The Fan Hitch PostScript
Number 5, posted January 2020
In this Post

From the Editor

The Aboriginal Dog as a Domesticate

Neuroanatomy and Behavior Correlations

Specialized Sledge Dogs Accompanied Inuit Dispersal Across the North American Arctic

Cold Case Reopened and Other QIMMEQ News

Langsomt på Svalbard (Slowly on Svalbard)

Frossen Frihe (Frozen Freedom)

Restoring a Historic Nansen Sledge

Media Review: Kamik, an NFB documentary

IMHO: A View from Across the Divide

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Kugluktuk (formerly Coppermine, NWT), Nunavut] 1949
Richard Harrington / Library and Archives Canada / PA-129937
    
“Specialized sledge dogs accompanied Inuit dispersal
across the North American Arctic”

a summary presented by Sarah K. Brown, Ph.D.

Evolutionary biologist Dr. Brown is one of the authors of this ten year multinational research project and was the lead scientist at the University of California, Davis where a portion of the work was performed. The paper was published in 2019 by the Royal Society under the terms of the Creative Commons Attribution License. You can access the full paper here. Ed.
Abstract from the published paper:
Domestic dogs have been central to life in the North American Arctic for millennia. The ancestors of the Inuit were the first to introduce the widespread usage of dog sledge transportation technology to the Americas, but whether the Inuit adopted local Palaeo-Inuit dogs or introduced a new dog population to the region remains unknown. To test these hypotheses, we generated mitochondrial DNA and geometric morphometric data of skull and dental elements from a total of 922 North American Arctic dogs and wolves spanning over 4500 years. Our analyses revealed that dogs from Inuit sites dating from 2000 BP possess morphological and genetic signatures that distinguish them from earlier Palaeo-Inuit dogs, and identified a novel mitochondrial clade in eastern Siberia and Alaska. The genetic legacy of these Inuit dogs survives today in modern Arctic sledge dogs despite phenotypic differences between archaeological and modern Arctic dogs. Together, our data reveal that Inuit dogs derive from a secondary pre-contact migration of dogs distinct from Palaeo-Inuit dogs, and probably aided the Inuit expansion across the North American Arctic beginning around 1000 BP.
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The earliest evidence of domestic dogs in North America dates to ~10,000 years ago. Until recently, not much was known about how, when, and if domestic dogs were introduced to North America, or if they were domesticated from North American wolves. Ancient DNA and archaeological data from a 2018 study (Leathlobhair et al. 2018) suggests that North American pre-contact dogs (PCD) belonged to a now extinct lineage, are most closely related to ancient Siberian populations, and did not derive from North American wolves. This data suggests that pre-contact dogs were introduced to the Americas by people from Siberia/Beringia, and subsequently went extinct after the introduction of European breeds. Interestingly, this now extinct lineage of pre-contact dogs appears to be closely related to modern Arctic dogs (Alaskan Malamutes, Inuit Dogs and Siberian Huskies). However, PCD are not the direct ancestor of these modern breeds. This leads to questions about the origin of Arctic breeds, both modern and ancient.

Leathlobhair et al. (2018) hypothesized that four waves of dogs (associated with humans) were introduced to North America; 1) the pre-contact dogs from Beringia/Siberia, approximately 17,500 – 13,000 years ago, 2) a potential introduction by Thule/Inuit or the Paleo-Inuit, into the Arctic ~4,500 – 2,000 years ago, 3) European breed dogs from settlers, as early as 1492 AD and 4) dogs during the gold rush, ~1900 AD. Leathlobhair et al. (2018) did not have enough ancient Thule/Inuit and Paleo-Inuit dog samples to address the question of how Arctic dogs arrived in the North American Arctic, and their relationship to the pre-contact dogs.

The North American Arctic was initially colonized by two waves of humans. The first, the Paleo-Inuit, who arrived in the North American Arctic roughly 4,500 years ago from Siberia, and show little evidence for extensive use of dogs. The second wave occurred ~2,000 years ago, with the arrival of the Thule people (ancestors of the Inuit), who heavily utilized dogs for sled traction. It is unclear what the relationship is between Paleo-Inuit and Thule dogs, and pre-contact dogs. To this end, our team investigated the following questions, 1) What is the relationship of ancient Thule/Inuit dogs and Paleo-Inuit dogs to pre-contact dogs? 2) Did the ancient Thule/Inuit dogs derive from the Paleo-Inuit dogs?, and 3) Are modern Inuit Dogs descended from ancient Thule dogs?

The Mammalian Ecology and Conservation Unit (MECU) in conjunction with the Department of Anthropology at the University of California, Davis, started investigating these questions through genetic and archaeological analysis nearly a decade ago (Brown et al. 2013; Brown et al. 2015). These efforts culminated in the most recent paper "Specialized sledge dogs accompanied Inuit dispersal across the North American Arctic." (Ameen, Feuerborn, Brown, Linderholm et al. 2019), which is the result of two initially independent efforts, one led by our team from UC Davis, and the other by researchers from the University of Exeter (and many other institutions).

Team members traveled to museums around the world to collect ancient dog remains from various Inuit/Thule and Paleo-Inuit Archaeological sites throughout Alaska, Canada, Siberia and Greenland. A total of 628 samples from the following time periods were collected, and extracted for DNA; Paleo-Inuit (5,500 – 500 Before Present (BP), Archaeological Inuit/Thule (2,000 – 200 BP), Historical and present (300 BP – present). Because DNA degrades over time, eventually leaving only trace amounts behind, archaeological DNA is especially vulnerable to contamination from modern sources. Therefore, we took special precautions when extracting ancient DNA to safeguard against contamination. We extracted all ancient samples in a special, isolated laboratory where no modern DNA has ever been present. After DNA was extracted from the ancient dog teeth and bones, the entire mitochondrial DNA genome was amplified and sequenced. By sequencing the entire ~16,000 base pairs (bp) (e.g. letters) of the mitochondrial genome, this allowed much deeper resolution than previous studies that utilized ~300 bp (Brown et al. 2013). These samples were analyzed and compared to 1) each other, 2) other published dog and wolf sequences, and 3) 71 North American dog mitochondrial genomes, previously published in Leathlobhair et al. (2018).

What is the relationship of ancient Thule/Inuit dogs and Paleo-Inuit dogs to pre-contact dogs?
Sequencing success of the oldest samples (Paleo-Inuit) was poor, with only 12 out of 92 samples producing data. The results of our genetic data indicate that ancient Paleo-Inuit dogs largely (83% of samples) shared the same genetic signature as pre-contact dogs (genotype A2b). However, the Thule/Inuit Dogs did not possess the pre-contact dog and Paleo-Inuit genetic profile (genotype A2b). The ancient Thule/Inuit dogs harbored another genetic signature (genotype A2a), which was common in ancient Siberian samples.

Did the ancient Thule/Inuit dogs derive from the Paleo-Inuit dogs?
Most likely, no. The majority of the ancient Thule/Inuit Dogs sampled have the A2a haplotype, which was common in Siberia, but was not common in Paleo-Inuit dogs. Only two of the Paleo-Inuit dogs possessed the A2a genotype, suggesting that it was present in the North American Arctic before the Thule/Inuit Dogs arrived. However, it is likely that the presence of this genotype in Paleo-Inuit dogs is due to Siberian ancestry. The Thule/Inuit people were responsible for spreading the Thule/Inuit Dog genotype (A2a) throughout the Arctic, causing a near replacement of the PCD/Paleo-Inuit genotype (A2b).

Additionally, we performed morphometric analysis (Geometric Morphometrics; GMM) on mandibles and crania from these specimens. We found that Thule/Inuit Dogs tended to be larger, with a more narrow cranium than Paleo-Inuit Dogs, suggesting that these populations are morphologically different, supporting the genetic data.

Are modern Inuit Dogs descended from ancient Thule dogs?
Yes. Historical and modern dogs still maintain the Thule/Inuit dog genotype (A2a), making modern Inuit dogs one of the last true Native American dogs remaining. However, roughly 37% of modern dogs sampled possessed the A1a genotype, potentially indicating European breed introgression.

The arrival of the third and fourth wave of dogs (European breeds arriving with settlers and the gold rush) resulted in the replacement and complete loss of the pre-contact dog lineage in North America. The genetic signature of the Thule/Inuit Dog (the second wave of dog migrations) remains in the North American Arctic.  The Inuit Dog may be one of the last descendants of pre-European dogs left in North America.
   
References:
Ameen, Carly, Tatiana R. Feuerborn, Sarah K. Brown, Anna Linderholm, Ardern Hulme-Beaman, Ophélie Lebrasseur, Mikkel-Holger S. Sinding et al. "Specialized sledge dogs accompanied Inuit dispersal across the North American Arctic." Proceedings of the Royal Society B 286, no. 1916 (2019): 20191929.

Brown, Sarah K., Christyann M. Darwent, and Benjamin N. Sacks. "Ancient DNA evidence for genetic continuity in arctic dogs." Journal of Archaeological Science 40, no. 2 (2013): 1279-1288.

Brown, S.K., Darwent, C.M., Wictum, E.J. and Sacks, B.N., 2015. Using multiple markers to elucidate the ancient, historical and modern relationships among North American Arctic dog breeds. Heredity, 115(6), p.488.

Leathlobhair, Máire Ní, Angela R. Perri, Evan K. Irving-Pease, Kelsey E. Witt, Anna Linderholm, James Haile, Ophelie Lebrasseur et al. "The evolutionary history of dogs in the Americas." Science 361, no. 6397 (2018): 81-85.

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Below is a Q & A with Dr. Brown and The Fan Hitch

TFH: How long has the Inuit Dog as we now know it, existed in the circumpolar north?
SB:   Our research has shown that the majority of the Paleo-Inuit Dogs (present in archeological sites starting ~4,500 YA) had a different genotype than the Thule/Inuit Dogs, and that there was a near replacement of genotypes when the Thule arrived in North America. The earliest that we see this Thule/Inuit Dog genotype is around 2,000 years ago. So, no, the Inuit Dog that was utilized by the Thule and later Inuit has not been in the arctic for ~4,000 years.

TFH: Pre-contact dogs (PCDs) are dated prior to 1492 AD, yes? And this applies only to North America, not Greenland?
SB:   Pre-contact dogs (PCDs) are prior to 1492 AD in North America, but PCD is also defined as the genetic group/clade (the A2b that is also found in the Paleo-Inuit). The Leathlobhair 2018 paper did not sample Greenland, and this is where the term was developed.

TFH: Is Ken MacRury’s conclusion about wolf DNA in Inuit Dogs still valid?
SB:    Leathlobhair et al. (2018) showed that North American dogs did not derive from North American canids. In our archaeological assemblages, we often found larger canid bones mixed in with dog bones, and assumed them to be wolf. In the example in the paper (sample AL2797), was one such mandible that appeared larger than the dog specimens it was found near. It’s mitochondrial type was wolf, and the nuclear genome confirmed that it was indeed a wolf. It is known that the Thule used wolves for pelts/fur, and could have hunted them as a predator control. Therefore, it seems more likely that the wolf bones found in the sites was due to utilization of wolves, rather than hybridization.

TFH:  About your Figure 1 and Table 2 and other comments on page 3 of your 2019 paper, going back as far as Ken’s 1991 thesis and more recent DNA findings, is it still correct to say that the Inuit Dog of Greenland is the same landrace as the Inuit Dog of Canada?
SB:    We use “Inuit Dog” to differentiate the archaeological samples from modern samples, which we termed “Greenland dogs.” However, this research confirms the result from our 2015 paper (Brown et al. 2015), which finds no difference between the Canadian Inuit dogs and Greenland Dogs. They should all be called Inuit Dogs.

TFH: How do you define an aboriginal landrace to be “authentic” (for lack of a descriptive other than the word  “pure” which is used in terms of cultured breeds).
SB:    I think my definition of purity/authentic/native is defined by whether or not the dog has the original Thule/Inuit dog haplotype (A2a). Really, my definition is the genetic continuity from the ancient genetic signature (A2a) from the ancient dogs to the modern dogs. This obviously is not complete, as a mitochondrial haplotype does not represent the whole genome, but it is good as we can get right now.