Wednesday, 26 October 2016

News from the Auerrindprojekt

After the depressing news of Johnny's death in August, there are some good news of the Auerrind Project again.
Domka, one of the three Hungarian Steppe cows of the project, is highly pregnant from Johnny the young Sayaguesa bull. This calf will be the first one of this combination in breeding back (and perhaps world wide) and I am curious on its looks. In sum, it will probably resemble some Taurus and Tauros cows a lot, but it will be interesting to see the amount of dilution in its coat, sexual dichromatism, horn orientation and so on. Of course one individual alone is not representative for the mean average of a certain breed combination.
Pregnant Domka
Currently, the Steppe cows are also grazing with a Cachena x Steppe cross calf. Its father is of Felix Hohmeyer's black Cachena variant and it is not going to be used for the Auerrindprojekt. 
Cachena x Hungarian Steppe
Another interesting photo was recently presented on auerrind.wordpress.com. It shows the breeding bull of Lorsch, Thando, a young Watussi bull. Thando is going to cover the young Sayaguesa and Grey cattle cows and the new photo reveals his impressive horn volume. I am especially looking forward to see the combination of Sayaguesa and Watussi; I think that a 25% Watussi and 75% Sayaguesa or a good "true F2" has the potential to look very impressive already. The winter coat would probably be sufficient, but might need a boost by Grey cattle genes subsequent generations. 
Thando, the young Watussi bull at Lorsch

Sunday, 24 July 2016

An aurochs-coloured Sayaguesa x Chianina bull

Several Sayaguesa x Chianina cross animals have been born in the Lippeaue so far, when the Sayaguesa bull Churro covered the Chianina cows at Klostermersch. Two cows of that combination are still in the Lippeaue population (Bionade and 79 810). Both cows have a diluted coat colour, which is to be expected since Chianina has mutated alleles on at least two dilution loci (Agouti and Dun, Olson 1999), and at least one of them is semi-dominant. Heck x Chianina cows are diluted as well. 
Heck x Chianina bulls had a diluted fur colour as well, but to varying extent. Leonardo, the half-Chianina bull that was sold to Denmark, had a light colour saddle but had black-coloured areas too. Luca, a half-Chianina bull that was covered here extensively already, was mostly coloured in dark brown with much beige areas (mind that winter coat is always darker than summer coat in cattle). 

So we would expect a Sayaguesa x Chianina bull to have a colour similar to that of Heck x Chianina bulls. In 2014, I was sent a photo of a young bull of that combination by Matthias Scharf. That individual was of a greyish tone and more diluted than the cows of the same combination, and even way more diluted than the half-Heck bulls. This was surprising to me, especially since the colour of Sayaguesa seemed to be more intense and dominant in crossbreeds with Heck. The bull was slaughtered because of its colour and the tiny horn in autumn of the same year. 

Nevertheless, while searching through my Lippeaue photo archive I was provided by Matthias Scharf, I discovered another young Sayaguesa x Chianina bull. Its number was 42 624 (born 2010, dispeared from the stock list after 2012), and like Bionade its parents were Churro and Eloisa, so they where fullblood siblings. Interestingly, this bull's coat colour was not diluted at all, it even had only a very faint saddle that was not visible in its winter coat and might have gotten outgrown if it would have reached full age. 

The photos show him at the age of one  (uppermost) and two years. As you see, the horns of this individuals were still meagre, but that is simply what you get from that combination. It seems that its hump was not that developed as in its grey coloured halfblood brother of the same combination, but it was still high on the legs at this age at least. I don't know whether 42 624 was slaughtered or sold alive, in any way it was not kept in the herd. 

Whether or not this bull might have grown tall or well-shaped, it shows two facts: 1) looking at just one individual of a breed combination is by far not enough to judge the potential in it, especially for F1 individuals, which are, after all, genetically of little relevance; 2) there is potential for accurately coloured Sayaguesa-Chianina crosses even in the first generation. Combining those two breeds might result in very useful animals for further breeding in being large, tall, muscular and well-shaped, with a good skull shape and colour. But of course the whole spectrum from that to disappointing is possible, and most results would be inbetween. In any case, if 42 624 would have been kept for a few years more he might have produced interesting results with his (half-blood-)sisters of the same combination. However, I understand that this animal was removed because of the negative influence on horn shape and size it might have had since its horns are far away from the breeding objective. 

As I recently reported, the Auerrindprojekt is now planning to produce some animals of that combination. Maybe we are going to see some F2 Sayaguesa-Chianina in the future. Of course either large quantity or simply luck is needed to unite the full potential of both breeds within early cross generations. 

Friday, 22 July 2016

Daniel Brühl to play Lutz Heck in a 2017 film

German actor Daniel Brühl is to play Lutz Heck in a war drama film scheduled to be released next year. 
The film, directed by Nici Caro, is named "The Zookeper's Wife" and covers the story of Dr. Jan Żabiński, director of the Warsaw zoo and his wife Antonia, who saved many lives during the German occupation of Poland (see here). Apparently, Lutz Heck's character is to be in this movie too. 

Daniel Brühl recently appeared in movies like Captain America: Civil War (2016), or Rush (2013), where he stars as the Austrian Forumula 1 driver Niki Lauda. 

I guess Lutz Heck never thought that his person is going to appear in a British-American movie production, neither did I. I am looking forward to see Brühl as Heck, since his portrayal of Lauda on screen was awesomely authentic; and maybe one or two times the aurochs might be mentioned as well. 

Thursday, 21 July 2016

New breeding herd at Lorsch, Germany

Although there has been a setback for the project recently, there are good news from the Auerrind project. They just set up a breeding herd composed of Thando, the young Watussi bull, and three young Sayaguesa cows plus two young Maremmana cows at Lorsch. 

The reason behind that is that Thando is still not ready to cover the Chianina cows, as originally planned, but the time can be seized to produce some experimental crosses. I am happy because the combination Watussi x Sayaguesa will be interesting to see; I think that a cross animal with a high portion of Sayaguesa and Watussi influence has the potential to optically resemble the aurochs to a large extent, especially because the Sayaguesa the Auerrindprojekt is working with are from a good herd. Of course the work would not be done with that alone, especially because the sensitivity to cold climate of Watussi needs to be compensated, but without question the results will be very interesting to look at. 
Photo owned by auerrind.wordpress.com
As I wrote above, these combinations are experimental crosses to get more clues on the heredity of certain traits like horn size or fur colour. The original plan of a Watussi x Chianina strain is still a goal of the project. 

EDIT: I was just informed by Claus Kropp that it is also planned to let the Chianina cows be covered by the young Sayaguesa bull in the meantime - that means more of the combination Sayaguesa x Chianina which we already know from the Lippeaue, and I am happy about that. A "true F2" of that combination has the potential to result in large, well-built animals with an acceptable colour. The horns would be rather small, but since the project is also working with Watussi, I do not worry that much about that aspect.

Sunday, 17 July 2016

A very unusual aurochs skull at Cambridge

This post has been made possible by Peter Stockwell from the UK who addressed me to this specimen and provided me with interesting photos and information – many thanks for all the effort!

The Sedgwick Museum of Earth Sciences, Cambridge, UK, has several skulls or skull fragments on display labelled as aurochs. But one of them looks really atypical in having upright horns of a comparably weak curvature. The orientation of aurochs horns in relation to the skull usually varies from 70° to 50° according to Van Vuure 2005, some skulls might slightly brake the rule but it is apparent even from that broken frontal bone that the horns of this individuals seem to have an angle of beyond 90°, perhaps even 100°. Actually, these horns are barely like those of any other known aurochs crania but resemble those of many domestic cattle forms.
Some aurochs skulls, such as that of the Vig specimen, have horns that are more upright than the average. And the curvature varies from tight and narrow to more wide-ranging. But this skull definitely is a big leap from the end of the spectrum with no intermediate forms that I know of.
The photos are owned by Peter Stockwell.

The question is, then, if the skull fragment is that of an aurochs at all. The horns are, despite being atypical in orientation and curvature, still large and thick compared to the frontal bone carrying them. Unfortunately, much of the cranium is not preserved so we cannot check it for other diagnostic wild type traits, such as a large braincase, elongated skull (especially nasal bones), comparably small orbitals, straight to slightly convex profile and other features. The frontal bones, however, are obviously broad and well developed and the measurements I was provided with show that the specimen was in the size range of large domestic bulls at least (the distance between the horns on the specimen is 23cm, which is between those I find my two Taurus bull skulls; looking at the skulls and the individuals they are from I expect some variation on this metrical trait and I have no measurements from aurochs at hand) and therefore compatible with the aurochs (not all aurochs were giants, and the skull does not seem to be significantly smaller than the more typical skulls next to it on the photos). The horns are large also in absolute size. The distance between the complete tip and the broken tip is 83 cm, the circumference at the base 43 cm, which is well within the aurochs size range and well larger than in domestic cattle.
But equally as important as physical traits to find out the true nature of the skull fragment are location and age. I was told that the specimen was excavated at Barrington, Cambridgeshire, UK, in the year 1900. The exact age of the material was unfortunately not to be found out. But judging by its state, it is very plausible that it is older, or even way older, than mere two or three millennia, so it is definitely possible that the individual belonged to the predomestic British aurochs population. If the skull fragment is as old or younger than the arrival of domestic cattle on the British isles in the Neolithic, it is possible that the atypical horn shape of this individual is the result of interbreeding with domestic cattle. It has been supported by genetic data recently that local aurochs left a genetic trace in domestic cattle of Europe in several cases, but the reverse is possible as well – domestic genes may on occasion have left a trace in local wild populations, as it also happens between wolves and dogs or pigs and wild boar. It is likely that these domestic alleles are not that successful in the wild gene pool, but may produce variations visible in single animals, and this atypical aurochs skull might be one example if it is geologically possible. Precise dating and/or an aDNA test could resolve that question.

But let us assume this individual was a pure, predomestic aurochs. Should this deviant skull allow a broader range for what is permitted in breeding-back? I would say no: this skull is obviously a unique, atypical one, one of those very rare cases in a wild population. Furthermore, all existing breeding-back strains are rather variable concerning horn curvature and it is apparent from existing breeding projects that removing all those variants from the pool takes a rather long time. Allowing that kind of horns in a herd would make it even more difficult to establish the typical primigenius curvature, especially since we do not know the particular genes that play a role in the development of the shape of the horns. Besides that, undesired traits are going to reappear on occasion anyway, so this kind of upright, not tightly curved horns will be probably among them because of its common presence in domestic cattle.

Saturday, 9 July 2016

News from the Auerrindprojekt, Germany

It has been long ago since my last post. The reason behind that is that I had a busy semester at university, but now the summer has started and I have some time for the blog and artworks. At the moment I am working on two new aurochs models made from polymer clay, since I sold my first ones to the Alpenzoo Innsbruck. I hope that my new models, again male and female and to same scale, will be more artistically refined and more anatomically correct. 

The Auerrind Project has started a new breeding herd in Einhausen, Germany, containing two Hungarian Grey cows plus a young Sayaguesa bull named Johnny. One young H. Grey cow will follow next year, as much the two Maremmana cows. 
It is likely that Johnny will become a very useful breeding bull, since he is from the herd of Peter van Geneijgen which has rather beautiful animals (see this post on the Auerrindprojekt's webpage for pictures), and the other Sayaguesa the Auerrind Project purchased from that herd look very good as well. Unless I am wrong, part of the Sayaguesa stock of the Tauros Project and the Lippeaue are of Peter van Geneijgen's stock too. 
Johnny will probably cover the cows this summer, so the first calves of this combination should be born in spring 2017. 

What can we expect from the combination Sayaguesa x Grey/Maremmana? The results are surely going to be well-suited ecologically with an effective winter fur, because Sayaguesa does well in German winters at least and Grey cattle are adapted to the harsh conditions of the Puszta and other Eastern European landscapes. Sayaguesa is a comparably large breed, and Grey cattle medium-sized, therefore the cross results will be somewhere in between (actually they will vary along this spectrum), so most of the results should become larger than Heck cattle. Stature and body shape should be good as well. The fur colour will probably have the usual spectrum we have in breeding back herds: some individuals show a diluted colour, others do not, some have sexual dichromatism to a varying degree, others do not. I cannot predict what type will be predominant. The horns will probably show a size range between what we see in the pure breeds. Since truly inwards-curving horn tips are rare in Sayaguesa, especially in cows, most cows will probably have more or less outwards-facing horn tips. I am looking forward to see the first cross results of the Auerrind Project. 

A while ago I did some coloured sketches illustrating my thoughts on what early Auerrind crosses might look like. They are just guesses that show the countless combinations of traits that could be possible. 

The uppermost drawing is what I think a 75% Watussi and 25% Chianina bull might look like, or a more progressed animal with a high Watussi portion. I cannot predict how dark bulls of that combinations might get, so I took dark Watussi bulls as a reference. The next drawing is a prognosis for a first generation Watussi x Chianina cow. I used a Grey x Watussi cow from Hortobagy and a half-Chianina bull from the Lippeaue as a reference. Perhaps the horn size is a bit too optimistic. The third picture is my prognosis for a first generation Sayaguesa x Grey cattle bull. I used pure bulls of both breeds as a reference and theorized that Grey cattle would cause a saddle but not completely remove all the red pigment from the fur, but there are many possibilities. I did not illustrate a cow of that combination, because I think that many existing Taurus and Tauros cows are perfect models for what a cross of Grey cattle and Sayaguesa might look like. The lowermost drawing shows what I imagine a more progressed Chianina-Watussi mix would be, or maybe a "true" F2. 
Of course dozens of such prognoses and trait combinations are possible, ranging from superior animals to "oddities", so my sketches purely arbitrary, but maybe some of the future Auerrind animals might resemble one or two of those. 

In any case, I am very much looking forward to seeing how all these combinations will work out. 

Friday, 1 April 2016

Uncertain aspects of the aurochs' live appearance

Most of the external and morphological traits of the aurochs are well known to us thanks to numerous superb skeletal remains and also contemporaneous artworks and written sources. While the skeletal material is complete, well-preserved and numerous enough to give us a very precise picture of the aurochs’ morphology, its dimorphism in sex and also, to a certain degree, variation along time and region, artistic impressions and written accounts are sometimes not comprehensive or unambiguous enough to clear all open questions. Do not get me wrong, the picture of the aurochs we have is very precise. But there are some aspects that are questionable and still leave some room for speculation. In this post I give you overview over all those points that come to my mind.

The length of the dewlap

It is unquestionable that the aurochs had a dewlap of a certain size. Tropical bovines tend to have a large one (banteng, gaur, kouprey), more northern ones not so much (bison). Zebuine cattle have a large dewlap as well, probably both due to domestication and climate. The European aurochs, on the other hand, probably could not effort too much heat loss caused by a large dewlap, and historic references probably would have mentioned it if it was an obvious trait. But they do not, and all artistic depictions, including cave paintings, show short dewlaps. Most likely the dewlap of European aurochs was about as short as in cold-adapted taurine cattle (yakut cattle, Highland cattle, Heck cattle at the Oostvaardersplassen). It is possible that it was even shorter, but we cannot know because of the absence of soft tissue aurochs mummies (there was, actually, a complete skin of an aurochs found in Germany at the beginning of the 20th century, but its soft tissue has not been preserved [Frisch, 2010]; there might be more cases like that).

Winter fur length

The aurochs most certainly had a bi-layered winter fur. The undercoat was likely short and dense, but the question is how long the outer coat was. Surely the length and density of the winter coat was dictated by the requirements given by the climate, so it was dependent on region. I have been wondering about the winter fur of central and northern European aurochs. Since there are no preserved aurochs skins, we can only guess how long the winter fur of these kind of aurochs actually was – what was the minimum, what was the maximum of what is physiologically useful and what was the actual case?
Schneeberger states that the aurochs was covered in longer fur than domestic cattle, but does not clarify whether this was generally the case or during a specific season, and what kind of domestic cattle he was thinking of is unclear either (hardy landraces do have a more efficient insulating coat than more derived cattle). Looking at hardy taurine landraces that are adapted to living outside all year round in temperate climate could provide an useful aurochs analogue. HungarianGrey cattle and Heck cattle sometimes grow a comparably long (but still not as overlong as in Highland cattle) woolly winter fur that gives them a shaggy appearance. In my subjective, google-based perception this is particularly apparent in some individuals from the Oostvaardersplassen, but I might be wrong. Many Grey and Heck cattle however have a winter fur that is a little shorter and similar to that of Betizu and Yakutian cattle, two breeds which are good models as well – the former having a feral history, the latter being well-adapted to harsh cold. However, Chianina have a comparably short winter coat yet they still do just as well in central European winters as do Heck cattle and Sayaguesa living in the same area. So was the aurochs’ winter coat minimalistic and as long as in Chianina, was it long and woolly as in some Heck and Grey cattle individuals or somewhere in between like in the majority of Heck cattle, Betizu, Yakutian cattle and other landraces? Size could be a factor: a large animal has, in relation to its size, lesser heat loss than a smaller one of the same morphology. Therefore a 180cm tall aurochs bull might not need a winter fur as long and shaggy as a 140cm tall Oostvaardersplassen Heck bull (I am not saying that this is the reason why Chianina have a comparably short winter coat, I consider that a coincidence that is the result of how Chianina have been husbanded as a breed). But it is questionable how influential that factor really is here, especially because domestic cattle and aurochs even at that height difference might still have a similar mass due to their different morphology. Another factor is the ability to store fat, which contributes to insulation as well and therefore influences the need for an insulating coat.
While it would be nice to know how long and shaggy the winter coat of central- and northern European aurochs was exactly, I think it is not something that breeding projects have to worry about in particular, as long as the animals do well during winter and seem to be fit for survival under natural circumstances.

The colour of the dorsal stripe

The immediate answer you get is “light-coloured”. But extant wildtype coloured cattle actually display dorsal stripes of various colour shades, ranging from yellowish-white over yellowish-red, golden, shiny red to brown. The English translation of Schneeberger’s report in Gesner 1602 says the dorsal stripe was of a lighter colour as translated in van Vuure 2005, originally it says “subnigra” in Latin. This is a bit imprecise but must have meant some kind of grey. Sigismund von Herberstein was in possession of an aurochs skin, and termed colour of the dorsal stripe “grahlat”, what is to be translated with grey. More precisely, this grey colour was the result of white hairs mixing with the black hairs that covered the rest of the body. The only cattle having a grey eel stripe without any shades of red or yellow are those that display a diluted coat colour, such as Podolian cattle. Contemporaneous artworks give no useful clue on the colour of the dorsal stripe; some of paintings at Lauscaux and Chauvet cave implicate that it was there, but do not point to a specific colour.
This opens the following questions: did all wild aurochs have a dorsal stripe of the same colour or colour variation spectrum, and was it as wide as in wildtype coloured domestic cattle? Or were wild aurochs, at least the central-northern European ones, different in having a greyish eel stripe without having diluted colour shades at the same time? It might also be the case that, assuming the colour of Herberstein’s skin was not artificial (due to damaging or discolouring), the dorsal stripe in that individual was simply reduced, perhaps due to aging. I have been playing with the thought that not all wild aurochs always had a visible dorsal stripe for quite some time, in the same way the extent of the muzzle ring in gaurs and banteng is variable. But the question is, was Herberstein’s individual an exception, or representative for the rule? And if the latter was the case, was this true for all wild aurochs or just the central-northern European group?
Because of these uncertainties, I don’t stick to a certain colour shade for the eel stripe, because evidence is simply not compelling enough neither as the rule for central-northern European aurochs nor all wild aurochs. Intuitively I think that the aurochs, at least the African, Near Eastern, Asian and probably also the southern European populations, might well have displayed the diversity of colour shades for the eel stripe that domestic cattle with an un-diluted, wildtype colour do. If I had to pick only one colour for the dorsal stripe of a central-northern European male aurochs, I would take a very pale, greyish one because of the literature references – but other colours might do it just as well.

Width of the dorsal stripe

The dorsal stripe is usually described as “narrow”. To be exact, Schneeberger stated it was “about two fingers wide”, therefore a few centimetres. Von Herberstein, who owned an aurochs hide, made no mention of the width of the stripe. The “two fingers wide” condition is most common among wildtype coloured domestic cattle. However, in some cases, the eel stripe is rather broad, such as in some Lidia, Maronesa or Heck bulls. In these individuals the anterior end of the stripe is V-shaped, meaning it starts rather broad at the shoulder area (this area is usually where the dorsal stripe is widest, it is narrowest in the lumbar region or at its posterior end). The opposite is the case as well. There are bulls which have a rather narrow, reduced eel stripe, for example some Sayaguesa or, again, Lidia bulls.
Due to the scarcity of data, we do not know whether there was variation in the width of the dorsal stripe or not, and if that variation was as big as in domestic cattle or less.

The colour of the forelocks
This is a question that has been puzzling me for years now. We know from historic references and also artworks (heraldry, Smith’s painting) that the European aurochs had frizzy, curly forelocks that we also see in many taurine cattle. In wildtype coloured domestic cattle, these forelocks (or more precisely, the area between the horns on the front head) show a colour that ranges from light blond over orange-reddish, reddish-brown, dark brown to completely black as the rest of the facial hair. Cis van Vuure considers lightly coloured forelocks in bulls a discolouration that occurred after domestication (in wildtype coloured cows, a lighter coloured area between the horns is a standard feature and also supported by a cave painting in Lascaux). Written contemporaneous accounts suggest that this area was coloured just like the rest of the face: Schneeberger mentions the eel stripe, so he might have mentioned a special colour of the forelocks as well if there was one (however, he did not mention the muzzle ring either, although cave painting suggest its presence in wild European aurochs). Von Herberstein was in possession of two belts made from aurochs forelocks, and the colour of these forelocks was supposedly black based on his descriptions. North African engravings showing the African subspecies depict bulls with a light colour saddle on its back, in at least one case even the light muzzle ring (see Frisch 2010; I am not referring to those tomb paintings that most likely show domestic cattle described in van Vuure 2005). Again, if these bulls had forelocks of a special colour, they might have drawn them just as the colour saddle and muzzle ring. Furthermore, the aurochs, or at least black bulls, are popular symbols in heraldry all over Europe, but not a single emblem shows a bull with lightly coloured forelocks.

Furthermore, lightly coloured forelocks might correlate with a reduced sexual dichromatism or at least a reduction of melanisation. Light forelocks often correlate with a colour saddle in bulls, f.e. in less melanised breeds such as Alistana-Sanabresa and Cachena, while blond forelocks are never to be found in a dark breed such as Sayaguesa. All cases of bulls with a colour saddle that I have seen so far show lightly coloured forelocks, whereas in Maronesa, one of the few breeds with a well marked sexual dimorphism, most bulls have either black or dark forelocks, and never a saddle. Also Heck lineages with an (by the breed’s standard) acceptable dichromatism show bulls with mostly black or dark forelocks (I am thinking of the herds at Hellabrunn, Tierpark Haag, the Neandertal and former Wörth herd), with some exceptions. There are a number of Lidia bulls that show blond forelocks, and Lidia is, just like Heck cattle, heterogeneous also in terms of sexual dichromatism.

Gaurs and Banteng also a have lighter coloured area between the horns. In the Java Banteng, which is the extant bovini with the most strongly marked sexual dichromatism, the light area is not part of the coat colour, but actually a keratinized area. The gaur has a very reduced sexual dimorphism – but not completely absent, you can still see that cows usually tend do be not quite as dark and blackish as the bulls are. Both sexes have lightly coloured hair between the horns. One could argue now that having a light area between the horns therefore is a plesiomorphic condition that the wild ancestors of taurine and zebuine cattle must have had as well. But this must not necessarily be the case; white muzzles, or eye rings as much as dorsal stripes are all characters that originated multiple times but perhaps have the same underlying genetic mechanisms (note however that the dorsal stripe in horses and bovines are different).

While it is just a speculation that lightly coloured forelocks in bulls are the result of reduced sexual dimorphism, evidence does not support this trait for wild aurochs bulls. All sources suggest that the forelocks of the aurochs were of a dark or black colour, so van Vuure might well be right with considering lightly coloured forelocks a discolouration that occurred after domestication. But the data is in my opinion not comprehensive and precise enough to rule out this trait for wild aurochs, due to the uncertainty it should also not be bred against in breeding projects.
But if I had to pick only one colour for the restoration of the forelocks of a wild European bull, I would take black or dark forelocks, which is the colour I usually use.


Some of you might already be familiar with my idea that at least the European aurochs had a kind of “mane” additionally to the frizzy, curly forelocks between the horns because of a post in 2015. By “mane” I do not mean an opulent mass of hair such as bison have, but rather just curly, perhaps only slightly longer, hair on the head, neck and perhaps also shoulder area. This trait would be present in bulls only, while curly forelocks can be present in both sexes. Actually a lot of domestic bulls show such a “mane”, though to a variable extent. I came up with the idea that it might be a functional aurochs trait when I read that this “mane” (which is also where I got the name for it from) is suspected to protect a Chillingham bulls’ skin during fight from the horns of the opponent, because these areas are the most exposed during such a combat. Then I noticed that a number of Oostvaardersplassen bulls actually have this kind of mane too, while I have seen barely one such a case for Heck cattle outside the OVP. So this might suggest that there is some kind of selective advantage for this trait, although it is only speculative. Lidia is another example for a breed were we find some bulls with a prominent “mane”.
Curly hair is a typical bull feature, and the mane was not necessarily that eye-catching that historic witnesses must have mentioned it. That is also why I gave my aurochs bull model from 2014 curly hair on face, neck and throat, and I think I will do so again when doing my next aurochs model. However, only preserved aurochs skin, a notion in historic references or contemporaneous artistic impressions can proof if some bull aurochs had such a mane. The various spots on neck and face of two of the Lascaux bulls (the line drawings) might indicate such curly hair, but that is only a guess of mine.

Aureole around the eye
Ocular aureoles, a white ring around the eye, are common in all kinds of ruminants. It is also common in Bos. One subspecies of Banteng, Bos javanicus birmanicus, have them on regular basis, other subspecies of Banteng and also gaurs have them on occasion. Aureoles are a common trait in domestic cattle as well, at least those having wildtype colour expressed. It is a trait often shown by calves that disappears later on, but there are many grown cows that show this trait. Grown bulls usually do not, except for breeds and types of cattle that are rather small and/or display a number of neotenic features, such as Cachena and (miniature) zebu. Aureoles in grown individuals do not correlate with diluted colour variants, because cows with a colour rich of red pheomelanin can have these rings too.

This provokes the following questions: Were aureoles present in wild aurochs too? If so, was it a general trait or dependent on regional variation, and was it limited to one sex in adulthood?

Artistic depictions of aurochs do not suggest the presence of ocular aureoles. There is just one case, the painting of a black male aurochs at Lascaux, that shows a white ring around the eye. But it is unknown whether the artist really wanted to depict an eye ring or merely intended to implicate the presence of the eye. CharlesHamilton Smith’s aurochs painting has a slim white ring around the eye that might be interpreted as an aureole, but this is only a second-hand artwork based on an artwork itself. Apart from those two cases, no artistic representation of the aurochs indicates the presence of eye rings, neither do any historic references.

This and the fact that only bulls that otherwise display neotenic traits and very small size have eye rings as adults makes it unlikely to me that at least the European and African subspecies had such aureoles as grown males. But for cows, it seems like an open question to me whether at least some of them had them or not. I tend to think that grown European female aurochs usually did not have light eye rings, mainly because there is no mention of it and none of the artistic depictions show them (I am thinking of coloured cave paintings).  In the end, we cannot know whether some wild aurochs cows (and also bulls) had this trait or not, and if it even was a general characteristic for certain populations or subspecies.

It turns out that even the white muzzle ring or mealy mouth is not that solidly supported by the data either. There are no written accounts for it that I know of. However, a number of cave paintings show it, or at least point to it, and also Smith’s aurochs shows a reduced one (OK, here goes the same as for the eye ring). But what is also striking is that it is a fixed, permanent trait of nearly all wildtype coloured cattle on this world; there are only a few exceptions where the muzzle ring is either strongly reduced or barely visible due to some modifiers, but basically it is always present in E+//E+ cattle. So it should have been in wild aurochs too. However, it could have been the case some individuals, especially bulls, that it was reduced or even very reduced when they aged, so that the white area might have been restricted to the upper lips and chin – the way we see it in a number of taurine bulls, Gaurs and Banteng. In such individuals, the “mealy mouth” would not have been such an obvious trait that it would be considered worth mentioning anymore. Perhaps such a reduced white muzzle was even the case in fully grown (European?) aurochs bulls. But that is speculation.

Colour shades in cows

This covers two aspects: the amount of eumelanisation (how much black is in the coat colour) and pheomelanisation (when is the colour so faint that it has to be considered a domestic dilution).
Wildtype coloured cows show a spectrum from almost uniformly light brown over darkening from head, neck and legs and subsequently the lateral side of the trunk until the whole body is dark brown/black except for a more or less large light colour saddle – if that saddle is absent, the cow is coloured like a bull should be.
So, which state in this continuum is the “correct” one, at least for European aurochs? We cannot say. Artistic depictions, even the Lascaux cave alone, support the whole spectrum. It shows, among more detailed figures, cows that are drawn uniformly brown, sometimes only as lines, which might either indicate that the individuals they are based on were of that uniformly brown colour or the artist did not intend to reproduce its colour more detailed. The will to represent the animals accurately is always a problem for artistic references; if someone, say, spotted a mufflon and drew or described it being just “brown”, he might consider that sufficient, although the coat colour of a mufflon is of course more facetted. That’s also the case with Schneeberger’s report, where the cows are simply said being of the colour of calves (“chestnut”, “blackish brown”, “dark brown” etc.). There are a number of Heck and Maronesa cows those colour is of a more or less uniformly dark chestnut brown colour, which might fit what Schneeberger had in mind.
Maronesa and a Heck cow from Wörth; photo of the Maronesa cow
by Goncalo Figueira, Wikipedia
Lascaux however also includes more defined representations of cow colours, such as very dark cows with a red colour saddle. And just as there are cow depictions completely lacking black, there is also one that is almost completely black with a very narrow colour saddle at Lascaux. This colour is perfectly represented by the Sayaguesa cow posted above (bottom row centre), or the Taurus cow at its left. Schneeberger also mentions that very rarely, black cows appeared. This notion is one of the reasons why black cows are usually tolerated in breeding projects. A bull-coloured female aurochs might have been either the result of the genetic diversity within the population, or of a twin birth. When a cow is the twin of a bull, it might get infertile and develop the colour of the male because it gets more male hormones than it should (I am not a developmental biologist, so please excuse me for this imprecise language). If Schneeberger was referring to results of such “accidents”, the legitimation for black cows that are the result of lessened sexual dichromatism in breeding-back herds would be gone. But in any case, I have always thought that in an ideal case the portion of bull-coloured cows in such a herd should not be larger than 5-10%.
Regarding all the non-bull-coloured shades that seemingly were present in the population, it is the question whether they were distributed equally in the population or if there was a prevalent type, if there was a regional gradient (the cave paintings are all from southern Europe, but not the same time; so they might represent aurochs of different genetic clusters).  Artistic interpretations and written accounts are too scarce, regionally limited and not precise enough to be nailed down to one colour shade only, so that we can probably permit the full range for breeding, or at least can not rule out one particular type.

What is also striking is that there is a continuum from a rich, shiny red pheomelanin-caused portion in the coat colour and the leucistic, colour-less grey that results from the lack of pheomelanin. Alleles that cause these state are called “dilution factors”. But since there is a continuum in crossbreed populations at least, the question is: at which point are we dealing with the presence of a dilution factor and which colour shade is still wildtype?
Historic references as much as contemporaneous paintings all suggest shades like dark brown, reddish brown, chestnut, light brown, ashy for aurochs cows. There is no mention or depiction of beige tones or greyish tints, for example. The only way to tell the individuals apart that carry the probable wildtype homozygous would be to identify the modifier alleles that are responsible for the amount of pheomelanin in the coat, because we are most likely dealing with more than one locus and intermediate states, that should be the reason why we see a continuum.

Colour saddle in bulls

This is another aspect of the aurochs’ coat colouration that can be a matter of debate. Some argue that a light colour saddle in bulls was present or might have been present in (European) aurochs, at least that we cannot rule it out. I use to regard this trait as a sign of reduced sexual dimorphism just like a bull-coloured cow, and there are no written or artistic references that support the presence of a colour saddle in male European aurochs. On the contrary, most references describe the aurochs as a black or at least very dark animal – be it Plinius (“bos sylvestris niger”), the classification of the aurochs as “suarzwild” (“black game”, together with wild boar) in the Lex Baiuvariorum AD 800, a figure on the Carta Marina by Olaus Magnus (who probably saw living aurochs at Jaktorow), or von Herberstein, who held an aurochs skin in his hand and described it “entirely black” except the dorsal stripe as much as Schneeberger’s description and all cave paintings that both include only completely black bulls with the dorsal stripe.
So the body of evidence suggests that wild European bulls did not have a colour saddle, at least there is no reason to think otherwise.
However, as mentioned above, there are artistic depictions from Northern Africa that suggest at least the African subspecies had a colour saddle.

Of course it is harder to prove absence than it is to prove presence, but until there is some evidence that European aurochs bulls were not always completely black (except for the dorsal stripe and muzzle ring, and maybe forelocks), I assume that a colour saddle in a bull is indicative of reduced sexual dimorphism and something that should be avoided in breeding – at least for breeding bulls. So far, a saddle in bulls is permitted in all breeding-back projects and breeds. I think this is not effective for breeding bulls, but of course such a bull can still be desirable because of other traits, it is always about balancing traits.

As you see, a large part of these open questions or uncertainties concern coat colour, since the morphology of the aurochs is well-known thanks to the comprehensive bone material. I have also speculated that some colour types we see in domestic cattle might have evolved in the aurochs already, such as solid black all over the body (Ed allele) in wild B. primigenius primigenius, or it might have gotten transferred from domestic populations into wild ones, or that the dilution alleles that cause the greyish colour of many zebuine cattle as much as in Podolian cattle might be a wildtype trait of B. p. namadicus. There is no evidence pointing to that, these are just thoughts that I consider not totally implausible.  
One could, just like a number of authors did it with horses, test aurochs bone material for coat colour alleles. As a test to see whether those alleles we consider wildtype were indeed present in the wildtype and to see whether unexpected variants, like Ed, were present or not in wild populations. It could also be used to rule out the proposal of a Chillingham cattle-like spotted pattern for predomestic aurochs (a suspicion that is based on line drawings from Lascaux – but I consider this evidence not compelling at all).
Since a number of cattle modifier loci and alleles are only speculative, some of them would have to be identified before looking for them in aurochs material. I am not that optimistic that such a study would be conducted in the near future however, since the interest in cattle and cattle coat colours is not nearly as big as the interest for horses. Nevertheless, the data gained from such a study could be helpful for breeding projects. For example, once all the dilution modifier alleles are identified, cattle from breeding projects could be tested for those. In this way, breeding bulls carrying recessive dilution alleles could be avoided.

A while ago, I illustrated what a Chillingham-coloured aurochs, solid black aurochs and my Indian aurochs bull with Agouti dilutions seen in Podolian cattle would look like. Again, I am neither proposing that aurochs of these colour variants existed nor am I “believing” it, I am just illustrating possibilities.


Walter Frisch: Der Auerochs: Das europäische Rind. 2010.
Cis van Vuure: Retracing the Aurochs - History, Morphology and Ecology of an extinct wild Ox. 2005 

Saturday, 26 March 2016

The horses in the Lippeaue

With a major delay, I finally do my post on the horses in the Lippeaue reserve, Germany, today. I already covered them in my 2013 post, but on my trip in 2015 I gathered some more material, and the herds changed a bit as well.

As it is the case in so many reserves for landscape conservation, horses and cattle share the area because they complement each other in their grazing activity and both are native species. I don't know the total number of horses in the Lippeaue, perhaps about 40 individuals but that is only a guess.

The horse herd at Disselmersch consists of mostly pure Koniks, plus one Konik x Przewalski hybrid. The herd at Hellinghauser Mersch is, as far as I can see, completely mixed. They had a pure Konik stallion in the herd until at least 2013, but that one seems to be gone now. Now there are 50% Przewalski hybrids, 25% Przewalski and 75% Konik hybrids as much as a pure Przewalski mare. The 50-50 hybrids resemble a pure Przewalski rather close, except that the short mane is semi-erect and their head looks somehow more caballine. The colour of these Konik x Przewalski is phenotypically bay dun with countershading and a white muzzle (which seems, however, to be slightly reduced in these hybrids), which makes sense, since black (a) is recessive under bay (A+), which are the base colours for black dun and bay dun that we see in the Konik and Przewalski, respectively. The colour of the back-crosses with Koniks on the other hand is interesting to me. They should end up as either a//A+ or a//a with a 50% probability, so either the same colour as the first-generation hybrids or a pure Konik, yet at least a number of them has a colour that is somewhat more reddish and sooty than those of the first-generation hybrids. So there might be more than just one locus involved. Countershading and white muzzle are absent or strongly reduced in the Konik-backcrosses. The new dominant stallion seems to be one of those; the new generations will produce some further interesting animals because genetics will cause individuals that show mosaic traits of both horse types.
Mostly pure Koniks in the Disselmersch.
Konik x Przewalski mare in the Disselmersch.
Konik x Przewalski as much as 25% Przewalski 75% Konik crosses plus the pure Przewalski in the background.
Hellinghauser Mersch.
The pure Przewalski mare at the far left.
A 25% Przewalski 75% Konik (?)stallion. Hellinghauser Mersch.
Konik x Przewalski mare. 
The pure Przewalski mare, Hellinghauser Mersch.
One might ask why crossing Konik and Przewalski's horses at all. As far as I understood, it was more of a coincidence that the ABU had the opportunity to use Konik and Przewalski's horses at the same time (I don't know if more than the one pure Przewalski mare were used). So it is kind of an experiment, and since the number of pure P. used is very low, I don't consider it a loss for the conservation of the subspecies. 
It is an open question whether or not Przewalski's horses can or should be introduced into European natural areas - it seems that the common opinion in rewilding projects and organizations is that European domestic horses are to be preferred over the P. horse. There are several reasons in favour of using domestic horses instead of the P. horse for rewilding: 
- The Przewalski's horse is not the western, but Asian subspecies, whereas domestic horses are descendants of the western wild horse subspecies 
- The P. horse is an animal of the steppe and adapted to arid climate, shown also by its standing mane (historic references for the European wild horse describe a short but never a standing mane), whereas native European landraces are adapted to the local climate. 
- Przewalski's horses and caballine horses separated between 120.000 and 240.000 years ago, the P. horse has one more pair of chromosomes and one more thoracic vertebrae (see Wikipedia) - so whether or not you regard both these types of equines separate species or not, they diverged quite some time ago and developed some evident differences. Mixing both types, or introducing one into the range of the other, would eradicate these results of evolution. 

But there are also arguments in favour of using the Przewalski's horse for European rewilding: 
- The P. horse is a wild animal, exhibiting a behaviour that is authentic for a wild horse, including a high intra- and interspecific aggression potential but also shyness towards humans in nature, whereas domestic horses often still remain tame after several generations, with their obtrusive behaviour causing troubles with workers and visitors (see Bunzel-Drüke et al. 2008). 
- With the possible exception of the standing mane as much as the lack of the allele for a black base colour (a), the Przewalski's horse itself matches the morphology and looks of the western European wild horse quite well, as much as a number of primitive landraces do, including the "small" size, sturdy body, robust head et cetera. 
- Although it seems that the P. is a different ecotype than European wild horses and European landraces are, they seemingly do well in the climate of this continent. In grazing projects like the Lippeaue or Döberitzer Heide they show no sign of discomfiture. 
- As long as pure animals are used, wild European populations of the Przewalski's horse would help preserving the subspecies. 

That's why I proposed that a mix of local or suitable landraces combined with influx from Przewalski's horses (by using crossbreeds, not pure individuals) would be a perspective for re-stocking Europe's natural area with authentic horse populations in a 2013 post. Nowadays I am not so convinced of that proposal anymore. As outlined above, the Przewalski's horse and caballine horses developed some recognizable genetic, ecologic (including geographic) and osteologic differences during about 200.000 years of separate evolution. 
Nevertheless, Przewalski's horses have already been introduced in European natural areas such as Tschernobyl or the Puszta in Hortobagy (the latter is especially reminiscent of their natural habitat in Asia), where they do well. 
The stock at the Lippeaue, however, is not supposed to become a large, self-sustaining population in a barely regulated area one day. Neither did someone propose that Konik x Przewalski is the one and only equine model for Europe's natural areas. Instead, they are the "grazing tools" of a conservation project for the Lippeaue reserve, and these experimental hybrids fulfill their job well. 

Whether you are in favour of Przewalski influence in grazing projects or not, it seems that many Koniks might have P. introgression already due to intermixing with Heck horses, which are kind of diluted Koniks with Przewalski influence and also look that way. In the 1970s, Przewalski's horses were crossed in again at the Wildpark Hardehausen, so that about the half of their stock has a standing mane today (see Bunzel-Drüke et al. 2008). Heck horses and Koniks are often mixed indiscriminately with each other, so that they are hard to distinguish at least in German grazing projects today. For a Konik/Heck horse with a standing mane revealing its Przewalski influence, go here for example. 

Bunzel-Drüke, Finck, Kämmer, Luick, Reisinger, Riecken, Riedl, Scharf & Zimball: „Wilde Weiden: Praxisleitfaden für Ganzjahresbeweidung in Naturschutz und Landschaftsentwicklung“ 2008.