SummaryIn morphological terms, “form” is used to describe an object’s shape and size. In dogs, facial form is stunningly diverse. Facial retrusion, the proximodistal shortening of the snout and widening of the hard palate is common to brachycephalic dogs and is a welfare concern, as the incidence of respiratory distress and ocular trauma observed in this class of dogs is highly correlated with their skull form. Progress to identify the molecular underpinnings of facial retrusion is limited to association of a missense mutation in BMP3 among small brachycephalic dogs. Here, we used morphometrics of skull isosurfaces derived from 374 pedigree and mixed-breed dogs to dissect the genetics of skull form. Through deconvolution of facial forms, we identified quantitative trait loci that are responsible for canine facial shapes and sizes. Our novel insights include recognition that the FGF4 retrogene insertion, previously associated with appendicular chondrodysplasia, also reduces neurocranium size. Focusing on facial shape, we resolved a quantitative trait locus on canine chromosome 1 to a 188-kb critical interval that encompasses SMOC2. An intronic, transposable element within SMOC2 promotes the utilization of cryptic splice sites, causing its incorporation into transcripts, and drastically reduces SMOC2 gene expression in brachycephalic dogs. SMOC2 disruption affects the facial skeleton in a dose-dependent manner. The size effects of the associated SMOC2 haplotype are profound, accounting for 36% of facial length variation in the dogs we tested. Our data bring new focus to SMOC2 by highlighting its clinical implications in both human and veterinary medicine.

In most animal taxa, longevity increases with body size across species, as predicted by the oxidative stress theory of aging. In contrast, in within-species comparisons of mammals and especially domestic dogs (e.g. Patronek et al., '97; Michell, '99; Egenvall et al., 2000; Speakman et al., 2003), longevity decreases with body size. We explore two datasets for dogs and find support for a negative relationship between size and longevity if we consider variation across breeds. Within breeds, however, the relationship is not negative and is slightly, but significantly, positive in the larger of the two datasets. The negative across-breed relationship is probably the consequence of short life spans in large breeds. Artificial selection for extremely high growth rates in large breeds appears to have led to developmental diseases that seriously diminish longevity.

The grey wolf (Canis lupus) was the first species to give rise to a domestic population, and they remained widespread throughout the last Ice Age when many other large mammal species went extinct. Little is known, however, about the history and possible extinction of past wolf populations or when and where the wolf progenitors of the present-day dog lineage (Canis familiaris) lived1–8. Here we analysed 72 ancient wolf genomes spanning the last 100,000 years from Europe, Siberia and North America. We found that wolf populations were highly connected throughout the Late Pleistocene, with levels of differentiation an order of magnitude lower than they are today. This population connectivity allowed us to detect natural selection across the time series, including rapid fixation of mutations in the gene IFT88 40,000–30,000 years ago. We show that dogs are overall more closely related to ancient wolves from eastern Eurasia than to those from western Eurasia, suggesting a domestication process in the east. However, we also found that dogs in the Near East and Africa derive up to half of their ancestry from a distinct population related to modern southwest Eurasian wolves, reflecting either an independent domestication process or admixture from local wolves. None of the analysed ancient wolf genomes is a direct match for either of these dog ancestries, meaning that the exact progenitor populations remain to be located.

A 19 th -century dog burial uncovered from a historical homelot in Toronto, Canada, provided a unique opportunity to reconstruct the individual's osteobiography. Of particular interest are the dog's very large size and a suite of skeletal pathologies. Recovery of a nearly complete skeleton combined with the use of x-rays and micro-computed tomography (micro-CT) allowed for a discriminating differential diagnoses. Stable isotope analyses were applied to investigate questions of diet. Results reveal an individual who suffered greatly from disease towards the end of his life and hint at its owners attitudes towards dogs. The interdisciplinary approach applied to this case study highlights the potential information obtainable from pet burials. We argue that better analyses and reporting of pet burials will help address research questions targeting broader themes related to human-animal relationships.2

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