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ADDITIONS TO WIKIPEDIA PAGES

1st page: https://en.wikipedia.org/wiki/Origin_of_the_domestic_dog

Origin of the Domestic Dog -> DNA Evidence -> The Dog is a descendant of a wolf that exists today

In 2002, this amount of populations was narrowed down to a possibility of at least five separate female lines when different dog haplotypes were split into five differing clades.

Origin of the Domestic Dog -> DNA Evidence -> The Dog is a descendant of a wolf that no longer exists today -> The location of the first domestication debate

In 1997, a study determined that domestic dogs could have originated as much as 135,000 years ago. This was determined by using mitochondrial DNA (mtDNA). However, this number could potentially be overinflated due to the possibility of unobserved multiple substitutions at hypervariable sites. These hypervariable sites are mutational hotspots where somatic mitochondrial DNA and germline mutations usually take place. Though this number of 135,000 years may not be completely accurate, it still gives the indication that domesticated dogs have been around for longer than 14,000 years, which was previously suggested by archaeological evidence.

2nd page: https://en.wikipedia.org/wiki/Dog#Differences_from_wolves

Dog -> Differences from wolves -> Behavioral differences

In 1982, a study to observe the differences between dogs and wolves raised in similar conditions took place. The dog puppies preferred large amounts of sleep at the beginning of their lives, while the wolf puppies were much more active. The dog puppies also preferred the company of humans, rather than their canine foster mother, though the wolf puppies were the exact opposite, spending more time with their foster mother. The dogs also showed a greater interest in the food given to them and paid little attention to their surroundings, while the wolf puppies found their surroundings to be much more intriguing than their food or food bowl. The wolf puppies were observed taking part in agonistic play at a younger age, while the dog puppies did not display dominant/submissive roles until they were much older. The wolf puppies were rarely seen as being aggressive to each other or towards the other canines. On the other hand, the dog puppies were much more aggressive to each other and other canines, often seen full-on attacking their foster mother or one another.

FINAL DRAFT STARTS HERE

Place and Time of Origin for Domestic Dogs from Wolves

The exact circumstances surrounding the domestication of dogs has remained shrouded in mystery for some time now. The geographic location and time period of origin have yet to be uncovered, and even the amount of wolf lineages that dogs derive from is uncertain. Having this information would allow researchers to delve further into the history of dog domestication and to see specifics as to how this evolution took place. There are several different theories about the location and time in which domestic dogs split from the wolf population. This information comes from the evaluation of mitochondrial DNA in comparison with archaeological evidence. Using the former data allows a large collection of dogs to be separated into multiple clades, and then further examined to better determine a probable time and place of origin. Also, the origination of domestic dogs from wolves can be explored by looking at MHC loci, mtDNA sequences, behavior, and morphology.

The similarities between domestic dogs and wolves can be seen through their morphologies, behavior, and DNA. DNA gives the most amount of evidence that dogs split off from wolves some time ago. When looking at the MHC loci, also known as the major histocompatibility complex where variation is caused by powerful genetic mechanisms, it is easy to see where dogs and wolves are similar, genetically speaking (Janeway et al. 2001). The MHC loci consists of the class II DQA, DQB, and DRB1 loci (Seddon and Ellegren 2002). Each of these class II loci refer to a different region of the MHC where proteins interact with promoters (Ting and Trowsdale 2002). In one study, at the DQA, all of the alleles in European wolves were found to be shared with either North American wolves, dogs, or both (Seddon and Ellegren 2002). At the same time, there were six DQB and nine DRB1 alleles that were only discovered in European wolves, and not shared with the other two previously mentioned groups (Seddon and Ellegren 2002). There are several ways in which this lack of shared DQB and DRB1 alleles can be explained, the first being that as the domestic dog population has increased with strong genetic drift in dog breeds and reduced intra-breed variability, the wolf population has decreased along with genetic drift and a drop in alleles (Seddon and Ellegren 2002). The second explanation goes along with the first in that the MHC alleles for dogs in this experiment came from more than 800 dogs, and the number of specimens from wolves was less than a quarter of that amount (Seddon and Ellegren 2002). The next cause could have come from a bias in the wolf sampling sites that seemed to favor northern Europe, which would have led to the lack of allelic overlap that was seen (Seddon and Ellegren 2002). The reduced overlap of alleles that was seen could indicate non-stochastic factors (Seddon and Ellegren 2002). The last way that the absence of shared DQB and DRB1 alleles can be described looks at what exactly domestic dogs come into contact with that wolves do not. Dogs spend much of their time around humans and other domesticated animals, whereas wolves do not as frequently see such organisms (Seddon and Ellegren 2002). Being in contact with such creatures would cause changes in domesticated dogs that could have a disruptive effect on certain pressures that maintain the polymorphism that is seen in the MHC loci, such as infectious diseases that are seen in dogs but not in wolves (Seddon and Ellegren 2002).

The DNA taken from dogs has also been compared to that of jackals and coyotes, though the results were not nearly as definitive as when they were put up against wolf DNA. It has been seen that dog mitochondrial DNA sequences did not differ by more than twelve substitutions from any wolf sequence, while dogs differed by at least twenty substitutions from coyotes and jackals (Vilà et al. 1997). Coyotes have not been shown to share many genetic aspects with dogs at all. When looking at the major histocompatibility loci this is also displayed. The DRB1 locus has shown that out of the five alleles coyotes have in this region they share only one with wolves or dogs (Seddon and Ellegren 2002). At the DQA locus coyotes even have an allele that is completely unique from anything that wolves or dogs display (Seddon and Ellegren 2002). This evidence cannot be taken as completely definitive proof of how distantly related dogs are to coyotes since the data for coyotes is much more limited than that of wolves and dogs, which could possibly explain these differences (Seddon and Ellegren 2002). The mtDNA that was used to determine the wolf ancestry for dogs could also be called into question when it comes to this matter. Mitochondrial DNA is inherited from the mother, making it possible that some dog traits could have been received from male jackals and coyotes and this would not have been able to be detected (Vilà et al. 1997). Even with these possibilities, wolves are still seen as being more closely related to dogs than any other type of canid.

Wolves and domesticated dogs share a large amount of morphological aspects. They do have their differences, but overall, it is clear that the two groups are closely related. Although, there are many contrasting appearances that dogs have, depending on what they have been bred for. This gradual change in morphology could have originated when humans first created agricultural settlements (Vilà et al. 1997). This change in civilization type would mean that certain aspects were no longer needed like they were in the previous hunting and gathering societies (Vilà et al. 1997). It is possible that before this, domestic dogs did not vary as much physically from wolves as they do now (Vilà et al. 1997). The likelihood that dogs and wolves continued to interbreed could also have contributed to the artificial selection that began to take place in domestic dogs, which can be seen today in the extensive physical variations between dog species (Vilà et al. 1997). The genetic material provided by wolves in the case of backcrossing events would have allowed for the preservation of specific genes displayed in certain dogs more than in others (Vilà et al. 1997).

The behaviors displayed by wolves and dogs vary as much as their morphology. In a study that observed wolf puppies and Malamute puppies growing up in the same circumstances, these contrasting actions were made more obvious. The wolf puppies were much more active at the beginning of their lives than the Malamute puppies, and they preferred the company of their canine foster mother while the Malamutes were more excited about the presence of humans (Frank and Frank 1982). The Malamutes showed a greater interest in their food and the space around the food bowl, while the wolf puppies were more intrigued by their surroundings rather than the food given to them (Frank and Frank 1982). The wolf puppies were also observed taking part in agonistic play at a younger age, while the Malamute puppies did not display dominant/submissive roles until they were older (Frank and Frank 1982). The most interesting fact is that the wolf puppies were rarely seen as aggressive, either towards one another or towards other canines (Frank and Frank 1982). On the other hand, the Malamute puppies were much more aggressive to each other and other canines, often seen full-on attacking their foster mother or one another (Frank and Frank 1982). It is possible that the wolf puppies have an innate knowledge that constantly being aggressive towards others will result in a waste of precious energy, and that they should follow the pack dynamic instead of continually fighting about it. The determination of exactly when the origin of domestic dogs took place has proven to be extremely difficult. Archaeological evidence shows that the earliest findings of domestic dogs are a group of small canids found in Israel 12,000 years ago and a single jaw found in Germany 14,000 years ago (Seddon and Ellegren 2002; Savolainen et al. 2002). Even with these specimens, it is difficult to tell if these are from small wolves or actual domestic dogs, since they were so morphologically similar until relatively recently (Savolainen et al. 2002). Evidence from mtDNA now suggests that dogs could have originated as much as 135,000 years ago (Vilà et al. 1997; Seddon and Ellegren 2002). However, this number could be overinflated due to the possibility of unobserved multiple substitutions at hypervariable sites (Vilà et al. 1997). These sites are mutational hotspots where somatic mtDNA and germline mutations usually occur (Stoneking 2000). Even if this amount of 135,000 years ago is not completely accurate, it still gives the indication that domesticated dogs have been around for longer than 14,000 years, as was previously suggested by archaeological remains (Vilà et al. 1997). The values of 135,000 years and 14,000 years are both recent enough to state that many of the alleles found in modern dogs are derived from wolves (Seddon and Ellegren 2002). The extreme variety of the alleles found in dogs suggests a large founding population, in order to account for this diversity (Seddon and Ellegren 2002).

Figuring out the exact geographic position of the origins for domestic dogs is also a challenge. The extreme mobility of wolf populations makes it difficult to determine, since the geographic paths of wolves cannot be seen when looking at their DNA (Savolainen et al. 2002).When different dog haplotypes were split into five differing clades, they were found to be of equal distance from the cenancestor of all wolves and dogs, which leads to the belief that the domestic dog population originated from at least five separate female lines (Savolainen et al. 2002). A cenancestor is described as the last universal common ancestor (Delaye and Becerra 2012). The largest of these aforementioned clades had three haplotypes from China and Mongolia, which would suggest an origin in East Asia, with this claim having support from other research (Savolainen et al. 2002; Tanabe et al. 1991). More haplotypes were discovered among this clade than the other two major clades and a large amount of these haplotypes were found to be unique to East Asia (Savolainen et al. 2002). One of the other major clades was shown to originate from the introduction of a subset of East Asian types, even though it was found to have types in the western part of the world (Savolainen et al. 2002). No archaeological evidence has been found to give support to this claim that domestic dogs originated in East Asia, but the archaeological work done in the area has been very limited for the late Paleolithic time period (Savolainen et al. 2002). There is evidence of domestic dogs that has been found in the earliest Neolithic which dates 7,500 years ago (Savolainen et al. 2002).

The origin of dogs from wolves has been known for a time, and this can be clearly seen by comparing their MHC loci and the alleles found there, as well as their morphologies. Their similar appearances can cause problems when looking at archaeological evidence, because it is often difficult to tell the difference between ancient wolves and dogs. The behavior of domesticated dogs has shifted a bit from that of wolves, but similarities can still be seen. It is much less likely that dogs are as closely related to coyotes and jackals as they are to wolves, but there is not enough evidence to completely disprove this. Even though the fact that dogs are derived from wolves is apparent no one has been entirely sure as to where and when this event took place. The data expressed in the paragraphs above states that the origin of domestic dogs from wolves took place in East Asia around 135,000 years ago. The mitochondrial DNA evidence gives a large amount of support to these claims, but it cannot account for wolf mobility or the fact that no archaeological data has yet been found to support these theories.

References

Delaye, L., A. Becerra. 2012. Cenancestor, the last universal common ancestor. Evolution: Education and Outreach 5(3): 382-388.

Frank, H., M. G. Frank. 1982. On the effects of domestication on canine social development and behavior. Applied Animal Ethology 8: 507-525.

Janeway, C. A., P. Travers, M. Walport, M. J. Shlomchik. 2001. The major histocompatibility complex and its functions. Garland Science, New York.

Savolainen, P., Y. Zhang, J. Luo, J. Lundeberg, T. Leitner. 2002. Genetic evidence for an east Asian origin of domestic dogs. Science 22: 1610-1613.

Seddon, J. M., H. Ellegren. 2002. MHC class II genes in European wolves: a comparison with dogs. Immunogenetics 54: 490-500.

Stoneking, M. 2000. Hypervariable sites in the mtDNA control region are mutational hotspots. American Journal of Human Genetics 67(4): 1029-1032.

Tanabe, Y., K. Ôta, S. Ito, Y. Hashimoto, Y. Y. Sung, J. K. Ryu, M. O. Faruque. 1991. Biochemical-genetic relationships among Asian and European dogs and the ancestry of the Japanese native dog. Journal of Animal Breeding and Genetics 108: 455–478.

Ting, J. P. Y., J. Trowsdale. 2002. Genetic control of MHC class II expression. Cell 109: 21-33.

Vilà, C., P. Savolainen, J. E. Maldonado, I. R. Amorim, J. E. Rice, R. L. Honeycutt, K. A. Crandall, J. Lundeberg, R. K. Wayne. 1997. Multiple and ancient origins of the domestic dog. Science 276: 1687– 1689.

Domestication of Dogs

Blood samples were taken from 2,959 dogs of 40 different breeds found in Japan, Korea, Taiwan, Bangladesh, and Europe. These blood samples were found to state that the native dogs were a crossbreed from a breed of dog in Southeast Asia and another dog from the Korean peninsula.

Tanabe Y., Ôta K., Ito S., Hashimoto Y., Sung Y. Y., Ryu J. K. and Faruque M. O. 1991. Biochemical-genetic relationships among Asian and European dogs and the ancestry of the Japanese native dog. Journal of Animal Breeding and Genetics 108, 455–478.

Blood and hair samples from 19 different dogs were taken and compared with samples from 27 populations of wolves from Europe, Asia, and North America and 67 different breeds. The samples taken from Xoloitzcuintlis (or Mexican hairless) suggest that it is derived from a very large and diverse population of dogs.

Vila C, Savolainen P, Maldonado JE, Amorim IR, Rice JE, Honeycutt RL, Crandall KA, Lundeberg J, Wayne RK. 1997. Multiple and ancient origins of the domestic dog. Science 276, 1687– 1689.

The large number of alleles found in dogs suggests that the founding population was very large and genetically diverse. The organisms that dogs come into contact with (and wolves do not) have been found to correspond with certain infectious diseases.

Seddon JM, Ellegren H. 2002. MHC class II genes in European wolves: a comparison with dogs. Immunogenetics 54, 490-500.

Two groups of puppies (one consisting of 4 Eastern timber wolves, and the other of 4 Alaskan Malamutes) were reared in identical conditions, and their development and behavior were compared. It was observed that the wolf pups developed more quickly than the Malamute pups, in social and behavioral aspects.

Frank H, Frank MG. 1982. On the effects of domestication on canine social development and behavior. Applied Animal Ethology 8, 507-525.

The mitochondrial DNA of 654 domestic dogs was tested in order to determine when and where the domestic dog was first founded. Although many of the dogs were found to be related to the wolf, there was still a large amount that seems to have an East Asian origin, dating back around 15,000 years ago.

Savolainen P, Zhang Y, Luo J, Lundeberg J, Leitner T. 2002. Genetic evidence for an east Asian origin of domestic dogs. Science 22, 1610-1613.

Wikipedia Assignment 2
https://en.wikipedia.org/wiki/Origin_of_the_domestic_dog

It has been seen that these traits can even prompt an adult female wolf to act more defensively of dog puppies than of wolf puppies.

Adding to Specialization
A type of breed of dog called "lupine", describing dogs that still have the same size and morphology as a wolf, could be added to the list of dog breed types (On the effects of domestication on canine social development and behavior 1982)

Adding Comparison of Social and Behavioral Development in Dogs and Wolves
This article does not extensively discuss how the behaviors of domestic dogs and wild wolves different. The morphological aspects are mentioned, but there is so much more that could be added. From an early age, wolves and dogs differ very differently, in ways such as submission, activeness, and aggression (On the effects of domestication on canine social development and behavior 1982)

Adding to DNA Evidence
A possible way to determine the geographic arrangement of the origin for dog populations in latitudinal and longitudinal directions could be determined using the frequencies at various loci (Biochemical-genetic relationships among Asian and European dogs and the ancestry of the Japanese native dog 1991)