Saturday, 28 March 2015

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, 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.


  1. I'm glad I'm not the only one who is extremely disappointed with the attitude of the QP. In my opinion they still have a mountain to climb, but when you listen to them talk they truly think that their mission has already been accomplished successfully. It is sad to see such delusion because it means that the end result will fall short of what it otherwise could be. I'd be much happier, and have more respect for them, if i heard them give an honest self assessment and say that they have plans for addressing the remaining phenotypic differences you correctly identified than to hear them ignore those things and proclaim success.

    1. I see it think in a similar manner. But my point is, at most, that their attitude is not possible this way (recreating the Quagga).

  2. Marcel_Hendrik21 July 2015 at 07:28

    I'm quite fond of your overall work, although from a South African perspective I would however like to make a few comments concerning the "extinct" Quagga and some points of controversy.

    Firstly, with regards to the true identity of the so-called 'extinct' Quagga (in a great sense) we need to look no further than the narratives of the early European colonists and especially the many literate naturalists (i.e. John Barrow, William Burchell, Francois le Vaillant, Martin Hinrich Carl Lichtenstein and many others) that had great exploratory journeys through the South African "veld" where the Quagga and Burchell's Zebra were most common. So now I might add that the Quagga is NOT an extinct subspecies of Plains Zebra, rather the Quagga was the southernmost population of Plains Zebra representing a natural cline. In other words the southernmost "Quagga" were least striped (i.e. most typical "Quagga"), while the populations further north towards the Orange River became ever more striped until the "fully-striped" Burchell's Zebra persisted.

    But here lies the confusion! In the earliest days of European colonization and exploration in the Cape of Good Hope region (now Cape Town), the new colonists were presented with two very much different Equus species - namely the "Quagga" (Equus quagga) and the smaller Cape Mountain Zebra (Equus zebra). But as the colony expanded ever further into the hinterland of Southern Africa, a seemingly "new" species came onto the scene - and so William Burchell dubbed the "newly discovered" animal was named the "Burchell's Zebra". Note all three these so-called "species" were called "Quagga" or rather "Kwagga" often without differentiation in the mother tongue of the locals - Afrikaans.

    So in conclusion the Quagga was by no means a distinct species from the Plains Zebra and even to call it a subspecies would in fact be pushing it. The southernmost animals (south of the prominent Cape mountains) were the least striped, while further north towards the Orange River the animals fashioned ever more stripes. So the thoughts that the Quagga was indeed a distinct species originated from the Afrikaans language which described the different colour variations by distinct names (i.e. "kwagga," "bontkwagga," "Kaapse kwagga," "Karoo kwagga " "vlaktekwagga;" Note all indeed the same species).

    The quagga did not diverge from the plains zebra 200 000 years ago, they are conclusively plains zebra, and massive zebra herds that once ranged across the Karoo contained both "unstriped" animals (rendered as essentially quagga) and "more-striped" animals rendered as (bontkwagga; later Burchell's zebra [note no distinction was ever noted apart from "colour morphs"]). So the comparison of the Quagga project to the work of the Heck brothers are most definitely not the same. The Heck brothers tried to recreate an extinct species from domesticated descendants, while the quagga project only captured and consolidated zebras that were essentially quagga into a single population.

    I am the author of "Rewilding The Lost Wilderness" - a book about the larger mammals that inhabited the southernmost parts of Africa. I can send you more info and parts of my book if you would like to read more about the Quagga.

    1. Hi,
      first of all, thanks for your offer of sending me more info, here is my email adress: daniel.foidl( )itmed.at. Where the space is there should be an "@" of course.

      But I feel that you raised no argument that I haven't covered in my article yet, except perhaps that there was a cline of striping within the Quagga itself. Is there direct evidence for that? I know that there was variation within the degree of striping within the quagga population, but I haven't read yet that this variation was a geographic cline within the Quagga. Not that I would doubt that.