How large were aurochs cows?

When talking about the size of the aurochs, we mainly talk about the size of male individual, because the upper size limit of the species is of particular interest. The size of female aurochs, however, should not be overlooked. The size of aurochs cows also tells us about the extent of the sexual dimorphism in the aurochs. 

Aurochs bull and cow. Based on the Lund bull and the Cambridge specimen. It is a rather old reconstruction of mine, I still consider it accurate except for the raised pelvis in the bull and that the horn tips do not face inwards enough

I already did a post on the size of the aurochs in 2013 which was very male-based. It has to be noted that the size of the aurochs varied along its geographic range and also geologic age. Focusing only on the European subspecies, Bos primigenius primigenius, the individuals from the north were larger than from the south, and Pleistocene specimen were larger than Holocene specimen (which might be of anthropogenic cause). The largest credible size estimate for males I found in the literature is 2 metres (Boessneck 1956). Calculating the withers height of the individual with the largest skull ever found (91,2 cm, see Frisch 2010), based on the more or less complete Sassenberg bull (which already is a large-headed specimen) results in a height of roughly 215 cm. The smallest size given for a European aurochs bull that I found in the literature was 154 cm withers height, for a specimen from Denmark (Kysely 2008). Considering soft tissue and hooves in the living animal, we can add about 5 to 10 centimetres to the height of the skeleton. Thus, the size of aurochs bulls in Europe may have varied from 160-200cm or even more. 

But how large were the cows? 

Van Vuure 2002 and 2005 gives an average cow size of 150 cm. The Cambridge specimen, which may or may not be a cow, measures 145 cm (Frisch 2010), what means that in live it must have been between 150 and 155 centimetres tall at the withers. The average size for Danish cows is given 145 cm, accompanied by 160 cm for the bulls (Kysely 2008). The smallest Danish cow was 139 cm tall, the largest 153 cm (Kysely 2008). The smallest cow specimen morphologically assigned to the aurochs is a cow with a withers height of 132 cm (Schibler et al. 2014) meaning 137 to 142 cm in life. For Polish aurochs, a size of 170-185 cm in bulls and 165 cm in cows is given (Kysely 2008). This means that the cow might have been 170-175 cm tall at the withers in life.

So the height of European aurochs cows found so far varied between 137 cm (minimum) and 175 cm (maximum) in life, while the bulls varied from 160 cm to 200 cm or more. Van Vuure gives an average size as it follows: 160-180 cm in bulls and 150 cm for cows (van Vuure 2002, 2005). While this is probably an accurate average, this range however does not represent the variation found in the European aurochs well enough. Also I don't know if this is an estimation for living specimen or skeletons. 

While both sexes varied considerably in size, the bulls of the respective populations were always at least one or two decimetres larger than the cows. This is a dimorphism similar to what we see in other wild bovines. It would be interesting to know how large the variation was within a population - if large aurochs cows reached the size of small aurochs bulls or if bulls were always and under any circumstances larger than cows. 

What is worth noticing is that both sexes apparently varied by 40 cm. This is considerable. A reason for that might be that not all individuals found were fully grown. A strong regional gradient is not detectable since the smallest Danish bulls apparently reached 154 cm (skeleton), while the largest Danish skeleton that is mounted measures 190 cm according to various sources from the internet (the Preljerup bull). So the aurochs probably was simply diverse in size. 

Literature 

Boessneck, 1956: Funde des Ures, Bos primigenius Bojanus 1827, aus den alluvialen Schichten Bayerns. 

Kysely, 2008: Aurochs and potential crossbreeding with domestic cattle from the Eneolithic period. A metric analysis of bones from the archeological site of Kutna Hora-Denemark (Czech Republic). 

Frisch, 2010: Der Auerochs - das europäische Rind. 

Schibler et al. 2014: Incorporation of aurochs into a cattle herd in Neolithic Europe: single event or breeding? 

Van Vuure 2005: Retracing the aurochs - history, morphology and ecology of an extinct wild ox. 

Van Vuure 2002: History, morphology and ecology of the aurochs. 

How many times did the primigenius spiral evolve?

The horn shape of the aurochs can be compared to a spiral, which is called the "primigenius spiral" in the literature (see van Vuure 2005). It is universal to all aurochs specimen known, what probably had functional reasons: this horn shape is simply practical in intraspecific fights, as they enable to push and pull the opponent. 

Looking at other bovine species, it becomes obvious that the primigenius spiral is found also in species other than Bos primigenius. Wild yaks have horns identical to that of the aurochs in both dimensions and curvature (see here or here). It is also found in male koupreys (see here). Occasionally, the primigenius spiral also appears in wisent (see here).

Now the question is: did the primigenius spiral evolve independently in these species, or was it already found in their common ancestor? 

In order to resolve that question, we have to look at the phylogeny of the Bos-Bison clade. 

+--- Bos primigenius (aurochs, including taurine and zebuine cattle) 

 `-+---+--- Bos sauveli (kouprey)

    |       `+--- Bos javanicus (banteng)

    |          `--- Bos gaurus (gaur)

     `+--- Poephagus mutus (yak)

        `---- Bison  

This phylogeny is based on mitochondrial data from three studies [1,2,3]. The wisent, as it is a special case, is not included in my cladogram. 

If the primigenius spiral evolved independently in the three species aurochs, yak and kouprey, it must have evolved three times according to the cladogram. If it goes back to a common ancestor, which would be the common ancestor of the Bos-Bison clade, it must have been lost two times again, once in the bison clade and once in the banteng-gaur clade which possess more crescent-shaped horns. But did banteng and gaur really lose the primigenius spiral? It depends on the angle you are looking at the horns. When looking from above, it becomes obvious that even banteng still have the primigenius spiral (see the horns of this cow), the horns are just more upright. And considering that this shape still can occur in wisent, it might indeed be possible that the primigenius spiral is basal for all members of the Bos-Bison clade. 

The primigenius spiral is even found in Leptobos, a possible ancestor of the Bos-Bison clade, at least in the species Leptobos stenometopon (see here). 

Putting everything together it is not unlikely that the basic horn shape of the aurochs, the primigenius spiral, was basal to all members of the Bos-Bison clade. 

[1] Hassanin et al.: Has the kouprey (Bos sauveli Urbain, 1937) been domesticated in Cambodia? 2006.

[2] Verkaar et al.: Maternal and Paternal lineages in cross-breeding bovine species. Has the wisent a hybrid origin? 2004.

[3] Wang et al.: Incomplete lineage sorting rather than hybridization explains the inconsistent phylogeny of the wisent. 2018. 

The lost Berlin skull, the Sassenberg cow and the Vig bull

I am constantly trying to reconstruct as many aurochs specimen as possible to get an impression of the variation in the European subspecies, B. p. primigenius. Recently I did another three artworks, as usual by reconstructing directly over photos of the actual skeletal material, to be as precise as possible. 

The lost skull from Berlin 

There is a very large aurochs skull on display at the Natural History Museum of Berlin. But the museum also had another skull which is now lost. It was part of a complete skeleton which was on display at the museum and may be now in the collection. Here you can see the photo of the skeleton which my reconstruction is based on. 

All rights reserved. 

The horns seem to be oriented in an angle larger than 60° to the skull (which is the average according to van Vuure 2005). The horns are also not that strongly inwards-curving, at least not the bony cores. The keratinous sheath adds about 10% to the length of the horn in my reconstruction, which may even be conservative. 

The horns resemble those of some Watussi and some Heck cattle of the Steinberg/Wörth lineage, at least from this angle. 

The Sassenberg cow 

The Sassenberg cow specimen was the subject of the previous post. Now I also did a reconstruction of the head in profile view. 

All rights reserved. 

The skull of aurochs is usually more elongated than in most domestic cattle, but this is more apparent in bulls than in cows. The skull shape of cows of some landraces, for example Sayaguesa, might be similar to that of the Sassenberg cow. Note that my reconstruction of the cow is not supposed to be completely black, but only neck and face as in many wildtype coloured cows. 

The Vig bull 

I reconstructed the Vig bull already several times, here is my most recent reconstruction. Using a photo by Markus Bühler I did another one recently: 

© Markus Bühler

All rights reserved. 

I think the mount is anatomically correct enough to be used for a reconstruction, I just slightly altered the position of the hind legs. The horns of the Vig bull are comparably upright, which is apparent in this view (I estimate 85° relative to the skull). 

The Sassenberg cow

The Sassenberg cow specimen is the only mounted skeleton that is certainly a cow of Bos primigenius primigenius that I know of (the Cambridge specimen is dubious to me, it's not 100% certain that it is a cow), the other definite cow skeletons that I know belong to B. p. suxianensis (see here). It was found in Sassenberg, Germany, in the 19th century. 

Skeletal material of aurochs cows is much rarer on display than that of the bulls, probably because they are "less impressive". I assume it is unlikely that less female skeletons have been found, most of them are probably in collections. 

Due to copyright issues I cannot post the photos of the skeleton that I have, but my skeletal reconstruction down below was done by tracking out a photo of the skeleton and therefore should give an accurate impression. 

Skeleton, muscle reconstruction and life reconstruction of the Sassenberg cow specimen © All rights reserved, please do not use without permission. 

The skeleton shows that cows had a shorter trunk than bull skeletons, which is congruent with the description of live aurochs in Anton Schneeberger's letter to Gesner (1602), who wrote that cows are shorter than the bulls. It also shows that also the cows had humps in the shoulder region (not only the bulls), indicated by the elongated processus spinosi in the shoulder region.  Humps in cows is rare in domestic cattle, it is found in draft breeds like Sayaguesa or Maronesa, but to the largest extent in Lidia. My life reconstruction bears some resemblance to what a cross of Lidia, Maronesa and Sayaguesa would look like, what fits my proposal of a breeding project with those three breeds. 

Here a Sayaguesa cow for comparison (the cow Dona-Urraca from the Lippeaue): 

The reconstruction above shows the cow in its summer coat, but I also did one of the individual in a longer coat during fall: 

The Sassenberg cow with a shagger coat during fall. Please do not use without permission. 

It is not known how long the winter coat of the aurochs was. Schneeberger wrote that the aurochs was covered with longer hair than domestic cattle. Whether this was generally the case, or referred to the coat during winter and fall, is unknown. 

Some Lidia bulls with long legs

Lidia, the Spanish fighting bull, is probably one of the least-derived cattle breeds on this world. Their morphology is much like that of a wild bovine, except for the short legs that most individuals have. While browsing on youtube, I found a video that shows Lidia bulls with comparably long legs. Especially in the light brown bull at 8:48 it looks like the withers height equals the trunk length due to the long legs, which was also the case in the aurochs. 

Here is the video. 

All of this bulls have a very good physique. While they are probably young and would grow a bit heftier when fully grown, they would be very valuable for "breeding-back". 

Auerrind project update

Claus Kropp recently announced an update for the breeding plan for the Auerrind project on Facebook. 

The herd at Bielefeld on Felix Hohmeyer's farm will consist of Apollo the Watussi x Maremmana bull that will cover two purebred Sayaguesa and a purebred Pajuna cow. Another herd at Lorsch will consist of the Sayaguesa x Maremmana bull Benito which will cover two Sayaguesa x Chianina and one Chianina x Watussi cow. 
So that we can expect the following crosses for the next few years: 

Sayaguesa x (Watussi x Maremmana) 

Pajuna x (Watussi x Maremmana) 

(Sayaguesa x Maremmana) x (Sayaguesa x Chianina) 

(Sayaguesa x Maremmana) x (Chianina x Watussi) 

Apollo (Watussi x Maremmana) © Claus Kropp

Benito (Maremmana x Sayaguesa) © Claus Kropp

I am looking forward to these crosses very much. The Sayaguesa x (Watussi x Maremmana) and the (Sayaguesa x Maremmana) x (Chianina x Watussi) will be particularly interesting to me. In the latter combination, the chance is 1/3 that the recessive colour dilution found in Maremmana and Chianina will be visible, but colour is a comparably easy fix. Maybe the second generation will include the first animals that have both a spectacular body size and horn size at the same time. 

How to limit undesired traits in "breeding-back"?

Modern “breeding-back” projects such as Taurus cattle and the Auerrind project work with a breed selection that comprises all achievable morphologic aurochs traits. Size, proportions, colour, horn shape and size and body morphology to a certain degree. But the problem is that the sets of breeds also include a lot of undesired traits. Recessive colour variants, as well as alleles for very small horns (which might be recessive too, see here) and for different horn shapes. It requires a lot of breeding work to purge those unwanted traits from the population. Recessive alleles in particular are very difficult to breed out. While it is well-possible to achieve very good animals with the sets of breeds chosen, it will be difficult and longsome to truly stabilize the breed for the traits desired because of the high number of undesired traits. 
Therefore, I have been making some thoughts on a project that minimizes the amount of undesired traits but still is able to achieve all the aurochs traits that are achievable with domestic cattle. 
 
Such a project would have to chose breeds that already resemble the aurochs to a large degree and do not have any undesired recessive colour variants or different horn shapes or sizes. I think that a combination of wildtype-coloured Lidia, Maronesa and Sayaguesa would be suitable for this purpose. Lidia would contribute a very aurochs-like body morphology, Maronesa has the sexual dimorphism and horn shape (in good individuals) and Sayaguesa would contribute large size, long snouts (at least in cows) and long legs. The horn shape of this combination would be good to very good, no deviant colour variants would be present and since some Sayaguesa grow up to 170 cm withers height the right size would also be found in the gene pool. The bulls might end up a bit short-legged, however, and the horn size might not be that impressive, but overall it would be possible to breed a good result quite fast and would be a lot easier to stabilize. I would love to see Lidia x Maronesa, Lidia x Sayaguesa or Sayaguesa x (Lidia x Maronesa). 

 

The genes influencing horn size

While the genetics of coat colour of domestic animals are comparably well-studied, the genetic background of other aspects, such as the horn shape and size of cattle, remains nebulous. Only the genes for the polled and scurred conditions are resolved, while the genes that determinate the shape of the horns as well as the size are unresolved. 

 

Horn size is a quantitative trait. That means it is influenced by a larger quantity of genes and shows a continuum. The identity and influence of those individual genes is probably largely unknown, but crossbreeding results can provide a clue for speculations. 

 

The idea for this post came to my mind when I saw photos of the Taurus cow “Lippe”, which is an F2 (Sayaguesa x Heck) individual. 

 

F2 Heck x Sayaguesa cow Lippe in the Lippeaue © Matthias Scharf

It is notable that the cow has very small horns like a Chianina. Yet it has no Chianina in its ancestry, only Heck and Sayaguesa. Two breeds that have horns that can be described as at least medium sized – the horns of the Heck cattle used in the Lippeaue are actually comparably large. So it is possible to breed individuals with tiny horns from two breeds that have at least medium-sized horns within only two generations. 

How is that possible? One possibility is that the gene or genes for this tiny horn size is or are recessive, and that the F2 carries two of the recessive alleles and is thus homozygous. However, in this case we would also see tiny-horned Sayaguesa and Heck cattle on occasion. The other explanation, which is much more plausible to me, is that this tiny horn size is the result of a cumulative effect. Sayaguesa may have alleles for small horn size one the one locus, and Heck cattle on another locus. In the F1 generation these loci would be heterozygous, thus the horn size would still be medium-sized. But in the F2 generation, coincidentally, the cow might be homozygous for the alleles causing small horn size on both loci, resulting in the very small horns not seen in the parental breeds or F1 animals. It might involve even more loci, two would be the minimum. 

 

The horn size we see in Chianina might be caused by different loci or alleles, we cannot know without resolving the alleles and testing it. Crossbreeds suggest that at least some of the alleles causing the small horn size in Chianina are recessive. The crossbred Taurus bull 01 856 was the son of the bull Laokoon and the cow Larissa, two individuals with medium-sized horns. However, both parents were part Chianina (25% respectively 62,5%). 01 856 happened to have rather small horns, not larger than in Chianina. This suggests that at least some of the alleles causing the horn size in Chianina are recessive. It would also explain why many half-Chianina individuals had horns of medium size (such as the bull Luca or the cow Larissa). 

If the small horn size of Chianina is indeed recessive, this is bad news. Recessive alleles are difficult to purge effectively from the population. 

 

The bull 01 856 

If it is possible to breed horns smaller than in both parental breeds within two generations, as the cow Lippe demonstrates, it might also be possible to breed large horns out of two breeds with medium-sized horns in few generations. This would depend on which alleles the parental breeds have. 

 

The Prejlerup aurochs

The Prejlerup aurochs is from the same island as the Vig bull, namely Zealand, Denmark. It is also from about the same time (early Holocene) and thus from the same population. 

Some sources on the web claim it is about 190cm tall, and based on photos that seems realistic. That means that in life the bull might have approached 2 meters withers height. 

I have reconstructed that skeleton previously, see here. Recently I did another reconstruction, also by directly reconstructing from a photograph. This is the result: 

You can see the typical morphology of the aurochs very clearly. The trunk is much shorter, the legs longer and the skull larger than in domestic cattle. The body is, as suggested by the skeleton, very muscular and athletic. The shoulder hump is very large. Of modern cattle, only some Lidia (Spanish fighting) bulls come close to this morphology. Interestingly, the horns of the Prejlerup bull are quite the opposite of that of the Vig specimen, despite being from the same population. The horns of the Vig bull are comparably upright and not that strongly curved, while the horns of the Prejlerup bull are forwards-curving and with a strong inwards curve. 

Looking at the reconstructed morphology of the bull, it is very plausible that it was a very swift and agile animal, as historic written sources suggest. 

Ancient Europe's landscape: grassland savannah or forest?

There is an on-going controversy on what Europe’s original landscape, untouched by human influence, was like: either heavily forested, a grassland savannah or a mix of open forests, park-like landscapes and grassland. 
The traditional view is that of Europe being a heavily forested continent. This view has been challenged in recent decades. It has been proposed that herbivores prevent open landscapes from becoming forested by damaging forest growth with their feeding, and are even able to turn forests into open landscapes this way. As a result, Europe’s original landscape would not have been one big forest but a mix of open landscapes, park-like landscapes and forests [1]. Some even claim Europe was a large grassland savannah. 
Indeed no large mammal in Europe is dependent on forests while they need at least some open landscape in their habitat [2,3]. Africa is used as an analogue, where large herbivores are claimed to create the open landscape we are familiar with. Especially elephants, which uproot trees are suspected to create open landscapes [3]. 
There is conflicting data, however. Studies suggest that the uprooting of trees by elephants does not diminish the forest but instead speeds up the forest rejuvenation [4]. And it is overlooked that not only the savannah but also the deep rainforests are home to large herbivores, such as the forest elephant (Loxodonta cyclotis), the forest buffalo (Syncerus nanus) and the okapi (Okapia johnstoni) [4]. There are also several species of large herbivores living in the rain forests of South-East Asia (banteng, gaur, Asian elephant). If large herbivores create open landscapes, elephants in particular, the forest elephant would be a contradiction in itself: a forest-adapted elephant (f.e. the smaller size) would not exist if elephants created open landscapes with their feeding and uprooting of trees. 
Furthermore, and more importantly, palynologic data suggests that Europe was densely forested until very recent millennia when man started agriculture in Europe [4]. Also the insect fauna shows that forest-dwelling species were very common in Europe until recently [4]. It is argued, however, that also an intensely grazed grassland area has the same palynologic signature as a closed forest (see for example the works of Frans Vera). But this apparently is only the case at a very high herbivore density [4], and it cannot explain why the data from insects suggest high forestation. 
Another argument against ancient Europe being a grassland savannah and for a strongly forested continent is the distribution of the European wild horse in the Holocene. The horse is heavily dependent on a grassy diet and is an open land animal. It got very rare in Europe after the last glacial [5]. It virtually disappeared from Central Europe [5]. When agriculture began, the equine remains increased again, possibly due to an increase in open landscapes [5]. These distribution patterns contradict the hypothesis that Europe would have been a grassland savannah due to grazing and instead supports the hypothesis of Europe without human influence being a mostly forested continent. 
 
Europe’s original landscape continues to be a subject of debate. The data that is available to me suggests that the idea of ancient Europe being a grassland savannah with large herds of horses is not the likeliest scenario. 
 
Literature 
 
[1] Bunzel-Drüke et al.: Der Einfluss von Großherbivoren auf die Naturlandschaft  Mitteleuropas. 2001. 
[2] Beutler: Die Großtierfauna Europas und ihr Einfluss auf Vegetation und Landschaft. 1996.
[3] Bunzel-Drüke et al.: Überlegungen zu Wald, Mensch und Megafauna. 1994.  
[4] van Vuure: Retracing the aurochs – history, morphology and ecology of an extinct wild ox. 2005. 
[5] Sommer et al.: Holocene survival of the wild horse in Europe – a matter of open landscape? 2010.
 

Life reconstruction of the Vig aurochs

The Vig skeleton is one of two more or less complete aurochs skeletons found on the island Zealand in Denmark. It is remarkable for its size (described as nearly two meters in the literature [1]) and for the fact that it shows several damages from arrows. It was even found with two arrowheads. Probably the individual was wounded by several arrows, fled into a marsh and died there. It is now exhibited in the National museum of Denmark. 

Recently I did a life reconstruction based on a photo I was sent by Markus Bühler. The result is down below. 

The Vig specimen at the National museum of Denmark, © Markus Bühler

Life reconstruction of the Vig bull

I did the life reconstruction by drawing over the skeleton, so it should be as accurate as possible. As it was most likely a male, I reconstructed it as a bull. The eel stripe is grey, as historic references such as von Herberstein or Anton Schneeberger suggest. 

The Vig bull has comparably upright and weakly curved horns. The horns are not really typical for an aurochs bull. The horns of the Prejlerup aurochs, which is the second complete skeleton from Zealand, is quite the contrary: it has smaller, forwards-facing and inwards-curving horns. Some of the better Heck cattle bulls have horns reminiscent of that of the Vig bull. 

[1] Frisch: Der Auerochs - das europäische Rind. 2010. 

Three new videos of Taurus cattle from the Lippeaue

Recently the youtube channel Auerochsen Weidelandschaften has released some new videos. It shows the breeding bull Darth Vader III (which is 1/32 Lidia) with his cows at Disselmersch in the Lippeaue reserve. Here they are: 

I think the bull and also the cows are very good. Honestly I believe that the Lippeaue animals already come close to the top level of aurochs-likeness that is achievable with modern domestic cattle. 

New photos from the Auerrind project

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

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

Maremmana x Watussi (© Claus Kropp)

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

Maremmana x Sayaguesa (©Claus Kropp)

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

The Holocene European wild horse

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

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

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

Why do some Polish koniks have erect manes?

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

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

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

Probably only a genetic test could bring clarification. 

Genetic breeding-back?

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

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

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

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

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

Leptobos and the origin of Bos and Bison

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

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

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

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

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

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

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

A zebuine ancestry for Chianina?

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

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

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

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

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

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

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

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

Three more Tauros bulls

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

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

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

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

K. L. Hartig's aurochs painting

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

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

Here is the result: 

Literature 

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

The Tauros cattle of Kettingdijk, Netherlands

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

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

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

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