Sayaguesa arrived at Kloster Lorsch

On February 12th, a group of Sayaguesa arrived at the herd in Kloster Lorsch, Hessen (Germany). They number four individuals, three young cows born in 2015 and a one year old bull and are part of the Lorsch-Bielefeld project. The Sayaguesa were imported from the Netherlands and are, unless I am wrong, from the same breeder as those of the Tauros Project and recent imports for the Lippeaue. 

The young bull is going to form a new herd together with Hungarian Grey cows expected for summer 2016. The young cows, however, will not partake in breeding until 2017 because of their young age. 

As you can see in the news entry of the Auerrind Project's webpage, they are beautiful animals. You can see that the cows are not totally black but have shades of brown instead, some more than others, probably thanks to Alistana-Sanabresa influence. I expect them to become good-looking animals, like their relatives at other locations are. And I am very much looking forward to see the first cross results. 

Genealogical trees for some Taurus cattle from the Lippeaue

Using stock lists of the Taurus cattle population in the nature reserve Lippeaue, Germany, of the years 2015, 2013 and 2011 I was able to track down the full genealogy of a number of Taurus individuals to their last un-crossed ancestors. I have been planning to visualize that since a few days after my return from the Lippeaue in 2015 (see here).

Thanks to an extensive photo archive that I was provided by Matthias Scharf from the ABU, I was able to do five complete genealogical tables showing the ancestry of 24 crossbred animals, about the half of which are still present in the herd.

I used the program GIMP to make these tables simply by arranging them in the correct order and putting lines on them, since I don’t know of a program that would be better for visualizing such genealogical trees.  

Now I am going to cover the respective trees and give some information on those individuals that have not been covered on the blog here. I have been planning to do a post on older, not formerly presented individuals from the Lippeaue of past years here for a long time, so more information is about to come and I don’t want to give away too much for now.

Mind that similar looking siblings or parents and offspring do not necessarily imply that those traits shared are genetically stable. Just a few animals are way too few to determine the genotype of an individual. Equally, a sibling pair made of a red cow and a black bull does not imply that both inherit a well-marked sexual dimorphism. They are just two possible outcomes of genetically heterogeneous parents.

The breed identity of a pure founding individual is marked with a letter in brackets, H standing for Heck, C for Chianina, S for Sayaguesa and L for Lidia. Only those individuals whose deaths I know for sure have a †.

42 604, 79 289, 42 650 

Photos:

Eloisa, Morena, Lusitana, Luca, Dona-Urraca, Lola: Matthias Scharf

Julia, Lamarck, Lirgit, 42 604, 79 289, 42 650: me

Mator, Lancelot: Either Margret Bunzel-Drüke or Matthias Scharf, I’m not sure.

Lancelot was a Heck breeding bull of Neandertal descend (Eiszeitliches Wildgehege Neandertal; a large-horned Heck cattle lineage; almost all founding Heck individuals of the Lippeaue were from the Neandertal). Mator was a Dutch Heck bull imported from Slikken van Flakkee. Both Lusitana and Lola are not present in the herd anymore, but I do not know if they are still alive or dead.

The sisters 42 604 and 79 289 look quite alike, but as I just outlined this does not necessarily imply they are homogeneous for those traits they share.

Laniana

Photos:

Besucona, Lucio, Limette, Sinnombre, Lombritz, Lani: Matthias Scharf

Nestor: Either Margret Bunzel-Drüke or Matthias Scharf

Laniana: me

Nestor was, as the “N” implies, born in the Neandertal and is the father of Lancelot. He is also the father of Lucio, a large Heck x Sayaguesa that left many descendants. Besucona was one of the ABU’s three Lidia cows. Lombritz was used as a breeding bull at Klostermersch-Süd until he died when he broke into the ice. Limette and Lani are not in the herds anymore, but I don’t know if they are used somewhere else or have died already.

Lena

Photos:

Churro: Matthias Scharf

Laura, Larissa, Lena: me

Lena is the daughter of Larissa, which probably measures about 160cm at the withers and is their largest individual at the moment. The father, Churro, was a beautiful Sayaguesa bull that produced a lot of good-looking offspring.

Larwin, Liberta

Photos:

Locusta, Leila, Lissy: Matthias Scharf

Larwin, Liberta: me

Morica was the only individual for which I was not able to find a photo, but I didn’t want to throw away the whole tree. The light colour of Locusta is very surprising for a half-Sayaguesa, but it seems that her father Lancelot carried some dilution in its genome (see the light brown colour) that might have left a mark in his daughter’s coat colour. Larwin was used as breeding bull at Disselmersch for a short time, but was slaughtered for his small head, long dewlap and heavy body. Liberta measures at least 153 cm at the shoulders, many of the other cows at Klostermersch-Süd are about the same size. Locusta, Leila and Lissy are not in the herds anymore, but I don’t know about their status.

01 862, Londo, 55 392

Photos:

Aguaclara: Matthias Scharf

Londo: Margret Bunzel-Drüke

Lepisma, 84 024, 01 862, 55 392: me

Neither of those young unnamed bulls will be kept in the herd, but it was still interesting to show their ancestry, especially because 01 862 contains all four breeds. Londo is, for now, used as a breeding bull at Klostermersch-Süd despite its longish trunk and the small size. He resembles his father, which is larger and one of their best bulls.

Dun factor identified - reveals surprises and confirms old suspicions

Finally I have the time and mood to write this post. Regarding the coat colour of Eurasian wild horses, historic reports give us a clue but are not always unambiguous. For example, there is room for interpretation what “tan” or “mouse-coloured” is supposed to mean, because the authors of former centuries certainly did not use the words in the sense of modern horse coat colour terminology. In previous posts, I summarized and analysed all historic texts on wild horse exterieurs available to me:

What theTarpan looked like

Westernwild horse exterieurs: what does the evidence actually say?

C.H.Smith’s description of the Tarpan

WereEuropean wild horses dun-coloured or not?

Due to the ambiguity of historic accounts, genetic research identifying coat colour genes from ancient DNA of predomestic horses provides substantial additional clues on the actual colour of European wild horses. The last one of the posts linked covers a crucial question. Previously it has been resolved that Pleistocene and Holocene European wild horses had both the Agouti A and a allele, which produce either a bay or black base colour. But these two loci do not determine the final colour that is expressed; the so-called dun factor works upon these two base colours, and produces the phenotypes bay dun (a colour that is certainly wild type since Przewalski’s horses and kulan-onagers show it as well), black dun (aka mouse dun, blue dun or grullo), plus the non dun versions of the Agouti colours. Sorrel is not relevant for us, because it is based on a mutation of the Extension locus that was seemingly not present in predomestic horses. The problem is that the dun factor was not identified until recently, so that the actual phenotype of these horses was not determinable. This table below shows all four possible colours based on the Agouti locus (never mind the leopard spotted, that’s another story) that I did for Wikipedia: 

Now, the dun factor has been identified in a paper by Imsland et al. a few weeks ago [2]. It was found that the Dun factor sits on a locus for the TBX3 transcription factor. Loss-of-function mutations on this locus cause developmental defects in humans and mice in the development of limbs, apocrine gland, tooth and genitals. This shows once again that “colour genes” are not only responsible for coat colour but a wide set of developmental factors (for more, see the Dedomestication series). The study used domestic horses, Przewalski’s horses, other equids and two ancient horses. One ancient horses is from Yakutia and 4,400 years old, while the other one is from Russia as well, but 42,700 years old. Now what is interesting is that the Dun locus houses three alleles: Dun (D), non-dun1 (d1) and non-dun2 (d2). Non-dun1 still shows vestiges of the wild-type markings, you can still see a dorsal stripe although the contrast to the surrounding hair is not that big. In non-dun2 horses, on the other hand, primitive markings are totally invisible. What is even more interesting is that non-dun1 was found to be a wild type allele too, besides Dun. The Holocene Russian wild horse examined was found to be homozygous for d1, so it might have been either bay or black in life (the Agouti locus was seemingly not tested, alas). The Pleistocene horse was found to be heterozygous D/d1. So that not only means that non-dun equines existed already back 43,000 years ago, but also that both these alleles were present in one population at the same time.

As an example for a d1//d1 horse, see this photo of an Exmoor pony that I took at the Exmoor Pony Centre EDIT: Exmoor ponies have been found to be d2//d2 according to the supplement of the study. Exmoor ponies are, as far as I know, neither a nor A, but At/At, causing a condition called seal brown or dark brown. This allele has not been found in ancient populations yet. So if I interpret this correctly, considering the Exmoor pony’s colour a fully wild type colour would be speculative. For a d2//d2 horse, see this Noriker on Wikimedia commons.

Additionally to that, a considerable number of the predomestic horses tested in Pruvost et al. were found to be heterozygous A/a on the Agouti locus as well. This might implicate that in many predomestic Eurasian wild horse populations could have displayed all four possible colour morphs at the same time, the dominant ones possible more frequently than others (bay is dominant over black, dun is dominant over non-dun).

Although I suspected that, I find it surprising. I was actually hoping that the identification of the dun factor gives us a correlation between dun/non dun and a certain geologic age or habitat type, because it would make sense according to the camouflage effects the different colour morphs have (dun intuitively seems more suited to open habitats while non-dun fits forested, bushy landscapes in my subjective perception). However, if it is indeed true that there is no regional and geological correlation between the alleles A, a, and D and d1, wild horses of the ferus subspecies would be the only large herbivores displaying more than one colour morph in one population. It has been assumed that the homogeneity of wild animals, especially prey animals, is due to selection by predators because single individuals being coloured differently from the majority of the herd might be more attractive. Based on the current data, predomestic horses seem to violate this suspicion (I consider it merely a suspicion, I don’t know if it has been tested empirically).

However, there is another equine species that is known for occasionally showing deviant colour morphs, the Plains zebra (see this post).

I have been collecting a number of wild horse depictions in prehistoric art. I am going to present and analyse them here when I have the time to. Art, of course, leaves a much greater room for interpretation than genetic data. 

The authors also consider it likely that the zebra coat pattern is an extreme expression of dun plus wildtype markings [2].

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

[2] Imslandet al.: Regulatory mutations in TBX3disrupt asymmetric hair pigmentation that underlies Dun camouflage colour inhorses. 2015.

New website and name for the Lorsch-Bielefeld project

Originally, the breeding sites at Kloster Lorsch and Bielefeld were part of the Uruz Project until a split last year. Now it is a project on its own run by the Freilichtlabor Lauresham and Förderkreis Große Pflanzenfresser Kreis Bergstraße (Kloster Lorsch) and Landschaftspflegebetrieb Hohmeyer (Bielefeld), and they still follow the plan of two-line breeding (Chianina x Watussi, Sayaguesa x Hungarian Grey/Maremmana). They are also thinking about experimenting with other combinations of these five breeds. A third herd is planned. 

The project has its own name now, Auerrindprojekt. Auerrind in German means basically the same as aurochs (Auer-ochse), just with "cattle" (-rind) instead of "ochs" as a suffix. Auerrind is not used as frequently as Auerochse, but it exists. 
The project also has a new website, in German: www.auerrind.wordpress.com. They are working on other language versions as well. 

This website includes a news section where they will regularly inform on the progress of the project, monitoring, introduction of individual animals and breeding sites, also with pictures. They started a herd book, so that the relationships of each animal to each other is documented right from the beginning.  

As I reported in the post linked above, the young Chianina bull "Bruno" covered at least one Watussi cow at Bielefeld last year, what means that we can expect the first Chianina x Watussi crosses to be born this year. 

Last week, two Maremmana cows arrived at Lorsch. The import of three Sayaguesa cows plus one young bull from the Netherlands is in progress. The Chianina herd at Lorsch is also going to receive a new young Watussi bull this year. 

So the Auerrindprojekt is really getting going, I am so much looking forward to see the first cross results. Sayaguesa x Maremmana/Grey might resemble some Tauros and Taurus crosses, while the appearance of a Chianina x Watussi combination is something that I am really curious on. 

Aurochs-coloured Chianina, once again

I did a post on the idea of how aurochs-coloured Chianina might look like a while ago. I did drawings by tracking out photos of a Chianina bull and a cow and coloured them in the wild type manner. But the drawings in the first post were rather sloppy. So I did new ones a few days ago:

Original photo of the bull and the cow. 
Colour is one of the most prominent traits of an animal that leap to the eyes when having the first glance at an animal. A completely striped donkey would look more like a zebra to us than a donkey, and vice versa. And an animal with aurochs-like colour but otherwise not many aurochs traits might look quite satisfying because that trait gets much attention. This is the case in for many Heck cattle that would otherwise look nothing like the aurochs. 
And the reverse is the case as well. A "wrong" colour might make the animal appear less aurochs-like than it is, because it distorts the image as a whole. So I tried to imagine what wild type-coloured Chianina would look like, and how it would accompany with the aurochs-like proportions and slender body shape the breed usually has. 
I think that wild type coloured Chianina would be quite aurochs-like. What would still be needed of course are horns of desired shape and size, they would also need larger heads with more elongated snouts (usually), as much as bigger humps. For cattle to be rewilded in colder parts of Europe, they would also need longer and denser winter coat. The winter coat of pure Chianina is sufficient for Central Europe (the Chianina in the Lippeaue do just as well as the other cattle in the reserve), but I think that a winter coat like Hungarian Grey, Heck cattle and others have is what can be considered the ideal case. 

The full aurochs genome and inclusion of local aurochs into European cattle stock

With the full genome of a 6700 year old male aurochs being sequenced since 2013, it is possible to get better insight on the genetic relationships between wild aurochs populations and domestic cattle, if there was local introgression from wild individuals and which genome regions were particularly influenced by domestication. Now a summary of some recent papers.  

The whole-genome sequencing data placed this British aurochs as an outgroup to all modern European cattle [1,2]. mtDNA suggested that Southern European and North-Central European aurochs formed different genetic groups, the latter one being closer to domestic cattle than North-Central European aurochs are [3]. Perhaps hence the genetic distance of the British aurochs, but I tend to think that a southern European aurochs would be an outgroup to cattle as well.

Nevertheless, it seems confirmed that farmers did consciously breed wild aurochs into their stock. Orlando 2015 found that British cattle breeds (in particular: Highland, Dexter, Welsh Black, Kerry, White Park [2]) show substantial amount of admixture with British aurochs, sharing many polymorphisms [1]. This suggests that Neolithic farmers consciously bred aurochs into their stock, perhaps to gain local climatic and immunologic adaptions for their cattle (those which, after all, originated in the Near East) [1,2].

Orlando 2015 concluded: „Most European breeds apparently developed in situ with no mitochondrial influence from local aurochs, except perhaps Italy, Poland and Switzerland where B. primigenius mtDNA variants can be occasionally found in modern and/or ancient cattle. “

The case from Switzerland that he mentioned is described in a 2014 paper that reported the skeleton of a small female bovine standing only about 1,10 meters high at the withers, therefore being undoubtedly a domestic cow, dated to 5300-5000 years BP, but possessed a mtDNA P-haplotype variant of the European aurochs. Therefore this individual is the result of local admixture – and further not a first-generation hybrid because of its size. It again suggests intentional breeding with (female) aurochs [4].  According to Park et al. 2015, the Q haplogroup suggests limited local admixture as well [1]. It is important to note that no modern domestic cattle have the P mtDNA haplotype, which does not imply that all the nuclear genes introduced by the interbreeding were lost as well, as long the lineage did not vanish.

So now we have it confirmed that local aurochs did leave a genetic trace in European domestic cattle. Evidence indicates that it happened only rarely, but in my opinion this kind of evidence and the material we have is not able to determine the quantity of such events.

However, I see no reason to be euphoric over the results and draw conclusions like Italian or British cattle being more of an European aurochs than other cattle.  

An interesting side note: Park et al. 2015 detected traces of zebuine components in some Italian cattle (Chianina, Marchigiana and Romagnola) and East Asian cattle (Hanwoo and Wagyu). But it is also possible that those are alleles that other taurine cattle have lost [2].

It was found that domestication affected genes for neurobiology, growth, muscle development, metabolism and immunology [2].

Last but not least, an interesting passage from Orlando 2015 that brings up some aspects of domestication that might not be that often considered:

„Animal domestication is, however, likely to not just have remodeled the sequence of the

 genome. Micro- biomes, for example, might also have changed in relation with dietary

 shifts, which possibly affected important phenotypic traits, ranging from the physiological 

to the behavioral. As wild and domestic animals show subtle changes in brain gene 

expression networks, transcrip- tional changes are also likely to have been an early

 component of domestication.“

References

[1] Orlando, L.: First aurochs genome reveals the breeding history of British and European cattle, 2015.

[2] Park et al.: Genome sequencing of the extinct Eurasian wild aurochs, Bos primigenius, illuminates the phylogeography and evolution of cattle. 2015.

[3] Lari et al.: The complete Mitochondrial genome of an 11,450-year-old Auerochsen (Bos primigenius) from Central Italy. 2011.

[4] Schibler, Elsner & Schlumbaum.: Incorporation of aurochs into a cattle herd in Neolithic Europe: single event or breeding? 2014.