Were European wild horses dun-coloured or not?

The European wild horse probably was of a similar build like the Przewalski’s horse, that means a 130-140 cm tall stocky body with a large and robust head. Numerous eyewitness accounts confirm that, as well as that it had a short but falling mane. But what about the exact colour? Genetic information tells us about the two basic colours that were present in European wild horse populations, but the phenotype of these horses is uncertain as long as we don’t know if the basic colour was diluted by dun or not.

To make more clear what I am about, let’s have a deeper look into the coat colour genetics of horses. Horses have two alleles on their Extension locus, which defines the basic colour: E, the wild type allele, resulting in the production both red and black pigmentation, and the domestic mutation e, which is recessive and produces totally red colours such as chestnut or sorrel. The amount of black pigmentation in wildtype coloured horses is controlled by the Agouti locus. The allele A restricts black to the legs, mane and tail and results in bay horses; the allele a on the other hand produces totally black horses. We know that both A and a are wild type alleles, they were present in undomesticated European horse populations [1]. There is another variant called A^t, producing colours like seal brown. This allele is not mentioned by Pruvost et al., but it might indeed be another wildtype allele; according to this website, it might even be present in donkeys as well (if that’s the case, it’s a very old allele). Countershading, also called pangare, which produces the mealy mouth and light belly in bay and bay dun (but not black and black dun) horses, serves the so-called somatolysis and is present in a variety of animals of all sizes and habitat preferences, so it doesn't say much on whether we look at a forest-dwelling animal or open land animal. 

Dun is the only dilution locus which is wildtype. Dun reduces the intensity of pigmentation; bay and seal brown horses with dun are sand-coloured (bay dun) while black horses with dun are grayish or tan-coloured (black dun), also called grullo. The so-called wild markings including the dark dorsal stripe and the leg striping are certain indication of the presence of dun since the former trait is barely visible and the latter invisible without dun. All living equines are dun-coloured (bay dun in particular), so it sounds logic that European wild horses were too. On the other hand, all living wild equines live in open habitats, while Europe likely had more closed, forest-based natural landscapes. Therefore the loss of dun resulting in a darker colour (a more efficient camouflage in forests) seems conclusive as well. And indeed bay/brown and black ponies like the Exmoor, Asturcon or Fell pony seem to fit closed or bushy environment much better than do bay dun and black dun Przewalskis or Koniks. Since there are no preserved skins of undomesticated native European horses, we have to ascertain that question using two sources of information: genetic information and historic accounts. Cave paintings could provide a clue as well, but they are not very reliable for two reasons: 1) they stem from the age of the last glacial, and therefore the horses depicted were most likely adapted to more steppe-like habitat than that of modern Europe, and 2) probably not all colours on the cave walls are an accurate representation of what was found within the population because the cave artists had to work with what was available.

Black dun Konik: perfectly camouflaged in open, grassy landscapes

Dark bay Exmoor: perfectly camouflaged in more closed environments

Genetic data is by far most reliable because it is unambiguous. Pruvost et al. 2011 tested predomestic wild horse remains from Europe of both the Holocene and Pleistocene for a number of colour alleles. It showed that the basic colours all were bay, black and leopard spotted (the latter is not relevant for the topic of this post, so let’s ignore it) [1]. No dilution allele was detected within the wild horse populations, but the dun mutation was not yet resolved, therefore it was impossible to distinguish between dun and non-dun horses [1].

Historic references describe a variety of colours. They are less clear because there are numerous words to describe one and the same colour, translation errors are another factor. Eyewitness accounts often are ambiguous because the colour terms we use colloquially are not congruent with the terminology of coat colour genetics: mouse-coloured might refer to shades of bay, bay dun or black dun, but in coat colour genetics it is used only for black dun. That means we have to be cautious when interpreting historic accounts.

Another problem is that we don’t know for sure the status of the free-roaming horses described as “wild”. As I wrote in previous posts, there are good reasons to believe that many of the texts actually describe wild horses, but domestic introgression can never be ruled out. Therefore we should stick with what is likely to be a variation of those colour genetically proven to be predomestic, and consider other colours to be the result of intermixture.

So let’s have a look on what historic references actually say.

Herodot describes lightly coloured wild horses on the area of the Ukraine in the 5^th century before Christ. “light” is quite vague, but it most likely means that the horses were dun-coloured. Albertus Magnus writes of mouse-coloured horses with a dark eel stripe in Germany of the 13^th century. Balthazar Hacquet saw wild horses at the Zoo of Zamosc, Poland, in the 18^th century and reported that they were of a blackish-brown colour. Kajetan Kozmian saw the wild horses of the same park and same century, and described them being uniformly of a dark mouse-colour. The reports of S. G. Gmelin and Peter Pallas refer to the wild horses of the Russian steppe (which also showed clear signs of intermixture), so they are of little value for the European forest horses [2].

The naturalist C. H. Smith described free-roaming “sturdy mountain-forest ponies” in his 19^th century work The Natural history of horses, with memoir of Gesner (see here for his full description of western wild horses). These wild ponies were found in western Europe and according to him had “a livery in general pale dun, yellowish brown and a streak along the spine and cross bars on the limbs, or the limbs entirely black, as well as all the long hair and mostly having a tendency to ashy and gray, often dappled on the quarter and shoulders” [3]. Genuine wild horses extant in western Europe during the 19^th century sound a bit far-fetched, but all the behavioural and phenotypic features Smith describes for these horses imply that they were wild or at least primitive feral ponies. The colour he describes certainly refers to bay dun (“yellowish brown and streak along the spine”) and black dun (“tendency to ashy and gray”), and also Herodot’s text supports that European wild horses were dun-coloured.

Dark brown might either refer to dark expressions of black dun or to the seal brown (bay) colour of ponies like the Exmoor. We probably will never know which colour that actually was, but the mealy mouth of bay horses could be a hint – this white area around the mouth is pretty apparent on an otherwise brownish coat, therefore it might have been mentioned if it was there. No historic reference mentions it, what might support the view that these horses had a brownish expression of black dun; but this is just a pure speculation of mine. The term “mouse coloured” also leaves big room for speculation. It is probably hard to figure out which colour they actually intended to describe with that term. European mice species display various shades of brown and gray, and no-one can know if the writer actually had any of these particular species in mind when speaking of mouse-coloured horses or just used it in the colloquial sense, and this colloquial sense might also be influenced by time and region. So we really have to guess.

All in all, it seems that historic references suggest European wild horses were dun-coloured, so both the colour of the Przewalski’s horse and brownish Koniks appeared. Intuitionally I consider it quite likely that there were non-dun wild horses in Europe as well because they are perfectly camouflaged in forested and bushy landscapes. But to be sure we have to wait until the mutations on the dun locus are identified so that dun and non-dun wild horses can be genetically distinguished. It would also be very interesting to know if the A^t allele was present in predomestic horse populations as well. 

Literature 

[1] Pruvost et al.: Genotypes of predomestic horses match phenotypes painted in paleolithic works of cave art. 2011

[2] Tadeusz Jezierski, Zbigniew Jaworski: Das Polnische Konik. 2008.

[3] Smith, Charles Hamilton: The Natural history of Horses, with Memoir of Gesner.(1814/1866)

Book review: "The Aurochs - Born to be wild"

In October this year, the third recent book on the aurochs was presented at the World Wildlife Congress in Salamanca, Spain. It was written by Ronald Goderie, initiator of the Tauros Programme, Henri Kerkdijk-Otten, former manager of the project, Wouter Helmer, managing director of ARK Nature and Staffan Widstrand, manager of Wild Wonders of Europe and published by Roodbont Publishers B.V.

The book is written more in mass-market style (and I guess also to attract possible sponsors) than technical, what makes it easy to read for starters. The book gives you general information on the biology of the aurochs and tells everything important about its appearance: colour, sexual dimorphism, body shape and proportions, horns, skull. It makes clear that the Aurochs is similar to domestic cattle in looks, but also showed some clear differences.  

The book emphasizes the impact on the European culture that aurochs and cattle had, especially during the antiquary. Aurochs (or actually unspecific bulls) were in the focus of numerous legends and cults, and aurochs also were used in roman arena fights.

The ecologic context of the species is portrayed as well. It gives you an introduction into what species it shared its habitat with, during the Pleistocene as much as the Holocene. It shows that Europe was inhabited by a number of large grazers which are now either extinct or confined to single nature reserves – the book claims that the masses of herbivores in Europe once were comparable with hat we find in the African Serengeti today, although this picture is questioned in the literature. In fact, the book portrays the very controversial megaherbivore hypothesis* and overkill hypothesis as consensual mainstream opinions, but in fact there is still much debate about these.

* The megaherbivore hypothesis suggests that megaherbivores play a crucial role in shaping natural landscapes and create open land where otherwise forests would grow.

The book has a lot of nice pictures, also including two artworks of mine and of Tom Hammond (I am going to cover his aurochs art here on this blog as well). I am also very fond of the wonderful illustrations by Jeroen Helmer, which are very inspiring for me. I love the painting down below because it shows probably all aspects of the aurochs’ daily behaviour repertoire.

Aurochs and their natural behaviour, © Jeroen Helmer

“The Aurochs – Born to be wild” also introduces the Iberian primitive breeds. I think it is very good that it does not spend too much attention on Heck cattle but instead focuses on less-derived or feral breeds as those living animals closest to the aurochs.

It gives a good overview on what Tauros Project is doing; still not the “material and methods” presentation I’d like to see, but certainly useful. It also reveals a bit of the results of all the genetic research that is done under the frame of the Tauros Programme that will hopefully bring us a better view on primitive cattle phylogeny, how much of the aurochs is left and where the local hybridization events with wild aurochs happened. I actually hoped to find some recent photos of Tauros crossbreeds, but the book only shows Manolo Uno. However, I was very happy to find the photo of one of his sons at the end of the book. It’s Maremmana x (Pajuna x Maremmana). It’s clearly visible that this bull calf is going to be grayish as an adult (there was a 1:1 probability that it inherits the dilution allele from its father so this F2 is probably homozygous for this feature), but maybe it will grow larger horns than its father. There are about 150 Tauros crossbreeds now, unfortunately only few of them were presented to the public yet.

All in all, it’s definitely worth buying this book if you are interested in the aurochs, European big game or rewilding. You can purchase it here. 

Coming soon: Taurus bull skulls!

I am the proud owner of two beautifully aurochs-like skull bones of nice Taurus bulls since yesterday. I'll post a lot of photos from all possible angles and measurements here.
I also know photos of the two living bulls in flesh. This enables me to do direct comparisons by overlaying photos, and consequently, after evaluating the differences in proportions between the Taurus skull and aurochs skulls, to transfer these proportional differences on the photo of the living bull and therefore doing a lively reconstruction what living aurochs skulls might have looked like, based directly on osteological information. This will be probably the most precise skull reconstructions I did so far. 
A little book review on the new aurochs book "The Aurochs: Born to be wild" is also about to come. 
Stay tuned!

Some slender cows

I have already mentioned the Taurus cattle herd at Schmidtenhöhe, a little nature reserve in Rheinland-Pfalz, Germany. The herd includes some usual, "un-improved" Heck cows that I would not use in any breeding-back attempt, some good and slender Taurus cows, a Sayaguesa bull named "Napoleon" as breeding bull, plus their offspring. The video down below, which I already posted here some time ago, shows the herd 4 years ago.

I post it here because I am really struck by the beauty of some of the cows in that video - long-legged, very slender, long snout, shiny reddish to black fur. This is what I imagine wild cattle to look like. If they'd have the horns of good Wörth Heck cattle and a larger shoulder area, they'd almost look perfect. I took some screen shots, all credits of course go to Elke Führer.

Note the dark brown cow at your right...

...and the cow with the saddle at your left

I am also very fond of the Pajuna in this video (below), they have a body just as slender and long-legged, and a more reddish coat. Their skull is, as usual for this breed, longish and aurochs-like as well, better than in the cows above. Unfortunately most Pajuna are rather small, but crossing them with Maremmana or Podolica, like Tauros Project does, will resolve that.

(Again, I would insert the video here, but Blogger does not find it)

https://www.youtube.com/watch?v=KCM-3kVLTCA

The horns of the Aurochs

The aurochs is often remarked for its impressive horns, which were larger than those of most domestic cattle. Contrary to some other claims, the horns of the aurochs were – just like those of other wild cattle – more or less uniform so there are clear objectives that breeding-back should focus on.

Curvature

Some authors compare the curvature of the aurochs’ horns to a spiral, which they call the “primigenius spiral” [1]. Because this is a useful term, I’ll continue to use it when referring to their characteristic shape here, which consists of a curve going outwards and upwards at the base, then curving downwards again, forwards and inwards, and finally inwards and upwards at the tip.

Image by Miguel Omar Ruiz Hernandez showing the 3-dimensional shape of the aurochs horn

The base was the thickest part of the horns and the tip usually ended slim and pointy, what surely was intensified by the keratinous horn sheath in life.

There are a lot, possibly hundreds, of preserved bony aurochs horn cores found in- and outside Europe. This amount of material tells us that the primigenius spiral shape was always present [1], but there was variation in the way the horns exactly showed that curve. Some individuals had more wide-ranging horns, such as a skull displayed in Stuttgart, while others had very inwards-facing ones, such as the bull from Prejlerup. 

Pleistocene skull with wide-ranging horns found, Stuttgart (Photo: ©Markus Bühler)

Life reconstruction of the Prejlerup bull, Denmark, with very inwards-curving horns. All rights reserved. 

One aurochs might have had comparably slim horns while another one had very thick ones. Also the angle in which the horns were attached to the skull varied. According to Cis van Vuure 2005, the angle varied between 50° and 70° and also other authors found 60° to be the arithmetic mean. However, I have seen photos of skulls which show an angle either larger (Vig) than 70° so the total scope might be slightly larger.

Bull skull from Italy with sharp angle between snout and horns

Bull skull from Vig (Denmark) with more upright horns

The position of the horns relative to the skull changes the way how the horns look in frontal view. The larger the angle to the snout is, the more < > -shaped the horns appear (I don’t have word for that, some call it “lyre shaped”, but a lyre looks more like the horns of Steppe cows to me). The lower the angle is, the more the horns look like that of good Maronesa or Lidia bulls. My aurochs horn model, made from DAS clay and painted with acrylics, photographed from different angles, illustrates that very well.

For anyone interested, the model measures 78 cm in length and has a circumference of 40 cm at the base.

Reconstruction of a lost skull from Berlin. All rights reserved.

Size

According to Cis van Vuure, the smallest and largest horn diameters at the base are 7 and 18 cm, the smallest and largest length 40 and 107 cm. These measurements were obtained from bony cores, so we have to consider the keratinous sheath, which adds about 10-20 cm in length [1] and 8% (therefore a few centimetres) to the circumference at the base [2]. A large Pleistocene skull that was found at Mannheim, Germany, had horn cores with a length of 93 cm and a horn span (from the outermost point of one horn to the other) of 123. Large skulls like these are no exceptions, the probably largest known aurochs skull is exhibited at the British Museum, London, and measures incredible 91,2 cm in profile. Surely this must have been a huge bovine. Other Pleistocene samples from Italy, which was a refuge for aurochs and other mammals of the interglacial fauna during the Würm glacial, show aurochs of very large size as well. Some horns of the Chiana Valley have a circumference of ~43 cm, the largest specimen are known from an area near to Rome, which had a length from 69,2 to unbelievable 120 cm (still without keratin) [2]. The horns of the Indian aurochs where about the same size, but since the B. p. namadicus was slightly smaller overall than B. p. primigenius, they were proportionally larger. With the end of the ice age the horns of the aurochs tended to be less huge and wide-ranging [2,3], but were still impressive. This reduction of size is probably connected to the disappearance of large feline predators, and the same happened to Bison as well after Panthera leo atrox died out, when you compare the horn dimensions of Bison priscus with those of extant B. bison. However, aurochs still had to deal with wolves in the Near and Middle east and North Africa.

Ridiculously huge pleistocene horns, found in Germany. 

Very large skull on display at Lodon. Photo © Marie Griggs

Sexual dimorphism

The horns of the aurochs are said to display a clear sexual dimorphism [1,2,3]. Usually the horns are described as being larger than those of the cows, which sounds plausible (but read on). Of the skulls I had a look on (and I know really a lot of photos of aurochs skulls), cows seem to have more wide-ranging, more upright and less strongly curved horns than the bulls. A considerable size difference is not really apparent for me, but I trust the experts. In domestic cattle, the bulls usually have more forwards-pointing horns while those of the cows are longer but thinner – how can that be? First of all, who knows how much keratin plays a role here. Furthermore, it is not always easy to determine the sex only from fragmentary skeletons, and there were probably exceptions regarding the horns within one sex of the aurochs as well. The Vig specimen for example, which is thought to be a male because of its large size and massive bones, has very female horns (they appear female in being very upright by aurochs standards, thin and do not curve inwards very strongly). What is strange is that in domestic cattle of the same breed the horns of the bulls are usually straighter and more outwards than in cows, you can see that in a number of breeds (Sayaguesa or Heck, for example).

Horn sheaths

A number of keratinous horn sheaths still exist; their either survived in turf or as ornamented (drinking) horns which where owned by the European nobility. One of the best-preserved and best-known aurochs horn sheaths is that of the last aurochs bull which died in 1620, which is now kept at the armoury of Stockholm. It shows the least discolouration because it is the youngest, and also is very slim compared other horn sheaths of the middle ages or older bony cores, what probably shows that the last aurochs at Jaktorow, Poland, had slightly diminished horns due to their limited range and trophy hunting [2]. The same is also known from elks but I have no reference for that at hand at the moment. The Danish national museum also has an impressive collection of a number of ornamented aurochs horn sheaths. Most of them are discoloured.

Horn of the last living aurochs bull which died in 1620 at Jaktorow

Another horn which shows only slight discolouration and mightier dimensions than the one from Jaktorow is displayed at Brussels. When looking at the photos of all those beautiful sheaths you’ll see that the base is rather massive and the tip is very slim  (consider that most of the ornamented horn tips have their horn tips cut off), more so than in the bony cores. This tells you that the sheath adds pretty much to the length of the complete horn. The photos of some horn sheaths give you the impression that the horns have a rather two-dimensional crescent-shape, instead of the classic three-dimensional horn shape of the aurochs. This might be just an optic effect of the photo, or can be explained by the fact that both their tips and bases are cut in order to attach the ornamentation.

Horn sheaths at Servaasbasiliek, Netherlands

Compared to other wild bovines

The horn curvature of the aurochs is said to be characteristic for the species, probably because most domestic cattle lost it. But is it really an apomorphy, which is a derived state which characterises the species or group you are looking at, or is it a plesiomorphy, which means that it is inherited from its ancestors? The easiest way to solve such a question is to look at close relatives of the aurochs.

At first glance, the horns of Bantengs and Gaurs do not resemble those of the aurochs; they are very vertical and the only similarity seems to be the well-pronounced inwards-curve. But taking a closer look, especially on moving pictures, shows that they actually do have the primigenius spiral, just less strongly expressed. The same is also true of the horns of many wisents. The photo below shows an individual with particularly aurochs-like horns that I photographed at the Hellabrunn Zoo at Munich. The males of the probably extinct Kouprey had horns that resemble the aurochs closely, the most obvious difference being only bristles at the tips and wrinkles at the base. But the biggest similarity in terms of horns we can find in wild yaks – some individuals have horns that are virtually indistinguishable from those of the aurochs.

Yak skull with horn curvature identical to that of the aurochs

Wisent with aurochs-like horn curvature at Zoo Hellabrunn, Munich

This tells us that the horns of the aurochs are clearly not a defining character, an apomorphy, of the species Bos primigenius. The fact that closely related species display the same horn curvature implies that their common ancestor, the ancestor of both Bos and Bison, must have had the same horn curvature. This holy concept of parsimony in my opinion also rules out Bos acutifrons as a possible aurochs ancestor because it had very long and wide-ranging horns, unlike all other extant Bos members.

In modern cattle

As I wrote above, most domestic cattle did not retain the horn characteristics of the aurochs over the millennia of domestication. Therefore the horns distinguish the wild type from the domestic type very well, and it’s important for breeding-back to achieve “good” aurochs-like horns – not only for our satisfaction, but also because horns play an important role in the life of wild cattle.

Is there any breed that has horns identical or nearly identical with those of the aurochs? To answer that question properly, horn size and horn shape, consisting of curvature and orientation relative to the skull, should be viewed separately.

Horn size

Domestication tended to reduce the horn size of cattle. Only few cattle breeds, which mostly were artificially selected for horn size like Texas Longhorn, Barrosa/Cachena, Watussi and some Heck cattle have horns the size of what we see in large-horned aurochs. Steppe-type cattle like Maremmana, Boskarin and Hungarian gray also have horns matching the size of those of the aurochs, especially in length. Some Maremmana primitivo bulls display very thick horns. Highlands have the right horn length, but their horns are mostly – with some exceptions – rather thin. The horns of Maronesa also fit into the aurochs horn size range, but a bit more on the smaller than on the larger end I would say. Most Maronesa cows also have considerably thinner horns than in the aurochs.

Maremmana bull

Cachena cow

Curvature

Dairy cow in Brazil with perfectly aurochs-like horn curvature

Surprisingly, a number of derived dairy and milk breeds occasionally show horns with a virtually perfect aurochs curvature, which was also recognized by Cis van Vuure [1]. With some exceptions, like the Eringer breed, these horn shapes are never fixed and found only in single individuals. Of the primitive breeds, many Maronesa bulls and also some cows show horns with a curvature almost identical to that of many aurochs skulls, although the orientation relative to the skull is smaller in Maronesa on average. In many cows however, the horn tips face too much outwards, in some individuals to such an extent that they look corkscrew-like. The same is true for the related breed Barrosa, where the horns of cows are quite vertical but some bulls of this breed have a very aurochs-like curvature. 

Maronesa bull with perfectly aurochs-like horns, albeit oriented slightly too low

Barrosa bull with aurochs-like horns

The Spanish fighting cattle have horns of variable curvature, and some of them display a very good resemblance to those of the aurochs. Many other Iberian breeds have horns which curve in an aurochs-like manner but show some more or less clear differences. For example, outwards-curving horn tips in cows are quite common, but that does not rule out that some single individuals of breeds like Sayaguesa show decently inwards-curving horns.

As portrayed in a previous post, single Highland cattle individuals show a very aurochs-like horn curvature. The range of horn curvatures we see in Heck cattle are extremely variable too, and in improved lineages the Wörth lineage or Taurus cattle we see individuals with horn curvatures resembling the aurochs very closely as well, although in the former the horns tend to face too much outwards and the latter needs more inwards-curving horn tips as well. Even some Watussi cattle display horn shapes reminiscent of the aurochs, although often curving not strongly inwards enough and being to upright.

Sayaguesa cow with horns facing inwards

Taking both horn size and curvature, the number of breeds which fit the bill gets considerably smaller. Actually there is no breed in which the horns always fit the aurochs both in size and curvature. Some primitive Barrosa bulls come very close in absolute dimensions and curvature, the same goes for primitive Maronesa bulls albeit the horns are oriented lower in this breed than in the aurochs. The horns of most of the cows however do not resemble the aurochs that much; in female Barrosa they tend to be more upwards and lyre-shaped, while in Maronesa the often show a thinner, corkscrew-like shape and are considerably lower than in aurochs cows.

Wörth Heck cow "Erni" with slightly oversized horns (in relation to the animal) which are perfect in absolute size and curvature

The overwhelming majority of Heck cattle of this world have horns far from being identical with those of the aurochs. Yet there are a few individuals of improved lineages that have horns that resemble their wild type very closely. My favourite example is the cow Erni of the Wörth breeding lineage. Surely her horns are slightly too large in relation to the animal, but the absolute dimensions are absolutely authentic. I saw that cow in real and her horns give a lively impression of what an impressive animal the aurochs was. Apart from that, the curvature of the horns are absolutely correct. The Wörth lineage has some more examples of correctly aurochs-like horns, but they are not stabilized even in this lineage.

Literature

[1] van Vuure, Cis: Retracing the Aurochs - History, Morphology and Ecology of an extinct wild Ox. 2005.

[3] Frisch, Walter: Der Auerochs – das europäische Rind. 2010.

[3] Hans-Peter Uerpmann: Der Rückzucht-Auerochse und sein ausgestorbenes Vorbild. Neandertal Museum 1999.