Possible evidence of bulls with a saddle from Europe

 While browsing the web looking for new evidence on the aurochs, I did indeed find something this time. Cave paintings from Roca dels Moros from El Cogul, Catalonia, seem to show aurochs of different colours, two males and one possible female, one with what seems to be what I call the "lateral saddle" we see in indicine cattle. 

 

As you can see, the cave paintings are badly weathered, but it seems to show two dark bulls and one possible reddish female. The male in the middle has, astoundingly, a light area on the side of the trunk that looks like what I call the "lateral zebu saddle". This trait seems to be the same as the dorsal saddle in taurine breeds, just on the side of the trunk instead of the dorsal region in indicine breeds. Which one of the traits is the ancient and which one the derived ones I cannot tell based on the evidence, maybe there even is not a general rule but it is part of the genetic variation of the wildtype (this pattern is most certainly regulated by the Agouti gene). 

This mesolithic cave painting, however, seems to suggest the presence of a "lateral zebu saddle" in an aurochs bull from this particular region and time. I think this is exciting, but I would not jump to conclusions to easily. However, the colour of the bull portrayed might have looked something like in this reconstruction of mine I did for the Wadi Sarrat cranium (just without the light areas on the ventral body side): 

 

 

 

The "Przewalski's horses are just feral Botai horses" hypothesis is outdated

Often in science, there is the problem that a wide-ranging proposal that has a lot of echo in and outside the scientific community (often more in the latter) is not based on a very solid ground. But because of its wide-ranging nature and – I dare not to say sensational – implications, it gets a lot of attention and becomes adopted in many further sources even if the wide-ranging conclusion was premature. But unfortunately, when one or several new studies come out that refute the original proposal, it does not get nearly the same resonance. This way, exciting but not factually correct “new discoveries” become mainstream in and outside the scientific community although they have been refuted already in sometimes several works. One example might be the hypothesis that the wisent is an aurochs hybrid, which is not correct according to recent research. Another example is in the focus of this post, namely the hypothesis put forward by Gaunitz et al. (2018) that the Przewalski’s horse is not a genuinely wild animal but the feralized descendant of early domesticated horses from Botai site, Kazakhstan. For brevity, I call it the Gaunitz hypothesis in this post.

 

The hypothesis

 

The 2018 Gaunitz et al. study looked at 42 ancient horse genomes, including 20 from the eneolithic Botai site. This site was of particular interest for the authors because Botai was or is considered the earliest archaeological evidence for horse husbandy (note: not necessarily domestication in a genetic sense). The big surprise of the study was that all Przewalski’s horses examined in the analysis cluster within the Botai-Borly complex, therefore the authors jumped to the conclusion that the Przewalski’s horse is not a genuine wild animal but a feral descendant of the earliest horse domestication event [1]. Further studies seem to confirm the deep nesting of the PH within the Botai-Borly complex, most notably Librado et al. [2].

Why should a domestic history for the PH surprise us that much? Obviously, because the PH does not show any obvious signs typically associated with domestication in mammals: it has a larger brain volume than domestic horses, a skull that is not paedomorphic at all with small eyes and a large viscerocranium, it has a much higher aggression potential than domestic horses even in captivity and does not have the same mutation accumulation as domestic horses have developed during domestication (known as the costs of domestication [3]). Ever since the 20^th century, the PH was used as a model for a wild horse opposed to the domestic horse. Apart from that: if the PH is not genuinely wild, it would mean that there are no genuinely wild, never-domesticated horses left on earth. So it is well worth investigating whether they truly have a domestic past or not. And it did not take long until the conclusion by Gaunitz et al. was called into question.

 

Were Botai horses domesticated?

 

In 2021, a study by Taylor & Barron-Ortiz was published that called the domestication of the Botai horses into question. They argued that the tooth recovered from the site that seems to show bit wear does not do so, as teeth of Pleistocene wild horses show the same wear pattern [4]. Furthermore, they argue that the equine lipids found on pottery from Botai do not necessarily have to be milk lipids (which would indicate that the mares there would have been tame enough to be milked) but could also be from meat or fat [4]. Also, the human teeth from the site do not show traces of milk proteins, which would be expected if they consumed milk on a regular basis [4]. Lastly, the authors argue that the age structure of the horse remains at the site is more consistent with mass harvesting of wild horses rather than raising horses under human custody, and they mention a source describing horse remains of signs of being hunted (see below). Thus, the authors conclude that Botai is not the earliest site with evidence of horse domestication but rather shows a culture that was heavily dependent on mass harvesting of wild horses present in the region [4]. Therefore, the hypothesis that the PH is a feral descendant of once domesticated horses loses its basis.

However, the paper by Taylor and Barron-Ortiz did not remain uncontested. In the same year, Outram et al. published a preprint that took their argumentation apart [5]. The Botai tooth does show a unique abrasion pattern consistent with bit wear, namely parallel abrasion at the front of the second premolar of the lower jaw, while similar wear is absent from other teeth in the mouth. This pattern was not found in the wild horses examined by Taylor and Barron-Ortiz, what actually endorses and not refutes the assumption that Botai horses were at least tamed [5].

The lipids from the pottery, so the rebuttal, are more likely to be from milk than from meat or fat because horse milk is produced during summer and therefore has a different deuterium signature compared to meat and fat, which take longer to build up in the body [5]. Furthermore, Outram and others argue that the sample size of two human teeth is not enough to refute milk consumption for the Botai people [5].

What about the signs of hunting on horse remains? The study referred to by Taylor and Barron-Ortiz was by Olsen (2005) and shows rather unambiguously that horses were hunted at the Botai site [6]. Harpoon projectiles were found and many horse remains at Botai show wounds consistent with these on rib bones and skulls, which indicates that these horses were hunted [6]. This, according to Outram et al., does not refute that there were domesticated horses at Botai, only that wild horses were hunted there too [5]. While that may be true, it has massive implications for the Gaunitz hypothesis: if not all horse remains found at Botai were from early domesticated ones but also from genuine wild horses that were hunted, the conclusion that the PH necessarily descendants from these early domestic horses loses its ground, assuming that the horses at Botai were all from the same region and thus genetically similar.

 

New data: it was a sample size problem

 

The last puzzle piece that refutes the Gaunitz hypothesis was a study by Lira Garrado et al. (2025). It actually focused mainly on the phylogenomics of the Iberian wild horse, including 87 genomes of this variant, but also included 261 other ancient horse genomes, which is a rather large sample size [7]. As it turns out, the PH is not nested in a monophyletic Botai-Borly clade, but rather both PH as much as Botai-Borly horses cluster with other old, predomestic lineages from the Ural and Central Asia; with these other old lineages, they form a much broader Central Asian/Uralic horse cluster [7]. This means that Botai/Borly horses and the PH do not form an exclusive monophyletic clade together, but rather descend from a common, large Central Asian wild horse population. There is therefore no evidence that the PH was ever domesticated; rather, it is the last remnant population of this large Central Asian wild horse population.

 

 

Literature

[1] Gaunitz, C., Fages, A., Hanghøj, K. E., Albrechtsen, A., Khan, N., Schubert, M., et al. (2018). Ancient genomes revisit the ancestry of domestic and Przewalski’s horses. Science, 360(6384), 111–114.

[2] Librado, P., Khan, N., Fages, A., Kusliy, M. A., Suchan, T., Tonasso-Calvière, L., et al. (2021). The origins and spread of domestic horses from the Western Eurasian steppes. Nature, 598, 634–640.

[3] Moyers, B. T., Morrell, P. L., & McKay, J. K. (2018). Genetic costs of domestication and improvement. Journal of Heredity, 109(2), 103–116.

[4] Taylor, W. T. T., & Barrón-Ortiz, C. I. (2021). Rethinking the evidence for early horse domestication at Botai. Scientific Reports, 11, 7440.

[5] Outram, A. K., Bendrey, R., Evershed, R. P., Orlando, L., & Zaibert, V. F. (2021). Rebuttal of Taylor and Barrón-Ortiz 2021 Rethinking the evidence for early horse domestication at Botai. Zenodo.

[6] Olsen, S. L. (2006). Early horse domestication: weighing the evidence. In S. L. Olsen, S. Grant, A. M. Choyke, & L. Bartosiewicz (Eds.), Horses and Humans: The Evolution of Human–Equine Relationships. BAR International Series 1560 (pp. 81–113). Oxford: BAR Publishing.

[7] Lira Garrido, J., Tressières, G., Chauvey, L., Schiavinato, S., Calvière-Tonasso, L., Seguin-Orlando, A., et al. (2025). The genomic history of Iberian horses since the last Ice Age. Nature Communications, 16, 7098.

 

Visiting Hortobagy in 2026

The weekend after my visit to the Lippeaue and the Auerrindprojekt, I visited the Hortobagyi National Park, where the largest continuous population of aurochs-like cattle in the world are grazing together with the largest population of Przewalski’s horses in Europe.

 

I met with Istvan Sandor and Peter Czoban to visit the herds. It was a very enjoyable and highly interesting trip, a great nature area with a diverse fauna and a megafauna experience quite like in the Pleistocene. Many thanks to all the people involved for making this possible!

 

The Hortobagyi National Park is located in the Puszta, which is basically an exclave of the Eurasian steppe, but it also has rivers and swamps flowing through it, which is what enabled the presence of aurochs several thousand years ago. Bos taurus (aurochs + cattle) are not a steppe species, so they would have stuck to the wet areas flowing through the Puszta back in their days. Horses on the other hand, the PH in particular, have their prime habitat in the hot and dry grasslands of the Puszta.

The cattle are kept in several paddocks. There is a herd for demonstration for the visitors, a bull herd, a cow paddock, a heifer herd and a paddock for the most precious bulls. I go over them one by one. The bulls chosen to be breeding bulls are allowed to join the cow herds during august and September, which is when aurochs probably mated so that the calves get born during spring. Without this intervention, calves would be born all year round, which could lead to handling problems. There are several steers in all of the herds. Steers have a calming effect on the herds, and they also make an impressive sight for the visitors because of their larger size.

 

The demonstration herd

 

The most eye-catching individual in this herd is a steer that is one quarter Watussi. The cattle are accompanied by Przewalski colts that are noticeably lighter in morphology than functional wild stallions. 

The steer

The steer 

Przewalski colt

The three precious bulls + one steer

 

Three bulls, sons of the Lippeaue bull Felipe, and one steer are kept separate from the other bulls because they are too precious to accept the risk of them getting injured or killed by other bulls on the pasture (this happened to the great bull Köpcös). Felipe was a Taurus bull, a son of Lamarck and a Sayaguesa cow, hence the bulls here are grandsons of Lamarck.

 

Steer on the right of the foto

1323, Fülöpke, is a grandson of Lamarck on the patrilinear side and a grandson of Rimu on the matrilinear side. Thus, he is roughly 6% Watussi, at least genealogically.

1444, Florian

1323, Fülöpke

 

There were three bulls in Hortobagy that left a significant influence on the herds: Anno, a Heck bull from the Steinberg/Wörth lineage; Felipe; Rimu, a (Sayaguesa x Heck) x (Hungarian Grey x Watussi). A pure Watussi bull was also briefly used, as much as the Taurus bull Lasso (Heck x (Heck x Sayaguesa)), a son of Lucio from the ABU. Other bulls were used occasionally. They also imported the Heck x Lidia bull Larus from the Lippeaue, but he was not used for breeding because of excessive aggression. Istvan told me while the other bulls might attack because they want to be left alone, but Larus really tried to kill. It was not possible to handle this bull, but his skull now makes an impressive sight in the visitor center.

 

Larus' skull

Lasso (bottom), Toldi? (top)

There is also the skull of Lasso, which is significantly shorter, being three quarters Heck.

 

In the visitor center there is also what I have been looking for for years: a complete, mounted skeleton of a Taurus bull (or steer). It is rather small, which is why Istvan does not like it, but I think it looks great apart from the body size: the legs are really long (note that the forelimb and shoulder blade are not positioned as in life), the trunk short. The skull is comparably short, but not much shorter than in the Cambridge or Torsac-dirac aurochs. The tips of the spinal processes in the shoulder area are unfortunately broken off.

Regarding the size of the bulls in the corral, I was able to measure one of them very indirectly. When it was standing next to the fence with not much hay beneath it, its withers were reaching half the distance between the two uppermost steps of the fence – which would be 155 cm. That’s 20 cm shorter than Dominator, but not a problem, since Hungarian aurochs were not much taller.

 

The main bull herd

 

I was amazed by the bull herd. There are many rather long-legged and short-trunked individuals that have the “nearly squarely built” morphology that aurochs-like cattle are supposed to have. Most of them are black, bulls with a saddle get selected out. One bull, named Turkana, sticks out. He is Felipe x a half-Watussi cow, therefore quarter Watussi and will be a future breeding bull. I think it is the strong Watussi influence that resulted in the desired short trunk of the bulls because indicine cattle usually have a shorter trunk than taurine cattle. Istvan likes the Watussi influence very much for their elegant morphology and also their extreme heat tolerance. The summers get pretty hot in the Puszta and due to recent warming, the Heck and Hungarian Grey influence, which was important to cope with cold winters in the beginning in the 1990s, becomes less important than the heat-tolerance of Watussi in recent years. So far, the zebu hump of Watussi did not reappear in later generations, also not in crosses that have Watussi heritage on both parental sides. In first-cross generations, the zebu hump is almost gone, and when crossed with another animal that does not have the zebu hump, it is phenotypically completely gone. 

Turkana

Turkana

Turkana

 

 

1543

1542

1542

1533

1533, 1543 and 1542 are sons of Köpcös.

 

Look at those long-legged, short-trunked bulls!

Steer

 

Istvan demonstrated a funny little trick to get the bulls stopping and looking at us – grunting like a dominant bull. This makes the bulls freeze and look what is going on. 

 

Me with an aurochs horn fossil at the visitor center

 

The cow herds

 

The cows live on a very large area and interestingly, they themselves split up into two herds, in sum counting around 150 animals. That is still much larger herds than what is usual for Bos taurus, but I was told that is because of the lack of bulls in the herd. With the presence of bulls, they would split up into subgroups.

 

There once also was a Grey x Holstein cow in the herd named Xena. She had a phenotypically wildtype colouration, except for ventral white spots. Holstein, however, turned out to be not that beneficial for the breeding, Istvan told me. One of her sons, Zseuz, looked good – but only on the photo, Istvan said.

 

The cow with the horns I like the most is Damalisz, her horns are simply fantastic and resemble those of my reconstruction of the Gramsbergen skull fragment very closely. I hope she passes on this horn shape to her offspring.

 

 

Delphi, quarter Watussi

Damalisz

Reconstruction of the Gramsbergen skull fragment

Damalisz

Damalisz

Damalisz

Cintia

Liszi

Szepasszony (daughter of Felipe)

Pure Heck from Hellabrunn Zoo

Half-Watussi 1

Half-Watussi 2

Half-Watussi 3

Half-Watussi 3

Young steer

Damalisz

The wild horses

 

As mentioned, Hortobagy has the largest PH herd in Europe, and possibly also the most diverse European herd – not only genetically, but also phenotypically, with some individuals being very light in colour and one stallion having a much darker colour that got passed on to both of its offspring. I speculated that this stallion might be dun/nondun1 genotypically because it still has the white muzzle, but I am guessing here. It is also possible that it has a nonpangare/pangare genotype – the nonpangare allele was pretty common in the wild catches of PH from the 19^th century, especially those at Askania Nova.  

 

The slightly darker mare

Quite similar to P. Pallas' 1771 tarpan 

Contrary to cattle, they form one very large herd when there are a lot of horses in the given area. But something interesting happens with the social structure in these super-herds: there seems to be an either intentional or unintentional stallion coalition. The dominant stallions that already have a harem group stay together in the centre of the herd, reducing the borders that are exposed to and must be defended from the bachelor groups, which are at the outside of the herd.

 

Standing surrounded by 250 wild horses in the steppe was an amazing experience – I felt like I did a time travel into the Pleistocene.

 

Water buffalo

 

We also had a look at the Water buffaloes in the reserve. They are put on the swampy areas with lots of reef. The ecological impact of water buffaloes is extremely beneficial for the wet ecosystems, since their foraging and wallowing behaviour creates microhabitats for many species. Habitat disturbance often enables more diversity than the absence of disturbance and Bubalus is a native genus in Europe. However, the native European water buffalo species, B. murrensis, is lost and the Carpathian mountain buffalo is a descendant of the Asiatic wild water buffalo. They do look like they are perfectly adapted to European winters with their long winter coat, but Istvan told me that they lack the underwool – during snowy winters, they are freezing and shaking, spend most of the time in the barn and would probably not survive without human intervention.

 

We also talked about Hortobagy as a potential bison habitat. Istvan told me that wisent would probably not do all too well in the reserve because it is a lowland habitat with salty ground, but American plains bison probably would – they are kept in America in habitats that are essentially ecologically identical to Hortobagy.

 

Conclusion

 

How do the Hortobagy Taurus cattle compare to the Lippeaue ones? The latter seem to be larger on average, although the height data for the Hungarian Taurus is very sparse. On the other hand, they have much larger horns on average compared to the Lippeaue ones and some of the bulls have a very good morphology (long legs + short trunk). So the Lippeaue cattle are, due to Chianina, larger but have smaller horns compared to the Hortobagy ones, which are, due to Watussi and no Chianina, larger-horned but not quite as large. So it really depends on which traits you prioritize and I think both populations are wonderful.