How aggressive was the aurochs?

As my readers will know, many of my blog posts on the aurochs focus on aspects of its morphology or appearance, because that is what we know most about. But I also made posts covering social behaviour patterns, like this one. Today I want to cover a behavioural aspect of the aurochs that I am sometimes asked about: its aggressiveness. More precisely, its aggressiveness towards humans. Was the aurochs an aggressive animal towards humans, or was it gentle and docile? 

Regarding the aggression of the animal, we have to differentiate between intraspecific aggression, aggression towards other animals (predators in particular), and aggression towards humans. That is important because domestication drastically changed the latter factor, while the other two factors seem to be somewhat independent from it. Cattle that are very agreeable and docile towards humans still can become rather aggressive against opponents in intraspecific combat or towards possible predators. The question how aggressive the aurochs was is also important for “breeding-back” as it raises the question what kind of behaviour the cattle need in order to survive in nature, or to be aurochs-like, and what to expect from dedomesticated cattle living in the wild. 

 

Historic evidence on the behaviour of the aurochs towards people

 

The most famous notion on the behaviour of the aurochs is in Caesar’s Commentarii de bello gallico, writing that aurochs “will spare neither humans nor animals at sight”. Caesar also wrote that aurochs never get used to humans, even if raised as calves. Also, Anton Schneeberger who visited the last Polish aurochs in Jaktorow wrote that aurochs will get very hot-tempered when challenged. Andrea Swiecicki reported from the 16^th century that aurochs will get very aggressive when being tormented and attack humans and horses, throwing them in the air with their horns [1]. Schneeberger further writes that aurochs are not afraid of people and will not run away if they stand in the way of the humans. As the Jaktorow herd was managed by humans and encountered people on a regular basis, it is possible that they were tamed to some degree [1]. These are the only general notions on the behaviour of aurochs towards people that have been preserved, at least as far as I know.

Caesar’s notion can be interpreted in a way that aurochs were generally aggressive animals, but Schneeberger’s and Swiecicki’s seem to be more differentiated, stating that they can become very aggressive when challenged or being hunted. Caesar’s impression that the aurochs will not spare any living being might come from the behaviour of the bovine in situations of threat. If one knows the animal only in the context of hunting it, one might assume it is generally a very dangerous animal. Schneeberger’s report clearly states that aurochs would not attack without a reason whenever they encounter a human, in fact he writes that one could easily approach it. If the aurochs at Jaktorow were indeed tamed to some degree, this could imply that the behaviour of aurochs towards humans was plastic and partly depended on socialization. However, Caesar’s notion that they never get used to humans even when raised as calves indicates that there were genetic limits to this. This is likely, as the difference in behaviour between domestic and wild animals is due to genetic differences to a certain degree. Curiously, Vergilius mentions that in the Po River area in Northern Italy aurochs were caught and tamed to be used as draft animals, in the lack of domestic oxen [1]. Either it was indeed possible to use grown, wild-caught aurochs for draft work or Vergilius was misinformed. It is even possible that these animals were not aurochs but feral cattle living in the region. Personally, I cannot imagine that it would be possible to use an undomesticated wild bovine for draft work, even if they were tamed to some degree. I think Caesar’s notion is more credible than that of Vergilius, both authors were basing themselves on second-hand information. 

 

The behaviour of domestic cattle 

 

For the social behaviour of the aurochs, cattle living under natural circumstances are a pretty good model. What we know of the social behaviour of the aurochs is congruent with that of cattle, and since domestication likely did not affect the social behaviour patterns of cattle, all the other aspects of their social behaviour are likely shared between aurochs and domestic cattle. One difference is, however, that cattle tend to mate all year round while the reproductive circle of the aurochs was adapted to that of the seasons, with a mating season during fall and the birth of the calves in spring. 

But that domestic cattle are a model for the aggression level of the aurochs is very unlikely. First of all, domestication likely drastically changed the hormonal system of cattle, particularly the corticosteroid hormones, what has an influence on the aggression and stress response of the animals. Furthermore, there are considerable differences between breeds in their behaviour towards humans. Highly derived breeds in particular are very agreeable, docile and have a very minor stress response. In some breeds the bulls can even be ridden. The Spanish fighting bull, on the other hand, is the complete opposite. Lidia have a very intense stress response and attack readily, although there is individual variation in “fighting spirit” within the breed. Most domestic breeds were artificially selected for agreeableness and less intense stress response, Lidia was artificially selected for an intense stress response, so that it may be the case that neither of them is a model for the behaviour of the aurochs towards humans. 

The potential for aggression, particularly towards humans, must have a genetic component otherwise it would not be heritable. The MAO-A gene seems to be different in Lidia compared to other taurine cattle breeds (go here). This gene plays a role in the endocrinology in the brain of mammals, and mutations on this gene are linked to increased aggression in other species, suggesting that this plays a role in the aggression of Spanish fighting cattle. It would be interesting to have a look at this gene in the aurochs. Since the full genome of the aurochs has been resolved, this should be possible. The MAO-A gene is probably one of many genes that influence the potential for aggression in cattle. 

 

To sum it up, aurochs likely would have had the potential to become very aggressive and energetic when defending itself, like other wild bovines. We do not know, however, how readily wild aurochs would have attacked if they encountered a human being that they consider threatening to them. Some aspects, like flight distance, might have partially depended on socialization. However, as Caesar writes, an individual raised under human custody likely would never lose its wild nature as they were undomesticated animals. 

 

All current “breeding-back” projects select against aggressive behaviour towards humans, which is understandable as they have to work with the cattle as the law dictates. As long as the animals have to be handled, it is better for the cattle and the people handling them that they do not have an extreme stress response. As we do not precisely know how aggressive aurochs were, I think it would be most sensible to let “rewilded” cattle develop their own aggression level that is formed by natural selection and thus what is best for them in order to survive in nature.

 

[1] van Vuure, 2005: Retracing the aurochs: history, morphology and ecology of an extinct wild ox. 

 

 

 

Reconstruction of a complete Indian aurochs skull

The Indian aurochs, Bos primigenius namadicus, is enigmatic compared to the European subspecies. Not a single complete skeleton has been found yet, and I have not seen a complete cranium either. I wanted to get an idea how the complete skull of that subspecies might have looked like, so I tracked out the namadicus skull presented in [1] (which might or might not be the same skull that is on display at the Geological Survey of India) on paper and reconstructed the complete skull using the fragmentary cranium. 

 

The trickiest part was to estimate how long the snout would have been. I suspect that the skull broke off right before where the nasal bone and the premaxillary bone touch each other, and that the toothless part of the upper jaw is roughly the same length in lateral view as the toothed part. The lower jaw is based on those of European skulls. This is the result:

I think the result is very plausible, it does not look proportionally weird concerning the snout length. It also looks credible for a type of aurochs that was the predecessor of indicine cattle. For comparison, here is a zebu skull. 

 

As no postcranial material that is worth mentioning has been published so far, I cannot do this with a complete skeleton, unfortunately. However, I think there must be enough fragmentary postcranial material of that subspecies to make a composite skeleton, so that it is possible to get an idea what the morphology of namadicus was like. A rigorous description of the postcranial skeleton of the Indian aurochs is lacking so far. 

 

[1] Gregoire Metairs: Evolutionary history of the large herbivores of South and Southeast Asia (Indomayalan Realm).2016. 

 

Video of fully grown Lidia bulls

Most Lidia bulls we see are young bulls at the age of three or four, because that is when they have their full body size but are comparably slender and most athletic. After that age, they become heavier, as all bulls do. It is rare that fully grown Lidia bulls are presented on the media as they are not as athletic and swift than young bulls. But here is a video of Lidia bulls that are most likely fully grown:

They are still pretty muscular, as typical for the breed, but heavier than their younger counterparts. What is most interesting to me is that a number of individuals have much more aurochs-like horns than young bulls. At the age of three, the horns are not yet fully developed and can change quite noticeably. Many young Lidia bulls have a somewhat two-dimensional horn curvature, while some of the bulls in this video have a nice primigenius spiral, in particular the bull at 5:07 and the one at 9:34. I think that supports the idea that the horns of Lidia are more often aurochs-like than what the young bulls seem to suggest, if they only get the opportunity to grow to full adulthood their horns will be more developed and that can result in a primigenius spiral. 

A plea for more Maronesa in "breeding-back"

Maronesa is an awesome breed that I always enjoy looking at. Like all aurochs-like breeds, they have their pros and con’s. To sum them up: 

Con’s: 

- small or at least not large body size 

- short skull shape 

- bulls can get rather short-legged and heavy 

Pros: 

- the coat colour is absolutely identical to that of the European aurochs 

- the sexual dichromatism is nearly always present and well-marked, identical to that of the European aurochs 

- the horns can face inwards in a very aurochs-like manner 

 

Although the number of pros and con’s that come to my mind is the same, I think the benefits of that breed outweigh the undesired traits. That is, for once, because it is very, very rare that a cattle breed has a colour that is truly identical to that of the European aurochs – actually the only other European breed that I know of is the old lineage of Corsican cattle, which is critically endangered if not has already disappeared. It’s impressing to imagine that during roughly 10.000 years of domestication, not a single domestic colour mutation has found its way into the Maronesa genome. But even more important is the fact that the sexual dichromatism is nearly always present and as well-marked as in the European aurochs, because sexual dichromatism is a complex trait that is very difficult to breed for in domestic cattle. I did a post on that, coming to the conclusion that the only way to achieve an authentic dichromatism in “breeding-back” is to rely on a breed that already has it to the desired extent. That would be Maronesa. At least I know of no other breed that has an authentic dichromatism except for the old lineage of Corsican cattle. Also, the horn curvature of some Maronesa is very useful as the horns curve very strongly inwards in an aurochs-like manner. This is very rare even among aurochs-like cattle. Just look at this cow. Overall it has great potential for aurochs-like offspring. The colour is perfectly identical to that of the aurochs, the horns curve inwards in an aurochs-like manner and it has a perfect dichromatism (I assume so because it is the rule in the breed and there are no lightly coloured Maronesa bulls). 

 

Considering the potential of the breed, I think it is dramatically underused in “breeding-back”. The only current project that is using Maronesa is the TaurOs Programme. That is problematic because they use only a small number of Maronesa individuals, they do not execute selective breeding but let the cattle breed for themselves instead, they crossbred them with breeds that are not really beneficial from the perspective of aurochs-likeness (f.e. Maremmana) and the results are modest. And that is although the project would badly need good Maronesa to improve the horn shape of their cattle, as the horns of most TaurOs cattle face outwards, and the sexual dichromatism which is completely absent in some herds (f.e. Milovice). I would highly recommend the Tauros Programme to try to achieve another herd of Maronesa from Portugal, this time better individuals (that have truly inwards-curving horns and a good morphology etc.), also including grown bulls, as bulls have a greater influence on the herds than single cows. 

Also, Maronesa could be beneficial for the Auerrind project in the future. That would depend on how good the dichromatism is going to get with the set of breeds currently used – Maremmana, Sayaguesa, Watussi and Grey cattle have a rather reduced dichromatism, Pajuna can be good in this respect, and in Chianina a dichromatism is possible but masked beneath their colour dilution if present. If it turns out in the future that Auerrind crosses do not have a well-marked dichromatism and inwards-facing horns, including good Maronesa could be beneficial. But as for now it is too early to judge that, as the second-generation crossbreeds are not fully grown yet. 

Maronesa would even be an option for Heck cattle breeders to increase the aurochs-likeness of their cattle. Some Heck cattle breeders don’t like large individuals, or simply like the looks of Heck cattle regardless of their aurochs-likeness. Maronesa would improve the horn shape and sexual dichromatism without altering the looks and body size of Heck cattle dramatically. But Heck cattle would not be my first choice to crossbreed good Maronesa with, to be honest. 

Would TaurUs cattle benefit from Maronesa? I am not sure about that. Many Taurus cattle in the Lippeaue already have inwards-facing horns, and the sexual dichromatism is good in the herds as well – I examined it for the year 2015 using a photo archive, and it turned out that more than 80% of the individuals have the “right” colour. Go here for the post. Apart from that, they have done a good job at creating truly large cattle, Maronesa might diminish that. As most of the achievable aurochs-like traits are already present in the Lippeaue, I do not think that Maronesa would be necessary or, considering their con’s, worth the effort in Taurus cattle. 

 

Another option to seize the potential of Maronesa would be conducting a new, heavily Maronesa-based breeding project. I would use Maronesa, Chianina and Watussi for such a project. Precisely, I would create a number of F2 Maronesa x Chianina, and F2 (Maronesa x Watussi) x Maronesa, and then create an F2 from the combination of both lineages. That would take five breeding generations, and thus roughly ten to 15 years, but the results could be quite qualitative. 

 

I think it would be a shame if Maronesa was not used on a larger scale in “breeding-back”. That is also because it seems that the aurochs-like less-derived Maronesa lineages are endangered. From what I have heard, the sexual dichromatism is not always appreciated by Maronesa breeders and there are already some almost black Maronesa cows. If that trend continues the breed might lose its dichromatism and end up looking like Sayaguesa in that respect. Also, some Maronesa are bred for an extremely massive body with a bulldog face, and many Maronesa cows have corkscrew-like horns (I do not know what is the preference of Maronesa breeders regarding horn shape). The less-derived type of all aurochs-like landraces is endangered because of crossbreeding with more economically productive breeds and/or selection towards a more derived appearance, and so is that of Maronesa, at least because of the latter factor. Thus I really hope that “breeding-back” will seize the potential of that breed, and that the less-derived aurochs-like representatives of Maronesa do not disappear without contributing noticeably to the “breeding-back” gene pool. 

 

The colour of the African aurochs

I did a post on the African aurochs a few years ago. In that post, I outline that it is likely that the bulls of this subspecies had a light colour saddle on their back, instead of being entirely black except for the dorsal stripe as the European aurochs evidently was. This is because at least three depictions of African aurochs bulls show this trait very clearly. This might not be the full story, however. 

 

Part of the reason why I am saying this is the ancient Egyptian aurochs depiction described in Beierkuhnlein 2015: 

from [1]

It shows an aurochs bull, a cow and a calf. It is somewhat stylized (see the horns), but rather detailed. It shows the white muzzle in the cow (the mouth of the bull and calf are not preserved), it shows either a rather broad dorsal stripe in the bull or a colour saddle (that is probably open to interpretation), a narrow dorsal stripe in the cow and a possible dorsal stripe in the calf similar to what is depicted in the bull. The colour of the bull seems to be a little bit darker than that of the cow, but still is brown. However, there is also a depiction of a dark brown aurochs bull from Europe that is also very stylized, and as I mentioned, the depiction is a bit stylized. So it’s still possible that the bull this artwork was based on was black. But what is most interesting to me is that the bull has light areas on the dewlap, belly and inner sides of the legs. Also the cow and the calf have a lightly coloured belly. 

When I first noticed that I did not really care about that because I thought the artwork is stylized anyway, so it could be artistic license. However, these light areas are a trait often seen in many zebus. In fact, I have not yet seen a male wildtype-coloured zebu without taurine influence that does not at least have lightly coloured “armpits”, often the light area extends to the dewlap and entire belly and the inner sides of the limbs. This Sahiwal bull shows this kind of colour: 

If these light areas were present in the Indian aurochs, which was the wildtype of the zebu, it might not be that far-fetched to assume their presence also in the African aurochs. This would be in agreement with this ancient artistic depiction. Regarding the width of the dorsal stripe, I think it is possible that the African aurochs had a broad dorsal stripe in the shoulder area while the European aurochs had a narrow one (about “two fingers wide” as Schneeberger reported). I have seen dorsal stripes that get rather broad in the shoulder area in one Heck bull, one Pajuna bull and one Maronesa bull. African aurochs likely influenced African taurine cattle, which left a mark in some Southern European breeds, so it is not impossible that a broad dorsal stripe is a legacy of African aurochs introgression. This is speculative, especially as it is not clear if the artist of the depiction intended to draw a broad dorsal stripe or a saddle. I am inclined to think the artist did not differentiate between both structures. 

I also found a depiction of a bull with a very similar or possibly the same colour. It’s from a mural in the tomb of Nefertari. It is clearly black, has light areas on the ventral side of the trunk and the inner area of the legs, and also has this either broad dorsal stripe or colour saddle (go here). I am pretty sure it shows a domestic bull because it is shown along other clearly domestic bovines based on their colour (go here). But it could be a hint that this colour was found in the local aurochs type, as there likely was introgression from mauretanicus into North African taurine cattle. It could be a hint, but it certainly does not have to be. 

If the African aurochs had this lightly coloured area on the ventral side of the body plus a broad dorsal stripe, it might have looked like this: 

Personally, I find this colour scheme incredibly beautiful and also rather colourful and rich in contrast. But it is just as plausible that North African aurochs did not have these lightly coloured areas and were all black except for the saddle, dorsal stripe and muzzle ring. The evidence simply is not clear or plentiful enough to tell the exact colour of Bos primigenius mauretanicus. 

 

[1] Beierkuhnlein: Bos primigenius in ancient Egyptian art – historic evidence for the continuity of occurrence and ecology of an extinct key species. 2015. 

 

 

 

Horn size #2: aurochs with smaller horns

In the previous post, I presented a number of aurochs specimen with very large horns. This post is going to focus on the lower end of the horn size spectrum. Although the aurochs was a large-horned bovine in general, some individuals had surprisingly small horns – these small-horned aurochs are, however, not very numerous in the fossil and subfossil record, indicating that they were not common in the species either. 

 

-) The Prejlerup bull 

This skeleton is possibly the largest more or less complete aurochs skeleton that is on display, with a withers height of roughly 190 cm. Its horns, however, are not very large. They seem to be shorter than the length of the skull, what makes them “smaller” to me by aurochs standards. Surely the keratinous sheath would have added to the length, but the curvature shows that this would not have doubled the length of the horns, otherwise they would have crossed each other as the horns are not wide-ranging. 

 

-) The Önnarp skeleton 

This bull specimen has horns that are smaller than average for the aurochs. I know no measurements for the horn cores of that bull, unfortunately. Go here for the skeleton. 

 

-) The Himmelev bull 

Although the horns of the Himmelev bull are not tiny, they are quite a bit smaller than what is average for the aurochs. No measurments for the horn cores that I found. See here and here. 

 

-) The skull from Bauges 

In a cave in Bauges, France, a skull was found that has truly tiny horns by aurochs standards. No measurements for the horn cores of this skull are given, but there can be no question that they are significantly shorter than the craniocaudal length of the skull. Those are the shortest horn cores I have ever seen in an aurochs skull. The skull can be seen in this video. 

 

Still shot from the video linked above

-) The last Jaktorow bull 

We know exactly how long the horns of the last bull from the Jaktorow forest in Poland were because the sheath and not the core has been preserved. It turns out that it is merely 46 cm long, which is very small compared to horns of other, earlier specimen. Those were up to three times as long than this sheath, speaking of only the cores. The Jaktorow horn does not look like it is not yet fully developed, so it is probably from a grown bull. Overall, the horn is rather meagre, also compared to earlier sheaths which are thicker and more strongly curved. 

Those were the smallest-horned aurochs specimen preserved that I know of. They all have something in common: they are exclusively from the European subspecies, and they are all from the Holocene. And that might explain why aurochs with smaller horns existed: the presence of humans. I explain in the post linked in the post on large-horned aurochs why I think that anthropogenic influence is the reason for small-horned aurochs to appear. First of all, the aurochs suffered from a fragmented and confined habitat as civilization competed with the aurochs, there likely was hybridization with domestic cattle in the wild in Europe, and the aurochs was hunted for its horns. Trophy hunting is known to affect the morphology of animals. For example, large-tusked elephants have become rare in both Asia and Africa and tuskless elephants became more frequent. Also, the cape buffalo used to have much larger horns in the past than it does today, go here or here and compare with what is average for this species today. I think those factors are the most credible explanations for why smaller-horned aurochs appeared in Holocene Europe, at least I am unable to come up with any other explanation that is plausible for why the European aurochs decreased in horn and body size during the Holocene, as I write in the post linked at the beginning of this post.