Please don't call it quagga

Don’t get me wrong, I am happy that there is something like the Quagga Project, and I like their current breeding results. But they make a number of claims that I simply cannot agree with. The Quagga Project pretends that they are able to – or even did already – recreate the quagga by selective breeding with extant Plains zebras.

They argue that since the quagga nowadays is classified as a subspecies of the Plains zebra it might even be only a colour variety with the only features making it distinct being the coat pattern. Furthermore, they say that since it was not a species on its own, the genes responsible for their colour scheme might still be present in extant Plains zebras just have to be united. And since the quagga displayed individual variation regarding the intensity of their stripe pattern, a rough overlap with the original and not necessarily a total match is required to call their results “quagga”.

“However, the extinct quagga was not a zebra species of its own but one of several subspecies or local forms of the Plains Zebra. This fact makes a big difference - the quagga’s extinction may not be forever!

[The Quagga was]A variety of zebra […]

However, since the coat -pattern characteristics are the only criteria by which the Quagga is identified, re-bred animals that demonstrate these coat-pattern characteristics could justifiably be called quaggas.

[…] Therefore the Quagga and other Plains Zebras belong to the same species and consequently the Quagga should be considered merely a different population (or deme), of the Plains Zebra.

[…]There are a few Zebras which qualify - not as true quaggas, but as Rau Quaggas in the project. From our point of view they are "real" quaggas, but since there has always been the possibility that there might have been other features of the original quagga that we have not selected for (because we do not know what those features, if any, might have been), we have chosen the term "Rau quagga" to describe our recovered phenotype. Rau quagga is the name we chose to apply to animals which meet the criteria of no scorable stripes on the hind body and effectively none on the legs (Darwin recorded occasional stripes on the hocks in some quaggas, so we allow that in the hind legs). More will doubtless come along as time goes by...”

The project claims that the quagga was merely a colour variant of the plains zebra as they see no differences between the existing subspecies other than the colour. However, there never was an osteologic study comparing the morphological similarity between the plains zebra subspecies including the quagga. There were craniometric measurements with conflicting results (I assume that the authors of these studies took care of whether the skulls were from quaggas indeed or from horses and donkeys for taxidermies) – some studies suggested that the skulls of the quagga were nearly identical to that of other plains zebras, while others suggested they differed from those of horses, donkeys and zebras to the same extent. I learned from a skull gallery by Vera Eisenmann that there is quite some variation regarding skull shape within an equine species, so that factor might not be that important.

As to genetics, the quagga is nested within the plains zebra. That means, it is either a member of the species, or, if you consider it a separate species, descended from the plains zebra at least. It seems that the quagga diverged from the other subspecies in the middle Pleistocene [1], which is about the same time distance as between domestic horses and Przewalski’s horses. The quagga does not share any mitochondrial haplotypes with other plains zebras [1], but the sample size is probably too small to ascertain whether it indicates that it was a reproductively isolated clade on its own or did indeed represent the end of a cline as the coat patterns within the species suggest.

Nevertheless, I think that the divergence of about 200.000 years is too long to regard the quagga merely as a colour variant. Genetic drift inevitably leads to changes. Apart from that, you could argue in the same way that the hooded crow is “just a colour variant” – we have exactly the same situation here. Furthermore, the sparce molecular data does not give us any clue on the karyotype of the quagga – it might have one or more chromosomes more or less than other plains zebras, as it is the case in the Przewalski’s horse compared with the domestic horse.

We cannot say whether the quagga had any ecological differences to other plains zebras, but I don’t think it was necessarily the case. Not much is reported on their behaviour, apart from that they had an idiosyncratic call was an inspiration for the animal’s name (f.e. “kwahaa”), remotely resembling a donkey’s. 

My belief is that there is also a certain misunderstanding of the concept of a subspecies when they say: “the extinct quagga was not a zebra species of its own but one of several subspecies or local forms of the Plains Zebra. This fact makes a big difference”. A subspecies is, as the term implies, a less-marked form of a species, or as Darwin called it, an incoming species. Everyone who has basic biological knowledge will know that giving a clear, unambiguous definition of a species and therefore also a subspecies is highly problematic. But what can be said is that a subspecies is not just a result of taxonomical boredom or nit-picking, but an actual clade and defined by biologic criteria, and this is clearly the case in the quagga. As outlined above, there are more differences between the quagga and the remaining subspecies than pelage characteristics, such as genetic distinctions that inevitably must have arisen during about 200.000 years of separation, and its distinct call. Currently there is not evidence for more as far as I know (as if it was not enough). But the project argues that if there is no further evidence, no further defining characters should be assumed and therefore something that shows all traits that we know is automatically a quagga. In my opinion, the assumption that only those distinct features that we know of through our limited access were the only ones is ultimately destined to fail. However, the QP does not deny that the quagga was different (and it was certainly the most distinct of the plains zebra subspecies), but their assumption is that mimicking those traits that were unique to it and eradicating those that differed should lead to a true quagga.

Different genetics and different descent, different animals. No living herd of plains zebras do descend from the quagga in any way. The QP’s point is that all the traits that defined the quagga might still be present in the extant population thanks to the fact that they belonged to the same species and it likely represented one end of a cline. I see problems in this guideline: We have no clue on the factors that control the pelage characteristics in the plains zebra, which is the only trait the QP is focusing on. Is it polygenetic, or regulated by one gene only? Which characters is it connected with? No extant plains zebra has a stripe pattern that is as reduced as in most of the quaggas, not to speak of the brown background colour, so the selective breeding program is not working on a fixation of a trait but on trying to achieve it in the first place from what is not there yet. It’s like selecting for cattle with 100 cm horn length when starting with horn lengths of 50 and 70 cm. But if I interpret the QP’s argumentation correctly, they assume the amount of striping in a zebra’s coat is regulated by a number of genes, and that those alleles that worked in the quagga are split up among those we see among other subspecies, present somewhere towards the middle of the cline.  That’s a legit hypothesis, although a speculative one.

This provides a good opportunity to test the guideline of the QP. One could take some strongly striped zebras from Etosha and select them for a stronger stripe pattern. Would anybody claim that, if after some decades of breeding such a herd is as strongly striped as possible, a Grant’s zebra (E. q. boehmi) comes out? Or taking wolves, would picking a bunch of white or crème-coloured Northwestern wolves (C. l. occidentalis) result in a polar wolf? I think these examples show how simplified this concept is.  

However, let’s take the QP’s argumentation for granted for a moment and say that a Plains zebra that looks like the quagga automatically is a quagga, and that there are no external differences other than the pelage characteristics. Do the most advanced current results of the project look like quaggas in this respect?  

Judging from the remaining quagga skins and those individuals of the QP that I know, it is true that indeed a number of those zebras show a stripe pattern that is as reduced as in the quagga (apart from the legs, which are not yet totally stripeless). However, I noticed a not negligible difference regarding the stripes on head and neck. In the quagga, they are packed closely together, being broad while the white space between them is thin. And there are no thin stripes between broad stripes as we see it in many extant zebras. In the zebras of the QP, the exact opposite is usually the case. I even get the impression that as stronger reduced the striping on the rear is, the more it is on the face as well. The facial stripes in the quagga on the other hand were very broad and intense, giving the head a dark aspect overall.

The QP admits that there has been only little progress in achieving a brown background colour on the trunk. This might change in future generations, but maybe not. Maybe the last few stripes on the leg will disappear, but maybe not. Perhaps the climax is reached already, only future will tell. But I see no trend towards the quagga regarding the neck and facial stripes. And it brings us to an empirical problem: when is the breeding program progressed enough to say: “ok, this and that trait have not been achieved”? The predictions made have to be falsifiable, otherwise they are of no empiric value. No such timeline has been given, and apart from that, how stable have the characteristics to be in a “recreated quagga population”?

So my conclusion is: Their claim that the quagga was merely a colour variant of “the” plains zebra is likely to be erroneous, their assumption on the genetic background of the stripe pattern is speculative, their guideline that one subspecies can be turned into another by selective breeding on external traits is simplified and probably wrong as my examples have shown, the resemblance of their current results to the quagga are not satisfying yet (contrary to what the QP says) and the resemblance to the quagga is only limitedly falsifiable following their concept.

Perhaps the reason for them considering their results recreated quaggas is just for public relations, but I fear that it is the same kind of self-delusion as it was the case with the Heck brothers. They officially call them Rau quaggas to appreciate that there “might” have been further differences, just as some people call Heck cattle a “model” of the aurochs. But this is not legitimate in my view: There can be no man-made alternative versions of wild animals by definition. Heck cattle is not a model of true wild-type aurochs (not more than any domestic cattle breed is), and the zebras of the QP are not more of a quagga than other Burchell’s zebras are, despite some resemblance in the coat pattern.

This is why I call those zebras “Rau zebras” instead of “Rau quaggas”, as you might have already noticed.

Literature 

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

Mammoth DNA inserted into elephant cells, and function normally

The tempting idea of cloning a woolly mammoth, Mammuthus primigenius, inspired by some exceptionally well preserved specimens in the arctic permafrost, is always causing a lot of media attention. Rumors are making the round, of alleged break-throughs, claims that it is all a hoax and scientists that are either very confident or very skeptical on cloning this magnificant and iconic elephant that was no more ancient than the extant three species.

Several ways have been proposed on how it could be done. For example, inseminating an Asian elephant cow with a reconstructed mammoth sperm and subsequent absorptive breeding. A more effective and modern idea is the CRISPR method that is favourised by a number of scientists today. To put it simply, CRISPR is about cutting (splicing) a DNA strand (in this case, an Asian elephant's) at the loci where it differs from the template (mammoth) and to exchange the original base pairs with the ancient ones to create an ever increasingly mammoth-like functional DNA strand. 
Renomed geneticist George M. Church and his lab at the Harvard University are involved in a project that tries to genetically reconstruct a mammoth on long-term sight this way. Now they managed to splice ancient mammoth genes into the genome of an Asian elephant and the ancient genes did indeed show normal function in the A. elephant cells. Allegedly these genes are involved in typical mammoth characteristics, such as subcutaneous fat, small ears and hair growth, but I don't know how reliable that claim is. The results have not been published in a peer-reviewed paper yet because there is more work to do, Church says.


Something similar has already been achieved with a gene of the Thylacine responsible for cartilage formation. Go here for the paper.

Does this bring us closer to seeing a living woolly mammoth again? Not necessarily. But at least it has been shown that it is possible to insert some mammoth genes into the genome of an extant elephant and to have them working normally. Of course it will be possible to create a full mammoth genome this way, as long it is fully resolved, but there are still practical issues, such as using a female elephant as a surrogate and perhaps also epigenetics. We have to be patient. 

Further read: 
http://www.telegraph.co.uk/news/science/science-news/11488404/Woolly-mammoth-could-roam-again-as-extinct-DNA-merged-with-elephant.html
http://www.iflscience.com/plants-and-animals/scientists-successfully-insert-woolly-mammoth-dna-elephant-genome
http://www.popsci.com/woolly-mammoth-dna-brought-life-elephant-cells
http://www.popsci.com/scitech/article/2009-06/shark-factory

For more article on cloning extinct animals, go here: 

• Clong as a chance for the Wisent? 
• Extinct species/subspecies that could be revived through cloning 
• Complete genome of an aurochs sequenced 

Cloning as a chance for the Wisent

As everybody should know, the extremely low diversity of the contemporary gene pool of the Wisent after the severe bottleneck event during the 1920s and 30s is the most immediate danger for the species’ long-term existence. In this post I outlined how the high degree of inbreeding affects the health and fertility of the global population. I proposed careful, controlled introgression of the American bison as a probable way to add more genetic diversity and resistance to diseases without affecting looks, behaviour and ecology of the Wisent too much, documented in an own breeding book.

When writing my post on extinct species that might one day be revived throughcloning, I came up with another idea helping the Wisent to get out of its genetic misère.   

A well-preserved bone from the early Holocene made it possible to fully sequence the genome of a 9,000 years old aurochs bull. If this is possible, it must be feasible to do the same with the genome of an ancient Wisent. There must be plenty of well-preserved Wisent bones or even soft tissues from early Holocene to the 19^th century onwards. Turf remains for example. Even more promising might be remains from historic times, such as hunting trophies in form of skulls and skins.

Once a full genome is recovered, either a complete set of chromosomes could be reconstructed (for which, as far as I know, the technique has not been developed yet), or the genome of a living Wisent could be used as a template and edited according to the ancient nucleotide sequence by genome editing. The latter method should be easier and more feasible. I think that there is a good chance to recover the whole genome of not only one but several ancient Wisents. Acquiring a surrogate would be no problem of course. Any specimen that lived prior to the bottleneck event would be a precious gain of diversity, and five individuals or so might even multiply it. You might be wondering how a small group of Wisent should distribute their genetic material on the whole global population. But one and the same individual can be cloned several times. Cloning as many as possible individuals, both bulls and cows, and adding them to herds in various regions. But adding only bulls, or replacing as much inbreed bulls with cloned bulls as possible would not be ideal in my opinion. The Y types of the cloned individuals have to be added to the population, but should not replace the old ones.

One of the advantages of cloning pre-bottleneck Wisents over the cloning of extinct species is that people won’t raise those annoying “ethical” non-issues and they will see the good in it more immediately than in cloning aurochs, Quagga and so on.

Even better: if it succeeds, those cloned wisents could serve as flagships for the good in cloning ancient animals that might help to get public acceptance.

Maybe the idea of cloning “ancient” wisent as a genetic long-term solution for the conservation of the wisent sounds unconventional. And yes, I am fully aware of the fact that it would face the same general problems of cloning just as any other project does (although, as far as my knowledge does, the offspring of cloned ancient wisents and modern ones would have the developmental problems clones have to a much lesser extent). But if we are honest, this concept is the only way to considerably increase the genetic diversity of the Wisent and therefore to solve its major threat as a species, without affecting its genetic integrity by crossing-in another species.

If you agree with me, feel free to spread this idea. I really hope that people who have the right connections are going to see this and maybe such a project might be realized in near future.

Newborn calves at Faia Brava, Portugal

Faia Brava is a reserve in Portugal that houses one of the two Iberian Tauros cattle herds. So far, this herd is composed of rather nice-looking Maronesa cows and two Sayaguesa bulls that have been added last year. Now, a few newborn calves have been spotted. It is unclear whether the Sayaguesa bulls are the fathers of the new Tauros cattle yet. 
Go here for the article on Rewilding Europe.
The Maronesa cows have a good build and the horns are ok as well, but their colour is pretty dark. I think that they will have to add a breed that contributes a more definite sexual dichromatism in the future, and they might also need more horn thickness. I don't know how large the bulls and cows are, but I estimate the cross bulls might reach a size of 150 cm or so, but this is just a wild guess.

Feral pigs: a correction

A while ago in my dedomestication series, I wrote that the feral pigs in the South of the USA are a good example for a regression towards the wild type through natural selection.

"Although not identical, they bear a considerable resemblance to wild boars in looks, behaviour and movement. They have a body build for agility and strength, and that's how they move. Their tusks are well-pronounced as they have a social and defensive function. The skull is very elongated, as much as in the wild boar – perhaps this is an example of a “reversal” of paedomorphism as described above through developmental cascades [UPDATE: I was pointed out to a paper that suggests that the elongated snout of feral pigs is a result of phenotypic plasticity due to the chewing mechanism]. What is also striking is their (with a few exceptions) uniform fur colour, beautiful mud-coloured brown or very dark, almost black, brown (not as greyish as in the European wild boar) – very likely camouflage in forested environment."

However, Markus Bühler from the Bestiarium provided me with additional facts that force me to revise my statements a bit. 

The fact that those feral pigs resemble wild boar is actually less surprising when considering which kind of farm pigs they descend. Pigs of former centuries cannot simply be equalled with the typical farm pigs of modern times, because they were less productive, less paedomorph, and overall less derived. Some of these populations even descend from those of early Spanish settlers which brought their pigs from the Iberian peninsular, and as everybody who reads this blog will know that Iberia is a hotspot for primitive landraces today. 
So hypothesizing that the wild boar-like apparence of these feral pigs in the South of the USA is mainly due to dedomestication is like claiming the same for an aurochs-like population of feral cattle descending from primitive cattle landraces. Apart from that, these pigs are not as homogeneously coloured as the videos on youtube suggest. In fact, there is stil a variety of colours present in their gene pool.

Therefore, the dedomestication concept, as logical as it is, looses another empirical example.

First fifth-generation Rau zebra

A little bit delayed news. It was born on 10th of december 2013. See here.
It has a clear brownish tint, but I am not sure if one could say that it is significantly more prominent than in non-selected plains zebras. What is interesting is that the stripes seem to fuse to a dark brown colour on the upper arms in this individual; I have noticed that in several other Rau zebras as well. 
I think I'm going to draw another prediction of how Rau zebras might look like by 2025, additionally to my first one.

Aurochs bull by Joschua Knüppe

There really are not many qualitative, anatomically correct aurochs reconstructions. Most artists make typical mistakes or just do not research properly. For example, there are artworks for which their creators certainly didn't care much on what the horns actually looked like, or which proportions the animal had. There are numerous very good illustration of other extinct bovines, such as extinct bison, Pelorovis, Leptobos and so on. But not even artists like Mauricio Anton, one of the greatest artists for extinct mammals of our time, seem to get the aurochs right. I think the reason for that might be that there are living models for the other taxa (living bison and buffaloes), while domestic cattle are used for the aurochs. Using cattle as models is sensible of course, I do it all the time, but it is a bad mistake to use their body shape for the aurochs. That is a typical pitfall for many artists. Except it is a breed that either lives in the wild or has a body conformation that resembles that of wild bovines anyway (f.e. Lidia). 

As there are only few artists that do qualitative aurochs ilustrations, I asked Joschua Knüppe to draw an aurochs for me. He is a student at te Kunstakademie Münster and a brilliant paleoartist (a paleoartist is, as you probably guessed, someone how illustrates paleontological themes; I consider myself an amateur paleoartist). Go to his deviantart page and have a look at his gallery:  http://hyrotrioskjan.deviantart.com/
Joschua drew an aurochs bull for me after I sent him some photos of aurochs material, life restorations and aurochs-like cattle. I like the result very much: 

Copyright by Joschua Knüppe. Used with permission.

It has all the important aurochs features: the tight, athletic body with a hum, a straight skull and the right colour. The eel stripe is not visible, what may be partly due to the perspective but I consider it possible that some aurochs bulls lacked the dorsal stripe overall anyway. The gentleman next to the aurochs is Ernst Stromer von Reichenbach, an important historic German paleontologist.

Thank you very much!

Some size comparisons

It is interesting how easy we might fail when estimating the size of an animal without appropriate metric reference objects. I experienced it myself when I stroke the Heck bull at the Tierpark Haag and thought that one might be around 150cm tall at the shoulders. I measured him and he turned out to be mere 140cm, the typical size for Heck bulls. It gets even more difficult when we encounter the animal moving in the field, and often they get overestimated. That’s why sometimes crocodiles or komodo dragons are believed to have astounding lengths like eight or six metres, but once shot and measured they turn out to be within the normal size range of their species. But I was surprised that even guessing the size of an animal next to a person on a photo can be rather difficult. I realized that when had a look at this photo of a Heck bull standing next to a man (http://www.morgenweb.de/polopoly_fs/1.2091254.1423066752!/image/image.jpg_gen/derivatives/galerie_940q/image.jpg). The bull looks large and massive, and I estimated it might be well over 145 cm at the shoulders. Then I did calculations assuming that the man next to the bull is 175 and 180cm tall, respectively. And the result was that this bull is 135 cm tall at best.

 I overlaid Wisent and Aurochs at the exact same shoulder height:

The aurochs seems to be the slightly larger animal, which is not surprising since the Wisent is has a short and high body. The largest aurochs bulls therefore were likely heavier than the largest Wisent bulls.

The last one is a comparison of aurochs of three different sizes: 180 cm, 170 cm and 160 cm.

Piebald deer

It was not new to me that the same spotted patterns we see in domestic cattle sometimes also occur in cervids. But until I did a quick google seach I didn't know how widespread it actually is. Spotted patterns can be found in red deer, roe deer, white-tailed deer and elk. At first I thought that these colour variants are the result of incipient domestication. There are deer populations that have been kept in game parks for many generations and it is likely that they have also been selected for tameness to a certain degree, simply for practical reasons. I think it is unlikely that these populations have been totally reproductively isolated, but they were probably isolated enough to develop the typical "novel traits" of domestication: new colour variants (f.e. totally white red deer, or fallow deer with very dark shades), piebald patterns, or shortened skulls (fallow deer). However, the presence of these piebald patterns in elk and white-tailed deer, which have certainly not been kept under the same conditions as those semi-domestic deer populations (at least elk), suggests to me that these traits are not necessarily connected to incipient domestications. Rather it seems that the same mutations that cause piebald patterns in domestic animals also occur "naturally" in wild animals, and sometimes not even rare. For example, there are skandinavian elk populations in which such deviant colour variants are particularly common. Although it has been suggested by scientists involved in the farm fox experiment that such colour mutations are pleiotropically coupled with mutations in the nervous system responsible for tame behaviour, we maybe should not connect such patterns too much with domestication. For example, the African wild dog is piebald in a certain way as well, certain bovids have white "socks" too, and the leopard-like patterns we see in some seals resemble those of some spotted horses as well. 
So perhaps a number of those spotted patterns of domestic animals is not as tightly related to behavioural modifications during domestication as I previously suggested on this blog. I don't know why deer show them particularly often compared to other wild animals.

Now here are some links of piebald deer. 

Elk: 
http://www.smokyriverexpress.com/newsroom/vol45/120314/frontpage.jpg
http://retrieverman.files.wordpress.com/2012/09/leopard-complex-moose.jpg
White-tailed deer: 
http://illaheecommunity.com/wp-content/uploads/2011/06/whtdr372.jpg?w=300
Roe deer: 
http://static1.fnp.de/storage/scl/importe/fnpartikel/rhein-main/limburg-lahn/843440_m3w605h320q75v27933_nps_geschecktes_reh_0912_4c.jpg?version=1418089918
http://bestiarium.kryptozoologie.net/wp-content/uploads/2010/08/PICT5026.jpg

There is even a stuffed hare with a streak along its face and sockings, photographed by Markus Bühler: 
http://bestiarium.kryptozoologie.net/wp-content/uploads/2010/08/PICT5043.jpg

Forelocks and manes

It is well supported that the aurochs had curly, frizzy forelocks on its forehead. It is reported from Anton Schneeberger who wrote: “The forehead, because of the curly, frizzy hair, makes them terrible to behold”. The posthumously (1634) published report by a certain Swiecicki mentions this feature as well. Also, belts were made from a bull aurochs’ forelocks in historic times. They were said to increase the fertility of women or help women having difficulties giving birth (see van Vuure, 2005). As a brutal fact, those forelocks were peeled together with the facial skin from the skull of the captured aurochs when it was still alive. Not to forget, Charles Hamilton Smith’s famouspainting of a bull aurochs from 1836, which is drawn from an oil painting dating back to the 16^th century, shows these curly forelocks very clear.

 

Forelocks of a Hereford bull

Such forelocks are very widespread among domestic cattle. I think their function might be display, apart from their probable protective function for the skin between the horns during combats. Bovids that live in hot regions tend to have fleshy structures for display, mostly dewlaps. Those in cold, northern regions cannot effort the heat lost caused by such appendages and therefore often have hairy display structures; such as the mane of the Barbary sheep or the beards of musk ox and bison. The European aurochs is climatically and geographically in between and funnily so are those structures. Those forelocks give the bulls indeed a fierce look, what could be useful in scaring off rivals – the mealy mouth, which is widespread among Bovidae, might have the same function. While the intensity of the forelocks of taurine cattle varies from breed to breed (Chianina, for example, doesn’t have any at all), zebuine cattle never have such (at least not that I know of). This brought me the idea that it might be a legacy of hybridization with bison. When I then got to know that such hybridizations did take place (see Verkaar et al.) I felt that this thought might have some plausibility in it. Although only introgression from Bos to Bison and not reverse is proven so far, I think it is likely that there was mutual hybridization because only sex-based markers (Y and mt) were used in this study.

Interesting side fact: The forelocks, or actually massive bulk of hair, in Bison is not only used for display and as a bumper during combat, but also to remove snow from the ground in order to reach the grass beneath it. Horses do that with their hooves, while cattle, and probably also aurochs, have no such abilities.

Forelocks in cattle are present in both sexes, but many domestic bulls that have forelocks also have a kind of “mane”. Actually it is not a mane as a lion or a Barbary sheep has, but it is merely the same kind of locks that is present all over the neck, parts of the shoulders and often also the entire face. You find that in a lot of breeds. Some Lidia have it to a very prominent extent (here), and also Chillingham cattle have it. But what is most interesting to me is that a number of Heck bulls in Oostvaardersplassen exhibit such a “mane” (f.e. here), while virtually no Heck bull outside the reserve does. Richard Marsh, the cattle warden of Chillingham, believes that this mane serves to protect the skin on face, neck and shoulders of the bulls. So if Lidia and Chillingham cattle, both breeds in which combats often (in the case of the latter always) decide on reproductive success, show that trait, and if Heck cattle in OVP suddenly developed it, it might indeed have such a function. I know that this is based on a weak ground: man plays a way more important rule in reproductive success of Lidia bulls than combat does, many cattle which are totally man-selected do have that trait too, it might have become coincidentally fixed in Chillingham cattle due to all the bottlenecks, and the “mane” is not all that common among OVP bulls. But, on the other hand, the presence of the forelocks themselves in non-primitive breeds is not a prove against their presence in the aurochs either, and the OVP population has been exposed to natural selection for only 10 generations yet. Anyway, it is just a thought. 

The "mane" of a Chillingham bull

Historic reports don’t say anything about such a mane. But the question is, would it be such an eye-catching feature that it would have been considered to be worth mentioning? On the other hand, if Schneeberger mentioned the forelocks he might have mentioned the mane as well. We can only speculate. C.H. Smith’s aurochs painting clearly does not show curly hair on neck or shoulders. The silhouette drawings at the Lascaux cave however do show agglomerations of dots on head, neck and shoulders. While some interpret it as an indication of Chillingham-coloured aurochs, I consider it more likely that they represent curly hair.

While the curly hair on face, neck and shoulders always has the same colour as the rest of the body, the colour of the forelocks varies in wild type-coloured bulls. The forelocks of cows are almost always of a lighter colour than the rest of the head, with a dark shade “coming from above”. In bulls however, the forelocks can be of an either black, reddish brown, orange or blond colour (in de-phaeomelanised cattle like Podolian cattle, the colour turns grey of course). There is no definite clue on what the colour of the aurochs’ forelocks exactly was. Black is the best-supported colour for the simple fact that forelocks of a colour different from the rest of the head are never mentioned or depicted anywhere – neither by any historic reports nor cave paintings or Smith’s aurochs. If the forelocks had been of a special colour, Schneeberger would have probably mentioned it since he also mentioned the muzzle ring and the eel stripe which contrast with the black base colour. There is also a Libyan petroglyph showing a North African bull aurochs, in which both a muzzle ring and a light colour saddle are indicated, but no bright forelocks. Gaurs, on the other hand, have bright blond hair between the horns and so do zebus. So I assumed bright forelocks were a basal state and dark ones the derived state of northern Aurochs. However, Tom Hammond pointed out to me that bright forelocks could be the result of reduced sexual dichromatism which is the case in Gaur and most of the aurochs-like breeds. And indeed the first breed with a clear sexual dichromatism that comes to my mind has mostly dark forelocks in bulls: Maronesa. But this breed displays all possible colours of forelocks in bulls. Bright forelocks in cows are confirmed through a painting at Lascaux.

Therefore, my opinion on the colour of the forelocks in aurochs is that cows’ always were of a blond, orange or reddish colour while that of the bulls were most likely black in most cases, but perhaps there was geographic variation that allowed the presence of brighter forelocks as well. Although we have no evidence for that, I certainly would not make bright forelocks in bulls a negative selection criterion in effigy breeding.

A number of breeds, especially many Heck cattle, do have elongate hair between their horns but they are not curled. Instead they look like the fringes of a carpet. This is probably not what aurochs forelocks should look like. 

Literature

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

Verkaar, Nijman, Beeke, Hanekamp, Lenstra: Maternal and Paternal Lineages in Cross-breeding bovine species. Has Wisent a Hybrid Origin?. 2004.

A Wörth cow is going to join the herd at Lippeaue!

Ever since I got interested in Taurus cattle, I wished to see a cross with Hecks from the Wörth-Steinberg line. Taurus cattle have the long legs and good proportions, often a satisfying skull length, way better size, better stature and often a hump, forwards facing horns and so on. But they often lack the desirable horn size. Wörth Heck cattle would be a desirable option to boost the horn size, and in a number of individuals the undesirable Heck cattle features like the paedomorphic skull and the elongated, heavy body, are not present to the same extent as in many usual "unimproved" members of the breed. 

Margret Bunzel-Drüke from the ABU told me that they have been interested in buying a Wörth cow indeed, "Erni" is their favorite. Luckily, the owner Walter Frisch announced on the VFA's webpage that he is going to sell three pregnant (!) cows and his current breeding bull (I don't know why since this bull looks very nice, or which bull is to follow). I immediately contacted MBD to inform her about that in the hope that one of the cows will appeal them despite Erni is not among them. 
You can imagine how pleased I was when I received the answer that they are going to buy one of the cows. It will be taken to the Lippeaue in March. 

I am pretty sure that the cow is Nostra, a full-blood sister of Arizona and Aretto (the former breeding bull on Wörth), daughter of Albatros and Noriga, which all have good horns. It would be interesting to know which bull is the father of the calf she is carrying, I hope Aribo (the new breeding bull that is going to be sold now) and not one of those with the beige colour saddle. 
Here are some photos of Nostra that I took in 2013: 

I hope her will be a good cow, because a pure Heck bull is unlikely to be used at the Lippeaue. I am very much looking forward to the results this cow will bring, and I am also curious to which bull's herd she will be moved. 
As a foretaste to what we might enjoy to see within even only one generation, here:

I simply took a photo of Lamarck's fullblood sister at the Lippeaue and enlarged her horns with GIMP. 

I am very happy that crossbreeding between Taurus and Wörth cattle is finally going to happen, and that I played even a little role in it. 

What would aurochs-coloured Chianina look like?

They would look awesome. Colour is one of the most prominent aspects of an animal’s look, and the first one that catches our eyes. I realized might also blur our perception of aurochs-likeness: a cow with aurochs-like colour that otherwise looks like a farm cow (f.e. many Heck cattle) might appear to be more aurochs-like than a Chianina with way more “primitive” proportions, body conformation and size.

That’s why I hesitated to consider Chianina a primitive breed until one and a half year ago or so – I thought the “wrong colour” is a considerable malus. But first of all, since there is not simply one colour locus that decides whether a colour is wild type or not, the colour of a breed is probably unlikely to be mutated on each locus. For example, Highland cattle are either E^d or e, so the production of eumelanin in the coat is either over-expressed or totally disabled, so we cannot say if they have the genetic make up for sexual dichromatism. Only crossbreeding or genome editing would provide a clue. The shiny red colour of some Highlands implies that they would have probably a nice and un-diluted aurochs colour (independent of sexual dichromatism) if they had the E⁺ allele. And so forth.

Chianina has the wild type base colour E+ and very likely also sexual dichromatism masked beneath the dilution factors (see the previous post).

Furthermore, and what is way more important, colour is regulated only by a few loci which follow the Mendelian rules quite clear (ignore the transposons that cause some colour variants which are not relevant for aurochs effigy breeding anyway), so it is way easier to breed for or against colour variants than it is for traits like size, proportions, appendages, horn size, skull shape and any other polygenic traits. This should be known to any animal breeder. Therefore, I prefer an aurochs-sized and aurochs-proportioned Chianina over a mediocrely aurochs-coloured Heck.

Anyway, to see how colour affects our perception of the looks of an animal, and also to see which results crossbreeding with Chianina can achieve, I did this drawing of hypothetical aurochs-coloured Chianina using the photo of a cow from Wikipedia and that of a bull I found via google (I would like to put the link there, but I couldn't find it anymore):

I admit the drawing is sloppy. But it was not my intention to fabricate a masterpiece, but merely to give an impression how such Chianina would look like. It would probably take a while to breed them, because all those annoying dilutions are at least partly recessive. All aurochs projects and Heck cattle have this problem.

Aurochs-coloured Highlands might be easier to breed because they do not have such dilutions, brindle is widespread but dominant and therefore easier to eradicate from the population. Taking black Highlands would be wise to avoid the recessive red variant, and crossbreeding with Maronesa might be the fastest way because they have the desired sexual dichromatism and long horns as well. It would be pointless from the breeding-back perspective, but fun. Initially I intended to draw hypothetical aurochs-coloured Highlands as well, but was too lazy in the end.

News from the True Nature Foundation

The TNF, the foundation that runs the Uruz Project, has posted the news that they received a number of Watussi cows from the Brüggen Zoo and they were moved to Bielefeld, where they will form a new breeding herd together with at least one Chianina bull. For a photo of these Watussi, go here. I was told that they are about 130 cm tall at the withers. That bull is a very young one, a son of one of the Chianina at Kloster Lorsch, and will turn one year old in April.
Claus Kropp kindly provided me with a photo of this bull calf from December 2014: 

It can't be judged how it looks will be like yet, but it seems that it is going to be a long legged and short trunked critter, also when looking at its mother. Considering that his mother is 165 cm tall according to C. Kropp, I am really curious on how large he is going to get. 

I was happy to see that this is his mother, as she is my favorite Chianina at Lorsch. Her build is very good with a decent hump, and her horn curvature is very desirable as well (the right one was accidentely deformed at young age, I was told). If you compare the colour of the mother below and the bull calf above, you might see that the first one has a beige/light brown tint while the latter one has a dark greyish tint, which is a clear sign of sexual dichromatism masked beneath all those dilution factors. The Sayaguesa x Chianina crossbreeds at the Lippeaue show the same. Therefore I am convinced that the chance is high that Chianina have retained sexual dichromatism. This will be helpful to get a good colour out of the Chianina x Watussi combination.

Dedomestication series pt. IV: Implications for "breeding-back"

The aurochs was, per definitionem, the wild type of Bos primigenius. If “breeding back” aims to approach the aurochs as close as possible, the result has, ultimately, to be a wild animal. Artificial breeding with domestic breeds can only result in an animal that is itself domestic, no matter how aurochs-like it looks, and it will still display a number of domestic traits – be it optic, behavioural or whatever.

For part I, for part II, for part III, for part IV.

Nature knows better

Our knowledge of the developmental cascades of many domestic traits is probably not deep enough to select against them and reconstitute the original, wild-type state. See for example the fact that selective breeding in farm foxes for earlier maturity and more offspring directly was not fruitful, but selection for tameness alone brought exactly that result. How to select on the genes coding for endocrinologic cascades that regulate the amount of hormones that control how bulky or muscular the body is, or those that regulate the timing of developmental mechanisms that result in an either elongate or paedomorphic skull? I am sure that just selection for long snouts or an athletic body directly in a population where these features are present only to a mediocre extent (f.e. uncrossed Heck cattle) will not be fully successful. The same goes for achieving a well-developed sexual dimorphism both in size and colour. Just selecting for large bulls or small cows certainly would not do it, breeding only with bulls that mature late would slow down the whole breeding process which is slow enough already and it even is not sure whether this is the right way or not. If just always taking out all black cows is the effective way to achieve a fixated, well-pronounced sexual colour dimorphism is dubious as well, because always even in Heck cattle herds with a good sexual dimorphism half of the cows are coloured like bulls.

Artificial selection might even be counteracting itself. I already expressed my thought that the muscular, athletic body of Lidia might be linked to its temperament. For example, the True Nature Foundation plans to do a project trying to collect suitably aurochs-like Lidia and breed them for a greater resemblance of their ancestor (which is awesome, I have been dreaming of such a project for a long time). But the selection program will also include selection against aggressiveness to make them easier to handle. That is fully understandable, grazing projects do the same, nobody wants dangerous cattle that are impossible to handle. But actually this is the same breeding for tameness and docility as in the farm fox experiment, so maybe it will result in a further domestication of fighting cattle and therefore reduce the athletic, muscular body and cause more paedomorphic features? This is just an idea, future might show if there is truth in my guess. The same goes for the pleiotropic connections between traits, such as those that cause white spots. I think that most of those connections are hardly accessible to us and that hinders us from effectively selecting for the desired phenotypic traits. For example, domestication in bovines almost always results in the horns getting “pulled” outwards or up/downwards. This is apparent in a lot of cattle and sheep breeds. Many breeds used in breeding-back have such horns and it proved to be difficult to select for a fully aurochs-like horn shape, but in OVP it seemingly developed by itself in a few individuals. The next problem is that we do not know how large exactly the impact of phenotypic plasticity is (f.e. on muscularity of the body, horn and body size, skull shape etc.; in the case of behaviour it is clear that environment is a crucial factor). To evaluate that, extensive comparative studies would be necessary which would take several years.

Probably all cattle would show herding and defensive behaviour in the wilderness. Free-ranging cattle do and also farm cows defend their calves aggressively. It is open whether the reproductive circle of cattle adapts to the seasons due to environmental impulses or natural selection, but I think the latter is the case because cattle in grazing projects are not fully adapted to the seasons in this respect. Perhaps natural selection could be mimicked by taking out cows and their calves that calved during winter or fall, but I am not sure if this would work.

All in all I would simply say that “nature knows better” in any case. This does not only apply to ecologic and immunological capacities, which are not visible and already present in primitive landraces to a certain(!) extent. The fact that certain wild traits appeared in free-ranging populations whose founding individuals certainly did not have them (f.e. the hump and body conformation in OVP, S.v.F. and Amsterdam cattle, or the long snout in some cows at OVP) and probably would have been hardly achieved by selective breeding implicates to me that natural selection “re-develops” aurochs-traits in the way outlined in the previous posts and better than artificial selection ever could. Artificial selection probably would be more of a mimic of adaptive wild type (=aurochs) traits, while natural selection produces truly adaptive traits. Natural selection knows better how to eradicate domestic traits such as paedomorphy or a reduced sexual dimorphism because we have no exact clue of the connections and interactions of development, pleiotropy and environment in morphology, behaviour and ecologic and immunologic capacities.

How to do it

Nevertheless, I think that simply tossing some bunch of any cattle into nature will not result in the dedomesticated near-aurochs that we want to see. The existing free-ranging cattle populations demonstrate that. If the founding individuals do not have horns that are either already aurochs-like in certain aspects or at least very diverse, it would take either very long for aurochs-like horns to evolve or they would end up in a more or less different shape. Aurochs colour (E+) won’t evolve if not present. Colour probably only has a weak influence on evolutionary fitness in cattle and probably coincidence will play a large role in when a certain colour variant becomes fixated in the population. So releasing just any cattle of any colours will not result in an authentic, aurochs-like colour – just as an example.  

As outlined in pt. III, we cannot simply expect that “nature” always changes the cattle in the way the aurochs was. Evolution is not static but dynamic and animals always adapt to the current circumstances. If we want a wild, near-aurochs, we have to simulate the evolutional adaptive environment the original lived in. The European aurochs evolved under the predative pressure not only of wolves (as juveniles also lynxes, foxes and bears), but also big cats (not only until the end of the Pleistocene; lions and leopards still lived in parts of southern Europe into the antiquary). They had way more space to live, graze and migrate and had to compete with a whole range of other herbivores. For that, a large reserve would be necessary to avoid the island effect and introduce competing herbivores (the most important competitors probably are deer and horses, as realised in the OVP) and carnivores. The introduction of big cats is a topic that I do not want to open here but let’s assume it won’t happen and there will be only fox, lynx and wolves (bears are problematic as well). No medical care or supplementary feeding will cause legal problems which should be solved like in the OVP and what the Tauros Project is trying to achieve, i.e. to classify the cattle as res nullius and legally wild animals. The cattle would have to be totally reproductively isolated, otherwise the dedomestication process would slow down.

And that’s how I would do it:

At first I would try to produce an aurochs effigy that is as authentic as possible by selective breeding (“breeding-back”). All or most desired traits that can be achieved should be present and as be stable as possible. Then I would introduce them together with individuals of primitive cattle like good Sayaguesa, Lidia, perhaps Maronesa and good Chianina, Castellana Axarquica, good Boskarin, Betizu for their feral history and Yakut cattle for their great adaptions to cold and their genetic distance to European breeds. The result would be a genetically diverse population that has all aurochs features and all founding breeds well-adapted to harsh climate, sparse vegetation and resistant to diseases. But I would not include too many individuals of breeds with strong dilution factors and/or short horns. The heterogeneity in the first generations would be high, and it would certainly be very interesting to see how the frequency of the single features is going to evolve. The traits should be categorized and their frequency be evaluated all three years or so, to document the evolutional shift in the population. Selective culling should only be used as a tool when it is apparent that natural selection has left a strong mark in the population already. Culling should focus primarily on fur colour, and only be carried out if the population is in a good state. For example, the Heck cattle at OVP are in a crisis at the moment and the last thing that I would do now is selective culling. I think an ideal population size would be 500 individuals at least. In any case more than 100.

But let’s be realistic, an area large enough to sustain a viable population of all the three herbivores plus wolves and perhaps also lynx that also can remain totally untouched by human management is not easy to achieve for conservation. But also without predators, most of the selective pressures described in part II would still be there, and if you have read carefully you probably noticed that predators are likely not among the most important factors acting upon such a free-ranging population.

The concept I described is basically what the Polish naturalist Feliks Pawel Jarocki suggested as early as 1835 (without the selective breeding part), only eight years after the aurochs was formerly described. He proposed that the release of cattle into wilderness so that they would live under the same circumstances as the aurochs did, would result in a “revival” of the original form. Note that I am not claiming that the original European aurochs can be revived the way described here.

Therefore what I am suggesting is a mix of breeding-back and dedomestication. My opinion is that dedomestication inevitably has to be the end phase of creating a near-aurochs. “Natural selection with a kick-start”, as the Tauros Project would call it.

Feral, wild and dedomesticated

At which point should be call such a dedomesticated aurochs-like population “wild”? Actually I have been dealing with terms like “dedomesticated”, “feral” and “wild” without defining them appropriately all the time. What I do now is what I should have done right at the beginning of the dedomestication series, I apologize. Parts of my definitions are inspired by those given by the user “Roberta” in the Carnivora Forum thread on the aurochs.

While genetics use the “wt” terminology mainly for alleles or any phenotypic features, zoology considers all aspects of the entire species, also including its history and interaction with the environment. Both feral and wild populations have in common that they are not enclosed (on small scale) and not dependent on active human help (this does not apply if the species is a commensal, where its ecological niche is living from and around human civilization). A wild species evolved/-s with and within its ecosystem and therefore does not change it rapid and dramatically. A feral species, on the other hand, is sometimes invasive and sometimes not.

Another distinction between feral and domestic might be the amount of genetic structure that was either influenced by evolution or by man. Considering that there are actually a lot of populations of wild animals are managed to some extent and are physically limited, this aspect might be of much greater importance. A wild type trait is defined as a trait that is shaped by evolution, not by man, and occurs in nature. As you have probably noticed, this applies to both a pre-domestic and a (I think hereby I introduce a new term) post-domestic phase. Classic zoological terminology only recognized pre-domestic wild animals as “wild-types” and therefore a wild type is usually also understood as the wild forerunner of the domestic type, and it never was considered that there can be a post-domestic wild-type as well. If you will, the Dingo or the European mufflon are post-domestic wild animals to me, or at least wild animals. They have been living in the wild for long enough, are adapted to their environment (with the exception of those mufflon populations that have been introduced where this species simply is not native, f.e. wet Central European lowland forests) and are an integral part of the ecosystem without causing changes that we call “damage”. But as you see, the line between feral and wild becomes arbitrary at this point. I would call Chillingham cattle and the Heck cattle at OVP not even truly feral, because both live on a rather small, enclosed area and furthermore, the Heck cattle have not been living there for all too long, and the genetic structure of Chillingham cattle is highly influenced by man (see previous part). In this case, I prefer to call the state they are living in “free-ranging”. Amsterdam Island cattle and Betizu on the other hand are/were feral in my understanding of the word.

No question that a post-domestic wild animal has to be a dedomesticated animal. Dedomestication is, as the word implies, the “reversal” of domestication. However, I prefer to call it the loss of domestic traits, as “reversal” implies the full re-emergence of the original state, which is not the case. Those domestic traits are, to put it in a nutshell:

- Paedomorphic features both in behaviour and morphology.

- reduced sexual dimorphism and usually also brain volume

- novel morphological traits (colour variants, horn shapes/sizes, change of body size, appendages etc.) due to pleiotropy, developmental cascades and relaxed selection; sometimes exaggerated due to artificial selection

- reduction of traits crucial for reproductive success – behavioural, immunological, sensory, ecological or morphological – due to relaxed selection

- loss or reduction of traits linked to sexual selection due to artificial selection

- Heterogeneity due to genetic drift and artificial selection

I would not say that a post-domestic or secondary wild type has to have lost all of those domestic traits; only those that lower the reproductive success. If natural selection or genetic drift allows or even fixates novel traits like some colour or horn variants or structures like a large dewlap, I think it is OK. In my opinion, they don’t even have to be completely homogeneous in terms of horn shapes and colour. First of all, the aurochs was quite diverse regarding the pronunciation of the “primigenius spiral”, size and orientation relative to the snout, and the snout length itself varied slightly as well. Although most wild species are highly uniform, there are some displaying more than one colour variants. Northern wolves for example are more or less diverse in that respect, partly also thanks to domestic dog introgression.

How to scientifically categorize the such a type of cattle? Regardless of if you regard domestic animals taxonomically relevant or not, these cattle would not be domestic, so why not classifying them? If so, I would simply tag them as Bos primigenius taurus because they will be neither genetically nor phenotypically 100% identical to any of the three aurochs subspecies, and the epitheton “taurus” has already been given for the domestic cattle clade. However, a prerequisite for a taxonomic status is that these post-domestic cattle form one reproductive (meta)population.

All in all, the line between feral and wild is fluid and the distinction is subjective. I doubt that we will live long enough to see a fully dedomesticated near-aurochs, but at least there are good chances to see an aurochs-like cattle population in a process of dedomestication which shows clear signs of evolutionary changes. The challenge is to find a suitable area of sufficient size and overcome legal and public issues.