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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 c ...
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 dif ...
between a
donkey The domestic donkey is a hoofed mammal in the family Equidae, the same family as the horse. It derives from the African wild ass, ''Equus africanus'', and may be classified either as a subspecies thereof, ''Equus africanus asinus'', or as a ...
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 million y ...
. It is the
offspring In biology, offspring are the young creation of living organisms, produced either by a single organism or, in the case of sexual reproduction, two organisms. Collective offspring may be known as a brood or progeny in a more general way. This ca ...
of a male donkey (a jack) and a female horse (a
mare A mare is an adult female horse or other 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 than four ...
). The horse and the donkey are different species, with different numbers of
chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
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). The hi ...
, which is the offspring of 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 (gorilla), the oldest gorilla in captivity at the time of ...
) and a male horse (a
stallion A stallion is a 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, "cresty" nec ...
). Mules vary widely in size, and may be of any color. 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 that has
oestrus The estrous cycle (, originally ) is the set of recurring physiological changes that are induced by reproductive hormones in most mammalian therian females. Estrous cycles start after sexual maturity in females and are interrupted by anestrous ...
cycles, and which could thus in theory carry a foetus, is called a "molly" or "Molly mule", though the term is sometimes used to refer to female mules in general. A male mule is properly called a "horse mule", though often called a "john mule", which is the correct term for a
gelded A gelding is a castrated male horse or other equine, such as a pony, donkey or a mule. Castration, as well as the elimination of hormonally driven behavior associated with a stallion, allows a male equine to be calmer and better-behaved, making th ...
mule. A young male mule is called a "mule colt", and a young female is called a "mule filly".


History

Breeding of mules became possible only when the range of the domestic horse, which originated in
Central Asia Central Asia, also known as Middle Asia, is a subregion, region of Asia that stretches from the Caspian Sea in the west to western China and Mongolia in the east, and from Afghanistan and Iran in the south to Russia in the north. It includes t ...
in about , extended into that of the domestic ass, which originated in north-eastern Africa. This overlap probably occurred in
Anatolia Anatolia, tr, Anadolu Yarımadası), and the Anatolian plateau, also known as Asia Minor, is a large peninsula in Western Asia and the westernmost protrusion of the Asian continent. It constitutes the major part of modern-day Turkey. The re ...
and
Mesopotamia Mesopotamia ''Mesopotamíā''; ar, بِلَاد ٱلرَّافِدَيْن or ; syc, ܐܪܡ ܢܗܪ̈ܝܢ, or , ) is a historical region of Western Asia situated within the Tigris–Euphrates river system, in the northern part of the F ...
in Western Asia, and mules were bred there before . 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 decora ...
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'' ), A new species called the kiang (''E. kiang''), a Tibetan relative, was previously considered to be a subspecies of the onager as ''E. hemionus kiang'', but recent molecular studies indicate it to be a distinct ...
s, as mules or as hinnies. Mules were present in
Israel and Judah The history of ancient Israel and Judah begins in the Southern Levant during the Late Bronze Age and Early Iron Age. "Israel" as a people or tribal confederation (see Israelites) appears for the first time in the Merneptah Stele, an inscripti ...
in the time of
King David David (; , "beloved one") (traditional spelling), , ''Dāwūd''; grc-koi, Δαυΐδ, Dauíd; la, Davidus, David; gez , ዳዊት, ''Dawit''; xcl, Դաւիթ, ''Dawitʿ''; cu, Давíдъ, ''Davidŭ''; possibly meaning "beloved one". w ...
. There are many representations of them in
Mesopotamian Mesopotamia ''Mesopotamíā''; ar, بِلَاد ٱلرَّافِدَيْن or ; syc, ܐܪܡ ܢܗܪ̈ܝܢ, or , ) is a historical region of Western Asia situated within the Tigris–Euphrates river system, in the northern part of the F ...
works of art dating from the first millennium BC. Among the
bas-relief Relief is a sculptural method in which the sculpted pieces are bonded to a solid background of the same material. The term ''relief'' is from the Latin verb ''relevo'', to raise. To create a sculpture in relief is to give the impression that the ...
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 Assyrian ...
from the North Palace of
Nineveh Nineveh (; akk, ; Biblical Hebrew: '; ar, نَيْنَوَىٰ '; syr, ܢܝܼܢܘܹܐ, Nīnwē) was an ancient Assyrian city of Upper Mesopotamia, located in the modern-day city of Mosul in northern Iraq. It is located on the eastern ban ...
is a clear and detailed image of two mules loaded with nets for hunting.
Homer Homer (; grc, Ὅμηρος , ''Hómēros'') (born ) was a 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. Homer is considered one of the ...
noted their arrival in Asia Minor in the ''
Iliad The ''Iliad'' (; grc, Ἰλιάς, Iliás, ; "a poem about Ilium") 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 ''Odysse ...
'' in 800 BC. Christopher Columbus brought mules to the New World.
George Washington George Washington (February 22, 1732, 1799) was an American military officer, statesman, and Founding Father who served as the first president of the United States from 1789 to 1797. Appointed by the Continental Congress as commander of th ...
bred mules at his home at Mount Vernon. At the time, they were not common in the United States, but Washington understood their value, as they were "more docile than donkeys and cheap to maintain." In the nineteenth century, they were used in various capacities as draught animals - on farms, especially where clay made the soil slippery and sticky; pulling canal boats; and famously for pulling, often in teams of 20 or more animals, wagonloads of
borax Borax is a salt (ionic compound), a hydrated borate of sodium, with chemical formula often written . It is a colorless crystalline solid, that dissolves in water to make a basic solution. It is commonly available in powder or granular form, ...
out of
Death Valley Death Valley is a desert valley in Eastern California, in the northern Mojave Desert, bordering the Great Basin Desert. During summer, it is the Highest temperature recorded on Earth, hottest place on Earth. Death Valley's Badwater Basin is the ...
, California from 1883 to 1889. The wagons were among the largest ever pulled by draught animals, designed to carry 10
short ton The short ton (symbol tn) is a measurement unit equal to . It is commonly used in the United States, where it is known simply as a ton, although the term is ambiguous, the single word being variously used for short, long, and metric ton. The vari ...
s (9 metric tons) of borax ore at a time. 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, ps, د افغان-انګرېز دويمه جګړه) was a military conflict fought between the British Raj and the Emirate of Afghanistan from 1878 to 1880, when the l ...
. 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 steam-, then 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 or machinery such as that used in agriculture, mining or construction. Most common ...
and lorries. On 5 May 2003
Idaho Gem The mule Idaho Gem (born May 4, 2003) is the first cloned equine and first cloned mule. He is the result of the collaboration of Dr. Gordon Woods and Dr. Dirk Vanderwall of the Northwest Equine Reproduction Laboratory at the University of Idaho a ...
, a mule foal
cloned Cloning is the process of producing individual organisms with identical or virtually identical DNA, either by natural or artificial means. In nature, some organisms produce clones through asexual reproduction. In the field of biotechnology, c ...
by
nuclear transfer Nuclear transfer is a form of cloning. The step involves removing the DNA from an oocyte (unfertilised egg), and injecting the nucleus which contains the DNA to be cloned. In rare instances, the newly constructed cell will divide normally, rep ...
of cells from
foetal A fetus or foetus (; plural fetuses, feti, foetuses, or foeti) is the unborn offspring that develops from an animal embryo. Following embryonic development the fetal stage of development takes place. In human prenatal development, fetal develo ...
material, 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. It is the state's land-grant and primary research university,, and the lead university in the Idaho Space Grant Consortium. The University ...
in
Moscow, Idaho Moscow ( ) is a city in North Central Idaho, United States. Located along the state border with Washington, it had a population of 25,435 at the 2020 census. The county seat and largest city of Latah County, Moscow is the home of the Universit ...
. Neither an equid nor a hybrid animal had been cloned before.


Characteristics

In general terms, in both the mule and the hinny the foreparts and head of the animal are similar to those of the sire, while the hindparts and tail tend to resemble those of the dam. A mule is generally larger than a hinny, with longer ears and a heavier head; the tail is usually covered with long hair like that of its mare mother. A mule has the thin limbs, small narrow hooves and short mane of the donkey, while its height, the shape of the neck and body, the uniformity of its coat and its teeth are more similar to those of the horse. Mules vary widely in size, from small miniature mules under to large and powerful draught mules standing up to at the
withers The withers is the ridge between the shoulder blades of an animal, typically a quadruped. In many species, it is the tallest point of the body. In horses and dogs, it is the standard place to measure the animal's height. In contrast, cattle ar ...
. The median weight range is between about . The coat may be of any colour seen in the horse or in the 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 round the eyes. They often have
primitive markings Primitive markings are a group of hair coat markings and qualities seen in several equine species, including horses, donkeys, and asses. In horses, they are associated with primitive breeds, though not limited to such breeds. The markings are ...
such as dorsal stripe, shoulder stripe or zebra stripes on the legs. The mule exhibits
hybrid vigour Heterosis, hybrid vigor, or outbreeding enhancement is the improved or increased function of any biological quality in a hybrid offspring. An offspring is heterotic if its traits are enhanced as a result of mixing the genetic contributions of ...
. Charles Darwin 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". The mule inherits from its sire the traits of intelligence, sure-footedness, toughness, endurance, disposition, and natural cautiousness. From its dam it inherits speed, conformation, and agility. Mules are reputed to exhibit a higher cognitive intelligence than their parent species, but robust scientific evidence to back up these claims is lacking. Preliminary data exist from at least two evidence-based studies, but they rely on a limited set of specialised cognitive tests and a small number of subjects. Mules are generally taller at the shoulder than donkeys and have better endurance than horses, although a lower top speed. In the early twentieth century the mule was preferred to the horse as a
pack animal A pack animal, also known as a sumpter animal or beast of burden, is an individual or type of working animal used by humans as means of transporting materials by attaching them so their weight bears on the animal's back, in contrast to draft anim ...
– its skin is harder and less sensitive than that of a horse, and it is better able to bear heavy weights.


Fertility

A mule has 63
chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
s, intermediate between the 64 of the horse and the 62 of the donkey. Mules are usually
infertile Infertility is the inability of a person, animal or plant to reproduce by natural means. It is usually not the natural state of a healthy adult, except notably among certain eusocial species (mostly haplodiploid insects). It is the normal state ...
for this reason. 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 fertilizati ...
. A few mare mules have produced offspring when mated with a horse or donkey stallion.
Herodotus Herodotus ( ; grc, , }; BC) was an ancient Greek historian and geographer from the Greek city of Halicarnassus, part of the Persian Empire (now Bodrum, Turkey) and a later citizen of Thurii in modern Calabria ( Italy). He is known f ...
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 (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of phil ...
, ''Historia animalium'', 6.24;
Varro Marcus Terentius Varro (; 116–27 BC) 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 Vergil and Cicero). He is sometimes calle ...
, ''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 the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
in early 2002 and
Colorado Colorado (, other variants) is a state in the Mountain West subregion of the Western United States. It encompasses most of the Southern Rocky Mountains, as well as the northeastern portion of the Colorado Plateau and the western edge of t ...
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, or TAMU) is a public, land-grant, research university in College Station, Texas. It was founded in 1876 and became the flagship institution of the Texas A&M University System in 1948. As of late 2021, T ...
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. This breed is referred to as the "Horse America Made". Descended from riding-type horses bred at the time of the American Revolution, the American Saddlebred includes the Nar ...
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 a more recent instance, a group from the
Federal University of Minas Gerais The Federal University of Minas Gerais ( pt, Universidade Federal de 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 sim ...
in 1995 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 , the superiority of the mule becomes apparent in their additional endurance. In general, a mule can be packed with dead weight up to 20% of its body weight, or around . Although it depends on the individual animal, mules trained by the
Army of Pakistan The Pakistan Army (, ) is the land service branch of the Pakistan Armed Forces. The roots of its modern existence trace back to the British Indian Army that ceased to exist following the Partition of British India, which occurred as a result ...
are reported to be able to carry up to and walk without resting. The average equine in general can carry up to roughly 30% of its body weight in live weight, such as a rider. About 3.5 million donkeys and mules are slaughtered each year for meat worldwide. Mule trains have been part of working portions of transportation links as recently as 2005 by the
World Food Programme The World Food Programme; it, Programma alimentare mondiale; es, Programa Mundial de Alimentos; ar, برنامج الأغذية العالمي, translit=barnamaj al'aghdhiat alealami; russian: Всемирная продовольствен ...
, and are still used extensively to transport cargo in rugged, roadless regions. The
Food and Agriculture Organization of the United Nations The Food and Agriculture Organization of the United Nations (FAO)french: link=no, Organisation des Nations unies pour l'alimentation et l'agriculture; it, Organizzazione delle Nazioni Unite per l'Alimentazione e l'Agricoltura is an intern ...
reports that China was the top market for mules in 2003, closely followed by Mexico and many Central and South American nations.


Gallery

File:Peter Birmann - The Devil%27s Bridge in the Schöllenen Gorge on the Way across the St. Gotthard Pass with a Mule Train, before 1805 - Google Art Project.jpg, ''The Devil's Bridge'' at
Schöllenen Gorge Schöllenen Gorge (german: Schöllenenschlucht; Schöllenen) is a gorge formed by the upper Reuss in the Swiss canton of Uri between the towns of Göschenen to the north and Andermatt to the south. It provides access to the St Gotthard Pass. Enc ...
, drawing by Peter Birmann File:Mule Train at Quesnel River (cropped).gif, Mule train fording the
Quesnel River The Quesnel River is a major tributary of the Fraser River in the Cariboo District of central British Columbia. It begins at the outflow of Quesnel Lake, at the town of Likely, British Columbia, Likely and flows for about northwest to its conflu ...
, 1868 File:Mule Battery WDL11495 crop.png, Mule battery in the
Second Anglo-Afghan War The Second Anglo-Afghan War (Dari: جنگ دوم افغان و انگلیس, ps, د افغان-انګرېز دويمه جګړه) was a military conflict fought between the British Raj and the Emirate of Afghanistan from 1878 to 1880, when the l ...
, 1879–1880 File:The Second Anglo - Boer War, South Africa 1899 - 1902 Q72044.jpg, British mule train during the
Second Anglo-Boer War The Second Boer War ( af, Tweede Vryheidsoorlog, , 11 October 189931 May 1902), also known as the Boer War, the Anglo–Boer War, or the South African War, was a conflict fought between the British Empire and the two Boer Republics (the South ...
in South Africa File:Cataline's Mule Train at Hazelton (cropped).gif, Mule train in
British Columbia British Columbia (commonly abbreviated as BC) is the westernmost province of Canada, situated between the Pacific Ocean and the Rocky Mountains. It has a diverse geography, with rugged landscapes that include rocky coastlines, sandy beaches, ...
, 1911 File:Transporte de mulas celaya.jpg, Mule cars in
Celaya Celaya (; ) is a city and its surrounding municipalities of Mexico, municipality in the state of Guanajuato, Mexico, located in the southeast quadrant of the state. It is the third most populous city in the state, with a 2005 census population of 3 ...
,
Guanajuato Guanajuato (), officially the Free and Sovereign State of Guanajuato ( es, Estado Libre y Soberano de Guanajuato), is one of the 32 states that make up the Federal Entities of Mexico. It is divided into 46 municipalities and its capital city i ...
, Mexico, 1930s File:USSMexicanID1655 Mule 1944.jpg, Unloading US Army mules in Naples, Italy, in September, 1944 File:Plowing ca 1947.jpg, Ploughing with a mule,
Saluda, South Carolina Saluda is a town in Saluda County, South Carolina, United States, along the Little Saluda River. The population was 3,565 at the 2010 census. It is the county seat of Saluda County. Geography Saluda is located at (34.003131, -81.771691). Accordi ...
, 1947 File:SupaiUSMailMules.jpg, Mules delivering mail,
Supai, Arizona Supai ( yuf-x-hav, Havasuuw) is a census-designated place (CDP) in Coconino County, Arizona, Coconino County, Arizona, United States, within the Grand Canyon. As of the 2010 United States Census, 2010 census, the CDP had a population of 208. Th ...
, 2008 File:Mule train.jpg, On the
South Kaibab trail The South Kaibab Trail is a hiking trail in Grand Canyon National Park, located in the U.S. state of Arizona. Unlike the Bright Angel Trail which also begins at the south rim of the Grand Canyon and leads to the Colorado River, the South Kaibab ...
in the
Grand Canyon The Grand Canyon (, yuf-x-yav, Wi:kaʼi:la, , Southern Paiute language: Paxa’uipi, ) 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 m ...


See also

*
Zebroid A zebroid is the offspring of any cross between a zebra and any other equine to create a hybrid. In most cases, the sire is a zebra stallion. Offspring of a donkey sire and zebra dam are called a donkra and offspring of a horse sire and a zebra ...


References


Further reading

* * * * * * * * Hosted by the UNT Digital Library. 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 information ...
. {{Authority control Equid hybrids Livestock Livestock guardians Pack animals