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This document provides an overview of dogs. It discusses how dogs were domesticated from wolves, with some evidence suggesting domestication occurred as early as 33,000 years ago. Dogs spread widely as human companions and workers, being important for hunting and sled pulling. Genetic studies show dogs diverged from wolves 15,000 years ago, though domestication may have begun earlier. The taxonomy of dogs is discussed, noting they are classified as Canis lupus familiaris as a subspecies of gray wolves.
This document provides an overview of dogs. It discusses how dogs were domesticated from wolves, with some evidence suggesting domestication occurred as early as 33,000 years ago. Dogs spread widely as human companions and workers, being important for hunting and sled pulling. Genetic studies show dogs diverged from wolves 15,000 years ago, though domestication may have begun earlier. The taxonomy of dogs is discussed, noting they are classified as Canis lupus familiaris as a subspecies of gray wolves.
This document provides an overview of dogs. It discusses how dogs were domesticated from wolves, with some evidence suggesting domestication occurred as early as 33,000 years ago. Dogs spread widely as human companions and workers, being important for hunting and sled pulling. Genetic studies show dogs diverged from wolves 15,000 years ago, though domestication may have begun earlier. The taxonomy of dogs is discussed, noting they are classified as Canis lupus familiaris as a subspecies of gray wolves.
This article is about the domestic dog. For related
species known as dogs, see Canidae. For other uses, see Dog (disambiguation). The domestic dog (Canis lupus familiaris) is a member of the Canidae family of the mammalian order Carnivora. The term domestic dog is generally used for both of the domesticated and feral varieties. The word dog can also refer to the male of a canine species as opposed to the word bitch, which refers to the female of the species. [2] The dog was the rst domesticated animal [3][4] and has been widely kept as a working, hunting, and pet compan- ion. In 2001, there were estimated to be 400 million dogs in the world. [5] 1 Etymology and related terminol- ogy Dog is the common use term that refers to members of the subspecies Canis lupus familiaris (canis, dog"; lupus, wolf"; familiaris, of a household or domestic). The term can also be used to refer to a wider range of related species, such as the members of the genus Canis, or true dogs, including the wolf, coyote, and jackals, or it can refer to the members of the tribe Canini, which would also include the African wild dog, or it can be used to refer to any member of the family Canidae, which would also include the foxes, bush dog, raccoon dog, and others. [6] Some members of the family have dog in their common names, such as the raccoon dog and the African wild dog. A few animals have dog in their common names but are not canids, such as the prairie dog. The English word dog comes from Middle English dogge, from Old English docga, a powerful dog breed. [7] The term may possibly derive from Proto- Germanic *dukkn, represented in Old English nger- docce (nger-muscle). [8] The word also shows the fa- miliar petname diminutive -ga also seen in frogga frog, picga pig, stagga stag, wicga beetle, worm, among others. [9] The term dog may ultimately derive from the earliest layer of Proto-Indo-European vocabulary, reect- ing the role of the dog as the earliest domesticated ani- mal. [10] In 14th-century England, hound (from Old English: hund) was the general word for all domestic canines, and dog referred to a subtype of hound, a group includ- ing the masti. It is believed this dog type was so common, it eventually became the prototype of the cat- egory hound. [11] By the 16th century, dog had become the general word, and hound had begun to refer only to types used for hunting. [12] Hound, cognate to German Hund, Dutch hond, common Scandinavian hund, and Icelandic hundur, is ultimately derived from the Proto- Indo-European *kwon- dog, found in Sanskrit kukuur ( ), [13] Welsh ci (plural cwn), Latin canis, Greek kn, and Lithuanian u. [14] In breeding circles, a male canine is referred to as a dog, while a female is called a bitch (Middle English bic- che, from Old English bicce, ultimately from Old Norse bikkja). A group of ospring is a litter. The father of a litter is called the sire, and the mother is called the dam. Ospring are, in general, called pups or puppies, from French poupe, until they are about a year old. The process of birth is whelping, from the Old English word hwelp (cf. German Welpe, Dutch welp, Swedish valpa, Icelandic hvelpur). [15] The term whelp can also be used to refer to the young of any canid, or as a (somewhat ar- chaic) alternative to puppy. 2 Taxonomy See also: Subspecies of Canis lupus In 1753, the father of modern biological taxonomy, Carl Linnaeus, listed among the types of quadrupeds famil- iar to him, the Latin word for dog, canis. Among the species within this genus, Linnaeus listed the fox, as Ca- nis vulpes, wolves (Canis lupus), and the domestic dog, (Canis canis). In later editions, Linnaeus dropped Canis canis and greatly expanded his list of the Canis genus of quadrupeds, and by 1758 included alongside the foxes, wolves, and jackals and many more terms that are now listed as synonyms for domestic dog, including aegyptius (hairless dog), aquaticus, (water dog), and mustelinus (lit- erally "badger dog"). Among these were two that later experts have been widely used for domestic dogs as a species: Canis domesticus and, most predominantly, Ca- nis familiaris, the common or familiar dog. [16] By 1995 with advancements in molecular biology, the DNA analysis of extant (i.e. living today) Canidea species indicated that the wolf and the domestic dog were so genetically similar that the wolf may have been the an- cestor of the dog. [17] By 1999, further genetic analysis indicated that the domestic dog may have emerged from 1 2 3 HISTORY AND EVOLUTION multiple wolf populations. [18][19] Based on these indica- tions, the domestic dog was reclassied in 2005 as Ca- nis lupus familiaris, a subspecies of the Gray Wolf Ca- nis lupus in a professional reference. [20] Based on these pieces of research and the reference reclassication, ca- nis lupis familiaris is the name for the taxon listed by ITIS. [21] However, canis familiaris is also accepted due to a nomenclature debate regarding wild and domestic sub- species. [22] 3 History and evolution Domestic dogs are descended from wolves. Main articles: Origin of the domestic dog and Gray wolf Domestic dogs inherited complex behaviors from their wolf ancestors, which would have been pack hunters with complex body language. These sophisticated forms of social cognition and communication may account for their trainability, playfulness, and ability to t into human households and social situations, and these attributes have given dogs a relationship with humans that has enabled them to become one of the most successful species on the planet today. [23] Recent studies of well-preserved remains of a dog-like canid from the Razboinichya Cave in the Altai Moun- tains of southern Siberia concluded that a particular in- stance of early wolf domestication approximately 33,000 years ago did not result in modern dog lineages, possi- bly because of climate disruption during the Last Glacial Maximum. [3][24] The authors postulate that at least sev- eral such incipient events have occurred. A study of fossil dogs and wolves in Belgium, Ukraine, and Rus- sia tentatively dates domestication from 14,000 years ago to more than 31,700 years ago. [25] Another recent study has found support for claims of dog domestication be- tween 14,000 and 16,000 years ago, with a range between 9,000 and 34,000 years ago, depending on mutation rate assumptions. [26] Although experts largely disagree over the details of dog domestication, it is agreed that human interaction played a signicant role in shaping the subspecies. [27] Domesti- cation may have occurred initially in separate areas, par- ticularly Siberia and Europe. It is thought that the current lineage of dogs were domesticated between 15,000 years and 8,500 years ago. Shortly after the latest domestica- tion, dogs became ubiquitous in human populations, and spread throughout the world. Emigrants from Siberia likely crossed the Bering Strait with dogs in their company, and some experts [28] sug- gest the use of sled dogs may have been critical to the success of the waves that entered North America roughly 12,000 years ago, [28] although the earliest archaeological evidence of dog-like canids in North America dates from about 9,400 years ago. [29][30] Dogs were an important part of life for the Athabascan population in North America, and were their only domesticated animal. Dogs also car- ried much of the load in the migration of the Apache and Navajo tribes 1,400 years ago. Use of dogs as pack ani- mals in these cultures often persisted after the introduc- tion of the horse to North America. [31] The current consensus among biologists and archae- ologists is that the dating of rst domestication is indeterminate, [27][31] although more recent evidence shows isolated domestication events as early as 33,000 years ago. [32][33] There is conclusive evidence the present lineage of dogs genetically diverged from their wolf ancestors at least 15,000 years ago, [34][35][36][37][38] but some believe domestication to have occurred earlier. [27] Evidence is accruing that there were previous domestica- tion events, but that those lineages died out. [39] It is not known whether humans domesticated the wolf as such to initiate dogs divergence from its ancestors, or whether dogs evolutionary path had already taken a dierent course prior to domestication. For example, it is hypothesized that some wolves gathered around the campsites of paleolithic camps to scavenge refuse, and associated evolutionary pressure developed that favored those who were less frightened by, and keener in ap- proaching, humans. [40] The bulk of the scientic evidence for the evolution of the domestic dog stems from morphological studies of archaeological ndings and mitochondrial DNA studies. The divergence date of roughly 15,000 years ago is based in part on archaeological evidence that demonstrates the domestication of dogs occurred more than 15,000 years ago, [23][31] and some genetic evidence indicates the do- mestication of dogs fromtheir wolf ancestors began in the late Upper Paleolithic close to the Pleistocene/Holocene boundary, between 17,000 and 14,000 years ago. [41] But there is a wide range of other, contradictory ndings that make this issue controversial. There are ndings begin- ning currently at 33,000 years ago distinctly placing them as domesticated dogs evidenced not only by shortening of the muzzle but widening as well as crowding of teeth. Archaeological evidence suggests that the latest point at which dogs could have diverged from wolves was roughly 3.1 DNA studies 3 Tesem, an old Egyptian sighthound-like dog. 15,000 years ago, although it is possible they diverged much earlier. [23] In 2008, a team of international scien- tists released ndings froman excavation at Goyet Cave in Belgium declaring a large, toothy canine existed 31,700 years ago and ate a diet of horse, musk ox and reindeer. [42] Prior to this Belgian discovery, the earliest dog bones found were two large skulls from Russia and a mandible fromGermany dated fromroughly 14,000 years ago. [23][36] Remains of smaller dogs from Natuan cave deposits in the Middle East, including the earliest burial of a human being with a domestic dog, have been dated to around 10,000 to 12,000 years ago. [36][43] There is a great deal of archaeological evidence for dogs through- out Europe and Asia around this period and through the next two thousand years (roughly 8,000 to 10,000 years ago), with specimens uncovered in Germany, the French Alps, and Iraq, and cave paintings in Turkey. [23] The old- est remains of a domesticated dog in the Americas were found in Texas and have been dated to about 9,400 years ago. [44] 3.1 DNA studies A basenji, one of the earliest domesticated breeds. DNA studies have provided a wide range of possible di- vergence dates, from15,000 to 40,000 years ago, [36] to as much as 100,000 to 140,000 years ago. [45] These results depend on a number of assumptions. [23] Genetic studies are based on comparisons of genetic diversity between species, and depend on a calibration date. Some esti- mates of divergence dates from DNA evidence use an es- timated wolfcoyote divergence date of roughly 700,000 years ago as a calibration. [46] If this estimate is incorrect, and the actual wolfcoyote divergence is closer to one or two million years ago, or more, [47] than the DNA ev- idence that supports specic dogwolf divergence dates would be interpreted very dierently. Furthermore, it is believed 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 signicantly 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 hybridization does not occur, and that breeds remain geographically localized. Although these assump- tions hold for many species, there is good reason to be- lieve that they do not hold for canines. [23] Genetic analyses indicate all dogs are likely descended from a handful of domestication events with a small num- ber of founding females, [23][41] although there is evidence domesticated dogs interbred with local populations of wild wolves on several occasions. [36] Data suggest dogs rst diverged from wolves in East Asia, and these do- mesticated dogs then quickly migrated throughout the world, reaching the North American continent around 8000 BC. [36] 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. [48] Some breeds thought to be very old, such as the Pharaoh Hound, Ibizan Hound, and Norwegian Elkhound, are now known to have been created more recently. [48] A great deal of controversy surrounds the evolutionary framework for the domestication of dogs. [23] Although it is widely claimed that man domesticated the wolf, [49] man might not have taken such a proactive role in the process. [23] The nature of the interaction between man and wolf that led to domestication is unknown and con- troversial. 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 time in contact with ca- nine species. [23] 4 4 ROLES WITH HUMANS Ancient Greek rhyton in the shape of a dogs head, made by Brygos, early 5th century BC. Jrme Carcopino Museum, Department of Archaeology, Aleria Despite this, there is no evidence of any adaptation of ca- nine species to the presence of the close relatives of mod- ern man. If dogs were domesticated, as believed, roughly 15,000 years ago, the event (or events) would have coin- cided with a large expansion in human territory and the development of agriculture. This has led some biologists to suggest one of the forces that led to the domestication of dogs was a shift in human lifestyle in the formof estab- lished human settlements. Permanent settlements would have coincided with a greater amount of disposable food and would have created a barrier between wild and an- thropogenic canine populations. [23] In 2013 Thalmann, Krause and coworkers revised the view that dog ancestors came from East Asia and showed using DNA analysis that all dogs living today go back to four genetic lineages, all of which originate in Eu- rope. [50] Their data indicated that bonding between hu- mans and dog occurred between 19,000 and 30,000 years ago, likely in the context of hunting. [50] The Zarzian cul- ture, an archaeological culture of late Paleolithic and Mesolithic (18,000-8,000 years BC) in Iraq, Iran, Central Asia is associated with remains of the domesticated dog. [51] 4 Roles with humans A Siberian Husky used as a pack animal A German Shepherd with a football The dogs value to early human hunter-gatherers led to them quickly becoming ubiquitous across world cul- tures. Dogs perform many roles for people, such as hunting, herding, pulling loads, protection, assisting po- lice and military, companionship, and, more recently, aiding handicapped individuals. This impact on human society has given them the nickname "mans best friend" in the Western world. In some cultures, however, dogs are also a source of meat. [52][53] 4.1 Early roles Wolves, and their dog descendants, would have derived signicant benets from living in human campsmore safety, more reliable food, lesser caloric needs, and more chance to breed. [54] They would have beneted from hu- mans upright gait that gives themlarger range over which to see potential predators and prey, as well as color vi- sion that, at least by day, gives humans better visual 4.2 As pets 5 discrimination. [54] Camp dogs would also have benet- ted from human tool use, as in bringing down larger prey and controlling re for a range of purposes. [54] Humans would also have derived enormous benet from the dogs associated with their camps. [55] For instance, dogs would have improved sanitation by cleaning up food scraps. [55] Dogs may have provided warmth, as referred to in the Australian Aboriginal expression three dog night (an exceptionally cold night), and they would have alerted the camp to the presence of predators or strangers, using their acute hearing to provide an early warning. [55] Anthropologists believe the most signicant benet would have been the use of dogs sensitive sense of smell to assist with the hunt. [55] The relationship between the presence of a dog and success in the hunt is often men- tioned as a primary reason for the domestication of the wolf, and a 2004 study of hunter groups with and with- out a dog gives quantitative support to the hypothesis that the benets of cooperative hunting was an important fac- tor in wolf domestication. [56] The cohabitation of dogs and humans would have greatly improved the chances of survival for early human groups, and the domestication of dogs may have been one of the key forces that led to human success. [57] 4.2 As pets A British Bulldog shares a day at the park. The most widespread form of interspecies bonding oc- curs between humans and dogs [55] and the keeping of dogs as companions, particularly by elites, has a long history. [58] (As a possible example, at the Natuan cul- ture site of Ain Mallaha in Israel, dated to 12,000 BC, the remains of an elderly human and a four-to-ve-month- old puppy were found buried together). [59] However, pet dog populations grew signicantly after World War II as suburbanization increased. [58] In the 1950s and 1960s, dogs were kept outside more often than they tend to be A young male border terrier with a raccoon toy. today [60] (using the expression in the doghouse to de- scribe exclusion from the group signies the distance be- tween the doghouse and the home) and were still primar- ily functional, acting as a guard, childrens playmate, or walking companion. From the 1980s, there have been changes in the role of the pet dog, such as the increased role of dogs in the emotional support of their human guardians. [61] People and dogs have become increasingly integrated and implicated in each others lives, [62] to the point where pet dogs actively shape the way a family and home are experienced. [63] There have been two major trends in the changing sta- tus of pet dogs. The rst has been the 'commodica- tion' of the dog, shaping it to conform to human expec- tations of personality and behaviour. [63] The second has been the broadening of the concept of the family and the home to include dogs-as-dogs within everyday routines and practices. [63] There are a vast range of commodity forms available to transform a pet dog into an ideal companion. [64] The list of goods, services and places available is enormous: from dog perfumes, couture, furniture and housing, to dog groomers, therapists, trainers and caretakers, dog cafes, spas, parks and beaches, and dog hotels, airlines and cemeteries. [64] While dog training as an organized activity can be traced back to the 18th century, in the last decades of the 20th century it became a high pro- le issue as many normal dog behaviors such as barking, jumping up, digging, rolling in dung, ghting, and urine marking became increasingly incompatible with the new role of a pet dog. [65] Dog training books, classes and tele- vision programs proliferated as the process of commodi- fying the pet dog continued. [66] The majority of contemporary people with dogs describe their pet as part of the family, [63] although some am- bivalence about the relationship is evident in the pop- ular reconceptualization of the doghuman family as a pack. [63] A dominance model of doghuman relation- ships has been promoted by some dog trainers, such as on the television program Dog Whisperer. However it 6 4 ROLES WITH HUMANS A pet dog taking part in Christmas traditions has been disputed that trying to achieve status is char- acteristic of doghuman interactions. [67] Pet dogs play an active role in family life; for example, a study of conver- sations in doghuman families showed how family mem- bers use the dog as a resource, talking to the dog, or talk- ing through the dog, to mediate their interactions with each other. [68] Another study of dogs roles in families showed many dogs have set tasks or routines undertaken as family mem- bers, the most common of which was helping with the washing-up by licking the plates in the dishwasher, and bringing in the newspaper fromthe lawn. [63] Increasingly, human family members are engaging in activities cen- tered on the perceived needs and interests of the dog, or in which the dog is an integral partner, such as Dog Dancing and Doga. [64] According to statistics published by the American Pet Products Manufacturers Association in the National Pet Owner Survey in 20092010, it is estimated there are 77.5 million people with pet dogs in the United States. [69] The same survey shows nearly 40% of American house- holds own at least one dog, of which 67% own just one dog, 25% two dogs and nearly 9% more than two dogs. There does not seem to be any gender preference among dogs as pets, as the statistical data reveal an equal number of female and male dog pets. Yet, although several pro- grams are undergoing to promote pet adoption, less than a fth of the owned dogs come from a shelter. The latest study using Magnetic resonance imaging (MRI) to humans and dogs together proved that dogs have same response to voices and use the same parts of the brain as humans to do so. This gives dogs the abil- ity to recognize emotional human sounds, making them friendly social pets to humans. [70] 4.3 Work Dogs have lived and worked with humans in so many roles that they have earned the unique nickname, mans best friend, [71] a phrase used in other languages as well. They have been bred for herding livestock, [72] hunting (e.g. pointers and hounds), [73] rodent control, [74] guard- ing, helping shermen with nets, detection dogs, and pulling loads, in addition to their roles as companions. [74] In 1957, a husky-terrier mix named Laika became the rst animal to orbit the Earth. [75][76] Book of the Hunt, Gaston III, Count of Foix, 138788. Service dogs such as guide dogs, utility dogs, assistance dogs, hearing dogs, and psychological therapy dogs pro- vide assistance to individuals with physical or mental disabilities. [77][78] Some dogs owned by epileptics have been shown to alert their handler when the handler shows signs of an impending seizure, sometimes well in advance of onset, allowing the guardian to seek safety, medica- tion, or medical care. [79] Dogs included in human activities in terms of help- ing out humans are usually called working dogs. Dogs of several breeds are considered working dogs. Some working dog breeds include Akita, Alaskan Malamute, Anatolian Shepherd Dog, Bernese Mountain Dog, Black Russian Terrier, Boxer, Bullmasti, Doberman Pinscher, Dogue de Bordeaux, German Pinscher, German Shep- herd, [80] Giant Schnauzer, Great Dane, Great Pyrenees, Great Swiss Mountain Dog, Komondor, Kuvasz, Masti, Neapolitan Masti, Newfoundland, Portuguese Water Dog, Rottweiler, Saint Bernard, Samoyed, Siberian Husky, Standard Schnauzer, and Tibetan Masti. 4.4 Sports and shows See also: Conformation show People often enter their dogs in competitions [81] such 4.6 Health risks to humans 7 as breed-conformation shows or sports, including racing, sledding and agility competitions. In conformation shows, also referred to as breed shows, a judge familiar with the specic dog breed evaluates in- dividual purebred dogs for conformity with their estab- lished breed type as described in the breed standard. As the breed standard only deals with the externally observ- able qualities of the dog (such as appearance, movement, and temperament), separately tested qualities (such as ability or health) are not part of the judging in confor- mation shows. 4.5 As a food source Main article: Dog meat Dog meat is consumed in some East Asian countries, in- Gaegogi (dog meat) stew being served in a Korean restaurant cluding Korea, China, and Vietnam, a practice that dates back to antiquity. [82] It is estimated that 1316 million dogs are killed and consumed in Asia every year. [83] The BBC claims that, in 1999, more than 6,000 restaurants served soups made from dog meat in South Korea. [84] In Korea, the primary dog breed raised for meat, the nureongi (), diers from those breeds raised for pets that Koreans may keep in their homes. [85] The most popular Korean dog dish is gaejang-guk (also called bosintang), a spicy stewmeant to balance the bodys heat during the summer months; followers of the custom claimthis is done to ensure good health by balancing ones gi, or vital energy of the body. A 19th century version of gaejang-guk explains that the dish is prepared by boil- ing dog meat with scallions and chili powder. Variations of the dish contain chicken and bamboo shoots. While the dishes are still popular in Korea with a segment of the population, dog is not as widely consumed as beef, chicken, and pork. [86] A CNN report in China dated March 2010 includes an interview with a dog meat vendor who stated that most of the dogs that are available for selling to restaurants are raised in special farms but that there is always a chance that a sold dog is someones lost pet, although dog pet breeds are not considered edible. [87] Other cultures, such as Polynesia and pre-Columbian Mexico, also consumed dog meat in their history. How- ever, Western, South Asian, African, and Middle Eastern cultures, in general, regard consumption of dog meat as taboo. In some places, however, such as in rural areas of Poland, dog fat is believed to have medicinal properties being good for the lungs for instance. [88] Dog meat is also consumed in some parts of Switzerland. [89] 4.6 Health risks to humans Main article: Dog attack It is estimated that 4.7 million people are bitten by dogs Pet waste station at a government building. each year. [90] In the 1980s and 1990s the US averaged 17 fatalities per year, while in the 2000s this has increased to 26. [91] 77% of dog bites are from the pet of family or friends, and 50% of attacks occur on the property of the dogs legal owner. [91] A Colorado study found bites in children were less severe than bites in adults. [92] The incidence of dog bites in the US is 12.9 per 10,000 inhabitants, but for boys aged 5 to 9, the incidence rate is 60.7 per 10,000. Moreover, children have a much higher chance to be bitten in the face or neck. [93] Sharp claws with powerful muscles be- hind them can lacerate esh in a scratch that can lead to serious infections. [94] In the UK between 2003 and 2004, there were 5,868 dog 8 4 ROLES WITH HUMANS attacks on humans, resulting in 5,770 working days lost in sick leave. [95] In the United States, cats and dogs are a factor in more than 86,000 falls each year. [96] It has been estimated around 2% of dog-related injuries treated in UK hospitals are domestic accidents. The same study found that while dog involvement in road trac accidents was dicult to quantify, dog-associated road accidents involving injury more commonly involved two-wheeled vehicles. [97] Toxocara canis (dog roundworm) eggs in dog feces can cause toxocariasis. In the United States, about 10,000 cases of Toxocara infection are reported in hu- mans each year, and almost 14% of the U.S. pop- ulation is infected. [98] In Great Britain, 24% of soil samples taken from public parks contained T. canis eggs. [99] Untreated toxocariasis can cause retinal dam- age and decreased vision. [99] Dog feces can also con- tain hookworms that cause cutaneous larva migrans in humans. [100][101][102][103] 4.7 Health benets for humans A human cuddles a Doberman puppy. The scientic evidence is mixed as to whether compan- ionship of a dog can enhance human physical health and psychological wellbeing. [104] Studies suggesting that there are benets to physical health and psychologi- cal wellbeing [105] have been criticised for being poorly controlled, [106] and nding that "[t]he health of elderly people is related to their health habits and social supports but not to their ownership of, or attachment to, a com- panion animal. Earlier studies have shown that people who keep pet dogs or cats exhibit better mental and phys- ical health than those who do not, making fewer visits to the doctor and being less likely to be on medication than non-guardians. [107] A 2005 paper states recent research has failed to sup- port earlier ndings that pet ownership is associated with a reduced risk of cardiovascular disease, a reduced use of general practitioner services, or any psychological or physical benets on health for community dwelling older people. Research has, however, pointed to sig- nicantly less absenteeism from school through sickness among children who live with pets. [104] In one study, new guardians reported a highly signicant reduction in minor health problems during the rst month following pet ac- quisition, and this eect was sustained in those with dogs through to the end of the study. [108] In addition, people with pet dogs took considerably more physical exercise than those with cats and those with- out pets. The group without pets exhibited no statisti- cally signicant changes in health or behaviour. The re- sults provide evidence that keeping pets may have pos- itive eects on human health and behaviour, and that for guardians of dogs these eects are relatively long- term. [108] Pet guardianship has also been associated with increased coronary artery disease survival, with human guardians being signicantly less likely to die within one year of an acute myocardial infarction than those who did not own dogs. [109] Gunnar Kaasen and Balto, the lead dog on the last relay team of the 1925 serum run to Nome. The health benets of dogs can result from contact with dogs in general, and not solely from having dogs as pets. For example, when in the presence of a pet dog, people show reductions in cardiovascular, behavioral, and psy- chological indicators of anxiety. [110] Other health bene- ts are gained from exposure to immune-stimulating mi- croorganisms, which, according to the hygiene hypoth- esis, can protect against allergies and autoimmune dis- eases. The benets of contact with a dog also include social support, as dogs are able to not only provide com- panionship and social support themselves, but also to act as facilitators of social interactions between humans. [111] 5.1 Senses 9 One study indicated that wheelchair users experience more positive social interactions with strangers when they are accompanied by a dog than when they are not. [112] The practice of using dogs and other animals as a part of therapy dates back to the late 18th century, when animals were introduced into mental institutions to help social- ize patients with mental disorders. [113] Animal-assisted intervention research has shown that animal-assisted ther- apy with a dog can increase social behaviors, such as smiling and laughing, among people with Alzheimers disease. [114] One study demonstrated that children with ADHD and conduct disorders who participated in an ed- ucation program with dogs and other animals showed in- creased attendance, increased knowledge and skill ob- jectives, and decreased antisocial and violent behavior compared to those who were not in an animal-assisted program. [115] 4.8 Shelters Main article: Animal shelter Every year, between 6 and 8 million dogs and cats en- ter US animal shelters. [116] The Humane Society of the United States (HSUS) estimates that approximately 3 to 4 million of those dogs and cats are euthanized yearly in the United States. [117] However, the percentage of dogs in US animal shelters that are eventually adopted and removed from the shelters by their new legal owners has increased since the mid-1990s from around 25% to a 2012 average of 40% among reporting shelters [118] (and many shelters reporting 6075%). [119] 5 Biology Main article: Dog anatomy Domestic dogs have been selectively bred for millen- Lateral view of a dogs bone structure. nia for various behaviors, sensory capabilities, and phys- ical attributes. [74] Modern dog breeds show more varia- tion in size, appearance, and behavior than any other do- mestic animal. Nevertheless, their morphology is based on that of their wild ancestors, gray wolves. [74] Dogs are predators and scavengers, and like many other preda- tory mammals, the dog has powerful muscles, fused wrist bones, a cardiovascular system that supports both sprint- ing and endurance, and teeth for catching and tearing. Dogs are highly variable in height and weight. The small- est known adult dog was a Yorkshire Terrier, that stood only 6.3 cm (2.5 in) at the shoulder, 9.5 cm (3.7 in) in length along the head-and-body, and weighed only 113 grams (4.0 oz). The largest known dog was an English Masti which weighed 155.6 kg (343 lb) and was 250 cm (98 in) from the snout to the tail. [120] The tallest dog is a Great Dane that stands 106.7 cm (42.0 in) at the shoulder. [121] 5.1 Senses 5.1.1 Vision Dogs visual colour perception compared with humans. Like most mammals, dogs are dichromats and have color vision equivalent to redgreen color blindness in humans (deuteranopia). [122][123][124][125] So, dogs can see blue and yellow, but have diculty dierentiating red and green because they only have two spectral types of cone pho- toreceptor, while normal humans have three. And dogs use color instead of brightness to dierentiate light or dark blue/yellow. [126][127][128][129] Dogs are less sensitive to dierences in grey shades than humans and also can de- tect brightness at about half the accuracy of humans. [130] The dogs visual system has evolved to aid procient hunting. [122] While a dogs visual acuity is poor (that of a poodle's has been estimated to translate to a Snellen rat- ing of 20/75 [122] ), their visual discrimination for moving objects is very high; dogs have been shown to be able to discriminate between humans (e.g., identifying their human guardian) at a range of between 800 and 900 m, however this range decreases to 500600 m if the object is stationary. [122] Dogs have a temporal resolution of between 60 and 70 Hz, which explains why many dogs struggle to watch tele- vision, as most such modern screens are optimized for 10 5 BIOLOGY humans at 5060 Hz. [130] Dogs can detect a change in movement that exists in a single diopter of space within their eye. Humans, by comparison, require a change of between 10 and 20 diopters to detect movement. [131] As crepuscular hunters, dogs often rely on their vision in low light situations: They have very large pupils, a high density of rods in the fovea, an increased icker rate, and a tapetum lucidum. [122] The tapetum is a reective sur- face behind the retina that reects light to give the pho- toreceptors a second chance to catch the photons. There is also a relationship between body size and overall diam- eter of the eye. A range of 9.5 and 11.6 mm can be found between various breeds of dogs. This 20% variance can be substantial and is associated as an adaptation toward superior night vision. [132] The eyes of dierent breeds of dogs have dierent shapes, dimensions, and retina congurations. [133] Many long-nosed breeds have a visual streaka wide foveal region that runs across the width of the retina and gives them a very wide eld of excellent vision. Some long- muzzled breeds, in particular, the sighthounds, have a eld of vision up to 270 (compared to 180 for humans). Short-nosed breeds, on the other hand, have an area cen- tralis": a central patch with up to three times the density of nerve endings as the visual streak, giving themdetailed sight much more like a humans. Some broad-headed breeds with short noses have a eld of vision similar to that of humans. [123][124] Most breeds have good vision, but some show a genetic predisposition for myopia such as Rottweilers, with which one out of every two has been found to be myopic. [122] Dogs also have a greater divergence of the eye axis than humans, enabling themto rotate their pupils farther in any direction. The divergence of the eye axis of dogs ranges from 1225 depending on the breed. [131] Experimentation has proven that dogs can distinguish be- tween complex visual images such as that of a cube or a prism. Dogs also show attraction to static visual im- ages such as the silhouette of a dog on a screen, their own reections, or videos of dogs; however, their interest de- clines sharply once they are unable to make social contact with the image. [134] 5.1.2 Hearing The frequency range of dog hearing is approximately 40 Hz to 60,000 Hz, [135] which means that dogs can detect sounds far beyond the upper limit of the human auditory spectrum. [124][135][136] In addition, dogs have ear mobil- ity, which allows them to rapidly pinpoint the exact lo- cation of a sound. [137] Eighteen or more muscles can tilt, rotate, raise, or lower a dogs ear. A dog can identify a sounds location much faster than a human can, as well as hear sounds at four times the distance. [137] The physiology of a dog ear. Transformation of the ears of a huskamute puppy in 6 days 5.1.3 Smell The wet, textured nose of a dog While the human brain is dominated by a large visual cortex, the dog brain is dominated by an olfactory cor- tex. [122] The olfactory bulb in dogs is roughly forty times bigger than the olfactory bulb in humans, relative to to- tal brain size, with 125 to 220 million smell-sensitive receptors. [122] The bloodhound exceeds this standard with nearly 300 million receptors. [122] Consequently, it has been estimated that dogs, in gen- eral, have an olfactory sense ranging from one hundred thousand to one million times more sensitive than a hu- mans. In some dog breeds, such as bloodhounds, the ol- factory sense may be up to 100 million times greater than a humans. [138] The wet nose, or rhinarium, is essential for determining the direction of the air current contain- 5.3 Types and breeds 11 ing the smell. Cold receptors in the skin are sensitive to the cooling of the skin by evaporation of the moisture by air currents. [139] 5.2 Physical characteristics Main article: Dog anatomy 5.2.1 Coat Main article: Coat (dog) The coats of domestic dogs are of two varieties: double A heavy winter coat with countershading in a mixed-breed dog being common with dogs (as well as wolves) originating from colder climates, made up of a coarse guard hair and a soft down hair, or single, with the topcoat only. Domestic dogs often display the remnants of countershading, a common natural camouage pat- tern. A countershaded animal will have dark coloring on its upper surfaces and light coloring below, [140] which reduces its general visibility. Thus, many breeds will have an occasional blaze, stripe, or star of white fur on their chest or underside. [141] 5.2.2 Tail See also: Docking There are many dierent shapes for dog tails: straight, straight up, sickle, curled, or cork-screw. As with many canids, one of the primary functions of a dogs tail is to communicate their emotional state, which can be im- portant in getting along with others. In some hunting dogs, however, the tail is traditionally docked to avoid injuries. [142] In some breeds, such as the Braque du Bour- bonnais, puppies can be born with a short tail or no tail at all. [143] 5.3 Types and breeds Main article: Dog breed Further information: Dog type Most breeds of dog are at most a few hundred Cavalier King Charles Spaniels demonstrate within-breed varia- tion. years old, having been articially selected for particu- lar morphologies and behaviors by people for specic functional roles. Through this selective breeding, the dog has developed into hundreds of varied breeds, and shows more behavioral and morphological variation than any other land mammal. [144] For example, height mea- sured to the withers ranges from 15.2 centimetres (6.0 in) in the Chihuahua to about 76 cm (30 in) in the Irish Wolfhound; color varies from white through grays (usu- ally called blue) to black, and browns from light (tan) to dark (red or chocolate) in a wide variation of pat- terns; coats can be short or long, coarse-haired to wool- like, straight, curly, or smooth. [145] It is common for most breeds to shed this coat. While all dogs are genetically very similar, [36] natural selection and selective breeding have reinforced certain characteristics in certain populations of dogs, giving rise to dog types and dog breeds. Dog types are broad cate- gories based on function, genetics, or characteristics. [146] Dog breeds are groups of animals that possess a set of in- herited characteristics that distinguishes them from other animals within the same species. Modern dog breeds are non-scientic classications of dogs kept by modern kennel clubs. Purebred dogs of one breed are genetically distinguish- able from purebred dogs of other breeds, [48] but the 12 5 BIOLOGY means by which kennel clubs classify dogs is unsystem- atic. Systematic analyses of the dog genome has re- vealed only four major types of dogs that can be said to be statistically distinct. [48] These include the old world dogs (e.g., Malamute and Shar Pei), Masti"-type (e.g., English Masti), herding"-type (e.g., Border Collie), and all others (also called modern"- or hunting"- type). [48][147] 5.4 Health Main articles: Dog health and CVBD Dogs are susceptible to various diseases, ailments, and poisons, some of which can aect humans. To de- fend against many common diseases, dogs are often vaccinated. There are many household plants that are poisonous to dogs, begonia and aloe vera. [148] Poinsettia is often claimed to be toxic but this is untrue. The LD50 for rats was determined to be greater than 25g/kg. [149] This would mean that a 50 pound dog would have to eat 500- 600 leaves to have a 50% chance of death. Some breeds of dogs are prone to certain genetic ail- ments such as elbowor hip dysplasia, blindness, deafness, pulmonic stenosis, cleft palate, and trick knees. Two se- rious medical conditions particularly aecting dogs are pyometra, aecting unspayed females of all types and ages, and bloat, which aects the larger breeds or deep- chested dogs. Both of these are acute conditions, and can kill rapidly. Dogs are also susceptible to parasites such as eas, ticks, and mites, as well as hookworm, tapeworm, roundworm, and heartworm. Dogs are highly susceptible to theobromine poisoning, typically from ingestion of chocolate. Theobromine is toxic to dogs because, although the dogs metabolism is capable of breaking down the chemical, the process is so slow that even small amounts of chocolate can be fatal, especially dark chocolate. Dogs are also vulnerable to some of the same health conditions as humans, including diabetes, dental and heart disease, epilepsy, cancer, hypothyroidism, and arthritis. [150] 5.4.1 Mortality Main article: Aging in dogs The typical lifespan of dogs varies widely among breeds, but for most the median longevity, the age at which half the dogs in a population have died and half are still alive, ranges from 10 to 13 years. [151][152][153][154] Individual dogs may live well beyond the median of their breed. The breed with the shortest lifespan (among breeds for A mixed-breed terrier. Mixed-breed dogs are generally healthier than pure-breds. which there is a questionnaire survey with a reasonable sample size) is the Dogue de Bordeaux, with a me- dian longevity of about 5.2 years, but several breeds, including Miniature Bull Terriers, Bloodhounds, and Irish Wolfhounds are nearly as short-lived, with median longevities of 6 to 7 years. [154] The longest-lived breeds, including Toy Poodles, Japanese Spitz, Border Terriers, and Tibetan Spaniels, have median longevities of 14 to 15 years. [154] The me- dian longevity of mixed-breed dogs, taken as an average of all sizes, is one or more years longer than that of pure- bred dogs when all breeds are averaged. [152][153][154][155] The dog widely reported to be the longest-lived is "Bluey", who died in 1939 and was claimed to be 29.5 years old at the time of his death; however, the Bluey record is anecdotal and unveried. [156] On 5 December 2011, Pusuke, the worlds oldest living dog recognized by Guinness Book of World Records, died aged 26 years and 9 months. [157] 5.4.2 Predation Although wild dogs, like wolves, are apex predators, they can be killed in territory disputes with wild animals. [158] Furthermore, in areas where both dogs and other large predators live, dogs can be a major food source for big cats or canines. Reports from Croatia indicate wolves kill dogs more frequently than they kill sheep. Wolves in Russia apparently limit feral dog populations. In Wiscon- sin, more compensation has been paid for dog losses than livestock. [158] Some wolf pairs have been reported to prey on dogs by having one wolf lure the dog out into heavy brush where the second animal waits in ambush. [159] In some instances, wolves have displayed an uncharacter- istic fearlessness of humans and buildings when attack- ing dogs, to the extent that they have to be beaten o or killed. [160] 5.6 Reproduction 13 Coyotes and big cats have also been known to attack dogs. Leopards in particular are known to have a predilection for dogs, and have been recorded to kill and consume them regardless of the dogs size or ferocity. [161] Tigers in Manchuria, Indochina, Indonesia, and Malaysia are re- puted to kill dogs with the same vigor as leopards. [162] Striped Hyenas are major predators of village dogs in Turkmenistan, India, and the Caucasus. [163] Reptiles such as alligators and pythons have been known to kill and eat dogs. 5.5 Diet See also: Dog food Despite their descent from wolves and classica- Golden Retriever gnawing a pigs foot tion as Carnivora, dogs are variously described in scholarly and other writings as carnivores [164][165] or omnivores. [74][166][167][168] Unlike obligate carnivores, such as the cat family with its shorter small intestine, dogs can adapt to a wide-ranging diet, and are not dependent on meat-specic protein nor a very high level of protein in order to fulll their basic dietary requirements. Dogs will healthily digest a variety of foods, including vegeta- bles and grains, and can consume a large proportion of these in their diet. [74] Compared to their wolf ancestors, dogs have adaptations in genes involved in starch diges- tion that contribute to an increased ability to thrive on a starch-rich diet. [169] 5.5.1 Foods toxic to dogs A number of common human foods and household in- gestibles are toxic to dogs, including chocolate solids (theobromine poisoning), onion and garlic (thiosulphate, sulfoxide or disulde poisoning), [170] grapes and raisins, macadamia nuts, xylitol, [171] as well as various plants and other potentially ingested materials. [172][173] The nicotine in tobacco can also be dangerous. Dogs can get it by scav- enging in garbage or ashtrays; eating cigars and cigarettes. Signs can be vomiting of large amounts (e.g., from eating cigar butts) or diarrhea. Some other signs are abdominal pain, loss of coordination, collapse, or death. To solve, soothe the stomach irritation by giving charcoal tablets. For severe signs, get immediate veterinary attention. [174] 5.6 Reproduction Main article: Canine reproduction In domestic dogs, sexual maturity begins to happen around age six to twelve months for both males and females, [74][175] although this can be delayed until up to two years old for some large breeds. This is the time at which female dogs will have their rst estrous cycle. They will experience subsequent estrous cycles biannually, dur- ing which the body prepares for pregnancy. At the peak of the cycle, females will come into estrus, being men- tally and physically receptive to copulation. [74] Because the ova survive and are capable of being fertilized for a week after ovulation, it is possible for a female to mate with more than one male. [74] 25 days post conception fertilization occurs, 1416 days embryo attaches to uterus 2223 days heart beat is detectable. [176][177] Dogs bear their litters roughly 58 to 68 days after fertilization, [74][178] with an average of 63 days, although the length of gestation can vary. An average litter consists of about six puppies, [179] though this number may vary widely based on the breed of dog. In general, toy dogs produce from one to four puppies in each litter, while much larger breeds may average as many as twelve. Some dog breeds have acquired traits through selective breeding that interfere with reproduction. Male French Bulldogs, for instance, are incapable of mounting the fe- male. For many dogs of this breed, the female must be articially inseminated in order to reproduce. [180] 5.6.1 Neutering Neutering refers to the sterilization of animals, usually by removal of the males testicles or the females ovaries and uterus, in order to eliminate the ability to procre- ate and reduce sex drive. Because of the overpopulation of dogs in some countries, many animal control agen- 14 6 INTELLIGENCE AND BEHAVIOR A feral dog from Sri Lanka nursing her four puppies cies, such as the American Society for the Prevention of Cruelty to Animals (ASPCA), advise that dogs not in- tended for further breeding should be neutered, so that they do not have undesired puppies that may have to later be euthanized. [181] According to the Humane Society of the United States, 34 million dogs and cats are put down each year in the United States and many more are conned to cages in shelters because there are many more animals than there are homes. Spaying or castrating dogs helps keep over- population down. [182] Local humane societies, SPCAs, and other animal protection organizations urge people to neuter their pets and to adopt animals from shelters in- stead of purchasing them. Neutering reduces problems caused by hypersexuality, especially in male dogs. [183] Spayed female dogs are less likely to develop some forms of cancer, aecting mam- mary glands, ovaries, and other reproductive organs. [184] However, neutering increases the risk of urinary in- continence in female dogs, [185] and prostate cancer in males, [186] as well as osteosarcoma, hemangiosarcoma, cruciate ligament rupture, obesity, and diabetes mellitus in either sex. [187] 5.7 Communication By the age of four weeks, the dog has developed the ma- jority of its vocalizations. [188] The dog is the most vocal canid and is unique in its tendency to bark in a myriad of situations. [188] Barking appears to have little more communication func- tions than excitement, ghting, the presence of a human, or simply because other dogs are barking. [188] Subtler signs such as discreet bodily and facial movements, body odors, whines, yelps, and growls are the main sources of actual communication. [188] The majority of these subtle communication techniques are employed at a close prox- imity to another, but for long-range communication only barking and howling are employed. [188] 6 Intelligence and behavior 6.1 Intelligence Main article: Dog intelligence The domestic dog has a predisposition to exhibit a social The Border Collie is considered to be one of the most intelligent breeds. [189] intelligence that is uncommon in the animal world. [122] Dogs are capable of learning in a number of ways, such as through simple reinforcement (e.g., classical or operant conditioning) and by observation. [190][122] Dogs go through a series of stages of cognitive develop- ment. As with humans, the understanding that objects not being actively perceived still remain in existence (called object permanence) is not present at birth. It develops as the young dog learns to interact intentionally with objects around it, at roughly 8 weeks of age. [122] Puppies learn behaviors quickly by following examples set by experienced dogs. [122] This form of intelligence is not peculiar to those tasks dogs have been bred to per- form, but can be generalized to myriad abstract problems. For example, Dachshund puppies that watched an expe- rienced dog pull a cart by tugging on an attached piece of ribbon in order to get a reward from inside the cart learned the task fteen times faster than those left to solve the problem on their own. [122][191] Dogs can also learn by mimicking human behaviors. In one study, puppies were presented with a box, and shown that, when a handler pressed a lever, a ball would roll out of the box. The handler then allowed the puppy to play with the ball, making it an intrinsic reward. The pups were then allowed to interact with the box. Roughly three quarters of the puppies subsequently touched the lever, and over half successfully released the ball, compared to only 6% in a control group that did not watch the hu- man manipulate the lever. [192] Another study found that 6.2 Behavior 15 handing an object between experimenters who then used the objects name in a sentence successfully taught an ob- serving dog each objects name, allowing the dog to sub- sequently retrieve the item. [193] Sergeant Stubby wearing his uniform and medals. Stubby partic- ipated in four oensives and 17 battles. Dogs also demonstrate sophisticated social cognition by associating behavioral cues with abstract meanings. [122] One such class of social cognition involves the under- standing that others are conscious agents. Research has shown that dogs are capable of interpreting subtle social cues, and appear to recognize when a human or dogs at- tention is focused on them. To test this, researchers de- vised a task in which a reward was hidden underneath one of two buckets. The experimenter then attempted to communicate with the dog to indicate the location of the reward by using a wide range of signals: tapping the bucket, pointing to the bucket, nodding to the bucket, or simply looking at the bucket. [194] The results showed that domestic dogs were better than chimpanzees, wolves, and human infants at this task, and even young puppies with limited exposure to humans performed well. [122] Psychology research has shown that humans gaze instinc- tively moves to the left in order to watch the right side of a persons face, which is related to use of right hemi- sphere brain for facial recognition, including human facial emotions. Research at the University of Lincoln (2008) shows that dogs share this instinct when meeting a hu- man being, and only when meeting a human being (i.e., not other animals or other dogs). As such they are the only non-primate species known to do so. [195][196] Stanley Coren, an expert on dog psychology, states that these results demonstrated the social cognition of dogs can exceed that of even our closest genetic relatives, and that this capacity is a recent genetic acquisition that dis- tinguishes the dog from its ancestor, the wolf. [122] Studies have also investigated whether dogs engaged in partnered play change their behavior depending on the attention- state of their partner. [197] Those studies showed that play signals were only sent when the dog was holding the at- tention of its partner. If the partner was distracted, the dog instead engaged in attention-getting behavior before sending a play signal. [197] Coren has also argued that dogs demonstrate a sophisti- cated theory of mind by engaging in deception, which he supports with a number of anecdotes, including one ex- ample wherein a dog hid a stolen treat by sitting on it until the rightful owner of the treat left the room. [122] Although this could have been accidental, Coren suggests that the thief understood that the treats owner would be unable to nd the treat if it were out of view. Together, the empiri- cal data and anecdotal evidence points to dogs possessing at least a limited form of theory of mind. [122][197] Similar research has been performed by Brian Hare of Duke Uni- versity, who has shown that dogs outperform both great apes as well as wolves raised by humans in reading human communicative signals. [198][199] A study found a third of dogs suered from anxiety when separated from others. [200] A border collie named Chaser has learned the names for 1,022 toys after three years of training, so many that her trainers have had to mark the names of the objects lest they forget themselves. This is higher than Rico, another border collie who could remember at least 200 objects. [201] 6.2 Behavior Main article: Dog behavior Although dogs have been the subject of a great deal of behaviorist psychology (e.g. Pavlovs dog), they do not enter the world with a psychological blank slate. [122] Rather, dog behavior is aected by genetic factors as well as environmental factors. [122] Domestic dogs exhibit a number of behaviors and predispositions that were in- herited from wolves. [122] Given that dogs abilities to use human social cues origi- nated during the process of domestication, it is likely that individual dogs that were able to use social cues to predict the behavior of humans more exibly than could their last common wolf ancestor were at a selective advantage. [194] Properly socialized dogs can interact with unfamiliar dogs of any size and shape and understand how to communicate. The existence and nature of personality traits in dogs 16 7 DIFFERENCES FROM WOLVES have been studied (15329 dogs of 164 dierent breeds) and ve consistent and stable narrow traits iden- tied, described as playfulness, curiosity/fearlessness, chase-proneness, sociability and aggressiveness. A fur- ther higher order axis for shynessboldness was also identied. [202][203] The average sleep time of a dog is said to be 10.1 hours per day. [204] Like humans, dogs have two main types of sleep: Slow-wave sleep, then Rapid eye movement sleep, the state in which dreams occur. [205] Dogs prefer, when they are o the leash and Earths mag- netic eld is calm, to urinate and defecate with their bod- ies aligned on a north-south axis. [206] 6.3 Dog growl A new study in Budapest, Hungary, has found that dogs are able to tell how big another dog is just by listening to its growl. A specic growl is used by dogs to protect their food. The research also shows that dogs do not lie about their size, and this is the rst time research has shown an- imals can determine anothers size by the sound it makes. The test, using images of many kinds of dogs, showed a small and big dog and played a growl. The result showed that 20 of the 24 test dogs looked at the image of the ap- propriately sized dog rst and looked at it longest. [207] 7 Dierences from wolves Some dogs, like this Tamaskan Dog, look very much like wolves. 7.1 Physical characteristics Further information: Wolf Compared to equally sized wolves, dogs tend to have 20% smaller skulls, 30% smaller brains, [208] as well as pro- portionately smaller teeth than other canid species. [209] Dogs require fewer calories to function than wolves. It is thought by certain experts that the dogs limp ears are a result of atrophy of the jaw muscles. [209] The skin of do- mestic dogs tends to be thicker than that of wolves, with some Inuit tribes favoring the former for use as cloth- ing due to its greater resistance to wear and tear in harsh weather. [209] 7.2 Behavioral dierences Dogs tend to be poorer than wolves at observational learn- ing, being more responsive to instrumental condition- ing. [209] Feral dogs showlittle of the complex social struc- ture or dominance hierarchy present in wolf packs. For example, unlike wolves, the dominant alpha pairs of a feral dog pack do not force the other members to wait for their turn on a meal when scavenging o a dead un- gulate as the whole family is free to join in. For dogs, other members of their kind are of no help in locating food items, and are more like competitors. [209] Feral dogs are primarily scavengers, with studies show- ing that unlike their wild cousins, they are poor ungulate hunters, having little impact on wildlife populations where they are sympatric. However, feral dogs have been reported to be eective hunters of reptiles in the Galpagos Islands, [210] and free ranging pet dogs are more prone to predatory behavior toward wild animals. Domestic dogs can be monogamous. [211] Breeding in feral packs can be, but does not have to be restricted to a domi- nant alpha pair (such things also occur in wolf packs). [212] Male dogs are unusual among canids by the fact that they mostly seem to play no role in raising their puppies, and do not kill the young of other females to increase their own reproductive success. [210] Some sources say that dogs dier from wolves and most other large canid species by the fact that they do not regurgitate food for their young, nor the young of other dogs in the same territory. [209] However, this dierence was not observed in all domestic dogs. Regurgitating of food by the females for the young as well as care for the young by the males has been ob- served in domestic dogs, dingos as well as in other feral or semi-feral dogs. Regurgitating of food by the females and direct choosing of only one mate has been observed even in those semi-feral dogs of direct domestic dog ancestry. Also regurgitating of food by males has been observed in free-ranging domestic dogs. [211][213] 7.3 Trainability Dogs display much greater tractability than tame wolves, and are, in general, much more responsive to coercive techniques involving fear, aversive stimuli, and force than 17 wolves, which are most responsive toward positive condi- tioning and rewards. [214] Unlike tame wolves, dogs tend to respond more to voice than hand signals. [215] 8 Cultural depictions Main article: Cultural depictions of dogs 8.1 Mythology In mythology, dogs often serve as pets or as watchdogs. [216] In Greek mythology, Cerberus is a three-headed watch- dog who guards the gates of Hades. [216] In Norse mythol- ogy, a bloody, four-eyed dog called Garmr guards Helheim. [216] In Persian mythology, two four-eyed dogs guard the Chinvat Bridge. [216] In Philippine mythology, Kimat who is the pet of Tadaklan, god of thunder, is re- sponsible for lightning. In Welsh mythology, Annwn is guarded by Cn Annwn. [216] In Hindu mythology, Yama, the god of death owns two watch dogs who have four eyes. They are said to watch over the gates of Naraka. [217] Hunter god Muthappan from North Malabar region of Kerala has hunting Dog as his mount. Dogs are found in and out of the Muthappan Temple and oerings at the shrine take the form of bronze dog gurines. [218] In Judaism and Islam, dogs are viewed as un- clean scavengers. [216] In Christianity, dogs represent faithfulness. [216] In Asian countries such as China, Korea, and Japan, dogs are viewed as kind protectors. [216] The role of the dog in Chinese mythology includes a position as one of the twelve animals which cyclically represent years (the zodiacal dog). 8.2 Gallery of dogs in art 9 See also Animal track Argos (dog) Dog in Chinese mythology Dogs in art Dog odor Dognapping Ethnocynology Hachika notable example of dog loyalty Lost pet services Wolfdog Lists: List of dog breeds List of ctional dogs List of individual dogs List of most popular dog breeds 10 References [1] Mammal Species of the World Browse: Canis lupus fa- miliaris. Bucknell.edu. 2005. Retrieved 12 March 2012. [2] Dog. Dictionary.com. [3] Nikolai D. Ovodov, Susan J. Crockford, Yaroslav V. Kuzmin, Thomas F. G. Higham, Gregory W. L. Hodgins, Johannes van der Plicht. (2011). A 33,000-Year-Old In- cipient Dog from the Altai Mountains of Siberia: Evidence of the Earliest Domestication Disrupted by the Last Glacial Maximum. 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FuzzyPants, Rupz112, The Nut, Xxxbel- lacutie998, Lastitem, Dgd, Bohan6637, Agreimann, AvicAWB, H3llBot, Jimbo123454321, SporkBot, Erianna, FroBro13, Stealth3861, Brutusbull, Anonwin6, Iamcoolerer, Greatboing3, Potpie22, Augurar, Kenndawg12345677789, Thompsfred32, Prettyponies 2, Leenlol, Alborzagros, Gsarwa, Ltcorpse, Usb10, MikeNicho231, Moshi Monster Fan303, Damirgrati, Ego White Tray, Monteitho, Chuispas- 28 13 TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES tonBot, Gcjdavid, SYTYCSM, Freemankevin15, ClueBot NG, Michaelmas1957, Breguet14, Jacksoncw, Ron keiser, Raghith, LittleJerry, Dav-FL-IN-AZ-id, Hazhk, Tfouche, PukiBunz81, KirtanDevotee, CaroleHenson, CopperSquare, GlassLadyBug, Hamnrye, DrChrissy, Crazymonkey1123, Pluma, Jk2q3jrklse, North Atlanticist Usonian, RakiSykes, Helpful Pixie Bot, Westnest, Legoless, Wichmer1, Bib- code Bot, ZoomanSP, Jackebower, XiaomingWang, Hodeken, MKar, Skullcandle, Trigg hound 2, Prisinor108, ElphiBot, JohnChrysostom, MusikAnimal, AvocatoBot, Samuel Rosenbalm, Earthsbuddy, Karbont, Muneeb2000, Frankonno, Iamthecheese44, Sridhar1000, Tanger- inehistry, Jeancey, Moulnj00, NotWith, Dotbrodu, Rwc2118, Checkb4uwreck, Jawadreventon, Tigris35711, Ashleyrms22, Peacock28, Carsenegame, BattyBot, AaMaCo13, HappyFridays, PanShiBo, Electricmun11, Khazar2, Rhlozier, Onepebble, Dexbot, Aborichan, JPollock412, Vidmas258, Webclient101, Urville86, Alphama, ChalkboardCowboy, Bhaskar Bhatt Joshi, KayakDave, Elekebia, Jochen Burghardt, Coonaldo, Reatlas, Donfbreed2, , Global Microscope, Andrew.mantz, Priyanshu7, SamX, Rejancar, Pepesia, Cscott79, Cen- soredScribe, WorldTraveller101, Shiningroad, Birhanb, Badger2424, Nicoderno1, Wpzhiyilee, Mevagiss, Berglopen, Monkbot, William of Aragon, Horselover Fats, Dodam.a.ih, Gunduu, TerryAlex, Dodynamic, Do The Roar and Anonymous: 2016 13.2 Images File:Acd_santa.jpg Source: http://upload.wikimedia.org/wikipedia/commons/2/28/Acd_santa.jpg License: CC-BY-SA-3.0 Contributors: Own work Original artist: Mdk572 File:Aleria,_Rhyton,_tte_de_chien.jpg Source: http://upload.wikimedia.org/wikipedia/commons/b/bc/Aleria%2C_Rhyton%2C_t% C3%AAte_de_chien.jpg License: CC-BY-SA-3.0-2.5-2.0-1.0 Contributors: Clio20, Own work, 2006-09-04 Original artist: Peintre de Brygos File:Ambox_globe_content.svg Source: http://upload.wikimedia.org/wikipedia/commons/b/bd/Ambox_globe_content.svg License: Public domain Contributors: Own work, using File:Information icon3.svg and File:Earth clip art.svg Original artist: penubag File:Anatomy_and_physiology_of_animals_The_ear.jpg Source: http://upload.wikimedia.org/wikipedia/commons/3/33/Anatomy_ and_physiology_of_animals_The_ear.jpg License: CC-BY-3.0 Contributors: Transferred from en.wikibooks; transferred to Commons by User:Adrignola using CommonsHelper. 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