A new video of Darth Vader III

Recently a new video of Darth Vader III, breeding bull of the Taurus cattle herd at Disselmersch, Lippeaue, has been posted: 

The colour is phenotypically perfect, the body shape is quite good as well, as much as the horns. The trunk should be shorter, however. 

The past 10 years in "breeding-back"

I have been researching on the aurochs, wild horse and “breeding-back” for exactly ten years now. It was in 2011, when I worked on a drawing of Europe’s large mammals that I looked at the Wikipedia page in order to get some knowledge on the aurochs’ physical appearance. I had been aware of “breeding-back” previously, but did not pay much attention to it. The photograph of the Heck bull “Ari” from the Wörth/Steinberg lineage immediately created my fascination for this subject. 

This was in a time when Heck cattle had a kind of monopoly for being an aurochs-like cattle breed, as the primitive landraces from Southern Europe were barely known outside the Iberian peninsular. But I learned of these breeds quite early because I joined the aurochs thread in the old Carnivora Forum, where a lot of photos of primitive Iberian breeds were shared. In the same year, I visited the Hellabrunn Zoo in Munich, which is where modern Heck cattle originated. The Hellabrunn Heck cattle herd is remarkable for its perfectly aurochs-like colour with a good sexual dimorphism. Being aware of the fact that Heck cattle is only one of many aurochs-like breeds, I enjoyed seeing living cattle with the colouration of an actually extinct animal very much. Thanks to the Carnivora thread, I also learned about the Tauros Project (today: Tauros Programme), which had only one crossbred individual at that time that was actually still a calf (the bull Manolo Uno). There also was barely any information on Taurus cattle on the web, with only two PDFs showing first-generation crosses at young age. I had no idea that by that time, animals like the bull Lamarck were thriving in the Lippeaue reserve. So there was Heck cattle, by many considered to be the only aurochs-like breed, Taurus cattle which were pretty much incognito back this time and the Tauros Project which only had one calf. 

I felt that the monopoly of Heck cattle was unjustified, and the Wikipedia articles in German and English were pretty “pro-Heck cattle” back this time, with little well-grounded information and unempirical claims. The Wikipedia article on the aurochs was not satisfying either, with some inaccuracies and no precise information on the aurochs’ physical appearance. And it was extremely difficult to find information and photos of the primitive Southern European breeds. So I started to rewrite the Heck cattle and aurochs articles in both the German and English Wikipedia, mainly using Cis van Vuure’s 2005 work as a reference. Also, I created Wikipedia articles for some of the primitive European aurochs-like landraces, with photos, because I thought this would be the most effective way to make the knowledge on these valuable breeds more accessible to a larger-scale audience. Also, I thought that by precisely describing the aurochs’ physical characteristics on Wikipedia that the differences between Heck cattle and the aurochs would become apparent. 

In 2013, I started the Breeding-back Blog. Back this time, there were not many people interested in the aurochs and “breeding-back” on the web. There was the Carnivora thread, which was mainly attended by four or five users on a regular basis. In the following years, the situation began to change. 

 

For once, “breeding-back” itself began to grow. The Tauros Project expanded to multiple countries and acquired quantity quite fast, and a new project, the Auerrind project came into existence. Taurus cattle finally found their way into the internet, also because – excuse my self-praise here – I went to the Lippeaue reserve several times and covered them on my blog, with many photos (now Taurus cattle also have a very good web presence on the webpage of the ABU). Also, the number of rewilding projects increased, and both practices got considerable media-coverage (compared to the years before 2011). And the number of people interested in “breeding-back” and the aurochs and also of course rewilding multiplied and multiplied. 

I see that every time I look at my blog’s statistics. My blog has a total of more than 900.000 clicks, with 200-400 clicks per day, some posts have over 1000 clicks and it has 2000 comments in sum (a big thanks to all my readers by the way!). I would have never expected this to happen, as “breeding-back” and the aurochs is a very small niche, so to say. But the number of people interested in this subject is constantly increasing. I see a lot of people who are not only interested in the aurochs, but also know a lot about this subject. They precisely know what an aurochs looked like, know all the projects and special herds/lineages in great detail. Also, the primitive aurochs-like breeds are now well-known to a lot of people. Back in 2011, all of this was not the case. 

I think that of all the animals killed off by mankind in historical times, the aurochs is perhaps the species that currently gets the most attention. The Thylacine also has a huge fan club (a fan club that also has a longer history), but I get the feeling that the “aurochs fan community” got ahead of that of the Thylacine. The mere fact that there is such a community now is worth noticing, because it did not exist back in 2011. 

 

Why did that change so dramatically? I think this has two main reasons: a) there are now more “breeding-back” projects than back then, with a lot more animals and also media-coverage b) the information on “breeding-back” and the aurochs is now easily accessible. The Wikipedia articles are pretty good (I just initiated the changes, a lot of people have helped to progress the pages since 2011), you can easily learn about the life appearance and biology of the aurochs, learn about the primitive aurochs-like breeds and learn about the “breeding-back” projects on the web. I hope that my blog contributes to that as well. 

 

The fact that people are increasingly aware of the anatomy and biology of the aurochs also has a kind of positive feedback effect on “breeding-back”. I get the impression that Heck cattle has slowly increased in quality during the last 10 years because breeders are increasingly aware of the anatomy of the aurochs. The slow process of improvement is not true for all Heck herds, but for some. Also, Heck cattle lost its monopoly on being “the” aurochs-like breed. This was necessary to seize the potential we have in modern day cattle. 

While the myth that Heck cattle are “recreated aurochs” is a thing of the past now, the acceptance of the myths concerning the European wild horse and its purported descendants has not changed yet. Still a lot of people believe the “the Konik is the Tarpan descendant” myth, the Sorraia myth and the Exmoor pony myth (go here for a review of these stories). These stories sometimes are even repeated in technical, peer-reviewed papers, which is concerning. But maybe this might change in the future, as fact-based information on these three breeds and on the European wild horse are at least available in the internet now. 

 

I think that we live in the prime time of “breeding-back” today. There are several projects, people now are aware of the anatomy of the aurochs and of suitable breeds (which was not the case, f.e., when the Heck brothers did their breeding experiments) and they have more attention than ever since there is a large number of people interested in this subject. “Breeding-back” already has produced a number of great animals, and there will be much more in the future. I am looking forward to that, and I am also looking forward to cover them on my blog as I always try to do. 

 

Differences between the quagga and other zebras

It has been established that the quagga is a subspecies of the plains zebra because it clusters genetically within this clade. This, however, does not imply that it does not differ considerably from other zebra species or subspecies. It diverged from the other plains zebra subspecies between 290 000 or 120 000 years ago [1], what is considerably longer ago than the differentiation between the lineage leading to domestic horses and Przewalski’s horse and definitely long enough to develop its own distinct traits. 

The most obvious difference between the quagga and other zebras is the fact that only the anterior part of the body was striped, while the posterior half was stripeless and of a brownish colour. The coat colour, or rather the extent of the striping, varies considerably in the 24 individuals of which the skin was preserved. I illustrated and described this variation in this post. Alas, the locality of the specimen has not been documented, so that the claim that the stripe reduction of the quagga represented a cline cannot be verified for at least within the quagga itself. 

Also, the quagga apparently had an idiosyncratic call (sounding like “kwa-ha-ha”, other variations are documented as well), of which the name of the animal is said to be an onomatopoeic derivation (Wikipedia). EDIT: It is possible that the quagga's call was identical with that of other plains zebras. 

Another very important distinct trait of the quagga is its sexual dimorphism. Preserved quagga skins suggest that the mares were longer and taller and thus larger than the males, which is the opposite of what we see in other zebras, including the plains zebra, where the males are larger than the females [2]. This is unique among extant equines. 

Another possible difference between the quagga and other plains zebra subspecies is the mane length. Looking at photographs of the 24 quagga skins preserved plus the photos of the only living mare to be photographed in the 19^th century, it appears to me that the mane is shorter than in other plains zebras. Of course this has to be verified by actually measuring the mane hair and comparing it to the other subspecies, but it seems rather apparent to me. 

 

My coloured version of the photo of the only quagga to be photographed, drawn with GIMP

Thus, it has to be concluded that the quagga was, despite being nested genetically within the plains zebra species, a distinct type of zebra with unique autapomorphies. This has consequences for the guideline of the Quagga Project. The quagga is evidently more than a colour variant, even though its range might have been continuous with that of other plains zebra subspecies (we find this also in species that form a hybrid zone, such as species within Bombina, Corvus, or Loxodonta to name a few). The similarity between the quagga and the zebras of the Quagga Project will merely be superficial. But, so claims the Quagga Project, the quagga was defined only based on external features, thus making that argument invalid. However, the overwhelming majority of animal (and plant) species on this planet is described on a phenotypic basis, it’s part of the requirements of the ICZN for a valid species description. Furthermore, the differences in the fur colour are not the only defining quagga characters, as outlined above. The zebras of the Quagga Project do not exhibit the idiosyncratic call of the quagga, they do not have the reverse sexual dimorphism and the (possibly) shorter mane. Apart from the phenotype, there are genetic differences as well. The quagga has unique mitochondrial haplotypes and does not share any haplotypes with the plains zebra [1], indicating reproductive isolation for a considerable time span. Thus, the quagga was also genetically distinct and the zebras of the Quagga Project will necessarily differ also genetically. 

Another problem is that the zebras of the quagga project only have a raw similarity to the quagga based on the coat colouration. While the amount of stripe reduction that was achieved is impressive, the animals lack the deep brown base colour on the trunk and also the stripe pattern is different: the white space between the stripes on the face and neck is rather white because the stripes are comparably thin, sometimes with faint brown stripes between the black stripes, while in all the preserved quagga skins the stripes on face and neck are rather broad, with a narrow white space between them and no faint stripes between the solid stripes. Therefore, the only similarity between the zebras of the quagga project is that the stripes are reduced on trunk and legs. Everything else is different. That is why I wrote the article Please don’t call it quagga in 2015, suggesting that these zebras should neither be called “quagga” nor “Rau quagga”. 

Perhaps the quagga project is not even a “breeding-back” project in the strict sense, since it does not work with living descendants of the quagga but merely more or less related animals of a completely different subspecies. 

Nevertheless, I am happy that there is the Quagga Project. While they cannot recreate the quagga, they have produced animals that could be useful for outbreeding a genetically resurrected quagga, for the case that the quagga one day will be recreated by using genome editing. 

 

Literature 

 

[1] Hofreiter et al.: A rapid loss of stripes: the evolutionary history of the extinct quagga. 2005. 

[2] Heywood: Sexual dimorphism of body size in taxidermy specimens of Equus quagga quagga Boddeart (Equidae).2019. 

 

A Pleistocene wild horse from Denmark

A few years ago, I did a life restoration of this wild horse skeleton from Denmark. But it wasn't that good, I found. So I did a new one, and finally had the time to finish it: 

In order to be as precise as possible, I tracked the original horse skeletons out using a pencil and paper. For the reconstruction of the horse, I used genetic information and the horse depictions from the Ekain cave. Genetic information tells us that most Pleistocene wild horses had the bay allele A+ [1], and both the dun allele and the wildtype non-dun (nondun1) allele was present in the population [2]. The horse therefore was either bay or bay dun. I chose bay dun, because it probably was more common as non-dun is recessive. The horse depictions at Ekain tell us that the striping was pretty pronounced, more so than in extant Przewalski's horses and domestic horses. The mane of all Pleistocene wild horses was erect. All in all, the Pleistocene wild horse was rather similar to the Przewalski's horse in appearance, although the head not quite as large, apparently. 

Horses like this one probably were the ancestors of Holocene European wild horses, according to a 2019 study [3]. They their colour shifted from mostly bay dun to mostly black (or maybe additionally black dun) during the Holocene [2]. For a reconstruction of the Holocene wild horse, go here. 

Literature

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

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

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

The skeleton of a Heck bull

When comparing living cattle to the aurochs, there is the danger of comparing apples with bananas when examining skeletons versus living animals. We do not know the aurochs' life appearance for sure, we only know the skeletons for sure. Thus, a fruitful approach could be comparing the skeletons of aurochs-like cattle to that of aurochs. I have seen some Heck bull skulls previously, but never a complete skeleton. Recently, I found the photo of a Heck bull skeleton mounted in France. It is from the publication Remaques sur l'osteometrie de l'aurochs-reconstitue: interet pour la selection de la race (code 30) en vue de se rapprocher du Bos primigenius by Claude Guintard and Quentin Blond. 

© Claude Guintard

Guintard has done a number of publications on Heck cattle, comparing it to the aurochs. The paper in particular looks at this skeleton plus a skull of the Heck bull Axis from the Wörth/Steinberg lineage. 

Go here for a skeleton of an aurochs bull. As you see, the trunk of the Heck bull is longer. If the trunk was shorter, the legs would be longer in relation to the overall size and would fit the aurochs better. The Heck bull has a hump, but smaller as in the aurochs. Also, the skull is shorter but not much compared to that particular specimen. There are aurochs bulls with much longer snouts (such as the Sassenberg bull), and also comparably short-snouted individuals (such as the Torsac dirac bull skeleton) which are comparable to average domestic cattle in skull shape. 

Besides the differences in proportions, there are also differences in the horn shape. The horns of the Heck bull would have to curve inwards more strongly in order to fit the aurochs. The dimensions of the horns are within the European aurochs' range of variation. 

References 

Claude Guintard, Quentin Blond: Remaques sur l'osteometrie de l'aurochs-reconstitue: interet pour la selection de la race (code 30) en vue de se rapprocher du Bos primigenius. 2018.