Articles on the aurochs and the European wild horse

About a month ago, I was invited to write some guest articles for the website www.theextinctions.com, a website on recently extinct animals. I wrote an article on the aurochs, and one for the European wild horse (which I do not call "tarpan"). 

Here are the articles: 

The aurochs 

The European wild horse 

With the articles I intended to do a comprehensive summary on what we know about these two animals. Long-term readers of my blog will be familiar with the knowledge presented in the articles. 

Video of Hungarian Taurus cattle

Recently I found a video of Hungarian Taurus cattle in the Hortobagy National Park: 

It shows bulls as well as steers. They seem to be strongly influenced by Heck cattle, probably the breeding bull Anno from the Wörth breeding line (hence the large horns), perhaps also Watussi. 

The ecologic niche of the aurochs

The ecologic niche of the aurochs and where it differed from that of the wisent is hard to ascertain as the aurochs is extinct. Right? Not exactly. There actually a lot of things on the ecology of the aurochs and its interplay with other European herbivores that we can infer from various sources. 

The aurochs and other European herbivores of the Holocene

1. The aurochs was a grazer and probably much like cattle ecologically

 

The aurochs had the hypsodont denture of a grazer, like cattle today [1]. Anton Schneeberger writes that aurochs fed on acorns, mown grain and hay they were provided, and on branches of shrubs and trees during winter [2]. It has been reported that aurochs grazed in the same places as cattle and horses [2]. Since cattle descend from aurochs, and domestication probably did influence the basic diet in cattle [2], it can be assumed that aurochs and cattle were very much alike in food choice. There are differences within cattle, but that is mainly between highly derived breeds and landraces, since the former do not consume plants that are less digestible while the latter do [3]. 

Isotope analyses suggest that aurochs were found in more forested and wetter habitats than cattle, which mainly grazed on pastures [1]. It is of course possible that this is a difference between the wildtype and the domestic form, but it is more likely that the aurochs was pushed into these less accessible regions because open habitats were reserved as pastures for domestic cattle. Also, free-roaming cattle have been found to prefer wet habitats as well [5]. The continuous habitat loss was one of the driving factors for the aurochs’ extinction [2].  

 

2. The aurochs preferred lowlands and wetter habitat than the wisent 

 

The distribution of fossil and subfossil remains of aurochs and wisent have shown that aurochs preferred plain habitat and lowlands while wisent were found much more often in mountainous habitat [4]. This is also in line with what is known of the historic distribution of both species [2]. While it is true that both the wisent and the aurochs have been limited to hideaway regions due to habitat loss because of the human civilization and hunting, it probably has a reason why the hideaway regions of aurochs were marshes and lowlands and that of the wisent was mountainous habitat, and this region were the ecologic adaptions of the respective species. The wisent avoids areas which are too wet [4], which is why a replacement of the cattle at Oostvaardersplassen with wisent, as it has been discussed on occasion, is not feasible (Bunzel-Drüke, pers. comm.)

Nevertheless, both bovines must have met each other [2]. 

 

3. There was niche partitioning among Europe’s large herbivores 

 

According to the competitive exclusion principle in ecology, two or more species cannot exist in one and the same ecologic niche. Therefore, there must have been niche partitioning among Europe’s large herbivores and apparently this was the case. Cattle consume more wooden material than horses, which are strict grazers. However, cattle do not include as much wooden material in their diet as the wisent, which is a so-called semi-intermediate feeder. So there is a graduation on how much the respective herbivore relies on either grasses or wooden material, with horses relying the most on grasses and wisents consuming the most wooden material, and cattle being intermediate [4]. Since this niche partitioning is the result of evolutionary adaptions of these species, it is likely that this would also be the niche partitioning between wild horse, aurochs and wisent.  

Therefore, considering the differing habitat and food preferences, cattle and wisent would concur only to a limited extent in Europe’s nature systems. So the fear that wisent and cattle cannot be kept in the same reserve is baseless as long as the reserve provides habitats for both species. Another reason why some people have objections against (re)introducing cattle into European nature systems is that they are concerned that wisent and cattle would hybridize. However, a work by Frans Vera has concluded that cattle and wisent do not hybridize spontaneously and only under artificial conditions, which also often lead to the assault of the domestic individual by the wisent [6]. Furthermore, parturition does not occur in those hybrid pregnancies [6], what makes human assistance necessary. Thus, cattle and wisent would not produce hybrid populations in the wild even if mating between both species would occur. 

 

 

[1] Lynch et al.: Where the wild things are: aurochs and cattle in England. 2008. 

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

[3] Poettinger, J.: Vergleichende Studie zur Haltung und zum Verhalten des Wisents und des Heckrinds. 2011. 

[4] Bunzel-Drüke, Finck, Kämmer, Luick, Reisinger, Riecken, Riedl, Scharf & Zimball: „Wilde Weiden: Praxisleitfaden für Ganzjahresbeweidung in Naturschutz und Landschaftsentwicklung“. 2011

[5] Gander et al.: Habitat use of Scottish Highland cattle in a lakeshore wetland. 2003. 

[6] Vera: Do European bison and domestic cattle hybridize spontaneously? 2002. 

 

 

 

European wild horse colour phenotypes

Back in 2013, I did an illustration of the five colour phenotypes found in European wild horses according to genetic research and presented it in a post (back in the time when I called the European wild horse "tarpan", which I nowadays do not). I also put it on Wikipedia, but it was removed by some know-it-alls editing the German page. 

I recently did an updated version of that illustration. Back in 2013 I considered it likely that the European wild horse had a falling mane, which I no longer do (see here). I corrected it, and here is the result: 

It shows all the colour phenotypes confirmed for Equus ferus ferus so far, including: bay dun (top left), bay + nondun1 (top right), black dun (middle left), black + nondun1 (middle right), leopard spotted (bottom). For the references for the colour phenotypes see [1,2]. Black and black dun are caused by the allele a, which became the dominant allele later in the Holocene (whether the phenotype is black or black dun depends on the alleles on the Dun locus) [3], while Pleistocene horses all had the A+ allele, and thus were either bay or bay dun. A 2015 study [4] examined the alleles on the dun locus in a late Pleistocene (42.700 years old) wild horse from Russia and that of a 4000 years old horse (also from Russia). It turned out that the Pleistocene wild horse was heterozygous for dun and nondun1, thus had a dun phenotype. And since all wild horses from that time were A+//A+, it must have been bay dun and thus looked like the drawing at the top left. The 4000 years old horse (whether it was a wild horse or domestic horse was not tested), was homozygous for nondun1. Thus, if it was a wild horse, it looked like the illustration at the top right, since the a allele was not found in Russian samples. 

I also gave the leopard spotted wild horse a dark neck and face, matching the cave art (see here). 

Literature 

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

 [2] Ludwig et al.: Coat color variation at the beginning of horse domestication. 2009.

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

[4] Imsland et al.: Regulatory mutations in TBX3disrupt asymmetric hair pigmentation that underlies Dun camouflage colour in horses. 2015.

"Breeding-back" with horses?

When talking about "breeding-back", mostly the aurochs comes to mind. Currently, there is no "breeding-back" project for horses. However, reconstructing the life appearance of European wild horses could be worth a try. 

There have been such projects in the past. The most widely known "breeding-back" project for horses was that of Lutz Heck in Berlin, Germany, in the 1930s and 1940s. He used a Przewalski's stallion to cover mares of the Dülmen pony, Icelandic horse and Gotland pony, and subsequently mixed the results with Koniks. The resulting Heck horse has been repeatedly crossed with Koniks after the Second World War so that it is virtually indistinguishable from this breed today [1]. The Konik itself is not the result of a "breeding-back" project, but a landrace (see here) instead. 

A Heck horse mare in the Tierpark Hellabrunn. The Heck horse is a well-known "breeding-back" result with horses. 

Another "breeding-back" project for horses focused on Koniks exclusively. It was that of Tadeusz Vetulani, started in the 1920s. He thought that keeping Koniks in a semi-feral state would re-develop what he considered wildtype traits, such as an upright mane and a white winter coat [2]. Actually, this was more of a dedomestication project rather than selective breeding. 

From 1960 onwards, Konik and Fjord horses have been crossed in the Bayerischer Wald, Germany, in order to achieve a more wild horse-like exterieur (or what the breeders considered wild horse-like). The result is called the Liebenthaler horse [1]. 

In the 1970s, the Wildpark Hardehausen crossed Heck horses with Przewalski's horses again on order to achieve an upright mane [1]. 

In the Lippeaue, Germany, several Przewalski-Konik crosses can be found. This is not really a "breeding-back" project - a Przewalski's mare was included in a Konik herd, and now there are Przewalski x Konik as well as 75% Konik 25% Przewalski's horses in the herd, and possibly even younger offspring. See here for my post on these horses.  

A Przewalksi x Konik mare in the Lippeaue

How about a new "breeding-back" project for European wild horses?

So far, there is no breed that matches European wild horses in "phenotype". Admittedly, the reconstruction of the life appearance of these horses is speculative, but a few things can be inferred. Holocene European wild horses probably were predominantly black at a certain time of their evolution, as genetic studies suggest (see here). Whether their mane was erect as in extant wild equines or falling as in domestic horses is unknown as there are no specimen with soft tissue preserved and no historic descriptions that unquestionably describe European wild horses. But it is not unlikely that they had an erect mane (see here). Concerning the body and its proportions, there is not much known as there are no articulated skeletons that have unambiguously been referred to wild horses in Europe. But in general, a "pony type" with a robust head is assumed for the European wild horse. 

A reconstruction of the Holocene wild horse as suggested by recent evidence (the mane is still speculative). 

With that in mind, one can dream of a "breeding-back" project for horses with the modern information we have. The breeding goal would be a black pony-like horse with an upright mane. For that, either Przewalski's horses or some of the hybrids of the Lippeaue (as long as they have an upright mane) could be used and crossed with primitive black ponies such as the Asturcon from Spain or black individuals of the Bosnian Mountain pony. Since introgression from the Przewalski's horse has been found in a European wild horse [3], using Przewalski's horses definitely would do no harm. The resulting horse breed can be used as an alternative to Koniks and other horses that are currently used in rewilding projects. 

Literature

 

[1] Bunzel-Drüke, Finck, Kämmer, Luick, Reisinger, Riecken, Riedl, Scharf & Zimball: „Wilde Weiden: Praxisleitfaden für Ganzjahresbeweidung in Naturschutz und Landschaftsentwicklung“. 2011 

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

[3] Wutke et al.: Decline of genetic diversity in ancient domestic stallions in Europe. 2018. 

European wild horse mane: erect or falling?

The Holocene European wild horse is enigmatic. It seemingly was not common in the probably predominantly forested Europe of the Holocene [1] and it is not known when it actually died out, as the numerous accounts of free-ranging horse populations in Europe during the last two millennia might just as well describe feral horses, or hybrids. Consequently, its life appearance leaves room for speculation. For a take on this subject based on recent genetic data, see this post. 

The most puzzling question concerning the life appearance of the European wild horse is the mane. For a pretty long time, the falling mane of the domestic horse has been considered a result of domestication, as all extant wild equines have an erect mane. In domestic donkeys, domestication also resulted in a falling mane in some breeds (see here, for example). 

It has been claimed in recent years that European wild horses or the wild progenitors of the domestic horse might indeed have had a falling mane. The falling mane allegedly helps to divert rain from neck and face in a more humid environment, and indeed all extant wild equines are found in arid ecosystems. However, it has to be kept in mind that the horse was domesticated in the arid steppe, therefore this is not a compelling argument for the western wild horse having a falling mane. Another argument is that an Equus lambei carcass found in North America allegedly shows a falling mane (I have been unable to find quality photos of the carcass, unfortunately). It has to be kept in mind though that Equus lambei, the North American form, is probably more distantly related to the domestic horse than Przewalski's horses, or Siberian Pleistocene horses, which have/had short erect manes of about 15 cm length [2]. Therefore, the falling mane would have to have evolved at least two times in wild equines and such a mane cannot be inferred by phylogenetic bracketing for European/western wild horses. 

Then there are the manifold Pleistocene cave paintings which all show horses with an erect mane very clearly (see here and here, for example). Since many of the depicted horses show a strong resemblance to the Przewalski's horse, it has been assumed that the horses present in Europe during the late Pleistocene must have been of the przewalskii lineage. However, it has been found that a horse remain from Belgium from approximately 36.000 years ago is genetically closer to Iberian wild horses and domestic horses than to Przewalski's horses [3]. It is thus assumed that these short-maned horses are the ancestors of European wild horses (which the authors of Fages et al. call "tarpans") [3]. Interestingly, Siberian Pleistocene wild horses (referred to as Equus lenensis), which were short-maned to according to a carcass [2], are actually closer to domestic horses than Holocene Iberian wild horses. And the Przewalski's horse, which has an upright mane as well, is closer to the domestic horse than both the Holocene Iberian and Pleistocene Siberian wild horse according to Fages et al. [3]. 

Therefore, it seems highly likely that at least Holocene Iberian wild horses had an upright mane, and probably the (other) ancestors of domestic horses in the Eurasian steppe as well. 

Literature: 

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

[2] Lazarev, P.: Large mammals of the Anthropogene of Yakutia. 2005. 

[3] Fages et al.: Tracking five millennia of horse managment with extensive ancient genome time series. 2019. 

The Taurus cow Lerida

Lerida, a Heck x Sayaguesa, in young years © Matthias Scharf

Today I want to cover one of the very good Taurus cattle individuals from the Lippeaue named Lerida. This cow is the result of the cross of the Heck bull Lancelot and the Sayaguesa cow Dona-Urraca, hence Heck x Sayaguesa. With this cross, a quite good resemblance to the European aurochs was achieved in the first generation of Taurus cattle breeding already. 

Her withers height was not measured yet, but she might be of the same size as the average for Taurus cows, that is 150-155 cm at the withers. Her colour fits a European aurochs cow perfectly, and also her horns are of an aurochs-like curvature. They would need to be a little bit bigger to fit European aurochs cows. The proportions and body shape are OK. The trunk needs to be slightly shorter (the trunk length of aurochs cows was slightly less than 100% of the withers height, see the Sassenberg aurochs cow). Also, the hump would need to be bigger and the udder smaller. If those "flaws" would be fixed, Lerida would be a perfect or near-perfect optic/morphologic copy of a European aurochs cow. This is why she has always been one of my favourite Taurus cows. Her behaviour is unproblematic. 

She left a number of descendants in the herd. One of them was Lippe, a F2 Heck x Sayaguesa, which happened to be rather small horned,  or 84 037, which inherited the hanging spine of Sayaguesa (the father was Lombriz, a Sayaguesa x (Heck x Chianina) bull). Another daughter of Lerida was the very good cow Lisette, which was half Heck x Lidia. 

Here are two photos more: 

Lerida a little bit older © Matthias Scharf

Photographed in the Lippeaue 2013, all rights reserved. 

A Sayaguesa x Chianina with excellent body shape

Recently another photo of one of the Sayaguesa x Chianina cows was published on Facebook by Claus Kropp: 

© Claus Kropp

The body shape of the cow looks excellent. The waist is slender and the hump is large. As it is not in profile view, I cannot judge if the trunk should be shorter (it was quite short in aurochs cows), but the legs are long nevertheless. Something intermediate between this and a good Lidia cow would be a perfect aurochs cow-lookalike in terms of body shape. Horn shape and size is not there yet, and the coat shows the dilution factor of Chianina, but this cow is a very good basis for further breeding. 

© Claus Kropp

This guy will cover her, or maybe has already covered her. It is Alvarez the Sayaguesa x Watussi bull. I am really looking forward to this offspring. It will be 50% Sayaguesa, 25% Chianina and 25% Watussi. It might not have a perfect primigenius spiral yet but otherwise has the chance to become very good. 

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.