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Aurochs - Bos primigenius; ancestor of domestic cattle
Topic Started: Apr 3 2012, 01:08 AM (484,007 Views)
phantom
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Aurochs - Bos primigenius

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Temporal range: Late Pliocene to Holocene

Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Artiodactyla
Family: Bovidae
Subfamily: Bovinae
Genus: Bos
Species: Bos primigenius

The aurochs (Bos primigenius), the ancestor of domestic cattle, was a type of large wild cattle which inhabited Europe, Asia and North Africa, but which is now extinct; it survived in Europe until the last recorded aurochs, a female, died in the Jaktorów Forest, Poland in 1627. Her skull is now the property of the Livrustkammaren ("Royal Armory") museum in Stockholm, Sweden.

During the early Holocene in course of the Neolithic Revolution, aurochs were domesticated in at least two domestication events: one concerning the Indian subspecies, leading to Zebu cattle, the other one concerning the Eurasian subspecies leading to taurine cattle. Other species of wild bovines were domesticated as well, such as the Wild water buffalo, Gaur and Banteng. In modern cattle, there are numerous breeds that share characteristics of the aurochs, such as a dark colour of the bulls with a light eel stripe and light colour in cows, or a typical aurochs-like horn shape.

Taxonomy and Etymology
The animal's original scientific name was Bos primigenius. This scientific name is now considered invalid by the Integrated Taxonomic Information System (ITIS), which classifies aurochs under Bos taurus – the same species as domestic cattle. In 2003, however, the International Commission on Zoological Nomenclature "conserved the usage of 17 specific names based on wild species, which are pre-dated by or contemporary with those based on domestic forms", confirming Bos primigenius for the Aurochs. Taxonomists who consider domesticated cattle a subspecies of the wild Aurochs should use B. primigenius taurus; the name B. taurus remains available for domestic cattle where it is considered to be a separate species.

The words "aurochs", "urus", and "wisent" have all been used synonymously in English. However, the extinct aurochs/urus is a completely separate species from the still-extant wisent. The two were often confused, and some 16th century illustrations of aurochs and wisents have hybrid features. The word urus (/ˈjʊərəs/) comes to English from Latin, but may have come to Latin from Germanic origins. It declines in English as urus (singular), uruses (plural). In the German language, Ur derived to Auer in course of a diphthongization in the language during the 13th century. Later, "-ochs" as added, which is meant to refer to a wild bovine. This is how the German name of the animal turned to Auerochs/Auerochse.

The word "aurochs" comes to English from German, where its normal spelling and declension today is Auerochs/Auerochse (singular), Auerochsen (genitive), Auerochsen (plural). The declension in English varies, being either "auroch" (singular), "aurochs" (plural) or "aurochs" (singular), "aurochses" (plural). The declension "auroch" (singular), "aurochs" (plural), acknowledged by MWU, is a back-formation analogous to "pea"-from-"pease" derived from a misinterpretation of the singular form ending in the /s/ sound (being cognate to "ox/Ochs(e)"). The use in English of the plural form "aurochsen" is not acknowledged by AHD4 or MWU, but is mentioned in The Cambridge Encyclopedia of the English Language. It is directly parallel to the German plural and analogous (and cognate) to English "ox" (singular), "oxen" (plural).

The name of the aurochs in other languages seems to be derived by "urus" as well. Such as uro (spanish language) or urokse (danish language).

Evolution
During the pliocene, the colder climate caused an extension of open grassland, which increased the evolution of large grazers, such as wild bovines. Bos acutifrons is an extinct species of cattle sometimes claimed to be the ancestor of aurochs, but it was a species with very long, outwards-facing horns. The oldest aurochs remains come from about 2 million years, India. Therefore, the Indian subspecies was the first aurochs subspecies to appear. During the pleistocene, the species migrated into the Middle East and further into Asia, and reached Europe about 270,000 years ago. The South Asian domestic cattle, or zebu, descended from Indian aurochs at the edge of the Thar Desert; this would explain the zebu's resistance to drought. Domestic yak, gayal and Javan cattle do not descend from aurochs.

The first complete mitochondrial genome (16,338 base pairs) DNA sequence analysis of Bos primigenius from an archaeologically verified and exceptionally well preserved aurochs bone sample was published in 2010.

Subspecies

Three wild subspecies of aurochs are recognized. Only the Eurasian subspecies survived until recent times.:

The Indian Aurochs (Bos primigenius namadicus), the Indian Aurochs, once inhabited India. It was the first subspecies of the aurochs to appear, at 2 million years ago, and from about 9000 years ago (BP) it was domesticated as zebu cattle. Fossil remains indicate, that there were wild Indian aurochs besides domesticated zebu cattle in Gujarat and the Ganges area until about 4,000-5,000 years ago. Remains from wild aurochs, 4,400 years old, are clearly identified from Karnataka in south India.

The Eurasian Aurochs (Bos primigenius primigenius) once ranged across the steppes and taigas of Europe, Siberia, and Central Asia. It is noted as part of the Pleistocene megafauna, and declined in numbers along with other megafauna species by the end of Pleistocene. The Eurasian aurochs were domesticated into modern taurine cattle breeds around the 6th millennium BC, in the Middle East, and possibly also at about the same time in the Far East. Aurochs were still widespread in Europe during the time of the Roman Empire, when they were widely popular as a battle beast in Roman arenas, and excessive hunting began and continued until it was nearly extinct. By the 13th century, aurochs existed only in low numbers in Eastern Europe, and hunting of aurochs became a privilege of nobles, and later royal households. The decreased hunting did not save the aurochs from extinction, and the last recorded live aurochs, a female, died in 1627 in the Jaktorów Forest, Poland from natural causes. Aurochs were found to have lived on the island of Sicily, where once there was a land bridge to Italy. After the disappearance of the land bridge, Sicilian aurochs evolved to a size 20% smaller than their mainland relatives.

The North African Aurochs (Bos primigenius africanus = mauretanicus), the African Aurochs, once lived in the woodland and shrubland of North Africa. It descended from Aurochs populations migrating from the Middle East. The North African aurochs was morphologically very similar to the Eurasian subspecies, therefore this taxon maybe stands on a biogeographical ground only. However, there is evidence that it was genetically distinct from the eurasian subspecies. North African aurochsen possibly had a light saddle on their backs, as depictions indicate. This subspecies possibly disappeared during the antiquity.

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Description
The appearance of the aurochs is reconstructed by skeletal material, historical descriptions or contemporaneous depictions, like cave paintings, engravings or Sigismund von Herberstein’s illustration. Another drawing often referred to is the work by Charles Hamilton Smith, which is a copy of a painting owned by a merchant in Augsburg, which may go back to the 16th century. However, it has been proposed that this last illustration was actually drawn after a cattle/aurochs hybrid, or even just an aurochs-like breed. Other depictions and descriptions of aurochs appear in prehistoric cave paintings and in Julius Caesar's The Gallic War.

The aurochs was one of the biggest herbivores in the postglacial Europe, only comparable to the Wisent, the European bison. The actual size of aurochs seemingly was dependent on the region. In Europe, aurochs populations of the north were bigger on average than those from the south. For example, during the Holocene aurochs from Denmark and Germany had an average height at the shoulders of 155–180 cm in bulls and 135 to 155 cm in cows, while aurochs populations in Hungary had bulls reaching 155–160 cm. The body mass of aurochs probably was comparable to the Wisent or Banteng, reaching 700 kg on average. The sexual dimorphism between bull and cow was, as it was concerning other aspects as well, strongly expressed, with the cows being several decimetres shorter than bulls on average.
Life restoration of the aurochs based on a bull skeleton from Lund and a cow skeleton from Copenhagen

Because of the massive horns, the frontals of aurochs were elongated and broad. The horns of the aurochs were characteristic for the species concerning size, curvature and orientation. They were curved in three directions: upwards-outwards at the base, then swung forwards-inwards, then inwards-upwards. Aurochs horns could reach 80 cm in length and between 10 and 20 cm in diameter. The horns of bulls were larger with the curvature more strongly expressed than in cows. The horns grew from the skull in a 60° angle to the snout, they are clearly facing forwards. The horn shape of aurochs was an advantage because of the way intraspecific fights were carried out: in contrast to bison, they do not butt their heads against each other, instead they hook themselves with their horns and try to push away the adversary. Modern cattle breeds still exhibit this way of combat fight. As a protection for their eyes, the eye sockets of bulls were much stronger pronounced than in most contemporary cattle bulls.

The body shape of the aurochs was strikingly different from many modern cattle breeds. For example, the legs were considerably longer and slender, resulting in a shoulder height that nearly equalled the trunk length. The skull, carrying the large horns, was substantially larger and more elongated than in most cattle breeds. Only some primitive Mediterranean cattle breeds like Maremmana primitivo or Pajuna Cattle exhibit those traits. The body shape of the aurochs was, like in other wild bovines, athletic and, especially in bulls, showed a strongly expressed neck- and shoulder musculature. Even in carrying cows, the udder was small and hardly visible from the side; this feature is equal to that of other wild bovines.

The coat colour of the aurochs can be reconstructed by using historical depictions and contemporary depictions. In his letter to Conrad Gesner (1602), Anton Schneeberger delivers one of the most precise descriptions of the aurochs, which agrees with cave paintings like those in Lascaux and Chauvet. Calves were born in chestnut colour, young bulls changed their coat colour during few months into a very deep brown or black with a white eel stripe running down the spine. Cows retained the reddish-brown colour. Typical for both sexes was the lightly coloured mouth, which is also found in some bantengs. Except some north African engravings showing aurochs with a light saddle on the back, there is no evidence of a different coat colour of aurochsen in or outside Europe. A passage from Mucante (1596), describing the “wild ox” as gray, is ambiguous and may actually refer to the wisent. Egyptian grave paintings show cattle with a reddish brown coat colour in both sexes with a light saddle, but the horn shape of these suggest they depict domestic cattle. Remains of aurochs hair were not known until the early 80s.

Some primitive cattle breeds still display the coat colour characteristics of the aurochs, like the black colour in bulls with a light eel stripe, a white mouth or the typical coloural sexual dimorphism. A feature that is often attributed to the aurochs are blond fronthead hairs. Historical descriptions tell that the aurochs had long and curly fronthead hair, but none of them mentions a certain colour for it. Cis van Vuure (2005) therefore calls it, although it is present in a variety of primitive cattle breeds, a discolouration after domestication. But he mentions, that Gaurs have a lightly coloured fronthead as well. A gene which is responsible for this feature is not known yet. Zebuine breeds show lightly coloured inner sides of the legs and belly, caused by the so-called Zebu-tipping gene. It has not been tested if this gene is present in the wild form of the zebu, the Indian aurochs.

Habitat, ecology and behaviour
Concerning the habitat of the aurochs there is no consensus. While some authors think that the habitat selection of the aurochs was comparable to the African Forest Buffalo, others describe it as an inhabitant of open grassland helping maintaining open areas by grazing, together with other large herbivores. With its hypsodont jaw, the aurochs likely was a grazer and had a food selection very similar to domestic cattle. Therefore, it was no browser like deer or a semi-intermediary feeder like the wisent. Comparisons of the isotope levels of Mesolithic aurochs and domestic cattle bones showed that aurochs likely inhabited wetter areas than domestic cattle. The description of Schneeberger tells that during winter, along with grasses, twigs and acorns were an additional part of the aurochs’ diet.

After the beginning of the Common Era, the habitat of aurochs became more and more fragmented because of the steadily growing human population. During the last centuries of its existence, the aurochs was limited to hideaway regions such as floodplain forests or marshes, where there were no competing domestic herbivores and less hunting pressure.

Like many bovids, aurochs formed herds for at least one part of the year. They probably did not number much more than thirty individuals, which likely were composed of cows with their calves and some young bulls. Older bulls probably wandered solely or in small bull herds outside the mating season. Assuming that the social behaviour between aurochs and descended domestic cattle is roughly the same, social status was evaluated through gestures and fights, which can occur in cows as well as in bulls. Like in other ungulates which form unisexual herds, there was a considerable dimorphism, because bulls which wander around solely can feed them selves better than within a whole herd, which is why selection forced for especially large bulls. In ungulates which form herds containing animals of both sexes, like horses, sexual dimorphism is weakly developed.
A painting by Heinrich Harder showing an aurochs fighting off a Eurasian Wolf pack

During mating season, which likely took place during the late summer or early autumn, the bulls had severe fights. It is known from the forest of Jaktorów, that fatalities could occur. During autumn, aurochs fed for the winter and got fatter and shinier than during the rest of the year, according to Schneeberger. During spring, the cows gave birth to the calves, for which they went into the forest. The mother stayed at its side as long it was not strong enough to join the herd on the feeding grounds.

Calves were especially vulnerable to wolves, while healthy adult aurochs probably did not have to fear these predators. In prehistoric Europe, North Africa and Asia, big cats like lions or tigers and hyenas were additional predators which likely preyed on aurochs.

Historical descriptions, like Caesar’s De Bello Gallico or Schneeberger, tell that aurochs were swift and fast, and could be very aggressive on occasion. According to Schneeberger, aurochs did not care much about an approaching man. But, teased or hunted, an aurochs could get very aggressive and dangerous, and threw the teasing person into the air, as Schneeberger wrote in his letter to Gesner in the year 1602. But basically, the aurochs likely must have been a peaceful animal to humans, otherwise it would have been hardly usable for domestication.
Domestication and extinction
This specimen is from around 7500 BC and is one of two very well preserved aurochs skeletons found in Denmark. The Vig-aurochs can be seen at The National Museum of Denmark. The circles indicate where the animal was wounded by arrows.

The aurochs, which ranged throughout much of Eurasia and Northern Africa during the late Pleistocene and early Holocene, is widely accepted as the wild ancestor of modern cattle. Archaeological evidence shows that domestication occurred independently in the Near East and the Indian subcontinent between 10,000–8,000 years ago, giving rise to the two major domestic taxa observed today — humpless Bos taurus (taurine) and humped Bos indicus (zebu), respectively. This is confirmed by genetic analyses of matrilineal mitochondrial DNA sequences, which reveal a marked differentiation between modern Bos taurus and Bos indicus haplotypes, demonstrating their derivation from two geographically and genetically divergent wild populations.

Domestication of the aurochs began in the southern Caucasus and northern Mesopotamia from about the 6th millennium BC, while genetic evidence suggests that aurochs were independently domesticated in India and possibly in northern Africa. Domesticated cattle and aurochs are so different in size that they have been regarded as separate species; however, large ancient cattle and aurochs "are difficult to classify because morphological traits have overlapping distributions in cattle and aurochs and diagnostic features are identified only in horn and some cranial element."

However, a DNA study suggests that all the domesticated cattles originate from a group of 80 wild aurochs. Those animals lived in Iran 10,500 years ago.

Comparison of aurochs bones with those of modern cattle has provided many insights about the aurochs. Remains of the beast, from specimens believed to have weighed more than a ton, have been found in Mesolithic sites around Goldcliff, Wales. Though aurochs became extinct in Britain during the Bronze age, analysis of bones from aurochs that lived in the same age as domesticated cattle there showed no genetic contribution to modern breeds. As a result of this study, modern European cattle were thought to have descended directly from the Near East domestication. Another study, however, found distinct similarities between modern breeds and Italian aurochs specimens suggesting that the previously tested British aurochs were not a good model of the diversity of aurochs genetics and suggesting possible North African and European aurochs input to domestic breeds. Further genetic tests have shown that domestic cattle in Europe is of Near Eastern origin, and that the European aurochs was not domesticated, nor did it interbreed with the imported Near Eastern cattle.
Petroglyphs in Fourneau-du-Diable (Devil's Furnace) rock in Bourdeilles, Dordogne, France. Solutrean period, 18,000 BP

Indian cattle (zebu), although domesticated eight to ten thousand years ago, are related to aurochs which diverged from the Near Eastern ones some 200,000 years ago. African cattle are thought to descend from aurochs more closely related to the Near Eastern ones. The Near East and African aurochs groups are thought to have split some 25,000 years ago, probably 15,000 years before domestication. The "Turano-Mongolian" type of cattle now found in Northern China, Mongolia, Korea and Japan may represent a fourth domestication event (and a third event among Bos taurus–type aurochs). This group may have diverged from the Near East group some 35,000 years ago. Whether these separate genetic populations would have equated to separate subspecies is unclear.

The maximum range of the aurochs was from Europe (excluding Ireland and northern Scandinavia), to northern Africa, the Middle East, India and Central Asia. Until at least 3000 years ago the aurochs was also found in Eastern China, where it is recorded at the Dingjiabao Reservoir in Yangyuan County. Most remains in China are known from the area east of 105° E, but the species has been also reported from the eastern margin of the Tibetan plateau close to Heihe River.

By the 13th century A.D., the aurochs' range was restricted to Poland, Lithuania, Moldavia, Transylvania and East Prussia. The right to hunt large animals on any land was restricted to nobles and gradually to the royal household. As the population of aurochs declined, hunting ceased but the royal court still required gamekeepers to provide open fields for the aurochs to graze in. The gamekeepers were exempted from local taxes in exchange for their service and a decree made poaching an aurochs punishable by death. In 1564, the gamekeepers knew of only 38 animals, according to the royal survey. The last recorded live aurochs, a female, died in 1627 in the Jaktorów Forest, Poland from natural causes. The skull was later looted by the Swedish Army during the Swedish invasion of Poland (1655–1660) and is now the property of Livrustkammaren in Stockholm. The causes of extinction were hunting, a narrowing of habitat due to the development of farming, climatic changes, and diseases transmitted by domestic cattle.

Breeding of aurochs-like cattle
In the 1920s two German zoo directors (in Berlin and Munich), the brothers Heinz and Lutz Heck, began a selective breeding program in the attempt to breed back the aurochs into existence from the domestic cattle that were their descendants. Their plan was based on the concept that a species is not extinct as long as all its genes are still present in a living population. The result is the breed called Heck cattle. Heck cattle breeders sometimes refer to it as "Recreated Aurochs", or "Heck Aurochs". It is claimed to bear some resemblance to what is known about the appearance of the wild aurochs in colour and in some cases also horn shape.

Scientists of the Polish Foundation for Recreating the Aurochs (PFOT) in Poland want to use DNA from bones in museums to recreate the aurochs and return this animal to the forests of Poland. The project has gained the support of the Polish Ministry of the Environment. They plan research on ancient preserved DNA. Other research projects have extracted "ancient" DNA over the past twenty years and their results published in such periodicals as Nature and Proceedings of the National Academy of Sciences USA. Polish scientists believe that modern genetics and biotechnology make recreating an animal almost identical to aurochs possible. They say this research will lead to examining the causes of the extinction of the aurochs, and help prevent a similar occurrence with domestic cattle.
The most primitive lineages of Maremmana, called "Maremmana primitivo", are used in TaurOs Project

In a similar program, TaurOs Project is trying to DNA sequence breeds of primitive cattle to find gene sequences which match those found in "ancient DNA" from aurochs samples. The modern cattle would then be selectively bred to try to bring the aurochs-type genes back into a single animal. TaurOs project selected a number of very primitive breeds mainly from Iberia and Italy, such as Sayaguesa Cattle, Maremmana primitivo, Pajuna Cattle, Limia Cattle, Maronesa, Tudanca Cattle and others, which already bear considerable resemblance to the aurochs in certain features. Numerous crossbreed calves have been born already.

-like cattle breeds (selection)
Because some cattle breeds have been changed more considerably than other breeds, there are certain breeds left which still bear a greater resemblance to the aurochs. These breeds are not very productive from the economical point of view, they do not give as much milk or meat as others. Most of the primitive phenotypes are facing extinction because farmers give them up or crossbreed them with more derived dairy and meat cattle. But, because they are very hardy and robust, they sometimes are used in nature conservation programs, where they can fill the place of their wild ancestor. Moreover, some of them are integrated in TaurOs Project, which aims to breed a cattle type phenotyically, genotypically and ecologically as close to the aurochs as possible.

  • Caldela
  • Heck cattle
  • Limia Cattle
  • Maremmana primitivo
  • Maronesa
  • Pajuna Cattle
  • Rhodopian Shorthorn
  • Sayaguesa Cattle
  • Spanish Fighting Bull
  • Tudanca Cattle


Edited by Taipan, Apr 11 2012, 03:21 PM.
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phantom
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Edited by phantom, Apr 3 2012, 02:35 PM.
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phantom
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Finally got around to trying out my experiment.
Here's the result. A very rough drawing of an aurochs over a reconstructed skeleton.
I used the skeleton of a Chillingham bull over the top of a photo of a live Chillingham bull and outlined the outer areas.
Then place the skeleton of the Chillingham, and the outline of the live Chillingham bull, and manipulated it until the Chillingham
lined up with the aurouchs skeleton. This was the result.
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Edited by phantom, Apr 3 2012, 11:33 PM.
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Scalesofanubis
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This needs actual species information and stuff.
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Arctodus Simus
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Cool! Can you do this with a male Lion?
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firefly
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Good work, Phantom.
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phantom
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Arctodus Simus
Apr 5 2012, 03:20 AM
Cool! Can you do this with a male Lion?


I'm pretty sure I could.
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phantom
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firefly
Apr 5 2012, 06:46 AM
Good work, Phantom.
Thanks, Firefly.
Good to see you here.
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phantom
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A while back I posted this picture of an aurochs skull. There was some question of whether this skull was from the Holocene,
or the Pleistocene.
There may be an answer.
You can read clearly the word Holozän, which is german for Holocene.
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Edited by phantom, Apr 8 2012, 03:42 AM.
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Dfoidl
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Hi Phantom,

can't believe I've overlooked that thread! Your restoration of the Kopenhagen skeleton is very, very nice! The only point I'd address is that this Kopenhagen skeleton is likely to be female, based on proportions, and horn size and shape.
Could you please post that Chillingham bull skeleton here? That sounds very interesting.

Interestingly I haven't yet realized that in fact "Holozän" is written below that aurochs skull, although I visited it two times in the Berlin museum... My first thought was, maybe it only addresses when the animal died out, but than I had a look at the steppe bison skull next to it, under which is written "Jungpleistozän", which means "lower Pleistocene", and Bison priscus died out in the upper pleistocene as we know. It seems that the skull is in fact of holocene origin, or is at least tagged to be, although I remember some sources to state that the "huge skull from Berlin" is from the pleistocene.

This is a new restoration done by me which I used for Wikipedia, based on the bull skeleton from Lund and another cow skeleton from Kopenhagen:

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Edited by Dfoidl, May 1 2012, 04:34 AM.
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firefly
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Spectacular work, Dfoidl!

These are true Degerbol aurochsen, right?
They represent some of the biggest aurochsen in Europe, that I´m aware about.
You are putting them, at which size ( at the shoulder)?
Edited by firefly, May 1 2012, 07:43 AM.
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Dfoidl
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Thanks! Yes, the individuals probably belong to the biggest aurochsen, therefore I made the bull around 170 cm at the shoulders and the cow 150 cm tall.
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phantom
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Dfoidl
May 1 2012, 04:34 AM
Hi Phantom,

can't believe I've overlooked that thread! Your restoration of the Kopenhagen skeleton is very, very nice! The only point I'd address is that this Kopenhagen skeleton is likely to be female, based on proportions, and horn size and shape.
Could you please post that Chillingham bull skeleton here? That sounds very interesting.


Hi Dfoidl. I'm glad you found this thread. The Chillingham Bull skeleton can be found here:
http://www.flickr.com/photos/browniebear/4734037591/lightbox/

If the Copenhagen aurochs is a female, I might have to start all over again.

I like your illustration for Wikipedia.
Edited by phantom, May 10 2012, 11:35 AM.
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phantom
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Looks like the Lund skeleton may have been revised to have more of a flat back.
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Edited by phantom, May 13 2012, 03:18 PM.
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Dfoidl
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Thanks for the link to the Chillingham skeleton, very interesting!

Hm, strange how the Lund skeleton has been re-arranged. Instead of correcting the fact that the hind legs are too much bent, they now bend the forelegs too much as well. Too bad that the designers who arrange museum mounts often do not have an idea of the anatomy of the animals - a correct stance would make the skeleton more dynamic, natural, taller and more impressive. It's the same case in many dinosaur mounts.
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