletons, the event most likely took place elsewhere upstream. A similar, not quite as extensive 'cemetery' came to light on the banks of the Seva south of Moscow in 1988 , where a total of 4,000 bone fragments from 10 to 15 individuals of all ages were found. The age of the site is dated to 13,950 years.

Numerous sites are also known from Germany, but only six complete skeletons are available. The oldest in terms of research history comes from Klinge near Cottbus ( Brandenburg ) and was found in 1903. Another was recovered in 1909 near Borna near Leipzig ( Saxony ). It was exhibited in the local ethnological museum, but was destroyed during the Second World War . The skeleton of a 3.2 m high woolly mammoth came to light near Ahlen ( North Rhine-Westphalia ) in 1910 and is now in the Geological and Paleontological Museum of the Westphalian Wilhelms University in Münster , while another was excavated in 1936 on the Koblenz - Trier motorway near Polch . The Pfännerhall mammoth came to light in 1953 during opencast mining in the Geiseltal and is now exhibited in the State Museum of Prehistory in Halle (Saxony-Anhalt). The latest find of a 3.6 m high woolly mammoth comes from Siegsdorf near Traunstein ( Bavaria ) and was discovered in 1975, but only fully excavated ten years later. It is now in the Southeast Bavarian Natural History and Mammoth Museum in Siegsdorf .

Finds of mummified woolly mammoth carcasses

The beginnings in the 18th and 19th centuries

Eugen Pfitzenmayer (left) with the Berjosowka mammoth at the place of discovery, 1901

Dermoplasty of the Beryosovka mammoth in the found position, exhibited in the Zoological Museum in Saint Petersburg

The mammoth calf "Dima" at the place where it was found in the former Beringia mammoth steppe

Preserved woolly mammoths have been found in the ice of the permafrost, which has not or hardly thawed since the last freezing phase, beyond the 60th degree of north latitude, both in northern and northeastern Siberia (mainly in Yakutia) and in northern North America (Alaska, Canada). Such finds are particularly important in order to reconstruct the appearance, the diet and the way of life of this Pleistocene species of proboscis. One of the earliest scientifically investigated carcasses was found in 1799 by the Tungusian ivory dealer Osip Schumachow in the Lena delta . It was partially exposed during a landslide, but not fully exposed until the summer of 1806 through further erosion . Schumachow stripped the carcass of the tusks and sold them to the Yakut dealer Roman Boltunow. He traveled to the site and made a sketch of the carcass, which was complete, apart from the predatory damage on the trunk and ears . This sketch finally came to the Russian Academy of Sciences in Saint Petersburg , where the German-Russian botanist and naturalist Michael Friedrich Adams (1780–1838) saw it. Adams organized an expedition to the find area, to which Schumachow led him in 1806. At this point, half of the mammoth carcass had already fallen victim to predators, but the left side of the body still had good hair and skin preservation. When the skeleton was recovered, however, all soft tissues were lost except for the scalp and the soles of the feet. The skeleton was then brought to Saint Petersburg and has been in the Kunstkammer since 1808 . It is thus the first assembled woolly mammoth skeleton worldwide and is called the Adam's mammoth after its excavator .

It was not until 1846 that Russian surveyors found a well-preserved carcass on the Indigirka in Siberia, on which there were still remains of fur and the stomach contents were preserved. The carcass had been washed away by the spring floods and its hind legs were still in the ground. During the recovery, however, a large part of the soft tissue was lost due to the decomposition process that began immediately . In 1900 the Berjosowka mammoth, named after its location on a tributary of the Kolyma in north-eastern Siberia, was found . An expedition equipped by the Academy of Sciences the following year was able to successfully recover the mammoth, which was buried in the ground in a sitting position. Except for the trunk and parts of the head that had been eaten by wolves , bears , red or arctic foxes , it was complete. In addition to the tongue, it also had up to 15 kg of food residues in its stomach. A dermoplasty and the skeleton of the mammoth bull are now accessible in the Zoological Museum of the Russian Academy of Sciences in Saint Petersburg .

Important finds of the 20th century

In the following years, several mammoth carcasses were found. One was recovered on the Lyakhov Islands between 1901 and 1903 and, shortly before an imperial decree banned the export of mammoth bones from Russia, was taken to Paris , where it is on display in the Jardin des Plantes . Eugen Wilhelm Pfizenmayer (1869–1941), who was also involved in the recovery and subsequent restoration of the Berjosowka mammoth , found another, but severely damaged, carcass on the Sanga Jurach in Yakutia in 1908. This specimen is the first with a completely preserved trunk. Just one year later, the discovery of a complete carcass in the Mochowaja river valley on the Taimyr Peninsula was reported and some skin scraps were recovered. An expedition sent there in 1913 led by GN Kutomanow found only half of them, the other had previously been used as dog food by local hunters. The remainder of this find, known today as the Kutomanov mammoth , was recovered and brought to Saint Petersburg.

An excellently preserved, partially mummified skeleton, in which only insignificant bones were missing, was discovered in 1948 on a left tributary ( Reka Mamonta "River of the Mammoth") of the Schrenk in the northeast of the Taimyr Peninsula and recovered the following year. Because the type material on which Blumenbach's first description was based was no longer available, this specimen, known as the Tajmyr mammoth , was declared a neotype of Mammuthus primigenius because of its good traditional quality and typical morphology . In the same year one of the best preserved young woolly mammoths was discovered in Alaska near Fairbanks . Only the head, front leg and shoulder of the animal were preserved, but the skin and muscles in particular were very well preserved. DNA examinations on this find have contributed significantly to determining the evolution of this animal species. The specimen, known as Effie , is now on display in the American Museum of Natural History in New York .

The almost complete skeleton of the Shandrin mammoth , discovered in 1972 at the Schandrin , a tributary of the Indigirka, did not have a good preservation of the outer soft tissues, but the internal organs were well preserved and allowed for the first time an insight into the internal structure of the mammoth and a comparison to the recent elephants. Furthermore, the carcass, which belonged to an old but not very large bull, contained 291 kg of food remains. In 1977 the carcass of a young female animal was found on the right bank of the Juribei on the Gydan Peninsula , while in the same year reindeer herders also discovered a carcass known as the Chatanga mammoth in alluvial sands on the left bank of the Khatanga in northern Siberia and in a two-year excavation campaign.

In the Kolyma Basin on the Kirgiljach River in Sussuman District in Magadan Oblast , a worker discovered the ice mummy of a fully preserved male mammoth calf while mining for gold. The famous Kirgiljach or Magadan mammoth baby , which was named "Dima", died around 39,000 years ago at the age of six to eight months. The calf was 115 cm long and 104 cm high and weighed about 100 kg at the time of death. Remnants of hair were still preserved at the distal ends of the legs because these parts of the body are the first to freeze after death. The internal organs of "Dimas" hardly differ from those of recent elephants, but its auricles are only a tenth of the area of ​​an African elephant of the same age ( Loxodonta africana ).

The mammoth calf Mascha , which was found on the banks of the Juribetejach on the Jamal Peninsula in 1988, should also be emphasized , which is comparable in age and size to Dima , but which only lacks its trunk. A calf, only three months old, with a preserved skull and skin, small tusks, trunk and ears and parts of its limbs was discovered by hunters in 1990 on the banks of the Indigirka. It was named the Abyi Mammoth after the province it came from . In 1997 members of the Dolganen found two mammoth tusks on the Taimyr Peninsula near the Great Balachnija. During excavations in 1998 and 1999 at the site, a mammoth carcass with good soft tissue conservation was discovered. This was then recovered in a complete block and flown to an artificial cave on Chatanga in order to completely expose the mammoth there at controlled temperatures of −15 ° C and to also thoroughly examine the bottom sediment. The find was named Jarkow mammoth after its discoverers .

The 21st century

As early as 1990, a cadaver had become known on the banks of the Upper Taimyra in the central part of the Taimyr Peninsula, which was immediately flooded again. Ten years later, however, S. Pankewitsch rediscovered it while fishing, which is why it is known as the fish-hook mammoth . The find was then partially excavated, the rear part together with the internal organs and the stomach contents, however, recovered in a block and flown to the same artificial and cooled cave on the Chatanga as the Jarkow mammoth for study purposes . The Jukagir mammoth was discovered in 2002 by V. Gorochow on the Maksunuocha in northern Yakutia and excavated in the summer of 2004 by an international team consisting of Russians, Yakuts and Japanese. It is characterized above all by an excellently preserved skull with soft tissue covering and both completely preserved ears. The carcass of a calf from a mine in the highlands of Oymyakon also dates from 2004 . The animal was a good one and a half years old at the time of death and probably died from head trauma.

One of the most recent finds dates back to 2007 and was recovered by a reindeer herder on the upper reaches of the Juribei on the Yamal Peninsula. It is a one-month-old calf that was named Ljuba and is extremely well preserved. The yuka mammoth was found by locals in 2009 on the coast of Laptev Strait, about 30 km west of the mouth of the Kondratievo River ( Sakha Republic ). The female young animal died at the age of about 9 years and was about 205 cm long with a shoulder height of 160 cm. Only the left hind foot and parts of the back and neck were missing from the carcass. As the only find so far, it also contained remains of the brain, the reconstructed volume of which was around 4100 cm³. According to radiometric data, the find is between 28,000 and 39,000 years old. In 2012, the discovery of a carcass of a full-grown male individual, which was named the Zhenya mammoth , was reported. This came to light near the mouth of the Yenisei on the bank of the Sopochnaya . In addition to the almost complete skeleton, part of the right skin covering including the ear and various internal organs have survived. During its lifetime the animal was around 2.3 m tall and weighed around 2.5 t. It was believed to have reached sexual maturity and may have died as a result of a rivalry with another mammoth bull.

Other cadaver finds

Outside the permafrost areas, preserved mammoth finds come from Starunia ( Ukraine ). These were found in 1907 together with the mummified carcass of a woolly rhinoceros in an oil shale pit, in which they had been preserved in the earth's wax interspersed with salts at a depth of 12 to 17 m below the surface of the earth. However, the fur was already gone.

Appearance and way of life

Appearance

Skeleton reconstruction of a woolly mammoth in the Anthropos Museum Brno ( Czech Republic )

Trunks of various elephant species. Left: African elephant ( Loxodonta africana ), middle: Asian elephant ( Elephas maximus ), right: Woolly mammoth ( Mammuthus primigenius ).

Replica of a woolly mammoth in the Royal British Columbia Museum in Victoria

Due to the widespread distribution of the woolly mammoth and the associated high number of finds that include bones, teeth or complete skeletons, but also the preserved mummified carcasses or the representations in cave paintings by the Upper Paleolithic man, this animal species is one of the best known and best studied from the Pleistocene . Therefore the appearance and the way of life of the woolly mammoth have been extensively reconstructed. Woolly mammoths weren't as huge as is often thought. Adult animals reached a height of 2.8 to 3.75 m at the withers , not much more than today's elephants. A relatively small adult individual from Rottweil ( Baden-Württemberg ) was only 2.5 m high. As with the recent elephants, however, there was a pronounced gender dimorphism . Male animals averaged 2.7 to 3.4 m shoulder height, while those of the cows were often only between 2.6 and 2.9 m. However, the animals were more compact and bulky than elephants, so that the weight was between 5 and 6 t, and occasionally up to 8 t. The younger representatives of the species were usually smaller than the older ones, whose body size was somewhat reminiscent of the mighty steppe mammoth from which the species emerged. Nevertheless, the woolly mammoth was the largest animal of the mammoth steppe in the Upper Pleistocene . The smallest representatives of the woolly mammoths were the last of their kind and lived on Wrangel Island. Due to the dwarfing of the islands, they reached just 1.8 m shoulder height with a weight of around 2 t.

In contrast to the recent elephants, the woolly mammoth had shorter and massive legs and was overall longer. The head was very high with a clear cranial dome, behind which a neck indentation lay. On its front back it had a hump, which is often interpreted as a pad of fat. The further line of the back was markedly steep.

This species is named after the coarse outer fur, which in winter consisted of hair up to 90 cm long. Similar to the musk ox , this fur formed an apron on the belly and flank. The trunk, tail and ears were also covered with fur to protect against the cold, while the hair on the head formed a characteristic pony . Under these outer hairs there was initially a not so coarse, fluffy layer of fur with 40 to 50 cm long hair, while the dense undercoat consisted of 10 to 20 cm long hair. The summer fur was much shorter and less dense than the winter fur . The fur color of the egg corpses of the woolly mammoth varies from blond, reddish, yellowish to brownish to black, also in patterns, which is primarily due to a polymorphism in the gene of the melanocortin type 1 receptor . In addition, color tones may have been changed by post-mortem oxidation processes, possibly the reddish ones.

The skin of the woolly mammoth reached an average thickness of three centimeters, but on the soles of the feet it was significantly thicker with five to six centimeters. There were also thick pads of skin between the toes. Under the skin was a layer of fat almost ten centimeters thick, which provided additional heat insulation. However, since the animal species lacked sebum glands with which it could have greased its fur, it was defenseless in damp weather conditions.

The trunk of the woolly mammoth was relatively short as a cold climatic adaptation, but much thicker and more massive than that of today's elephants. The end of the trunk, which consisted of a "finger" and a shovel-shaped tip, is striking. The African elephant, on the other hand, has two "fingers" at the end of its trunk, while the Indian elephant ( Elephas maximus ) only has one. The fingers were largely used to grasp the food and functioned similarly to the human hand . In some individuals, a widening of the trunk towards the lower end has been observed, as a result of which the diameter doubles and the otherwise oval cross-section changes to an elliptical one. The very small ears are also a climate adaptation. They had an oval shape with a length of 38 cm and a width of 18 to 28 cm. As a result, the ears of adult animals only reached about a twentieth of those of their current African relatives, and in young animals like Dima it was still a tenth compared to elephant calves of the same age. The tail was also relatively short at around 40 cm and had a skin flap at the base that served as an anal valve . The tail end had a pronounced tassel made of hair up to 60 cm long.

Skull, dentition, and postcranial skeletal features

The mammoth "Millie" in the "Celtic and Prehistoric Museum", Ventry, Ireland

The skull was large and very high with a partly pronounced transverse crest and a massive lower jaw . The crest served as a starting point on the one hand for the required strong back muscles, which had to hold the skull with the tusks, and on the other hand for the powerful chewing apparatus. As with all elephants, the interior consisted of a honeycomb structure with thin bone lamellae, which led to a low skull weight in relation to its volume.

Characteristic are the alveoli of the tusks, which in the woolly mammoth ran almost parallel or at a small angular distance from one another. This distinguishes it, for example, from the Middle and Young Pleistocene European forest elephants, where they were at a stronger angle of up to 80 ° to each other.

Like all real elephants, the woolly mammoth had a greatly reduced set of teeth that only comprised the molars and the tusks formed from the upper incisors . The woolly mammoth had one molar tooth per arch, which, starting with the first deciduous molar, could be exchanged five times: a total of three deciduous molars and three permanent molars. The molars were long-oval, often over 20 cm long, and reached the highest crowns of all elephants . The high number of enamel lamellae, which was between 21 and 30 in the third molar, with an average enamel fold thickness of 0.9 to 2 mm is striking. This is the highest number among all known representatives of the elephants. In part, this value serves as a taxonomic distinguishing feature, whereby high degrees of chewing can impair an exact identification of the species.

The postcranial skeleton of the woolly mammoth has only a few differences from that of other elephants. The columnar limbs are characteristic. Also striking is the short tail, which is formed from only 21 vertebrae , while that of the recent proboscis ranges between 28 and 33. For a long time, paleontologists hoped to be able to assign individual or isolated skeletal elements to the various elephant species, but this has not been fulfilled. Only on the first cervical vertebra, the atlas , does the woolly mammoth show a lower and wider cranial joint than the European forest elephant .

tusks

Cross section of a mammoth tusk. The ring-shaped lamellae and the Schreger lines are clearly visible, especially in the second and third outer lamellae.

The tusks are one of the most striking external features of the woolly mammoth and, in contrast to the elephant species still alive today, are clearly twisted upwards in a spiral, with the pointed ends mostly pointing towards one another. By rotating the defenses, they are very wide at the sides with a span of up to 1.7 m. The tusks of old bulls could reach a length of more than 4.50 m, about a quarter of which was in the alveoli , and a weight of more than 100 kg. Measured across the curvature, the largest known specimen is 4.9 m long. On average, however, the tusks found are significantly smaller, measuring 2.50 m in length and 45 kg in weight. At the exit point from the alveoli, they have a diameter of up to 20 cm. Cows had shorter and thinner tusks - the diameter at the alveoli averages 9 to 10 cm - that were not so clearly spiral, but rather curved like a saber. In general, the woolly mammoth's defenses were longer than those of the elephants living today.

The tusks are made of ivory, a mixture of carbonate - hydroxyl - apatite crystals, which are fused with collagen fibers in an oriented manner, whereby the crystals are responsible for the necessary hardness, the collagen for the elasticity. Due to the differing degrees of crystallization in the ivory of the various elephant species, the tusks can be distinguished in thin section samples with the help of infrared spectroscopy . Likewise, the collagen shows variations in the impact of the individual teeth elephant species, so that by using the ultraviolet - Fluorescence - spectrophotometry these can be assigned.

In the internal structure, the tusks have three different positions. The outermost thin layer consists of cement , with dentin as the main component below . This has a fibrous structure and is interspersed with innumerable tubules that are filled with collagen. The innermost area forms the pulp , in which blood vessels and nerves are located. Numerous dentinal tubules also extend outwards from it. Unlike other teeth, tusks do not have tooth enamel . New tusk material is mainly formed in the alveoli and occurs from the inside out. As a result, the tusk is made up of concentric rings in cross section , which represent the respective growth spurts and are comparable to annual rings in the trunks of trees . The innermost rings are the youngest. In the longitudinal section, however, there are parallel, alternating light and dark lines, which also indicate growth rates.

In addition, structures can be observed in cross-section that extend beyond the growth rings and form a rosette-like grain of alternating light and dark lines. These formations, known as Schreger lines , are the result of a regular change in the collagen content in the dentin. Due to the strong curvature of the mammoth tusk, the angles at which the lines regularly meet are more acute than those of the more recent elephants and vary between 25 and 40 degrees depending on their position in the tusk.

The function of the tusks was certainly varied. On the one hand, they served to depict the dominance of individual animals in mating or ritual combat. Here, however, impaling was not possible as with the recent elephant species. Furthermore, they could have been a warning signal to food competitors in interspecific competition or to large predators. The tusks were very often used for foraging. This is shown by grinding marks, which are mainly located on the lower edge, but occasionally also on the top and the tip. Most of these grinding marks are explained by the shoveling of the snow-covered ground in search of food, but they could also have arisen in general when digging out plants or bending over bushes. Such tracks are also common in recent elephants.

Diet

The woolly mammoth was an animal adapted to steppe areas, as shown by its compact physique and, above all, its strong limbs, which specialized in long migrations. Since grasses were mainly available as a source of food in the steppes , it was suspected very early on in research history that the proboscis was a proven grass-eater. There are also anatomical features, such as the very high-crowned molars with their numerous enamel folds. Since grasses are not very nutritious, the woolly mammoth had to eat very large quantities. The high tooth crowns compensated for the greater chewing caused by the high amount of food. Since grasses also contain silica and are therefore very hard, the thick enamel folds also prevented greater abrasion.

Some of the mammoth carcasses discovered in the Siberian permafrost still contained stomach contents, such as the Shandrin mammoth or the Berjosowka mammoth ; the latter even had food residues on the tongue or in the mouth. The examinations of these food residues confirmed the anatomical evidence. Predominate, especially grasses, such as foxtail grass , barley , Agrostis , puccinellia and sedges , in addition, are buttercups and moss proven. To a lesser extent, there were also remains of willow and larch branches , as well as parts of willow , alder and pine trees . The occurrence of woody plants in the woolly mammoth's diet shows that the steppe was not completely free of shrubs and trees. The ingestion of such plant species was also important in order to get the nutrients needed for growth. The large variety of plants consumed also covered the amino acid requirement , which the woolly mammoth, in contrast to the large ruminants, had to regulate completely through food intake. The preserved intestinal contents of the Yukagir mammoth show, however, that the animals could also live in completely tree-free cold steppes. The necessary nutrients here apparently came mainly from dwarf subarctic willow species.

Depending on the season, a single animal required 150 to 300 kg of food per day, the amount of plants ingested depending on the degree of dryness, i.e. H. in spring with freshly grown grass an animal needed less than in summer and autumn with dry vegetation. In addition to comparisons with recent elephants, the stomach contents of the Shandrin mammoth , which weighed more than 290 kg, indicate the generally high need for food . The degradation of the vegetable cellulose took place in the woolly mammoth in the appendix . This enabled him to eat larger quantities with decreasing protein and increasing fiber content and thus to spend most of the day eating.

In addition to the vegetable diet, the woolly mammoth was heavily dependent on water. On average, it used around 70 to 90 liters a day. The amount of water required was in turn dependent on the nature of the plant food. With lush fresh grass, the woolly mammoth managed with less water. This also allowed him to move further away from the water points such as river and lake banks in the surrounding areas. In summer and autumn, when most of the plants were dried out, the water requirement was correspondingly higher. It is possible that during these times of the year the woolly mammoths were concentrated at the water points, which meant that the migration of the species was highly seasonal.

Individual way of life

Reconstruction of the woolly mammoth and its environment

Woolly mammoths reached an age of 60 to 65 years. This is evident on the one hand from the size of the animals and on the other hand from the characteristic "horizontal" tooth change that is common to all elephant species. The last molar, the third, is chewed off around the age of 60 and falls out. Old animals can no longer take in enough food and eventually die.

It is assumed that woolly mammoths, similar to today's elephant species, lived together in matriarchal family groups. The oldest dam led the group, which usually consisted of two to a maximum of 20 animals. Bulls, on the other hand, led a largely solitary life or, if necessary, formed into "bachelor groups", they only came together with the cows during the rut . This was probably seasonal due to the strong seasonal temperature fluctuations and took place in summer, in contrast to today's elephants , which are adapted to the tropical climate and are ready to mate all year round. After a 22-month gestation period, the young animal was born in spring, and it probably weighed around 90 kg. Young bulls were then evicted from the herd at 9 to 12 years of age. As is also the woolly mammoth in the recent elephant subcutaneously at the head located between the eye and ear musth - gland demonstrated in the annual cycle secretions eliminated, which increased aggressiveness caused the dominance and mating behavior.

Adult woolly mammoths had no natural enemies, but young animals could be separated from the herd by large predators such as the cave lion and then torn down. Only early humans, as active hunters with long-range weapons, were dangerous to the woolly mammoth. In addition, there were various dangers that affected the life of the woolly mammoth. Slippery slopes, flood-bearing rivers in spring or collapsing thermal karsts could lead to serious falls or even death of individual individuals. Occasionally, individual broken bones, for example on the shoulder blade , testify to such events. It is possible that a large part of the ice mummies can be traced back to such random events.

As with the recent elephants, it can be assumed that the woolly mammoth reacted very emotionally to the death of its conspecifics, mainly members of the herd, and often stayed at the place of death for up to several hours. Since the animals that stayed behind mostly avoided these places later on, this behavior may also have had an impact on the hunting strategies and collecting activities of early humans.

Pathologies and parasites

Furthermore, individual symptoms of illness can also be observed. Molar and tusk anomalies in the form of growth disorders occur relatively frequently, as do periodontitis and, in some cases, tooth decay . The isolated occurrence of surplus molars in older individuals is also remarkable. Occasionally, canker sores on teeth have also been found. In terms of bone diseases, arthritis of the vertebrae or even entire sections of the spine have grown together. Osteomyelitis could also be proven in some finds .

Investigations on the Berjosowka , Kirgiljach and Shandrin mammoths, among other things , revealed indications of several parasites . Cobboldia and Protophormia , for example, are two external parasites from the two-winged group . Stomach debris also contained remnants of roundworms and tapeworms as internal parasites.

Woolly mammoth and human

The woolly mammoth as a supplier of food and raw materials

As with many large Pleistocene mammals, active hunting by early humans is difficult to prove, since hunting weapons , which are mostly made of organic material, are extremely rarely found. In principle, all remains of a woolly mammoth could be used, for example meat, fat and bone marrow as food, bones and ivory as raw materials for equipment, tools and jewelry , the fur as clothing or covering for residential buildings such as tents , and sinews for threads and cords. In November 2012, the almost complete skeleton of a woolly mammoth was excavated in Changis-sur-Marne in the Seine-et-Marne department . Apparently flint fragments were also discovered. Whether the animal was hunted or used after it was possibly submerged in the mud is still being researched.

Early finds of human remains and woolly mammoths come from the Middle Paleolithic (300,000 to 40,000 years ago) of the first half of the Vistula glacial period. At the 60,000-year-old site in Salzgitter-Lebenstedt (Lower Saxony), bones and teeth from at least 16 mammoth individuals along with around 1,000 flint artifacts have been identified, along with other animal remains . It is not known whether the mammoths were also shot by the Neanderthals of that time . Since the reindeer dominates the site with around 80 individuals, it is more likely a group of specialized reindeer hunters. In the 90,000 year old layer A of Königsaue (Saxony-Anhalt) in the northern Harz foothills, remains of at least four younger mammoth individuals were found that were associated with devices from the Micoquien . A possible direct use of a woolly mammoth on site or even active hunting could be the finding from Asolo (Italy), which includes more than 50 bone fragments from an adult female mammoth in direct connection with five flint artifacts, including two Levallois points . In Siberia, too, several sites from the Middle Paleolithic are known where mammoth bones are associated with human remains, eight in the northern part alone and at least two in the southern part on the upper reaches of the Yenisei . The use of mammoth bones as raw material is demonstrated by the well-known bony, around 15 cm long hand ax from Rhede (North Rhine-Westphalia) or individual processed bones from the Kůlna Cave in Moravia . A unique find comes from Tata ( Hungary ), where a lamella of a lower mammoth molar was completely polished by Neanderthals, the edges rounded and the surfaces covered with ocher. Since this object was reworked but not usable, it is one of the rare finds of non-utilitarian objects (objects that have not been used), which show a possible early emergence of artistic expression in the Neanderthals.

In the subsequent Upper Palaeolithic (40,000 to 11,600 years ago), which was carried by anatomically modern humans, the woolly mammoth was also an important raw material resource. But even here, despite the huge distribution area, there is only very little evidence of active hunting of this large mammal. More than 3500 remains of at least 28 mammoth individuals are known from the layers of finds of the Vogelherd cave (Baden-Württemberg) from the Aurignacia period. Not only was the meat eaten, but bones and ivory were also processed, as finds of ivory rods show, which - based on the natural curvature of the tusks - were straightened. A similar, approximately simultaneous finding comes from the Istállóskö Cave in the Bükk Mountains (Hungary). During the following Gravettia , the use of woolly mammoths as a resource in southern and southeastern Central Europe (also called Pavlovia here ) increased significantly. At the Krems-Wachtberg station in Lower Austria , the animal species is the dominant species. Particularly on the vertebrae and ribs, there are clear cuts that were obviously made when the meat was removed. Individual ribs have also been processed into devices. Particularly noteworthy are the accumulations of woolly mammoth bones at the Dolní Věstonice (Moravia) site, which also belongs to Gravettia , where over 6300 bone and tooth remains of more than 156 mammoth individuals were found on an area of ​​around 12 by 45 m. There are similar findings from Predmosti and Milovice (both also Moravia). The use of the bones and tusks of the woolly mammoth was particularly important in Eastern Europe. In Mesyn alone , the remains of more than 100 mammoths were found, while there were 110 in Meschyritsch (also Ukraine). Both sites belong to the Eastern European Epigravettia (corresponds to the Magdalenian Central Europe) and date around 15,000 BP. The mammoth ossuary from both sites are spectacular, but also from Dobranichevka and Kiev, Kirillovskaja Ulica (all Ukraine). In Mezin alone there are five remains of huts. the best preserved remnant was 5 m in diameter. On its periphery there were 14 skulls of the proboscis, along with long and pelvic bones, while on the inside there were mainly shoulder blades and lower jaw. These obviously formed the walls of the round hut, which later collapsed. The woolly mammoth was also important as a supplier of raw materials in eastern Eurasia, albeit not as prominently as in eastern Europe. In northern Siberia, woolly mammoths and humans occur together at at least ten sites.

After the last Ice Age maximum advance 20,000 to 16,000 years ago and in the subsequent Magdalenian period , the woolly mammoth appears only rarely in Western and Central Europe. For example, the remains of a single individual have come down to us at the important camp site in Gönnersdorf ( Rhineland-Palatinate ). In eastern Eurasia, too, where the woolly mammoth was more common at that time, its importance as a source of raw materials is gradually declining. In the late phase of the Vistula Glaciation , mammoth remains are only found at every third archaeological site in the Yenisei region , while in the previous period it was still detectable in almost two thirds of all human settlement areas. Relatively few finds are known from other regions of Siberia. A settlement site for human hunter-gatherer groups on the shores of Bjorjoljoch is important here, only about 100 m from the famous and contemporary mammoth cemetery , which suggests that the members of these groups used this cemetery as a source of raw materials.

The woolly mammoth in Upper Paleolithic art

Mammoth representation from the cave of Les Combarelles (France)

Fully plastic mammoth representation from the Vogelherd cave (near Niederstotzingen , Swabian Alb ), approx. 40,000 years old ( Aurignacien ), mammoth ivory, UNESCO World Heritage " Caves and Ice Age Art in the Swabian Jura ", Museum of the University of Tübingen MUT

The fact that the woolly mammoth was one of the most impressive animals of the mammoth steppe is shown by the cave paintings and small art of the Upper Paleolithic (from around 37,000 years ago). The oldest mammoth representations of the Franco- Cantabrian cave art from the Chauvet grotto (France) are assigned to the Aurignacien . At least 34 images, alternately in red or black pigments or engraved in the rock, have survived and represent the third most common animal species after the woolly rhinoceros and the cave lion . Characteristic is the almost horseshoe- shaped line that shows the belly and the inside of the legs. The woolly mammoth is particularly often depicted in the southern French cave of Rouffignac , the images of which belong to the Magdalenian cultural stage and where there are at least 150 drawings of this proboscis. Some of the woolly mammoths are reproduced very realistically, with high skull humps, sloping back line and strongly curved tusks. For the recognition of the authenticity of the cave art of Rouffignac, the fact that the depiction of the dorsal valve as an anatomical detail was only rediscovered in paleozoology after the discovery of the drawings was important. With regard to anatomical details, the bull , known as the patriarch , with striking tusks and two mammoths, which face each other forehead to forehead and seem to be celebrating a ritual fight or a welcoming ceremony, should be emphasized . Other frequent representations of the woolly mammoth can be found in Font-de-Gaume and Pech Merle with 23 images each and in Les Combarelles with 14 images, which is why these are also referred to as "mammoth shrines". Overall, the woolly mammoth comprises six to seven percent of all animal and human representations in the caves of Frankokantabria. Outside of this cultural area, the red depictions of the animal species from the Kapova cave in the Urals are known.

The woolly mammoth reproduces the cabaret of the Upper Palaeolithic as full and half sculptures. Among the best known are those from the Aurignacian settlement phase of the Vogelherd cave and the Geißenklösterle (both in Baden-Württemberg); they are among the oldest works of art known to man. During archaeological excavations in the Vogelherd cave (Swabian Alb) in 1931 and in the spoil in front of the cave from 2006, a total of five mammoth sculptures - some of them fragmented - were discovered. A relief representation of a mammoth on a bone fragment from the Vogelherd cave is significant. The palm-sized figures made of mammoth ivory and bone are part of the UNESCO World Heritage " Caves and Ice Age Art in the Swabian Jura ". A somewhat more recent depiction comes from Sungir (Russia). From the following Gravettia, the mammoth figures from Pavlov and Dolní Věstonice (both Czech Republic ) should be emphasized, which are made of fired clay and are among the oldest ceramic finds in the world. More functional were spear throwers in the form of mammoth representations, as they have come from Bruniquel or Canecaude (both France) and belong to the Magdalenian.

Thrower of Bruniquel (France) from the Magdalenian

In addition, the more than 60 depictions of the proboscis from the Magdalenian settlement of Gönnersdorf, which are carved into slate, are outstanding . Two different groups can be highlighted here: animals with a saddle behind the hump of the skull and sloping back line and those with a convex back, the highest point of which is roughly in the middle. While the former are interpreted as fully grown animals, the latter are intended to represent young animals. Also from the “Devil's Bridge” near Saalfeld (Thuringia) comes a piece of clay slate with a mammoth depiction showing only the head and trunk. Other images of this proboscis were found in La Madeleine (France), a site that helped define Magdalenian. The animal, which appears excited here with its tail raised, is executed with numerous details such as the characteristic pony or throat hair. Representations on bone plates are also known from Malta near Lake Baikal (Russia).

Originally no representations of the mammoth were known from North America. Recently, however, such a find was reported, which shows a clearly reproduced proboscis with a high head hump, sloping back line and twisted tusks. The depiction, which is only 7.5 cm long, is carved into the long bones of a large mammal (mammoth, mastod or giant sloth ) and is around 13,000 years old. Since the find comes from Florida (main channel of Vero Beach ), where the woolly mammoth itself has not been detected, it is obviously an image of a prairie mammoth that occurred here.

The Lady of Brassempouy (France) from Gravettia

Bones and tusks were also used as raw materials for Upper Paleolithic art, and some of the most important artifacts of the period are made from these organic materials. Some of the mammoth figurative sculptures already mentioned are made of ivory, such as those from the Vogelherd cave, but also other animal figures, including a horse, a big cat, a bear or a bison from the same cave. The outstanding works of art also include the Aurignacia- era lion man from the Hohlenstein- Stadel, the Venus vom Hohlefels or the adorant figure from the Geißenklösterle (all in Baden-Württemberg), as well as the head depictions of the Lady of Brassempouy (France) and a similar figurine from Dolní Věstonice. Painted bones from Mezin (Ukraine) or those with complex incised decorations in tusks from Předmostí (Czech Republic) have also survived.

Some musical instruments were also made of ivory, as the finds of flute fragments from the Vogelherd Cave , Geißenklösterle and the Hohlen Fels , all in the valley of the Ach near Blaubeuren , prove. Finally, jewelry and art objects were also used in the Upper Paleolithic funeral rite. The grave find of a man and the double burial of two children from the early to middle Young Paleolithic station Sungir are outstanding. In addition to pierced ivory discs, over 3500 ivory beads were found distributed over the body of the deceased, especially in the men's grave, which made it possible to reconstruct the clothing precisely, while an ivory lance around 2.4 m long was added to each of the two young people. In addition, complete bones from woolly mammoths were also placed in the graves. In the Gravettia, shoulder blades were used to cover burials. Examples of this are the graves of Dolni Vestonice, Pavlov or Předmostí, but also the children's double grave in Krems-Wachtberg, which was discovered only a few years ago.

Tribal history

Origin and development

Schematic representation of the change in skull morphology in different mammoth species. Left: southern elephant ( Mammuthus meridionalis ), middle: steppe mammoth ( Mammuthus trogontherii ), right: woolly mammoth ( Mammuthus primigenius ).

The genus Mammuthus originates from Africa , where it is recorded as Mammuthus subplanifrons in the early Pliocene and as Mammuthus africanavus in the Pleistocene as well. From Mammuthus subplanifrons the southern elephant ( Mammuthus meridionalis ) developed, which almost three million years ago also set foot on Eurasian soil as one of the first representatives of the elephants. The steppe mammoth split off from this around 1.5 million years ago. The individual evolutionary stages were accompanied by changes in the skeletal anatomy, including the elongation of the occiput and thus the formation of a high cranium , but also the gradual curvature of the tusks and changes in the molar structure. The latter show a significant increase in the enamel lamellae with a simultaneous decrease in the thickness of the tooth enamel. While the southern elephant only had 13 to 18 enamel lamellae with an average thickness of 2.0 to 3.9 mm, its phylogenetic successor, the steppe mammoth, already had between 17 and 23 with a thickness of 1.0 to 3.5 mm. The increase in melting folds is a sign of greater adaptation to open landscape conditions and increasing specialization in the resulting grass food.

The steppe mammoth reached its largest area in the late Old Pleistocene around 800,000 years ago and at that time came from East Asia to Western Europe. With a shoulder height of up to 4.5 m and a weight of 10 t, it was one of the largest representatives of the elephants. The woolly mammoth with its characteristic properties ultimately developed from the steppe mammoth. The exact process has not yet been finally clarified. The starting point for the development of the woolly mammoth is probably in eastern Eurasia. In northeast Siberia, in the Olyor fauna complex , the first advanced mammoths appeared 800,000 to 600,000 years ago, which are characterized by a higher number of lamellae (22 to 24) in the molar teeth and a greater tooth height. The process is then completed 200,000 years ago with the appearance of the typical woolly mammoth. This region seems to be one of the centers of the development of the woolly mammoth.

In western Eurasia the steppe mammoth undergoes a significant reduction in body size at the same time. While relatively large individuals were found in Süßenborn (Thuringia) and Mosbach 500,000 to 600,000 years ago, in the late Middle Pleistocene 200,000 years ago they were much smaller, as finds from Ilford (England) and Ehringsdorf (Thuringia) show. As a result, there is also a slight increase in the number of lamellae, but this has not yet reached the typical high number as in the later woolly mammoths and averages 19 lamellae. The tooth crowns do not yet achieve the values ​​of the woolly mammoth that will later be used.

The morphological change of this kind in the steppe mammoth led many researchers in western Eurasia to suspect that this was a mixed or transitional form to the woolly mammoth. Therefore, the designation Mammuthus trogontherii-primigenius was often used for the mammoth finds from the late Middle Pleistocene to indicate this intermediate position; The term Mammuthus primigenius fraasi was even introduced for the remains from Steinheim , which are around 300,000 years old and have been reconstructed to form a 3.7 m high skeleton . Some researchers even questioned the species status of the steppe mammoth, but this was strictly rejected by most. Today the older finds of the steppe mammoth are called Mammuthus trogontherii trogontherii and the younger ones as Mammuthus trogontherii chosaricus .

However, the first woolly mammoths appeared in Europe as early as the early Saale Glaciation around 250,000 years ago and lived together with steppe mammoths , such as the mammoth from Pfännerhall from the Geiseltal. In the late Saale Glaciation around 200,000 years ago, only mammoth forms can be detected in western Eurasia that can no longer be separated from the typical Weichsel glaciation. Balderton (England) and Zemst ( Belgium ) belong to such late hall cold period woolly mammoth finds . In the course of the Glaciation of the Vistula, there is another size reduction, which is usually understood as a further climatic adaptation. Some researchers also assume a fluctuating body size, which includes larger individuals in warm climatic phases ( interstadials ) and smaller individuals in cooler phases ( stadials ), but there is no correlation with absolute age values. Since the woolly mammoths sometimes show clear regional population differences, there is a confusing large number of subspecies in the literature.

die out

→ Main article: Quaternary extinction wave

Period of extinction

For a long time it was assumed that the last woolly mammoths in Eurasia became extinct at the end of the last glacial period around 12,000 years ago. However, the woolly mammoth had its most frequent appearance before the last glacial cold maximum. During the cold maximum it disappeared from large parts of western Eurasia, only to return during the subsequent rewarming ( Bölling-Interstadial ). The population density was now much lower, and it no longer reached southern Europe either. After this warm phase ended, it was extremely rare in Western and Central Europe. The most recent data so far come from Gough's Cave (England) and Les Coudrays (France) and are 14,600 and 14,700 (calibrated) years ago ( BP ) respectively . However, it must have been found here around 1000 years later, since Gönnersdorf not only has mammoth remains, but also numerous pictures of this species. In Northern Europe, the woolly mammoth survived until the last cooling phase ( Younger Dryas ) at the end of the Vistula glacial period, while in Eastern Europe it apparently survived until the beginning of the Holocene. Young finds here come from Puurmani ( Estonia ) with around 10,100 to 10,200 BP and Tscherepowez in northern Russia with around 9800 BP.

In western Siberia the species also appeared in the Allerød-Interstadial , but then also disappeared during the Younger Dryas. The most recent date comes from finds from the Juribei and is 11,700 years ago in the transition from the Pleistocene to the Holocene. Populations from the Taimyr Peninsula and on the Chukchi Peninsula can be detected extending further into the Holocene. There the most recent data of finds from the Lower Taimyra are about 9600 years ago.

The last members of the species lived on Wrangel Island until 3700 years ago , so survived comparable to the time of the Egyptian pharaohs or the Central European Bronze Age . The oldest human remains on this island also date from this time, leading to the assumption that humans exterminated the animals through heavy hunting. Genetic studies from 2015 and 2020 showed that the population on Wrangel Island, which possibly consisted of no more than 300 to 500, a maximum of 800 individuals due to the size of the island, had numerous function- changing mutations , especially in the final phase , caused by inbreeding . This includes, among other things, a decreased fertility of the bulls, an increased susceptibility to disease and developmental disorders. The animals also obviously suffered from a diminished sense of smell, which for example prevented the perception of certain plant scents. But this was essential for the search for food. On the North American continent, the woolly mammoth apparently also survived into the early Holocene. The last representatives on the mainland can be found around 10,700 years ago in the area of ​​the Yukon . On St. Paul Island, which is part of the Pribilof Islands and was separated from Alaska by rising sea water 13,000 years ago, a population was discovered that survived into the mid-Holocene. They are relatively small woolly mammoths, but their small size is not considered to be a complete island dwarf like on Wrangel Island. The most recent data from the Qagnax Cave of Sankt Paul is 5725 BP ( ¹⁴ C years).

Extinction theories

• The extermination hypothesis, also known in its extreme form as the overkill hypothesis or blitzkrieg hypothesis , primarily makes humans responsible for the decline of the Pleistocene megafauna. The woolly mammoth was one of the hunting animals of the people in the Upper Pistocene . This is documented by numerous cave paintings and a large number of mammoth bones accumulations in archaeological sites of the Aurignacia, Gravettia and Epigravettia. Whether excessive hunting (“overkill hypothesis”) caused the animals to become extinct or rapid climatic changes at the end of the Ice Age (warming in the Allerød Interstadial ) is still a matter of dispute. Since the mammoth and other Ice Age mammals had survived many violent climatic fluctuations before, a human influence on the complete disappearance is plausible. A study by C. Johnson suggests that the extinction of the woolly mammoth and other Pleistocene species was accompanied by a rapid decline in fertility. According to these analyzes, it was not the largest species of the Pleistocene that became extinct, but those with the lowest reproduction rates. Consequently, the Blitzkrieg hypothesis should be rejected as a particularly rapid extinction wave through targeted hunting of the largest species. However, the extermination of the megafauna, including the woolly mammoth, by growing hunter-gatherer populations is consistent with the results of the analysis.
• In addition to the extermination hypothesis, the climate hypothesis is the most frequently cited explanation for the extinction of large animal fauna at the end of the Pleistocene. The gradual disappearance of the woolly mammoth partly coincides with the warming of the climate at the end of the Pleistocene. A particularly problematic aspect of this hypothesis is the fact that there were numerous climatic fluctuations, some of which were significantly greater, throughout the Pleistocene, which did not lead to mass extinctions. As a variant of the climate hypothesis postulated the so-called impact hypothesis as the cause of climate fluctuations the alleged impact ( impact ) of one or more large astronomical objects sparse approximately 12,900 years (± 100 years) in the northern North America. The impact was the cause of the sudden strong cooling of the Younger Dryas , which in turn was the cause of the extinction of the Pleistocene megafauna in North America (including the woolly mammoth) and the extinction of the prehistoric Clovis culture of the Paleo-Indians . The hypothesis was confirmed in 2007 by Richard B. Firestone et al. published . However, a study published in 2008 of the demographic development of the Paleo-Indians for the period in question found no indication of the population decline mentioned in the hypothesis. The alleged signs of the impact could also not be confirmed by an independent research group.

A recent study found that a combination of both factors, climate and humans, is the most likely cause of the woolly mammoth's extinction. The climatic changes between the late Pleistocene and the early Holocene led to the fact that the woolly mammoth lost large parts of its range. In the end it was restricted to the arctic regions of Siberia. This was accompanied by a sharp decline in the population, which made the species vulnerable to human hunting. This eventually apparently led to the species' ultimate extinction.

Systematics and genetics

Internal systematics of the elephants according to Meyer et al. 2017.

Elephantidae	Loxodonta      Paleoloxodon       Elephas      Mammothus

Reconstruction of woolly mammoth (left) and American mastodon (right)

The woolly mammoth as a representative of the genus Mammuthus belongs to the family of actual elephants, to which the more recent genera Elephas with the Asian elephant, Loxodonta with the African elephant and the forest elephant ( Loxodonta cyclotis ) and the extinct genus Primelephas are assigned. Common to all representatives of the elephants is the lamellar structure and the pronounced high crown ( hypsodontia ) of the molars and the formation of the tusks from the upper incisors.

The woolly mammoth emerged from the steppe mammoth and forms with it a sister line to the prairie mammoth and the dwarf mammoth ( Mammuthus exilis ) on the American continent. According to molecular genetic studies, the two lines separated about two million years ago. Also the genome of that species, consisting of about 4 Gb ( Giga - base pairs ) which has been studied and new using DNA sequencing methods was a large proportion (3.3 Gb) to decrypt it. The resulting estimated genetic difference between woolly mammoths and African elephants was less than 0.6% and is only about half the size of that between chimpanzees and humans. The woolly mammoth is even more closely related to the Asian elephant. Expressed in absolute age data, this means that the lines Elephas and Mammuthus separated 6.7 million years ago, while Loxodonta had split off 7.6 million years ago. The mammoths, a primeval species of proboscis, including the American mastodon, which survived into the Upper Pleistocene, went their own evolutionary path as early as 26 million years ago.

As early as the beginning of the 21st century, DNA studies on at least six different individuals showed that the woolly mammoth apparently consists of two genetic groups. Later analyzes of at least 18 Siberian finds - for example the Jarkow mammoth , the fish hook mammoth , Dima and the Adams mammoth - confirmed this view. The two woolly mammoth lines were called clade I and clade II. While representatives of Klade I were widespread across northern Eurasia and North America, those of Klade II have so far been limited to the region between the Lena and Kolyma. It also seems to have died out at least 33,000 years ago. What caused the separation of the two groups, whether selection or genetic drift , has not yet been clarified; anatomical or functional changes do not seem to have caused them. According to previous studies, the splitting of the two clades happened a million years ago.

In addition, analyzes of the mitochondrial DNA of 160 mammoth individuals from the entire Holarctic showed that the woolly mammoth apparently not only spread via the Bering Strait to North America, but also occasionally returned to Eurasia and thus exhibited dynamic colonization behavior. Five haplogroups (A to E) were identified, of which group C was originally only native to North America until 47,000 years ago and apparently represented the original wave of immigration. Until 22,000 years ago, the representatives of the four other groups also migrated via the Bering Strait to North America, while the population there was now also to be found in northern Eurasia. The mixed groups then survived until the end of the last freezing phase, while the representatives of Group E were the last woolly mammoths on Wrangel Island.

Furthermore, the DNA code for the hemoglobin of a 43,000-year-old woolly mammoth had three sequences that differed from the hemoglobin of an Asian elephant. The hemoglobin gene of the woolly mammoth was introduced into bacteria in 2010, whereupon they produced the mammoth protein. It is hoped that the different oxygen saturation between the hemoglobin of the woolly mammoth and that of the Asian elephant will provide information about the mammoth's adaptation to cold.

So far, no genes have been identified that cause the woolly mammoth's thick hair in comparison to today's elephant forms. The Fgf5 gene is responsible for elongated hair in many mammals. Long-haired mice, cats and dogs often have a destroyed Fgf5 gene. Sequence comparisons between woolly mammoths and various modern elephant forms revealed hardly any mammoth-specific peculiarities in the Fgf5 gene. The same is true for the gene KRTHAP1 which at least the different between humans and other very hairy part apes effected. African elephants and woolly mammoths each have an intact KRTHAP1 gene. The keratin genes KRT25, KRT27 and KRT83 do not seem to be responsible for the difference between today's elephants and woolly mammoths. The most likely candidates for the abnormal hair development in woolly mammoths are likely to be differences in other keratin genes or keratin-associated protein (KRTAP) genes.

Recreation of the woolly mammoth

For a long time there have been considerations to create a woolly mammoth using genetic engineering methods. Concrete plans to clone the woolly mammoth with the help of frozen cells have so far failed due to the strong fragmentation of the DNA . At the beginning of 2011, the retired professor Akira Iritani ( University of Kyoto ) announced that he would start another attempt to extract intact genetic material from the remains of frozen mammoths. The team wanted to use a technique that would enable researcher Teruhiko Wakayama to clone a mouse that had been frozen for 16 years. Another possible approach is to artificially create the woolly mammoth's chromosomes on the basis of sequence data, to pack them into a cell nucleus and finally to smuggle them into an elephant egg cell. This egg cell could then be planted in a cow elephant, which, if successful, would carry a mammoth. However, the methods required for this are currently not advanced enough to create a mammoth in this way. So far, you have not been able to artificially recreate entire chromosomes. Another way to create the woolly mammoth would be to gradually modify the elephant's genetic makeup through gene targeting in order to get closer to the mammoth with every step. This technique is currently already established in other animals, but has not yet been tested in the case of the elephant. When it comes to the implantation of egg cells or embryos, the natural anatomy of the elephant in particular presents considerable obstacles. The very long generation of elephants would also make this approach a real long-term project. In the medium term, at least one animal could emerge in this way that is very similar to the woolly mammoth and could possibly even take on its ecological role. In the long term, it would theoretically also be possible in this way to generate an animal that genetically corresponds almost completely to the woolly mammoth.

In 2019, a team of researchers working with Kazuo Yamagata succeeded for the first time in extracting cell nucleus- like structures from the bone marrow and muscle meat of the Yuka mammoth, discovered in 2009, and implanting them in the egg cells of mice. The mammoth cell nuclei began to take over proteins from the mouse cells and develop further nucleus-like structures. However, cells did not divide because the mammoth cell nuclei were obviously too badly damaged. The study's scientists emphasize that it is not possible to clone a mammoth with current nucleotide technology, but they see their results as an important step in this direction.

Research history

Johann Friedrich Blumenbach

Finds of woolly mammoths were discovered very early in western Eurasia, but often not recognized as a fossil elephant species. The 1577 the monastery Reyden ( Lucerne discovered) mammoth bones were from the former doctor F. Plater as remnants of a 19  foot high giant interpreted. Likewise, the skeleton of a woolly mammoth unearthed in 1663 near Seveckenberg near Quedlinburg (Saxony-Anhalt) in the presence of the natural scientist Otto von Guericke (1602–1686) was regarded as a remnant of the mythical Unicornu fossile - it was similar with the first discoveries recovered around 33 years later European forest elephants near Tonna (Thuringia) - and later also reconstructed as such. In 1696, the Russian Ludloff described the remains of an animal found in Siberia, which the local Yakuts and Tungus called Mamantu and, in their opinion, resembled a giant mole that would die as soon as it saw the sunlight.

The origin of the term Mamantu or Maimanto is not clear. Often an origin in the Nenet or Estonian language area is sought. In the latter language maa means "earth" and mutt means "mole". However, a connection is occasionally made to the Arabic word Behemoth - a huge monster with curved horns and tusks - and justified with the Siberian ivory trade by Arabs, which has been documented since the 9th century. Even who ultimately introduced the name in Europe is not sufficiently certain.

The woolly mammoth was scientifically described for the first time in 1799 by the Göttingen natural scientist and anatomist Johann Friedrich Blumenbach (1752-1840) as Elephas primigenius based on finds near Osterode am Harz . In the same year the carcass of the Adam's mammoth was discovered on the Lena. The sketch of this find made by Roman Boltunow in 1806 was later passed on by Henry Michail Adams to Blumenbach in Göttingen , who recognized it as his recently described species of elephant. The genus name Mammuthus comes from the English biologist Joshua Brookes (1761-1833), who he introduced in 1828.

Cultural history

As an extinct animal, the woolly mammoth was of great importance early in human history of the post-Ice Age. The indigenous peoples of Siberia and northern North America mainly used mammoth ivory for carvings. The trade in fossil ivory from Yakutia by Arabs has been documented since the 9th century, and was later continued by the Russian Empire . From 1800 to 1914 it is estimated that between 20 and 32 tons of ivory were traded annually, which corresponds to a total of 25,000 to 46,000 mammoth individuals. Since the trade in ivory from living elephants was banned in 1989, fossil ivory is once again of considerable importance, for example for ivory carving . In China, fossil mammoth tusks were seen as the teeth of "ice rats" and, similar to the Pleistocene animal remains that are estimated as dragon bones , were ground into powder and sold as medicine . In Europe, they were sometimes considered to be the remains of the unicorn until modern times and, when crushed to powder, were also important as medicinal products.

But even according to its scientific description, the mammoth has entered human culture. Today, the term mammoth stands for something big or difficult to manage (such as the “mammoth task”). It also found its way into heraldry and is represented in the coats of arms of Srednekolymsk (Republic of Sakha, Russia) or of Seedorf (Lower Saxony). It is also a popular element in modern pop culture and appears in films (including In the Beginning Was The Fire by Jean-Jacques Annaud , Ice Age and sequels) and novels (such as in the cycle Children of the Earth by Jean M. Auel ) . The adventure series Syberia also takes hold , v. a. in the second part, the mammoths as a leitmotif.

literature

• Miles Barton: Wild America. Witnesses to the Ice Age. Vgs, Cologne 2003, ISBN 3-8025-1558-7 .
• Robert Darga and Hans Steiner: Southeast Bavarian Natural History and Mammoth Museum Siegsdorf. (= Bavarian Museums. Volume 25). Weltkunst, Munich 1998, ISBN 3-921669-25-1 . For the history of the mammoth's discovery, see pp. 121 to 126.
• Wighart von Koenigswald: Living Ice Age. Theiss, Stuttgart / Sigmaringen 2002, ISBN 3-8062-1734-3 .
• Adrian Lister and Paul Bahn: Mammoths - The Giants of the Ice Age. Thorbecke, Sigmaringen 1997, ISBN 3-7995-9050-1 .
• Ronald M. Nowak: Walker's Mammals of the World. 6th edition, Johns Hopkins University Press, Baltimore MD et al. a. 1999, ISBN 0-8018-5789-9 .
• Reinhard Ziegler: The mammoth (Mammuthus primigenius Blumenbach) from Siegsdorf near Traunstein (Bavaria) and its accompanying fauna. Munich Geoscientific Treatises Series A: Geologie und Paläontologie 26, Munich 1994, pp. 49–80.

Individual evidence

1. ↑ ^{a b c d e f g h} Ralf-Dietrich Kahlke: The origin, development and distribution history of the Upper Pleistozönen Mammuthus-Coelodonta Faunenkomplexes in Eurasia (large mammals). Treatises of the Senckenbergische Naturforschenden Gesellschaft 546, Frankfurt am Main 1994.
2. ↑ ^{a b c d e} Diego J. Álvarez-Lao, Ralf-Dietrich Kahlke, Nuria García, Dick Mol: The Padul mammoth finds - On the southernmost record of Mammuthus primigenius in Europe and its southern spread during the Late Pleistocene. Palaeogeography, Palaeoclimatology, Palaeoecology. 278, 2009, pp. 57-70.
3. ↑ Diego J. Álvarez-Lao and Nuria García: Geographical distribution of Pleistocene cold-adapted large mammal faunas in the Iberian Peninsula. Quaternary International 233, 2011, pp. 159-170.
4. ↑ ^{a b} Ralf-Dietrich Kahlke: The maximum geographic extension of Late Pleistocene Mammuthus primigenius (Proboscidea, Mammalia) and its limiting factors. Quaternary International 379, 2015, pp. 147–154, doi: 10.1016 / j.quaint.2015.03.023 .
5. ↑ ^{a b c d e f} Dick Mol, Alexei N. Tikhonov, Johannes van derpflicht and Dmitry Yu. Bolshiyanov: Discoveries of woolly mammoth, Mammuthus primigenius (Proboscidea: Elephantidae) and some other Pleistocene mammals on the Taimyr Peninsula. Russian Journal of Theriology 2 (2), 2003, pp. 77-95.
6. ↑ ^{a b c d e f g h i j k l m n} Adrian Lister and Paul Bahn: Mammuts - The Giants of the Ice Age . Sigmaringen, 1997.
7. ↑ ^{a b c} Yaroslav V. Kuzmin: Extinction of the woolly mammoth (Mammuthus primigenius) and woolly rhinoceros (Coelodonta antiquitatis) in Eurasia: Review of chronological and environmental issues. Boreas 39, 2010, pp. 247-261.
8. ↑ ^{a b} Ute Koch: Mammoth steppe habitat. In: Ulrich Joger and Ute Koch (eds.): Mammuts from Siberia. Darmstadt, 1994, pp. 55-73.
9. ↑ ^{a b c d e f g h} Rudolf Musil: Survival strategies of elephants. In: Harald Meller (Hrsg.): Elefantenreich - Eine Fossilwelt in Europa. Halle / Saale, 2010, pp. 323–336.
10. ^ AK Markova, AN Simakova, A. Yu. Puzachenko and LM Kitaev: Eastern European mammoth distribution and environments during the Middle Valdai Briansk Interstade (33,000-24,000 BP). In: G. Cavarretta, P. Gioia, M. Mussi and MR Palombo: The World of Elephants - International Congress. Consiglio Nazionale delle Ricerche Rome, 2001, pp. 299-304.
11. ↑ ^{a b c d e f g} Wighart von Koenigswald: Lebendige Eiszeit. Climate and fauna in transition. Stuttgart, 2002, pp. 54-61.
12. ↑ Wei Guang Biao, Hu Song Mei, Yu Ke Fu, Hou Ya Mei, Li Xin, Jin Chang Zhu, Wang Yuan, Zhao Jian Xin and Wang Wen Hua: New materials of the steppe mammoth, Mammuthus trogontherii, with discussion on the origin and evolutionary patterns of mammoths. Science China - Earth sciences 53 (7), 2010, pp. 956-963.
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14. ↑ ^{a b c d e f} Ulrich Joger: History (s) of the mammoth finds. In: Ulrich Joger and Ute Koch (eds.): Mammuts from Siberia. Darmstadt, 1994, pp. 9-23.
15. ↑ ^{a b} Gennady G. Boeskorov and Dick Mol: Quaternary Mammal Collections in the Museums of Yakutsk (Eastern Siberia, Yakutia, Russia). Cranium 21 (1-2), 2004, pp. 19-32.
16. ↑ Volker Toepfer: The mammoth finds from Pfännerhall in the Geiseltal . Publications of the State Museum for Prehistory in Hall 16, Hall / Saale, 1957.
17. ^ Wighart von Koenigswald: Lebendige Eiszeit. Climate and fauna in transition. Stuttgart, 2002, p. 47.
18. ↑ ^{a b c} Ralf-Dietrich Kahlke and Dick Mol: Ice Age Large Mammals of the Siberian Arctic. The Cerpolex / Mammuthus expeditions on Tajmyr. E. Schweizerbart'sche Verlagbuchhandlung (Nägele and Obermiller), Stuttgart 2005.
19. ↑ CR Harington: Woolly mammoth. From: Canadian Museum of Nature in Yukon Beringia Interpretive Center ( online ) and ( PDF ).
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21. ↑ Dick Mol and Ralf-Dietrich Kahlke: The fishhook mammoth - a Late Pleistocene (Early Sartanian) individual of Mammuthus primigenius with preserved gut contents from the Upper Taimyra River (Taimyr Peninsula, Arctic Siberia). Abstracts of the 18th International Senckenberg Conference April 25-30, 2004 in Weimar
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23. ↑ ^{a b c} Dick Mol, Jeheskel (Hezy) Shoshani, Alexei Tikhonov, Bas van Geel, Shin-ichi Sano, Peter Lazarev, Gennady Boeskorov and Larry D. Agenbroad: The Yukagir Mammoth: Brief history, 14C Dates, individual age, gender , size, physical and environmental conditions and storage. Scientific Annals, School of Geology Aristotle University of Thessaloniki (AUTH), special volume 98, 2006, pp. 310-313.
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27. ↑ Eugene Maschenko, Olga Potapova, Gennady Boeskorov, Valery Plotnikov and Larry Agenbroad: Preliminary data on the new partial carcass of the Woolly mammoth, Mammuthus primigenius, from Yakutia, Russia. Journal of Vertebrate Paleontology 32 (suppl. 2), 2012, p. 137.
28. ↑ Gennady G. Boeskorov, Olga R. Potapova, Eugeny N. Mashchenko, Albert V. Protopopov, Tatyana V. Kuznetsova, Larry Agenbroad and Alexey N. Tikhonov: Preliminary analyzes of the frozen mummies of mammoth (Mammuthus primigenius), bison (Bison priscus) and horse (Equus sp.) from the Yana-Indigirka Lowland, Yakutia, Russia. Integrative Zoology 9, 2014, pp. 471-480.
29. ↑ Anastasia S. Kharlamova, Sergei V. Saveliev, Albert V. Protopopov, Busisiwe C. Maseko, Adhil Bhagwandin, and Paul R. Manger: The Mummified Brain of a Pleistocene Woolly Mammoth (Mammuthus primigenius) Compared With the Brain of the Extant African Elephant (Loxodonta africana). The Journal of Comparative Neurology 523, 2015, pp. 2326-2343.
30. ↑ Evgeny N. Maschenko, Olga R. Potapova, Alisa Vershinina, Beth Shapiro, Irina D. Streletskaya, Alexander A. Vasiliev, Gleb E. Oblogov, Anastasia S. Kharlamova, Eugene Potapov, J. van der Plicht, Alexey N. Tikhonov , Natalia V. Serdyuk and Konstantin K. Tarasenko: The Zhenya Mammoth (Mammuthus primigenius (Blum.)): Taphonomy, geology, age, morphology and ancient DNA of a 48,000 year old frozen mummy from western Taimyr, Russia. Quaternary International 445, 2017, pp. 104-134, doi: 10.1016 / j.quaint.2017.06.055 .
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110. ^ Spiegel Online - Wissenschaft: Japanese want to clone mammoth. ( [4] ) January 17, 2011.
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• Mammoth Museum: The woolly mammoth