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The mule is a
domestic Domestic may refer to: In the home * Anything relating to the human home or family ** A domestic animal, one that has undergone domestication ** A domestic appliance, or home appliance ** A domestic partnership ** Domestic science, sometimes cal ...
equine
hybrid Hybrid may refer to: Science * Hybrid (biology), an offspring resulting from cross-breeding ** Hybrid grape, grape varieties produced by cross-breeding two ''Vitis'' species ** Hybridity, the property of a hybrid plant which is a union of two diff ...
between a
donkey The donkey or ass is a domesticated equine. It derives from the African wild ass, ''Equus africanus'', and may be classified either as a subspecies thereof, ''Equus africanus asinus'', or as a separate species, ''Equus asinus''. It was domes ...
, and a
horse The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 mi ...
. It is the
offspring In biology, offspring are the young creation of living organisms, produced either by sexual reproduction, sexual or asexual reproduction. Collective offspring may be known as a brood or progeny. This can refer to a set of simultaneous offspring ...
of a male donkey (a
jack Jack may refer to: Places * Jack, Alabama, US, an unincorporated community * Jack, Missouri, US, an unincorporated community * Jack County, Texas, a county in Texas People and fictional characters * Jack (given name), a male given name, incl ...
) and a female horse (a
mare A mare is an adult female horse or other equidae, equine. In most cases, a mare is a female horse over the age of three, and a filly is a female horse three and younger. In Thoroughbred horse racing, a mare is defined as a female horse more th ...
). The horse and the donkey are different species, with different numbers of
chromosome A chromosome is a package of DNA containing part or all of the genetic material of an organism. In most chromosomes, the very long thin DNA fibers are coated with nucleosome-forming packaging proteins; in eukaryotic cells, the most import ...
s; of the two possible first-generation hybrids between them, the mule is easier to obtain and more common than the
hinny A hinny is a domestic equine hybrid, the offspring of a male horse (a stallion) and a female donkey (a jenny). It is the reciprocal cross to the more common mule, which is the product of a male donkey (a jack) and a female horse (a mare). ...
, which is the offspring of a male horse (a
stallion A stallion is an adult male horse that has not been gelded ( castrated). Stallions follow the conformation and phenotype of their breed, but within that standard, the presence of hormones such as testosterone may give stallions a thicker, "cre ...
) and a female donkey (a
jenny Jenny may refer to: * Jenny (given name), a popular feminine name and list of real and fictional people * Jenny (surname), a family name Animals * Jenny (donkey), a female donkey * Jenny (elephant), a female elephant in the German Army in Worl ...
). Mules vary widely in size, and may be of any color seen in horses or donkeys. They are more patient, hardier and longer-lived than horses, and are perceived as less obstinate and more intelligent than donkeys.


Terminology

A female mule is often called a "molly" or "Molly mule," though the correct term is "mare mule." A male mule is called a "john" or "John mule," though the correct term is "horse mule." A young male mule is called a "mule colt," and a young female is called a "mule filly." The donkey used to produce mules is called a "mule jack," and a group of mules is often called a pack, though the words barren and span also apply.


History

Breeding of mules became possible only when the range of the domestic horse, which originated in
Central Asia Central Asia is a region of Asia consisting of Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, and Uzbekistan. The countries as a group are also colloquially referred to as the "-stans" as all have names ending with the Persian language, Pers ...
in about , extended into that of the domestic ass, which originated in north-eastern Africa. This overlap probably occurred in
Anatolia Anatolia (), also known as Asia Minor, is a peninsula in West Asia that makes up the majority of the land area of Turkey. It is the westernmost protrusion of Asia and is geographically bounded by the Mediterranean Sea to the south, the Aegean ...
and
Mesopotamia Mesopotamia is a historical region of West Asia situated within the Tigris–Euphrates river system, in the northern part of the Fertile Crescent. Today, Mesopotamia is known as present-day Iraq and forms the eastern geographic boundary of ...
in Western Asia, and mules were bred there before . The Hittites, a people of Asia Minor known for their horse-riding skills, held mules in higher esteem than their best horses. The price of a mule was three times that of a good horse for the Hittites. Similarly, mules were seven times as valuable as donkeys to the Sumerians. A painting in the
Tomb of Nebamun The lost Tomb of Nebamun was an ancient Egyptian tomb from the Eighteenth Dynasty located in the Theban Necropolis located on the west bank of the Nile at Thebes (present-day Luxor) in Egypt. The tomb was the source of a number of famous decorate ...
at Thebes, dating from approximately , shows a chariot drawn by a pair of animals which have been variously identified as
onager The onager (, ) (''Equus hemionus''), also known as hemione or Asiatic wild ass, is a species of the family Equidae native to Asia. A member of the subgenus ''Asinus'', the onager was Scientific description, described and given its binomial name ...
s, mules or hinnies. Mules were present in Israel and Judah in the time of
King David David (; , "beloved one") was a king of ancient Israel and Judah and the third king of the United Monarchy, according to the Hebrew Bible and Old Testament. The Tel Dan stele, an Aramaic-inscribed stone erected by a king of Aram-Damas ...
. There are many representations of them in
Mesopotamian Mesopotamia is a historical region of West Asia situated within the Tigris–Euphrates river system, in the northern part of the Fertile Crescent. Today, Mesopotamia is known as present-day Iraq and forms the eastern geographic boundary o ...
works of art dating from the first millennium BC. Among the
bas-relief Relief is a sculptural method in which the sculpted pieces remain attached to a solid background of the same material. The term ''relief'' is from the Latin verb , to raise (). To create a sculpture in relief is to give the impression that th ...
s depicting the
Lion Hunt of Ashurbanipal The royal Lion Hunt of Ashurbanipal is shown on a famous group of Assyrian palace reliefs from the North Palace of Nineveh that are now displayed in room 10a of the British Museum. They are widely regarded as "the supreme masterpieces of Assyria ...
from the North Palace of
Nineveh Nineveh ( ; , ''URUNI.NU.A, Ninua''; , ''Nīnəwē''; , ''Nīnawā''; , ''Nīnwē''), was an ancient Assyrian city of Upper Mesopotamia, located in the modern-day city of Mosul (itself built out of the Assyrian town of Mepsila) in northern ...
is a clear and detailed image of two mules loaded with nets for hunting.
Homer Homer (; , ; possibly born ) was an Ancient Greece, Ancient Greek poet who is credited as the author of the ''Iliad'' and the ''Odyssey'', two epic poems that are foundational works of ancient Greek literature. Despite doubts about his autho ...
noted their arrival in Asia Minor in the ''
Iliad The ''Iliad'' (; , ; ) is one of two major Ancient Greek epic poems attributed to Homer. It is one of the oldest extant works of literature still widely read by modern audiences. As with the ''Odyssey'', the poem is divided into 24 books and ...
'' in 800 BC. Mules contributed to the development of the Americas when the Italian explorer for Spain,
Christopher Columbus Christopher Columbus (; between 25 August and 31 October 1451 – 20 May 1506) was an Italians, Italian explorer and navigator from the Republic of Genoa who completed Voyages of Christopher Columbus, four Spanish-based voyages across the At ...
, introduced donkeys and horses to the continent during his expeditions in 1495. Bringing four male and two female donkeys in addition to horses allowed for the production of mules, which assisted the Spanish in their conquest of the continent. The defeat of the
Aztecs The Aztecs ( ) were a Mesoamerican civilization that flourished in central Mexico in the post-classic period from 1300 to 1521. The Aztec people included different ethnic groups of central Mexico, particularly those groups who spoke the ...
, for example, opened up Mexico as a mule breeding ground, and the presence of mules in military operations and watch duties soon became commonplace.
George Washington George Washington (, 1799) was a Founding Fathers of the United States, Founding Father and the first president of the United States, serving from 1789 to 1797. As commander of the Continental Army, Washington led Patriot (American Revoluti ...
bred mules at his
Mount Vernon Mount Vernon is the former residence and plantation of George Washington, a Founding Father, commander of the Continental Army in the Revolutionary War, and the first president of the United States, and his wife, Martha. An American landmar ...
estate in America. Often called the “father of the American mule,” George Washington sent his donkeys to a multitude of American farms to breed with horses and create mules. Washington found that mules work harder and longer than horses at the same tasks. They also need less food and water than horses. At the time, they were not common in the United States, but Washington understood their value, as they were "more docile than donkeys and cheaper to maintain." By the nineteenth century, mules had become favored draught animals on farms and for pulling boats. Their numbers in the US reached 885,000, though they remained more popular in the south than the north. Most notably, the Twenty Mule Teams pulled, often in teams of 20 or more animals, wagonloads of
borax The BORAX Experiments were a series of safety experiments on boiling water nuclear reactors conducted by Argonne National Laboratory in the 1950s and 1960s at the National Reactor Testing Station in eastern Idaho.
out of
Death Valley Death Valley is a desert valley in Eastern California, in the northern Mojave Desert, bordering the Great Basin Desert. It is thought to be the Highest temperature recorded on Earth, hottest place on Earth during summer. Death Valley's Badwat ...
, California from 1883 to 1889. They pulled wagons carrying 10
short tons The short ton (abbreviation: tn or st), also known as the US ton, is a measurement unit equal to . It is commonly used in the United States, where it is known simply as a ton; however, the term is ambiguous, the single word "ton" being variously ...
(9 metric tons) of borax ore during trips to the Borate mines before being replaced by railroads. Mules were used by armies to transport supplies, occasionally as mobile firing platforms for smaller cannons, and to pull heavier field guns with wheels over mountainous trails such as in Afghanistan during the
Second Anglo-Afghan War The Second Anglo-Afghan War (Dari: جنگ دوم افغان و انگلیس, ) was a military conflict fought between the British Raj and the Emirate of Afghanistan from 1878 to 1880, when the latter was ruled by Sher Ali Khan of the Barakzai dy ...
. In the second half of the twentieth century, widespread use of mules declined in industrialised countries. The use of mules for farming and for transportation of agricultural products largely gave way to diesel-powered
tractors A tractor is an engineering vehicle specifically designed to deliver a high tractive effort (or torque) at slow speeds, for the purposes of hauling a Trailer (vehicle), trailer or machinery such as that used in agriculture, mining or constructio ...
and transportation. The first cloned equine was a mule foal,
Idaho Gem The mule Idaho Gem (born May 4, 2003) is the first cloned equine and first cloned mule. Background and cloning Cloning equines has historically posed challenges, as horse oocytes do not mature efficiently ''in vitro'', and equine embryos are ...
, who was
cloned Cloning is the process of producing individual organisms with identical genomes, either by natural or artificial means. In nature, some organisms produce clones through asexual reproduction; this reproduction of an organism by itself without ...
by nuclear transfer of cells from
foetal A fetus or foetus (; : fetuses, foetuses, rarely feti or foeti) is the unborn offspring of a viviparous animal that develops from an embryo. Following the embryonic development, embryonic stage, the fetal stage of development takes place. Pren ...
material, and was born at the
University of Idaho The University of Idaho (U of I, or UIdaho) is a public land-grant research university in Moscow, Idaho, United States. Established in 1889 and opened three years later, it was the state's sole university for 71 years, until 1963. The un ...
in
Moscow, Idaho Moscow ( ) is a city and the county seat of Latah County, Idaho. Located in the North Central Idaho, North Central region of the state along the border with Washington (state), Washington, it had a population of 25,435 at the 2020 United States ...
, United States on May 5, 2003. Neither an equid nor a hybrid animal had been cloned before that time. As of 2018, Mexico had the largest population of mules in the world at 3,287,449.


Characteristics

In general terms, in both mules and hinnies, the foreparts and head of the animal are similar to those of the
sire Sire is an archaic respectful form of address to reigning kings in Europe. In French and other languages it is less archaic and relatively more current. In Belgium, the king is addressed as "Sire..." in both Dutch and French. The words "sire" an ...
(father), while the hindparts and tail tend to resemble those of the
dam A dam is a barrier that stops or restricts the flow of surface water or underground streams. Reservoirs created by dams not only suppress floods but also provide water for activities such as irrigation, human consumption, industrial use, aqua ...
(mother). Mules are generally larger than hinnies, with longer ears and a heavier head like donkeys, while their tails are usually covered with long hair like that of horses. Mules have thin limbs, small narrow hooves, and short manes like donkeys, while their height, shape of neck and body, and uniformity of their coat and teeth are more similar to what is seen in horses. Mules vary widely in size, from small miniature mules under to large and powerful draught mules standing up to at the
withers Withers are the ridge between the shoulder blades of an animal, typically a quadruped. In many species, this ridge is the tallest point of the body. In horses and dogs, it is the standard place to measure the animal's height. In contrast, catt ...
. The median weight range is between about . Mules’ coats may be of any color seen in the horse or donkey. Mules usually display the light points commonly seen in donkeys: pale or mealy areas on the belly and the insides of the thighs, on the muzzle, and around the eyes. They often have
primitive markings Primitive markings are a group of equine coat color, hair coat markings and qualities seen in several equine species, including horses, donkey, donkeys, and asses. In horses, they are associated with primitive breeds, though not limited to such ...
such as dorsal stripes, shoulder stripes, or zebra stripes on the legs. Mules exhibit
hybrid vigor Heterosis, hybrid vigor, or outbreeding enhancement is the improved or increased function of any biological quality in a Hybrid (biology), hybrid offspring. An offspring is heterotic if its trait (biology), traits are enhanced as a result of m ...
.
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English Natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
wrote: "The mule always appears to me a most surprising animal. That a hybrid should possess more reason, memory, obstinacy, social affection, powers of muscular endurance, and length of life, than either of its parents, seems to indicate that art has here outdone nature." Mules inherit from donkeys the traits of intelligence, sure-footedness, toughness, endurance, disposition, and natural cautiousness; from horses, they inherit speed, conformation, and agility. They grow faster and live longer, giving them a larger interval of productivity compared to horses or donkeys. They rarely become ill, unlike horses, who often need to be checked for worms, and like their donkey fathers, mules can see their hind legs and have hooves better suited to dry climates. Additionally, the skin of mules is tougher than that of horses or donkeys, and they tend to live longer on fewer resources than do horses. Mules are reputed to exhibit a higher cognitive intelligence than both of their parent species, but robust scientific evidence to back up these claims is lacking. Preliminary data exists from at least two evidence-based studies, but it relies on a limited set of specialized cognitive tests and a small number of subjects. Similar to other equine animals, mules can sleep while standing (mostly to protect themselves against danger). To prevent predation, a group of mules may select members of the pack to "stand watch" while sleeping upright as the others sleep on the ground.


Fertility

The speciation of horses and donkeys from their common ancestor happened sometime between 7.7 and 15.4 million years ago. Today, they are phenotypically and genetically different, with a horse having 64 chromosomes and a donkey having 62. A mule has 63
chromosomes A chromosome is a package of DNA containing part or all of the genetic material of an organism. In most chromosomes, the very long thin DNA fibers are coated with nucleosome-forming packaging proteins; in eukaryotic cells, the most importa ...
, 32 from the horse and 31 from the donkey. Its odd number of chromosomes makes gamete formation difficult, often leaving mules
infertile In biology, infertility is the inability of a male and female organism to reproduce. It is usually not the natural state of a healthy organism that has reached sexual maturity, so children who have not undergone puberty, which is the body's sta ...
. This also disqualifies them as a species under the biological species model. The conception of a mule is difficult due to the differences in behavior and mating patterns between donkeys and horses. When in close proximity, groups of horses and groups of donkeys do not interact with each other often, and donkeys remain at the bottom of the equine social hierarchy while horses rule the pasture and mules are left in a mid-tier social caste. Mule pregnancy is rare, but can occasionally occur naturally, as well as through
embryo transfer Embryo transfer refers to a step in the process of assisted reproduction in which embryos are placed into the uterus of a female with the intent to establish a pregnancy. This technique - which is often used in connection with in vitro fertili ...
. A few mare mules have produced offspring when mated with a horse or a jack.
Herodotus Herodotus (; BC) was a Greek historian and geographer from the Greek city of Halicarnassus (now Bodrum, Turkey), under Persian control in the 5th century BC, and a later citizen of Thurii in modern Calabria, Italy. He wrote the '' Histori ...
gives an account of such an event as an ill omen of Xerxes' invasion of Greece in 480 BC: "There happened also a portent of another kind while he was still at Sardis—a mule brought forth young and gave birth to a mule" (Herodotus '' The Histories'' 7:57), and a mule's giving birth was a frequently recorded portent in antiquity, although scientific writers also doubted whether it was really possible (see e.g.
Aristotle Aristotle (; 384–322 BC) was an Ancient Greek philosophy, Ancient Greek philosopher and polymath. His writings cover a broad range of subjects spanning the natural sciences, philosophy, linguistics, economics, politics, psychology, a ...
, ''Historia animalium'', 6.24;
Varro Marcus Terentius Varro (116–27 BCE) was a Roman polymath and a prolific author. He is regarded as ancient Rome's greatest scholar, and was described by Petrarch as "the third great light of Rome" (after Virgil and Cicero). He is sometimes call ...
, ''De re rustica'', 2.1.28). Between 1527 and 2002, approximately sixty such births were reported. In
Morocco Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines on the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to Algeria–Morocc ...
in early 2002 and
Colorado Colorado is a U.S. state, state in the Western United States. It is one of the Mountain states, sharing the Four Corners region with Arizona, New Mexico, and Utah. It is also bordered by Wyoming to the north, Nebraska to the northeast, Kansas ...
in 2007, mare mules produced colts. Blood and hair samples from the Colorado birth verified that the mother was indeed a mule and the foal was indeed her offspring. A 1939 article in the ''Journal of Heredity'' describes two offspring of a fertile mare mule named "Old Bec," which was owned at the time by
Texas A&M University Texas A&M University (Texas A&M, A&M, TA&M, or TAMU) is a public university, public, Land-grant university, land-grant, research university in College Station, Texas, United States. It was founded in 1876 and became the flagship institution of ...
in the late 1920s. One of the foals was a female, sired by a jack. Unlike her mother, she was sterile. The other, sired by a five-gaited
Saddlebred The American Saddlebred is a horse breed from the United States. Descended from riding-type horses bred at the time of the American Revolution, the American Saddlebred includes the Narragansett Pacer, Canadian Pacer, Morgan and Thoroughb ...
stallion, exhibited no characteristics of any donkey. That horse, a stallion, was bred to several mares, which gave birth to live foals that showed no characteristics of the donkey. In 1995, a group from the
Federal University of Minas Gerais The Federal University of Minas Gerais (, UFMG) is a federalIn the Brazilian Higher Education context, ''Federal'' does not mean ''collegiate'' (even though most Federal Universities in Brazil enjoy a similarly collegiate system), but it means ...
described a female mule that was pregnant for a seventh time, having previously produced two donkey sires, two foals with the typical 63 chromosomes of mules, and several horse stallions that had produced four foals. The three of the latter available for testing each bore 64 horse-like chromosomes. These foals phenotypically resembled horses, though they bore markings absent from the sire's known lineages, and one had ears noticeably longer than those typical of her sire's breed. The elder two horse-like foals had proved fertile at the time of publication, with their progeny being typical of horses.


Use

While a few mules can carry live weight up to , mules can generally be packed with dead weight up to around 90 kg (198 lb). Although it depends on the individual animal, mules trained by the Army of Pakistan are reported to be able to carry up to and walk without resting. Mules have uses beyond heavy lifting. In the Abruzzo region of Italy, for example, mules are used to defend herds of animals against predators like wolves and feral dogs. Horse breeders in Italy used to add a female mule to their flocks, finding that the mule tended to protect the foals with even more vigor than the actual mother. Mule trains are also used to deliver food to remote areas of the world. In Nepal, for example, the World Food Programme provides food for children in the Sudurpaschim province with the help of mule drivers. The most recent use for the mule is as a research specimen, especially in studies regarding the slicing, rearrangement, and compatibility of chromosomes. File:Wagon mules, Talkeetna, Alaska.jpg, Pair, pulling a wagon (2008) File:20151016-ESP A.MARKUS CJOPCOMMTF-08 (22338729192).jpg, Army mule (2015) File:Arando con mula en Candeleda.jpg, Plowing (2012) File:Mule train.jpg, Supply train in the
Grand Canyon The Grand Canyon is a steep-sided canyon carved by the Colorado River in Arizona, United States. The Grand Canyon is long, up to wide and attains a depth of over a mile (). The canyon and adjacent rim are contained within Grand Canyon Nati ...
(2009) File:SupaiUSMailMules.jpg, Carrying the mail in rural Arizona (2008)


Care and management

A mule's diet is more similar to that of a donkey than a horse. They store water more efficiently, for example, and can consume 15 gallons of it daily, making them suitable for more desert-like climates. Their diet is also herbivorous, mainly consisting of grains, hay and greens, though they can also enjoy fruits and vegetables. Mules can have dietary preferences based on taste and texture. Additionally, protein intake can be an issue with mules, so monitoring the amount of essential amino acids in their diet can be helpful. A mule's hooves should be cleaned regularly to remove debris and should be shortened at least every two months to prevent overgrowth, pain, and discomfort. If necessary, mules may also wear muleshoes to protect their hooves, and generally wear a smaller
horseshoe A horseshoe is a product designed to protect a horse hoof from wear. Shoes are attached on the palmar surface (ground side) of the hooves, usually nailed through the insensitive hoof wall that is anatomically akin to the human toenail, altho ...
than a horse of the same size, due to their smaller and narrower hooves. A mule's coat needs to be groomed regularly. An adult mule's temperature should remain within the range of 37.5°C – 38.5°C, their pulse has a healthy range of 26-40 bpm, and they should take 8-16 breaths per minute.


References


Further reading

* * * * * * * Renner, G. K. "The Mule in Missouri Agriculture, 1821-1950." ''Missouri Historical Review'' 74 (July 1980): 433–457
online
* * Hosted by the
UNT Digital Library The University of North Texas Libraries is an American academic research library system that serves the constituent colleges and schools of University of North Texas in Denton. The phrase "University of North Texas Libraries" encompasses thre ...
. Originally published by the
U.S. Government Printing Office The United States Government Publishing Office (USGPO or GPO), formerly the United States Government Printing Office, is an agency of the legislative branch of the United States federal government. The office produces and distributes informatio ...
.


External links

* * * {{Authority control Equid hybrids Livestock Livestock guardians Pack animals