Fig 1.
Three contemporary hunting spitz breeds representing breeds with Finnish origins.
The Nordic Spitz (left), although officially Swedish, the breed originates from feral dogs typical for norther Finland and was brought to Sweden by Finnish settlers. The breed almost went to extinction prior to its recognition in 1966. Finnish Spitz (center) is the Finnish national dog and was established as a breed already in 1892. Karelian Bear Dog (right), a much larger spitz type used for large game hunting, was recognized as a breed in 1936 but registered only in 1946. All photos by the first author.
Table 1.
Sample sizes and median heterozygosity frequencies as indicated. Notice the low heterozygosity in the endangered Norwegian Lundehund [14].
Fig 2.
Multidimensional scale (MDS) plots of genetic similarities among the analyzed breeds.
(A) MDS of Northern hemisphere hunting spitz and Laika breeds. The plots are three-dimensional (3D) and the size of each dot (sample) is relative to its position on the z-axis. Pannable, interactive 3D plot can be accessed at https://www.mydogdna.com/crm/index.html#en/breeds/519248a83cd390a0520000ce/norrbottenspitz/relationships (tab “Nordic Hunting Dogs” or “Nordic Spitz Breeds” to include also the herding breeds). As MDS (as well as other PCA applications) find the best possible fit for all samples in the dataset, the separation of the different clusters is dependent on which samples are included. This can be demonstrated by the separation of closely related Lapphound and Lapponian Herder when viewed together with other breeds (B) or when only the two breeds are compared (C). Therefore, also the position of the clusters does not represent true genetic distances (e.g. % of genetic difference) between the breeds, as can be depicted on dendrograms.
Fig 3.
Phylogeny of 13 contemporary northern Eurasian spitz breeds, based on 1319 SNPs.
(A) Phylogeny based on coalescent. Maximum clade credibility tree drawn in black, with posterior probabilities shown at each node. Tree cloud shows the range of alternative consensus tree topologies and was produced using DensiTree visualization of SNAPP results, with samples taken every 1000 MCMC repetitions from 1M iterations. (B) SNP phylogeny obtained from MrBayes, which ignores possible coalescent variance. Maximum clade credibility and DensiTree visualization of consensus tree topologies as previously. Black arrowheads point to frequent alternative topologies linking Karelian Bear Dog with Finnish and Nordic Spitz clade (A) or Norwegian and Swedish Elkhound as sibling breeds (B). Posterior probabilities under 0.4 were omitted.
Fig 4.
Maximum-likelihood tree obtained from TreeMix analysis, based on the most common alleles within each population.
Branch lengths represent the amount of genetic drift that has occurred in the breeds and arrows depict the degree as well as direction of ancestral admixture. Yellow-red arrows between the branches depict the direction and degree of migration (gene flow).
Fig 5.
Population structures for assorted northern Eurasian breeds.
(A) Population structure for Finnish spitz breeds and Laikas. Note the ancestral variability within the Laika breeds. (B) Population structure of selected breeds showing influence to or from the Laika breeds. (C) STRUCTURE analysis also detects low levels of gene flow from Finnish Hound to Nordic and Finnish Spitz breeds. A column represents an individual. For simplicity, each breed was assumed as an independent population (K). Population substructures within East-Siberian Laika and Lapphound, revealed by STRUCTURE analysis, are also evident in MDS plots (D, E) where samples from these breeds form separate clusters (dotted line). Different dot colors mark countries of origin (Finland blue, others vary). No obvious geographical differentiation can be observed.
Fig 6.
A possible scenario for the origin of some Fennoscandian hunting spitz breeds as interpreted from the TreeMix analysis.
Finnish Spitz, Nordic Spitz and the Karelian Bear Dog share rather recent common ancestry and are closely related with the Laika and the reindeer herding breeds. While it is possible that Karelian Bear Dog has contributed to the founders of the contemporary Russian-European Laika, it shows no evidence of admixture from the other analyzed breeds (Fig 5). Instead, both the western Laika breed as well as the herder breeds have influenced the ancestral population of Finnish and Nordic Spitz.