Friday, August 24, 2012

The Sorraia's Prehistoric Relatives / Countering a Historian's Critique

The colour of the lake shore clay
Mouse-dun (grulla) Sorraia filly, Altavida
Ravenseyrie Sorraia Mustang Preserve
Manitoulin Island, Ontario, Canada

Note:  All photos taken by Lynne Gerard unless otherwise noted.  If you find them useful, please feel free to copy and share giving credit to the author/photographer.  Remember, by double-clicking on an image you can view it in a larger format.

Several weeks ago a Portuguese researcher with an interest in rewilding landscapes in the Iberian Peninsula sent me a document written by an historian who serves as a consultant on several such projects underway in Europe.  As the document is specifically meant to discredit the ancestral heritage of the Sorraia horses and provides references to published research articles to support this historian's opinion, the Portuguese researcher asked for me to review the document and provide my thoughts regarding the rather damning claims being asserted within it.

I received this document second hand and at the time of this writing the Portuguese researcher who sent it has not responded to my inquiry to refer to him by name, nor has he provided me the email of the historian so that I might obtain his permission to publish his article here in the Journal of Ravenseyrie.  Nevertheless, since I am aware of several individuals who have received already a copy of this historian's unofficial paper on the Sorraia--and surmise there are other recipients who will take this misinformation as definitive truth--I propose to provide short quotes and summaries of the key points being raised in an attempt to offer a fuller perspective on this very important topic.  Many readers of the Journal of Ravenseyrie will find this lengthy "point and counter point" entry to be not nearly as interesting as stories of the Sorraia horses here and their wilderness living, but I consider it my duty to include the practical as well as the poetic aspects of these primitive horses.  One need not undertake musical theory to be moved by a richly evocative symphony, but such educative explorations (especially when controversy continues) can bring a surprising enhancement to what we already feel an affinity for.  So--I hope followers of the this blog will weave their way through this long exchange as best as can be.

The issues raised by the historian will be identified by a different font and colour than what I will use for my comments.  My comments will be preceded with my initials.  Likewise, any quotes from the historian's supporting sources (and mine) will have a different font and colour.  Hopefully this will assist in clarity and comprehension of all the material being discussed.

[new! 28Aug12  one reader emailed me relaying that there was a little difficulty still in distinguishing what is attributed to the historian and what is my commentary, so I am going one step further and putting anything that isn't my own commentary in a different colour and font with highlighting.]  

-My comments will look like this:  Times font in black

-Things attributed to the historian will look like this:  Verdana font in purple with highlight

-Quotes from other sources will look like this:  Courier font in blue-green with highlight

Hope this helps!  I hope I didn't miss or error on the necessary highlighting..if so, let me know.  I will continue to edit as necessary to provide the most accurate data I can and make it as easy to digest as possible, and appreciate assistance from readers--Thank you!]

I am presenting the historian's contentious writings and my commentary to his various points in the manner which he delivers them. First there is his general discredidation, followed by a more academic approach wherein he provides scientific references and lastly he offers some final words that I felt deserved review and commentary as well.

Starting then from the beginning issues raised...

The historian says "no substantial evidence has ever been delivered" for the Sorraia "story".

LG:  Ample evidence has been given (D'Andrade, Oelke, Oom, Gerard) which has been often times misunderstood or other times deemed unacceptable because researchers have determined without definitive fossil or genetic material identical to the present day Sorraia, no other proofs of prehistoric ancestry can be considered.

The fossil record for all European horses who lived during the Late Pleistocene and early Holocene is most of the time limited to isolated bones and teeth.  Rarely are complete skeletal remains discovered intact and it is also rare to find full skulls and, specifically, the remains of ice age caballine horses in the Iberian Peninsula are especially scanty, more so in the southwestern region where the ancestors of today's Sorraia horses had their last foothold.  The words of archaeologist Marsha Levine fit well here:

"Moreover, it should not be assumed that the absence of horses from archaeological assemblages is evidence that they were not present.  They might well have been available, but not hunted for either logistic or cultural reasons."  (Levine, mtDNA and horse domestication:  the archaeologist's cut 2002)

Advances in genetic testing provide interesting and additional data, but unfortunately (especially in the case of the Sorraia) sample materials being used in testing are of a limited nature (typically from 18 individuals who are reduced to just two maternal lineages) and is often compared to results obtained from fossil specimens that are themselves marginal, potentially misidentified and representative of equids from a different habitat region altogether.  As such, conclusions and statements made from scientific research are frequently coupled with cautionary notes.  All must be kept in perspective and reviewed alongside historical accounts as well as modern day examples.  Perhaps no one knows this more than the preeminent specialist in equid archaeology, Véra Eisenmann (whom our historian repeatedly uses to verify his points).   Eisenmann's multiplicity of published works typically are injected with reminders that the limitations of these fossils certainly temper what definitive conclusions can be made, for example:

"The story of equids is still tantalizing and frustrating after 150 years of study, during which time it has attracted scientists and fed their debates. The fossil material is rich enough to raise hopes, but not rich enough to fulfill them. The variety and abundance of data exercises such constraints on interpretations that they rarely remain simple and conclusive. " (Equus: an evolution without lineages? 2004)

Seeming almost like apparitions, the Sorraia stallion, Altamiro, and the Sorraia Mustang mares, Zorita and Bella each have coat colours of varying shades of mouse-gey that are perfectly suited to their environment.
Ravenseyrie Sorraia Mustang Preserve
 Manitoulin Island, Ontario Canada

The historian discredits Dr. Ruy d'Andrade's discovery and conservation of the Sorraia, citing the following points of contention:

--Archeology during d'Andrade's day was in its infancy or in some regions non-existent
--knowledge of "true wild horses" was rudimentary
--d'Andrade built a breed based on "stylized sketches from cave paintings"

LG:  Dr. Ruy d'Andrade published his first studies of wild Iberian horses in the late 1920's and was much more thorough in his investigations than relying upon prehistoric art as the only clues to what wild horses were like and he was quite adept at carrying out appropriate field study and documentation.  One need only research the man himself a bit to learn what a huge figure he was and what a legacy he left not only in the field of hippology, but in archeology as well, to name just a few interests he invested his time in.  Archeological study during d'Andrade's era already had a rich history to draw upon (as the field is reported to have developed its scientific structure in the 1700's).  It probably is a habit of many modern scientists to belittle the work of those who came before.  Regardless of what advances have been made in the various fields of science, it is worthwhile to appreciate much of the findings from earlier research.

 Examples of some of the equine cranial studies undertaken Dr. Ruy d'Andrade and presented to the Zoological Congress in Lisbon back in 1935 and reproduced in the Studbook da Raca Sorraia 

In 1939 d'Andrade published an exhaustive paper titled Equideos do fim do Paleolitico e inicio do Neolitico, wherein he provides detailed drawings and measurements of skulls, limb bones and teeth of Equus asinus and what he suspected might be Equus caballus Celticus--his data, reproduced in the Studbook da Raca Sorraia is handled in a manner of which someone like Vera Eisenmann would applaud...fastidious measurements and drawings , very scientifically presented, along with very good quality photos.  Dr. d'Andrade was no amateur as can be attested in that he presented his findings and theories on the Sorraia at the 13th International Zoological Congress in Lisbon in 1935.  In his later work, *O cavalo do Sorraia* 1945, d'Andrade more comprehensively presents his evidence for the prehistoric heritage of the Sorraia, provides a thorough geographic sketch of the habitat, defines the differences between the horses that developed in the north, versus those from the south and includes photos of a complete skull of one of the founding stallions along with measurements, sketches and photographs of living specimens.  I have to rely upon those portions of *O cavalo do Sorraia* which have been translated into English.  Those who can read the original Portuguese ought to do so and stand up for the quality of d'Andrade's work.  It is sound evidence, and if some of it has not proven theories in precisely the manner which d'Andrade had anticipated, this is no reason to completely discard such a comprehensive body of work.  Researchers would do well to revive his skeletal comparisons with new fossils that may have been recovered since d'Andrade's day, explore more deeply historical records that offer insights into the wild fauna of the region, appraise surviving specimens for their capacity for continued wilderness fitness (proven in the Vale de Zebro and Ravenseyrie preservation projects) and not rely solely upon genetic comparisons.

The historian states that Dr. Ruy d'Andrade's sighting of wild horses while hunting in Coruche during the 1920's occurred in "one of the most densely populated areas of Portugal" and believes it is "inconceivable" they represented a pure wild horse.

LG:  Not anywhere have I found researchers claiming the Sorraia to be free of domesticated genes or that it is a totally pure wild form.  All describe the Sorraia as a surviving remnant worthy of recovery and consolidation of the remaining primitive characteristics (genotype and phenotype) which persisted despite whatever "contamination" the original wild type suffered due to various domestic influences.  Back in 2010 when I was doing some research on these matters I had actually wondered the same thing myself (whether such uninhabited regions existed) and asked Constança d'Andrade de Oliveira de Sousa (Dr. Ruy d'Andrade's great-granddaughter) if she could help clarify.  She actually took great interest in helping my research and did some historic digging into old records providing me with some census data from Coruche county.  In 1920 the population density of that region was recorded as 13.1 hab/km2.  In 1911 that region was reported to be 100% rural.  In 1940 it was reported to be 87.1% rural.  The sesmaria land d'Andrade was hunting on in 1920 was in the marginal wetlands of the Sorraia Valley, he described this region as "totally uninhabited".  This was prior to the damming of the Sorraia River which saw an expansion of agricultural pursuits, which further reduced the type of habitat where wild horses would have been able to survive.  The region where d'Andrade first saw a wild-living group of these horses was indeed remote enough and inaccessible enough for herds of horses to eek out an existence undetected, or ignored by humans.

The historian states that Dr. Ruy d'Andrade thought the horses he saw were wild because they looked like the Tarpan.

LG:  D'Andrade's description did not use the word "tarpan" but wrote "They looked totally wild or primitive, as if they were a type of zebra or hemiones." (Arsenio Raposo Cordeiro, translation)

 Legado and Zorita
Ravenseyrie Sorraia Mustang Preserve, Manitoulin Island, Ontario Canada

The historian has further comments regarding the Tarpan and whatever comparisons one might wish to see with the Sorraia:

--sums up the Gmelini description of the Tarpan as "a very small stocky pony from the steppes of the Ukraine, with uni-colored short curly manes, uni-colored black tail" which he states according to personal communications with Véra Eisenmann were confirmed by her own studies

--states the Sorraia doesn't look like the "original description of the Tarpan" and "ecoregion, climate, vegetation etcetera" of the Ukrainian steppes "can not that easily be transported to a horse breed from Portugal"

--states that the Tarpan's status as a "truly wild species" is doubtful because it consisted of escaped domestic stock, "confirmed by statements made by the Cossack people in the 19th century"

LG:  The recorded descriptions of the Tarpan are mostly from the 18th and 19th centuries and often provide ambiguous and/or conflicting details, yet within them we note enough characteristics shared with the Sorraia (some descriptions more than others) to give one pause and not completely discount obvious connections.  Also the skeletal remains that have been examined  are limited to those remains that reside in Russia (one complete skeleton and one mandible).  Unfortunately by the time the last Tarpans became of interest to zoologists...the remaining sightings and specimens were likely no longer "pure".  Nevertheless, it would be appropriate to consider that of those wild horses throughout various regions of Europe and Asia, many would have morphological adaptations related to the habitats were they lived.  (Later we will read a quote mentioning the range of the Tarpan extended to Spain and France.)  Whether the Tarpans were feral or truly wild has not been proven one way or the other.  Later in this article, I will quote from the variety of sources that gave descriptions of the Tarpan (including Eisenmann's study) and also relay sources that show Cossacks knew well the difference between feral and true wild horses, even having different names to describe them, reserving the word "tarpan" for the wild horses and referring to the feral horses as "takja" or "muzin".

The historian discounts Dr. Ruy d'Andrade's claim that the Sorraia is a remnant of Pleistocene horses because the last surviving specimens he chose to preserve (saving them from sure extinction) belonged to regional farmers and were pastured with domestic stock.  The historian intimates that d'Andrade's selection was based on phenotype with no regard for the genotype or ecotype that would directly link to prehistoric horses.

LG:  Dr. d'Andrade did a laudable job selecting his founders working with what he knew of their phenotype, ecotype and surmised genotype, providing the best he could for conservation efforts considering how limited in numbers the remaining specimens were. With only rare exceptions (mostly the occasional incidence of white markings or -in the past- individuals who had the grey gene mutation) the offspring are notably homogeneous.  Whatever adulteration to the wild type by domestic genes was remarkably rarely passed on to the offspring, rather, it is more typical that the wild traits typified by the Sorraia persist in domestic breeds.  More on this phenomenon later in the article.

The historian considers the later incorporation of an Argentinian Criollo stallion to be a "major warning sign", again intimating Dr. d'Andrade's decision was based on coat colour instead of "proven (genetic) ancestry".

An historic image of a South American Criollo
photos courtesy of Wikimedia Commons

LG:  Considering the Criollo's heritage stems from 16th century Iberian stock, finding an appropriate candidate with Sorraia characteristics would - even today, in my opinion, be a worthy outcross for the Sorraia (like we are doing here with the Sorraia mustangs) to alleviate the severity of the inbreeding.  It was an important experiment for d'Andrade to undertake and took into account the appropriateness of the Criollo's genetic origins as well as its Sorraia-like phenotype.  Dr. d'Andrade's great-granddaughter informed me the experiment was discontinued due to domestic marks that emerged in the offspring.  What this tells us is that this one particular Criollo stallion was not able to assist in consolidation of the wild genetics, but this does not mean that all Sorraia-type horses from South America would prove equally ineffective.  My opinion is that these types of outcrosses should continue to be experimented with.  Fortunately our conservation efforts here at the Ravenseyrie which incorporates Sorraia-type mustang mares crossed with a purebred Sorraia stallion are much more successful and suggest it may be time for other Sorraia breeders to follow our example.

The historian points to Dr. d'Andrade's mention of convex equine profiles as being a primitive marker but mis-represents d'Andrade's words by summing them up thusly:  "The convex profile appeared for the first time on various Iberian coins of the 3rd and 1st centuries B.C. and later on the monument dedicated to Marcus Aurelius."  The historian follows this misconstrued summary of d'Andrade's words by stating that given the known timeline "you are not even near to prehistoric horses".

Convex profiles of the Sorraia stallion, Altamiro, and his 2012 colt, Fidalgo (out of Belina)

LG:  What d'Andrade wrote was "...the horse of convex profile and large size appeared for the first time on various Iberian coins of the 3rd and 1st centuries B.C. and later on the monument to Marcus Aurelius (2nd century A.D.).  This shows that the large horse evolved only later and hitherto was not the norm."  He was not saying this was the first time the convex profile was detected...just the first time it was detected on larger horses.  Dr. d'Andrade considered the convex profile to have been first recorded much earlier.  Arsenio Raposo Cordeiro writes about this:  "The rock engravings in the cave of Escoural, discovered as late as 1963, not only suggest the possibility of there still being much to be uncovered about this subject in Portugal; they also demonstrate that even before the last Ice Age, a swift horse, extremely similar to the Sorraia breed and present day Lusitano, already existed in this Atlantic region...The beautiful shape of the mare's head, depicted at Escoural as being long, lean, dry and slightly convex, tapering to a slender point at the muzzle, the whole profile in perfect harmony with the long line of the neck which arches to protect the colt, provides us with the most significant information about the consistent, hereditary characteristics of the Lusitano horse." (pg. 1 *Lusitano Horse / Son of the Wind 1997, Lisboa)

To conclude his opening dismissal of the Sorraia, the historian feels the slender jaws of the Sorraia cannot deal with poor and rough forage.  Once again, the historian wrote that Véra Eisenmann agrees with his assessment, stating that the "slender jaws of the Sorraia (and other horses) do not resemble that of prehistoric horses, which has thick jaws with teeth that were deeply rooted, just like present day Przewalski and Exmoor pony."

LG:  Historically the Sorraia thrived on poor and rough forage, and are still capable of this today.  I have photos and video clips of Sorraias in the Vale de Zebro preserve easily crunching and consuming acorns from Cork Oak trees (Quercus suber).

Despite extreme drought conditions, this Sorraia mare is in robust condition as she forages for acorns beneath an aged Cork Oak tree in the Vale de Zebro preserve in Portugal.

Likewise my own Sorraias browse readily on shrubs, twigs, bark, roots, woody plant stems, etc. in addition to grazing the grasslands.  Not only can these horses "deal with poor and rough forage" they often seek it out to supplement their diets.  Most modern caballine feral horses (who always seem to be pushed by humans to extreme, low quality environments) likewise demonstrate a capacity to manage subsisting on marginal grazing and browsing habitats.

 The Sorraia horses of Ravenseyrie supplement their diets by browsing on a variety of trees during all times of the year as seen in the photos above and below.

If there is documentation where Vera Eisenmann has published her measurements on the jaws of the Sorraia, I would very much like to read it.  I suspect Prof. Eisenmann has measured a great number of slender jawed prehistoric horses, whether they were from European specimens that existed during the Pleistocene or not, I will have to research to find out.  At any rate, M. Eisenmann is considered the premier authority on equine fossils, and her determinations have to be considered for sure, but alongside other evidences and with the understanding of how few full skulls suspected to belong to Equus ferus are available for comparative study, before making sweeping judgments.  We do know, of course, that she examined the Tarpan specimens from Russia and in her paper titled, "Discriminating Equus skulls:  the Franck's Index and the new Palatal Index", she wrote this about the Tarpan specimens:

"E. ferus is based on the recently extinct Russian Tarpan.  Osteologically it is represented by two skulls of two castrated males (the Moscow specimens S 94535 has extremely worn upper cheek teeth and lacks the mandible) and the associated skeleton of one of them (LG521).  The material is far from satisfactory (Gromova 1959, 1965).  Both skulls mainly differ from E. przewalskii by more caballine Franck's and Palatal indices.  On LG 521, the protocones are very short, like in E. missi.  The skeleton has exceptionally short metapodials and short first posterior phalanges.  Proportions and size are close to an Island pony.  To my knowledge there is no clear evidence of a similar conformation anywhere in the Late Pleistocene (no associated bones at Missy).  All the common late Pleistocene horses (E. latipes, E. gallicus, etc.) have completely different conformations and may not be reffered to E. ferus s.str.  However some authors use E. ferus in the meaning of 'wild caballine' to replace the 'domestic' E. caballus (although both Tarpans having been castrated, their status of 'wild' is somehow impaired)."

(I would like to point out that the castration of a wild equid doesn't infer that such a creature is no longer considered a representative of a wild specimen--that they were extracted from the wild and forced to accept certain human manipulation does not erase what value they represent as repositories of wild genetic material.)

The skeletal remains of our treasured mare, Ciente (Kiger Mustang of Sorraia type), ever beautiful

It has become pretty well accepted through the modern genetic analysis of fossil remains that there was more than one ancestral horse type, and certainly in his day Ewart seems to have been studying distinctions between the heavier jawed horses and those with more slender ones.  (1909  The Possible Ancestors of the Horses living under Domestication)  One of the specimens Dr. Ruy d'Andrade measured (photographs and drawings of which are included in the Studbook da raca Sorraia) shows a subconvex profile and slender jaw, which is listed as belonging to a Paleolithic horse fossil recovered from "de Vila Nova da Rainha, Matao".  Dr. d'Andrade determined the Sorraia (and Lusitano) horses had dentition more like Equus stenonis.  He wrote:

"Teeth, as we know, being among the most common archaeological finds, are the surest and most characteristic element when classifying animals.  On examining them we can note that those of the E. Stenonis are distinguishable for having simple estilideon; protocone without talon and off centre; an advanced and developed hipocone; prega cabalina.  Identical characters pertain to the horses of the south of the Peninsula and, among them, the Sorraias.  This means that the difference between the Sorraia Horse and the 'garrano' of the north lies in their dental characteristcs, and for this reason we believe it to belong to a different variety of the species E.c.caballus.  And in truth these teeth reoccur in both the Sorraia and the Andalucian horses of the purer breeds, with convex-shaped head profiles and the eliptical orbit shortened at the back, the base of the cranium at an angle to the forehead, the occipito-incisiva line below the orbit, the maxilla having a short ascending line at an obtuse angle."

Detail of the dentition of the lower jaw of Ciente, a 6 year old Kiger Mustang mare of Sorraia type

 Cranium of Ciente, a 6 year old Kiger Mustang of Sorraia type, what revelations might such skeletal remains have?

At the time of this writing, there is (available for purchase!) a slender-jawed complete mandible of a late Pleistocene horse from Poland as well as less complete, but equally slender specimens from Russia depicted here:
Of the specimen from Poland, this is written:   

"Despite what one might think, seldom are Ice Age horse fossils from the European continent available for purchase.  Horse fossils from North America and Russia (Siberia) are common but true European Pleistocene deposits provide few horse fossils for the market.  As a perfect compliment to European Ice Age fossil collections, this interesting specimen makes for an uncommon addition."

Scientists can look at these things and form all manner of conclusions and I happily leave that task to them.  For myself, what I can offer is living proof wild living slender jawed horses can manage to browse and graze and thrive upon a wide variety of rough forage as well as their heavier jawed counterparts.

The historian concludes his first series of disparaging statements by reiterating how wrong d'Andrade was about his belief in the prehistoric ancestry of the Sorraia and suggests d'Andrade's work is faulty due to his lack of expertise in the field of archaeozoology, which was anyhow "still virtually nonexistent in his time".

LG:  In addition to finding the historian's discredidation of the work of Dr. Ruy d'Andrade to be based on numerous mis-representations of d'Andrade's legacy, my opinion is that there are enough retained primitive and true wild horse characteristics present in the Sorraia and sufficient enough provocative historic evidence indicative of a wild heritage to prompt researchers to keep an open mind regarding its link to prehistoric horses.  When contemplating which horse types to "seed" rewilding conservation projects, the Sorraia, along with other wild horse types like the Exmoor, Konik, Przewalski, Garrano, etc. have already proven well suited to the task.  All surviving wild-type horses with potential links to prehistoric caballines should be given an opportunity to live truly wild again in habitats similar to those where remaining specimens (whether "pure" or not - for  "purity" cannot be guaranteed) have survived.  This would best reflect the multiple horse origins theories that present day research indicates.  Conservation projects should not be focused on just one wild horse type, rather for the protection of biodiversity, each variant form should be valued, protected and re-introduced to suitable landscapes.

 Sorraia stallion, Altamiro, damp from a summer rain, dozes nearby his harem of mares

Let's now take a look at the second portion of the historian's paper aimed at discrediting the Sorraia.

In this next segment, the historian wrote that he is presenting "some hard evidence 'against' the Sorraia" and cites seven previously published research papers along with his conclusions of what these papers mean.  For the purposes of my countering what the historian has written, I am presenting the title of each research paper and a short synopsis of the key points the historian has drawn from them, followed by my own comments. 

#1:  Jansen et al., Mitochondrial DNA and the origins of the domestic horse, 2002

-based on this paper, the historian states severe inbreeding points to the Sorraia being a breed and not wild, for "genetic diversity is the trademark of a truly wild species"

LG:  When a species suffers from various factors that affect the number of animals available for reproduction severe inbreeding and genetic erosion occur.  This has happened and continues to happen to "truly wild species".  (O'Brian 1996 ), (Keller 2002)  The degree of inbreeding within the Sorraia population, does not, on its own, disqualify it as being "truly wild".  It is  accepted, of course, that some amount of genetic adulteration occurred within the original wild stock of the Sorraian ancestors over the centuries due to the wild and domestic populations sometimes overlapping territories and or deliberate cross-breeding by humans, but (when appraising phenotype and behaviour) the strength of the wild genetics appears to mostly "trump" whatever domestic traits might be genetically passed on, especially if these horses are allowed to live as wild animals in an appropriate habitat.  As such the characteristics of the European wild horse retained within the Sorraia horses reflect its wild heritage and are not the product of manmade selection.

Sorraia and Sorraia Mustang horses have retained many characteristics of the Tarpan (here we show the Sorraia stallion, Altamiro) and may represent a regional variant of the extinct wild horse, Equus ferus gmelini

We find this phenomenon has been already discussed in the historic record of the Tarpan as relayed by Heptner, Nasimovich and Bannikov (1961 Mammals of the Soviet Union) who write:

--"Feralization does not impart a tarpan-like resemblance."

--"In general, it should be emphasized that we have exaggerated views on the admixture of blood of domestic horses in that of tarpan. In fact, such admixture has been very insignificant, if indeed it has occurred at all. The reverse phenomenon, i.e., the effect of tarpan on local breeds of domestic horses, could at places be notable. For details, see Heptner, 1955"

As will be noted later, descriptions of the Tarpan (Equus ferus gmelini) vary in some aspects but also share distinct commonalities, which one also finds in the Sorraia and Sorraia Mustang, suggesting the possibility it is a regional variant of the Tarpan.  It should be relayed, also that it is not a matter of shared coat colours, but also of certain morphological and behavioural elements and overall fitness adapted to survival in wilderness environments.

 Sorraia Mustang mares, Bella and Zorita at the Ravenseyrie Sorraia Mustang Preserve 
Manitoulin Island, Ontario Canada

--The historian's second point drawn from research paper #1 is that the Sorraia clusters neither with the Pleistocene or the Lusitano horse groups

LG:  The Sorraia presents quite a number of difficulties for conclusive determinations of genetic relatedness, due to its low variability, which is mentioned time and again in the various research papers.  In paper #1 it is demonstrated that the Sorraia does not genetically cluster with the Iberian group of horses but rather with the more distant, older A1 cluster:

All 18 sampled Sorraias have either of two A1 mtDNA types (61% A1 root type, 39% ancestral JSO41 type), which are quite unrelated to the D1 type predominant in the other Iberians. 

These results (from Jansen et al., 2002) were confirmed in another, more recent study carried out by Georgescu et al., in a paper titled, Phylogenetic relationships of the Hucul horse from Romania inferred from mitochondrial D-loop variation., published in 2011.  Here are some excerpts from their findings:
"Certain unique haplotypes were also present in the case of the Sorraia breed, a possible descendent of the Tarpan breed fro the Western European region."

"The separation of clusters in the network may represent additional proof of the evolution in particular conditions of these ancient horse breeds.  Thus, Sorraia horses originated from a group of 7 mares and 4 stallions obtained by R. d'Andrade near Coruche, Portugal around 1930, after he had seen a phenotypically identical wild population there in 1920 (Jansen et al., 2002).  Concurrently, just as in the case of the phylogenetic tree, the Przewalksi and Mongolian wild hrse breeds formed as we expected, a totally separate cluster.  The anaylsis of the phylogentic network led to observations regarding the clear separation of several breed groups, including the Sorraia, Przewalski, Mongolian wild horse and Hucul breeds."

We will see later that these assorted genetic research papers often reveal conflicting results which are possibly the result of the limited sample base, the unavoidable presence of both wild and domestic lineages present in the founder group used in the conservation effort and the type of markers used in identifying genetic relatedness.  Nevertheless, the overall impression one gains is that within these Sorraia horses there persists an ancestral genotype that needs continues consolidation and preservation.

[02Sept12 - It has been brought to my attention by a private email that my attempts to present a more rounded picture of what these various genetic studies mean in relation to the Sorraia horses is creating greater confusion rather than making these issues clearer, due to my lack of understanding of this field of science.  This means my perception of what these studies reveal is as misunderstood as that of the historian.  Until such time as someone with sound knowledge provides a better explanation of the genetic studies, I am withdrawing my own attempts to place them in appropriate context. --LG]  

#2:  Ludwig et al., Coat Color Variation at the Beginning of Horse Domestication, 2009
This paper has prompted the historian to draw novel conclusions and share his opinion on no small number of colour related issues:

-- prehistoric horses were bay/brown and black
--dun colour cannot be verified
--grulla/mouse-grey colour is not the "most natural colour" of the Sorraia, that such an idea  [that the Sorraia originated from grullo progenitors] originated with Hardy Oelke who believes Sorraias have a relation to Tarpans
--mouse-coloured references in historic sources meant brown not grey
--states that bay/brown colours have emerged in individual at the Sorraia reserve at Alpiarca, Portugal
--states the characteristics of the Sorraia are "not stable" and are "not a synthetic breed"
--misinterprets what coat colours the Portuguese words "baia" and "rato" stand for
--states that Dr. Ruy d'Andrade selected bay and bay dun horses as founders for his conservation of the Sorraia horses
--states that Dr. Michael Schaefer remarked that the Sorraias he imported to Germany were mostly bay or yellowish coloured
--states that the preservation of the Tarpan within the Polish Konik program initiated by Vetulani is based on a mis-translation of "the old Polish source", which according to the historian should have been "mouse-coated" and not "mouse-grey"
--states the description of the Konik horse is "blue-dun, as opposed to mouse-dun, which has some greyish undertones, but isn't grey itself"
--states that Vetulani started with bay and black horses
--states that the "actual colors of the Tarpan more resemble that of the Fjord horse and Przewalski horse" and that "nowhere is the tarpan described as 'grey'"

LG:  The perception given by the historian regarding selection for mouse-grey is quite conflicting and is providing a characterization of the colour that I've have not yet come upon in any other research.  It seems that the historian is making a distinction between "blue dun", "mouse-dun", "mouse-coated" and "mouse-grey", though my understanding is that these are the same basic black-base-dun-modified wild colour coats that are expressed in varying hues and which are not at all the same type of grey colour the greying gene puts on horses, but shares some commonalities with bay-base-dun-modified wild colour pelages.  I can see in my own horses their "mouse-grey" coat colours, depending on the time of year and the way the light hits them, at times is the same colour of rocks, tree bark, dried grass, shadows in the forest or clay on the beach.  At times the lighter grullas look nearly white and the darker grullas look tan or fact sometimes these black-based-dun horses look very much like bay-based-dun horses!

Mouse-grey mares and foals, above and below, illustrate how their wild coat colour changes hue depending on the time of year and the quality of the light.
Ravenseyrie Sorraia Mustang Preserve, Manitoulin Island, Ontario, Canada

I am not sure the historian is interpreting the research paper he cited correctly to draw the conclusion that prehistoric horses only had a "bay/brown color and also a black color".  As a simple layperson, having no affiliation with academic or scientific institutions, some research papers are not available for me to access the full version--which is the case with this Ludwig et al., 2009  paper the historian is referring to.  I have however read a later paper by some of the same authors (Ludwig included):  Pruvost, et. al, 2011, Genotypes of predomestic horses match phenotypes painted in Paleolithic works of cave art

In this paper, the two dun colours (bay dun and black dun) are discussed and images of Przewalski's horse and the Konik are shown as examples of the wild coat colour depicted in the cave paintings.  Some excerpts from that paper:

"Recently, studies of both maternal (mtDNA) (24–25) and paternal lineages (Y chromosomal DNA) (26) found that the Przewalski horse displays DNA haplotypes not present in modern or ancient domestic horses, suggesting that Przewalski horses are not directly ancestral to modern domestic horses. However, independently of its taxonomic status, several lines of evidence suggest that the bay phenotype of the Przewalski horse represents an ancestral character. Firstly, several wild ass species, which undoubtedly represent wild equids, also show a bay–dun phenotype; and secondly, horses of this phenotype are depicted in remarkable detail in Paleolithic cave paintings (e.g., in Chauvet). Whereas black or black–dun and leopard spotted phenotypes also occurred at measurable frequencies in Pleistocene and Copper Age wild horses, as shown by both contemporary depictions and our genotyping results, their absence in modern Przewalski's horses is probably explained by the severe population bottleneck that they have undergone (27), possibly in combination with the Asian origin of these horses, where LP seems to have been rarer, if not entirely absent."


"Predomestic horses inhabited, in vast numbers, large areas of Eurasia, and some extant species that still occupy a similarly large area, such as gray wolves, are also found in different color morphs. It is therefore not entirely surprising that not all wild horses shared the bay–dun or black–dun phenotypes."

This paper also includes an image comparing photos of present day examples of these coat colours to the images in cave paintings of particular interest to our discussion are the presence of a "Bay–dun Przewalski's horse", "a black–dun Konik with winter coat" and a "black–dun Konik with summer coat (same genotype)"

From the many examples provided below we will see that mouse-grey, mouse-dun, blue-dun, rato, dosinus, cineras, etc.  all describe the same black/dun colour and these terms were used for European wild horses prior to Vetulani's recovery of the Tarpan in Polish primitive horses (the Konik) and well before Hardy Oelke began to suggest the Sorraia was a remnant of a variant strain of Tarpanoid wild equine that developed in southwestern Iberia.

A large herd of Konik horses at the Oostvaarderplassen nature preserve.
Many in this photo appear to share common phenotypical characteristics with the Sorraia and Sorraia Mustangs.
(Photos: A. Stuart)

Some historic descriptions of the colour seen in the Sorraia and the Tarpan are:

--From M. Poliakof's monograph on Przewalski's horse (Supposed new Species of Horse from Central Asia published in The Annals and Magazine of Natural History Vol. VIII 1881, Chapter III pages 16-26) we are given the following descriptions attributed to the colour of the Tarpan.  These horses all had dorsal stripes and some had mention of lighter fringe hairs on mane and tail--for our purpose presently, I have focused mainly on the colour hues:
  -Gmelin, "the colour is invariably that of the mouse, with an ashy shade underneath the belly, whilst the legs, from the knee downwards, are black" ("Reise durch Russland in den Jahren 1768-1769', vol. i. p. 44)
  -Pallas, "Plerique sunt colore griseo-fusco val pallido, juba, loro dorsi, caudaque fuscis, rostro albido, circa os nigricante." (Zoographia R. -As. i. p. 260) [no free online translator sufficiently tackles this, but loosely one can find such translations for the words as grey, dusky, dark and pale as indicative of grullo shades, with a dorsal stripe, though the white around a black mouth is curious, not necessarily a "mealy mouth" description but perhaps, which on Koniks and Sorraias one only sees in very young foals and is soon rubbed off revealing a black muzzle. -LG] [new!  one of our readers offered this translation of Pallas' Latin sentence:  "most of them are grayish brown going paler, a stripe on the back and a brown tail, a lighter face towards the mouth becoming black" --LG]

 Newborn Sorraia foal, Fidalgo appears here to have the "mealy"mouth one associates with E. prezewalski, but the whitish hair is soon rubbed off revealing a dark muzzle, as we have already seen in the photo of him with his sire inserted earlier in this article.

   -Rytchkof, "colour dun or bluish, other shades exceptional" (Topography of Orenburg, pt. i. p. 290 St. Petersburg, 1762)  

 Sorraias at Ravenseryie, illustrate varying hues of mouse-dun, sometimes with yellow or brown, blue or grey tones depending on the time of year and the quality of the light.

Poliakof (who certainly had an agenda to pushPrzewalski's horse into the limelight as the last of the purely wild horses and may therefore have a prejudiced opinion) writes further about the descriptions of colour given by Gmelin, Pallas and Rytchkof:   

"From these descriptions of the tarpan or wild horse by Gmelin and Pallas, it is evident they were unacquainted with Equus Przewalksii; and Rytchkof had perhaps only accidentally heard of it when he mentioned a dun colour (lutescens).  As to tarpans of blue (caerulescens) and other colours mentioned by Rytchkof, they were such as had probably resumed a feral state in the same way as those described by Gmelin and Pallas."  

Given that Rytchkof was not just "passing through" but lived in the Orenburg region for 43 years, conducting steadfast research on the fauna of mammals in that and adjacent provinces, one can feel confident he, like the Cossacks Groves tells us about later, knew well the difference between feralized domestic breeds and true wild horses.

-St. Isidore  -Prior to those intrepid 18th century men exploring far away Eurasia landscapes we find an earlier (perhaps the first?) documentation of grullo wild horses in Iberia, given to us by St. Isidore, c. early 600's A.D. Etymologiae Book XII: de animalibus Chapt. 44 De Equo:  "Dosina autem dictus, quod sit color eius de asino: idem et cinereus. Sunt autem hi de agresti genere orti, quos equiferos dicimus, et proinde ad urbanam dignitatem transire non possunt."  English translation  "The dosinus horse is so called because its color is that of an ass (de asinus); it is also known as ash-colored (cinereus). These originate from wild stock, which we call equi-ferus, and therefore they cannot be used as city horses."  Curiously, after describing the colours most desired (in which dun is on the list), St. Isidore disparaged the various shades of ash-colour, saying they were the worst:  "reliquus varius color vel cinereus deterrimus" 

In wintertime, the mouse-grey (grulla) colour of the Ravenseyrie Sorraias exhibits a much lighter, "ash-coloured" hue

-Berenger, 1771 The History and Art of Horsemanship:  "The Tarpans are a kind of wild-horse, in the desert, east of the river Yaik.  They are of a middling size, roundish, short, generally a blueish-grey colour, with big heads and ewe-necked."

-Darwin 1868 The Variation of Animals and Plants Under Domestication: "...the term dun-coloured is vague, and includes three groups of colour, viz. that between cream-colour and reddish-brown, which graduates into light-bay or light-chesnut--this, I believe, is often called fallow-dun; secondly, leaden or slate-colour or mouse-dun, which graduates into an ash-colour; and, lastly, dark-dun, between brown and black...With respect to the primitive colour of the horse having been dun, Colonel Hamilton Smith[136] has collected a large body of evidence showing that this tint was common in the East as far back as the time of Alexander, and that the wild horses of Western Asia and Eastern Europe now are, or recently were, of various shades of dun. It seems that not very long ago a wild breed of dun-coloured horses with a spinal stripe was preserved in the royal parks in Prussia. I hear from Hungary that the inhabitants of that country look at the duns with a spinal stripe as the aboriginal stock, and so it is in Norway."

 These young Sorraia horses appear dun coloured in the late day sun, but are acutally mouse-grey (grulla).  
Ravenseyrie Sorraia Mustang Preserve, Manitoulin Island, Ontario, Canada

-Harper 1945 Extinct and vanishing mammals of the Old WorldSpeaking of Eruasian wild horses Harper relays the words of Antonius, "Only two, or at the utmost three, of the many local and geological races have survived until our days.  The first of these was a mouse-dun horse, which Albertus Magnus, the great interpreter of Aristotle, means when he calls the colour of the wild horse "cinereus," i.e., ash-coloured."  Harper later talks about the wild horses captured by Gmelin, "After him the author named these horses scientifically 'Equus gmelini' but perhaps there is an earlier name:  Equus silvestris v. Brincken, dedicated to the mouse-dun wild horses of Poland, surviving in the forest of Bialowieze until the middle of the eighteenth century and in another game park until 1812."  Harper retraces the Tarpan to even earlier times, "In the well-known forest of Bialowieza, Poland, the 'Tarpanis' were hunted as royal game in 1409". 

Grulla (mouse-dun) Sorraia youngsters looking quite lavender in the morning sun
Ravenseyrie Sorraia Mustang Preserve, Manitoulin Island, Ontario, Canada

-Groves 1994   The Przewalski Horse:  Morphology, Habitat and Taxonomy    

Groves provides a good synopsis of the conflicting descriptions of the Tarpan, but makes it pretty clear their coat colours were various hues of mouse dun, those that were creamy/yellow and those that were ashy/grey.  He also relays that the Cossacks knew the difference between feral and true wild horses:

 "Smith (1841) provided a wealth of interesting information on wild horses, admittedly second-hand from Tatars and Cossacks whom he had interviewed in 1814, but in the main quite consistent with other evidence. Referring to true wild horses as tarpan and tarpani, to distinguish them from feral horses (takja or muzin)"   

See also Hamilton-Smith's book, The Natural History of Horses 1841 and Lydekker's book, The Horse and its Relatives 1912 for further descriptions (both sometimes tangled, yet useful) of the Tarpan.

Mingling elementals of the Ravenseyrie Sorraia Mustang Preserve
Manitoulin Island, Ontario, Canada

Interestingly, it is the professional naturalists passing through on expeditions to Eurasia who typically believe the tarpans were feral and not representative of truly wild horses (most of which who did not necessarily "know" horses as well as insects and plant life).  It is the people living among the fauna in these landscapes day in and day out for centuries who, in my opinion, would have the more reliable determination regarding differences between feral (muzin) and wild (tarpan) horses, so I do not so readily dismiss the anecdotal information given by the Cossacks in this regard.  In fact in 1882 it is reported "there was a total of 1,300,000 Cossack horses broken to saddle or shaft, and a further 1,200,00 feral horses in the Cossack steppe herds." (Seaton 1972 / The Cossacks)  Seaton tells about the types of horses these Cossack militia were mounted on:  "Whereas part of the Guard squadrons might be mounted on the larger west European horses, the line Cossacks used the horse of the open steppe, a small rough and ugly pony, probably of Mongolian origin and often of the unmistakable 'tarpan' type, dun in colour with erect mane, big and long asinine head, and without a trace of hot blood in them." (Seaton later says these horses were nonetheless hardy and capable of great speed in "short bursts".)  Like Heptner pointed out (in the quote used earlier in this lengthy counterpoint to the historians attempt to dismiss the Sorraia) it is the stamp of the indigenous wild horses on local domestic stock that one notes more frequently and not the other way around.  Despite whatever primitive characteristics one could appraise in the mixed-blood feral steppe herds, apparently they were removed enough from the truly wild bands that the Cossack troops were keen to recognize the difference between these "muzin" and the "tarpans.

--Heptner and Nasimovich 1988 Mammals of the Soviet Union, Vol.1
This book offers a significant amount of information on the Tarpan and (right or wrong) defines three "subspecies":  1. Southern Russian tarpan, E.p. gmelini, Antonius 1912, 2.  Forest tarpan, E. p. silvaticus, Vetulani 1928 and 3.  Dzhungarian or eastern tarpan, E. p. przewalski, Poljakov 1881.   E. p. przewalski is described in the manner one would describe the reintroduced wild Asiatic horses of today saying their color was "yellow with a rusty tinge and tip of muzzle whiteish", while the descriptions E. p. gmelini and E. p. silvaticus are "dark gray" (which earlier in the general overview of the tarpans the Heptner distinguishes dark gray as "mouse").  The text offers descriptions of the variations of the  colours covering pretty much every hue and seasonal alterations that can be seen in the Przewalski's horses, the Konik and the Sorraia--not surprising since according to Heptner:

"Outside the Soviet Union tarpan were quite common over much of western Europe. Commencing from ancient times up to the nineteenth century one finds mention of their presence in Poland, several regions of Germany, including East Germany, Denmark and the Danish islands, Belgium, France, the Iberian Peninsula, Switzerland, and several other countries." [emphasis is mine. -LG]

The sampling of descriptions of what can be said is the "mouse grey" coat colour should be sufficient enough to dispel any further confusion over what colour the Tarpan (E. gmelini, not E. Przewalski) was/is.  Once again we see the shades of Sorraia and the Tarpan have a range of hue variation among individuals and these hues are further altered by age, seasons, perhaps also habitat and they have a long history of being associated with wild horses.  These colours are not the same as solid bay or solid black, and they are not the same as coat colours that are altered by the graying gene, and, according to the literature I have read, the dun/grulla colours are examples of wild colours in mammals.

Two mouse-grey (grullo) Sorraia bachelor stallions blend superbly with their environment in the Vale de Zebro, Portugal

Two solid bay coloured Alter Real bachelor stallions mostly stand in contrast to their environment on the island of Potril da Azambuja, along the River Tejo in Portugal.

 The "wild-bay" colouring of the Exmoor pony, like the "mouse-grey" of the Sorraia is much more capable of blending into the environment as we see in this photo from these members of the Anchor Herd, Winsford Hill, U.K.  If viewed from a bit more distance, these ponies would appear as shadows and shrubs.  (photo courtesy of Wikimedia Commons)

It seems to be quite appropriate to use the terms "mouse-grey" or "rato" to express the varying greyish hues in the Sorraias, in that the way the pigment is distributed in the various hues of wild rodents shares commonalities with the way such pigments are distributed in the different dun colours of horse hair coats--which are distinct from the pigment distribution of other colours, i.e the pigment of this type of wild colour is not uniformly filling the hair shaft as it does in solid colours. 

As mentioned before from personal observation I can vouch for the way the colours of the Sorraias living here at Ravenseyrie provide the capacity for blending in with their environment in the same way that other wild fauna does, i.e. like the field mice, shrews, Mourning Doves, Sandhill Cranes*, Great Blue Heron*, Whitetail Deer, coyotes, wolves, etc. that live in or migrate through our preserve.  (*It is not a coincidence that the colour "grulla" is the Spanish word for "crane".)  The grulla colour is truly an amazing pelage for horses, offering a wide range of camouflage capabilities for horses that made their homes in both grassland and forest habitats.

Two other grulla, mouse-grey coloured inhabitants of Ravenseyrie are the Sandhill Crane and the Canada Goose

For further information on these types of wild colours (though perhaps some of these references are not applicable to horse coat colours - or may be outdated - they nevertheless give insight into the differences in how pigment expresses itself in the hair shafts of wild colours versus domestic colours) see:

the A gene:
"This gene determines the distribution of pigment in the hair. The wild-type allele A produces a phenotype called agouti. Agouti is an overall grayish color with a brindled, or “salt and pepper,” appearance. It is a common color of mammals in nature. The effect is caused by a band of yellow on the otherwise dark hair shaft. In the nonagouti phenotype (determined by the allele a), the yellow band is absent, so there is solid dark pigment throughout (Figure 4-18)."

"The breeding of domestic horses seems to have eliminated the A allele that determines the agouti phenotype, although certain wild relatives of the horse do have this allele. The color that we have called brown in mice is called chestnut in horses, and this phenotype also is recessive to black."


"Although a smoky black that is almost impossible to tell by the eye will show well under the microscope as will a black dun. Below shows photos of a black dun shaft, showing the clear tip and follicle and pigment loading to one side that is consistent."

Ben K. Green, 1983 The Color of Horses, for diagrams of the hair shafts of dun and grullo

The historian's paper on the Sorraia mis-translates Baia and Rato.  There is no doubt among those involved with the Sorraia that these colours represent dun horses.  The word "baia" represents the yellow-dun hues.  The word "rato" represents the greyish, bluish, mousey, ashy colours that all are variants of "black-dun", i.e. the dun gene modifying a black base as opposed to the dun gene modifying a bay base.

 A dun (baia) Sorraia stallion
Herdade Font'Alva, Portugal

Grullo (rato) Sorraia stallions
Herdade Font'Alva, Portugal

The historian's statement regarding Dr. Michael Shaefer's remarks on the colours likewise are being mis-translated--suggesting he imported bays, when he imported baia and rato coloured Sorraias, which were dun and grulla.  All one needs to do is see photos and information in "Studbook da Raca Sorraia" to recognize the horses depicted as founders were not bay horses, but were dun and grulla.  Because some people continue to be confounded by the colour issue, I confirmed this with Constança d'Andrade Oliveira de Sousa, Dr. Ruy d'Andrade's great-granddaughter (who is fluent in both Portuguese and English) back in June of 2010, who replied via email:  "The horses chosen by Dr. Ruy d'Andrade, and which he baptized Sorraias, were only yellow dun and grullo, not grey.  They were horses with the Dun gene."

 A "baia" (dun) mare
Agolada de Baixo, Portugal

 A "rato" (grulla) Sorraia mare
Herdade do Azinhal, Portugal

Regarding the photo from the Reserva Natural do Cavalo do Sorraia which the historian used in his paper to back up his claim "there are also bay/brown colors popping up in the herd", this is misleading.  It is my understanding there are times at the Reserva Natural do Cavalo do Sorraia when lessons and riding demonstrations are being given and Lusitanos and Garranos are sometimes stabled and turned out with the Sorraias, but are not part of the breeding group.  You can see an example here, of a horse stabled at reserve that appears to be a Garrano and not a Sorraia.

#3: Luis et al., Genetic diversity and the relationships of Portuguese and other horse breeds based on protein and microsatellite loci variation, 2007

From this paper the historian has drawn the following conclusions:
-the Sorraia  is not the ancestor of the Lusitano
-the Sorraia is not a prehistoric breed
-the Sorraia's lack of genetic diversity makes it less able to adapt

LG:  As pointed out already, the genetic tests must be placed in perspective and given the limitations of the samples used for the Sorraia, the results do not represent a full picture.

[02Sept12 - It has been brought to my attention by a private email that my attempts to present a more rounded picture of what these various genetic studies mean in relation to the Sorraia horses is creating greater confusion rather than making these issues clearer, due to my lack of understanding of this field of science.  This means my perception of what these studies reveal is as misunderstood as that of the historian.  Until such time as someone with sound knowledge provides a better explanation of the genetic studies, I am withdrawing my own attempts to place them in appropriate context. --LG]  

I am in agreement, however, with the historian's point about the Sorraia's lack of genetic diversity, and believe it is indeed a potential handicap, at least when considering the fertility issues that have begun to undermine the present preservation efforts of the remaining populations in Germany and Portugal.  However, with the careful incorporation of those individuals genetically, morphologically and phenotypically associated with the Sorraia (select strains of North American Mustangs, South American Criollos and individual Lusitanos) crossed with European Sorraias, genetic diversity and vitality is restored (as quickly as one generation as appears to be occurring with the offspring at the Ravenseyrie Sorraia Mustang Preserve).

Two photos of the Sorraia Mustang mare, Ciente and her 2009 colt, Silvestre, by Altamiro, looking more lavender in the lower photo and more yellow tinted in the top photo

#4:  Luis et al., A lost Sorraia maternal lineage found in the Lusitano horse breed, 2006

From this paper, the historian concludes the following:

-the Sorraia has no connection with the Lusitano or any other Iberian horse breed
-Overlap between Sorraia and Lusitano suggests d'Andrade selected founders for the Sorraia preservation from among the Lusitano group

[02Sept12 - It has been brought to my attention by a private email that my attempts to present a more rounded picture of what these various genetic studies mean in relation to the Sorraia horses is creating greater confusion rather than making these issues clearer, due to my lack of understanding of this field of science.  This means my perception of what these studies reveal is as misunderstood as that of the historian.  Until such time as someone with sound knowledge provides a better explanation of the genetic studies, I am withdrawing my own attempts to place them in appropriate context. --LG]

#5:  J. Lira, Ancient DNA reveals traces of Iberian Neolithic and Bronze Age lineages in modern Iberian horses, 2009

From this paper the historian drew the following conclusions:
-no relationship found between ancient Iberian samples and Sorraia
-Data gives no support to d'Andrade's claim the Sorraia had a primitive, predomestic ancestry

LG:  What can be more accurately stated is that the 22 horse remains samples from Portalon and the two horse remain samples from Cova Fosca showed no relationship to the DNA samples of the Sorraia used in this research.   The Sorraia and Zebro were documented as being fauna of southewestern Iberia.  The prehistoric samples in the Lira study are from the northeast.  D'Andrade made it known that the horses of the north were not the same as those from the south.  The Lira et al., paper is a limited study (24 samples) and simply shows that these samples are not ancestral to the Sorraia.  It is inappropriate conjecture to use this study to claim the Sorraia have no prehistoric connection in Iberia, and this is not quite what the Lira paper is saying, rather they concluded:
"Lastly, our data do not support the Sorraia horses as a primitive predomestic lineage, in contrast to a previous claim (D'Andrade 1945). The only unique ancient Iberian haplotypes that are closely related to Sorraia sequences are not extremely old, but rather date to the Middle Ages (ATA07 near to Pomba, and HOR 23 near to Azambuja) (Fig. 2B and Fig. 5). However, it is worth noting that today, as a result of the bottleneck they underwent during the 20th century (Luıs et al. 2002), the only maternal lineages of this breed are constituted by Azambuja, Cigana and Pomba. Thus, the possibility that some of the Portalon haplotypes could represent extinct maternal lineages from Sorraia cannot be completely excluded." [underlined emphasis is mine. --LG] 

To which I would add that once again these tests are resting on mtDNA which tells only half the story of what potential relationships present day horses have with prehistoric horses.

The mouse-dun (grullo) hues of Altamiro's first colt, Animado--here he looks like the landscape itself!

#6:  Roya et al., The Origins of Iberian Horses Assessed via Mitochondrial DNA, 2005

The historian used this paper to conclude:
-Sorraia is not ancestral to Iberian horses

[02Sept12 - It has been brought to my attention by a private email that my attempts to present a more rounded picture of what these various genetic studies mean in relation to the Sorraia horses is creating greater confusion rather than making these issues clearer, due to my lack of understanding of this field of science.  This means my perception of what these studies reveal is as misunderstood as that of the historian.  Until such time as someone with sound knowledge provides a better explanation of the genetic studies, I am withdrawing my own attempts to place them in appropriate context. --LG]

#7:  Cieslak et al., Origin and History of Mitochondrial DNA Lineages in Domestic Horses, 2010

from this research article, the historian concluded:
-Sorraia doesn't possess ancient haplotypes
-Sorraia possesses haplotypes occuring in Asia, not Iberia
-No direct link between native Iberian prehistoric horses and Sorraia
-Lusitano different haplotypes than the Sorraia, no direct ancestry to Sorraia

LG:  See earlier responses to these types of mtDNA researches.  More precisely, the historian's determination should state that within the present day remaining genetic data from this particular research did not find any shared haplotypes between the two surviving maternal lineages of Sorraia and the prehistoric equine samples used for comparison.   It may be that the Sorraia does possess prehistoric as well as native Iberian relations on the paternal side, or that lost maternal lineages had such connections.  The results from Jansen et al., 2002 and Georgescu et al. 2011 certainly suggest the Sorraia indeed has distinct, haploytypes that are more closely aligned to wild horses types than modern domestic breeds.

Where does the grass end and the colour grulla (mouse-grey) begin?
Dearly missed, the lovely Ciente!

Item #8:  A last bit of "hard evidence" the historian presented he simply said came from "personal correspondence" with Professor Vera Eisenmann  (which he did not share or quote from)

from item #8 the historian concluded:
--Sorraia doesn't resemble the appearance of the original European wild horse

LG:  If Vera Eisenmann has limited wild horses of Europe to just one "original" type, I would need to see some supporting research to that end before commenting.  The scant remains of the Tarpan wouldn't give a complete picture of what the European wild horses looked like. There is enough incidence in prehistoric art, along with stenoid and caballoid fossils and surviving remnants of primitive horses living in a variety of regions in Europe to suggest more than one type of European wild horse was present in prehistoric eras and within these types regional variants would be expected.

(Double click on the above image to see the enlarged size)

It appears there is not yet sufficient equine caballoid fossil remains recovered from southern Iberia to provide a definitive determination of what variant of Equus ferus made its habitat there, but this doesn't mean there were not more than one type ("absence of evidence is not evidence of absence"), rather, it means that the limitations of the fossil record do not allow for sweeping judgments regarding what types of wild horses roamed Eurasia, it can only give us incomplete puzzle pieces to conjecture from.  When accompanied by what we can glean from historic texts as well as present day specimens like the Sorraia who have retained a large degree of prehistoric characteristics (both phenotypically and behaviourally) the conservation of the Sorraia and Sorraia Mustang is not only justified, but vital, for perhaps no other remnant of Equus ferus is as near to extinction are these "Iberian Tarpans". 

(Double click on the above image to see the enlarged size)

The historian ended his detraction of the Sorraia with several closing points and opinions that I would like to pass comment on as well, first of which is that he stated:

"I could also argue that the Sorraia phenotype isn’t the most efficient phenotype."

While the historian did not in that sentence define for what purpose he was measuring the efficiency of the Sorraia phenotype against, but knowing he has written numerous articles for conservancy and rewilding websites, we must suppose the historian is saying the Sorraia's phenotype would be inefficient for serving as large herbivores living a free range, wild existence in a variety of wilderness parks...if this is the case, I cannot imagine in what way the Sorraia would be considered inefficient.

 Sorraia stallion eating dirt for the mineral content
Ravenseyrie Sorraia Mustang Preserve 
Manitoulin Island, Ontario, Canada

In Portugal though some Sorraias are kept in stables and breed artificially, there remain some individual breeders who provide a semi-wild existence on traditional range lands that for large parts of the year are subjected to drought conditions and poor forage.  One private initiative has created the Vale de Zebro Sorraia refuge where these horses have been living without human assistance since 2004 (except for the rare removal of excess individuals to assure the Sorraia do not overpopulate in a manner detrimental to the overall health of the refuge).  But perhaps it is no surprise the Sorraia thrive in the region where they have developed.  Examples of phenotypical efficiency have been proven as well in less temperate zones as demonstrated by various breeders in Germany and France who are  keeping Sorraias more or less exposed to the elements in natural settings.

Sorraia foal browsing on a tree stump
Ravenseyrie Sorraia Mustang Preserve
Manitoulin Island, Ontario, Canada

And, perhaps even more illustrative are the examples of the Sorraia and Sorraia Mustangs living here at Ravenseyrie in Northern Ontario, where we can experience seasonal temperatures which can swing from -30°C in the winter to +30°C in the summer.  The Ravenseyrie horses range over a varied landscape of forests, grasslands, wetlands and cobble lakeshore.  Due to its limited acerage (360) the horses are supplemented with fodder over winter, but they nonetheless browse and forage their entire landscape and live semi-wildly, in natural family units with as much autonomy as we can afford them.  They are not on any maintenance programs for parasites, immunizations or hoof trimming as their environment provides for all their needs.  I have no doubt that were the whole island available to them, they would survive more easily than do the wild Whitetail Deer that populate the island...(in fact the deer manage winters much better where horses dwell, due to how adept our horses are at maintaining trails through the deep snow and dig large openings in the snow to graze - both aspects that deer take advantage of.  Efficient equines, I should say!

Sorraia mares and foals drink the last of the water in a favourite watering hole
Ravenseyrie Sorraia Mustang Preserve
Manitoulin Island, Ontario, Canada

    The Sorraia horses of Ravenseryie are very keen to find water in wintertime...they will dig through snowpack and ice to reach water flowing beneath

 Sorraia mare Zorita paws through deep snow to graze the grass beneath it.

The Ravenseyrie Sorraia horses are as adept at using the forest as well as the open grasslands to their best advantage. The copious photos used in this article should present a convincing picture of just how efficient the Sorraia horses phenotype is as well as reveal how good they are at making the most of living in the wilderness.

 Foals in the forest at Ravenseyrie

This Sorraia foal runs through the grassland almost completely indistinguishable from the landscape itself!
Ravenseryie Sorraia Mustang Preserve 
Manitoulin Island, Ontario, Canada

The variation in hues of the mouse-grey (grullo) coat colour and contrasting dorsal stripe adeptly mimic the colour tones in the landscape.
Ravenseyrie Sorraia Mustang Preserve
Manitoulin Island, Ontario, Canada

Likewise, the variety of striping that often accompanies the mouse-grey coat colours are useful in the forest, mimicking the trees, dried leaves and shadows, as shown here on two Ravenseyrie horses

As an "overall conclusion" the historian makes six statements which I am quoting directly from the paper and giving brief replies to:

1.  The breeding history of the Sorraia is vague and represents the very poor knowledge of the time about wild and prehistoric horses.

This history of the Sorraia may have a few questions presently unable to be answered, but it is nonetheless straight forward, with sufficient support from historic sources and morphological comparisons to provide a unique status for the Sorraias as a regional variant of the Tarpan (E. gmelini).

2.  D’Andrade did not select wild horses, but selected from domesticated horses of the region and also imported a stallion from Argentina.

The horses d'Andrade selected for preservation were examples of primitive phenotypes which persisted/survived in semi-wild non-pedigreed stock as well as in certain individuals of domestically bred Lusitanos.  Later the inclusion of the Argentinian Criollo of Sorraia phenotype was experimented with in order to alleviate the inbreeding resulting from such a limited founder group.  D'Andrade was not trying to "breed back" the wild type, rather he was attempting to recover and consolidate the genetics that had obviously been retained in certain individuals.

3.  All genetic researches do point to the very clear fact that the Sorraia is neither an ancestor to any Iberian horse, nor is it a direct descendant of prehistoric/original wild horses.

The genetic research submitted in the historian's dismissal of the Sorraia is of limited value and not sufficiently reliable to make definite claims that a prehistoric and/or Iberian ancestry is not present.

4.  When looking at phenotype, I get the feeling that it doesn’t represent the most efficient phenotype for living under natural conditions. It will survive, but I am sure that other proven primitive breeds (like the Garrano, Retuerta Pottoka, etc.) will perform better.

The Sorraia's history is that of an efficient phenotype that survives incredibly well in the most marginal of areas under the roughest conditions - it already is a "proven primitive breed", or rather "type".  In modern times, it continues to demonstrate dramatic efficiency and resiliency in a broad range of habits and climates, in wild and semi-wild settings.  The Sorraia is on par with other primitive wild types of horses and is well suited to conservation schemes if measures are further undertaken (like outcrossing to Sorraia types in North American Mustang and South American Criollo populations) to reduce the effects of their genetic bottleneck.

5.  Sorraia doesn’t look like prehistoric horses, including a wrong color.

The Sorraia is a classic representative of Ebhardt's ancestral form III and as demonstrated is of the colour of the European wild horse E. gmelini.

 Sorraia stallion, Altamiro, looking very much like a prehistoric horse of the right colour

6.  Most important is the fact that the diverse papers and researches all point into the same general direction. There is no escaping the inevitable conclusion.

The volume of papers and researches provide enough insight into the type of horse the Sorraia represents to merit its continued preservation and extend its conservation to rewilding projects in a number of appropriate places. 

Given all the points the historian used to build his case "against the Sorraia", it is quite provocative to read what he has written in his " strong plea to protect the wild herds of the Letea forest horse of Romania" which can be viewed in Romanian and English if you follow the link (by clicking on the underlined text) and scroll all the way through the web page.  The core information has been reprinted on the Large Herbivore Network page devoted to the Letea Forest Horse. I agree this Romanian wild horse type should be preserved, indeed...but when it shares so many similarities with the type of history on record for the Sorraia and is also of a more refined morphology than horses like the E. przewalskii  why is the Letea Forest horse's "story" determined to be more acceptable to the historian than that of the Sorraia, which has a much greater body of evidence pointing toward their wild heritage? 

With as much visibility and authority this historian is projecting into the places where wild horse preservation is finally beginning to coincide with habitat conservation, it is important to offer the public the full picture of the Sorraia lest the historian's misrepresentations and misunderstandings of Sorraias, Tarpans (and Koniks) undermine the essential and valuable work role these wild horses play alongside other wild horses.  It is my hope the historian will have a more expansive understanding of the Sorraia and rethink his dismissal of its prehistoric heritage.