Monday, 24 March 2014

Post #100! And: Is there a difference between effigy breeds and landraces?

This is my hundredth post on the breeding-back blog, and I am really happy how my blog has been growing – last week I cracked the 500 mark for views per day, and it seems that my blog is the second among the top Google results for “breeding back” already.  Thanks a lot to my awesome readers!!
I spent the last day clearing up older posts by correcting language mistakes, removing dead picture links and inserting a few updates.

Moving back to the topic, the question in the caption. As usual, I define an “effigy breed” as the result of breeding for the desired traits of the extinct animal you want to optically recreate à effigy breeding, a term I transferred from German Abbildzüchtung as a neutral alternative to “breeding back”.  I often thought about whether it is useful to make a sharp distinction between an effigy breed and a primitive landrace, and if we should consider an effigy breed “artificially/secondarily primitive” and a landrace “originally/primarily primitive” (note that by “primitive” I mean less-derived in a value-free sense. I am not happy with that term though, but I haven’t found a useful alternative yet). The arguments for such a distinction are
1) effigy breeds are aurochs-like or wild horse-like because they have been crossed and selected for uniting as much wanted traits as possible, while primitive landraces apparently remained in this state of primitiveness since the beginning of their domestication and therefore their resemblance with their wild types is the legacy of nature instead of artificial selection.
2) some effigy breeds/lineages are selectively breed for increasing the size of certain features without crossing by selecting within the breed only, for example large horns or long legs and snouts in some Heck cattle lineages, large horns in Watussi and large body size in Chianina. This might suggest that these traits should be considered secondarily primitive.

If that’s the case, what difference would it make, and are uncrossed primitive landraces more desirable than effigy breeds? I think this is a totally subjective question and everybody is free to form his own opinion, but I see problems with these two hypotheses outlined above.

1) Domestication is a process consisting exclusively of crossing and selecting over thousands of years. There are, above a certain level, no consistent lineages with a consistent set of features and purebreeding is a relatively recent phenomenon of the last few centuries. Phenotypic traits get transferred, altered or lost all over the phylogenetic tree of the domestic trait of a species (as long there are no isolated subclades). This explains why some cattle of central European dairy breeds like Braunvieh or Fleckvieh can show the upright, lyre-shaped horns of steppe cattle and zebus but also, surprisingly, horns with a perfectly aurochs-like shape. Or the Murnau-Werdenfelser, a central European dairy breed as well, is a morphologically unspectacular breed but yet has a quite aurochs-like coat colour. Each cattle breed has a history of being a mosaic of its neighbouring conspecifics, depending on how large the area of exchange and how intensive this exchange is.
The history of most Iberian primitive cattle is seemingly not very well-documented, but it is in some Italian and eastern European breeds. And it shows that aurochs-like landraces are no exception from having an origin of crossing and selecting and introgression of breeds differing to a varying extent. For example, Podolica was influenced by Braunvieh, Chianina, Romagnola, Maremmana, Simmentaler and other breeds during the 20th century. Boskarin from Croatia is a mix of Podolica, Maremmana, Chianina and others. Many of the small Italian aurochs-coloured breeds apparently are mixes of local landraces and Braunvieh-type cattle, just as many eastern European cattle breeds such as the Rhodopian shorthorn or Busha. Pisano looks quite aurochs-like with their primitive colour, the long legs and their – in some cases – aurochs-like trunk. If it was an Iberian breed with an undocumented history, an advocate of the distinction of original and artificial primitiveness probably would propose that this breed and its ancestors always possessed these features in this combination, and if their level of optic primitiveness had changed it more likely decreased rather than increased. However, we know that Pisano is a mix of Chianina, Braunvieh and Tuscan cattle – and since none of the breeds above has a slender, long-legged stature with an aurochs-like colour at the same time, it must be considered artificially primitive because it is the result of crossing and selection, just like f.e. Taurus cattle would be tagged this way. Whether it was the intention to breed aurochs-like cattle or not is irrelevant for the genetic make-up of the cattle and plays no role. The only thing the “genes care about” is whether or not the breeder selects for, say, long legs, or not and not what his intention behind is. There are more examples for this, like Sardo Bruna, and I bet that if we’d know more about the breeding history of the Iberian primitive breeds, we’d rub our eyes how many examples there are. Look at this Sayaguesa cow used in Tauros Project. It has a reddish brown back like an aurochs cow would have, while most other modern Sayaguesa cows are very dark brown to black all over their body. You might think we are looking at a case of a cow with a remaining sexual dichromatism that wasn’t diminished by selection yet, but in fact this cow is 25% Alistana-Sanabresa. Does that make this cow less aurochs-like? If we wouldn’t know, we wouldn’t even think that there is a difference. Another example: imagine Barrosa and Sayaguesa already existed 300 years ago, but not Maronesa. Some breeders had the idea to crossbreed these two for whatever reason, and the result after some selection is a new breed with horns intermediary between these two, sexual dichromatism (we know thanks to Taurus cattle that sexual dichromatism can surface in crosses between wildtype-coloured breeds of different tones), a short snout and comparably small body size. After a few genetic bottlenecks that always can occur, the modern Maronesa phenotype would have occurred. This is just an imaginary scenario and very likely totally incorrect, but considering that the creation of a new breed always is the mixing of other breeds, and that there is no reason to believe that there was less exchange between the breeders than in similarly civilized regions of Europe, we have to assume that each primitive breed is originally a mix from other more or less primitive breeds. The same goes for horses of course. And it is simply obvious that coincidental combinations of certain primitive breeds can result in cattle or horses that are either a) as much as, b) less or c) more like their ancestors in appearance than the founding breeds themselves. See the Barrosa-Sayaguesa example, or a mix of Maremmana-Pajuna. Whether it is a, b or c depends on the (coincidental) selection or bottlenecks as well of course. As an example, I wouldn’t be surprised if the very long snouts of some female Sayaguesa and perhaps some Pajuna are the results of in-crossing of Holstein-Frisian, which increasingly take place.
Furthermore, it is important to note that some wildtype traits simply cannot be reinvented just by selection. While this is true for quantitative traits like horn size, body size or leg length (which are influenced by environmental factors as well), it is simply impossible for certain colour aspects. Wild type base colour (E+) simply does not reappear in a population that doesn’t have it just because one “tries” to select for it somehow. It has to be present and become fixed, and it stays the same allele as in the founding breeds you got it from. You can’t “create” dun horses either. So a wholly aurochs-like or wild horse-like colour in an effigy breed that is the result of crossbreeding and selection is as authentic as in a primitive landrace, which itself was very likely originally a stabilized mix, and there is no evidence that all ancestors of that landrace always possessed the same colour setting.

2) It gets more complicated in features such as horn size and curvature, leg length or body or hump size – these traits probably are the result of the interaction of many genes we can’t track down yet, and are influenced by environmental factors and phenotypic plasticity as well.  But what we can say is that if you cross a large-horned with an aurochs-coloured breed and unite those traits in one herd, both the features are not anymore artificial than those of its founding breeds, because they passed on exactly the same genes to this herd. And whether or not the genes responsible for large horns are the same as in the aurochs we just can’t tell. Therefore I use “aurochs-like” instead of “primitive” when referring to phenotypic traits, because the latter word automatically implicates that it is a unchanged, genetically identical trait of the aurochs, which is baseless except in the case of colour, while “aurochs-like” simply is more neutral.
Not to forget, there are primitive southern landraces which were in fact selected for traits unintentionally resembling those of the aurochs – such as Barrosa, which has artificially large horns evidently created by selection within a population without crossing in other large-horned breeds (just compare modern Barrosa with those on that old photo UPDATE: I just realized that the link did not direct to an old photo of Barrosa cattle but to a Russian Death Metal song, sorry.). Does that make their large horns less desirable for breeding-back? I don’t think so. It is the goal to create an optical and functional imitation of the aurochs not only for fanciers and education but in fact primarily for its function and value as a native animal in natural areas, and nature does not care about such bagatelles we don’t even have a convincing clue about.

 To put it in a nutshell, I don’t think it is useful to regard effigy breeds as artificially aurochs-like and landraces as originally aurochs-like. After all, it sounds like an inconsequence in itself – a product of crossing aurochs-like breeds and selecting them for uniting their aurochs-like traits should be less original in the end? The history of modern domestic breeds is very dynamic and so probably was that of their phenotype. Before race breeding came up, farmers probably were very pragmatic and used those animals accessible to them and selected them on traits they needed for certain purposes. Either large udders for milk production, or a strong and large-humped body for draft work. The history of many of the Iberian breeds is undocumented, and that of Italian and neighbouring landraces shows that they are themselves mixes of a several breeds, so we should assume that the Iberian cattle breeds are of mixed origin like any other domestic breeds as well, and that their phenotypic resemblance to the aurochs is mostly coincidental. As a consequence, I don’t think that the aurochs-like phenotype of a good Taurus bull is less original than the phenotype of an aurochs-like Pajuna.

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