THE HISTORY OF DOGS. By Gareth Johnston - Johnston K9 - Sept 2011

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1 THE HISTORY OF DOGS By Gareth Johnston - Johnston K9 - Sept 2011 The History of Dogs The archaeological record cannot resolve whether domestic dogs originated from a single wolf population or arose from multiple populations at different times. However, evidence suggests that dogs may have diverse origins. Wolves During most of the late Pleistocene, humans and wolves coexisted over a wide geographic area, providing ample opportunity for independent domestication events and continued genetic exchange between wolves and dogs. The extreme phenotypic diversity of dogs, even during the early stages of domestication also suggests a varied genetic heritage. The earliest remains of the dog are found in the Upper Phocene deposits and at the end of the Tertiary period. Dog There have been five distinct varieties discovered at different times, but the three most important are the Canis familiar ispalustris, Canis familiar isanutschin and the Canis familiaris of the Bronze Age. The skull of the first-named (Canis familiar ispalustris) represents a smallish dog, and would seem to be the parent of the Chows, Pomeranian and Spitz dogs. The skull of the Bronze Age dog shows a much larger development, and is practically identical with the modern sheep dog. Some dog skulls, which were dug up in Roman remains, closely resemble those of the Bronze period, and also of our sheep dogs, and form an interesting link between the dog of prehistoric man and our own, and also show how true to type this particular dog seems to have remained, in essential characteristics. Early Dog Skull The actual origin of the dog, that is to say, whether it has always been a species by itself, or whether it is a descendant of a wild animal, such as the wolf or jackal, has never been satisfactorily decided.

2 Many naturalists and scientific men have come to the conclusion, that it has always been a distinct species, and certainly, the more you study the subject, the greater does the likelihood seem that they are right. Count Buffon, in his 1749 work "Natural History", gives an account of an experiment to breed a dog with a wolf: "In our climates the wolf and fox make the nearest approach to the dog, particularly the shepherd's dog, which I consider as the original stock of the species; and as their internal structure is almost entirely the same and their external differences very slight, I wished to try to see if they would intermix and produce together.... With this in view, I reared a she-wolf, taken in the woods at the age of three months, along with an Irish greyhound of the same age. They were shut up together in a pretty large court, to which no other beast could have access, and they were provided with shelter for their retirement. They were kept in this state three years, without the smallest restraint. During the first year they played perpetually, and seemed extremely fond of each other. Count Buffon (Georges-Louis Leclerc) French naturalist and philosopher The second year they began to quarrel about their food, though they were supplied in a plentiful manner. The wolf always began the dispute. Instead of seizing the meat, the wolf instantly drove off the dog, then laid hold of the edge of the plate so dexterously, as to allow nothing to fall, and carried off the whole. I have seen her run in this manner five or six times round the wall, and never set it down, save to take breath, to devour the meat, or to attack the dog when he approached. After the second year these quarrels and combats became more frequent and more serious. In the third year it was hoped to breed from them, but this condition, instead of softening them and making them approach each other, rendered them more intractable and ferocious. Nothing now was heard but dismal howling s and cries of resentment." This sad story is closed with an account of the destruction of the wolf by the dog. Count Buffon also tried similar experiments with foxes and dogs, but the result was the same strong antagonism or indifference. With this in mind it would seem that Count Buffon is saying that it seems impossible to credit any race, with such intensely strong natural antipathy to each other, can at any time have been united in a common ancestor. And the fact that, although so like each other in structure and appearance, this great antagonism does exist, seems to emphasize with corresponding significance the curious differences in the appearance of dogs themselves, and yet the entire absence of enmity. For instance, the English Mastiff (Top Right) and the Pomeranian (Bottom Right) are, in appearance, as opposite as possible, and yet they each recognize in each other the same species, and if introduced correctly will be happy to live together, unlike Count Buffon s She-Wolf and Irish Greyhound. The English Mastiff

3 Consequently, it may be correct to assume that once, a long time ago, all canines shared a common ancestor, however they quickly diversified and became different things, that look similar. It is possible to assume that the genetic diversity of The Pomeranian dogs may have been enriched by multiple founding events, possibly followed by occasional interbreeding with wild wolf populations. Just because Count Buffon couldn t mate two different species it does not mean that this cannot happen, it can and does. Indeed we are interbreeding dogs and wolves today so there is no reason to assume that this was not possible in the past. I think what Count Buffon s experiments show us, is that although some of these animals may look the same, they are different, and they know it. There is no doubt that scent enters largely into the question of species. As human beings, we have, to a great extent, lost all understanding of the properties of scent, as understood by dogs and the rest of the animal kingdom. They are far ahead of us in this respect. The greater part of their powers of recognition comes from this quality of scent, and they also use it as a means of communication from the one to the other. I believe that there is a great deal of separation that exists, between the dog, and those animals of similar appearance. The body scent is completely different, and we human beings are unable to explain the meaning of the difference, because we do not understand the language of scent. Count Buffon continued his experiments with wolves and wolf cubs discovering that the natural habits and instincts of the latter varied from the dog in many ways, from the selection of a mate to period of gestation and even the methods used to rear the young cubs. He summed up his research in the following downright sentence: The wolf and the dog have never been regarded as the same animal, but by the nomenclatures of Nature History, who, being acquainted with the surface of nature only, never extend their views beyond their own methods, which are always deceitful and often erroneous, even in the most obvious facts" (Count Buffon, 1749).

4 What Do The Genetic Scientists Say? This is quite interesting as test that have been undertaken around the world are showing that Count Buffon s claims made nearly 300 years ago may be correct. In an experiment called The Multiple and Ancient Origins of the Domestic Dog, conducted by; Caries Vila, Peter Savolainen, Jesus E. Maldonado, Isabel R. Amorim, John E. Rice, Rodney L. Honeycutt, Keith A. Crandall, Joakim Lundeberg, Robert K. Wayne. Scientists sequenced portions of the mitochondrial DNA of wolves and domestic dogs. Initially, 261 base pairs (bp) of the left domain of the mitochondrial control region were sequenced from 140 dogs representing 67 breeds and five cross-breeds and 162 wolves representing 27 populations from throughout Europe, Asia, and North America. Because all wild species of the genus Canis can interbreed and thus are potential ancestors of the domestic dog, five coyotes (Canis latrans), and two golden, two blackbacked, and eight Simien jackals (C. aureus, C. mesomelas, and C. simensis, respectively) were also sequenced. Some naturalists now believe that early domesticated dogs looked more like the Tibetan Mastiff (above) than the Siberian Husky (below). Personally I have no idea, and I don't think we will ever know, short of building a time machine. Until such time it will always make for interesting debate. They found the control region of wolves and dogs was highly polymorphic. They found 27 wolf haplotypes that differed on average by / (+/- SE) substitutions (2.10 +/- 0.04%), with a maximum of 10 substitutions (3.95%), and they found out that the distribution of wolf haplotypes demonstrated geographic specificity, with most localities containing haplotypes unique to a particular region. Four haplotypes (W2, W7, W14, and W22) had a widespread distribution. Click here to display their charted results and information on which breeds of dog where studied ---> Wolf / Dog Gene Chart Wolf / Dog Gene Chart Click to enlarge

5 In dogs, 26 haplotypes were found. Only haplotype D6 also occurred in some grey wolves from western Russia and Romania (W6). Sequence divergence among dogs was similar to that found among wolves. Dog haplotypes differed by an average of / substitutions, (2.06 +/- 0.07%), with a maximum divergence of 12 substitutions (4.67%). Mitochondrial haplotype diversity in dogs could not be partitioned according to breeds. For example, in eight German shepherds examined, five distinct sequences were found, and in six golden retrievers, four sequences were detected. Moreover, many breeds shared sequences with other breeds. For instance, dog haplotypes D4, D3, D5, and Dl were found in 14, 14, 9, and 7 breeds, respectively. They also discovered that no dog sequence differed from any wolf sequence by more than 12 substitutions; however dogs differed from coyotes and jackals by at least 20 substitutions and two insertions. These results clearly support wolf ancestry for dogs. However, because mitochondrial DNA is maternally inherited, interbreeding between female dogs and male coyotes or jackals would not be detected. More limited studies of nuclear markers support the conclusion that the wolf was the ancestor of the domestic dog. Several methods of phylogenetic analysis, including maximum likelihood, maximum parsimony, minimum spanning networks, and statistical parsimony, were used to investigate relationships among sequences. All analyses supported a grouping of dog haplotypes into four distinct clades, although the topology within and among clades differed among trees. The image to the right exemplifies this (Fig. A) Three of the four monophyletic clades defined a larger clade containing all but three dog haplotypes and a subset of wolf haplotypes (W4 and W5). Clade I included 19 of the 26 dog haplotypes. This group contained representatives of many common breeds as well as ancient breeds such as:

6 The Dingo New Guinea Singing Dog African Basenji The Greyhound Clade II included dog haplotype D8, from two Scandinavian breeds (Elkhound and the Jamthund), and was closely related to two wolf haplotypes found in Italy, France, Romania, and Greece (W4 and W5). Clade III contained three dog haplotypes (D7, D19, and D21) found in a variety of breeds such as the German Shepherd, Siberian Husky, and the Mexican Hairless. Clade IV contained three haplotypes (D6, D10, and D24) that were identical or very similar to a wolf haplotype (W6) found in Romania and western Russia, which suggests recent hybridization between dogs and wolves. Many breeds contained representatives of more than one dog haplotype grouping and this can be seen in the Wolf / Dog Gene Chart base pairs of the control region were sequenced for 24 Canids, including representatives of the four dog clades. Although the association of clades was different, the analyses of the longer sequences provided stronger support for the four monophyletic groupings of dog haplotypes. A Wilcoxon signed-rank test was used to assess the monophyly of dog clades. Monophyly of all dog haplotypes can be rejected, and monophyly of clades I, II, and III is marginally rejected (P = and P = 0.053, respectively). In both trees, dog haplotype clakes II and IV are most closely related to wolf sequences from eastern Europe (Greece, Italy, Romania, and western Russia). The coyote and wolf have a sequence divergence of / (17) and diverged about one million years ago, as estimated from the fossil record. Consequently, because the sequence divergence between the most different genotypes in clade I (the most diverse group of dog sequences) is no more than 0.010, this implies that dogs could have originated as much as 135,000 years ago. Although such estimates may be inflated by unobserved multiple substitutions at hyper variable sites, the sequence divergence within clade I clearly implies an origin more ancient than the 14,000 yearsago that is presently suggested by the archaeological record. The Multiple and Ancient Origins of the Domestic Dog experiment has shown that the Rhodesian Ridgeback (above) and the Pug (below) are more closely related to each other than, than they are the wolf.

7 Nevertheless, bones of wolves have been found in association with those of hominids from as early as the middle Pleistocene, up to 400,000 years ago. The ancient dates for domestication based on the control region sequences cannot be explained by the retention of ancestral wolf lineages, because clade I is exclusively monophyletic with respect to dog sequences and thus the separation between dogs and wolves has been long enough for coalescence to have occurred. To explain the discrepancy in dates, the scientists hypothesize that early domestic dogs may not have been morphologically distinct from their wild relatives. Conceivably, the change around 10,000 to 15,000 years ago from nomadic hunter-gatherer societies to more sedentary agricultural population centres may have imposed new selective regimes on dogs that resulted in marked phenotypic divergence from wild wolves. Although individual breeds show uniformity with respect to behaviour and morphology, most breeds show evidence of a genetically diverse heritage because they contain different haplotypes. Moreover, dog sequences cluster differ greatly from groups of wolf haplotypes. Therefore, after the origin of dogs from a wolf ancestor (i.e. after the two species diversified), dogs and wolves may have continued to exchange genes. Backcrossing events could have provided part of the raw material for artificial selection and for the extraordinary degree of phenotypic diversity in the domestic dog. This experiment has also shown just how different two dogs that share much of the same genetic makeup can look. According to their research the Papillion (above) and the Rottweiler (below) have much of the same genes, and just look how different they are, size, weight, tail and ear carriage, all very different. From my research I feel comfortable saying that I do not think that Dogs are the direct descendant of Wolves, rather that somewhere in there make up they share a common prehistoric ancestor, just like the Chimpanzee and the Bonobo share lineage. I feel that just like the Chimpanzee and the Bonobo, they diverted and became a new species in their own right, sharing some of the traits of one another but also having developed new ones of their own a very long time ago. Whilst we may learn new things all the time from observing wild packs of both dogs and wolves it should be remembered that whilst they share some similarities they are not the same.

8 I learnt a lot doing this research and this article is based on what I have read for myself, the information I assimilated and the conclusions I personally draw from it. If you are more interested start researching for yourself and post up your conclusions. Unless someone invents a time machine we will never know for sure, but until then the subject makes for great debate. What Do The History Books Say? Dogs feature in many legends and myths. For example, Argus - the old dog of Odysseus that recognized its master on his return to Ithaca after an absence of 19 years; Bouhaki the pet dog pictured in a tomb in the necropolis of Thebes; Cabal - King Arthur's dog and Cerberus the guard dog of Hades to name but a few. Early in recorded history there are records of dogs being trained for the purpose of attack. One of the most well recorded ancient uses of dogs in a battle was between the Greeks and Corinthians. Dogs were used to keep watch over a fortress and despite being unsuccessful in opposing an enemy attack, they were able to alert the sleeping soldiers to the attack. The history books accredit the Romans with being the first to use dogs in the military; but this might be because the books were written by Romans. However I would like to dispute that, and with the same bit of evidence that is used to support the Roman claims. It is said that the Rome started using dogs in their military after a hard fought battle at Versella. The Battle of Versella was fought in 101 BC, the Roman Empire had been expanding for many years and there latest campaign has them fighting in what is now parts of Germany and France. This large chunk of land was ruled in smaller pieces by Tribal War Lords, but collectively to the Romans it was known as Gaul. It was at The Battle of Verseela that Roman forces clashed with the tribal warriors of the Teutons (not to be confused with the Teutonic Order of Knights), a brave warrior society in which men and women fought alongside each other. It was a bloody and hard fought battle, and one which the Romans did not find easy to win. The Battle of Versella took place in 101BC and saw the Roman Empire (above) clash with Teuton Warriors (below). A hard won battle by the Romans made difficult by constant attacks by hordes of fierce dogs.

9 It was not the mighty Teuton men that gave the Romans so much more than they were reckoning on, that honour fell to the women of the tribe. The Roman accounts tell of a bloody battle made harder because they had to fight hordes of fierce dogs organised by the "blonde-haired women of Wagen-burg." who from out of nowhere, would appear and release hordes of attack dogs to harass the already engage Roman troops. Their stories portray the Teuton heroes as massive mountains of men, wielding large heavy axe and two handed swords, not to mention the rank and file armed with spears and shields. Many of their heroes would wear no armour and fight in just animal skins. This may have captured the Roman soldiers imagination and allowed for a little artistic licence. They give the same impression about their dogs too, large powerful dogs, thick of coat, and strong in jaw. No two share the same markings, lest they be all black. In fact The Romans documented that it was not uncommon for the Tribal Chiefs of Gaul to be accompanied by bodyguards with dogs. The Romans comment on how every effort seems to have been made to protect the fierce beasts as they covered them in leather coats pieced with large nails, and some Romans reports noted that some of the War Chiefs would armour their dogs rather than the men that accompanied them. An old pen and ink drawing of what a Roman Soldier and his war dog might of looked like. (Not drawn by a Roman Soldier) Personally I think the Romans did what the Romans always do, they saw a good idea and integrated it into their own society. I think the Romans should be accredited with being the first people to accredit in writing the Teutons for use of Dogs in the Military, after all they did steal the idea. A few years after the Battle of Verseela, somewhere between 60 to 80. The current emperor had a huge column erected. Marcus Aurelius commissioned the column to commemorate his recent victories in the Danubian or Marcomannic wars, waged by him from 166b.c. to his death.

10 The story begins with the army crossing the river Danube, probably at Carnuntum. A Victory separates the accounts of two expeditions. The exact chronology of the events is disputed; however, the latest theory states that the expeditions against the Marcomanni and Quadi in the years 172b.c. and 173b.c. are in the lower half and the successes of the Emperor over the Samaritans in the years 174b.c. and 175b.c. in the upper half. What is important to us is that on the column dogs are represented, adorned in mail with spiked collars, fighting beside men. What this shows us is that in a very short time dogs became an important part of Roman warfare, enough for them to be eternally commemorated in an epic column. Like the early stories of the Greeks and Corinthians using dogs in warfare, there are other early accounts, for example, when the King of Sparta was besieging Mantenea he discovered that there was treachery taking place in his own forces. Someone was taking food into the town at night. He had the town surrounded by a cordon of dogs, they prevented any help reaching the besieged town. The Column of Marcus Aurelius in Rome, built somewhere between 176b.c. and 180b.c. to commemorate several great military victories. The column shows men and dogs fighting alongside each other. I think it s safe to say that since we realised that we could influence dogs we have been doing so. I don't think any one nation can lay claim to being the fathers of dog training. I hope this has been helpful for anyone reading it, I certainly learnt a lot whilst researching this topic, and enjoyed the process. Like I have said before, unless some invents a time machine we will never know the truth about anything for sure, all we can do is theorise and debate over the information we have at hand and how we perceive it. References Vila, C et al. (1997) Multiple and Ancient Origins of the Domestic Dog In Science Vol.276, No.13 Images courtesy of: Wikipedia [WWW] Accessed: 09/11

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