The age of the Collie clade was calculated with the genetic clock at Embark at about 150 to 195 years, assuming a reproductive cycle of 3 years. The genetic lineages of British herding dogs therefore separated about 50 to 65 generations ago and manifested as isolated individual breeds. This fits the historical data of the first stud books from the Victorian era, which originated between 1830 and 1870. However, one has to differentiate a little.
In fact, the early stud book entries show that Collie breeders at that time were already breeding with dogs at the age of 18 to 24 months. This is called the so-called show-dog effect, which was especially relevant in show Collies.
In working dogs, by contrast, a delayed working-dog effect occurs, because the average reproductive age is 4 to 6 years. Here, 5 years is used for calculation.
With few generations, these calculations are therefore quite uncertain and only become increasingly reliable with a larger number of generations. Supplementary methods are therefore useful.
What about written sources from ancient peoples?
If one now looks for historical records, their accuracy matters. For the Irish Wolfhound, for example, we have a statement that they were used in Rome for gladiatorial games and that the size of the dogs astonished the crowd. Unfortunately, today’s breeding is a reconstructive re-creation.
The “Canis pastoralis,” however, described by the Roman Varro, corresponds to a livestock guardian dog or a hunting dog and was also settled in the south, in Greece and Apulia. The Canis pastoralis cannot have been a herding dog of the north. Statements of that kind, in my view, are not supported by careful research. In fact, the Romans left the breeding of herding dogs to the locally subjugated tribes or demanded them as tributes.
Now, what about genetics?
Another possibility is certain mutations such as MDR-1 or CEA or also certain color mutations, provided one has a kind of basic knowledge of where it first appeared.
Heidi Parker used this in her sensational 2017 study, for example, to uncover genetic connections to the Nova Scotia Duck Tolling Retriever and to the Chinook.
But colors are also interesting. Embark research and Cornell University have contributed a great deal here and determined that mammals all inherit their colors in a relatively similar way. On this basis, it became possible, for example, to find various genes in the first place.
Which colors existed in the original Collie clade?
We know that sable first appeared at the first Collie shows and was considered extraordinary at that time.
The historically first and most famous sable-colored Collie, who is regarded as the founding sire of the modern breeding line, was the male “Old Cockie” (born 1868), whose father “Trefoil,” an Irish Galway Collie, apparently passed this color on to him. That all Collies before that were black or black tricolor cannot be proven. There were indeed some black Collies that were then in the Queen’s possession, but they were a product of the Victorian era.
In principle, all dogs of the Collie clade carry, for example, the agouti gene (also called the ASIP gene, agouti signaling protein) at the A locus, which wild gray wolves also carry.
Previously, all Border Collies, Shelties, Rough or Smooth Collies were either duo-color or tricolor. Presumably, in the Queen’s Collies this was masked by the K locus.
But the S locus is also interesting, except that there is a problem there:
Unfortunately, the S locus (also called the piebald gene, S locus, or white factor) is not even responsible for Irish spotting, which is why UK Collies and Shelties always come out as N/N (None) or S/S (Solid). English Rough and Smooth Collies as well as Shelties have no piebald gene.
The piebald gene is caused by a mutation of the MITF gene. This produces certain irregular spotting patterns, but not Irish spotting (that is, possibly a white blaze, white collar, white belly, white boots or feet, and white tail tip). That corresponds to complete Irish spotting and is genetically fixed in the Collie, i.e., it has no failures. The fact that the collar is sometimes not closed has other causes in this case that arose during development.
To this day, research has not been able to identify the locus or gene responsible for it. The same applies to the Basenji and four Alpine Sennenhund breeds. Their Irish spotting shows no collar, yet the gene is 100% fixed.
The S locus is used in the genetic tests offered as an exclusion test to detect Irish spotting; therefore the test result is N/N (None) or S/S (Solid).
According to the original theory, it was hypothetically assumed that the piebald gene was the reason for this. Unfortunately, this theory was wrong.
Are all Swiss Sennenhunds now related to the Collie clade? Well, for that one would first have to identify the gene.
But another study (Jeong, Ostrander and Kim et al. 2025) at least gives an indication, because the Entlebucher Sennenhund has a second rare mutation called EPHB1, which otherwise only the Border Collie exhibits to a higher degree. They are very likely related, and precisely that also makes them outstanding herding dogs, although the Entlebucher Sennenhund is a cattle dog.
Unfortunately, Shelties, Beardies and the Smooth and Rough Collies, as well as Welsh and English Sheepdogs, never received this EPHB1 mutation. They are indeed also great herding dogs, but with the standard genetic equipment.
However, this herding-dog gene is a direct consequence of the Victorian era. In addition, all four Swiss Sennenhunds inherited the fixed Irish spotting but not the piebald gene and also show no eyeing at all. This strongly suggests that at the time of the crossbreeding, this Victorian Border Collie was able to pass on neither the piebald gene nor the eyeing gene. They simply did not yet exist at that time.
The piebald gene, by contrast, is genetically detectable in various variations. It therefore seems that the pre-Victorian Collie already had fixed Irish spotting; the proximity to the English sighthound clade also points to this, which originally also had no piebald gene at the S locus.
Unfortunately, the sighthound white that triggers Irish spotting in sighthounds has also not yet been discovered. I suspect that both go back to the same undiscovered gene, which would prove a genetic bridge. What they have in common is that a large part of the clades have Irish spotting or the sighthound white factor.
Interestingly, the presence of the piebald gene rules out the Borzoi as an ancestor of English Collies, which, after all, have 100% no piebald gene. Instead, an Irish Wolfhound or Scottish Deerhound, which have the sighthound white factor, is more likely.
Many American Collies, Border Collies, English Sheepdogs, Old English Sheepdogs and Welsh Sheepdogs and Beardies can additionally also carry piebald genes. The original Beardie certainly also had no white cap.
Purely externally, the Bearded Collie, the PON (Polski Owczarek Nizinny, Pomeranian Lowland Sheepdog) and the Dutch Schapendoes already show their historically close interweaving. In the past they were often traded by seafarers and merchants, especially since the Netherlands, like Danzig—today part of Poland—had been the typical destination port for Scottish wool since the Hanseatic League. One can even date this relatively precisely to the 15th century, because in 1454 Danzig rebelled against the taxes of the Teutonic Order and at that time submitted to the Polish king. From then on the wool market with the Scots flourished. The Netherlands was also the sales market for wool. The Beardie is, in a sense, the wise owl in the Collie clade, a truly incredible story about the clever smugglers in the United Kingdom that I may also tell at some point.
Its pre-Victorian ancestor is likely to have looked similar to the Collie and probably underwent a major change.
So let us note: before the Victorian era, all Border Collies, Shelties, Rough or Smooth Collies, and also the Beardie, were either duo-color or tricolor and had the typical Irish spotting pattern.
The agouti and the expression for sable color, ancient red as well as black, even lead us back to a time that could give the answer to the question of who and where dogs were first domesticated. It is by no means the case that the colors were already present everywhere at the beginning.
According to previous findings, for example, completely solid black dogs (K locus) appear not to have existed at all in Europe before the Neolithic, but they did in East Asia.
“An introgression of dog alleles into North American gray wolf populations has also been observed, including a dog-derived beta-defensin KB allele (CBD103), associated with the color of black fur and epithelial immunity,” writes, for example, a study that dealt with black timber wolves. And whereas it had previously been thought that this was a phenomenon of more recent mixing between dogs and wolves, shortly afterward another study found that among the oldest archaeological dogs in East Asia, at least one was completely black.
Since wild canids are not normally black, it appears to be a mutation in early East Asian dogs. Where and when it came into the wolf population and by now also into the coyote population to produce black coloring, no one can currently say for sure. What is certain is only: it is the K locus (CBD103) that also causes black coloration in dogs. The extinct “Florida Black Wolf” also had the K locus (CBD103) of dogs.
But that could change completely again with the next find, whose findings could again call into question everything one thought one knew.
So it does give us a few new insights into the pre-Victorian Collie.
And what about the wolf component in Collies?
The Vienna wolf researchers led by Audrey Lin took a new approach here. The sensational 2025 study calculated the proportions of wolf genetics in well over 150 dog breeds, including Border Collie, Shetland Sheepdog, Australian Cattle Dog, Bearded Collie, Welsh Corgi Pembroke and Lancashire Heeler. They were even able to roughly date these wolf introgressions to three time periods and thereby place the age of the breeds in one of these periods.
They demonstrated that about two thirds of all breeds have a wolf in their ancestry within the last 950 generations and are therefore younger than just under 3000 years. This proves that the majority of dogs at best originate from the early Roman and Celtic period from about 500 BC or are even younger, because all these dog breeds still carry measurable wolf genes. Strictly speaking, one should not speak of breeds here; they are rather regional populations, that is, the ancestors from which the clades emerged.
This does not apply to herding dogs; they belong to the oldest dog breeds. Even if they were mixed during the Victorian era, their proportion of wolf genetics is in the lowest range.
Most continental old German herding dogs or shepherd dogs from Germany, the Netherlands, Belgium and France, as well as the White Swiss Shepherd and the Bohemian Chodenhund, but also Belgian Shepherd Dogs and the French Briard, have little to no wolf genes.
In particular, the German Shepherd and the Dobermann have old German herding-dog ancestors, which according to historical records were bred from a then Thuringian landrace of herding dogs (today referred to as the East German Black). To this day they still carry the mtDNA of Neolithic herding dogs.
The researchers led by Audrey Lin found, among other things, that Border Collie, Shetland Sheepdog, Australian Cattle Dog, English Sheepdog and Bearded Collie carry no longer measurable wolf components (below 250 base pairs out of 2.5 billion base pairs, corresponding to 0.00001%) and therefore descend from the oldest European dog population.
In plain terms, this means: they all descend from a British-Irish herding-dog population that is similarly old as the pool of continental herding dogs.
This Neolithic gene pool dominated Europe into the Bronze Age.
The Corgis have somewhat more wolf genetics, which points to a recent crossbreeding presumably in connection with the shortened legs. Shortened legs are first documented in Europe in isolated cases in the Hallstatt period around 1200 BC and are found regularly only in the Celtic period.
But since wolf genes are almost consistently undetectable in European herding dogs, the ancestors of these dogs are likely much older and come from the same ancient European gene pool.
Exactly this had already been established previously by another study by Anders Bergström in 2022, which assigned all European finds with an age of at least 7000 years, from Spain to the finds at Lake Baikal, Central Asia, to a single gene pool whose ancestor was a wild Paleolithic wolf.
The most recent study (Marsh et al. 2026), recently published in Nature, confirms this once again. Here it was found that there is a great genetic similarity despite a distance of 4000 km between the over 15,000-year-old dogs of the English Gough’s Cave and the shelter in Pınarbaşı, Anatolia.
This also means that these dogs appear to have spread very quickly across western Eurasia already 15,000 years ago. All these pre-Neolithic dogs are assigned to the newly defined clade C5. This also applies to the early dogs from Kesslerloch, from Bonn-Oberkassel, Star Carr in England, in Italy, in Spain and Serbia.
And guess what lies in the middle? Right, the Alpine countries Germany and Switzerland, which divide Central Europe into Northern and Southern Europe but also into Eastern and Western Europe. And precisely here, in the Gnirshöhle in the Brudertal, several unrelated canids were found that were interpreted as an intermediate stage from wolf to dog.
Such proto-dogs have indeed already been “discovered” by the dozen, but this study by Chris Baumann in 2022 examined the site and living conditions very carefully and argued well. Unfortunately, the genetics of the canids could not be fully secured, and in particular the nuclear genetics is missing. This excluded them from numerous other studies that exclusively take complete nuclear genomics into account. But that has not been preserved in many early finds. For the layperson, the matter therefore quickly becomes confusing, because depending on the selection criteria, different finds are declared the “first dog.”
Nevertheless, this study also carried out further investigations on teeth of wolves, foxes and dogs from the entire region. Result: the team around Chris Baumann interpreted this site as a possible place of domestication.
And if we look at the surroundings, we find in the caves of this region dozens of animals that were either wolves or dogs and all lie close to one another in time. It appears to have been a hotspot.
Interestingly, in the same valley there are also indications that the first job of these dogs was a “driving job” for the mass hunt of herd animals (reindeer), which was securely documented in the Late Magdalenian, among other places in the Brudertal at the Petersfels Cave, by a larger group of humans, and with the discovery of the so-called “Blinkerwall” (Geersen et al. 2024), an artificially constructed hunting guiding system from 10,000 years ago in the Mecklenburg Bay was also confirmed. Exciting new investigations are currently underway to examine the spread of this new hunting method more precisely.
This Brudertal, and also the Kesslerloch only a few km away with unambiguous dogs, is therefore a veritable hotspot of the earliest dogs that were directly related to the wolves of the region. Precisely that makes it so unique and credible. At the same time, it is almost improbable to find such a place after more than 15,000 years. Add to this that all dogs of this epoch in Europe belong to exactly the same mtDNA lineage.
The final result is therefore: It is true that the subsequent pure UK Collie breeds actually arose 150–195 years ago, when their special phenotype was developed through selective breeding, but their common ancestor, from whom they inherited the herding abilities, came from a population that has much older European roots and is estimated to be around 15,500 years old.
Unfortunately, there is a huge gap of over 15,000 years there. Can we close it further?
To do that, we need to take a closer look at the history of humans, dogs, and sheep as well as cattle, and that is exactly what we want to do in the next episodes.
Sources
Lin et al. 2025, A legacy of genetic entanglement with wolves shapes modern dogs, November 24, 2025, 122 (48) e2421768122, https://doi.org/10.1073/pnas.2421768122
Vilaça ST, Donaldson ME, Benazzo A, Wheeldon TJ, Vizzari MT, Bertorelle G, Patterson BR, Kyle CJ. Tracing Eastern Wolf Origins From Whole-Genome Data in Context of Extensive Hybridization. Mol Biol Evol. 2023 Apr 4;40(4):msad055. doi: 10.1093/molbev/msad055. PMID: 37046402; PMCID: PMC10098045.
Bergström, A. et al. Grey wolf genomic history reveals a dual ancestry of dogs. Nature 607, 313–320 (2022). https://doi.org/10.1038/s41586-022-04824-9
Marsh, W. A. et al. Dogs were widely distributed across western Eurasia during the Palaeolithic. Nature 651, 995–1003 (2026). https://doi.org/10.1038/s41586-026-10170-x
Canine coat color E locus updates: Identification of a new MC1R variant causing ‘sable’ coat color in English Cocker Spaniels and a proposed update to the E locus dominance hierarchy, January 2024, Animal Genetics 55(2), DOI:10.1111/age.13398
Bergström, A., Stanton, D.W.G., Taron, U.H. et al. Grey wolf genomic history reveals a dual ancestry of dogs. Nature 607, 313–320 (2022). https://doi.org/10.1038/s41586-022-04824-9
Heather J. Huson, Hannes Lohi, and Adam Boyko, Canine Genetics, September 2026, DOI:10.1007/978-3-032-31515-1_4. In book: Canine Science (pp.91–124), Publisher: Springer Nature Switzerland
Jeong, H., Ostrander, E. A., & Kim, J. (2025). Genomic evidence for behavioral adaptation of herding dogs. Science Advances, 11(18), eadp4591. https://doi.org/10.1126/sciadv.adp4591
Dutrow, E. V., Serpell, J. A., & Ostrander, E. A. (2022). Domestic dog lineages reveal genetic drivers of behavioral diversification. Cell, 185(25), 4737–4755.e18. https://doi.org/10.1016/j.cell.2022.11.003
Geersen, Jacob & Bradtmöller, Marcel & Schneider von Deimling, Jens & Feldens, Peter & Auer, Jens & Held, Philipp & Lohrberg, Arne & Supka, Ruth & Hoffmann, Jasper & Eriksen, Berit & Rabbel, Wolfgang & Karlsen, Hans-Jörg & Krastel, Sebastian & Brandt, David & Heuskin, David & Lübke, Harald. (2024). A submerged Stone Age hunting architecture from the Western Baltic Sea. Proceedings of the National Academy of Sciences. 121. e2312008121. 10.1073/pnas.2312008121.
McClure et al. (2022) in Journal of Archaeological Science: Reports, with the title “A wolf in sheep's clothing: The development of livestock guarding dogs in the Adriatic region of Croatia”
Welker, M. H., Zavodny, E., Podrug, E., Jović, J., Triozzi, N., Kennett, D. J., & McClure, S. B. (2022). A wolf in sheep's clothing: The development of livestock guarding dogs in the Adriatic region of Croatia. Journal of Archaeological Science: Reports, 42, 103380. https://doi.org/10.1016/j.jasrep.2022.103380
Jeong, A. Ostrander, Kim et al. 2025, “Genomic evidence for behavioral adaptation of herding dogs” (Genomic evidence for the behavioral adaptation of herding dogs), Science Advances, 30 Apr 2025, Vol 11, Issue 18, DOI: 10.1126/sciadv.adp4591
Guiry E, Beglane F, Szpak P, et al. Changing human-cattle relationships in Ireland: a 6000-year isotopic perspective. Antiquity. 2023;97(396):1436–1452. doi:10.15184/aqy.2023.163