The dog (Canis familiaris[4][5] or Canis lupus familiaris[5]) is
a domesticated descendant of the wolf. Also called the domestic dog, it
is derived from the extinct Pleistocene wolf,[6][7] and the modern wolf is the dog's
nearest living relative.[8] Dogs were the first species to be domesticated[9][8] by huntergatherers over 15,000 years ago[7] before the development of agriculture.[1] Due to
their long association with humans, dogs have expanded to a large number of
domestic individuals[10] and gained the ability to thrive on a starch-rich diet that would
be inadequate for other canids.[11]
The dog has been selectively bred over millennia for various behaviors, sensory
capabilities, and physical attributes.[12] Dog breeds vary widely in shape, size, and
color. They perform many roles for humans, such as hunting, herding, pulling
loads, protection, assisting police and the military, companionship, therapy,
and aiding disabled people. Over the millennia, dogs became uniquely adapted to
human behavior, and the human-canine bond has been a topic of frequent
study.[13] This influence on human society has given them the sobriquet of "man's
best friend".[14]
Taxonomy
Further information: Canis lupus dingo § Taxonomic debate – the domestic dog,
dingo, and New Guinea singing dog
In 1758, the Swedish botanist and zoologist Carl Linnaeus published in his Systema
Naturae, the two-word naming of species (binomial nomenclature). Canis is
the Latin word meaning "dog",[15] and under this genus, he listed the domestic dog,
the wolf, and the golden jackal. He classified the domestic dog as Canis
familiaris and, on the next page, classified the grey wolf as Canis lupus.[2] Linnaeus
considered the dog to be a separate species from the wolf because of its upturning
tail (cauda recurvata), which is not found in any other canid.[16]
In 1999, a study of mitochondrial DNA (mtDNA) indicated that the domestic dog may
have originated from the grey wolf, with the dingo and New Guinea singing
dog breeds having developed at a time when human communities were more
isolated from each other.[17] In the third edition of Mammal Species of the
World published in 2005, the mammalogist W. Christopher Wozencraft listed under
the wolf Canis lupus its wild subspecies and proposed two additional subspecies,
which formed the domestic dog clade: familiaris, as named by Linnaeus in 1758
and, dingo named by Meyer in 1793. Wozencraft included hallstromi (the New
Guinea singing dog) as another name (junior synonym) for the dingo. Wozencraft
referred to the mtDNA study as one of the guides informing his
decision.[3] Mammalogists have noted the inclusion of familiaris and dingo together
under the "domestic dog" clade[18] with some debating it.[19]
In 2019, a workshop hosted by the IUCN/Species Survival Commission's Canid
Specialist Group considered the dingo and the New Guinea singing dog to
be feral Canis familiaris and therefore did not assess them for the IUCN Red List of
Threatened Species.[4]
Evolution
Main article: Evolution of the wolf
Location of a dog's carnassials; the inside of the 4th upper premolar aligns with the outside of the 1st
lower molar, working like scissor blades
The Cretaceous–Paleogene extinction event occurred 66 million years ago and
brought an end to the non-avian dinosaurs and the appearance of the first
carnivorans.[20] The name carnivoran is given to a member of the order Carnivora.
Carnivorans possess a common arrangement of teeth called carnassials, in which
the first lower molar and the last upper premolar possess blade-like
enamel crowns that act similar to a pair of shears for cutting meat. This dental
arrangement has been modified by adaptation over the past 60 million years for diets
composed of meat, for crushing vegetation, or for the loss of the carnassial function
altogether as in seals, sea lions, and walruses. Today, not all carnivorans
are carnivores, such as the insect-eating aardwolf.[5]
The carnivoran ancestors of the dog-like caniforms and the cat-like feliforms began
their separate evolutionary paths just after the end of the dinosaurs. The first
members of the dog family Canidae appeared 40 million years ago,[21] of which only
its subfamily the Caninae survives today in the form of the wolf-like and fox-like
canines. Within the Caninae, the first members of genus Canis appeared six million
years ago,[15] the ancestors of modern domestic dogs, wolves, coyotes, and golden
jackals.
Domestication
Main article: Domestication of the dog
The earliest remains generally accepted to be those of a domesticated dog were
discovered in Bonn-Oberkassel, Germany. Contextual, isotopic, genetic, and
morphological evidence shows that this dog was not a local wolf.[22] The dog was
dated to 14,223 years ago and was found buried along with a man and a woman, all
three having been sprayed with red hematite powder and buried under large, thick
basalt blocks. The dog had died of canine distemper.[23] Earlier remains dating back
to 30,000 years ago have been described as Paleolithic dogs, but their status as
dogs or wolves remains debated[24] because considerable morphological diversity
existed among wolves during the Late Pleistocene.[1]
This timing indicates that the dog was the first species to be domesticated[9][8] in the
time of hunter–gatherers,[7] which predates agriculture.[1] DNA sequences show that
all ancient and modern dogs share a common ancestry and descended from an
ancient, extinct wolf population which was distinct from the modern
wolf lineage.[6][7] Most dogs form a sister group to the remains of a Late Pleistocene
wolf found in the Kessleroch cave near Thayngen in the canton
of Schaffhausen, Switzerland, which dates to 14,500 years ago. The most recent
common ancestor of both is estimated to be from 32,100 years ago.[25] This indicates
that an extinct Late Pleistocene wolf may have been the ancestor of the
dog,[8][1][26] with the modern wolf being the dog's nearest living relative.[8]
The dog is a classic example of a domestic animal that likely travelled
a commensal pathway into domestication.[24][27] The questions of when and where
dogs were first domesticated have taxed geneticists and archaeologists for
decades.[9] Genetic studies suggest a domestication process commencing over
25,000 years ago, in one or several wolf populations in either Europe, the high
Arctic, or eastern Asia.[10] In 2021, a literature review of the current
evidence infers that the dog was domesticated in Siberia 23,000 years ago
by ancient North Siberians, then later dispersed eastward into the Americas and
westward across Eurasia.[22]
Breeds
Main article: Dog breed
Further information: Dog type
Dog breeds show a range of phenotypic variation
Dogs are the most variable mammal on earth with around 450 globally
recognized dog breeds.[10] In the Victorian era, directed human selection developed
the modern dog breeds, which resulted in a vast range of phenotypes.[8] Most breeds
were derived from small numbers of founders within the last 200 years, [8][10] and since
then dogs have undergone rapid phenotypic change and were formed into today's
modern breeds due to artificial selection imposed by humans. The skull, body, and
limb proportions vary significantly between breeds, with dogs displaying more
phenotypic diversity than can be found within the entire order of carnivores. These
breeds possess distinct traits related to morphology, which include body size, skull
shape, tail phenotype, fur type and colour.[8] Their behavioural traits include guarding,
herding, and hunting,[8] retrieving, and scent detection. Their personality traits include
hypersocial behavior, boldness, and aggression,[10] which demonstrates the functional
and behavioral diversity of dogs.[8] As a result, present day dogs are the most
abundant carnivore species and are dispersed around the world. [10] The most striking
example of this dispersal is that of the numerous modern breeds of European
lineage during the Victorian era.[7]
Bangladeshi Dog
Biology
Anatomy
Main article: Dog anatomy
Skeleton
A lateral view of a dog skeleton
All healthy dogs, regardless of their size and type, have an
identical skeletal structure with the exception of the number of bones in the tail,
although there is significant skeletal variation between dogs of different
types.[28][29] The dog's skeleton is well adapted for running; the vertebrae on the neck
and back have extensions for powerful back muscles to connect to, the long ribs
provide plenty of room for the heart and lungs, and the shoulders are unattached to
the skeleton allowing great flexibility.[28][29]
Compared to the dog's wolf-like ancestors, selective breeding since domestication
has seen the dog's skeleton greatly enhanced in size for larger types as mastiffs and
miniaturised for smaller types such as terriers; dwarfism has been selectively utilised
for some types where short legs are advantageous such
as dachshunds and corgis.[29] Most dogs naturally have 26 vertebrae in their tails, but
some with naturally short tails have as few as three.[28]
The dog's skull has identical components regardless of breed type, but there is
significant divergence in terms of skull shape between types.[29][30] The three basic
skull shapes are the elongated dolichocephalic type as seen in sighthounds, the
intermediate mesocephalic or mesaticephalic type, and the very short and broad
brachycephalic type exemplified by mastiff type skulls.[29][30]
Senses
Further information: Dog anatomy § Senses
A dog's senses include vision, hearing, smell, taste, touch. One study suggested that
dogs can feel Earth's magnetic field.[31]
Coat
Main article: Dog coat
Dogs display wide variation in coat type, density, length, color, and composition
The coats of domestic dogs are of two varieties: "double" being familiar 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. Breeds may have an occasional
"blaze", stripe, or "star" of white fur on their chest or underside. [32] Premature graying
can occur in dogs from as early as one year of age; this is associated with impulsive
behaviors, anxiety behaviors, fear of noise, and fear of unfamiliar people or
animals.[33]
Tail
There are many different shapes for dog tails: straight, straight up, sickle, curled, or
corkscrew. As with many canids, one of the primary functions of a dog's tail is to
communicate their emotional state, which can be crucial in getting along with others.
In some hunting dogs the tail is traditionally docked to avoid injuries.
Health
Main article: Dog health
Some breeds of dogs are prone to specific genetic ailments such as elbow and hip
dysplasia, blindness, deafness, pulmonic stenosis, cleft palate, and trick knees. Two
severe medical conditions significantly affecting dogs are pyometra,
affecting unspayed females of all breeds and ages, and Gastric dilatation
volvulus (bloat), which affects larger breeds or deep-chested dogs. Both of these are
acute conditions and can kill rapidly. Dogs are also susceptible to parasites such
as fleas, ticks, mites, hookworms, tapeworms, roundworms, and heartworms, which
is a roundworm species that lives in the hearts of dogs.
Several human foods and household ingestible are toxic to dogs, including chocolate
solids, causing theobromine poisoning, onions and garlic,
causing thiosulphate, sulfoxide or disulfide poisoning, grapes and raisins, macadami
a nuts, and xylitol.[34] The nicotine in tobacco can also be dangerous to dogs. Signs of
ingestion can include copious vomiting (e.g., from eating cigar butts) or diarrhea.
Some other symptoms are abdominal pain, loss of coordination, collapse, or
death.[35][page needed]
Dogs are also vulnerable to some of the same health conditions as humans,
including diabetes, dental and heart disease, epilepsy, cancer, hypothyroidism,
and arthritis.
Lifespan
Further information: 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.[36][37] The median longevity of mixed-breed dogs,
taken as an average of all sizes, is one or more years longer than that of purebred
dogs when all breeds are averaged.[36][37][38] For dogs in England, increased body
weight has been found to be negatively correlated with longevity (i.e., the heavier the
dog, the shorter its lifespan), and mixed-breed dogs live on average 1.2 years longer
than purebred dogs.[39]
Reproduction
Main article: Canine reproduction
A female dog nursing newborn puppies.
In domestic dogs, sexual maturity happens around six months to one year for both
males and females, although this can be delayed until up to two years of age for
some large breeds, and is the time at which female dogs will have their first estrous
cycle. They will experience subsequent estrous cycles semiannually, during which
the body prepares for pregnancy. At the peak of the cycle, females will become
estrous, mentally and physically receptive to copulation. Because the ova survive
and can be fertilized for a week after ovulation, more than one male can sire the
same litter.[12]
Fertilization typically occurs two to five days after ovulation; 14–16 days after
ovulation, the embryo attaches to the uterus and after seven to eight more days, a
heartbeat is detectable.[40][41]
Dogs bear their litters roughly 58 to 68 days after fertilization,[12][42] with an average of
63 days, although the length of gestation can vary. An average litter consists of
about six puppies.[43]
Neutering
Neutering is the sterilization of animals, usually by removing the male's testicles or
the female's ovaries and uterus, to eliminate the ability to procreate and reduce sex
drive. Because of dogs' overpopulation in some countries, many animal control
agencies, such as the American Society for the Prevention of Cruelty to
Animals (ASPCA), advise that dogs not intended for further breeding should be
neutered, so that they do not have undesired puppies that may later be euthanized. [44]
According to the Humane Society of the United States, three to four million dogs and
cats are euthanized each year.[45] Many more are confined to cages in shelters
because there are many more animals than there are homes. Spaying or castrating
dogs helps keep overpopulation down.[46]
Neutering reduces problems caused by hypersexuality, especially in male
dogs.[47] Spayed female dogs are less likely to develop cancers affecting the
mammary glands, ovaries, and other reproductive organs.[48][page needed] However,
neutering increases the risk of urinary incontinence in female dogs[49] and prostate
cancer in males[50] and osteosarcoma, hemangiosarcoma, cruciate ligament rupture,
obesity, and diabetes mellitus in either sex.[51]
Inbreeding depression
A common breeding practice for pet dogs is mating between close relatives (e.g.,
between half and full siblings).[52] Inbreeding depression is considered to be due
mainly to the expression of homozygous deleterious recessive
mutations.[53] Outcrossing between unrelated individuals, including dogs of different
breeds, results in the beneficial masking of deleterious recessive mutations in
progeny.[54]
In a study of seven dog breeds (the Bernese Mountain Dog, Basset Hound, Cairn
Terrier, Brittany, German Shepherd Dog, Leonberger, and West Highland White
Terrier), it was found that inbreeding decreases litter size and survival.[55] Another
analysis of data on 42,855 Dachshund litters found that as the inbreeding
coefficient increased, litter size decreased and the percentage of stillborn puppies
increased, thus indicating inbreeding depression.[56] In a study of Boxer litters, 22% of
puppies died before reaching 7 weeks of age. Stillbirth was the most frequent cause
of death, followed by infection. Mortality due to infection increased significantly with
increases in inbreeding.[57]
Behavior
Main article: Dog behavior
See also: Dog behavior § Behavior compared with other canids
Dog swimming over to catch a ball, pay attention to the leg and tail movements
Dog behavior is the internally coordinated responses (actions or inactions) of the
domestic dog (individuals or groups) to internal and external stimuli.[58] As the oldest
domesticated species, dogs' minds inevitably have been shaped by millennia of
contact with humans. As a result of this physical and social evolution, dogs have
acquired the ability to understand and communicate with humans more than any
other species and they are uniquely attuned to human behaviors.[13] Behavioral
scientists have uncovered a surprising set of social-cognitive abilities in domestic
dogs. These abilities are not possessed by the dog's closest canine relatives or other
highly intelligent mammals, such as great apes, but rather parallel to children's
social-cognitive skills.[59]
Unlike other domestic species selected for production-related traits, dogs were
initially selected for their behaviors.[60][61] In 2016, a study found that only 11 fixed
genes showed variation between wolves and dogs.[62] These gene variations were
unlikely to have been the result of natural evolution and indicate selection on both
morphology and behavior during dog domestication. These genes have been shown
to affect the catecholamine synthesis pathway, with the majority of the genes
affecting the fight-or-flight response[61][63] (i.e., selection for tameness) and emotional
processing.[61] Dogs generally show reduced fear and aggression compared with
wolves.[61][64] Some of these genes have been associated with aggression in some dog
breeds, indicating their importance in both the initial domestication and later in breed
formation.[61] Traits of high sociability and lack of fear in dogs may include genetic
modifications related to Williams-Beuren syndrome in humans, which
cause hypersociability at the expense of problem-solving ability.[65]
Intelligence
Main article: Dog intelligence
Dog intelligence is the dog's ability to perceive information and retain it as knowledge
for applying to solve problems. Studies of two dogs suggest that dogs can learn by
inference and have advanced memory skills. A study with Rico, a Border Collie,
showed that he knew the labels of over 200 different items. He inferred the names of
novel things by exclusion learning and correctly retrieved those new items
immediately and four weeks after the initial exposure. A study of another Border
Collie, Chaser, documented his learning and memory capabilities. He had learned
the names and could associate by verbal command over 1,000 words.[66] Dogs can
read and react appropriately to human body language such as gesturing and
pointing and human voice commands.
One study of canine cognitive abilities found that dogs' capabilities are no more
exceptional than those of other animals, such as horses, chimpanzees,
or cats.[67] One limited study of 18 household dogs found that they lacked spatial
memory, and were more focused on the "what" of a task rather than the "where".[68]
Dogs demonstrate a theory of mind by engaging in deception.[69] An experimental
study showed compelling evidence that Australian dingos can outperform domestic
dogs in non-social problem-solving, indicating that domestic dogs may have lost
much of their original problem-solving abilities once they joined humans.[70] Another
study revealed that after undergoing training to solve a simple manipulation task,
dogs faced with an unsolvable version of the same problem look at the human, while
socialized wolves do not.[71]
Communication
Main article: Dog communication
Dog sounds
0:15
A dog making noises and barking
Problems playing this file? See media help.
Dog communication is how dogs convey information to other dogs, understand
messages from humans and translate the information that dogs are
transmitting.[72]: xii Communication behaviors of dogs include eye gaze, facial
expression, vocalization, body posture (including movements of bodies and limbs),
and gustatory communication (scents, pheromones, and taste). Humans
communicate to dogs by using vocalization, hand signals, and body posture.
Ecology
Population
The dog is probably the most widely abundant large carnivoran living in the human
environment.[73][74] In 2013, the estimated global dog population was between 700
million[75] and 987 million.[76] About 20% of dogs live as pets in developed
countries.[77] In the developing world, dogs are typically feral or communally owned,
with pet dogs uncommon. Most of these dogs live their lives as scavengers and have
never been owned by humans, with one study showing their most common response
when approached by strangers is to run away (52%) or respond aggressively
(11%).[78] Little is known about these dogs, or the dogs in developed countries that
are feral, strays, or are in shelters because the great majority of modern research on
dog cognition has focused on pet dogs living in human homes.[79]
Competitors and predators
Although dogs are the most abundant and widely distributed terrestrial carnivores,
feral and free-ranging dogs' potential to compete with other large carnivores is
limited by their strong association with humans.[73] For example, a review of the
studies in dogs' competitive effects on sympatric carnivores did not mention any
research on competition between dogs and wolves.[80][81] Although wolves are known
to kill dogs, they tend to live in pairs or in small packs in areas where they are highly
persecuted, giving them a disadvantage facing large dog groups.[80][82]
Wolves kill dogs wherever they are found together.[83] In some instances, wolves
have displayed an uncharacteristic fearlessness of humans and buildings when
attacking dogs to the extent that they have to be beaten off or killed.[84] Although the
numbers of dogs killed each year are relatively low, it induces a fear of wolves
entering villages and farmyards to take dogs and losses of dogs to wolves have led
to demands for more liberal wolf hunting regulations.[80]
Coyotes and big cats have also been known to attack dogs. In
particular, leopards are known to have a preference for dogs and have been
recorded to kill and consume them, no matter what their size.[85] Siberian tigers in
the Amur River region have killed dogs in the middle of villages. This indicates that
the dogs were targeted. Amur tigers will not tolerate wolves as competitors within
their territories, and the tigers could be considering dogs in the same way.[86] Striped
hyenas are known to kill dogs in their range.[87]
Diet
See also: Dog food
A Golden Retriever gnawing on a pig's foot
Dogs have been described as omnivores.[12][88][89] Compared to wolves, dogs from
agricultural societies have extra copies of amylase and other genes involved in
starch digestion that contribute to an increased ability to thrive on a starch-rich
diet.[11] Similar to humans, some dog breeds produce amylase in their saliva and are
classified as having a high starch diet.[90] However, more like cats and less like other
omnivores, dogs can only produce bile acid with taurine and they cannot
produce vitamin D, which they obtain from animal flesh. Also more like cats, dogs
require arginine to maintain its nitrogen balance. These nutritional requirements
place dogs halfway between carnivores and omnivores.[91]
Range
As a domesticated or semi-domesticated animal, the dog is nearly universal among
human societies. Notable exceptions once included:




The Aboriginal Tasmanians, who were separated from Australia before the
arrival of dingos on that continent
The Andamanese peoples, who were isolated when rising sea
levels covered the land bridge to Myanmar
The Fuegians, who instead domesticated the Fuegian dog, a different
canid species
Individual Pacific islands whose maritime settlers did not bring dogs, or
where dogs died out after original settlement, notably the Mariana
Islands,[92] Palau[93] and most of the Caroline Islands with exceptions such
as Fais Island and Nukuoro,[94] the Marshall Islands,[95] the Gilbert
Islands,[95] New
Caledonia,[96] Vanuatu,[96][97] Tonga,[97] Marquesas,[97] Mangaia in the Cook
Islands, Rapa Iti in French Polynesia, Easter Island,[97] the Chatham
Islands[98] and Pitcairn Island (settled by the Bounty mutineers, who killed
off their dogs to escape discovery by passing ships).[99]
Dogs were introduced to Antarctica as sled dogs, but were later outlawed by
international agreement due to the possible risk of spreading infections. [100]
Roles with humans
Domestic dogs inherited complex behaviors, such as bite inhibition, from their wolf
ancestors, which would have been pack hunters with a complex body language.
These sophisticated forms of social cognition and communication may account for
their trainability, playfulness and ability to fit into human households and social
situations. These attributes have given dogs a relationship with humans that has
enabled them to become one of the most successful animals today.[101]
The dogs' value to early human hunter-gatherers led to them quickly becoming
ubiquitous across world cultures. Dogs perform many roles for people, such
as hunting, herding, pulling loads, protection, assisting police and
the military, companionship and aiding disabled individuals. This influence on human
society has given them the nickname "man's best friend" in the Western world. In
some cultures, however, dogs are also a source of meat.[102][103]
Pets
Siberian Huskies are pack animals that still enjoy some human companionship
It is estimated that three-quarters of the world's dog population lives in the
developing world as feral, village, or community dogs, with pet dogs
uncommon.[104][page needed]
"The most widespread form of interspecies bonding occurs between humans and
dogs"[105] and the keeping of dogs as companions, particularly by elites, has a long
history.[14] Pet dog populations grew significantly after World War II as
suburbanization increased.[14] In the 1950s and 1960s, dogs were kept outside more
often than they tend to be today[106] (the expression "in the doghouse" - recorded
since 1932[107] - to describe exclusion from the group implies a distance between
the doghouse and the home) and were still primarily functional, acting as a guard,
children's playmate, or walking companion. From the 1980s, there have been
changes in the pet dog's role, such as the increased role of dogs in the emotional
support of their human guardians.[108][page needed] People and their dogs have become
increasingly integrated and implicated in each other's lives[109][page needed] to the point
where pet dogs actively shape how a family and home are experienced.[110]
There have been two significant trends occurring within the second half of the 20th
century in pet dogs' changing status. The first has been "commodification", shaping it
to conform to social expectations of personality and behavior.[110] The second has
been the broadening of the family's concept and the home to include dogs-as-dogs
within everyday routines and practices.[110]
A vast range of commodity forms aims to transform a pet dog into an ideal
companion.[111] 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.[111] Dog training books, classes, and television programs proliferated as
the process of commodifying the pet dog continued.[112]