New photos from the Auerrind project

 Claus Kropp has recently posted some new photos of the cattle from the Auerrind project: 

left: Maremmana x Sayaguesa, right: Maremmana x Watussi (© Claus Kropp)

Maremmana x Watussi (© Claus Kropp)

(Maremmana x Watussi) x (Sayaguesa x Chianina) (© Claus Kropp)

Maremmana x Sayaguesa (©Claus Kropp)

The Maremmana x Sayaguesa bull seems to develop formidable horns. I love his curly hair on the forehead, a typical trait of the European aurochs. The Maremmana x Watussi bull has large horns as well. On Facebook, Claus Kropp wrote that a pure Sayaguesa or Sayaguesa x Chianina might be crossbreeding options for this bull, and I completely agree with that. The young (Maremmana x Watussi) x (Sayaguesa x Chianina) bull should be old enough to have its final colour, apparently it inherited some colour dilutions. However, if it gets large and well-proportioned I would still consider him a useful individual (colour is easy to breed), especially if he gets good horns. 

The Holocene European wild horse

Recently I did a life reconstruction for the Holocene European wild horse: 

Since there is not a single complete skeleton of a Holocene European wild horse, I based the body, head and proportions on Przewalski's horses and also primitive pony breeds. A recent study suggests that Holocene European wild horses were predominantly black [1]. As for the mane, it cannot be ascertained today if the western wild horse subspecies, Equus ferus ferus, had an erect mane or a falling mane. All cave paintings show erect manes, and all extant wild equines have an erect mane. It has been suggested that a falling mane would be advantageous in a more humid climate because it diverts rain water, but domestic horses were domesticated in the arid steppe. It is well possible that a falling mane is a trait that arose during domestication. Hence, I drew my horse with an erect mane. 

[1] Sandoval-Castellanos et al.: Coat colour adaption of post-glacial horses to increasing forest vegetation. 2017

Why do some Polish koniks have erect manes?

While searching on google for Konik horses from Poland, which is where the breed originated and should have the most genetic diversity, I found a number of individuals with an erect mane. See this young individual, this one from Popielno (one of the main Konik breeding sites), or this one from the Roztoczanski Park Narodowy. Why is that? 

Some Heck horses, which are almost one popualtion with the Konik in Germany, have erect manes because Lutz Heck bred with a Przewalski stallion and Przewalski's horses have been crossed-in again at the Wildpark Hardehausen in order to achieve an erect mane. The results can be seen here. Breeders in Germany often do not differentiate between Koniks and Heck horses, so that the breeds are often intermixed. 

However, the Koniks in Poland should be free of that comparably recent intermixture, as Heck horses are found mostly in Germany. Nevertheless it is possible that even "pure" modern Koniks have Przewalski introgression. The reason for that is that some of the Koniks that have been used in Lutz Heck's experiment were returned to Poland after the war. It is well possible that some of these Koniks were part Przewalski's horse due to Heck's breeding. The fact that some Polish koniks have erect manes endorses this assumption, at least in my opinion, as I find no other explanation for this trait in the Polish stock. 

Probably only a genetic test could bring clarification. 

Genetic breeding-back?

Some modern aurochs projects claim they carry out selective breeding on a genetic level. In an article from 2017, I outline why I doubt this. First of all, we have every reason to assume that a lot of, or at least some, key wildtype alleles have been lost during the process of domestication. Furthermore, although the full genome of an aurochs has been resolved, the wildtype alleles of this specimen have not been traced down in modern cattle individually so that selective breeding for those alleles would be possible. And lastly, the number of involved loci might be very, very high and the selection would probably be a century-long project. 

Another practical problem for "genetic selection" would be genetic linkage. Genes do not get passed on individually, but are inherited via chromosomes. Let us say we have ten key aurochs alleles on ten loci on the same chromosome. Breed A has five of those alleles on the loci 1-5, and mutated (=domestic) alleles on loci 6-10. Breed B the other five alleles on loci 6-10 and mutated alleles on loci 1-5. No matter how you crossbreed those breeds and select the offspring, it will not be possible to unite these ten alleles in one individual, unless you get very lucky with recombination, which is unlikely. This is a descriptive example of course, but it is very likely that in many cases wildtype and domestic alleles are linked on the same chromosome. This could even be a problem for traditional "breeding-back" as it is executed by all modern projects. 

Nevertheless, let us ignore those practical obstacles for now and assume that "genetic selection" is feasible. Which animals would be needed in order to unite as much genetic material of the aurochs present in living bovines? A set of primitive European taurine breeds will not be enough in order to seize the maximum potential. Zebus have been shown to share wildtype alleles with the British aurochs whose genome was sequenced that taurine cattle have lost [1]. Therefore, zebus would have to be included in a "genetic selection" project. It is likely, or perhaps very likely, that there are also alleles which taurine cattle lost and are not preserved in zebus, therefore have been lost in cattle altogether. Some of them, however, could be shared by closely related wild bovines, such as the banteng, gaur, yak, or the two bison species. This would have to be tested. Probably also a wider range than just Southern European primitive breeds would be needed in order to seize the genetic potential of taurine cattle. What about Near Eastern cattle, or Asiatic taurine cattle such as the Turano-Mongolian group? It is not unlikely that they have wildtype gene material not found in in European taurine cattle. 

Thus, if one really is to carry out a project that tries to get as much genetic material from the aurochs as possible, not only European taurine cattle, but maybe also taurine cattle of other genetic groups, surely zebuine cattle and perhaps even related bovine species would have to be used. I still think such a project is not practically feasible, but it would be interesting "how far" such a project would get to approximate the aurochs. However, it might be a lot easier to recreate a genuine aurochs via CRISPR-Cas9. For my thoughts on what to do with a genetically recreated aurochs, go here. 

[1] Orlando et al.: The first aurochs genome reveals the breeding history of British and European cattle. 2015. 

Leptobos and the origin of Bos and Bison

In the most recent phylogenies based on mitochondrial DNA, Bos turns out to be paraphyletic. The yak is grouped as a sister taxon to Bison, and banteng and gaur form a clade that is sister to yak and bison [1,2]. The wisent is a sister taxon to cattle in this phylogeny, was has been interpreted as the result of hybridization. The most recent study, however, considers it more likely that this is the result of incomplete lineage sorting [1].  

Regarding the origin of Bos, there are two conflicting hypothesis. One that suggests an African origin, and one that suggests an Asiatic origin. Advocates of the "out of Africa" hypothesis suggest an origin from Pelorovis oldowayensis [3]. Pelorovis, however, is considered to be too different by other authors. They suggest an origin from Leptobos instead, which is described as more similar to Bos [4]. 

Leptobos comprises a group of fossil bovine species with very variable horns. Leptobos is also suggested to be the direct ancestor of Bison. And indeed the cranium of L. vallisarni bears striking resemblance to bison skulls, so that some authors list it as Bison vallisarni instead. Leptobos stenometopon, on the other hand, has a horn curvature that is reminiscent of that of aurochs, kouprey and yak. This horn curvature could either be basal to the Bos-Bison clade or evolved several times. The fact some species of Leptobos resemble Bos species while others resemble bison in cranial anatomy might imply that different species of Leptobos gave rise to the modern Bos clades (aurochs/cattle, the banteng-gaur and possible also kouprey clade) and bison. In this case, Leptobos would be a genus that includes the basalmost members of the Bos-Bison clade. This is only my personal speculation. More complete skeletal material would be needed in order to better resolve the paleontological evidence for the origin of the Bos and bison species. 

Leptobos etruscus - is Leptobos the ancestor of the Bos-Bison clade?

[1] Wang et al.: Incomplete lineage sorting rather than hybridization explains the inconsistent phylogeny of the wisent. 2018. 
[2] Verkaar et al.: Maternal and paternal lineages in cross-breeding bovine species. Has the wisent a hybrid origin? 2004. 

[3] Martinez-Navarro et al.: The Olduvai buffalo Pelorovis and the origin of Bos. 2007. 

[4] Tong et al.: New fossils of Bos primigenius (Artiodactyla, Mammalia) from Nihewan and Longhua of Hebei, China. 2014. 

A zebuine ancestry for Chianina?

Genetic studies have helped a great deal to understand the history of domestic cattle populations in recent years. 
For example, it has been resolved that many Southern European cattle breeds have influence from North African taurine cattle [1], which is not surprising considering that there were trade routes in ancient times. North African taurine cattle are genetically distinct from other taurine cattle, which is interpreted as the result of significant introgression of African aurochs [1]. Therefore, many Iberian and some Italian breeds might have African aurochs in their ancestry. For Chianina (and related breeds such as Romagnola, Marchigiana and others) in particular, zebuine influence has been detected as well [1]. This does not surprise me that much, as I have been suspecting that the white colour of Chianina (produced by at least two different alleles, on the Agouti and Dun locus) is actually inherited from zebus. Some zebu breeds have exactly the same white colour as Chianina, for example see the Nelore breed. Also, the face of Chianina looks slightly zebuine to me, as well as the fact that it lacks curly hair on the front head (which is typical for zebuine cattle but rare in taurine cattle). 

So Chianina is influenced by zebuine cattle. This might be used as an argument against the use of Chianina in "breeding-back" by those who want to use Maremmana instead for large size. However, Podolian cattle - such as Maremmana - are significantly influenced by zebuine cattle as well [2,3]. This also shows in the phenotype: I suspect that the upright horns, large dewlap and Agouti dilution of Podolian cattle are derived from zebuine cattle. 

But I think this is neither an argument against Chianina or Maremmana. Zebuine influence is simply not all that uncommon in taurine cattle, and unavoidable for "breeding-back", as many breeds needed for certain traits, such as size, have zebuine influence. 

As an interesting side note, it has been recognized that zebus share some wildtype alleles with the British aurochs whose genome was fully sequenced, while taurine cattle have other alleles on these loci [4]. So zebus do have some alleles in common with the European aurochs. 

[1] Decker et al.: Worldwide patterns of ancestry, divergence and admixture in domestic cattle. 2014. 

[2] Papachristou et al.: Genomic diversity and population structure of the indigenous Greek and Cypriot cattle. 2020. 

[3] Upadhyay et al.: Genetic origin, admixture and population history of aurochs (Bos primigenius) and primitive European cattle. 

[4] Orlando et al.: The first aurochs genome reveals the breeding history of British and European cattle. 2015. 

Three more Tauros bulls

I have been covering Tauros cattle a lot here recently, and I have found some more photos of three Tauros bulls that are interesting. 
This includes a bull at Maashorst. The horns are really good. The size is ok, the curvature is basically right but should be more intense in order to be perfect. Actually the horns are better than in most Taurus cattle. The body shape and proportion cannot be judged because the bull is lying. 

Another bull is from Keent. I wonder what the breed combination of the bull is. I suspect that it is part Highland x Maremmana, the body shape reveals Highland influence and the long dewlap and the horns point to Maremmana. Since the bull is from Keent, Manolo Uno (Maremmana x Pajuna) might be the father. In this case the bull would be (Maremmana x Highland) x (Maremmana x Pajuna), what I consider plausible. 

The third bull is from Keent too. I have no idea what combination this one could be. 

Looking at the horns of the upper two bulls, I might reconsider my pessimism concerning the horns of Tauros cattle. The sizes of the horns are useful, and the curvature of the horns of the first bull is really good. 

K. L. Hartig's aurochs painting

There is an aurochs painting from 1955 that is worth noticing. It's the painting by the German artist K. L. Hartig, which can be seen here. It was done for a publication by the zoologist Hans von Lengerken in the same year. The original was coloured and is untraceable today, unfortunately. What is special about this painting is its accuracy. Hartig took measurements from a nearly complete skeleton and a skull from the collection of the natural history museum of Berlin, so the painting can arguably be called a reconstruction. The only aspects to criticize are that the shoulder hump is to low and the snout too short, perhaps also the head too small, but otherwise it is pretty accurate. The fact that the hump is too low was recognized in the literature as early as in 1957. While most historic aurochs reconstructions were rather imprecise or incorrect, with Hartig's painting there is at least one that gave an accurate impression of the aurochs' life appearance already back in the 1950s. 

The painting inspired me to do my own version of it. I based it on the Sassenberg bull skeleton, which I used quite often for my reconstructions. It is a large-horned Holocene specimen, but there were also Holocene aurochs bulls with comparably small horns such as the Himmelev specimen. 

Here is the result: 

Literature 

Walter Frisch: Der Auerochs - das europäische Rind. 2010. 

The Tauros cattle of Kettingdijk, Netherlands

On monday this week, Gerard vanne Smeed posted a blog article with many photos of the Tauros cattle from Kettingdijk in the Netherlands. Also with the link to the flickR page for more photos. It was awesome to the so many photos of a large Tauros cattle herd, as it is pretty hard to find good photos of the project on the web. 

The herd consists of crossbreeds and possibly also pure individuals of Limia and maybe also Maremmana as well as a pure Maronesa bull. The influence of Highland cattle still shows in a number of individuals that have the brindle coat colour pattern (see here or here for example). The Highland influence also shows in the colour and horns of this cow. Some individuals could easily be sold as Heck cattle, such as this bull or these cows. Interestingly, one bull is very pale-coloured, perhaps a combination of Maremmana and Highland colour alleles. There are also bulls with a correct colour such as this one. The adult bull on this photo also looks nice. Concerning the sexual dimorphism, there are lightly coloured cows but also many dark cows, bulls can be wholly black but about the half seems to have a colour saddle. So the sexual dichromatism is not very marked in the herd. Also, the bulls seem to be barely larger than the cows, which might be because they are not fully grown yet. All of the bulls seem to have a hump, which is good. The bulls are muscular overall, they just need longer legs. If I had to rank these Tauros cattle, I would say they are somewhere between Heck cattle and Taurus cattle. 

The Tauros Programme seems to have a different approach to the breeding. While most "breeding-back" projects breed by selecting a chief bull and selecting out individuals that do not fit the standards, the Tauros Programme seemingly let the cattle breed for themselves with many bulls in one herd, with barely any selection (the fact that there are still brindle individuals suggests to me that no selection has taken place yet, as this trait is dominant and easy to select out). Maybe the selection phase has not started yet. I think it is good that the cattle have bull competition in their breeding, as this enables natural selection, but I think this phase is too early for that. It will be very difficult to raise the quality* of the herds this way, except if all bulls were quality bulls. Again, maybe the selective breeding phase has not started yet and they will start selective breeding in the next months or years. 
 

*By quality I mean the resemblance to the aurochs in its morphological traits, not the surviving capacity, behaviour or other traits. 

Przewalski's horse cloned for genetic diversity

In 2015 I proposed cloning as a chance for the wisent's survival. Cloning pre-bottleneck wisents would greatly increase the genetic diversity of the species, since the modern population descends from a population of only about 50 individuals which itself descended from only 12 individuals. Adding the genetic diversity of wisents that lived before the dramatic bottleneck event in the 20th century would help the species to overcome its inbreeding depression. And if cloning is not possible, genome editing with CRISPR-Cas9 is a viable alternative. 

It seems that there are people who had the same idea for the Przewalski's horse, which descends from only 12 individuals as well. A stallion has been cloned from an individual that has been cryopreserved since 1980. For details, you can have a look at the article from Revive & Restore. 

I hope that cloning for conservation will not be restricted to this one individual. I hope this idea will be put into practice for other species as well, including the wisent. 

Reconstruction of aurochs cow horns

Recently I did a reconstruction of the horns of the Asti specimen at the Museo paleontologico territoriale dell'Astiagiano. 

Reconstructing the horn sheaths on the bony cores always requires speculation to a certain degree. The horn sheath continues the curvature of the bone, but there is no rule of thumb on how much the sheath increases length and thickness of the horn. Especially the length of the sheath may vary greatly in comparison to the bony core (see van Vuure 2005). So we can only guess how much the sheath added to the horns in the individual specimen. I usually reconstruct the horn sheath with 133% of the length of the bony core. 

Here is the result: 

Please do not use without permission. 

In modern cattle, Watussi, Heck cattle of the Wörth lineage and Maronesa may have horns similar to those above. A mix of all three might reach the goal both in curvature and dimensions. 

Another reconstruction of the Asti specimen is about to come. 

Tauros cattle arrived in Velebit, Croatia

As Rewilding Europe recently announced, 20 Tauros cattle (18 cows, two bulls) have been moved to Velebit, Croatia. They will join a herd of 120 individuals that consist of pure Sayaguesa and Maremmana as well as crosses of Sayaguesa, Maremmana and Tudanca that have been grazing there since 2016. The purebred bulls will be removed in turn if I understand correctly.
You can see photos of the herd here and here. 

Based on their looks, the herd might include Maremmana x Sayaguesa, Limia x Maremmana and Pajuna x Maremmana individuals. But that is only a guess. I doubt that these fully grown individuals are fifth-generation. First of all, the Tauros Programme counts the parental generation as first-generation, what means that their second-generation animals are actually the first cross generation. So that means they mean fourth-generation. Also that seems a far stretch for a breeding period of 11 years. Body shape and proportions are OK, they are pretty much the standard we see in Taurus cattle and primitive cattle breeds. So is the skull shape. The horns, however, are not that impressing, as in most Tauros cattle. They are to small and the curvature is not aurochs-like either. What I also noticed is that the cows are mostly pretty dark, what is true for most Tauros cattle. I wonder why. Surely, Sayaguesa contributes dark colour shades in cows but other projects achieve a nice reddish-brown coat colour even in half-Sayaguesas. Maremmana, Limia, Tudanca and Pajuna also have lightly coloured cows. So I wonder why most Tauros cows are that dark. 

The Tauros Programme, after 11 years of breeding, now has reached a level of good quantity. They have a total animal count in the three-figure range. What the programme needs is to improve the sexual dichromatism (i.e. achieve more lighter-coloured cows) and to improve the horns in both size and curvature. There are no size measurements for Tauros cattle (at least none that have been published) but the cattle at Keent (NL) do not look large. So they also might want to improve the size of the animals, considering that they are using a number of small breeds (Pajuna, Highland cattle, Maronesa). While selection can improve the sexual dichromatism and if they pay attention on only using bulls with good horns also the horn shape, I doubt that they can achieve the improvement of the horns and body size with the breeds they are currently using. The horn size of many Maremmana (the largest-horned breed they are using) is actually smaller than in the aurochs (see here, for example). Maremmana is also the largest breed they are using, and Maremmana does reach sizes of 170cm and allegedly more, but their Maremmana individuals don't look large (one of their Maremmana bulls was barely larger than the Pajuna bull, and the other one didn't look much taller than the Highland individuals). 
Summa summarum, I think the Tauros Programme needs breeds that add large size, truly large horn volume, and also a good horn curvature when starting the quality-building phase otherwise Tauros cattle might end up deficient in terms of horns and body size. 

A Watussi steer with perfectly aurochs-like horns

Recently I found a video of a Watussi individual with very aurochs-like horns. Normally, Watussi bulls' horns look more like that. This individual however has horns that have a perfectly aurochs-like curvature (yes the horns are very large, though there were Pleistocene aurochs bulls with horns only a little smaller). I was not surprised that this individual is obviously a steer. Steers often tend to have more aurochs-like horns. Why is that? I explain this in this post. Horn length and consequently shape is determined by developmental factors. Development is deeply changed in domestic animals, usually there is a premature stop in development, causing the animals to display paedomorphic features. Thus it is not far-fetched to assume that horns tend to stop growing earlier in domestic animals, producing the horn shapes they have (usually less curved than in the aurochs). Steers, however, have an atypical development as their gonads are removed. The body does not know when to stop development and thus they grow larger, taller and also the horns grow longer (for another example, go here). Coincidentally, this developmental elongation compensates the developmental delay that results from domestication, resulting in a wildtype-like outcome. This is what we see in the Watussi steer. 

This has two implications for "breeding-back". For once, it means that bulls with banana-shaped forwards-facing horns (as we see it in many primitive breeds and "breeding-back" cattle) actually have the right genes for an aurochs-like horn curvature (the horns would also end up longer). It are developmental factors that result in the domestic condition. Furthermore, the big question is then: can "breeding-back", with the cattle that we have, produce perfectly aurochs-like horns at all? Selecting just on the phenotype that we want would not reverse the developmental changes from wildtype to domestic that we have in cattle. However, there are occasionally bulls with horns very close to the original aurochs horn shape, such as some Maronesa bulls (and also cows). And maybe it is not a coincidence that this breed has also retained a substantial degree of colour dimorphism. 

Two Tauros bulls with good horns

Most examples of Tauros cattle from the Tauros Programme don't have that good horns. Mostly they face outwards too much and lack the inwards curve. This goes for cows as much as for bulls. However, today I was addressed to two photos of two Tauros bulls with comparably good horns from the Loozerheide, Netherlands. 

Go here and here. The head of the first bull resembles some Taurus bulls quite closely. The second bull might have Maronesa ancestry but I could be wrong on that. 

It's really good to see Tauros bulls with such nice horns, it makes me hope that future Tauros cattle will develop better horns than the current majority. However, I think the project still needs a breed that reliably and truly adds slenderness, long legs and large size. Maremmana alone probably will not do it. 

New photos from the Auerrind project

Claus Kropp recently sent me interesting photos of some of the Auerrind crosses, which I am going to present today. 

© Claus Kropp

This is the Chianina x Watussi cow born in 2019. Her horns are developing well. She will be covered in 2021. Here is another photo of her, next to a Sayaguesa x Chianina cow: 

© Claus Kropp

In sum, there are three Chianina x Watussi cows, two calves have been born this year: 

© Claus Kropp

I wonder what would be the ideal combination to cross them with. Claus Kropp told me one option is the Sayaguesa x Maremmana bull. There are also Sayaguesa x Maremmana cow calves, one of them down below: 

© Claus Kropp

It apparently inherited the colour of a Sayaguesa cow.  Here is a new photo of the Maremmana x Watussi bull: 

© Claus Kropp

There is no plan yet as to which combination it could be crossed with. 

More Sayaguesa x Chianina calves have been born: 

© Claus Kropp

© Claus Kropp

The cow Maxima on the upper photo has a very good and useful horn curvature. The cow in the back at the lower photo with the asymmetric horns (I think it is La Nova) is interesting as well; the question is which horn shape gets passed on. If she passes on the horn shape of her left horn it would be perfect. 

It is very exciting to see those interesting combinations growing up and I am looking forward to the second-generation animals they will produce. The second generation is also where the selection starts. Since both Leo the Sayaguesa bull and Luca the Maremmana bull are about 170cm tall, many Auerrind crosses might end up being on the larger side. 

Tauros bulls, a Taurus birth and a new Auerrind calf

Today I have news from all three main aurochs breeding back projects. 

Tauros bulls 
I found some photos of Tauros bulls on the web that I haven't posted here yet. Here, here and here. The first bull is from Herpeduin, the other two photos are from Maashorst. On the last photo, I particularly like the bull in the front. The colour is perfect and the horns look pretty good as well. They have the best curvature I have seen so far in a Tauros bull. 

Birth of a Taurus calf 
I found three videos showing the birth of a Taurus bull calf. Here, here and here. It is interesting that the cow gave birth in the open field. Normally they would look for a shelter and give birth there, but not all cows have the instinct to do that. 

A new Auerrind calf 
Claus Kropp posted a new photo of a second-generation cow calf today: 

© Claus Kropp

Its name is Doro, and it is (Sayaguesa x Watussi) x (Sayaguesa x Grey cattle). I am looking forward to see it growing, it has the potential to look very good. Claus Kropp also told me that they recently measured Leo the Sayaguesa bull. It is 171cm tall at the withers, which is a very good size. In about two weeks, Alvarez the Sayaguesa x Watussi bull is going to be moved to the herd with the Sayaguesa x Chianina cows. 

Aurochs cow artwork

This is a reconstruction of the female skull at the Gramsberg Museum in the Netherlands. I particularly like the horn shape of this specimen. A mix of Sayaguesa, Watussi and Maronesa might have the potential to result in horns like these. And Chianina for the body shape. Although they do not use Maronesa, let's see if the Auerrind project will achieve cows like that. Cows of the combination (Chianina x Watussi) x (Sayaguesa x Watussi) might have the potential to get close. 

Bos primigenius suxianensis

The classic convention is that there are three subspecies of aurochs – Bos primigenius primigenius, the European one, Bos primigenius namadicus, the Indian one and Bos primigenius africanus, the African one. Here is a chart for the distribution of the subspecies: 

As you see, material from Central and East Asia are included in the European subspecies primigenius. In western literature, the East Asian aurochs material is described as very similar to that of Europe (van Vuure, 2005). 

However, there is Asian literature that is barely recognized in the western scientific community that assigns the material to its own subspecies, Bos primigenius suxianensis [1]. A 2018 study found that aurochs remains were pretty abundant in Neolithic China, and that much of the material has been wrongly assigned to Bison previously. It was also found that East Asian aurochs belonged to a unique haplotype (haplotype C) and therefore were genetically distinct from other aurochs populations [2]. Chinese aurochs were probably hunted, and most likely also hunted to extinction as were the European populations. 

Not only were those East Asian populations genetically distinct, the bone material that is available on the web shows some morphological differences to the European subspecies. The horns are always long and more upright than what is average in the European population, and the shape is slightly different as well. They do not curve inwards that strongly, and they curve more upwards at the base. You see that in this and these specimen. Also, the nasal bones are somewhat raised and more convex than in the European subspecies. You see that in this Chinese specimen and the Baikal skull. 

I did a life reconstruction of the specimen linked above: 

 

Life reconstruction of Bos primigenius suxianensis

Note the different horn shape and the convex snout. Nothing is known about the coat colouration of East Asian aurochs, but for this drawing I assumed it had the same colour as the European subspecies. 

So it seems that Central and East Asian aurochs were genetically and also morphologically distinct populations, justifying its subspecies status. Therefore, there actually were four wild aurochs subspecies. The reason why the Asiatic subspecies is not part of the “basic aurochs knowledge” is probably that the Chinese literature is less recognized in the western scientific community, and that the bone material (which even includes nicely preserved complete specimen) is less noticed compared to the plentiful European material, except for the Baikal skull. It would be interesting to know if there was a continuum between primigenius and suxianensis, as the distribution area apparently was continuous at least some of the time of their existence. The Kiev specimen, which is the Easternmost specimen of the European subspecies that I know of, does have some similarities in horn shape. 

The aurochs apparently was a species with a wide geographical range and regional variations. The European subspecies which is the “standard” when we think of an aurochs, the African subspecies that apparently had a colour saddle, the Indian aurochs with its slender cranium and the long wide-ranging horns, and the East Asian aurochs with the slightly different horns and the convex snout. When we look at other bovines with a wide geographical range such as the Banteng or the Cape buffalo, there might have been more local colour variants that we do not know of. 

Literature 

[1] Xie: A skull of Bos primigenius suxianensis from Anhui. 1988.  

[2] Cai et al.: Ancient DNA reveals evidence of abundant aurochs (Bos primigenius) in Neolithic Northeast China. 2018. 

Quagga life restoration

Today I did another Quagga life restoration, based on the Amsterdam skin. 

I drew it in the same posture as this zebra of the Quagga Project so that both can be easily compared. 

The stripe pattern on the Quagga is different from those of the zebras of the project. The stripes are broader, with a much smaller space in between, especially on the head. Also, the brownish background colour of the trunk is not quite achieved yet. Furthermore, I suspect that there are more differences between Quaggas and Burchell's zebras. For example, the mane seems to be shorter in each of the skins and the photos of the London mare. Also, it might be that the ears are smaller. Although being nested within the Burchell's zebra, the quagga has unique haplotypes identifying the subspecies. 
The zebras of the Quagga project, on the other hand, are simply Burchell's zebras with a reduced stripe pattern, not more than that. This is why I wrote Please don't call it Quagga. 

Aurochs horn variation

I described the horns of the aurochs in a 2013 post already. The basic curvature of the horns was always the same in the aurochs, but there was considerable variation on the actual length, thickness, shape and orientation of the horns within a certain range. 

I tried to capture this variation on two drawings in 2015 where I reconstructed the horn sheaths onto 22 skulls in order to see what the horns might have looked like in life. I am going to repost them down below. We cannot be absolutely sure on the life appearance of the sheaths as there was no general rule of thumb of how much the sheath adds to length and thickness of the horn cure. There was quite some variation (van Vuure, 2005). So the reconstructions down below are an approximation. 

For the identity of the skulls, go to the 2015 post. 

Recently I did some more sketches, also including skulls that I already reconstructed in 2015: 

From left to right top down: Berlin skull, a skull of a location unknown to me, the Cambridge cow, Gramsberger Museum skull, Himmelev specimen, Horsholm specimen, a cow skull from Italy, the Stuttgart skull and the Vig bull. 

Having reconstructed about 30 specimen, I think this is a fairly representative sample for the variation of the horns of the European subspecies Bos primigenius primigenius. 

Orientation relative to the skull 

The literature states that the orientation of the horns relative to the skull varied from 50 to 70° (van Vuure, 2005). However, having had a look at so many skulls I find that the range is actually larger. The oldest aurochs skull which was discovered in 2014 had an orientation of 40°. The Vig specimen has an orientation of 90°, and the Horsholm specimen probably an even larger angle. 

Geographic variation

I see some sort of geographic correlation in the variation of the horn types. For example, the more Southern the skulls, the sharper is the angle between the horns and the snout. The more Northern and Eastern, the higher is the orientation of the horns. You see that very clearly in the Kiev specimen and the Eastern Asian aurochs Bos primigenius suxianensis (yes, apparently Eastern Asian aurochs were a distinct subspecies, more on that in an upcoming post). Small-horned aurochs seemingly only appeared in Northern Europe, all other locations (Southern Europe, Africa, Asia) had pretty large-horned aurochs.