How does evolutionary evaluation illuminate body size among canids?

In this review, we examine mammalian body size as it reflects life history and genomic composition, with a primary focus on canids and the domestication of the gray wolf. The range of variation in body size is greater among Carnivora than any other terrestrial order. In the Canidae, this range is some 2 orders of magnitude. Macroevolutionary patterns (eg, Bergmann's rule and Cope's rule) that have been proposed in the past often fail to comport with modern studies on this aspect of carnivoran evolution. Clades often begin with small to medium size (mesocarnivorans) and diversify mostly in a right-skewed (larger) direction. The observed variation in body size reflects phenotypic plasticity in response to life history. As with many Mammalia, historically high gene flow (hybridization and introgression) among canid lineages has been a crucial source of genomic variation (nuclear and mitochondrial), yielding the potential for high plasticity of phenotypes such as body size. In addition, epigenetic marks connect genetic expression with environmental conditions in the manifested phenotypes. Among Mammalia generally, a larger size is associated with a longer life span, reflecting the foregoing genomic composition and environmental influences over a long geological time. However, the larger modern domestic dog breeds trend toward shorter life spans. The latter appears to reflect genetically mediated phenotypes that emerged secondary to domestication but nonetheless against a background of broadly and deeply conserved developmental and physiological patterns and body plans.

Developmental features of the canid proximocaudal femur.

We continued direct morphological studies of the canid coxofemoral joint, considering early-life spatial relationships around the locus of the proximocaudal joint capsule insertion. Our primary goal was to elucidate the postnatal developmental gross anatomy of the proximocaudal femur, among juveniles across Canidae. From an original database of 267 independent (museum) specimens from 11 canid taxa and 1 hybrid taxon, we identified 29 ancient or modern candidate juvenile specimens (nine taxa and one hybrid taxon). Based on optimal ability to recognize landmarks, the best photographic data were categorized into five groups of four each (n = 20). The data groups approximated early juvenile, early-mid juvenile, mid-juvenile, mid-late juvenile; and young adult stages. In this descriptive photographic essay, we demonstrate the developmental spatial proximity among (a) the dorsal meeting of the respective lateral and medial extensions from the growth centers of the femoral head and greater trochanter; (b) the caudodorsal aspect of the coxofemoral joint capsule attachment; (c) a segment of the proximocaudal femoral shaft physis; and (d) an eventual associated mineralized prominence. The latter occurs frequently but not universally, suggesting natural population variability across taxa. Across taxa and juvenile age categories, the morphology thus supports developmental conservation among ancient and modern Canidae. The biomechanical and biological cause-effect implications are not yet clear. For zoological purposes, we apply the term postdevelopmental mineralized prominence to the residual caudolateral surface feature. We extend the original anatomical work of Morgan in zoological and phylogenic arenas, using direct observation of cleared skeletal specimens.

Osteoarthritis from evolutionary and mechanistic perspectives.

Developmental osteogenesis and the pathologies associated with tissues that normally are mineralized are active areas of research. All of the basic cell types of skeletal tissue evolved in early aquatic vertebrates. Their characteristics, transcription factors, and signaling pathways have been conserved, even as they adapted to the challenge imposed by gravity in the transition to terrestrial existence. The response to excess mechanical stress (among other factors) can be expressed in the pathologic phenotype described as osteoarthritis (OA). OA is mediated by epigenetic modification of the same conserved developmental gene networks, rather than by gene mutations or new chemical signaling pathways. Thus, these responses have their evolutionary roots in morphogenesis. Epigenetic channeling and heterochrony, orchestrated primarily by microRNAs, maintain the sequence of these responses, while allowing variation in their timing that depends at least partly on the life history of the individual.