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Treaty on the Functioning of the European Union
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Consolidated version of the Treaty on the Functioning of the European Union
The Treaty on the Functioning of the European Union (2007) is one of two primary Treaties of the European Union, alongside the Treaty on European Union (TEU). Originating as the Treaty of Rome, the TFEU forms the detailed basis of European Union law, by setting out the scope of the EU's authority to legislate and the principles of law in those areas where EU law operates.
Contents
[hide]
1 History
2 Contents
2.1 Part 1, Principles
2.2 Part 2, Non-discrimination and citizenship of the Union
2.3 Part 3, Union policies and internal actions
2.3.1 Title I: Internal market
2.3.2 Title II: Free movement of goods
2.3.3 Title III: Agriculture and Fisheries
2.3.4 Title IV: Free movement of workers, services and capital
2.3.5 Title V: Area of freedom, justice and security
2.3.6 Title VI: Transport
2.3.7 Title VII: Common Rules on Competition, Taxation and Approximation of Laws
2.3.8 Title VIII: Economic and monetary policy
2.3.9 Titles IX to XV: Employment, social and consumer policy
2.3.10 Titles XVI to XXIV Networks, industry, environment, energy, other
2.4 Part 4, Association of the overseas countries and territories
2.5 Part 5, External action by the Union
2.6 Part 6, Institutional and financial provisions
2.7 Part 7, General and final provisions
3 See also
4 References
History[edit]
The TFEU originated as the treaty establishing the European Economic Community (the EEC treaty), signed in Rome on 25 March 1957. On 7 February 1992, the Maastricht treaty, which led to the formation of the European Union, saw the EEC Treaty renamed as the Treaty establishing the European Community (TEC) and renumbered. The Maastricht reforms also saw the creation of the European Union's three pillar structure, of which the European Community was the major constituent part.
Following the 2005 referenda, which saw the failed attempt at launching a European Constitution, on 13 December 2007 the Lisbon treaty was signed. This saw the 'TEC' renamed as the Treaty on the Functioning of the European Union (TFEU) and, once again, renumbered. The Lisbon reforms resulted in the merging of the three pillars into the reformed European Union.
Contents[edit]
The TFEU consists of seven parts:
Part 1, Principles[edit]
In principles, article 1 establishes the basis of the treaty and its legal value. Articles 2 to 6 outline the competencies of the EU according to the level of powers accorded in each area. Articles 7 to 14 set out social principles, articles 15 and 16 set out public access to documents and meetings and article 17 states that the EU shall respect the status of religious, philosophical and non-confessional organisations under national law.[1]
Part 2, Non-discrimination and citizenship of the Union[edit]
The second part begins with article 18 which outlaws, within the limitations of the treaties, discrimination on the basis of nationality. Article 19 states the EU will "combat discrimination based on sex, racial or ethnic origin, religion or belief, disability, age or sexual orientation". Articles 20 to 24 establishes EU citizenship and accords rights to it; to free movement, consular protection from other states, vote and stand in local and European elections, right to petition Parliament and the European Ombudsman and to contact and receive a reply from EU institutions in their own language. Article 25 requires the Commission to report on the implementation of these rights every three years.[1]
Part 3, Union policies and internal actions[edit]
Part 3 is the largest in the TFEU. Articles 26 to 197 concern the substantive policies and actions of the EU.
Title I: Internal market[edit]
Title II: Free movement of goods[edit]
Including the customs union
Title III: Agriculture and Fisheries[edit]
Common Agricultural Policy and Common Fisheries Policy
Title IV: Free movement of workers, services and capital[edit]
free movement of people, services and capital
Title V: Area of freedom, justice and security[edit]
Including police and justice co-operation
Title VI: Transport[edit]
Title VII: Common Rules on Competition, Taxation and Approximation of Laws[edit]
European Union competition law, taxation and harmonisation of regulations (note Article 101 and Article 102)
Title VIII: Economic and monetary policy[edit]
Articles 119 to 144 concern economic and monetary policy, including articles on the euro. Chapter 1: Economic policy - Article 126 deals with how excessive member state debt is handled.[2] Chapter 2: Monetary policy - Article 127 outlines that the European System of Central Banks should maintain price stability and work with the principles of an open markets and free competition.[3]
Titles IX to XV: Employment, social and consumer policy[edit]
Main articles: European labour law and European consumer law
Title IX concerns employment policy, under articles 145–150. Title X concerns social policy, and with reference to the European Social Charter 1961 and the Community Charter of the Fundamental Social Rights of Workers 1989. This gives rise to the weight of European labour law.
Title XI establishes the European Social Fund under articles 162–164. Title XII, articles 165 and 166 concern education, vocational training, youth and sport policies. Title XIII concerns culture, in article 167. Title XIV allows measures for public health, under article 168. Title XV empowers the EU to act for consumer protection, in article 169.
Titles XVI to XXIV Networks, industry, environment, energy, other[edit]
Title XVI, articles 170–172 empower action to develop and integrate Trans-European Networks. Title XVII, article 173, regards the EU's industrial policy, to promote industry. Title XVIII, articles 174 to 178 concern economic, social and territorial cohesion (reducing disparities in development). Title XIX concerns research and development and space policy, under which the European Research Area and European Space Policy are developed.
Title XX concerns the increasingly important environmental policy, allowing action under articles 191 to 193. Title XXI, article 194, establishes the Energy policy of the European Union.
Title XXII, article 195 is tourism. Title XXIII, article 196 is civil protection. Title XXIV, article 197 is administrative co-operation.
Part 4, Association of the overseas countries and territories[edit]
Part 4, in articles 198 to 204, deals with association of overseas territories. Article 198 sets the objective of association as promoting the economic and social development of those associated territories as listed in annexe 2. The following articles elaborate on the form of association such as customs duties.[1]
Part 5, External action by the Union[edit]
Part 5, in articles 205 to 222, deals with EU foreign policy. Article 205 states that external actions must be in accordance with the principles laid out in Chapter 1 Title 5 of the Treaty on European Union. Article 206 and 207 establish the common commercial (external trade) policy of the EU. Articles 208 to 214 deal with co-operation on development and humanitarian aid for third countries. Article 215 deals with sanctions while articles 216 to 219 deal with procedures for establishing international treaties with third countries. Article 220 instructs the High Representative and Commission to engage in appropriate co-operation with other international organisations and article 221 establishes the EU delegations. Article 222, the Solidarity clause states that members shall come to the aid of a fellow member who is subject to a terrorist attack, natural disaster or man-made disaster. This includes the use of military force.[1]
Part 6, Institutional and financial provisions[edit]
Part 6, in articles 223 to 334, elaborates on the institutional provisions in the Treaty on European Union. As well as elaborating on the structures, articles 288 to 299 outline the forms of legislative acts and procedures of the EU. Articles 300 to 309 establish the European Economic and Social Committee, the Committee of the Regions and the European Investment Bank. Articles 310 to 325 outline the EU budget. Finally, articles 326 to 334 establishes provision for enhanced co-operation.[1]
Part 7, General and final provisions[edit]
Part 7, in articles 335 to 358, deals with final legal points, such as territorial and temporal application, the seat of institutions (to be decided by member states, but this is enacted by a protocol attached to the treaties), immunities and the effect on treaties signed before 1958 or the date of accession.[1]
See also[edit]
EU law
Treaty on European Union
References[edit]
^ Jump up to: a b c d e f "Treaty on the Functioning of the European Union" (PDF). Eur-lex.europa.eu. Retrieved 17 March 2016.
Jump up ^ "EUR-Lex - 12008E126 - EN - EUR-Lex".
Jump up ^ "EUR-Lex - 12008E127 - EN - EUR-Lex".
[show]
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Treaties of the European Union and related documents
Legal basis
Treaty on the Functioning of the European Union (2007)
Treaty on European Union (2007)
Treaty establishing the European Atomic Energy Community (1957)
Main treaties
Brussels (Western Union, 1948)
Paris (European Coal and Steel Community, 1951)
Modified Brussels (Western European Union, 1954)
Rome (European Economic Community, 1957)
Merger (1965)
Single European Act (1986)
Maastricht (1992)
Amsterdam (1997)
Nice (2001)
Lisbon (2007)
Accession Treaties
1972
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Minor treaties
Netherlands Antilles Association Convention (1962)
First Budgetary Treaty (1970)
Second Budgetary Treaty (1975)
Greenland Treaty (1984)
Minor amendments
Protocol 36 (2011)
Article 136 (2011)
Abandoned treaties and agreements
Treaty establishing the European Defence Community (1952)
Treaty establishing a Constitution for Europe (2004)
UK renegotiation of EU membership (2016)
Declarations
Schuman Declaration (1950)
Declarations at the London and Paris Conferences (1954)
Solemn Declaration (1983)
Petersberg Declaration (1992)
Charter of Fundamental Rights (2000)
Berlin Declaration (2007)
Other documents
Schengen Agreement (1985)
Schengen Convention (1990)
Schengen acquis of the EU (1999)
PFI Convention (2002)
Prüm Convention (2005)
Treaty Establishing the European Stability Mechanism (2012)
European Fiscal Compact (2012)
Agreement on a Unified Patent Court (2013)
Single Resolution Fund Agreement (2014)
European Union Portal
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text:
My Name Is Jack
From Wikipedia, the free encyclopedia
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"My Name Is Jack" is a song written by American record producer John Simon[1] and released as a single by British group Manfred Mann in 1968.[2] Their version reached number 8 on UK record charts.[3]
Contents
[hide]
1 Background
2 Cover versions
3 References
4 External links
Background[edit]
The lyrics and music were written by John Simon, and his own version was included on the soundtrack of the 1968 film You Are What You Eat.[1] The song tells the story of a resident of the " Greta Garbo Home for Wayward Boys and Girls", which was the nickname of a real hostel, the Kirkland Hotel, in San Francisco, where part of the movie was filmed.[4][5][6] Formerly the Kashu Hotel, the building became dilapidated and was demolished; the Christ United Presbyterian Church was opened on the site in 1975.[7]
The song was one of the first high-profile sessions at the newly constructed Trident Studios in London, which would later become renowned for its use by artists like The Beatles, David Bowie, Queen, and others.
Cover versions[edit]
1978 – Moonriders, Nouvelles Vagues[8][better source needed]
1994 – Pizzicato Five, Great White Wonder[9][better source needed]
References[edit]
^ Jump up to: a b "AFI Catalog of Feature Films - You Are What You Eat". American Film Institute. Retrieved 17 July 2014.
Jump up ^ Shinner, Merrill (13 April 1977). "Manfred Mann rewrites the past". St. Petersburg Times. Retrieved 17 July 2014.
Jump up ^ Betts, Graham (2004). Complete UK Hit Singles 1952-2004 (1st ed.). London: Collins. p. 488. ISBN 0-00-717931-6.
Jump up ^ GarboForever.com. Retrieved 17 July 2014
Jump up ^ Mark Christensen, Acid Christ: Ken Kesey, LSD, and the Politics of Ecstasy, IPG, 2010, p.203, Retrieved 17 July 2014
Jump up ^ Greta Garbo Home for Wayward Boys and Girls, discussion at HipForums.com. Retrieved 17 July 2014
Jump up ^ San Francisco's Japantown, Arcadia Publishing, 2005, p.35
Jump up ^ "Nouvelles Vagues", 'Discogs.com. Retrieved 1 December 2015
Jump up ^ "Great White Wonder", 'Discogs.com. Retrieved 1 December 2015
External links[edit]
Lyrics
[show]
v
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Manfred Mann
Manfred Mann
Mike Hugg
Mike Vickers
Tom McGuinness
Paul Jones
Mike d'Abo
Jack Bruce
Klaus Voormann
Dave Richmond
Studio albums
The Five Faces of Manfred Mann
Mann Made
As Is
Up the Junction
Mighty Garvey!
EPs
Manfred Mann's Cock-a-Hoop
Groovin' with Manfred Mann
The One in the Middle
No Living Without Loving
Machines
Instrumental Asylum
As Was
Instrumental Assassination
Compilations
Mann Made Hits
Soul of Mann
What A Mann
Chapter Two: The Best of the Fontana Years
Basic: Original Hits
The Best of Manfred Mann: The Definitive Collection
The Best of the EMI Years
Manfred Mann at Abbey Road, 1963-1966
BBC Sessions
Very Best of Manfred Mann
The Story
Classic Masters
The Evolution of Manfred Mann
Complete Greatest Hits
UK singles
"Why Should We Not"
"Cock-a-Hoop"
"5-4-3-2-1"
"Hubble Bubble (Toil and Trouble)"
"Ha! Ha! Said the Clown"
"Do Wah Diddy Diddy"
"Sha La La"
"Come Tomorrow"
"Oh No, Not My Baby"
"If You Gotta Go, Go Now"
"Pretty Flamingo"
"You Gave Me Somebody to Love"
"Just Like a Woman"
"Semi-Detached Suburban Mr James"
"Ha! Ha! Said the Clown"
"Sweet Pea"
"So Long, Dad"
"Mighty Quinn"
"Theme from "Up The Junction"
"My Name Is Jack"
"Fox on the Run"
"Ragamuffin Man"
Related articles
Discography
Manfred Mann Chapter Three
Manfred Mann's Earth Band
The Manfreds
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Evolution of the horse
From Wikipedia, the free encyclopedia
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This image shows a representative sequence, but should not be construed to represent a "straight-line" evolution of the horse. Reconstruction, left forefoot skeleton (third digit emphasized yellow) and longitudinal section of molars of selected prehistoric horses
Skeletal evolution
The evolution of the horse, a mammal of the family Equidae, occurred over a geologic time scale of 50 million years, transforming the small, dog-sized,[1] forest-dwelling Eohippus into the modern horse. Paleozoologists have been able to piece together a more complete outline of the evolutionary lineage of the modern horse than of any other animal. Much of this evolution took place in North America, where horses originated but became extinct about 10,000 years ago.[2]
The horse belongs to the order Perissodactyla (odd-toed ungulates), the members of which all share hooved feet and an odd number of toes on each foot, as well as mobile upper lips and a similar tooth structure. This means that horses share a common ancestry with tapirs and rhinoceroses. The perissodactyls arose in the late Paleocene, less than 10 million years after the Cretaceous–Paleogene extinction event. This group of animals appears to have been originally specialized for life in tropical forests, but whereas tapirs and, to some extent, rhinoceroses, retained their jungle specializations, modern horses are adapted to life on drier land, in the much harsher climatic conditions of the steppes. Other species of Equus are adapted to a variety of intermediate conditions.
The early ancestors of the modern horse walked on several spread-out toes, an accommodation to life spent walking on the soft, moist grounds of primeval forests. As grass species began to appear and flourish,[citation needed] the equids' diets shifted from foliage to grasses, leading to larger and more durable teeth. At the same time, as the steppes began to appear, the horse's predecessors needed to be capable of greater speeds to outrun predators. This was attained through the lengthening of limbs and the lifting of some toes from the ground in such a way that the weight of the body was gradually placed on one of the longest toes, the third.
Contents
[hide]
1 History of research
2 Before odd-toed ungulates
2.1 Phenacodontidae
3 Eocene and Oligocene: early equids
3.1 Eohippus
3.2 Orohippus
3.3 Epihippus
3.4 Mesohippus
3.5 Miohippus
4 Miocene and Pliocene: true equines
4.1 Kalobatippus
4.2 Parahippus
4.3 Merychippus
4.4 Hipparion
4.5 Pliohippus
4.6 Dinohippus
4.7 Plesippus
5 Modern horses
5.1 Equus
5.2 Genome sequencing
5.3 Pleistocene extinctions
5.4 Return to the Americas
6 Details
6.1 Toes
6.2 Teeth
6.3 Coat color
7 See also
8 Further reading
9 References
10 External links
History of research[edit]
Extinct equids restored to scale. Left to right: Mesohippus, Neohipparion, Eohippus, Equus scotti and Hypohippus.
Wild horses were known since prehistory from central Asia to Europe, with domestic horses and other equids being distributed more widely in the Old World, but no horses or equids of any type were found in the New World when European explorers reached the Americas. When the Spanish colonists brought domestic horses from Europe, beginning in 1493, escaped horses quickly established large feral herds. In the 1760s, the early naturalist Buffon suggested this was an indication of inferiority of the New World fauna, but later reconsidered this idea.[3] William Clark's 1807 expedition to Big Bone Lick found "leg and foot bones of the Horses", which were included with other fossils sent to Thomas Jefferson and evaluated by the anatomist Caspar Wistar, but neither commented on the significance of this find.[4]
The first Old World equid fossil was found in the gypsum quarries in Montmartre, Paris, in the 1820s. The tooth was sent to the Paris Conservatory, where it was identified by Georges Cuvier, who identified it as a browsing equine related to the tapir.[5] His sketch of the entire animal matched later skeletons found at the site.[6]
During the Beagle survey expedition, the young naturalist Charles Darwin had remarkable success with fossil hunting in Patagonia. On 10 October 1833, at Santa Fe, Argentina, he was "filled with astonishment" when he found a horse's tooth in the same stratum as fossil giant armadillos, and wondered if it might have been washed down from a later layer, but concluded this was "not very probable".[7] After the expedition returned in 1836, the anatomist Richard Owen confirmed the tooth was from an extinct species, which he subsequently named Equus curvidens, and remarked, "This evidence of the former existence of a genus, which, as regards South America, had become extinct, and has a second time been introduced into that Continent, is not one of the least interesting fruits of Mr. Darwin's palæontological discoveries."[4][8]
In 1848, a study On the fossil horses of America by Joseph Leidy systematically examined Pleistocene horse fossils from various collections, including that of the Academy of Natural Sciences, and concluded at least two ancient horse species had existed in North America: Equus curvidens and another, which he named Equus americanus. A decade later, however, he found the latter name had already been taken and renamed it Equus complicatus.[3] In the same year, he visited Europe and was introduced by Owen to Darwin.[9]
Restoration of Eurohippus parvulus, a mid- to late Eocene equid of Europe (Museum für Naturkunde, Berlin)
The original sequence of species believed to have evolved into the horse was based on fossils discovered in North America in the 1870s by paleontologist Othniel Charles Marsh. The sequence, from Eohippus to the modern horse (Equus), was popularized by Thomas Huxley and became one of the most widely known examples of a clear evolutionary progression. The horse's evolutionary lineage became a common feature of biology textbooks, and the sequence of transitional fossils was assembled by the American Museum of Natural History into an exhibit that emphasized the gradual, "straight-line" evolution of the horse.
Since then, as the number of equid fossils has increased, the actual evolutionary progression from Eohippus to Equus has been discovered to be much more complex and multibranched than was initially supposed. The straight, direct progression from the former to the latter has been replaced by a more elaborate model with numerous branches in different directions, of which the modern horse is only one of many. George Gaylord Simpson in 1951[10] first recognized that the modern horse was not the "goal" of the entire lineage of equids,[11] but is simply the only genus of the many horse lineages to survive.
Detailed fossil information on the distribution and rate of change of new equid species has also revealed that the progression between species was not as smooth and consistent as was once believed. Although some transitions, such as that of Dinohippus to Equus, were indeed gradual progressions, a number of others, such as that of Epihippus to Mesohippus, were relatively abrupt in geologic time, taking place over only a few million years. Both anagenesis (gradual change in an entire population's gene frequency) and cladogenesis (a population "splitting" into two distinct evolutionary branches) occurred, and many species coexisted with "ancestor" species at various times. The change in equids' traits was also not always a "straight line" from Eohippus to Equus: some traits reversed themselves at various points in the evolution of new equid species, such as size and the presence of facial fossae, and only in retrospect can certain evolutionary trends be recognized.[12]
Before odd-toed ungulates[edit]
Phenacodontidae[edit]
Restoration of Phenacodus
Phenacodontidae is the most recent family in the order Condylarthra believed to be the ancestral to the odd-toed ungulates.[citation needed] It contains the genera Almogaver, Copecion, Ectocion, Eodesmatodon, Meniscotherium, Ordathspidotherium, Phenacodus and Pleuraspidotherium. The family lived from the Early Paleocene to the Middle Eocene in Europe and were about the size of a sheep, with tails making slightly less than half of the length of their bodies and unlike their ancestors, good running skills for eluding predators.[citation needed]
Eocene and Oligocene: early equids[edit]
Eohippus[edit]
Main article: Eohippus
Eohippus appeared in the Ypresian (early Eocene), about 52 mya (million years ago). It was an animal approximately the size of a fox (250–450 mm in height), with a relatively short head and neck and a springy, arched back. It had 44 low-crowned teeth, in the typical arrangement of an omnivorous, browsing mammal: three incisors, one canine, four premolars, and three molars on each side of the jaw. Its molars were uneven, dull, and bumpy, and used primarily for grinding foliage. The cusps of the molars were slightly connected in low crests. Eohippus browsed on soft foliage and fruit, probably scampering between thickets in the mode of a modern muntjac. It had a small brain, and possessed especially small frontal lobes.[12]
Eohippus, with left forefoot (third metacarpal colored) and tooth (a enamel; b dentin; c cement) detailed
Its limbs were long relative to its body, already showing the beginnings of adaptations for running. However, all of the major leg bones were unfused, leaving the legs flexible and rotatable. Its wrist and hock joints were low to the ground. The forelimbs had developed five toes, of which four were equipped with small proto-hooves; the large fifth "toe-thumb" was off the ground. The hind limbs had small hooves on three out of the five toes, while the vestigial first and fifth toes did not touch the ground. Its feet were padded, much like a dog's, but with the small hooves in place of claws.[13]
For a span of about 20 million years, Eohippus thrived with few significant evolutionary changes.[12] The most significant change was in the teeth, which began to adapt to its changing diet, as these early Equidae shifted from a mixed diet of fruits and foliage to one focused increasingly on browsing foods. During the Eocene, an Eohippus species (most likely Eohippus angustidens) branched out into various new types of Equidae. Thousands of complete, fossilized skeletons of these animals have been found in the Eocene layers of North American strata, mainly in the Wind River basin in Wyoming. Similar fossils have also been discovered in Europe, such as Propalaeotherium (which is not considered ancestral to the modern horse).[14]
Orohippus[edit]
Approximately 50 million years ago, in the early-to-middle Eocene, Eohippus smoothly transitioned into Orohippus through a gradual series of changes.[14] Although its name means "mountain horse", Orohippus was not a true horse and did not live in the mountains. It resembled Eohippus in size, but had a slimmer body, an elongated head, slimmer forelimbs, and longer hind legs, all of which are characteristics of a good jumper. Although Orohippus was still pad-footed, the vestigial outer toes of Eohippus were not present in Orohippus; there were four toes on each fore leg, and three on each hind leg.
The most dramatic change between Eohippus and Orohippus was in the teeth: the first of the premolar teeth were dwarfed, the last premolar shifted in shape and function into a molar, and the crests on the teeth became more pronounced. Both of these factors gave the teeth of Orohippus greater grinding ability, suggesting Orohippus ate tougher plant material.
Epihippus[edit]
In the mid-Eocene, about 47 million years ago, Epihippus, a genus which continued the evolutionary trend of increasingly efficient grinding teeth, evolved from Orohippus. Epihippus had five grinding, low-crowned cheek teeth with well-formed crests. A late species of Epihippus, sometimes referred to as Duchesnehippus intermedius, had teeth similar to Oligocene equids, although slightly less developed. Whether Duchesnehippus was a subgenus of Epihippus or a distinct genus is disputed.[citation needed] Epihippus was only 2 feet tall.[citation needed]
Mesohippus[edit]
In the late Eocene and the early stages of the Oligocene epoch (32–24 mya), the climate of North America became drier, and the earliest grasses began to evolve. The forests were yielding to flatlands,[citation needed] home to grasses and various kinds of brush. In a few areas, these plains were covered in sand,[citation needed] creating the type of environment resembling the present-day prairies.
Restoration of Mesohippus
In response to the changing environment, the then-living species of Equidae also began to change. In the late Eocene, they began developing tougher teeth and becoming slightly larger and leggier, allowing for faster running speeds in open areas, and thus for evading predators in nonwooded areas[citation needed]. About 40 mya, Mesohippus ("middle horse") suddenly developed in response to strong new selective pressures to adapt, beginning with the species Mesohippus celer and soon followed by Mesohippus westoni.
In the early Oligocene, Mesohippus was one of the more widespread mammals in North America. It walked on three toes on each of its front and hind feet (the first and fifth toes remained, but were small and not used in walking). The third toe was stronger than the outer ones, and thus more weighted; the fourth front toe was diminished to a vestigial nub. Judging by its longer and slimmer limbs, Mesohippus was an agile animal.
Mesohippus was slightly larger than Epihippus, about 610 mm (24 in) at the shoulder. Its back was less arched, and its face, snout, and neck were somewhat longer. It had significantly larger cerebral hemispheres, and had a small, shallow depression on its skull called a fossa, which in modern horses is quite detailed. The fossa serves as a useful marker for identifying an equine fossil's species. Mesohippus had six grinding "cheek teeth", with a single premolar in front—a trait all descendant Equidae would retain. Mesohippus also had the sharp tooth crests of Epihippus, improving its ability to grind down tough vegetation.
Miohippus[edit]
Around 36 million years ago, soon after the development of Mesohippus, Miohippus ("lesser horse") emerged, the earliest species being Miohippus assiniboiensis. As with Mesohippus, the appearance of Miohippus was relatively abrupt, though a few transitional fossils linking the two genera have been found. Mesohippus was once believed to have anagenetically evolved into Miohippus by a gradual series of progressions, but new evidence has shown its evolution was cladogenetic: a Miohippus population split off from the main genus Mesohippus, coexisted with Mesohippus for around four million years, and then over time came to replace Mesohippus.[15]
Miohippus was significantly larger than its predecessors, and its ankle joints had subtly changed. Its facial fossa was larger and deeper, and it also began to show a variable extra crest in its upper cheek teeth, a trait that became a characteristic feature of equine teeth.
Miohippus ushered in a major new period of diversification in Equidae.[16] While Mesohippus died out in the mid-Oligocene, Miohippus continued to thrive, and in the early Miocene (24–5.3 mya), it began to rapidly diversify and speciate. It branched out into two major groups, one of which adjusted to the life in forests once again, while the other remained suited to life on the prairies.[citation needed]
Miocene and Pliocene: true equines[edit]
Kalobatippus[edit]
Fossil Megahippus mckennai
The forest-suited form was Kalobatippus (or Miohippus intermedius, depending on whether it was a new genus or species), whose second and fourth front toes were long, well-suited travel on the soft forest floors. Kalobatippus probably gave rise to Anchitherium, which travelled to Asia via the Bering Strait land bridge, and from there to Europe.[17] In both North America and Eurasia, larger-bodied genera evolved from Anchitherium: Sinohippus in Eurasia and Hypohippus and Megahippus in North America.[18] Hypohippus became extinct by the late Miocene.[19]
Parahippus[edit]
The Miohippus population that remained on the steppes is believed to be ancestral to Parahippus, a North American animal about the size of a small pony, with a prolonged skull and a facial structure resembling the horses of today. Its third toe was stronger and larger, and carried the main weight of the body. Its four premolars resembled the molar teeth; the first were small and almost nonexistent. The incisor teeth, like those of its predecessors, had a crown (like human incisors); however, the top incisors had a trace of a shallow crease marking the beginning of the core/cup.
Merychippus[edit]
Merychippus, an effective grazer and runner
In the middle of the Miocene epoch, the grazer Merychippus flourished. It had wider molars than its predecessors, which are believed to have been used for crunching the hard grasses of the steppes. The hind legs, which were relatively short, had side toes equipped with small hooves, but they probably only touched the ground when running.[16] Merychippus radiated into at least 19 additional grassland species.
Hipparion[edit]
Protohippus simus
Three lineages within Equidae are believed to be descended from the numerous varieties of Merychippus: Hipparion, Protohippus and Pliohippus. The most different from Merychippus was Hipparion, mainly in the structure of tooth enamel: in comparison with other Equidae, the inside, or tongue side, had a completely isolated parapet. A complete and well-preserved skeleton of the North American Hipparion shows an animal the size of a small pony. They were very slim, rather like antelopes, and were adapted to life on dry prairies. On its slim legs, Hipparion had three toes equipped with small hooves, but the side toes did not touch the ground.
In North America, Hipparion and its relatives (Cormohipparion, Nannippus, Neohipparion, and Pseudhipparion), proliferated into many kinds of equids, at least one of which managed to migrate to Asia and Europe during the Miocene epoch.[20] (European Hipparion differs from American Hipparion in its smaller body size – the best-known discovery of these fossils was near Athens.)
Pliohippus[edit]
Pliohippus pernix
Pliohippus arose from Callippus in the middle Miocene, around 12 mya. It was very similar in appearance to Equus, though it had two long extra toes on both sides of the hoof, externally barely visible as callused stubs. The long and slim limbs of Pliohippus reveal a quick-footed steppe animal.
Until recently, Pliohippus was believed to be the ancestor of present-day horses because of its many anatomical similarities. However, though Pliohippus was clearly a close relative of Equus, its skull had deep facial fossae, whereas Equus had no fossae at all. Additionally, its teeth were strongly curved, unlike the very straight teeth of modern horses. Consequently, it is unlikely to be the ancestor of the modern horse; instead, it is a likely candidate for the ancestor of Astrohippus.[21]
Dinohippus[edit]
Dinohippus was the most common species of Equidae in North America during the late Pliocene. It was originally thought to be monodactyl, but a 1981 fossil find in Nebraska shows some were tridactyl.
Plesippus[edit]
Mounted skeleton of Hagerman horse (Equus simplicidens)
Plesippus is often considered an intermediate stage between Dinohippus and the extant genus, Equus.
The famous fossils found near Hagerman, Idaho were originally thought to be a part of the genus Plesippus. Hagerman Fossil Beds (Idaho) is a Pliocene site, dating to about 3.5 mya. The fossilized remains were originally called Plesippus shoshonensis, but further study by paleontologists determined the fossils represented the oldest remains of the genus Equus.[22] Their estimated average weight was 425 kg, roughly the size of an Arabian horse.
At the end of the Pliocene, the climate in North America began to cool significantly and most of the animals were forced to move south. One population of Plesippus moved across the Bering land bridge into Eurasia around 2.5 mya.[23]
Modern horses[edit]
Equus[edit]
Main article: Equus (genus)
Skull of a giant extinct horse, Equus eisenmannae
The genus Equus, which includes all extant equines, is believed to have evolved from Dinohippus, via the intermediate form Plesippus. One of the oldest species is Equus simplicidens, described as zebra-like with a donkey-shaped head. The oldest fossil to date is ~3.5 million years old from Idaho, USA. The genus appears to have spread quickly into the Old World, with the similarly aged Equus livenzovensis documented from western Europe and Russia.[24]
Molecular phylogenies indicate the most recent common ancestor of all modern equids (members of the genus Equus) lived ~5.6 (3.9–7.8) mya. Direct paleogenomic sequencing of a 700,000-year-old middle Pleistocene horse metapodial bone from Canada implies a more recent 4.07 Myr before present date for the most recent common ancestor (MRCA) within the range of 4.0 to 4.5 Myr BP.[25] The oldest divergencies are the Asian hemiones (subgenus E. (Asinus), including the kulan, onager, and kiang), followed by the African zebras (subgenera E. (Dolichohippus), and E. (Hippotigris)). All other modern forms including the domesticated horse (and many fossil Pliocene and Pleistocene forms) belong to the subgenus E. (Equus) which diverged ~4.8 (3.2–6.5) million years ago.[26]
Pleistocene horse fossils have been assigned to a multitude of species, with over 50 species of equines described from the Pleistocene of North America alone, although the taxonomic validity of most of these has been called into question.[27] Recent genetic work on fossils has found evidence for only three genetically divergent equid lineages in Pleistocene North and South America.[26] These results suggest all North American fossils of caballine-type horses (which also include the domesticated horse and Przewalski's horse of Europe and Asia), as well as South American fossils traditionally placed in the subgenus E. (Amerhippus)[28] belong to the same species: E. ferus. Remains attributed to a variety of species and lumped as New World stilt-legged horses (including E. francisci, E. tau, E. quinni and potentially North American Pleistocene fossils previously attributed to E. cf. hemiones, and E. (Asinus) cf. kiang) probably all belong to a second species endemic to North America, which despite a superficial resemblance to species in the subgenus E. (Asinus) (and hence occasionally referred to as North American ass) is closely related to E. ferus.[26] Surprisingly, the third species, endemic to South America and traditionally referred to as Hippidion, originally believed to be descended from Pliohippus, was shown to be a third species in the genus Equus, closely related to the New World stilt-legged horse.[26] The temporal and regional variation in body size and morphological features within each lineage indicates extraordinary intraspecific plasticity. Such environment-driven adaptative changes would explain why the taxonomic diversity of Pleistocene equids has been overestimated on morphoanatomical grounds.[28]
According to these results, it appears the genus Equus evolved from a Dinohippus-like ancestor ~4–7 mya. It rapidly spread into the Old World and there diversified into the various species of asses and zebras. A North American lineage of the subgenus E. (Equus) evolved into the New World stilt-legged horse (NWSLH). Subsequently, populations of this species entered South America as part of the Great American Interchange shortly after the formation of the Isthmus of Panama, and evolved into the form currently referred to as Hippidion ~2.5 million years ago. Hippidion is thus only distantly related to the morphologically similar Pliohippus, which presumably became extinct during the Miocene. Both the NWSLH and Hippidium show adaptations to dry, barren ground, whereas the shortened legs of Hippidion may have been a response to sloped terrain.[28] In contrast, the geographic origin of the closely related modern E. ferus is not resolved. However, genetic results on extant and fossil material of Pleistocene age indicate two clades, potentially subspecies, one of which had a holarctic distribution spanning from Europe through Asia and across North America and would become the founding stock of the modern domesticated horse.[29][30] The other population appears to have been restricted to North America. However, one or more North American populations of E. ferus entered South America ~1.0–1.5 million years ago, leading to the forms currently known as E. (Amerhippus), which represent an extinct geographic variant or race of E. ferus.
Genome sequencing[edit]
In June 2013, a group of researchers announced that they had sequenced the DNA of a 560–780 thousand year old horse, using material extracted from a leg bone found buried in permafrost in Canada's Yukon territory.[31] Prior to this publication, the oldest nuclear genome that had been successfully sequenced was dated at 110–130 thousand years ago. For comparison, the researchers also sequenced the genomes of a 43,000-year-old Pleistocene horse, a Przewalski's horse, five modern horse breeds, and a donkey.[32] Analysis of differences between these genomes indicated that the last common ancestor of modern horses, donkeys, and zebras existed 4 to 4.5 million years ago.[31] The results also indicated that Przewalski's horse diverged from other modern types of horse about 43,000 years ago, and had never in its evolutionary history been domesticated.[25]
Pleistocene extinctions[edit]
Digs in western Canada have unearthed clear evidence horses existed in North America until about 12,000 years ago.[33] However, all Equidae in North America ultimately became extinct. The causes of this extinction (simultaneous with the extinctions of a variety of other American megafauna) have been a matter of debate. Given the suddenness of the event and because these mammals had been flourishing for millions of years previously, something quite unusual must have happened. The first main hypothesis attributes extinction to climate change. For example, in Alaska, beginning approximately 12,500 years ago, the grasses characteristic of a steppe ecosystem gave way to shrub tundra, which was covered with unpalatable plants.[34][35] The other hypothesis suggests extinction was linked to overexploitation by newly arrived humans of naive prey that were not habituated to their hunting methods. The extinctions were roughly simultaneous with the end of the most recent glacial advance and the appearance of the big game-hunting Clovis culture.[36][37] Several studies have indicated humans probably arrived in Alaska at the same time or shortly before the local extinction of horses.[37][38][39] Additionally, it has been proposed that the steppe-tundra vegetation transition in Beringia may have been a consequence, rather than a cause, of the extinction of megafaunal grazers.[40]
In Eurasia, horse fossils began occurring frequently again in archaeological sites in Kazakhstan and the southern Ukraine about 6,000 years ago.[29] From then on, domesticated horses, as well as the knowledge of capturing, taming, and rearing horses, probably spread relatively quickly, with wild mares from several wild populations being incorporated en route.[30][41]
Return to the Americas[edit]
Horses only returned to the Americas with Christopher Columbus in 1493. These were Iberian horses first brought to Hispaniola and later to Panama, Mexico, Brazil, Peru, Argentina, and, in 1538, Florida.[42] The first horses to return to the main continent were 16 specifically identified horses brought by Hernán Cortés. Subsequent explorers, such as Coronado and De Soto, brought ever-larger numbers, some from Spain and others from breeding establishments set up by the Spanish in the Caribbean. Later, as Spanish missions were founded on the mainland, horses would eventually be lost or stolen, and proliferated into large herds of feral horses that became known as mustangs.[citation needed]
The indigenous peoples of the Americas did not have a specific word for horses, and came to refer to them in various languages as a type of dog or deer (in one case, "elk-dog", in other cases "big dog" or "seven dogs", referring to the weight each animal could pull).[citation needed]
Details[edit]
Toes[edit]
The ancestors of the horse came to walk only on the end of the third toe and both side toes. Skeletal remnants show obvious wear on the back of both sides of metacarpal and metatarsal bones, commonly called the "splint bones". They are the remnants of the second and the fourth toe. Modern horses retain the splint bones; they are often believed to be useless attachments, but they in fact play an important role in supporting the carpal joints (front knees) and even the tarsal joints (hocks).[citation needed]
See also: Limbs of the horse
Teeth[edit]
Throughout the phylogenetic development, the teeth of the horse underwent significant changes. The type of the original omnivorous teeth with short, "bumpy" molars, with which the prime members of the evolutionary line distinguished themselves, gradually changed into the teeth common to herbivorous mammals. They became long (as much as 100 mm), roughly cubical molars equipped with flat grinding surfaces. In conjunction with the teeth, during the horse's evolution, the elongation of the facial part of the skull is apparent, and can also be observed in the backward-set eyeholes. In addition, the relatively short neck of the equine ancestors became longer, with equal elongation of the legs. Finally, the size of the body grew as well.[citation needed]
Further information: Horse teeth
Coat color[edit]
Reconstruction of possible ancestral coat colors[43]
The ancestral coat color of E. ferus was possible a uniform dun, consistent with modern populations of Przewalski's horses. Pre-domestication variants including black and spotted have been inferred from cave wall paintings and confirmed by genomic analysis.[44] Domestication may have also led to more varieties of coat colors.[45]
Further information: Equine coat color
See also[edit]
Evidence of common descent
Horses portal
Paleontology portal
Prehistoric mammals portal
Further reading[edit]
MacFadden, Bruce J (1994). Fossil Horses: Systematics, Paleobiology, and Evolution of the Family Equidae. Cambridge & New York: Cambridge University Press. ISBN 0-521-47708-5. Retrieved 6 June 2010.
References[edit]
Jump up ^ Legendre, Serge (1989). Les communautés de mammifères du Paléogène (Eocène supérieur et Oligocène) d'Europe occidentale : structures, milieux et évolution. München: F. Pfeil. p. 110. ISBN 978-3-923871-35-3.
Jump up ^ Singer, Ben (May 2005). A brief history of the horse in America. Canadian Geographic Magazine. Archived from the original on 2012-01-07. Retrieved 22 December 2017.
^ Jump up to: a b "Ancient American Horses". Joseph Leidy Online Exhibit. Academy of Natural Sciences of Drexel University. Archived from the original on 2012-03-05. Retrieved 2015-09-20.
^ Jump up to: a b "Ancient Horse (Equus cf. E. complicatus)". Academy of Natural Sciences. Archived from the original on 2008-08-29. Retrieved 2018-01-26.
Jump up ^ James Warren Evans (1992). Horse breeding and management.
Jump up ^ Knell, Simon J.; Suzanne Macleod; Sheila E. R. Watson (2007). Museum revolutions: how museums and change and are changed. Routledge. ISBN 978-0-415-44467-5.
Jump up ^ 'Filled with astonishment': an introduction to the St. Fe Notebook,
Barlow, Nora (ed. 1945) Charles Darwin and the voyage of the Beagle. London: Pilot Press, p. 210
Jump up ^ Darwin, C. R. (ed. 1840). Fossil Mammalia Part 1 No. 4 of The zoology of the voyage of H.M.S. Beagle. By Richard Owen. London: Smith Elder and Co. p. 108–109
Jump up ^ Academy of Natural Sciences - Joseph Leidy - Leidy and Darwin
Jump up ^ Simpson, George Gaylord (1951): Horses. Oxford University Press; New Impression edition. ISBN 0-19-500104-4 (1971 reprint)
Jump up ^ The notion of a goal would contradict modern evolutionary synthesis,
^ Jump up to: a b c Hunt, Kathleen (1995). "Horse Evolution". TalkOrigins Archive. Retrieved 2018-01-26. See also downloadable pdf version}
Jump up ^ MacFadden, B. J. (March 18, 2005). "Fossil Horses--Evidence for Evolution". Science. 307 (5716): 1728–1730. doi:10.1126/science.1105458. PMID 15774746.
^ Jump up to: a b MacFadden, B. J. (1976). "Cladistic analysis of primitive equids with notes on other perissodactyls". Syst. Zool. 25 (1): 1–14. doi:10.2307/2412774. JSTOR 2412774.
Jump up ^ Prothero, D. R. and Shubin, N. (1989). "The evolution of Oligocene horses". The Evolution of Perissodactyls (pp. 142–175). New York: Clarendon Press.
^ Jump up to: a b Fossil Horses In Cyberspace. Florida Museum of Natural History and the National Science Foundation.
Jump up ^ MacFadden, B. J. (2001). "Three-toed browsing horse Anchitherium clarencei from the early Miocene (Hemingfordian) Thomas Farm, Florida". Bulletin of the Florida Museum of Natural History. 43 (3): 79–109.
Jump up ^ Salesa, M. J.; Sánchez, I. M. & Morales, J. (2004). "Presence of the Asian horse Sinohippus in the Miocene of Europe" (PDF). Acta Palaeontologica Polonica. 49 (2): 189–196. Archived (PDF) from the original on 2005-12-24.
Jump up ^ Waring, George H (2003). Horse Behavior (2nd ed.). New York: Noyes Publications/William Andrew Publishing. p. 9. ISBN 0-8155-1484-0. Retrieved 6 June 2010.
Jump up ^ MacFadden, B. J. (1984). "Systematics and phylogeny of Hipparion, Neohipparion, Nannippus, and Cormohipparion (Mammalia, Equidae) from the Miocene and Pliocene of the New World". Bulletin of the American Museum of Natural History. 179 (1): 1–195.
Jump up ^ MacFadden, B. J. (1984). "Astrohippus and Dinohippus". J. Vert. Paleon. 4 (2): 273–283. doi:10.1080/02724634.1984.10012009.
Jump up ^ McDonald, G. (March 1993). "Hagerman "Horse" - Equus simplicidens". The Fossil Record. Archived from the original on January 3, 2007.
Jump up ^ Jens Lorenz Franzen (2007). Die Urpferde der Morgenröte. München: Elsevier, Spektrum Akademischer Verlag. ISBN 3-8274-1680-9.
Jump up ^ Azzaroli, A. (1992). "Ascent and decline of monodactyl equids: a case for prehistoric overkill" (PDF). Ann. Zool. Finnici. 28: 151–163.
^ Jump up to: a b Orlando, L.; Ginolhac, A.; Zhang, G.; Froese, D.; Albrechtsen, A.; Stiller, M.; Schubert, M.; Cappellini, E.; Petersen, B.; et al. (4 July 2013). "Recalibrating Equus evolution using the genome sequence of an early Middle Pleistocene horse". Nature. 499 (7456): 74–8. doi:10.1038/nature12323. PMID 23803765.
^ Jump up to: a b c d Weinstock, J.; et al. (2005). "Evolution, systematics, and phylogeography of Pleistocene horses in the New World: a molecular perspective". PLoS Biology. 3 (8): e241. doi:10.1371/journal.pbio.0030241. PMC 1159165  . PMID 15974804. Retrieved 2008-12-19.
Jump up ^ Azzaroli, A. (1998). "The genus Equus in North America". PalaeontographItal. 85: 1–60.
^ Jump up to: a b c Orlando, L.; Male, D.; Alberdi, M. T.; Prado, J. L.; Prieto, A.; Cooper, A.; Hänni, C.; et al. (May 2008). "Ancient DNA Clarifies the Evolutionary History of American Late Pleistocene Equids". Journal of Molecular Evolution. 66 (5): 533–538. doi:10.1007/s00239-008-9100-x. PMID 18398561.
^ Jump up to: a b Vila, C.; et al. (2001). "Widespread Origins of Domestic Horse Lineages" (PDF). Science. 291 (5503): 474–7. doi:10.1126/science.291.5503.474. PMID 11161199. Retrieved 2008-12-19.
^ Jump up to: a b Jansen, T.; et al. (July 2002). "Mitochondrial DNA and the origins of the domestic horse". Proceedings of the National Academy of Sciences. 99 (16): 10905–10910. doi:10.1073/pnas.152330099. PMC 125071  . PMID 12130666. Retrieved 2008-12-19.
^ Jump up to: a b Erika Check Hayden (26 June 2013). "First horses arose 4 million years ago". Nature. doi:10.1038/nature.2013.13261.
Jump up ^ Jane J Lee (26 June 2013). "World's Oldest Genome Sequenced From 700,000-Year-Old Horse DNA". National Geographic.
Jump up ^ Singer, Ben (May 2005). A brief history of the horse in America. Canadian Geographic Magazine. Archived from the original on 19 August 2014. Retrieved 16 October 2009.
Jump up ^ LeQuire, Elise (2004-01-04). "No Grass, No Horse". The Horse, online edition. Retrieved 2009-06-08.
Jump up ^ Guthrie, R. D. (2003-11-13). "Rapid body size decline in Alaskan Pleistocene horses before extinction". Nature. 426 (6963): 169–171. doi:10.1038/nature02098. PMID 14614503. Retrieved 2010-12-30.
Jump up ^ "Ice Age Horses May Have Been Killed Off by Humans" National Geographic News, May 1, 2006.
^ Jump up to: a b Buck, Caitlin E.; Bard, Edouard (2007). "A calendar chronology for Pleistocene mammoth and horse extinction in North America based on Bayesian radiocarbon calibration". Quaternary Science Reviews. 26 (17–18): 2031–2035. doi:10.1016/j.quascirev.2007.06.013.
Jump up ^ Solow, Andrew; Roberts, David; Robbirt, Karen (May 9, 2006). Haynes, C. Vance, ed. "On the Pleistocene extinctions of Alaskan mammoths and horses". Proceedings of the National Academy of Sciences of the United States of America (19 ed.). Proceedings of the National Academy of Sciences of the United States of America. 103 (19): 7351–3. doi:10.1073/pnas.0509480103. PMC 1464344  . PMID 16651534.
Jump up ^ Guthrie, R. D. (2006-05-11). "New carbon dates link climatic change with human colonization and Pleistocene extinctions". Nature. 441 (7090): 207–209. doi:10.1038/nature04604. PMID 16688174. Retrieved 2010-12-30.
Jump up ^ Zimov, S. A.; Chuprynin, V. I.; Oreshko, A. P.; Chapin, F. S.; Reynolds, J. F.; Chapin, M. C. (Nov 1995). "Steppe-tundra transition: a herbivore-driven biome shift at the end of the Pleistocene". The American Naturalist. 146 (5): 765–794. doi:10.1086/285824. JSTOR 2462990.
Jump up ^ Warmuth, V.; Eriksson, A.; Bower, M. A.; Barker, G.; Barrett, E.; Hanks, B. K.; Li, S.; Lomitashvili, D.; Ochir-Goryaeva, M.; Sizonov, G. V.; Soyonov, V.; Manica, A. (2012-05-07). "Reconstructing the origin and spread of horse domestication in the Eurasian steppe". Proceedings of the National Academy of Sciences. 109 (21): 8202–8206. doi:10.1073/pnas.1111122109. PMC 3361400  . PMID 22566639. Retrieved 2012-05-08.
Jump up ^ Luís, Cristina; et al. (2006). "Iberian Origins of New World Horse Breeds". Journal of Heredity. 97 (2): 107–113. doi:10.1093/jhered/esj020. PMID 16489143.
Jump up ^ http://www.pnas.org/content/108/46/18626.full.pdf
Jump up ^ Pruvost, M.; et al. (November 2011). "Genotypes of predomestic horses match phenotypes painted in Paleolithic works of cave art". Proceedings of the National Academy of Sciences. 108 (46): 18626–18630. doi:10.1073/pnas.1108982108. PMC 3219153  . PMID 22065780.
Jump up ^ Hofreiter, Michael; Ludwig, Arne; Pruvost, Melanie; Reissman, Monkia; Benecke, Norbert; Brockmann, Gudrun A; Castanos, Pedro; Cieslak, Michael; Lippold, Sebastian; Llorente, Laura; Malaspinas, Anna-Sapfo; Slatkin, Montgomery (2009). "Coat Color Variation at the Beginning of Horse Domestication". Science. 324 (5926): 485.
External links[edit]
Horse Evolution Over 55 Million Years. Tufts University. January 10, 1998. Retrieved July 11, 2007. : An excerpted collection of images of horse fossils
Evolution of Horses
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