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Friday, February 13, 2015

Whale


From Wikipedia, the free encyclopedia
Eubalaena glacialis with calf.jpg

North Atlantic right whales, mother and calf
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Subclass: Eutheria
Order: Cetacea

Whale (origin Old English hwæl from Proto-Germanic *hwalaz) is the common name for various marine mammals of the order Cetacea.[1] The term whale sometimes refers to all cetaceans, but more often it excludes dolphins and porpoises, which belong to the suborder Odontoceti (toothed whales). This suborder includes the sperm whale, killer whale, pilot whale, and beluga whale. The other cetacean suborder, Mysticeti (baleen whales), comprises filter feeders who eat small organisms caught by straining seawater through a comblike structure found in the mouth called baleen. This suborder includes the blue whale, the humpback whale, the bowhead whale and the minke whale. All cetaceans have forelimbs modified as fins, a tail with horizontal flukes, and nasal openings (blowholes) on top of the head.

Whales range in size from the blue whale, the largest animal known to have ever existed,[2] at 30 m (98 ft) and 180 tonnes (180 long tons; 200 short tons), to pygmy species such as the pygmy sperm whale at 3.5 m (11 ft). Whales inhabit all the world's oceans and number in the millions, with annual population growth rate estimates for various species ranging from 3% to 13%.[3] Whales are long-lived, humpback whales living for up to 77 years, while bowhead whales may live for more than a century.

Human hunting of whales from the seventeenth century until 1986 radically reduced the populations of some whale species.

Whales play a role in creation myths, for example among the Inuit, and they are revered by coastal people in countries such as Ghana and Vietnam.

Taxonomy

Cetaceans are divided into two suborders:
  • The largest suborder, Mysticeti (baleen whales), is characterized by the presence of baleen, a sieve-like structure in the upper jaw made of keratin, which it uses to filter plankton from the water.
  • Odontoceti (toothed whales) bear sharp teeth for hunting. Odontoceti includes dolphins and porpoises. If they were not considered to be whales, this would mean that the informal grouping 'whale' is not a clade.
Cetaceans and artiodactyls now are classified under the super-order Cetartiodactyla, which includes both whales and hippopotami. Whales are the hippopotamus's closest living relatives.[4]

Evolution


Ambulocetus natans – a primitive cetacean

All cetaceans, including whales, dolphins, and porpoises, are descendants of land-dwelling mammals of the artiodactyl order (even-toed ungulates). Both are related to the Indohyus, an extinct semi-aquatic deer-like ungulate, from which they split approximately 54 million years ago.[5][6]
These primitive cetaceans first took to the sea approximately 50 million years ago and became fully aquatic by 5–10 million years later.[7] Their features became adapted for living in the marine environment. Major anatomical changes include streamlining of the body, the migration of the nasal openings toward the top of the cranium, the shrinking and eventual disappearance of the hind limbs, the modification of the forelimbs into flippers, and the growth of flukes on the tail.

Anatomy

As with all mammals, whales breathe air, are warm-blooded, nurse their young with milk from mammary glands, and have body hair.[8] Beneath the skin lies a layer of fat called blubber, which stores energy and insulates the body. Whales have a spinal column, a vestigial pelvic bone, and a four-chambered heart. Typically, the neck vertebrae are fused, an adaptation trading flexibility for stability during swimming.[9][10]

Blowhole(s)


Features of a blue whale

Whales breathe via blowholes; baleen whales have two and toothed whales have one. These are located on the top of the head, allowing the animal to remain almost completely submerged while breathing. Breathing involves expelling stale air (which is warm and moist), as well as some mucus and excess water from the blowhole, forming an upward, steamy spout, followed by inhaling fresh air into the lungs.[11] Spout shapes differ among species, which facilitates identification.

Appendages

The body shape is fusiform and the modified forelimbs, or fins, are paddle-shaped. The end of the tail is composed of two flukes, which propel the animal by vertical movement, as opposed to the horizontal movement of a fish tail. Although whales do not possess fully developed hind limbs, some (such as sperm whales and baleen whales) possess discrete rudimentary appendages, which may contain feet and digits. Most species have a dorsal fin.[12][13]

During March 2014 in Japan a bottle-nosed dolphin with hind fins about as large as a human hand, was captured.[14]

Dentition

Toothed whales, such as the sperm whale, possess teeth with cementum cells overlying dentine cells. Unlike human teeth, which are composed mostly of enamel on the portion of the tooth outside of the gum, whale teeth have cementum outside the gum. Only in larger whales, where the cementum is worn away on the tip of the tooth, does enamel show.[15]

Instead of teeth, baleen whales have a row of baleen plates on the upper side of their jaws that resemble the teeth of a comb.

Ears

The whale ear has specific adaptations to the marine environment. In humans, the middle ear works as an impedance equalizer between the outside air's low impedance and the cochlear fluid's high impedance. In aquatic mammals, such as whales, however, there is no great difference between the outer and inner environments. Instead of sound passing through the outer ear to the middle ear, whales receive sound through the throat, from which it passes through a low-impedance fat-filled cavity to the inner ear.[16] The whale ear is acoustically isolated from the skull by air-filled sinus pockets, which allow for greater directional hearing underwater.[17]

Life history and behavior

Reproduction

Males are called 'bulls', females, 'cows' and all newborns, 'calves'. Most species do not maintain fixed reproductive partnerships. Females have several mates each season.[18][19]
The female usually delivers a single calf, which is birthed tail-first to minimize the risk of drowning. Whale cows nurse by squirting milk into the mouths of their young. This milk is so rich in fat that it has the consistency of toothpaste.[18] In many species, nursing continues for more than a year and is associated with a strong bond between mother and calf. Reproductive maturity typically occurs at seven to ten years. This mode of reproduction produces few offspring, but increases the survival probability of each one.

Socialization

Whales are known to teach, learn, cooperate, scheme, and even grieve.[20] The neocortex of many species of whale is home to elongated spindle neurons that, prior to 2007, were known only in hominids.[21] In humans these cells are involved in social conduct, emotions, judgment, and theory of mind.[22] Whale spindle neurons are found in areas of the brain that are homologous to where they are found in humans, suggesting that they perform a similar function.[23]

Sleep

Photo of humpback whale with most of its body out of the water and its pectoral fins extended
A humpback whale breaching

Unlike most animals, whales are conscious breathers. All mammals sleep, but whales cannot afford to become unconscious for long because they may drown. While knowledge of sleep in wild cetaceans is limited, toothed cetaceans in captivity have been recorded to sleep with one side of their brain at a time, ostensibly so that they may swim, breathe consciously, avoid both predators and social contact during their period of rest. It is thought that only one hemisphere of the whale's brain sleeps at a time, so that they rest, but are never completely asleep.[24]

A 2008 study found that wild sperm whales (Physeter macrocephalus) sleep in vertical postures just under the surface in passive shallow 'drift-dives', generally during the day, during which whales do not respond to passing vessels unless they are in contact, leading to the suggestion that whales possibly sleep during such dives.[25]

Surfacing behavior

Many whales exhibit behaviors that expose large parts of their bodies to the air, such as breaching and tail slapping.

Sounding

Sounding is a term used for whales diving. Typically it is only used for longer dives. Before sounding, whales typically stay close to the surface for a series of short, shallow dives while building their oxygen reserves. They then make a sounding dive.

Lifespan

Whale lifespans vary among species and are not well characterized. Whaling left few older individuals to observe directly. R.M. Nowak of Johns Hopkins University estimated that humpback whales may live as long as 77 years.[26] In 2007, a nineteenth-century lance fragment was found in a bowhead whale off Alaska, which suggests the individual could be between 115 and 130 years old.[27] Aspartic acid racemization in the whale eye, combined with a harpoon fragment, indicated an age of 211 years for another male, which, if true, would make bowheads the longest-lived extant mammal species.[28][29] The accuracy of this age determination method has been questioned because racemization does not correlate well with other dating methods.[30]

Vocalization

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Recording of Humpback Whales singing and Clicking.

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Some species, such as the humpback whale, communicate using melodic sounds, known as whale song. These sounds may be extremely loud, depending on the species. Sperm whales only have been heard making clicks, while toothed whales (Odontoceti) use echolocation that may generate approximately 20,000 watts of sound (+73 dBm or +43 dBw)[31] and be heard for many miles. Whale vocalization is likely to serve many purposes, including echolocation, mating, and identification.[32]

Captive whales occasionally have been known to mimic human speech. Scientists have suggested this indicates a strong desire on behalf of the whales to communicate with humans, as whales have a very different vocal mechanism, so producing human speech likely takes considerable effort.[33]

Ecology


"Whale pump" - the role played by whales in nutrient recycling in the oceans

Whales are considered as "marine ecosystem engineers" for the following reasons:[34]
  • Whales are major consumers of fish and oceanic invertebrates.
  • Whales act as reservoirs of nutrients, such as iron and nitrogen, and they recycle them both horizontally and vertically in the water column.
  • Whale detritus provides energy and habitat for deep sea organisms.

Feeding

Whales generally are classed as predators. Their food ranges from microscopic plankton to very large animals.

Toothed whales eat fish and squid, which they hunt by the use of echolocation. Killer whales sometimes eat other marine mammals, including whales.

Baleen whales, such as humpbacks and blues, mainly eat krill when feeding in the higher latitudes (such as the Southern Ocean). They take in enormous amounts of seawater that they expel through their baleen plates; the krill in the seawater are retained on the plates and then swallowed.[18] Whales do not drink seawater. They extract water indirectly from their food by metabolizing fat.[18]

Whale pump

A 2010 study considered whales to be a positive influence to the productivity of ocean fisheries, in what has been termed a "whale pump." Whales carry nutrients such as nitrogen from the depths back to the surface. This functions as an upward biological pump, reversing an earlier presumption that whales accelerate the loss of nutrients to the bottom. This nitrogen input in the Gulf of Maine is "more than the input of all rivers combined" emptying into the gulf, some 23,000 metric tons each year.[35][36] Whales defecate at the oceans surface and this excrement is important for fisheries because it is rich in iron and nitrogen. The whale feces are liquid and instead of sinking, they stay at the surface where phytoplankton feed off it.[37]

Whale fall

Upon death, whale carcasses fall to the deep ocean and being massive, with body weights of the range 30 to 160 tonnes (30,000 to 160,000 kg), provide a substantial habitat for marine creatures. Evidence of whale falls in present day and fossil records shows that deep sea whale falls support a rich assemblage of creatures, with a global diversity of 407 species as per Smith & Baco (2003), comparable to other neritic biodiversity hotspots, such as cold seeps and hydrothermal vents.[38]
Deterioration of whale carcasses happens though a series of three stages. Initially, moving organisms such as sharks and hagfish, scavenge the soft tissues at a rapid rate over a period of months, and as long as two years. This is followed by the colonisation of bones and surrounding sediments (which contain organic matter) by enrichment opportunists, such as crustaceans and polychaetes, throughout a period of years. Finally, sulfophilic bacteria reduce the bones releasing hydrogen sulfide enabling the growth of chemoautotrophic organisms, which in turn, support other organisms such as mussels, clams, limpets, and sea snails. This stage may last for decades and supports a rich assemblage of species, averaging 185 species per site as per Smith & Baco (2003).[38]

Interaction with humans

Whaling

Dutch whalers near Spitsbergen, Abraham Storck, 1690
Map showing IWC non-members such as Canada and most Middle Eastern and African countries in white
World map of International Whaling Commission (IWC) members/non-members(member countries in blue)
Diagram showing the pre-whaling of 275,000, 1930s population of 30–40,000, mid-60s population of 650–2,000 and 1994 population of fewer than 5,000
World population graph of blue whales (Balaenoptera musculus)

Some species of large whales are listed as endangered by multinational organizations, such as CITES, as well as governments and advocacy groups. This status is due primarily to the impact of whaling. Whales have been hunted commercially since the seventeenth century for whale oil, meat, baleen, and ambergris (a perfume ingredient from the intestine of sperm whales).[39] More than two million whales were taken during the twentieth century,[40] and by the middle of that century, many populations were severely depleted.

The International Whaling Commission banned commercial whaling in 1986.[41] The ban is not absolute, however, and some whaling continues under the auspices of scientific research[41] (sometimes not proved[42]) or aboriginal rights. Current whaling nations are Norway, Iceland, and Japan as well as the aboriginal communities of Siberia, Alaska, and northern Canada.

Captivity

Belugas and orcas have been kept in captivity since 1861[43] and 1961,[44][45] respectively, for public display in a few locations.[46][47] They are popular due to their intelligence, trainability,[48][49] striking appearance,[50] playfulness in captivity,[51][52] and size. Belugas have also been kept captive for naval research in the US[53] and Russia.[54]

Most current captives were caught in the wild, since captive breeding has had limited success.[55] There is controversy over captivity,[46][56] with limited enrichment activities and tank sizes, though defenders say easy access for research[49][56][57][58] and public viewing are beneficial.[59] The whales have far larger family groups and ranges in the wild[60] than in captivity.

Whale watching

An estimated 13 million people went whale watching globally in 2008, in all oceans except the Arctic.[61][62] There are numerous rules and codes of conduct to minimize harassment of the whales.[63] Iceland, Japan and Norway have both whaling and whale watching industries. Whale watching lobbyists are concerned that the most inquisitive whales, which approach boats very closely and provide much of the entertainment on whale-watching trips, will be the first to be taken if whaling is done in the same areas.[64]

Bycatch

Several species of small whales are caught as bycatch while fishing for other species. In the Eastern Tropical Pacific tuna fishery, thousands of dolphins drowned in purse-seine nets, until preventive measures were introduced. Gear and deployment modifications, and eco-labelling (dolphin-safe or dolphin-friendly brands of tuna), have contributed to a reduction in dolphin mortality by tuna vessels.[65]

Naval sonar

Environmentalists speculate that advanced naval sonar endangers some cetaceans, including whales. In 2003, British and Spanish scientists suggested in Nature that the effects of sonar trigger whale beachings and they point to signs that such whales have experienced decompression sickness.[66] 
Responses in Nature the following year discounted the explanation.[67]Mass beachings occur in many species, mostly beaked whales that use echolocation for deep diving. The frequency and size of beachings around the world, recorded throughout the last thousand years in religious tracts, and more recently in scientific surveys, have been used to estimate the population of various whale species by assuming that the proportion of the total whale population beaching in any one year is constant. Beached whales can give other clues about population conditions, especially health problems. For example, bleeding around ears, internal lesions, and nitrogen bubbles in organ tissue suggest decompression sickness.[20]

Following public concern, the U.S. Defense department was ordered by the 9th Circuit Court to strictly limit use of its Low Frequency Active Sonar during peacetime. Attempts by the UK-based Whale and Dolphin Conservation Society to obtain a public inquiry into the possible dangers of the Royal Navy's equivalent (the "2087" sonar launched in December 2004) failed as of 2008. The European Parliament has requested that EU members refrain from using the powerful sonar system until an environmental impact study has been carried out.[citation needed]

In fiction

The 1851 American novel, Moby-Dick by Herman Melville concerns a vexed captain's hunt for a gigantic white whale. Rudyard Kipling's 1902 Just So Stories includes the tale of "How the Whale got his Throat".[68] The film Whale Rider directed by Niki Caro has a Maori girl ride a whale in her quest to be a suitable heir to the chiefship.[69] An enormous whale called Monstro is the final antagonist featured in Walt Disney's 1940 animated film Pinocchio.

In mythology


Whale weather-vane atop the Nantucket Historical Association Whaling Museum displaying a sperm whale

Whales were little understood for most of human history as little of their lives could be seen from the surface of the ocean.[70] Many cultures, even those that have hunted them, hold whales in awe and feature them in their mythologies. In China, Yu-kiang, a whale with the hands and feet of a human was said to rule the ocean.[71] In the Tyrol region of Austria, it was said that if a sunbeam were to fall on a girl entering puberty, she would be carried away in the belly of a whale.[72] Paikea, the youngest and favourite son of the chief Uenuku from the island of Mangaia (Cook Islands), was said by the Kati Kuri people of Kaikoura to have traveled from the Pacific Islands on the back of a whale many centuries before.[73] The whale features in Inuit creation myths. When 'Big Raven', a deity in human form, found a stranded whale, he was told by the Great Spirit where to find special mushrooms that would give him the strength to drag the whale back to the sea and thus, return order to the world.[71] The Tlingit people of northern Canada said that the Orcas were created when the hunter Natsihlane carved eight fish from yellow cedar, sang his most powerful spirit song, and commanded the fish to leap into the water.[74] In an Icelandic legend a man threw a stone at a fin whale and hit the blowhole, causing the whale to burst. The man was told not to go to sea for twenty years, but during the nineteenth year he went fishing and a whale came and killed him.[75] In East African legend, King Sulemani asked God that he might permit him to feed all the beings on earth. A whale came and ate until there was no corn left and then told Sulemani that he still was hungry and that there were 70,000 more in his tribe. Sulemani then prayed to God for forgiveness and thanked the creature for teaching him a lesson in humility.[71]

Some cultures associate divinity with whales, such as among Ghanaians and Vietnamese, who occasionally hold funerals for beached whales, a throwback to Vietnam's ancient sea-based Austro-Asiatic culture.[76][77][78][79]

The Bible mentions whales in Genesis 1:21, Job 7:12, Ezekiel and 32:2. The "sea monsters" in Lamentations 4:3 have been taken by some commentators to refer to marine mammals, in particular whales, although most modern versions use the word "jackals" instead.[80] The story of Jonah being swallowed by a "big Fish" is told both in the Qur'an and in the Bible.[81] The Old Testament contains the Book of Jonah and in the New Testament, Jesus mentions this story in Matthew 12:40.[82]

The engraving by William van der Gouwen shows a 20 m (65.6 ft) long whale, stranded on the Dutch coast between Scheveningen and Katwijk on 3 February 1598

In music

And God Created Great Whales, written in 1970 by American composer Alan Hovhaness, is a work for orchestra and whale songs, including the recorded sounds of humpback, bowhead, and killer whales.[83] The song "Il n'y a plus rien", from French singer-songwriter Léo Ferré's eponymous album (1973), is an example of biomusic that begins and ends with recorded whale songs mixed with a symphonic orchestra and his voice.

Origin of the domestic dog


From Wikipedia, the free encyclopedia


DNA evidence indicates that the dog and the modern wolf (above) are both descendents of an extinct wolf-like canid that lived in Europe.

Some dog breeds, like this Tamaskan Dog, look very much like wolves due to admixture.

The 14,500 year old upper-right jaw found in Kesslerloch Cave, Switzerland is the sister to 2/3 of modern dogs (courtesy Hannes Napierala)

The domestic dog (Canis lupus familiaris or Canis familiaris), based on nuclear DNA evidence as of January 2014, began from a single domestication 11 to 16 thousand years ago that predates the rise of agriculture and implies that the earliest dogs arose along with hunter-gatherers and not agriculturists.[1] Mitochondrial DNA evidence as of November 2013 indicates that all modern dogs are most closely related to the extant and extinct canids of Europe[2][3] compared to earlier writers who proposed the origins from Eurasia as well as Eastern Asia.[4][5][6]

The analysis indicates that the dog is not a descendant of the extant (i.e. living) Gray wolf but forms a sister clade, and that dogs were originally domesticated from a now-extinct wolf population that was more genetically diverse than today’s wolf population. The dog's genetic closeness to modern wolves is due to admixture.[1][7]

Conceivably, proto-dogs might have taken advantage of carcasses left on site by early hunters, assisted in the capture of prey, or provided defense from large competing predators at kills. Furthermore, several ancient dogs may represent failed domestication events, such as the 36,000 year old Goyet specimen of Belgium and the 33,000 year old Altai Mountains specimen from Russia, as they have no descendents today.[2][8]

Genetic evidence

The development of molecular biology allows us to infer evolutionary relationships for species and to represent them in a phylogenetic tree, however these are not without their limitations.
There are three schools of thought on the origin of the dog.

Descendant of other canids that exist today

Within the Canidae, three distinct phylogenetic groupings are apparent:
  1. The fox-like canids, which include species closely related to the red fox (Vulpes vulpes), and the arctic fox (Vulpes lagopus), and the fennec fox (Vulpes zerda or Fennecus zerda, disputed).
  2. The wolf-like canids, including dog, wolf, coyote, Ethiopian wolf or Simien jackal, and three other species of jackals (genus Canis), and the African hunting dog (genus Lycaon) and the dhole (genus Cuon).
  3. The South American canids, including fox-sized canids, such as the pampus fox, crab-eating fox, and small-eared dog (genus Pseudalopex, Lycalopex, Atelocynus) and the maned wolf (genus Chrysocyon) and bushdog (genus Speothos).
Additionally, there are several canids that have no close living relatives and define distinct evolutionary lineages; such as the gray fox (genus Urocyon), the bat-eared fox (genus Otocyon), and the raccoon dog (genus Nyctereutes).

These phylogenetic relationships imply that the dog has several close relatives within its genus, in fact, all members of Canis can produce fertile hybrids and several species may have genomes that reflect hybridization in the wild.[9] Charles Darwin speculated in On the Origin of Species that given the vast morphological variation across numerous breeds, dogs must have had more than one wild ancestor.[10] Therefore, it was once thought possible that the dog could have been a descendant of a number of canids that exist today—similar to the Coyote-Wolf-Dog hybrid found among some Eastern Coyotes[11]—until DNA analysis indicated otherwise.

Descendant of a wolf that exists today

It has been thought for some time that "... the wolf is the most probable ancestor and closest relative of the domestic dog."[12]:54 By 1993 with advancements in molecular biology, the mitochondrial DNA mtDNA analysis of extant (i.e., living today) Canidea species indicated that "The domestic dog is an extremely close relative of the gray wolf, differing from it by at most 0.2% of mtDNA sequence.... In comparison, the gray wolf differs from its closest wild relative, the coyote, by about 4% of mitochondrial DNA sequence."[13] In the same year, the domestic dog Canis familiaris was reclassified as Canis lupus familiaris, a subspecies of the gray wolf Canis lupus in Mammal Species of the World.[14] In 1995, research indicated that the wolf may have been the ancestor of the dog.[15]
By 1999, further genetic analysis indicated that the domestic dog may have emerged from multiple wolf populations.[16][17] Based on these pieces of research and the reference reclassification, canis lupis familiaris is the name for the taxon listed by ITIS.[18] However, canis familiaris is also accepted due to a nomenclature debate regarding the naming of wild and domestic sub-species.[19]

Descendant of a wolf that no longer exists today

Location of the first domestication

In 1997, a study determined that domestic dogs could have originated as much as 135,000 years ago.[20] This was determined by using mitochondrial DNA (mtDNA). However, this number could potentially be overinflated due to the possibility of unobserved multiple substitutions at hypervariable sites.[20] These hypervariable sites are mutational hotspots where somatic mitochondrial DNA and germline mutations usually take place.[21] Though this number of 135,000 years may not be completely accurate, it still gives the indication that domesticated dogs have been around for longer than 14,000 years, which was previously suggested by archaeological evidence.[20]

In 2002, a study examined the mitochondrial DNA sequence variation among 654 domestic dogs representing all major dog populations worldwide. All sequences belonged to three phylogenetic groups universally represented at similar frequencies, suggesting a common origin from a single gene pool for all dog populations. A larger genetic variation in East Asia than in other regions and the pattern of phylogeographic variation suggested an East Asian origin for the domestic dog around 15,000 years ago.[22] In 2008, a study of several skulls of fossil large canids from sites in Belgium, Ukraine and Russia were examined to look for possible evidence of the presence of Palaeolithic dogs. The fossil large canid from Goyet (Belgium), dated at c. 31,700 BP was identified as a Palaeolithic dog, suggesting that dog domestication had already started in Europe during the Aurignacian.[23][24][25]

In 2009, a study of African dogs sampled 318 village dogs from 7 regions in Egypt, Uganda, and Namibia, measuring genetic diversity >680 bp of the mitochondrial D-loop, 300 SNPs, and 89 microsatellite markers. This study found a high level of mtDNA diversity in African dogs.[26] Later in 2009, a study analyzed entire mitochondrial genomes for 169 dogs to obtain maximal phylogenetic resolution and the control region for 1,543 dogs across the Old World for a comprehensive picture of geographical diversity. The analyses showed that dogs universally share a common homogenous gene pool containing 10 major haplogroups. However, the full range of genetic diversity, all 10 haplogroups, was found only in southeastern Asia south of Yangtze River, and diversity decreased following a gradient across Eurasia, through seven haplogroups in Central China and five in North China and Southwest Asia, down to only four haplogroups in Europe. The mean sequence distance to ancestral haplotypes indicates an origin 5,400–16,300 years ago (ya) from at least 51 female wolf founders.[27]

In 2010, a study conducted an extensive genome-wide survey of more than 48,000 single nucleotide polymorphisms in dogs and their wild progenitor, the grey wolf. It showed that dog breeds share a higher proportion of multi-locus haplotypes unique to grey wolves from the Middle East, indicating that they are a dominant source of genetic diversity for dogs rather than wolves from East Asia, as suggested by mitochondrial DNA sequence. This indicates the Middle East as the source of domestication.[28] In 2011, a study analyzed 14 437 bp of Y-chromosome DNA sequence in 151 dogs sampled worldwide. It found 28 haplotypes distributed in five haplogroups. Two haplogroups were universally shared and included three haplotypes carried by 46% of all dogs, but two other haplogroups were primarily restricted to East Asia. Highest genetic diversity and virtually complete phylogenetic coverage was found within Asia south of the Yangtze. The 151 dogs were estimated to originate from 13–24 wolf founders, but there was no indication of post-domestication dog–wolf hybridisations. Thus, Y-chromosome and mtDNA data give strikingly similar pictures of dog phylogeography, most importantly that roughly 50% of the gene pools are shared universally but only Asia south of the Yangtze has nearly the full range of genetic diversity, such that the gene pools in all other regions may derive from Asia south of the Yangtze. This corroborates that Asia south of the Yangtze was the principal, and possibly sole region of wolf domestication, that many wolves were domesticated, and that later dog–wolf hybridisation contributed modestly to the dog gene pool.[29]

In 2011, a study looked at skulls from the Gravettian Předmostí site in the Czech Republic. Three were dated to 24,000 and 27,000 years old and identified as Palaeolithic dogs, characterized by short skull lengths, short snouts, and wide palates and braincases relative to wolves. One complete skull could be assigned to the group of Pleistocene wolves. Three other skulls could not be assigned to a reference group; these might be remains from hybrids or captive wolves.[30]

"Wayne" studies


33,000 year old skull of a dog-like canid found in the Altai Mountains. It has no descendants today

In 1934, an eminent paleontologist indicated that the ancestor of the dog lineage may have been an extinct Canis lupus.[31] In 1999, Robert K Wayne emphasized that while molecular genetic data seem to support the origin of dogs from wolves, dogs may have descended from a now extinct species of canid whose closest living relative was the wolf.[16] Between March 2013 and January 2014, three studies were released that undertook similar investigation using the same or similar samples and each had Wayne as a senior author. These studies appear to have validated Wayne's original hypothesis.
Druzhkova et al.
In March 2013, a study[32] isolated, sequenced and analysed 413 nucleotides of the mitochondrial control region of ancient DNA from the well-preserved 33,000-year old skull of a dog-like canid that was excavated from Razboinichya Cave in the Altai Mountains of southern Siberia (Central Asia) in 1975. The sample was deposited in GenBank with accession number JX173682 and classified as Canis lupus familiaris (dog). The sample was compared with those of 72 extant dogs and 30 wolves (17 Old World and 13 New World), 35 prehistoric New World canids (including 2 from the Beringian wolf), and 4 coyotes. The sample was also compared to 3 ancient wolf teeth also found in the cave (32,500 BP, 48,000 BP and 50,000 BP). "The analyses revealed that the unique haplotype of the Altai dog is more closely related to modern dogs and prehistoric New World canids than it is to contemporary wolves... This preliminary analysis affirms the conclusion that the Altai specimen is likely an ancient dog with shallow divergence from ancient wolves. These results suggest a more ancient history of the dog outside of the Middle East or East Asia." The haplotype groups closest to the Altai dog included such diverse breeds as the Tibetan Mastiff, Newfoundland, Chinese Crested, Cocker Spaniel and Siberian Husky. The study also stated "We stress the point that these analyses were limited to a single, maternally inherited locus and more sequence data would be needed to obtain a statistically well-supported phylogeny and unambiguously resolve the genetic relationship of the Altai specimen....More data of prehistoric wolves from the same region are needed to estimate the population diversity and obtain a more comprehensive picture of genetic relationships of the Altai canid."

One of the team members of this study had conducted an earlier study,[33] which used Accelerator Mass Spectrometry radiocarbon dating independently conducted through 3 laboratories located in Tucson, Arizona (USA), Oxford (UK), and Groningen (the Netherlands) to derive an age of 33,000-years for the well-preserved skull and left mandible of the Altai dog. The morphology was compared to the skulls and mandibles of large Pleistocene wolves from Predmosti, Czech Republic dated 31,000 BP, modern wolves from Europe and North America, and prehistoric Greenland dogs from the Thule period (1,000 BP or later) to represent a large-sized but unimproved fully domestic dogs. The morphologies best matched the Greenland dogs and not the ancient nor modern wolves. However, the lower carnassial tooth fell within the lower range of values for prehistoric wolves and was only slightly smaller than modern European wolves, and the upper carnassial tooth fell within the range of modern wolves. "We conclude, therefore, that this specimen may represent a dog in the very early stages of domestication, i.e. an incipient dog, rather than an aberrant wolf... The Razboinichya Cave specimen appears to be an incipient dog that did not give rise to late Glacial – early Holocene lineages and probably represents wolf domestication disrupted by the climatic and cultural changes associated with the Last Glacial Maximum. The two earliest incipient dogs from Western Europe (Goyet, Belgium) and Siberia (Razboinichya), separated by thousands of kilometers, show that dog domestication was multiregional, and thus had no single place of origin (as some DNA data have suggested) and subsequent spread."
Thalmann et al.
In November 2013, a study[2] analysed the complete and partial mitochondrial genomes of 18 fossil canids dating from 1,000 to 36,000 years ago from the Old and New Worlds, and compared these with the complete mitochondrial genome sequences from 49 modern wolves and 77 modern dogs—including divergent dog breeds, such as the Basenji and Dingo—3 recently published Chinese indigenous dogs, and 4 coyotes totaling 148 mitochondrial genomes. Mitochondrial DNA was analyzed because there is a higher abundance of mitochondrial than nuclear DNA available from ancient specimens. The data indicates that 22% (17 of 77) of the dogs sampled are sister to modern wolves from Sweden and the Ukraine with a most recent common ancestor 9,200 years ago (else the mitochondrial genome introgressed from wolves, as dogs were clearly domesticated by this time), and 78% (60 of 77) are sister to one or more ancient canids from Europe. Some 12% (9 of 77) of the dogs are sister to two morphologically distinct ancient dogs from Germany, one 14,700 years old from Bonn-Oberkassel (GenBank accession number KF661093) and one 12,500 years old from the Kartstein cave (KF661094) with a most recent common ancestor 16,000-24,000 years ago. [Note: the study's Figure 1 shows the Phylogenetic relationships and indicates that this most recent common ancestor was also dog-like, and separated from what was to become the ancestor of many of the modern Eurasian wolf lines at 24,000-31,000 years ago.] Some 3% (2 of 77) of the dogs are sister to the 14,500 year old wolf sequence from the Kesslerloch cave in Switzerland (KF661087) with a most recent common ancestor 18,300 years ago. They are also distantly rooted in the same sequence as the Altai dog, however the study does not support its recent common ancestry with most modern dogs.
Some 64% (49 of 77) of the dogs are sister to another 14,500-year-old wolf sequence also from the Kesslerloch cave in Switzerland (KF661091) with a most recent common ancestor 32,100 years ago. This group of dogs matches 3 fossil pre-Columbian New World dogs between 1,000 and 8,500 years old. Matching these 3 to the 49 relatives indicates a most recent common ancestor 18,800 years ago, which supports the hypothesis that pre-Columbian dogs in the New World share ancestry with modern dogs. Thus, these dogs likely arrived with the first humans to the New World. The early dog population appears to have undergone a population bottle-neck (decrease) between 5,000 and 2,000 years ago, followed by a sharp increase that parallels the trajectory of the human population.

Belgium 26,000 - Pleistocene wolf skull from the Trou des Nutons cave in Belgium dated to be 26,000 years old (courtesy Royal Belgian Institute of Natural Sciences)

Three haplotypes from ancient Belgium canids (Belgium 36,000 years BP cataloged as Canis species KF661079 and Belgium 30,000 and 26,000 years BP cataloged as Canis lupus KF661080 KF661078) form the most diverging group. Although the cranial morphology of the Goyet dog (Belgium 36,000) has been interpreted as dog-like, it's mtDNA relation to other canids places it as an ancient sister-group to all modern dogs and wolves rather than a direct ancestor. Belgium 26,000 has been found to be uniquely large and could be related to a distinct genetically and morphologically form of wolves from Late Pleistocene deposits in the High Arctic permafrost. However, none of the sequences from three northerly permafrost wolves (dated at 28,000 years BP, 21,000 years BP, and 20,800 years BP and classified as Canis lupus KF661088, KF661089, KF661090) fall within or are sister to this Belgium clade. The Belgium canids may represent a phenotypically distinct and not previously recognized population of gray wolf, or the 36,000-year-old Goyet dog from Belgium and the 33,000-year-old Altai Mountain dog from Russia (catalogued as Canis lupus familiaris - dog JX173682), may represent aborted domestication episodes.

The data from this study indicates an association of modern dogs with ancient European canids and some with modern European wolves, with no close affinity to modern wolves from the Middle East or East Asia. This suggests the origin of dogs from Europe, rather than the Middle East or East Asia, as previously suggested. Additionally, there are no ancient dog remains from these regions older than 13,000 years. Divergence times implies a European origin of the domestic dog dating 18,800-32,100 years ago, which supports the hypothesis that dog domestication preceded the emergence of agriculture and occurred in the context of European hunter-gatherer cultures. An evolutionary scenario consistent with these results is that dog domestication was initiated close to the Last Glacial Maximum when hunter-gatherers preyed on megafauna.[34][8]

The co-senior author of the study, Robert K Wayne, stated that:
"But if domestication occurred in association with hunter-gatherers, one can imagine wolves first taking advantage of the carcasses that humans left behind - a natural role for any large carnivore - and then over time moving more closely into the human niche through a co-evolutionary process. The idea of wolves following hunter-gatherers also helps to explain the eventual genetic divergence that led to the appearance of dogs. Wolves following the migratory patterns of these early human groups would have given up their territoriality and would have been less likely to reproduce with resident territorial wolves. We have an analog of this process today, in the only migratory population of wolves known existing in the tundra and boreal forest of North America. This population follows the barren-ground caribou during their thousand-kilometer migration. When these wolves return from the tundra to the boreal forest during the winter, they do not reproduce with resident wolves there that never migrate. We feel this is a model for domestication and the reproductive divergence of the earliest dogs from wild wolves. We know also that there were distinct wolf populations existing ten of thousands of years ago. One such wolf, which we call the megafaunal wolf, preyed on large game such as horses, bison and perhaps very young mammoths. Isotope data show that they ate these species, and the dog may have been derived from a wolf similar to these ancient wolves in the late Pleistocene of Europe....A dog from Belgium dates back approximately 36,000 years, and a group of dogs from Western Russia is approximately 15,000 years old." [3]
Freedman et al.
In January 2014, a study[1] analysed using a number of tests the 10 million single-nucleotide variant sites from whole-genome data generated for six unique canid lineages. These data include whole-genome sequences of:
  • Three individual wolves (Canis lupus) to represent the broad regions of Eurasia where domestication has been hypothesized to have taken place - Croatia (Europe), Israel (Middle-East), and China (East/South-East Asia).
  • An Australian dingo and a basenji, being divergent lineages to the reference boxer genome (that is available from GenBank with accession number AAEX01000000) and so maximize the odds to capture distinct alleles present in the earliest dogs. These lineages are also geographically distinct, with modern basenjis tracing their ancestry to hunting dogs of western Africa, while dingoes are free-living semi-feral dogs of Australia that arrived there at least 3,500 years ago. The natural range of wolves has never extended this far south and due to geographic isolation, they are less likely to have overlapped and admixed with wolves in the recent past. For some analyses, data was leveraged from a companion study of 12 additional dog breeds.
  • A golden jackal (Canis aureus) to infer the ancestral state of variants arising in dogs and wolves.[1]
The data provided significant evidence of admixture between the Israeli wolf and the basenji, the Israeli wolf and the boxer, and between the Chinese wolf and dingo. The Chinese wolf with dingo likely represents ancient admixture in Eastern Eurasia, and the Israeli wolf with Basenji and Boxer likely represents ancient admixture in Western Eurasia. The fact that these lineages have been geographically isolated from wolves in the recent past suggests that this gene flow was ancestral and has likely impacted on most dog lineages. There was significant gene flow between the golden jackal and the Israeli wolf, as well as the population ancestral to the dog and wolf samples.[1]

One test indicated that dogs and modern wolves form monophyletic sister clades i.e. the dog is a sister to the modern wolf and they share a common ancestor. Supporting this, another test indicated that none of the sampled wolf populations is more closely related to dogs than any of the others, and dogs diverged from wolves at about the same time as wolves diverged from each other. This infers that the wolf population(s) from which dogs originated has gone extinct and the current wolf diversity from each region represents novel, younger wolf lineages.[1]

The data indicates that the golden jackal and the ancestor of the wolf/dog diverged 400,000 years ago. Dogs and wolves then diverged 14,900 years ago. The 3 wolf populations diverged from each other 13,400 years ago. The dog populations diverged from each other not long after, with the Dingo at 12,800 years ago and divergence between the boxer and the basenji at 12,100 years ago. There was a 16-fold population bottleneck for dogs since divergence.[1]

(Two studies have found evidence of a human population bottleneck (i.e. loss of genetic diversity) in Europe assessed as occurring somewhere between 17,000–43,000 years ago[35] and between 10,000-60,000 years ago[36] that the researchers attribute to a large human migration out of Europe towards Asia. This implies that humans migrated out of Europe at the end of the Last Glacial Maximum and may have taken dogs with them, resulting in a bottleneck being observed in both populations and the spread of dogs across Eurasia.)

There was a 3-fold population decline for the 3 wolf samples since divergence, and appears to have occurred well in advance of direct extermination campaigns by humans and within the timeframe of environmental and biotic changes associated with the ending of the Pleistocene era i.e. changes in climate and prey, including megafaunal extinctions. This indicates that before the divergence of dogs from wolves that there was much more wolf diversity. The results support a recent divergence between dogs and wolves followed by a dramatic reduction in population size.[1]

The AMY2B (Alpha-Amylase 2B) is a gene that codes a protein that assists with the first step in the digestion of dietary starch and glycogen. An expansion of this gene in dogs would enable early dogs to exploit a starch-rich diet as they fed on refuse from agriculture. Data indicated that the wolves and Dingo had just 2 copies of the gene, the Siberian Husky that is associated with hunter-gatherers had just 3-4 copies, whereas the Saluki that is associated with the Fertile Crescent where agriculture originated had 29 copies. The results show that on average, modern dogs have a high copy number of the gene whereas wolves and dingoes do not. The high copy number of AMY2B variants likely already existed as a standing variation in early domestic dogs, but expanded more recently with the development of large agriculturally based civilizations. This suggests that at the beginning of the domestication process, dogs may have been characterized by a more carnivorous diet than their modern-day counterparts, a diet held in common with early hunter-gatherers.[1]

The divergence time between dogs and wolves is estimated to have occurred between 11,000 and 16,000 years ago. Other recent studies place the divergence between 11,000 to 32,000 years ago (depending on the mutation-rate assumptions used), however all of the studies place the divergence prior to the adoption of extensive agriculture by humans. This finding raises questions regarding the hypothesis that the advent of agriculture created a novel niche that was the driving force for dog domestication.[1][37]

The co-senior author of the studies, Robert K Wayne, stated that:
"The common ancestor of dogs and wolves was a large, wolf-like animal that lived between 9,000 and 34,000 years ago. Based on DNA evidence, it lived in Europe."[7]

Archaeological evidence

Archaeology locates the first dog and human remains together in the Gravettian period (i.e. somewhere between 32,000–22,000 BP), however dog-like canid fossils have also been found at Goyet (36,000 BP) and Altai (33,500 BP) in the Aurignacian period.

Years BP Location Finding
200,000 Zhoukoudian cave system, China Small, extinct wolf skulls - Canis lupus variabilis. The skull differs from the typical wolf in much smaller size with a more slender muzzle and noticeably reduced or absent sagittal crest. In addition, the lower border of some Canis lupus variabilis mandibles is "strongly convex as in the dog".[31]:15 More recent researchers have proposed that Canis lupus variabilis may be an ancestor of the dog lineage.[38][39]:7 At the site, the small wolf's remains were in close proximity to Homo erectus pekinensis or Peking man.
36,000 Goyet Cave, Samson River Valley, Belgium Dog-like skull. The skull was found in a side gallery of the cave, together with remains from mammoth, lynx, red deer and large canids. The Goyet skull is very similar in shape to Eliseevich I dogs and to the Epigravettian Mezin 5490 and Mezhirich dog skulls, which are about 18,000 years younger. Palaeolithic artifacts in this system of caves date from the Mousterian, Aurignacian, Gravettian, and Magdalenian, which indicates recurrent occupations of the cave from the Pleniglacial until the Late Glacial.[23] No descendants, genetic classification of species is contentious.[2]
33,500 Razboinichya Cave, Altai Mountains, Russia Dog-like skull, mandibles (both sides) and teeth. Specimen is similar to fully domesticated dogs from Greenland (about 1000 years old), and unlike ancient and modern wolves, and putative dogs from Eliseevichi I site in central Russia.[32] No descendants, genetic classification of species is contentious.[2]
32,500 Western Europe Based on genetic analysis, the estimated date that the line of most modern dogs and the ancient wolf remains found in Kesslerloch Cave, near Switzerland’s northern border with Germany and dated 14,500 BP, parted from the most recent common ancestor - a large, wolf-like canid.[2]
32,000-22,000 Predmostí, Moravia, Czech Republic Three skulls. Predmostí is a Gravettian site (32,000–22,000 BP). The skulls were found in the human burial zone and identified as Palaeolithic dogs, characterized by - compared to wolves - short skulls, short snouts, widepalates and braincases, and even-sized carnassials. For one skull, "a large bone fragment is present between the upper and lower incisors that extends several cm into the mouth cavity. The size, thickness and shape of the fragment suggest that it could be a fragment of a bone of a large mammal, probably from a mammoth. The position of the bone fragment in the mouth and the articulated state of the lower jaw with the skull indicate that this mammoth bone fragment was inserted artificially into the mouth of the dog post-mortem." Wolf skulls were also found at the site. The presence of dogs buried with humans at this Gravettian site corroborates the hypothesis that domestication began long before the Late Glacial.[30][40]
26,000 Chauvet cave, Vallon-Pont-d'Arc, Ardèche region, France 50-metre trail of footprints made by a boy of about ten years of age alongside those of a large canid. The size and position of the canid's shortened middle toe in relation to its pads indicates a dog rather than a wolf. The footprints have been dated by soot deposited from the torch the child was carrying. The cave is famous for its cave paintings.[41]
14,700 Bonn-Oberkassel, Germany Ancient dog mandible. Directly associated with a human double-grave of a 50-year-old man and a 20-25-year-old woman.[42]
13,900 Eliseevich I site, Russian Plain, Russia Two ancient dog skulls. The most complete dog skull was found in a hearth deposit, near a concentration of mammoth skulls. Its braincase has been perforated on the left and right side. Cut marks are present on the zygomatic and frontal bones. With exception of the canines and some premolars, all its teeth are missing. In addition the left and right carnassials were apparently removed by damaging the alveoli. The remains of at least eight mammoth bone complexes and large quantities of worked ivory were discovered at this site.[23]
13,500 approx Mezin, Ukraine Ancient dog skull - Mezin 5490 - as well as ancient wolf specimens found at the site. Dated to the Epigravettian period (17,000-10,000 BP). The Epigravettian Mezin site is well known for its round mammoth bone dwelling.[23]
13,500 approx Mezhirich, Ukraine Ancient dog skull. Dated to the Epigravettian period (17,000-10,000 BP). The Epigravettian Mazhirich site has four mammoth bone dwellings present.[23]
12,000 Bin Mallaha (Eynan) and Hayonim terrace, Israel Three canid finds. A diminutive carnassial and a mandible, and a wolf or dog puppy skeleton buried with a human during the Natufian culture.[43]
9,200 Texas, USA Dog bone fragment. Found in Hinds Cave in southwest Texas. DNA analysis confirms the bone was from a dog whose ancestry was rooted in Eurasia.[44]
7,800 Jiahu site, China Eleven dog internments. Jaihu is a Neolithic site 22 kilometers north of Wuyang in Henan Province.[45]
7,425 Baikal region, Siberia, Russia Dog buried in a human burial ground. Additionally, a human skull was found buried between the legs of a "tundra wolf" dated 8,320 BP (but it does not match any known wolf DNA stored in Genbank). The evidence indicates that as soon as formal cemeteries developed in Baikal, some canids began to receive mortuary treatments that closely paralleled those of humans. One dog was found buried with four red deer canine pendants around its neck dated 5,770 BP. Many burials of dogs continued in this region with the latest finding at 3,760 BP, and they were all buried laying on their right side, and faced towards the east as did their humans. Some were buried with artifacts, e.g., stone blades, birch bark and antler bone.[46]
5,250 Skateholm, Sweden Cemeteries contained dogs among humans. A dog burial with an antler headdress and three flint blades was recovered at one of the sites.[47]

Domestication

Polychrome cave painting of a wolf-like canid painted 17,000 years ago, Font-de-Gaume, France.

Theory

The current theory of dog domestication is based on the comparison between the dog and extant gray wolves, and although it has been claimed that "man domesticated the wolf", one study highlighted a number of inconsistencies with this comparison and proposed that the ancestor of the dog appears more likely to have been a generalist canid and not the specialized gray wolf.[39] Another study proposed that the ancestor of Canis familiaris was the wild "Canis familiaris".[48] A genetic study recently found that the ancestor of the dog is not the extant gray wolf and the fossil remains of the "wolf-like canid" ancestor has yet to be found.[2] (Readers should be aware of this inconsistency with the theory before progressing through this section.)

Domestication of the wolf over time has produced a number of physical or morphological changes. These include: a reduction in overall size; changes in coat coloration and markings; a shorter jaw initially with crowding of the teeth and, later, with the shrinking in size of the teeth; a reduction in brain size and thus in cranial capacity (particularly those areas relating to alertness and sensory processing, necessary in the wild); and the development of a pronounced “stop”, or vertical drop in front of the forehead (brachycephaly). Certain wolf-like behaviors, such as the regurgitation of partially digested food for the young, have also disappeared.[49]

Although experts disagree over the details of domestication, it is agreed that human interaction played a significant role in shaping the subspecies. 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 canid species.[50]:pages95-136

Method

How domestication arose is unknown but the theories proposed include one, or all, of the following:

Free will and powerful emotions

"The dog could have arisen only from animals predisposed to human society by lack of fear, attentiveness, curiosity, necessity, and recognition of advantage gained through collaboration....were not biological Audio-Animatronics born with a preprogrammed response...It is fair, I think, to say that the humans and wolves involved in the conversion were sentient, observant beings constantly making decisions about how they lived and what they did, based on the perceived ability to obtain at a given time and place what the needed to survive and thrive. They were social animals willing, even eager, to join forces with another animal to merge their sense of group with the others' sense and create an expanded super-group that was beneficial to both in multiple ways. They were individual animals and people involved, from our perspective, in a biological and cultural process that involved linking not only their lives but the evolutionary fate of their heirs in ways, we must assume, they could never have imagined. Does this thesis project too much self-awareness into the past? I doubt it. Powerful emotions were in play that many observers today refer as love - boundless, unquestioning love."[51]:40

Cooperation

A genetic study has found a dog-wolf divergence time of greater than 15,000 years ago. An evolutionary scenario consistent with these results is that dog domestication was initiated close to the Last Glacial Maximum when hunter-gathers preyed on megafauna. Conceivably, proto-dogs might have taken advantage of carcasses left on site by early hunters, assisted in the capture of prey, or provided defense from large competing predators at kills.[2]

Access to food/self-domestication

Some researchers argue that those wolves that were more successful at interacting with humans would pass these traits on to their offspring, eventually creating wolves with a greater propensity to be domesticated. The behavioral characteristic called "flight distance" represents how close an animal will allow humans (or anything else it perceives as dangerous) to get before it runs away. Animals with shorter flight distances will linger and feed when humans are close by. This behavioral trait would have been passed on to successive generations, and amplified, creating animals that are increasingly more comfortable around humans. To be able to eat in the presence of human beings is something that wild wolves can't do.[52] The "most social and least fearful" wolves were the ones who were kept around the human living areas, helping to breed those traits that are still recognized in dogs today.[53][54]

Against this proposition, wolves have been scavenging around human living areas, noted in Israel and Italy for centuries, without demonstrating any move towards domestication.[51]

Orphaned wolf-pups

Studies have shown that some wolf pups taken at an early age and reared by humans are easily tamed and socialized,[12]:140 and one study has demonstrated that adult wolves can be socialized.[12]:141
Some researchers propose that humans adopted orphaned wolf pups and breastfed them alongside human babies.[53][55] In Alaska and other northern areas where people still live close to wolves, wolf pups are sometimes captured and some become acceptable as pets or sled dogs, and once these breeding over generations would become more dog-like.[12]:pages55-56

Against this proposition, other researchers attempting to socialize wolf pups after they reach 21 days of age found it very time-consuming and seldom practical or reliable in achieving success.[56]

Human selection

The "farm fox" experiment attempted to reenact how domestication may have occurred.[57] Researchers, working with wild silver foxes selectively bred over 35 generations and 40 years for the sole trait of friendliness to humans, created more dog-like animals. The "domestic elite" foxes are much more friendly to humans and actually seek human attention, but they also show new physical traits that parallel the selection for tameness, even though the physical traits were not originally selected for. They include spotted or black-and-white coats, floppy ears, tails that curl over their backs, the barking vocalization and earlier sexual maturity. It was reported "On average, the domestic foxes respond to sounds two days earlier and open their eyes one day earlier than their non-domesticated cousins. More striking is that their socialization period has greatly increased. 
Instead of developing a fear response at 6 weeks of age, the domesticated foxes don't show it until 9 weeks of age or later. The whimpering and tail wagging is a holdover from puppyhood, as are the foreshortened face and muzzle. Even the new coat colours can be explained by the altered timing of development. One researcher found that the migration of certain melanocytes (which determine colour) was delayed, resulting in a black and white 'star' pattern.
A criticism of this experiment that has been made based on information obtained after its publication is that the definition of "tame" was changed at least once during the experiment, and that some of the foxes that were classified as neither tame nor aggressive also exhibited these changes, indicating that some factor other than human selection for tameness may have been at work during domestication.[51] When humans restrict dog's breeding diversity, another variable also comes into play that may have contributed to the change - inbreeding.[53]:30

Inequality (mathematics)

From Wikipedia, the free encyclopedia https://en.wikipedia.org/wiki/Inequality...