Tuesday, 28 November 2017

Thoughts for a Fighting cattle project

In a previous post, I presented an aurochs reconstruction I did recently. I wrote that it matches 100% what I imagine a Holocene European aurochs to have looked like, and that there are obvious similarities to many Lidia bulls (Iberian fighting cattle). Of course there is the danger of a Pygmalion effect: I might draw my aurochs Lidia-like, because I consider Lidia aurochs-like. This cannot be ruled out completely. But I always try to follow the evidence and not to be guided by preconceptions when doing illustrations. In the post linked above, I explained why I drew the bull the way I did. Here is the illustration I am talking about:

Actually, the fact that Lidia is one of the least derived cattle breeds left today did not just come to my mind because of that drawing, but was apparent to me right from the beginning when I started to become interested in the aurochs. I use to explain the looks of the aurochs to people as “much like a big, long-legged and large-horned Iberian fighting bull”. Also, aurochs expert Cis van Vuure comes to the conclusion that this Iberian breed is the most aurochs-like breed in his 2005 book “Retracing the aurochs: History, morphology and ecology of an extinct wild ox”. And also Lutz Heck considered the breed very reminiscent of the extinct wild form and relied heavily on it in his breeding project (his lineage diminished, modern Heck cattle most likelyhas no influence from the Iberian fighting bull).

I am going to sum up the aurochs-like pro’s of Lidia cattle:
- Muscular, athletic body shape very reminiscent of wild bovines
- High processus spinosi in the shoulder region (“hump”), probably as large as in the aurochs
- Often elongated skull shape, straight to convex snout and often prominent forelocks, sometimes also on neck and face
- Horn shape reminiscent of that of the aurochs with an inwards curve, although usually not entirely as prominent as in the aurochs  
- Short dewlap and scrota and small udders in less derived individuals
- Wildtype colour plus sexual dichromatism (albeit reduced) present in the breed
- Robust landrace, used to live outside with little husbandry all year round

Especially the first four points are very advantageous for “breeding-back”, as those traits are rarely found in any other primitive breeds to the same extent. All of them give primitive Lidia individuals a very aurochs-like appearance, and I am going to link some of those less-derived aurochs-like Lidia individuals here:

Bulls:

Cows:

For a video of aurochs-like Lidia bulls, go here for example. Another video of bulls with an exquisite anatomy and also respectable horns is here.

It is not only their less-derived anatomy and looks that make Lidia special, but also their hardiness and independency. As I reported in 2013, the three Lidia cows in the German Lippeaue reserve were the only cattle that were never seen to make use of supplementary feeding.

Despite of their many advantages, this breed also has considerable disadvantages. First of all, Lidia are quite small cattle. According to the Domestic Animal Diversity Information System, bulls reach a withers height of 130cm, and cows 110, which is 10-15cm smaller than Heck cattle and almost half a meter smaller than an average Holocene European aurochs. But even more important, Lidia cattle are being bred for aggression and “fighting spirit” and consequently are very aggressive and nervous, difficult to handle and often attack easily. Additionally to these two major disadvantages, Lidia bulls also tend to be short-legged compared to aurochs.

But of course it is the task of crossbreeding and selection to eradicate such negative traits, just as too small horns or undesired colour schemes. This has been tried in the Lippeaue, where three pure Lidia cows have been used for breeding. But they were not really satisfied with the results. For once, the crossbreeds remained comparably small, even if they had portions of Chianina or Sayaguesa, two large breeds, in their genome. The crossbreeds also displayed overly nervous or aggressive behaviour. This goes for the half-Lidia bulls as much as those that were only quarter Lidia such as Latino. One half-Lidia bull was sold to Hortobagy, Larus (which looked quite good), and Istvan Sandor it was the most aggressive bull he ever experienced. Another half-Lidia bull was sold to Denmark and slaughtered because he was too difficult to handle. I had a look at many Lidia cross individuals in Matthias Scharf’s photo archive, and many of them did look more muscular than crossbreeds without Lidia, but they also tended to develop a rather elongated trunk and did not possess the slim athletic waist of pure Lidia. As a side note, even pure Lidia bulls often grow rather hefty at higher age, and the crossbreeds seemingly tended to go that direction right from the beginning. So crossing-in Lidia was not that successful as hoped, which is why the ABU diluted the influence of this breed in their herds, and Matthias Scharf even said that he considered most Lidia-crosses to be “small, ugly and mean”. Currently, they have one good half-Lidia cow in their herd that also has a comparably relaxed behaviour.
Larus, a cross between a Lidia cow and a Dutch Heck bull (photo by Istvan Sandor)
So does this mean that Lidia is, despite its obvious qualities, not a useful breed for “breeding-back”? The Tauros Programme, for example, will not use this breed, especially because of its behaviour. Considering the difficulties in handling Lidia crosses, and the disappointing looks of many of the cross results concerning size and body shape, I would say that Lidia is indeed perhaps not worth the effort in conventional crossbreeding and keeping them in grazing projects.
Lidia x (Heck x Chianina) cow
Nevertheless, I also think that it would be a shame not to make use of the potential that is in the breed. Maybe the crossbreeding results are too double-edged to speak of a “successful use” of this breed in crossbreeding, but perhaps a project working exclusively with good, useful individuals of that breed and trying to eradicate the negative traits of Fighting cattle would be fruitful. I am thinking of a project that would at first try to find a sufficient number of aurochs-like Lidia cattle such as those on the photos I linked above that have the right colour setting, an athletic body, elongate skulls and good horns, and trying to acquire individuals that are as large as possible, as long-legged as possible and perhaps not overly aggressive or nervous. For this, I would try to search breeding sites all over Spain and Portugal and also include the Casta-Navarra breed. Selecting them for couple of generations for the good traits and against the three undesirable Lidia traits (small size, short-leggedness, aggression/nervousness) might produce some good results. When selecting against their aggressive and nervous behaviour, I would try to pay attention to not domesticate them any further by selecting explicitly on tame and docile behaviour, as pleiotropic effects can affect other traits as well as we see in all domestic animals (see the Dedomestication series). The question is, then, how to handle the cattle. It would be very difficult to next to impossible to keep them like in grazing projects or private farms. It might be wisest to handle them just like other Lidia breeders do, or perhaps like American farmers keep their bison, also with the necessary equipment. After all, a true wild aurochs probably could not be handled much differently, and it is no coincidence either that most natural grazing projects work with cattle instead of wisent. Of course it would be more advantageous if their behaviour was suitable for grazing projects, as grazing projects provide the most abundant opportunities to spread the breed and keep the animals under semi-natural conditions in herds of sufficient size, but this post focuses only on a scheme to breed a very aurochs-like strain of Lidia and not how to spread it. If it is indeed possible to breed a more relaxed and less dangerous behaviour but maintaining the wild cattle-like body conformation, it would be pleasant of course and perhaps enable the use of these cattle in grazing projects. However, I think it is questionable if this is possible within a few generations as fighting cattle have been bred for aggression for dozens of generations.

The idea of establishing an aurochs-like cattle population using Iberian Fighting cattle only is not new. Lutz Heck considered this breed almost indistinguishable from the aurochs, so that it is surprising that he still bred with other breeds as well. Cis van Vuure also proposed a project working with the most primitive Lidia cattle only in order to achieve maximum resemblance to the aurochs. I also remember that in email conversations it was once considered to put such a project into practise, but as several years passed now and I discovered no mention of it on the web I assumed this idea has died a silent death.

Also, I have my doubt that it is possible to achieve all aurochs-like traits using Lidia only. While some Lidia grow larger than the norm, I would be very surprised if selective breeding is able to raise the size range from 130cm for bulls to 160cm at least within a couple of generations. The same goes for horn size – while Lidia are not small-horned, the largest horns of this breed are still only at the lower size range of aurochs horns, perhaps still a bit too small when considering the size of the horn sheath. Also, the breed might need a boost in leg length, even after selective breeding. Furthermore, the problem with breeding for quantitative traits is that it takes comparably long and other traits might get neglected, including also genetic diversity (Lidia is already inbred).
Thus, crossing-in breeds that might help to overcome these deficiencies and also to increase genetic diversity is suggestive. The first option that comes to my mind are Chianina or Maltese cattle that would add the large size, slender posture and long legs. Maltese cattle have the advantage of introducing colour dilution alleles to a lesser extent and having longer snouts, but are far less available, with not even hundred individuals and only on Malta. The big disadvantage both breeds have are the very small horns. The horn size of Lidia already has to be improved (although some individuals, like these young bulls, have comparably large horns), and crossing-in one of these breeds would introduce alleles for very small horns, and since horn size is probably controlled by several gene loci it would be even more difficult to breed for the desired horn size. So it would be advantageous to add another breed to increase horn volume. Which breed would be best to do this job? One idea would be very large-horned Texas Longhorn with the right colour setting. However, the disadvantage would be that they are small-sized as well and have very outwards-facing horns. Another possibility are large-horned Heck cattle from lineages with a comparably stable inheritance; their disadvantages are their very domestic, heavy body and paedomorphic skulls. Using Watussi would greatly increase horn size, but also introduce many undesired traits concerning body shape, dewlap, colour, fleshy hump, skull shape et cetera, which is why this breed would have to be used wisely and with patience. If one cannot decide among existing stable breeds which one to chose in order to increase horn size, one could wait for other projects that are already in progress now to produce results that can be used for this job. Such as the Auerrind project, which has just started and is using Watussi. Perhaps in 15 years, when also a Lidia project would see first results, there might already be good Watussi-influenced Auerrind cattle that could be used in order to increase horn size in an improved Lidia strain without introducing too many undesired traits. This is just brainstorming – in practice it depends on many factors of course.

I would not suggest to start crossbreeding right from the beginning and to a larger extent, otherwise I would not be talking about a Lidia project here. It is about preserving, concentrating and improving the aurochs-like traits present within the Lidia population, and using other breeds only after a phase of selection when selective breeding has been proven to not being able to reach body size, leg length and horn size to a desirable degree. Another positive side effect of crossing-in other breeds would be that at least some individuals might display a more relaxed behaviour. I would use other breeds only carefully and perhaps use only F1 crossbred individuals in order to not diminish the advantageous Lidia traits. The point at which I would start to make use of such crossbreeds is when selective breeding has been shown to not reach all traits to a desirable extent, which would be after a couple of generations and therefore about 15 years or so (which would equal about five generations).

How could such an aurochs-like Lidia strain be named? For Maremmana, some people like to speak of the optically less derived individuals or herds of “Maremmana primitiva”, although those Maremmana primitiva do not represent one gene pool or breeding line. In the case of this Lidia lineage, on the other side, it would be one population/lineage. So perhaps calling them “Lidia primitiva” would be an idea.
I think that such a Lidia project could achieve a high level of authenticity regarding looks and anatomy and perhaps also behaviour (yes, exaggerated aggression is not “natural” behaviour; but tameness and docility are not either; you certainly could not have used an aurochs for draft work or have milked them). There is also the chance that with selection against exaggerated aggression and crossing-in of other breeds Lidia primitiva might once have a behaviour that is suitable to be used in grazing projects and other aurochs projects.

This post is just a thought-experiment, but I hope that it inspires and perhaps other people have similar ideas in order to make use of the potential we find in Spanish fighting cattle.



Saturday, 18 November 2017

Wild horses pt. IV: The Przewalski's horse - is it still a wild horse?

This title is surely a bit provocative – it is of course zoological consensus that the Asiatic Przewalski’s horse, Equus ferus przewalskii, is the last living genuine wild horse that is extant today after the last western wild horses disappeared. However, advocates of a number of horse breeds that they purport as living remnants of the European wild horse or at least being strongly influenced by original wild horses, sometimes put the status of the Przewalski’s horse as a genuine wild animal into question. They argue that decades of breeding in captivity and introgression from domestic horses has altered the nature of the Przewalski’s horse, and claim that the situation is comparable to what has happened to their favoured “wild” breed: an original wild population has been influenced by domestic horses and artificial selection. As a consequence, they argue that if the Przewalski’s horse still deserves status as a genuine wild horse, which is zoological consensus, then so does their breed of choice. However, the often purported background stories for those breeds being “near-wild horses” are not tenable after objective examination for a number of reasons (see here), but what about the arguments against the original, wild status of the Przewalski’s horse?

The lineage of the Przewalski’s horse separated from that of domestic horses several millennia ago. The exact point of separation varies from study to study, depending on the molecular chronometer and its calibration. The maximum I found was 160.000 [1] and the minimum 38.000 [2] years ago. This comparably long reproductive separation resulted in a different karyotype, the Przewalski’s horse having 23 pairs of chromosomes and the domestic horse 24 due to a fission or fusion (depending on what is the plesiomorphic state), but they intermix readily and without fertility problems. In the millennia of living side by side in the Eurasian steppe, the Przewalski’s horse contributed genetically to the domestic stock (which is not only genetically [3] but also optically apparent, f.e. see some Mongolian horses), and vice versa. The Przewalski’s horse gene pool was introgressed by domestic horses, especially in the 20th century. Photographs of wild herds from 1954 showed individuals of divergent colours (Wikipedia), indicating admixture. The whole modern population descends from 13 founding individuals, one of them was a domestic Mongolian stallion [4]. Does this mean that the original, genuine Przewalski’s horse is lost and the modern population is an altered result of intermixture?
Orlando et al. 2015 made a genomic study compromised of a large sample of Przewalski’s horses, post and prior to the bottleneck (including the holotype specimen), domestic horses plus a late Pleistocene wild horse as outgroup. The result is that although there are genetic traces of intermixture, also including such having an effect on the phenotype such as an allele associated with increased withers height, there are still lineages in the population that are virtually free of admixture[3]. Also, height is a highly multifactorial trait, therefore it cannot be claimed that Przewalski’s horses are taller now due to admixture because of one allele – the average withers height is still between 122-142cm according to English Wikipedia, between 120-146 according to German Wikipedia (note that there is also sexual dimorphism in size). Przewalski’s horses are still uniform in their typical colour, sturdy build, robust head shape, erect and short mane, short-haired tail basis, a very lightly coloured almost white winter coat, and other typical morphologic differences to domestic horses such as thicker hooves (Wikipedia). I would even say that domestic cattle left a bigger trace on modern American bison than domestic horses did on the Przewalski’s horse, yet nobody is questioning the bison’s status as a wild animal. Also I found no source stating that Przewalski’s horses with a domestic karyotype have been observed.

Yes, the Przewalski’s horse seemingly intermixed with domestic horses continuously after their point of separation, but I see no compelling evidence that this fact altered the genetic integrity of this wild subspecies. Furthermore, domestic animals introgressed the gene pool of their wild counterparts everywhere they shared the habitat – this evident in European wild boar that show deviant, domestic colours and there is also the hypothesis that American wolves inherited black and other colour variants from domestic dog introgression several millennia ago (this might also explain blue-eyed wolves). Yet nobody is calling their wild animal status into question.

If Przewalski’s horses indeed lost part of their wild animal nature due to domestic introgression and being bred in captivity for a number of decades, it might be helpful to look at a checklist of aspects typical signs of domestication:

- Morphological paedomorphy
- Behavioural paedomorphy
- Reduced brain volume
- novel morphological/optical traits (very typical: colour variants, particularly white spots)
- Earlier maturity and increased litter size (the latter aspect is not true of domestic horses either, so let us ignore it for now)

Przewalski’s horses do not show any signs of morphological paedomorphy, not even if you compare photos of the early 1900s to modern individuals. Przewalski’s horses still always have the robust, donkey-like skull with small eyes and their proportions do not seem to be altered as well. I have not found any remarks in the literature stating that Przewalski’s horses as a whole lost brain volume; domestic horses have about 14% less brain volume than Przewalski’s horses [5]. Captive Przewalski’s horses also have 14% less brain capacity than wild counterparts [5]. Since there are no separate genetic lineage between wild and captive Przewalski’s horses, this should be applicable to phenotypic plasticity. According to Wikipedia, earlier maturity in captive Przewalski’s horses has been reported, but explained with better nutritional conditions in zoos than in the wilderness and as far as I know the same phenomenon can be observed in other zoo animals. The behaviour of Przewalski’s horses and domestic horses is well comparable, but Przewalski’s horses have a way higher aggression potential than domestic horses, especially the stallions. This is universal for this subspecies and evident in zoos as well as grazing projects. I once was told that zookeepers are more afraid of Przewalski’s horses than lions. Przewalski’s horses can be tamed and ridden to a certain degree, but this is also true for zebras, including the quagga.
There are not any novel traits found in any Przewalski’s horse, such as a new colour variant, or fur modifications. There are occasionally individuals showing a white streak along the face or white socks, which is applicable to introgression from domestic horses.
Looking at some deer populations which have been kept in game parks for many generations, we see incipient signs of domestications, such as new colour variants or typical domestic spotted patterns, or beginning paedomorphic skull shapes – you can find this in some roe deer, red deer and fallow deer in European game parks and this is what I would call an early state of slow domestication. But we do not see that at all in Przewalski’s horses.
All in all I think there is not one compelling reason to claim that the original Przewalski’s horse is gone, that it has been altered by man and hybridization, or that it is on the edge of domestication. I see nothing that calls their status as a genuine wild animal seriously into question, especially when we look at other wild animals. And even if the critics were right, it would not make any of the domestic horse breeds praised as near-wild horses “wilder” than they are (or not are, actually).

Literature

[1] Ryder et al.: A massively parallel sequencing approach uncovers ancient origins and high genetic variability of endangered Przewalski’s horses. 2011.
[2] Orlando et al.: Recalibrating Equus evolution using the genome sequence of an early Middle Pleistocene horse. 2013.
[3] Orlando et al.: Evolutionary genomics and preservation of the endangered Przewalski’s horse. 2015.
[4] Bunzel-Drüke, Finck, Kämmer, Luick, Reisinger, Riecken, Riedl, Scharf & Zimball: „Wilde Weiden: Praxisleitfaden für Ganzjahresbeweidung in Naturschutz und Landschaftsentwicklung“. 2010
[5] Röhrs, Ebinger: Are zoo Przewalski’s horses domesticated horses? 1998.



Friday, 27 October 2017

New aurochs bull reconstruction

About three weeks ago, I presented a recent drawing I did of Europe's megaherbivore fauna of the Holocene. I promised that I am going to present the aurochs of the drawing on a separate, refined version which is what I am going to do today. I do that because I think it is one of the most accurate aurochs artworks I did recently. 

It is not a reconstruction that is based on an actual skeleton, but it is based on such reconstructions. Actually, the basis for the drawing were the bull reconstructions that I did in June 2017. I did them by tracking out mounted skeletons (I corrected anatomical flaws in the mounts at first) to assure maximum precision: 
For my recent life illustration, I used the Vig bull and Kopenhagen bull (c and d) as a model, and decided to do it in a running pose. Here is the result (I decided to watermark it as I repeatedly made the experience that my drawings have been used without permission and crediting, which is not pleasant): 
While body shape and proportions are dictated by the skeletons and wild cattle anatomy, there are a few aspects that allow subjective decisions. For example, I could have given my aurochs larger horns. But as the horns of Holocene aurochs were smaller on average than those of Pleistocene ones (see here or here), I gave it medium-sized horns. The snout is slightly convex and the nose curves downwards a bit, which is a trait found in many Lidia bulls but also displayed by a skull of a British aurochs. Colouration also leaves room for speculation or subjective decisions (see here), such as the colour of the eel stripe, forelocks and extent of the lightly coloured muzzle. I gave my aurochs a yellowish eel stripe, as it is the most frequent colour in wildtype coloured bulls and written sources are not clear (they just speak of a "lightly coloured" stripe). The lightly coloured mouth is never mentioned in contemporaneous texts, and cave paintings show them only for cows. In wildtype coloured cattle, this trait is very prominent in almost all cows and also widespread in bulls, but aging bulls often have it reduced. In gaurs and bantengs, this trait does occur, but not in all individuals. So perhaps it was not that prominent in grown aurochs bulls as well. As for the colour of the forelocks, I decided to draw them black, as this is better supported by the evidence (see the post linked above). 

All in all, I am very happy with this aurochs drawing as it really shows a 100% of what I imagine a typical Holocene European aurochs bull to have looked like. Perhaps not surprising as it was me who did this drawing, but this is not always the case. The result also bears great similarity to many Lidia bulls (Spanish fighting bulls), but more on that on an upcoming post. 

My recent aurochs reconstructions and illustrations all have been rather bull-based, which is why cows are about to follow. 

Wednesday, 25 October 2017

The Heck bull at Lembruch, Germany

I have several posts in preparation at the moment, but for today, I have a short post. It is about a Heck bull I found recently on the web, and I think he is interesting for a number of reasons. It is or was (the article is from 2014, so I do not know if it is up to date) the breeding bull of a herd at Lembruch, Germany, owned by the breeder Martin Kockmeyer. At first I want to present some photos of the bull (photos owned by the Presse-Bild-Agentur Nokem Martin Kemper, I got them from this article): 
First of all, the upper two photos show nicely how Heck cattle is a mosaic of its founding breeds: the body and skull shape of a Highland bull (not desirable from an "aurochs point of view"), the colour of a Werdenfelser or Corsican bull, the horns very reminiscent of Watussi. 
Secondly, the bull's name is "Arak", thus I suspect he might be from the Wörth lineage, which is/was remarkable for being the only closed breeding line within Heck cattle and having established large, thick, comparably well-curved horns. The bull also resembles those of the lineage, like "Albatross" or "Aretto" quite well. It is always good to see bulls from that lineage being breeding bulls on other herd, as it means that they are spreading the genetic make-up for good horns among the population. As it is apparent from the photos, the bull is seemingly not very large, which is unfortunately true of many un-crossed Heck bulls. 

On the VFA's sale site, there is currently a bull for sale from the same herd, named "Delika", born in 2014. The horns are of a very useful dimension and curvature, it might be a son of Arak: 



Thursday, 19 October 2017

Wild horse series Pt.II: The Konik, Exmoor and Sorraia myths

In my 2017 wild horse summary I had a look at what the evidence suggests regarding the life appearance, extinction and population genetics of European wild horses. There are some open questions, but also some things that we can say with certainty. One of those things is that European wild horses are as extinct as the aurochs. Nevertheless, there are three horse breeds that are often referred to either as surviving wild horses or near-wild horses, having a special status among European horses in being particularly close to the original wild form. Those three breeds are the Polish Konik, the English Exmoor pony and the Portuguese Sorraia, and each one of those has its own proposed background story that is supposed to link them closely to European wild horses. Those stories, however, are only purported and believed by advocates of those breeds, people involved in various projects, and are spread exclusively in non-scientific text books, public relations articles, or on signs in zoos while nothing of them is replicated in scientific literature, and none of those breeds are considered zoologically special among domestic horses. Advocates of the breeds blame a “conservative scientific mainstream” that is biased against the idea that the European wild horse might still be extant in some form, while the truth is that the background stories for the purported special status of the Konik, Exmoor or Sorraia are largely based on wishful interpretations, Chinese whispers and fabrications, actually making those stories myths that do not withstand objective examination.
Here I want to present and subsequently dismantle those myths. When not relying exclusively on sources that repeat the same stories over and over, but actually provide a deeper background knowledge, backed up by more evidence than usually delivered, it becomes rather obvious that the usually purported background stories for those breeds are not tenable. Yet those stories are pretty persistent and widespread, and I hope this post might become a little contribution to clear things up. 

The Konik myth: The Polish game park at Zamosc was the place were Europe’s wild horses survived longest. In the year 1806, these wild horses were donated to local farmers of the Bilgoraj region and incorporated to their farm horse stock. In the 1920s, the Polish agriculturist Tadeusz Vetulani started a breeding-back project using wild horse-like individuals from the Bilgoraj region and bred them selectively for a wild horse-like nature, eliminating the domestic influence their genome experienced in the hundred years before. He called the result “Konik”.
The truth behind it: Whether or not the supposed wild horses at Zamosc were truly wild and not feral, it is unlikely that they left a noticeable trace in the local farm horses. This claim is based on a notion by Julius Brincken in a book from 1826, but this book is full of errors, misinterpretations and fabrications, so it is not a reliable source. Furthermore, the Zamoski family was at war with the local farmers due to social unrests in the 1780s and 1790s, so it is unlikely that they would have provided them with generous gifts in the form of livestock. And even if they did, it is unlikely that the farmers tolerated a strong wild influence in their horses as it would have been a throwback in their productivity and suitableness for agricultural work, as wild horse hybrids used to show intractable behaviour [2]. And even if they incorporated wild horses, and perhaps only mares, into their stock, not much would have been left after 100 years. So it is very unlikely that the horses of those particular rural regions of Poland had a notable wild horse influence in the 20th century [1,2].
In Poland, there was/is a landrace called Panje horses. Those horses were very robust, of a small and stocky body and varying in colour between black dun, black, sorrel and bay. In an 1921 article, Panje horses were considered as possible wild horse descendants (which is to be refuted for the reasons stated above) by the researchers Gabrowski and Schuch. This drew the attention of Polish agriculuturist Tadeusz Vetulani to the Panje horse as “wild horse relict”, he tried to back up this suspicion with cranial measurements (that have been questioned [1]), and coined the name “Konik”, which successively replaced the name Panje horse. Private and public studs were created in order to preserve and spread the landrace. In 1927, Vetulani started an experiment that is often considered a “breeding-back” experiment, but was actually more of a dedomestication attempt, using Konik/Panje horses that he considered wild horse-like. But Vetulani’s herd was only one of many Konik lineages back then, and the Second World War created a lot of confusion, as many Koniks were moved all over Poland and Germany during and after the war. After the war, the stud at Popielno was the most important Konik breeding site. They performed two separate ways of breeding: one to continue Vetulani’s way of few human interference, and one of traditional indoor breeding and commercial sales. Popielno was one of many breeding sites, and when other countries started to become interested in this breed, they purchased from any stud available [2].
Thus, neither was Vetulani’s experiement an experiment of selective breeding, nor was it the ancestral stock of the modern Konik population but merely one of many lineages [2]. He did not create the breed, but merely coined the modern name of the breed. The fact that 10-5% of the modern Konik still show a black or sorrel colour or white streaks on the forehead (respectively) reveal the mixed origin of this landrace [2], and there is no compelling, not even plausible, evidence that their ancestral stock was strongly influenced by wild horses [1,2].
The Heck horse, which is often presented as a “recreated Tarpan” in Germany and also other countries, is in a sense a washy Konik – it was created by the Heck brothers in the 1930s and 1940s by crossing Icelandic horses, Gotland ponies and Dülmen ponies with a Przewalski stallion. Later on, Koniks were used massively as breeding stallions on Heck horses, so that they are heavily influenced by that breed, hence the large optic resemblance [8]. The same was the case in the Dülmen pony, which is why the three breeds sometimes are regarded as the Konik group [8]. The Liebenthaler horse can also be regarded as a member of this group.

The Exmoor Pony myth: The Exmoor Pony is a remnant population of a wild horse type or at least a feral type of very original western European horses that once ranged on the entire British island. Several other populations of this primitive British horse type have been intermixed with derived horse breeds, creating the modern British pony breeds such as the Welsh Pony, New Forest Pony, Dartmoor Pony and others. The Exmoor Pony however is the only population that retains a stable, wild horse-like appearance with a brown colour, countershading + white muzzle and a sturdy body. The similarity to other northern ponies such as the Gotland or Faroe Pony, and especially some primitive Iberian breeds such as the Garrano and Pottoka, endorse the hypothesis of a north-western European wild horse/primitive horse type that once was found in this region and is most authentically represented in the form of the Exmoor pony.
The truth behind it: At first, this scenario sounded convincing to me. But what is most important to note first is that equines disappeared from the archaezoological record of the British isle at the end of the Mesolithic until domestic horses were introduced by the Celts [3,4,5]. Therefore, the Exmoor pony cannot be a remnant wild horse population or wild horses with domestic introgression, they have to be of domestic origin. But this alone does not rule out that they are the last representative of a homogeneous, feral and primitive horse population that once ranged across the entire British isle. One argument is their homogeneously small body, comparably short mane, the brown colour with countershading plus white muzzle. However, careful examination suggests that the population at the Exmoor never was homogeneous after all[4]. There are no helpful references prior to the 18th century on the colour variants found in the Exmoor population. Between 1805 and 1809, 81 Exmoor ponies were sold from the moor. Their colour was documented and included black, grey, bay, dun, chestnut and one piebald individual. There is no evidence that purposefully “non-pure” Exmoor ponies were caught in this case. Two illustrations in the Illustrated London News from 1835 clearly show horses with a long mane, their colours are probably implemented to be brown with a white blaze and sock, grey and a black one [4]. A notion by Worthley Axe in the year 1906 is even more revealing: “… the majority of the so-called Exmoors are simply mongrels” [4]. The Acland herd, which made a considerable contribution to the modern Exmoor population, also included a number of greys and blacks in 1900. There is a record that suggests that black Exmoor ponies were selected out because they lacked the expression of the white muzzle, indicating that artificial selection started back then. Furthermore, the Exmoor pony population went through several bottlenecks, the most severe in course of the Second World War. A stud book for the breed was set up in 1921, at first black and grey individuals were tolerated but selected out later. It is thus far more in line with the evidence that artificial selection and genetic bottlenecks created the homogeneous external appearance we see in the modern Exmoor pony, and there is no evidence that it was homogeneous prior to 1906. Most likely the Exmoor pony that is always brown in colour with countershading, a white muzzle and no white markings is an invention of the 20th century [2]. It would therefore also be more parsimonious to assume that other British pony landraces like the Welsh pony, Dartmoor pony or New Forest pony never were homogeneous either. Furthermore, the use of British ponies including the Exmoor pony on Iberian breeds like the Pottoka in the 20th century is well-documented[2], which is at least partially responsible for the optical resemblance. Thus there is no empiric basis for a once feral, primitive free-ranging horse type that looked like the Exmoor and ranged across Great Britain or even whole western Europe.
Also, which is important to note, Exmoor ponies most often display a colour variant that is called seal brown, At. This is an allele that has not been identified in wild horses yet, in contrast to the two wildtype alleles bay A+ and black a [6,7], and therefore most likely is a domestic colour. The black individuals, which have been actively purged from the population, however, would have displayed a wildtype colour variant [6,7].

The Sorraia myth: The Portuguese agriculturist Ruy d’Andrade spotted a herd of strongly striped, free-ranging horses in a remote region in Portugal. He considered them to be a remnant population of the zebro, a strongly striped Iberian wild horse type. He was unable to find them again, but collected a number of farm horses from that region that he considered to be closest to those horses, and started to breed them. This is the modern Sorraia horse, by some even communicated to be identical to the zebro and thus a wild horse.
The truth behind it: The claim that the Sorraia is a surviving Iberian wild horse is flawed by the story itself – d’Andrade did not catch any wild horses and started breeding them, but merely collected farm horses he considered to be reminiscent of the horses he spotted. He collected four local stallions and seven mares, and a Criollo stallion was added later on. Therefore, the Sorraia is a descendant of domestic farm horses. But is it possible that is particularly strongly influenced by an optically identical Iberian type of strongly striped wild horses, the zebro? First of all, there are not any references suggesting a survival of wild horses on Iberia into the beginning of the 20th century. C.H. Smith reports free-ranging horses of black dun and bay dun colour with wild markings that ranged from the Camargue, parts of Spain to the Ardennes, Great Britain and Scandinavia (see here), but also describes them as “sturdy mountain-forest ponies”, and it is not clear if those are wild or feral horses anyway. Regarding the actual zebro, it seems that this population of equines vanished in the 16th century [8] and many authors tend to consider them feral donkeys instead of horses. Indeed a genetic test of the skeleton of the supposedly last zebro turned out to be a donkey [9]. The strongly striped ashy grey colour of wildtype coloured donkeys is very reminiscent of the black dun colour scheme in horses.
The allegedly strong stripe pattern of the Sorraia is often purported as its trademark and an indicator of particular primitiveness that links it to the zebro. However, stripes are part of the bay dun and black dun colour scheme and found in many horse breeds. Therefore, it does not give the Sorraia any special status (see a very strongly striped Hucule here, for example).
Ferus-type wild horses are usually assumed to have been comparably small and sturdy in build. The Sorraia, however, is comparably large (140-150cm at the withers) and lanky. As there is no comprehensive archaezoological record of predomestic wild horse skeletons that has been osteometrically examined (only some single fragmentary specimen) it can neither be proven nor ruled out that Iberian wild horses were larger and more gracile in build than other Eurasian wild horse types, but it would be necessary to have evidence at hand for making such a claim.
More importantly, genetic information has revealed the Sorraia as a domestic horse [10].

I am going to come back to genetic studies more extensively in the next part of the wild horse series 2017. A closer look at the respective history of those three breeds that are often supposed to be wild horse relicts, near-wild horses or recreated wild horses alone is sufficient to show that there is actually nothing of substance that suggests that these three breeds deserve a special status among robust landraces of European domestic horses. But this of course provokes the question which horse breeds are best suited to be used as a substitute for the European wild horse in ecologic restoration. Thus, the upcoming post is on this question as much as on the domestication of the horse in general.

Similar but older posts: 

Literature

[1] Cis van Vuure: On the origin of the Polish Konik and its relation to Dutch nature management. 2014.
[2] Tadeusz Jezierski, Zbigniew Jaworski: Das Polnische Konik. 2008.
[3] Baker, Sue, 2008: Exmoor Ponies: Survival of the Fittest – A natural history.
Baker
[4] Peter Green, 2013: The free-living ponies within the Exmoor National Park: their status, welfare and future. A report to the Exmoor moorland landscape partnership. 
[5] Bunzel-Drüke, Finck, Kämmer, Luick, Reisinger, Riecken, Riedl, Scharf & Zimball: „Wilde Weiden: Praxisleitfaden für Ganzjahresbeweidung in Naturschutz und Landschaftsentwicklung“. 2010
[6] Pruvost et al.: Genotypes of predomestic horses match phenotypes painted in paleolithic works of cave art. 2011
[8] Bunzel-Drüke, Finck, Kämmer, Luick, Reisinger, Riecken, Riedl, Scharf & Zimball: „Wilde Weiden: Praxisleitfaden für Ganzjahresbeweidung in Naturschutz und Landschaftsentwicklung“. 2010
[9] L. Orlando et al.: Revising the recent evolutionary history of Equids using ancient DNA. 2009.
[10]  Lira et al.: Ancient DNA reveals traces of Iberian Neolithic and Bronze Age lineages in modern Iberian horses. 2009