วันอังคารที่ 19 พฤษภาคม พ.ศ. 2552

Taxonomy and evolution

In Jan van Eyck's famous Arnolfini Portrait (1434), care was taken to include the couple's little pet dog.

The domestic dog was originally classified as Canis familiaris and Canis familiarus domesticus by Linnaeus in 1758, and is currently classified as Canis lupus familiaris, a subspecies of the gray wolf Canis lupus, by the Smithsonian Institution and the American Society of Mammalogists. Overwhelming evidence from behavior, vocalizations, morphology, and molecular biology led to the contemporary scientific understanding that a single species, the gray wolf, is the common ancestor for all breeds of domestic dogs, however the timeframe and mechanisms by which dogs diverged are controversial.

The current consensus among biologists and archaeologists is that no one can be sure when dogs were domesticated. There is conclusive evidence that dogs genetically diverged from their wolf ancestors at least 15,000 years ago but some believe domestication to have occurred earlier. The bulk of the scientific evidence for the evolution of the domestic dog stems from archaeological findings and mitochondrial DNA studies. The divergence date of roughly 15000 years ago is based in part on archaeological evidence that demonstrates that the domestication of dogs occurred more than 15,000 years ago, and some genetic evidence indicates that the domestication of dogs from their wolf ancestors began in the late Upper Paleolithic close to the Pleistocene/Holocene boundary, between 17,000 and 14,000 years ago. But there is a wide range of other, contradictory findings that make this issue controversial.

Archaeological evidence plays a large role in this debate. In 2008, a team of international scientists released findings from an excavation at Goyet Cave in Belgium declaring that a large, toothy canine existed 31,700 years ago and ate a diet of horse, musk ox and reindeer. The morphology of the skeleton and the context in which it was found led researchers to hypothesize that this canine was an ancient domesticated dog. Prior to this Belgium discovery, the earliest dog fossils were two large skulls from Russia and a mandible from Germany, that dated from roughly 14,000 years ago. Remains of smaller dogs from Natufian cave deposits in the Middle East have been dated to around 10,000 to 12,000 years ago. There is a great deal of archaeological evidence for dogs throughout Europe and Asia around this period and through the next two thousand years (roughly 8,000 to 10,000 years ago), with fossils uncovered in Germany, the French Alps, and Iraq, and cave paintings in Turkey.

This ancient mosaic, likely Roman, shows a large dog with a collar hunting a lion.

Thus, the archaeological evidence suggests that the latest dogs could have diverged from wolves was roughly 15000 years ago, although it is possible that they diverged much earlier.
DNA studies have provided a wider range of possible divergence dates, from 15,000 to 40,000 years ago, to as much as 100,000 to 140,000 years ago. This evidence depends on a number of assumptions that may be violated. Genetic studies are based on comparisons of genetic diversity between species, and depend on a calibration date. Many estimates of divergence dates from DNA evidence use an estimated wolf-coyote divergence date (roughly 1 million years ago) as a calibration. If this estimate is incorrect, and the actual wolf-coyote divergence is closer to 750,000 or 2 million years ago, then the DNA evidence that supports specific dog-wolf divergence dates would be interpreted very differently. Furthermore, it is believed that the genetic diversity of wolves has been in decline for the last 200 years, and that the genetic diversity of dogs has been reduced by selective breeding. This could significantly bias DNA analyses to support an earlier divergence date. The genetic evidence for the domestication event occurring in East Asia is also subject to violations of assumptions. These conclusions are based on the location of maximal genetic divergence, and assume that hybridization does not occur, and that breeds remain geographically localized. Although these assumptions hold for many species, there is good reason to believe that they do not hold for canines.

Genetic analyses indicate all dogs are likely descended from a handful of domestication events with a small number of founding females, although there is evidence that domesticated dogs interbred with local populations of wild wolves on several occasions. Data suggests that dogs first diverged from wolves in East Asia, and that these domesticated dogs then quickly migrated throughout the world, reaching the North American continent around 8000 B.C. The oldest groups of dogs, which show the greatest genetic variability and are the most similar to their wolf ancestors, are primarily Asian and African breeds, including the Basenji, Lhasa Apso, and Siberian Husky. Some breeds that were thought to be very old, such as the Pharaoh Hound, Ibizan Hound, and Norwegian Elkhound, are now known to have been created more recently.

There is a great deal of controversy surrounding the evolutionary framework for the domestication of dogs. Although it is widely claimed that "man domesticated the wolf," man may not have taken such a proactive role in the process. The nature of the interaction between man and wolf that led to domestication is unknown and controversial. At least three early species of the Homo genus began spreading out of Africa roughly 400,000 years ago, and thus lived for a considerable period in contact with canine species. Despite this, there is no evidence of any adaptation of canine species to the presence of the close relatives of modern man. If dogs were domesticated, as believed, roughly 15,000 years ago, the event (or events) would have coincided with a large expansion in human territory and the development of agriculture. This has led some biologists to suggest that one of the forces that led to the domestication of dogs was a shift in human lifestyle in the form of established human settlements. Permanent settlements would have coincided with a greater amount of disposable food and would have created a barrier between wild and anthropogenic canine populations.

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