Genetics of Domesticated Behavior in Dogs and Foxes - Genetics and the Behavior of Domestic Animals - page 358

the differences in social behavior of dogs and wolves are undeniably genetic,
little is known about the evolutionary processes that led to these behavioral
modifications. In the mid-20th century, the Russian geneticist, Dmitry
Belyaev, argued for a primary role of selection for behavior in animal domes-
tication. To test this hypothesis, experimental selection of the silver fox (color
morph of the red fox, Vulpes vulpes), a close relative of the dog, was started in
1959 ( Belyaev, 1969, 1979 ; Trut, 1999, 2001, 2009 ). The experiment led to
the development of a strain of foxes that show friendly, dog-like behavior to
humans. Selecting foxes for behavior constituted a completely novel
long-range approach to the genetics of behavioral modification that preceded
the demonstration that dogs evolved from wolves ( Axelsson et al., 2013;
Leonard et al., 2002; Savolainen et al., 2002; Vil ` et al., 1997; vonHoldt
et al., 2010 ).

THE DOG IS THE FIRST DOMESTICATED SPECIES
The dog was the first species to be domesticated. In fact, this “event” took
place so early that it is arguable that the dog self-domesticated itself, and
provided the example that humans subsequently followed in deliberately
domesticating other species.
Archeological evidence for the coexistence of dogs with humans has been
identified from as early as 14,000 17,000 ybp in Russia ( Sablin and
Khlopachev, 2002 ); 14,000 ybp in Germany ( Nobis, 1979 ); 12,000 ybp in
Israel ( Dayan, 1994; Davis and Valla, 1978; Tchernov and Valla, 1997 ); and
5000 ybp ( Olsen, 1985 ) in China. Recent findings identified remains of dog-
like canids living prior to the Last Glacial Maximum (c. 26,500

19,000 cal
BP) in southern Siberia (Altai Mountains, Russia) dated c. 33,000 cal BP
( Ovodov et al., 2011 ), and Western Europe (Goyet, Belguim) dated 36,000 cal
BP ( Germonpr´ et al., 2009 ). Although these incipient dogs may represent
domestication events that were terminated by climatic changes associated
with the Last Glacial Maximum, and thus may have not given rise to modern
dogs, molecular analysis of these remains may still shed light on the evolution
of domestication in the dog. Identification of futher dog-like canid remains in
different parts of Eurasia supports a hypothesis of recurrent, multiregional dog
domestication ( Ovodov et al., 2011 ).
Molecular data suggest that the divergence of dogs from wolves took
place as recently as 15,000 years ago ( Savolainen et al., 2002; Skoglund
et al., 2011 ). Initial molecular analyses, based on mitochondrial DNA and
Y-chromosome markers from dog breeds and wolf populations from different
geographical locations, suggested an East-Asian origin for canine domestica-
tion ( Brown et al., 2011; Ding et al., 2012; Pang et al., 2009; Savolainen
et al., 2002 ; Oskarsson et al., 2012 ); while genomic DNA analysis supports
the Middle East ( vonHoldt et al., 2010 ) as the most likely center. Studies of
ancient dog remains from Latin America and Alaska, and dog breeds