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Bats are mammals of the order Chiroptera (/kaɪˈrɒptərə/; from the Greek χείρ - cheir, "hand"[1] and πτερόν - pteron, "wing"[2]) whose forelimbs form webbed wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums, and colugos, can only glide for short distances. Bats do not flap their entire forelimbs, as birds do, but instead flap their spread-out digits,[3] which are very long and covered with a thin membrane or patagium.

Bats are the second largest order of mammals (after the rodents), representing about 20% of all classified mammal species worldwide, with about 1,240 bat species divided into two suborders: the less specialized and largely fruit-eating megabats, or flying foxes, and the highly specialized and echolocating microbats.[4] About 70% of bat species are insectivores. Most of the rest are frugivores, or fruit eaters. A few species, such as the fish-eating bat, feed from animals other than insects, with the vampire bats being hematophagous, or feeding on blood.

Bats are present throughout most of the world, with the exception of extremely cold regions. They perform the vital ecological roles of pollinating flowers and dispersing fruit seeds; many tropical plant species depend entirely on bats for the distribution of their seeds. Bats are economically important, as they consume insect pests, reducing the need for pesticides. The smallest bat is the Kitti's hog-nosed bat, measuring 29–34 mm (1.14–1.34 in) in length, 15 cm (5.91 in) across the wings and 2–2.6 g (0.07–0.09 oz) in mass.[5][6] It is also arguably the smallest extant species of mammal, with the Etruscan shrew being the other contender.[7] The largest species of bat are a few species of Pteropus (fruit bats or flying foxes) and the giant golden-crowned flying fox with a weight up to 1.6 kg (4 lb) and wingspan up to 1.7 m (5 ft 7 in).[8]
Contents

    1 Etymology
    2 Classification and evolution
        2.1 Fossil bats
    3 Habitats
    4 Anatomy
        4.1 Wings
        4.2 Other
    5 Echolocation
        5.1 Other senses
    6 Behaviour
        6.1 Reproduction
        6.2 Life expectancy
        6.3 Hunting, feeding, and drinking
    7 Conservation efforts
        7.1 Artificial roosts
        7.2 Nectar-rich plants attracting moths
        7.3 Threats
    8 Bats in human culture
        8.1 Mythology
        8.2 Tongan
        8.3 Heraldry
    9 Depictions in art
        9.1 State symbols
    10 See also
    11 References
    12 Further reading
    13 External links

Etymology

In many languages, the word for "bat" is cognate with the word for "mouse": for example, chauve-souris ("bald-mouse") in French, murciélago ("blind mouse") in Spanish, saguzahar ("old mouse") in Basque, летучая мышь ("flying mouse") in Russian, slijepi miš ("blind mouse") in Bosnian, nahkhiir ("leather mouse") in Estonian, vlermuis (winged mouse) in Afrikaans, from the Dutch word vleermuis (from Middle Dutch "winged mouse").

An older English name for bats is flittermouse, which matches their name in other Germanic languages (for example German Fledermaus and Swedish fladdermus), related to fluttering of wings, as for Latin bakke, a "moth, nocturnal insect".[9]
Classification and evolution
Giant golden-crowned flying fox, Acerodon jubatus

Bats are placental mammals. Bats were formerly thought to have been most closely related to the flying lemurs, treeshrews, and primates,[10] but recent molecular cladistics research indicates that they actually belong to Laurasiatheria, a diverse group also containing Carnivora and Artiodactyla.[11][12]

The two traditionally recognized suborders of bats are:

    Megachiroptera (megabats)
    Microchiroptera (microbats/echolocating bats)

Not all megabats are larger than microbats. The major distinctions between the two suborders are:

    Microbats use echolocation; with the exception of the genus Rousettus, megabats do not.[13]
    Microbats lack the claw at the second finger of the forelimb.[14][15]
    The ears of microbats do not close to form a ring; the edges are separated from each other at the base of the ear.[15]
    Microbats lack underfur; they are either naked or have guard hairs.[citation needed]

Megabats eat fruit, nectar, or pollen. Most microbats eat insects; others may feed on fruit, nectar, pollen, fish, frogs, small mammals, or the blood of animals. Megabats have well-developed visual cortices and show good visual acuity, while microbats rely on echolocation for navigation and finding prey.

The phylogenetic relationships of the different groups of bats have been the subject of much debate. The traditional subdivision between Megachiroptera and Microchiroptera reflects the view that these groups of bats have evolved independently of each other for a long time, from a common ancestor already capable of flight. This hypothesis recognized differences between microbats and megabats and acknowledged that flight has only evolved once in mammals. Most molecular biological evidence supports the view that bats form a single or monophyletic group.[16]

Researchers have proposed alternative views of chiropteran phylogeny and classification, but more research is needed.

In the 1980s, a hypothesis based on morphological evidence was offered that stated the Megachiroptera evolved flight separately from the Microchiroptera. The so-called flying primates theory proposes that, when adaptations to flight are removed, the Megachiroptera are allied to primates by anatomical features not shared with Microchiroptera. One example is that the brains of megabats show a number of advanced characteristics that link them to primates. Although recent genetic studies strongly support the monophyly of bats,[17] debate continues as to the meaning of available genetic and morphological evidence.[18]

Genetic evidence indicates that megabats originated during the early Eocene and should be placed within the four major lines of microbats.[19]

Consequently, two new suborders based on molecular data have been proposed. The new suborder of Yinpterochiroptera includes the Pteropodidae, or megabat family, as well as the families Rhinolophidae, Hipposideridae, Craseonycteridae, Megadermatidae, and Rhinopomatidae[20] The other new suborder, Yangochiroptera, includes all of the remaining families of bats (all of which use laryngeal echolocation). These two new suborders are strongly supported by statistical tests. Teeling (2005) found 100% bootstrap support in all maximum likelihood analyses for the division of Chiroptera into these two modified suborders. This conclusion is further supported by a 15-base-pair deletion in BRCA1 and a seven-base-pair deletion in PLCB4 present in all Yangochiroptera and absent in all Yinpterochiroptera.[20] Perhaps most convincingly, a phylogenomic study by Tsagkogeorga et al (2013) showed that the two new proposed suborders were supported by analyses of thousands of genes.[19]

The chiropteran phylogeny based on molecular evidence is controversial because microbat paraphyly implies that one of two seemingly unlikely hypotheses occurred. The first suggests that laryngeal echolocation evolved twice in Chiroptera, once in Yangochiroptera and once in the rhinolophoids.[21][22] The second proposes that laryngeal echolocation had a single origin in Chiroptera, was subsequently lost in the family Pteropodidae (all megabats), and later evolved as a system of tongue-clicking in the genus Rousettus.[23]
Common pipistrelle, Pipistrellus pipistrellus

Analyses of the sequence of the "vocalization" gene, FoxP2, were inconclusive as to whether laryngeal echolocation was secondarily lost in the pteropodids or independently gained in the echolocating lineages.[24] However, analyses of the "hearing" gene, Prestin, seemed to favor the independent gain in echolocating species rather than a secondary loss in the pteropodids.[25]

In addition to Yinpterochiroptera and Yangochiroptera, the names Pteropodiformes and Vespertilioniformes have also been proposed for these suborders.[26][27] Under this new proposed nomenclature, the suborder Pteropodiformes includes all extant bat families more closely related to the genus Pteropus than the genus Vespertilio, while the suborder Vespertilioniformes includes all extant bat families more closely related to the genus Vespertilio than to the genus Pteropus.

Little fossil evidence is available to help map the evolution of bats, since their small, delicate skeletons do not fossilize very well. However, a Late Cretaceous tooth from South America resembles that of an early microchiropteran bat. Most of the oldest known, definitely identified bat fossils were already very similar to modern microbats. These fossils, Icaronycteris, Archaeonycteris, Palaeochiropteryx and Hassianycteris, are from the early Eocene period, 52.5 million years ago.[16] Archaeopteropus, formerly classified as the earliest known megachiropteran, is now classified as a microchiropteran.

Bats were formerly grouped in the superorder Archonta, along with the treeshrews (Scandentia), colugos (Dermoptera), and the primates, because of the apparent similarities between Megachiroptera and such mammals. Genetic studies have now placed bats in the superorder Laurasiatheria, along with carnivorans, pangolins, odd-toed ungulates, even-toed ungulates, and cetaceans.[28] A recent study by Zhang et al. places Chiroptera as a sister taxon to the clade Perissodactyla (which includes horses and other odd-toed ungulates).[29] However, the first phylogenomic analysis of bats shows that they are not sisters to Perissodactyla, instead they are sisters to a larger group that includes ungulates and carnivores.[19]


   Laurasiatheria      

  

 Eulipotyphla

   Scrotifera      

  

 Chiroptera

   Fereuungulata      
   Ferae      

  

 Pholidota


  

 Carnivora


   Euungulata      

  

 Perissodactyla  


  

 Cetartiodactyla (also called Artiodactyla)





"Chiroptera" from Ernst Haeckel's Kunstformen der Natur, 1904

The traditional classification of bats is:

    Order Chiroptera
        Suborder Megachiroptera (megabats)
            Pteropodidae
        Suborder Microchiroptera (microbats)
            Superfamily Emballonuroidea
                Emballonuridae (Sac-winged or sheath-tailed bats)
            Superfamily Molossoidea
                Molossidae (Free-tailed bats)
            Superfamily Nataloidea
                Furipteridae (Smoky bats)
                Myzopodidae (Sucker-footed bats)
                Natalidae (Funnel-eared bats)
                Thyropteridae (Disk-winged bats)
            Superfamily Noctilionoidea
                Mormoopidae (Ghost-faced or moustached bats)
                Mystacinidae (New Zealand short-tailed bats)
                Noctilionidae (Bulldog bats or fisherman bats)
                Phyllostomidae (Leaf-nosed bats)
            Superfamily Rhinolophoidea
                Megadermatidae (False vampires)
                Nycteridae (Hollow-faced or slit-faced bats)
                Rhinolophidae (Horseshoe bats)
                Hipposideridae (Old World leaf-nosed bats)
            Superfamily Rhinopomatoidea
                Craseonycteridae (Bumblebee bat or Kitti's hog-nosed bat)
                Rhinopomatidae (Mouse-tailed bats)
            Superfamily Vespertilionoidea
                Vespertilionidae (Vesper bats or evening bats)
                Antrozoidae (Pallid bat and Van Gelder's bat)

Megabats primarily eat fruit or nectar. In New Guinea, they are likely to have evolved for some time in the absence of microbats, which has resulted in some smaller megabats of the genus Nyctimene becoming (partly) insectivorous to fill the vacant microbat ecological niche. Furthermore, some evidence indicates that the fruit bat genus Pteralopex from the Solomon Islands, and its close relative Mirimiri from Fiji, have evolved to fill some niches that were open because there are no nonvolant or nonflying mammals on those islands.
Fossil bats

Fossilized remains of bats are few, as they are terrestrial and light-boned. Only an estimated 12% of the bat fossil record is complete at the genus level.[30] Fossil remains of an Eocene bat, Icaronycteris, were found in 1960. Another Eocene bat, Onychonycteris finneyi, was found in the 52-million-year-old Green River Formation in Wyoming, United States, in 2003.[31][32] This intermediate fossil has helped to resolve a long-standing disagreement regarding whether flight or echolocation developed first in bats. The shape of the rib cage, faceted infraspious fossa of the scapula, manus morphology, robust clavicle, and keeled sternum all indicated Onychonycteris was capable of powered flight. However, the well-preserved skeleton showed that the small cochlea of the inner ear did not have the morphology necessary to echolocate. O. finneyi lacked an enlarged orbical apophysis on the malleus, and a stylohyal element with an expanded paddle-like cranial tip—both of which are characteristics linked to echolocation in other prehistoric and extant bat species.[16] Because of these absences, and the presence of characteristics necessary for flight, Onychonycteris provides strong support for the “flight first” hypothesis in the evolution of flight and echolocation in bats.

The appearance and flight movement of bats 52.5 million years ago were different from those of bats today. Onychonycteris had claws on all five of its fingers, whereas modern bats have at most two claws appearing on two digits of each hand. It also had longer hind legs and shorter forearms, similar to climbing mammals that hang under branches, such as sloths and gibbons. This palm-sized bat had short, broad wings, suggesting that it could not fly as fast or as far as later bat species. Instead of flapping its wings continuously while flying, Onychonycteris likely alternated between flaps and glides while in the air.[16] Such physical characteristics suggest that this bat did not fly as much as modern bats do, rather flying from tree to tree and spending most of its waking day climbing or hanging on the branches of trees.[33] The distinctive features noted on the Onychonycteris fossil also support the claim that mammalian flight most likely evolved in arboreal gliders, rather than terrestrial runners. This model of flight development, commonly known as the "trees-down" theory, implies that bats attained powered flight by taking advantage of height and gravity, rather than relying on running speeds fast enough for a ground-level take off.[34]

The mid-Eocene genus Necromantis is one of the earliest examples of bats specialised to hunt vertebrate prey, as well as one of the largest bats of its epoch.
Habitats

Flight has enabled bats to become one of the most widely distributed groups of mammals.[35] Apart from the Arctic, the Antarctic and a few isolated oceanic islands, bats exist all over the world.[36] Bats are found in almost every habitat available on Earth. Different species select different habitats during different seasons, ranging from seasides to mountains and even deserts, but bat habitats have two basic requirements: roosts, where they spend the day or hibernate, and places for foraging. Most temperate species additionally need a relatively warm hibernation shelter. Bat roosts can be found in hollows, crevices, foliage, and even human-made structures, and include "tents" the bats construct by biting leaves.[37]

The United States is home to an estimated 45 to 48 species of bats.[38][39] The three most common species are Myotis lucifugus (little brown bat), Eptesicus fuscus (big brown bat), and Tadarida brasiliensis (Mexican free-tailed bat). The little and the big brown bats are common throughout the northern two-thirds of the country, while the Mexican free-tailed bat is the most common species in the southwest,[40] sometimes even appearing in portions of the Southeast.
Anatomy
Skeleton of a Myotis myotis (greater mouse-eared bat)
Scapulae, spine and ribs of Eptesicus fuscus (big brown bat)
Wings
A preserved fruit bat showing how the skeleton fits inside its skin
Thermographic image of a bat using trapped air as insulation

The finger bones of bats are much more flexible than those of other mammals, owing to their flattened cross-section and to low levels of minerals, such as calcium, near their tips. In 2006, Sears et al. published a study that traces the elongation of manual bat digits, a key feature required for wing development, to the upregulation of bone morphogenetic proteins (Bmps). During embryonic development, the gene controlling Bmp signaling, Bmp2, is subjected to increased expression in bat forelimbs - resulting in the extension of the offspring's manual digits. This crucial genetic alteration helps create the specialized limbs required for volant locomotion.[41] Sears et al. (2006) also studied the relative proportion of bat forelimb digits from several extant species and compared these with a fossil of Lcaronycteris index, an early extinct species from approximately 50 million years ago. The study found no significant differences in relative digit proportion, suggesting that bat wing morphology has been conserved for over 50 million years.[41]

The wings of bats are much thinner and consist of more bones than the wings of birds, allowing bats to maneuver more accurately than the latter, and fly with more lift and less drag.[42] By folding the wings in toward their bodies on the upstroke, they save 35 percent energy during flight.[43] The membranes are also delicate, ripping easily;[44] however, the tissue of the bat's membrane is able to regrow, such that small tears can heal quickly.[44][45] The surface of their wings is equipped with touch-sensitive receptors on small bumps called Merkel cells, also found on human fingertips. These sensitive areas are different in bats, as each bump has a tiny hair in the center,[46] making it even more sensitive and allowing the bat to detect and collect information about the air flowing over its wings, and to fly more efficiently by changing the shape of its wings in response.[46] An additional kind of receptor cell is found in the wing membrane of species that use their wings to catch prey. This receptor cell is sensitive to the stretching of the membrane.[46] The cells are concentrated in areas of the membrane where insects hit the wings when the bats capture them.
Other

The teeth of microbats resemble insectivorans. They are very sharp to bite through the hardened armor of insects or the skin of fruit.

Mammals have one-way valves in their veins to prevent the blood from flowing backwards, but bats also have one-way valves in their arteries.

The tube-lipped nectar bat (Anoura fistulata) has the longest tongue of any mammal relative to its body size. This is beneficial to them in terms of pollination and feeding. Their long, narrow tongues can reach deep into the long cup shape of some flowers. When the tongue retracts, it coils up inside its rib cage.[47]

Bats possess highly adapted lung systems to cope with the pressures of powered-flight. Flight is an energetically taxing aerobic activity and requires large amounts of oxygen to be sustained. In bats, the relative alveolar surface area and pulmonary capillary blood volume are significantly larger than most other small quadrupedal mammals.[48]
Echolocation
Main article: Animal echolocation § Bats
Spectrogram of Pipistrellus pipistrellus (common pipistrelle) bat vocalizations — one echolocation call, followed by series of multiharmonic pulses, forming a social call (here, precisely, an advertisement call, produced by territorial male during the mating season in autumn).[49]
  
Pipistrellus Pulses
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Recording of Pipistrellus pipistrellus bat time-expanded echolocation calls and social call.
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Bat echolocation is a perceptual system where ultrasonic sounds are emitted specifically to produce echoes. By comparing the outgoing pulse with the returning echoes, the brain and auditory nervous system can produce detailed images of the bat's surroundings. This allows bats to detect, localize, and even classify their prey in complete darkness. At 130 decibels in intensity, bat calls are some of the most intense, airborne animal sounds.[50]

To clearly distinguish returning information, bats must be able to separate their calls from the echoes that they receive. Microbats use two distinct approaches.

    Low duty cycle echolocation: Bats can separate their calls and returning echoes by time. Bats that use this approach time their short calls to finish before echoes return. This is important because these bats contract their middle ear muscles when emitting a call, so they can avoid deafening themselves. The time interval between the call and echo allows them to relax these muscles, so they can clearly hear the returning echo.[51] The delay of the returning echoes provides the bat with the ability to estimate the range to their prey.
    High duty cycle echolocation: Bats emit a continuous call and separate pulse and echo in frequency. The ears of these bats are sharply tuned to a specific frequency range. They emit calls outside of this range to avoid self-deafening. They then receive echoes back at the finely tuned frequency range by taking advantage of the Doppler shift of their motion in flight. The Doppler shift of the returning echoes yields information relating to the motion and location of the bat's prey. These bats must deal with changes in the Doppler shift due to changes in their flight speed. They have adapted to change their pulse emission frequency in relation to their flight speed so echoes still return in the optimal hearing range.[52]

The new Yinpterochiroptera and Yangochiroptera classification of bats, supported by molecular evidence, suggests two possibilities for the evolution of echolocation. It may have been gained once in a common ancestor of all bats and was then subsequently lost in the Old World fruit bats, only to be regained in the horseshoe bats, or echolocation evolved independently in both the Yinpterochiroptera and Yangochiroptera lineages.[53]

Two groups of moths exploit a bat sense to echolocate: tiger moths produce ultrasonic signals to warn the bats that they (the moths) are chemically protected or aposematic, other moth species produce signals to jam bat echolocation. Many moth species have a hearing organ called a tympanum, which responds to an incoming bat signal by causing the moth's flight muscles to twitch erratically, sending the moth into random evasive maneuvers.
Plecotus auritus, the brown long-eared bat

In addition to echolocating prey, bat ears are sensitive to the fluttering of moth wings, the sounds produced by tymbalate insects, and the movement of ground-dwelling prey, such as centipedes, earwigs, etc. The complex geometry of ridges on the inner surface of bat ears helps to sharply focus not only echolocation signals, but also to passively listen for any other sound produced by the prey. These ridges can be regarded as the acoustic equivalent of a Fresnel lens, and may be seen in a large variety of unrelated animals, such as the aye-aye, lesser galago, bat-eared fox, mouse lemur, and others.[54][55][56]

By repeated scanning, bats can mentally construct an accurate image of the environment in which they are moving and of their prey item.[57]
Other senses

Although the eyes of most microbat species are small and poorly developed, leading to poor visual acuity, no species is blind.[58] Microbats use vision to navigate, especially for long distances when beyond the range of echolocation,[59] and species that are gleaners—that is, ones that attempt to swoop down from above to ambush insects, like crickets on the ground or moths up a tree,often have eyesight about as good as a rat's. Some species have been shown to be able to detect ultraviolet light and most cave-dwelling species have developed the ability to utilize very dim light. They also have high-quality senses of smell and hearing. Bats hunt at night, reducing competition with birds, minimizing contact with certain predators, and travel large distances (up to 800 km) in their search for food.[3]

Megabat species often have excellent eyesight as good as, if not better than, human vision. This eyesight is, unlike its microbat relations, adapted to both night and daylight vision and enables the bat to have some colour vision whereas the microbat sees in blurred shades of grey.
Behaviour

Most microbats are nocturnal[60] and are active at twilight. A large portion of bats migrate hundreds of kilometres to winter hibernation dens,[61] while some pass into torpor in cold weather, rousing and feeding when warm weather allows for insects to be active.[62] Others retreat to caves for winter and hibernate for six months.[62] Bats rarely fly in rain, as the rain interferes with their echolocation, and they are unable to locate their food.

The social structure of bats varies, with some leading solitary lives and others living in caves colonized by more than a million bats.[63] The fission-fusion social structure is seen among several species of bats. The term "fusion" refers to a large numbers of bats that congregate in one roosting area, and "fission" refers to breaking up and the mixing of subgroups, with individual bats switching roosts with others and often ending up in different trees and with different roostmates.

Studies also show that bats make all kinds of sounds to communicate with others. Scientists in the field have listened to bats and have been able to associate certain sounds with certain behaviours that bats make after the sounds are made.[63]

Insectivores make up 70% of bat species and locate their prey by means of echolocation. Of the remainder, most feed on fruits.[64] Only three species sustain themselves with blood.

Some species even prey on vertebrates. The leaf-nosed bats (Phyllostomidae) of Central America and South America, and the two bulldog bat (Noctilionidae) species feed on fish. At least two species of bat are known to feed on other bats: the spectral bat, also known as the American false vampire bat, and the ghost bat of Australia.[64] One species, the greater noctule bat, catches and eats small birds in the air.

Predators of bats include bat hawks, bat falcons and even spiders.[65]
Reproduction
Newborn common pipistrelle, Pipistrellus pipistrellus
Colony of mouse-eared bats, Myotis myotis

Most bats have a breeding season, which is in the spring for species living in a temperate climate. Bats may have one to three litters in a season, depending on the species and on environmental conditions, such as the availability of food and roost sites. Females generally have one offspring at a time, which could be a result of the mother's need to fly to feed while pregnant. Female bats nurse their young until they are nearly adult size, because a young bat cannot forage on its own until its wings are fully developed.

Female bats use a variety of strategies to control the timing of pregnancy and the birth of young, to make delivery coincide with maximum food ability and other ecological factors. Females of some species have delayed fertilization, in which sperm is stored in the reproductive tract for several months after mating. In many such cases, mating occurs in the fall, and fertilization does not occur until the following spring. Other species exhibit delayed implantation, in which the egg is fertilized after mating, but remains free in the reproductive tract until external conditions become favorable for giving birth and caring for the offspring.

In yet another strategy, fertilization and implantation both occur, but development of the fetus is delayed until favorable conditions prevail, during the delayed development the mother still gives the fertilized egg nutrients, and oxygenated blood to keep it alive. However, this process can go for a long period of time, because of the advanced gas exchange system.[66] All of these adaptations result in the pup being born during a time of high local production of fruit or insects.

At birth, the wings are too small to be used for flight. Young microbats become independent at the age of six to eight weeks, while megabats do not until they are four months old.
Life expectancy

A single bat can live over 20 years, but bat population growth is limited by the slow birth rate.[67] Five species have been recorded living over 30 years in the wild: the brown long-eared bat (Plecotus auritus), little brown bat (Myotis lucifugus), Brandt's bat (Myotis brandti), lesser mouse-eared bat (Myotis blythii) and greater horseshoe bat (Rhinolophus ferrumequinum).[68]
Hunting, feeding, and drinking
File:BAT.young-Tamil Nadu-202.ogvPlay media
A very young bat in Tamil Nadu

Newborn bats rely on the milk from their mothers.[69] When they are a few weeks old, bats are expected to fly and hunt on their own. It is up to them to find and catch their prey, along with satisfying their thirst.[70]
Hunting
This Philippines bat circles the sweet manzanitas fruits of the Aratiles tree (Muntingia).

Most bats are nocturnal creatures. Their daylight hours are spent grooming and sleeping; they hunt during the night. The means by which bats navigate while finding and catching their prey in the dark was unknown until the 1790s, when Lazzaro Spallanzani conducted a series of experiments on a group of hooded and surgically blinded bats.[71] These bats were placed in a room in total darkness, with silk threads strung across the room. Even then, the bats were able to navigate their way through the room. Spallanzani concluded the bats were not using their eyes to fly through complete darkness, but something else.

Spallanzani decided the bats were able to catch and find their prey through the use of their ears. To prove this theory, Spallanzani plugged the ears of the bats in his experiment. To his pleasure, he found that the bats with plugged ears were not able to fly with the same amount of skill and precision as they were able to without their ears plugged. Unfortunately for Spallanzani, the twin concepts of sound waves and acoustics would not be understood for another century and he could not explain why specifically the bats were crashing into walls and the threads that he'd strung up around the room, and because of the methodology Spallanzani used, many of his test subjects died.

It was thus well known through the nineteenth century that the chiropteran ability to navigate had something to do with hearing, but how they accomplish this was not proven conclusively until the 1930s, by Donald R. Griffin, a biology student at Harvard University. Using a locally native species, the little brown bat, he discovered that bats use echolocation to locate and catch their prey. When bats fly, they produce a constant stream of high-pitched sounds. When the sound waves produced by these sounds hit an insect or other animal, the echoes bounce back to the bat, and guide them to the source.[70]
Feeding and diet

The majority of food consumed by bats includes insects, fruits and flower nectar, vertebrates and blood.[72] Almost three-fourths of the world's bats are insect eaters. Bats consume both aerial and ground-dwelling insects. Each bat is typically able to consume one-third of its body weight in insects each night, and several hundred insects in a few hours. This means that a group of a thousand bats could eat four tons of insects each year. If bats were to become extinct, it has been calculated that the insect population would reach an alarmingly high number.[73]
Vitamin C

In a test of 34 bat species from six major families of bats, including major insect- and fruit-eating bat families, all were found to have lost the ability to synthesize vitamin C, and this loss may derive from a common bat ancestor, as a single mutation.[74] However, recent results show that there are at least two species of bat, the frugivorous bat (Rousettus leschenaultii) and insectivorous bat (Hipposideros armiger), that have retained their ability to produce vitamin C.[75] In fact, the whole Chiroptera are in the process of losing the ability to synthesize Vc which most of them have already lost.[76]
Aerial insectivores

Watching a bat catch and eat an insect is difficult. The action is so fast that all one sees is a bat rapidly changing directions, and continuing on its way. Scientist Frederick A. Webster discovered how bats catch their prey. In 1960, Webster developed a high-speed camera that was able to take one thousand pictures per second. These photos revealed the fast and precise way in which bats catch insects.[70] Occasionally, a bat will catch an insect in mid-air with its mouth, and eat it in the air. However, more often than not, a bat will use its tail membrane or wings to scoop up the insect and trap it in a sort of "bug net".[69] Then, the bat will take the insect back to its roost. There, the bat will proceed to eat said insect, often using its tail membrane as a kind of napkin, to prevent its meal from falling to the ground.[72] One common insect prey is Helicoverpa zea, a moth that causes major agricultural damage.[77]
Forage gleaners

These bats typically fly down and grasp their prey off the ground with their teeth, and take it to a nearby perch to eat it. Generally, these bats do not use echolocation to locate their prey. Instead, they rely on the sounds produced by the insects. Some make unique sounds, and almost all make some noise while moving through the environment.[69]
Fruits and flower nectar
A colony of great fruit-eating bats

Fruit eating, or frugivory, is a specific habit found in two families of bats. Megachiropterans and microchiropterans both include species of bat that feed on fruits. These bats feed on the juices of sweet fruits, and fulfill the needs of some seeds to be dispersed. The fruits preferred by most fruit-eating bats are fleshy and sweet, but not particularly strong smelling or colorful.[69] To get the juice of these fruits, bats pull the fruit off the trees with their teeth, and fly back to their roosts with the fruit in their mouths. There, the bats will consume the fruit in a specific way. To do this, the bats crush open the fruit and eat the parts that satisfy their hunger. The remainder of the fruit, the seeds and pulp, are spat onto the ground. These seeds take root and begin to grow into new fruit trees.[73] Over 150 types of plants depend on bats in order to reproduce.[73]

Some bats prefer the nectar of flowers to insects or other animals. These bats have evolved specifically for this purpose. For example, these bats possess long muzzles and long, extensible tongues covered in fine bristles that aid them in feeding on particular flowers and plants.[69] When they sip the nectar from these flowers, pollen gets stuck to their fur, and is dusted off when the bats take flight, thus pollinating the plants below them.[73] The rainforest is said to be the most benefitted of all the biomes where bats live, because of the large variety of appealing plants.[78] Because of their specific eating habits, nectar-feeding bats are more prone to extinction than any other type of bat.[79] However, bats benefit from eating fruits and nectar just as much as from eating insects.[80]
Vertebrates

A small group of carnivorous bats feed on other vertebrates and are considered the top carnivores of the bat world.[69] These bats typically eat a variety of animals, but normally consume frogs, lizards, birds, and sometimes other bats.[73] For example, one vertebrate predator, Trachops cirrhosus, is particularly skilled at catching frogs. These bats locate large groups of frogs by distinguishing their mating calls from other sounds around them. They follow the sounds to the source and pluck them from the surface of the water with their sharp canine teeth.[69] Another example is the greater noctule bat, which is believed to catch birds on the wing.

Also, several species of bat feed on fish. These types of bats are found on almost all continents. They use echolocation to detect tiny ripples in the water's surface to locate fish. From there, the bats swoop down low, inches from the water, and use specially enlarged claws on their hind feet to grab the fish out of the water. The bats then take the fish to a feeding roost and consume the animal.[69]
Blood

A few species of bats exclusively consume blood as their diet. This type of diet is referred to as hematophagy, and three species of bats exhibit this behavior. These species are the common, the white-winged, and the hairy-legged vampire bats. The common vampire bat typically consumes the blood of mammals, while the hairy-legged and white-winged vampires feed on the blood of birds.[81] These species live only in Mexico, Central, and South America, with a presence also on the Island of Trinidad.
Defecation

Bat dung, or guano, is so rich in nutrients that it is mined from caves, bagged, and used by farmers to fertilize their crops. During the U.S. Civil War, guano was used to make gunpowder.[73]

To survive hibernation months, some species build up large reserves of body fat, both as fuel and as insulation.[70]
Drinking

In 1960, Frederic A. Webster discovered bats' method of drinking water using a high-speed camera and flashgun that could take 1,000 photos per second. Webster's camera captured a bat skimming the surface of a body of water, and lowering its jaw to get just one drop of water. It then skimmed again to get a second drop of water, and so on, until it has had its fill. A bat's precision and control during flight is very fine, and it almost never misses.[70] Other bats, such as the flying fox or fruit bat, gently skim the water's surface, then land nearby to lick water from chest fur.[82]
Conservation efforts
See also: List of bats by population

Through conservancy efforts of groups such as the Organization for Bat Conservation and Bat Conservation International, bats are becoming better understood and people beginning to understand the crucial role bats play in insect control and pollination.

In the United Kingdom, all bats are protected under the Wildlife and Countryside Acts, and even disturbing a bat or its roost can be punished with a heavy fine.

In Sarawak, Malaysia, bats are protected species under the Wildlife Protection Ordinance 1998 (see Malaysian Wildlife Law). The large Naked bat (see Mammals of Borneo) and Greater nectar bat are consumed by the local communities.

Bats can be a tourist attraction. The Congress Avenue Bridge in Austin, Texas is the summer home to North America's largest urban bat colony, an estimated 1,500,000 Mexican free-tailed bats, which eat an estimated 10,000 to 30,000 pounds of insects each night. An estimated 100,000 tourists per year visit the bridge at twilight to watch the bats leave the roost.
Artificial roosts
Very large bat house, Tallahassee, Florida, United States
Bat roost with Charles Augustus Rosenheimer Campbell in San Antonio, Texas in 1915

Many people put up bat houses to attract bats just as some put up birdhouses to attract birds. Reasons for this vary, but most revolve around the bats being the primary nocturnal insectivores in most, if not all, ecologies. Bat houses can be made from scratch or from kits, or bought ready made. Plans for bat houses exist on many web sites, as well as guidelines for designing a bat house.[83]

Constructed in March 1991, the University of Florida bat house is the largest occupied artificial roost in the world. The bat house has approximately 300,000 insect-eating residents. These bats can eat upwards of 2.5 billion bugs (2,500 pounds or 1,100 kilograms) per night.

A 1987 fire caused a colony of bats residing in Johnson Hall's attic to become homeless. This forced them to move to the James G. Pressly Stadium on the north side of campus. The odor and the guano from this newly arrived colony did not please spectators, thus furthering the movement for a new bat-ordained structure. In September 1991, thousands of bats were captured and transported to the newly built bat house. In the following evenings, these bats flew away, and found new homes. For three years the bat house remained empty. Finally, in 1995, the bats moved in permanently, and the colony continued to grow. The colony grew so much that, in 2009, part of the original bat house collapsed, and a new "Bat Barn" was constructed next to it. Along with that, the interior of the original was rebuilt.[84]

In Britain, British hardened field defences of World War II have been converted to make roosts for bats. Pillboxes that are well dug-in and thick-walled are naturally damp and provide the stable thermal environment required by bats that would otherwise hibernate in caves. With a few minor modifications, suitable pillboxes can be converted to artificial caves for bats.[85][86]

Again in the UK, purpose-built bat houses are occasionally built when existing roosts are destroyed by developments, such as new roads; one such has been built associated with bat bridges on the new (2008) A38 Dobwalls bypass.[citation needed]
Nectar-rich plants attracting moths
Further information: Biological_pest_control § Conservation

Besides mosquitoes, some species of bats also consume moths. Moths benefit from the availability of nectar-rich plants.[87]
Threats
A little brown bat with white nose syndrome

While conservation efforts are in place to protect bats, many threats still remain. Bats are known to carry diseases communicable to humans, such as rabies.
White nose syndrome
Main article: White nose syndrome

White nose syndrome is a condition associated with the deaths of millions of bats in the Eastern United States and Canada.[88] The disease is named after a white fungus, Pseudogymnoascus destructans, found growing on the muzzles, ears, and wings of afflicted bats. This fungus, which is mostly spread from bat to bat, is the sole cause of the disease.[89] The fungus was first discovered in central New York State in 2006 and spread quickly to the entire Eastern US north of Florida; mortality rates of 90–100% have been observed in most caves.[90] New England and the mid-Atlantic states have, since 2006, witnessed entire species completely extirpated and others with numbers that have gone from the hundreds of thousands, even millions, to a few hundred or less.[91] As of June 2014 this fungus is doing a large amount of damage to the populations of bats that use the caves of the Appalachian Mountains in the Southeastern United States, including the Mammoth Cave system in Kentucky, which was one of the largest hibernacula in the East as recently as 2010, able to hold over a million bats. In Canada, the provinces of Quebec, Ontario, and the Maritimes [92] have witnessed identical die offs, with the Canadian government making preparations to protect the remaining bat populations in Manitoba, Alberta, Saskatchewan, and British Columbia, since scientific evidence suggests that longer winters where the fungus has a longer period of time to infect bats is a threat to Canadian bats.[93]

Scientists in both countries are alarmed that the lack of bug-eating bats in summer could cause severe problems in the ecosystem of the Eastern forests and on farmland and beyond: nearly all of the bats Continental US and Canada are insectivores, and save the government billions in pesticides because they are natural predators of bugs that eat crops like cabbages, potatoes, pears, cherries, peaches, apples, wheat, and corn, consuming tons of them from late May-early October which coincides with the height of planting and harvest. They are also facing the possibility that once common species with robust numbers could go extinct given the rapid progress the disease has made in less than a decade. There is evidence that at least one species, the little brown bat, has started to change its roosting behavior, which scientists hope may yet save the species from extinction[94] Unfortunately, this does not negate the fact that their numbers have undergone a very steep crash. They were up until 2008 the most common bat in the East, and were often seen catching moths attracted to street lamps in New York City, Montreal, and the average small town; they were once so common in the average town that citizens often discovered them roosting in maternity colonies in late spring in their attics. As of 2014, scientists are reckoning that unless something changes soon, the subspecies myotis lucifugus lucifugus may face total population collapse since the population of survivors is dangerously low in an area nearly the size of France.[95] In July 2014 the US state of Maine recorded a population of this subspecies that is now so low that it and the northern long eared bat are candidates for the state endangered species list, after both being plentiful only seven years earlier.[96]

The most recent scholarship suggests that the pseudogymnoascus fungus is native to Europe where it is part of the naturally occurring biota found on the walls of caves, and is harmless to the myotis, bentwing, pipistrelle, and horseshoe bat species that live there. However, as a novel pathogen, it has proven to be lethal to North American bat species that utilise caves or abandoned mines. Unlike European species, most cave dwelling North American bat species form very large winter colonies that can number from a few hundred to many thousands, with bats forming tight clusters on the walls and ceilings of caves and mines. Most North American species shall even migrate many hundreds of miles in September from other states to a very specific type of cave or mine that has a constant temperature of 7.2 °C to 10 °C. They cluster together to conserve body heat and counteract the (compared to most of Europe) colder winter temperatures that may blow in from outer portions of the hibernaculum. Without the natural immunities of their European cousins, these North American bats go into hibernation in autumn a few weeks before Halloween, often forming huge swarms of males and females outside the cave or mine entrance that join up to mate before going off to sleep until spring comes in late March. Normally, they will have been eating moths, beetles, mosquitoes, potato bugs, and flies all summer long and have gained weight to prepare for winter. However, once they enter torpor, their immune system response is suppressed to conserve energy and hibernate through winter. Eventually the bat becomes infected by the fungus, the fungus desiccates the bat's wings, upsetting hydrostatic balance, and the bat, chronically thirsty, is forced to wake up more often than it must, using up its fat reserves, and eventually dying from dehydration and starvation. By then it will have infected several more of its neighbours in the cluster, over time causing hundreds or even thousands to die in a single winter. This number includes what would have been the following season’s offspring, dead with the pregnant mothers, and has a knock on effect on the populations in other states and provinces that never see the return of migratory bats in spring. Aeolus Cave in New England had a headcount numbering 300,000 in 2008. When scientists went to make their count in the spring of 2014, more than 270,000 had already perished and the floors of the caves were covered up to the ankles of researchers with the bones of thousands of bats.[97]

Species thus far that have been badly affected include the little brown bat, the endangered gray myotis, the endangered Indiana bat, the northern long-eared myotis, the eastern small-footed myotis, the tri-coloured bat, and two subspecies of Townsend's big-eared bat. More than half of all bat species that live in North America use caves at some point in their life cycle; in the East, though infected specimens have been caught, only tree dwelling bats of the Lasiurus family and Eptesicus fuscus, a relative of the serotine bat, seem to have been minimally affected: these species are far less dependent on caves for their winter survival and the last mentioned species does not cluster tightly if it is underground at all and will also accept manmade structures to hibernate in. As of 2014, the geographical spread of white nose syndrome in North American bats has worsened in the sense that infection has crossed the Mississippi River:[98] Species native to northern Mexico and the West have not yet been affected, but it remains unknown how or if white nose syndrome shall affect bat species here.[99] All North American bat species have a very slow reproductive rate: with the exception of a handful of species like the little brown bat, which occasionally gives birth to twins, most females give birth to only one pup a year. Though measures are being taken to slow the spread of the disease and a cure is being sought, the recovery of bats In North America may take decades if not centuries due to this fungus and the prognosis for the future of bats is grim.[citation needed]
Barotrauma and wind turbines

Evidence suggests that barotrauma is causing bat fatalities around wind farms.[100] The lungs of bats are typical mammalian lungs, and unlike the lungs of birds, they are thought to be more sensitive to sudden air pressure changes in their immediate vicinity, such as near wind turbines, and are more liable to rupture.[101] Bats suffer a higher death rate than birds in the neighborhood of wind turbines.[102][103][104] Since there are no signs of external trauma, the cause has been hypothesized to be a greater sensitivity to sudden pressure fluctuations in the mammalian lung than in that of birds.[105] In addition, it has been suggested that bats are attracted to these structures, perhaps seeking roosts, and thereby increasing the death rate.[101] Acoustic deterrents may prove beneficial in mitigating bat mortality at wind energy facilities.[106]
Pathogens and role in the transmission of zoonoses

Among ectoparasites, bats occasionally carry fleas, but are one of the few mammalian orders that cannot host lice (most of the others are water animals).

Bats are natural reservoirs for a large number of zoonotic pathogens,[107] including rabies,[108] severe acute respiratory syndrome (SARS),[109] Henipavirus (i.e. Nipah virus and Hendra virus)[110] and possibly ebola virus.[111][112] Their high mobility, broad distribution, and social behaviour (communal roosting and fission-fusion social structure) make bats favourable hosts and vectors of disease. Many species also appear to have a high tolerance for harbouring pathogens and often do not develop disease while infected. However, contrary to folklore, this is not true of rabies, which is as fatal to bats as it is to all other species. However, a bat may be ill with rabies for a longer time than other mammals.[citation needed]

In regions where rabies is endemic, only 0.5% of bats carry the disease. In the United States, bats typically constitute around a quarter of reported cases of rabies in wild animals. However, their bites account for the vast majority of cases of rabies in humans.[113] Of the 36 cases of domestically acquired rabies recorded in the country in 1995–2010, two were caused by dog bites and four patients were infected by receiving transplants from an organ donor who had previously died of rabies. All other cases were caused by bat bites.[114] Rabies is considered fully preventable if the patient is administered a vaccine prior to the onset of symptoms. However, unlike raccoon or skunk bites, bat bites may go ignored or unnoticed and hence untreated. Rabid bats are broadly distributed throughout the United States; in 2008–2010, cases were reported in every state except Alaska and Hawaii, and Puerto Rico.

Rabid bats may be clumsy, disoriented, and unable to fly, which makes it more likely they will come into contact with humans. One should avoid handling them or having them in one's living space, as with any wild animal. If a bat is found in living quarters near a child, mentally handicapped person, intoxicated person, sleeping person, or pet, living in an area where rabies is known to occur, the person or pet should receive immediate medical attention for rabies. Bats have very small teeth and can bite a sleeping person without being felt. There is evidence that bat rabies virus can infect victims purely through airborne transmission ("cryptic rabies"), without direct physical contact of the victim with the bat itself. This phenomenon has very rarely been reported, and has occurred among victims breathing virus-infected air in environments such as caves, after long exposures.[115][116]

If a bat is found in a house and the possibility of exposure cannot be ruled out, the bat should be sequestered and an animal control officer called immediately, so that the bat can be analysed. This also applies if the bat is found dead. If it is certain that nobody has been exposed to the bat, it should be removed from the house. The best way to do this with live bats is to close all the doors and windows to the room except one that opens to the outside. The bat should soon leave.

Due to the risk of rabies, and to health problems related to their faecal droppings in some regions, bats should be excluded from inhabited parts of houses. The Centers for Disease Control and Prevention (CDC) provide fully detailed information on all aspects of bat management in North America, including how to capture a bat, what to do in case of exposure, and how to bat-proof a house humanely.[117] In certain countries, such as the United Kingdom, it is illegal to handle bats without a license and advice should be sought from an expert organisation, such as the Bat Conservation Trust, if a trapped or injured bat is found.

Where rabies is not endemic, as throughout most of Western Europe, small bats can be considered harmless. Larger bats can give a nasty bite.
Bats in human culture
Mythology
    This section needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. (June 2013)
Common fruit bat (Rousettus aegyptiacus) flying

In European cultures, bats have long been associated with witchcraft, black magic and darkness. The Weird Sisters incorporate the fur of a bat in their brew in Shakespeare's Macbeth, written around 1603-1605.[118] Because bats are mammals, yet can fly, this gives them status as liminal beings in many cultural traditions.[119] In Western culture, the bat is often a symbol of the night and its foreboding nature. The bat is a primary animal associated with fictional characters of the night, both villains, such as Dracula, and heroes, such as Batman. The association of the fear of the night with the animal was treated as a literary challenge by Kenneth Oppel, who created a best-selling series of novels, beginning with Silverwing, which feature bats as the central heroic figures much as anthropomorphized rabbits were the central figures to the classic novel Watership Down.

Among some Native Americans, such as the Creek, Cherokee and Apache, the bat is a trickster spirit. In Tanzania, a winged bat cryptid known as Popobawa, is believed to be a shapeshifting evil spirit that assaults and sodomises its victims.[120]

Not all legends surrounding bats are negative, Chinese lore claims that the bat is a symbol of longevity and happiness, and is similarly lucky in Poland, geographical Macedonia and among the Kwakiutl and Arabs.

An old wives' tale has it that bats will entangle themselves in people's hair. One likely source of this belief is that insect-eating bats seeking prey may dive erratically toward people, who attract mosquitoes and gnats, leading the squeamish to believe the bats are trying to get in their hair.
Aztec

In Mesoamerican mythology during the Classic-Contemporary period, bats symbolized the land of the dead, which was considered to be the underworld.[121] They also symbolized destruction and decay. Bats may have symbolized in this way because they fly only at night and dwell in caves during the daytime and are associated with human skulls and bones by classic Maya ceramists. Central Mexicans sometimes depicted bats having snouts that looked like "sacrificial knives and carrying human head" in the Postclassic era.[122] Bat images were engraved onto funerary urns, and were emphasized with large claws and round ears by Zapotecs. They were commonly associated with death.[123] The depiction of bats on funeral urns and goods took on some the characteristics of the jaguar, which was, and still is, another entity of the night and the underworld. There have also been instances where bats are portrayed next to other animals portrayed negatively in Mesoamerica, including scorpions and other nocturnal animals such as owls. Pre-Columbian cultures associated animals with gods, and often displayed them in art. The Moche people depicted bats in their ceramics.[124]

A life-size, ceramic bat-man was discovered and dug up from the Templo Mayor.[125] The Templo Mayor is located in the center of the Mexica capital of Tenochtitlan. Known as a god of death, this statue has the clawed feet and hands of a bat, but the body of a man. The statue's human-like eyes bulged out from the bat-like head, making the Zapotec images very realistic and living. In the 1930s, the Kaqchikel Maya were said to have proclaimed the bat was the Devil's provider. Kaqchikel would leave the Devil's underworld home and collect blood from the animals to be used for scrumptious meals to feed the Devil. "In the myths, the beast of prey and the animal that is preyed upon play two significant roles. They represent two aspects of life—the aggressive, killing, conquering, creating aspect of life, and the one that is the matter or, you might say, the subject matter".[126] In the Devil's underworld, dead sinners would work off their sins to get to heaven, indicating that the bat, too, was a sinner and worked under the authority of the Devil.[127]
Oaxaca
Zapotec bat god

According to Oaxacan mythology, the bat's nocturnal nature can be traced back to its ancient jealousy of birds' feathers. One day, as the myth goes, the bat felt isolated and undesirable, and told God that he was cold. God, fair and just, turned to birds in the animal kingdom and asked if they would show compassion and donate a feather to the bat to help him keep warm. The birds all agreed and began to pluck one feather from their bodies to give to the bat. With all of these feathers, the bat became even more magnificent-looking than all birds, and was able to spread color to the night sky. During daylight, the bat created rainbows that reflected vibrant colors from the sun. With his new beauty and abilities, the bat soon became arrogant and conceited. The birds grew tired of the bat's self-glorification and decided to fly up to heaven and ask God to do something. When the birds told God of the bat's behaviour, He was surprised and decided to take a look Himself. When on earth, God called on the bat to show him what he was doing. The bat began to fly across the light blue sky where, one by one, each feather began to fall out, uncovering the bat's natural, ugly-looking body. When all his feathers were gone, the bat became distressed and ashamed of his appearance. He decided to hide in caves during the day and only come out during the night to search for his long-lost feathers.[127]
East Nigeria

According to a particular East Nigerian tale, the bat developed its nocturnal habits after causing the death of his partner, the bush-rat. The bat and the bush-rat would share activities, such as rummaging through the grass and trees, hunting, talking and bonding during the day. When at night, the bat and the bush-rat would alternate in cooking duties, cooking what was caught, and eat together. It appeared to be a dedicated partnership, but the bat hated the bush-rat immensely. The bush-rat always found the bat's soup more appetising, so when eating dinner one night, asked the bat why the soup tasted better than his own and also asked how it was made. The bat agreed to show him how to make it the next day, but instead was forming a malicious plan.

Next day, as the bat prepared his soup, the bush-rat came, greeting him and asking if he could be shown what was agreed yesterday. Earlier, the bat had found a pot looking exactly like the one he used usually, but it held warm water and so decided to use this instead. The bat explained to the bush-rat that to make his soup, he had to boil himself prior to serving the soup, where sweetness and flavor of the soup came from the flesh. The bat jumped in the pot seemingly excited, with the bush-rat mesmerised. After a few minutes, the bat climbed out and while the bush-rat was distracted, switched pots. The bat then served his soup out of the soup pot, both tasted it. Overanxious and eager, the bush-rat jumped into the pot of warm water. He stayed much longer in the pot, dying in the process.

When the bush-rat's wife returned that night to find her husband dead, she wept and ran to the chief of the land's house, telling him about what had happened and what she was sure the bat had done. In hearing this, the chief became angry, ordering for the immediate arrest of the bat. It just so happened that the bat was flying over the house and overheard what was just said. He quickly went into hiding high up in a tree. When the chief's men went looking for the bat, he could not be found. The search to arrest the bat carried on over several days, but he still could not be found. The bat needed to eat, so he flew out of hiding every night to hunt for food to avoid being arrested. This, according to Eastern Nigeria mythology, is why bats only fly at night.[128]
Tongan

The bat is sacred in Tonga and is often considered the physical manifestation of a separable soul.[129]
Heraldry
Main article: Bat (heraldry)
Burgee of the Royal Valencia Yacht Club

The bat is sometimes used as a heraldic symbol in Spain. The coats of arms of certain cities in eastern Spain, such as Valencia, Palma de Mallorca and Fraga, have the bat over the shield. Formerly, the Barcelona city coat of arms also had a bat crowning it, but the bat has been removed in the present-day versions. Heraldic use of the bat in Valencia, Catalonia and the Balearic Islands has its origins in a winged dragon (vibra or vibria), which featured in King James I of Aragon's helmet or cimera reial. This is the most widely accepted theory, although there is also a legend that says that, due to the intervention of a bat, King James was able to win a crucial battle against the Saracens that allowed him to win Valencia for his kingdom. The use of the bat as a heraldic symbol is prevalent in the territories of the former Crown of Aragon and it is little used elsewhere. However, it can be found in a few places, as in the coats of arms of the city of Albacete, in Spain, as well as the town of Montchauvet (Yvelines), in France.

Non heraldic organizations also use bats in their symbols. Certain Spanish football clubs including Valencia CF and Levante UD use bats in their badges. The burgee of th

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Football is a family of team sports that involve, to varying degrees, kicking a ball to score a goal. Unqualified, the word football is understood to refer to whichever form of football is the most popular in the regional context in which the word appears. Sports commonly called 'football' in certain places include: association football (known as soccer in some countries); gridiron football (specifically American football or Canadian football); Australian rules football; rugby football (either rugby league or rugby union); and Gaelic football.[1][2] These different variations of football are known as football codes.

Various forms of football can be identified in history, often as popular peasant games. Contemporary codes of football can be traced back to the codification of these games at English public schools during the nineteenth century.[3][4] The expanse of the British Empire allowed these rules of football to spread to areas of British influence outside of the directly controlled Empire,[5] though by the end of the nineteenth century, distinct regional codes were already developing: Gaelic football, for example, deliberately incorporated the rules of local traditional football games in order to maintain their heritage.[6] In 1888, The Football League was founded in England, becoming the first of many professional football competitions. During the twentieth century, several of the various kinds of football grew to become some of the most popular team sports in the world.[7]
Contents
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    1 Common elements
    2 Etymology
    3 Early history
        3.1 Ancient games
        3.2 Medieval and early modern Europe
        3.3 Calcio Fiorentino
        3.4 Official disapproval and attempts to ban football
    4 Establishment of modern codes
        4.1 English public schools
        4.2 Firsts
        4.3 Cambridge rules
        4.4 Sheffield rules
        4.5 Australian rules
        4.6 Football Association
        4.7 Rugby football
        4.8 North American football codes
        4.9 Gaelic football
        4.10 Schism in Rugby football
        4.11 Globalisation of association football
        4.12 Further divergence of the two rugby codes
    5 Use of the word "football"
    6 Popularity
    7 Football codes board
        7.1 Football codes development tree
    8 Present day codes and families
        8.1 Association football and descendants
        8.2 Rugby school football and descendants
        8.3 Irish and Australian varieties
        8.4 Surviving medieval ball games
        8.5 Surviving UK school games
        8.6 Recent inventions and hybrid games
        8.7 Tabletop games, video games and other recreations
    9 See also
    10 Notes
    11 References

Common elements

The various codes of football share certain common elements. Players in American football, Canadian football, rugby union and rugby league take up positions in a limited area of the field at the start of the game.[8] They tend to use throwing and running as the main ways of moving the ball, and only kick on certain limited occasions. Body tackling is a major skill, and games typically involve short passages of play of 5–90 seconds.[8] Association football, Australian rules football and Gaelic football tend to use kicking to move the ball around the pitch, with handling more limited. Body tackles are less central to the game, and players are freer to move around the field (offside laws are typically less strict).[8]

Common rules among the sports include:[9]

    Two teams of usually between 11 and 18 players; some variations that have fewer players (five or more per team) are also popular.
    A clearly defined area in which to play the game.
    Scoring goals or points, by moving the ball to an opposing team's end of the field and either into a goal area, or over a line.
    Goals or points resulting from players putting the ball between two goalposts.
    The goal or line being defended by the opposing team.
    Players being required to move the ball—depending on the code—by kicking, carrying, or hand-passing the ball.
    Players using only their body to move the ball.

In all codes, common skills include passing, tackling, evasion of tackles, catching and kicking.[8] In most codes, there are rules restricting the movement of players offside, and players scoring a goal must put the ball either under or over a crossbar between the goalposts.
Etymology
Main article: Football (word)

There are conflicting explanations of the origin of the word "football". It is widely assumed that the word "football" (or the phrase "foot ball") refers to the action of the foot kicking a ball.[10] There is an alternative explanation, which is that football originally referred to a variety of games in medieval Europe, which were played on foot. There is no conclusive evidence for either explanation.
Early history
Ancient games

The Ancient Greeks and Romans are known to have played many ball games, some of which involved the use of the feet. The Roman game harpastum is believed to have been adapted from a Greek team game known as "ἐπίσκυρος" (Episkyros)[11][12] or "φαινίνδα" (phaininda),[13] which is mentioned by a Greek playwright, Antiphanes (388–311 BC) and later referred to by the Christian theologian Clement of Alexandria (c. 150 – c. 215 AD). These games appear to have resembled rugby football.[14][15][16][17][18] The Roman politician Cicero (106–43 BC) describes the case of a man who was killed whilst having a shave when a ball was kicked into a barber's shop. Roman ball games already knew the air-filled ball, the follis.[19][20] Episkyros is recognised as an early form of football by FIFA.[21]

A Chinese game called Tsu' Chu, Cuju or Zuqiu has been recognised by FIFA as the first version of the game with regular rules.[22] It existed during the Han Dynasty, the second and third centuries BC. The Japanese version of cuju is kemari (蹴鞠), and was developed during the Asuka period.[23] This is known to have been played within the Japanese imperial court in Kyoto from about 600 AD. In kemari several people stand in a circle and kick a ball to each other, trying not to let the ball drop to the ground (much like keepie uppie). The game appears to have died out sometime before the mid-19th century. It was revived in 1903 and is now played at a number of festivals.[citation needed]

There are a number of references to traditional, ancient, or prehistoric ball games, played by indigenous peoples in many different parts of the world. For example, in 1586, men from a ship commanded by an English explorer named John Davis, went ashore to play a form of football with Inuit (Eskimo) people in Greenland.[24] There are later accounts of an Inuit game played on ice, called Aqsaqtuk. Each match began with two teams facing each other in parallel lines, before attempting to kick the ball through each other team's line and then at a goal. In 1610, William Strachey, a colonist at Jamestown, Virginia recorded a game played by Native Americans, called Pahsaheman.[citation needed] On the Australian continent several tribes of indigenous people played kicking and catching games with stuffed balls which have been generalised by historians as Marn Grook (Djab Wurrung for "game ball"). The earliest historical account is an anecdote from the 1878 book by Robert Brough-Smyth, The Aborigines of Victoria, in which a man called Richard Thomas is quoted as saying, in about 1841 in Victoria, Australia, that he had witnessed Aboriginal people playing the game: "Mr Thomas describes how the foremost player will drop kick a ball made from the skin of a possum and how other players leap into the air in order to catch it." Some historians have theorised that Marn Grook was one of the origins of Australian rules football.

The Māori in New Zealand played a game called Ki-o-rahi consisting of teams of seven players play on a circular field divided into zones, and score points by touching the 'pou' (boundary markers) and hitting a central 'tupu' or target.[citation needed]

Games played in Mesoamerica with rubber balls by indigenous peoples are also well-documented as existing since before this time, but these had more similarities to basketball or volleyball, and since their influence on modern football games is minimal, most do not class them as football.[citation needed]Northeastern American Indians, especially the Iroquois Confederation, played a game which made use of net racquets to throw and catch a small ball; however, although it is a ball-goal foot game, lacrosse (as its modern descendant is called) is likewise not usually classed as a form of "football."[citation needed]

These games and others may well go far back into antiquity. However, the main sources of modern football codes appear to lie in western Europe, especially England.

    Ancient Greek football player balancing the ball. Depiction on an Attic Lekythos.

    A Song Dynasty painting by Su Hanchen, depicting Chinese children playing cuju.

    Paint of a Mesoamerican ballgame player of the Tepantitla murals in Teotihuacan.

    A revived version of kemari being played at the Tanzan Shrine, Japan.

    An illustration from the 1850s of Australian Aboriginal hunter gatherers. File:Children in the background are playing a football game, possibly Woggabaliri.[25]

    A group of aborigines playing football in Guiana.

Medieval and early modern Europe
Further information: Medieval football

The Middle Ages saw a huge rise in popularity of annual Shrovetide football matches throughout Europe, particularly in England. An early reference to a ball game played in Britain comes from the 9th century Historia Brittonum, which describes "a party of boys ... playing at ball".[26] References to a ball game played in northern France known as La Soule or Choule, in which the ball was propelled by hands, feet, and sticks,[27] date from the 12th century.[28]
An illustration of so-called "mob football"

The early forms of football played in England, sometimes referred to as "mob football", would be played between neighbouring towns and villages, involving an unlimited number of players on opposing teams who would clash en masse,[29] struggling to move an item, such as inflated animal's bladder[30] to particular geographical points, such as their opponents' church, with play taking place in the open space between neighbouring parishes.[31] The game was played primarily during significant religious festivals, such as Shrovetide, Christmas, or Easter,[30] and Shrovetide games have survived into the modern era in a number of English towns (see below).

The first detailed description of what was almost certainly football in England was given by William FitzStephen in about 1174–1183. He described the activities of London youths during the annual festival of Shrove Tuesday:

    After lunch all the youth of the city go out into the fields to take part in a ball game. The students of each school have their own ball; the workers from each city craft are also carrying their balls. Older citizens, fathers, and wealthy citizens come on horseback to watch their juniors competing, and to relive their own youth vicariously: you can see their inner passions aroused as they watch the action and get caught up in the fun being had by the carefree adolescents.[32]

Most of the very early references to the game speak simply of "ball play" or "playing at ball". This reinforces the idea that the games played at the time did not necessarily involve a ball being kicked.

An early reference to a ball game that was probably football comes from 1280 at Ulgham, Northumberland, England: "Henry... while playing at ball.. ran against David".[33] Football was played in Ireland in 1308, with a documented reference to John McCrocan, a spectator at a "football game" at Newcastle, County Down being charged with accidentally stabbing a player named William Bernard.[34] Another reference to a football game comes in 1321 at Shouldham, Norfolk, England: "[d]uring the game at ball as he kicked the ball, a lay friend of his... ran against him and wounded himself".[33]

In 1314, Nicholas de Farndone, Lord Mayor of the City of London issued a decree banning football in the French used by the English upper classes at the time. A translation reads: "[f]orasmuch as there is great noise in the city caused by hustling over large foot balls [rageries de grosses pelotes de pee][35] in the fields of the public from which many evils might arise which God forbid: we command and forbid on behalf of the king, on pain of imprisonment, such game to be used in the city in the future." This is the earliest reference to football.

In 1363, King Edward III of England issued a proclamation banning "...handball, football, or hockey; coursing and cock-fighting, or other such idle games",[36] showing that "football" – whatever its exact form in this case – was being differentiated from games involving other parts of the body, such as handball.

A game known as "football" was played in Scotland as early as the 15th century: it was prohibited by the Football Act 1424 and although the law fell into disuse it was not repealed until 1906. There is evidence for schoolboys playing a "football" ball game in Aberdeen in 1633 (some references cite 1636) which is notable as an early allusion to what some have considered to be passing the ball. The word "pass" in the most recent translation is derived from "huc percute" (strike it here) and later "repercute pilam" (strike the ball again) in the original Latin. It is not certain that the ball was being struck between members of the same team. The original word translated as "goal" is "metum", literally meaning the "pillar at each end of the circus course" in a Roman chariot race. There is a reference to "get hold of the ball before [another player] does" (Praeripe illi pilam si possis agere) suggesting that handling of the ball was allowed. One sentence states in the original 1930 translation "Throw yourself against him" (Age, objice te illi).
France circa 1750

King Henry IV of England also presented one of the earliest documented uses of the English word "football", in 1409, when he issued a proclamation forbidding the levying of money for "foteball".[33][37]

There is also an account in Latin from the end of the 15th century of football being played at Cawston, Nottinghamshire. This is the first description of a "kicking game" and the first description of dribbling: "[t]he game at which they had met for common recreation is called by some the foot-ball game. It is one in which young men, in country sport, propel a huge ball not by throwing it into the air but by striking it and rolling it along the ground, and that not with their hands but with their feet... kicking in opposite directions" The chronicler gives the earliest reference to a football pitch, stating that: "[t]he boundaries have been marked and the game had started.[33]

Other firsts in the mediæval and early modern eras:

    "a football", in the sense of a ball rather than a game, was first mentioned in 1486.[37] This reference is in Dame Juliana Berners' Book of St Albans. It states: "a certain rounde instrument to play with ...it is an instrument for the foote and then it is calde in Latyn 'pila pedalis', a fotebal."[33]
    a pair of football boots was ordered by King Henry VIII of England in 1526.[38]
    women playing a form of football was first described in 1580 by Sir Philip Sidney in one of his poems: "[a] tyme there is for all, my mother often sayes, When she, with skirts tuckt very hy, with girles at football playes."[39]
    the first references to goals are in the late 16th and early 17th centuries. In 1584 and 1602 respectively, John Norden and Richard Carew referred to "goals" in Cornish hurling. Carew described how goals were made: "they pitch two bushes in the ground, some eight or ten foote asunder; and directly against them, ten or twelue [twelve] score off, other twayne in like distance, which they terme their Goales".[40] He is also the first to describe goalkeepers and passing of the ball between players.
    the first direct reference to scoring a goal is in John Day's play The Blind Beggar of Bethnal Green (performed circa 1600; published 1659): "I'll play a gole at camp-ball" (an extremely violent variety of football, which was popular in East Anglia). Similarly in a poem in 1613, Michael Drayton refers to "when the Ball to throw, And drive it to the Gole, in squadrons forth they goe".

Calcio Fiorentino
An illustration of the Calcio Fiorentino field and starting positions, from a 1688 book by Pietro di Lorenzo Bini.
Main article: Calcio Fiorentino

In the 16th century, the city of Florence celebrated the period between Epiphany and Lent by playing a game which today is known as "calcio storico" ("historic kickball") in the Piazza Santa Croce. The young aristocrats of the city would dress up in fine silk costumes and embroil themselves in a violent form of football. For example, calcio players could punch, shoulder charge, and kick opponents. Blows below the belt were allowed. The game is said to have originated as a military training exercise. In 1580, Count Giovanni de' Bardi di Vernio wrote Discorso sopra 'l giuoco del Calcio Fiorentino. This is sometimes said to be the earliest code of rules for any football game. The game was not played after January 1739 (until it was revived in May 1930).
Official disapproval and attempts to ban football
Main article: Attempts to ban football games

There have been many attempts to ban football, from the middle ages through to the modern day. The first such law was passed in England in 1314; it was followed by more than 30 in England alone between 1314 and 1667.[41]:6 Football faced armed opposition in the 18th Century when used as a cover for violent protest against the enclosure act. Women were banned from playing at English and Scottish Football League grounds in 1921, a ban that was only lifted in the 1970s. Female footballers still face similar problems in some parts of the world.
Establishment of modern codes
English public schools
Main article: English public school football games

While football continued to be played in various forms throughout Britain, its public schools (known as private schools in other countries) are widely credited with four key achievements in the creation of modern football codes. First of all, the evidence suggests that they were important in taking football away from its "mob" form and turning it into an organised team sport. Second, many early descriptions of football and references to it were recorded by people who had studied at these schools. Third, it was teachers, students and former students from these schools who first codified football games, to enable matches to be played between schools. Finally, it was at English public schools that the division between "kicking" and "running" (or "carrying") games first became clear.

The earliest evidence that games resembling football were being played at English public schools – mainly attended by boys from the upper, upper-middle and professional classes – comes from the Vulgaria by William Herman in 1519. Herman had been headmaster at Eton and Winchester colleges and his Latin textbook includes a translation exercise with the phrase "We wyll playe with a ball full of wynde".[42]

Richard Mulcaster, a student at Eton College in the early 16th century and later headmaster at other English schools, has been described as "the greatest sixteenth Century advocate of football".[43] Among his contributions are the earliest evidence of organised team football. Mulcaster's writings refer to teams ("sides" and "parties"), positions ("standings"), a referee ("judge over the parties") and a coach "(trayning maister)". Mulcaster's "footeball" had evolved from the disordered and violent forms of traditional football:

    [s]ome smaller number with such overlooking, sorted into sides and standings, not meeting with their bodies so boisterously to trie their strength: nor shouldring or shuffing one an other so barbarously ... may use footeball for as much good to the body, by the chiefe use of the legges.[44]

In 1633, David Wedderburn, a teacher from Aberdeen, mentioned elements of modern football games in a short Latin textbook called Vocabula. Wedderburn refers to what has been translated into modern English as "keeping goal" and makes an allusion to passing the ball ("strike it here"). There is a reference to "get hold of the ball", suggesting that some handling was allowed. It is clear that the tackles allowed included the charging and holding of opposing players ("drive that man back").[citation needed]

A more detailed description of football is given in Francis Willughby's Book of Games, written in about 1660.[45] Willughby, who had studied at Bishop Vesey's Grammar School, Sutton Coldfield, is the first to describe goals and a distinct playing field: "a close that has a gate at either end. The gates are called Goals." His book includes a diagram illustrating a football field. He also mentions tactics ("leaving some of their best players to guard the goal"); scoring ("they that can strike the ball through their opponents' goal first win") and the way teams were selected ("the players being equally divided according to their strength and nimbleness"). He is the first to describe a "law" of football: "they must not strike [an opponent's leg] higher than the ball".[citation needed]

English public schools were the first to codify football games. In particular, they devised the first offside rules, during the late 18th century.[46] In the earliest manifestations of these rules, players were "off their side" if they simply stood between the ball and the goal which was their objective. Players were not allowed to pass the ball forward, either by foot or by hand. They could only dribble with their feet, or advance the ball in a scrum or similar formation. However, offside laws began to diverge and develop differently at each school, as is shown by the rules of football from Winchester, Rugby, Harrow and Cheltenham, during between 1810 and 1850.[46] The first known codes – in the sense of a set of rules – were those of Eton in 1815 [47] and Aldenham in 1825.[47])

During the early 19th century, most working class people in Britain had to work six days a week, often for over twelve hours a day. They had neither the time nor the inclination to engage in sport for recreation and, at the time, many children were part of the labour force. Feast day football played on the streets was in decline. Public school boys, who enjoyed some freedom from work, became the inventors of organised football games with formal codes of rules.

Football was adopted by a number of public schools as a way of encouraging competitiveness and keeping youths fit. Each school drafted its own rules, which varied widely between different schools and were changed over time with each new intake of pupils. Two schools of thought developed regarding rules. Some schools favoured a game in which the ball could be carried (as at Rugby, Marlborough and Cheltenham), while others preferred a game where kicking and dribbling the ball was promoted (as at Eton, Harrow, Westminster and Charterhouse). The division into these two camps was partly the result of circumstances in which the games were played. For example, Charterhouse and Westminster at the time had restricted playing areas; the boys were confined to playing their ball game within the school cloisters, making it difficult for them to adopt rough and tumble running games.[citation needed]
Rugby School

William Webb Ellis, a pupil at Rugby School, is said to have "with a fine disregard for the rules of football, as played in his time [emphasis added], first took the ball in his arms and ran with it, thus creating the distinctive feature of the rugby game." in 1823. This act is usually said to be the beginning of Rugby football, but there is little evidence that it occurred, and most sports historians believe the story to be apocryphal. The act of 'taking the ball in his arms' is often misinterpreted as 'picking the ball up' as it is widely believed that Webb Ellis' 'crime' was handling the ball, as in modern soccer, however handling the ball at the time was often permitted and in some cases compulsory,[48] the rule for which Webb Ellis showed disregard was running forward with it as the rules of his time only allowed a player to retreat backwards or kick forwards.

The boom in rail transport in Britain during the 1840s meant that people were able to travel further and with less inconvenience than they ever had before. Inter-school sporting competitions became possible. However, it was difficult for schools to play each other at football, as each school played by its own rules. The solution to this problem was usually that the match be divided into two halves, one half played by the rules of the host "home" school, and the other half by the visiting "away" school.

The modern rules of many football codes were formulated during the mid- or late- 19th century. This also applies to other sports such as lawn bowls, lawn tennis, etc. The major impetus for this was the patenting of the world's first lawnmower in 1830. This allowed for the preparation of modern ovals, playing fields, pitches, grass courts, etc.[49]

Apart from Rugby football, the public school codes have barely been played beyond the confines of each school's playing fields. However, many of them are still played at the schools which created them (see Surviving UK school games below).

Public schools' dominance of sports in the UK began to wane after the Factory Act of 1850, which significantly increased the recreation time available to working class children. Before 1850, many British children had to work six days a week, for more than twelve hours a day. From 1850, they could not work before 6 a.m. (7 a.m. in winter) or after 6 p.m. on weekdays (7 p.m. in winter); on Saturdays they had to cease work at 2 p.m. These changes mean that working class children had more time for games, including various forms of football.
Firsts
Clubs
Main article: Oldest football clubs

Sports clubs dedicated to playing football began in the 18th century, for example London's Gymnastic Society which was founded in the mid-18th century and ceased playing matches in 1796.[50][51]

The first documented club to bear in the title a reference to being a 'football club' were called "The Foot-Ball Club" who were located in Edinburgh, Scotland, during the period 1824–41.[52][53] The club forbade tripping but allowed pushing and holding and the picking up of the ball.[53]

In 1845, three boys at Rugby school were tasked with codifying the rules then being used at the school. These were the first set of written rules (or code) for any form of football.[54] This further assisted the spread of the Rugby game.
Competitions
Main article: Oldest football competitions

One of the longest running football fixture is the Cordner-Eggleston Cup, contested between Melbourne Grammar School and Scotch College, Melbourne every year since 1858. It is believed by many to also be the first match of Australian rules football, although it was played under experimental rules in its first year. The first football trophy tournament was the Caledonian Challenge Cup, donated by the Royal Caledonian Society of Melbourne, played in 1861 under the Melbourne Rules.[55] The oldest football league is a rugby football competition, the United Hospitals Challenge Cup (1874), while the oldest rugby trophy is the Yorkshire Cup, contested since 1878. The South Australian Football Association (30 April 1877) is the oldest surviving Australian rules football competition. The oldest surviving soccer trophy is the Youdan Cup (1867) and the oldest national soccer competition is the English FA Cup (1871). The Football League (1888) is recognised as the longest running Association Football league. The first ever international football match took place between sides representing England and Scotland on March 5, 1870 at the Oval under the authority of the FA. The first Rugby international took place in 1871.
Modern balls
Main article: Football (ball)
Richard Lindon (seen in 1880) is believed to have invented the first footballs with rubber bladders.

In Europe, early footballs were made out of animal bladders, more specifically pig's bladders, which were inflated. Later leather coverings were introduced to allow the balls to keep their shape.[56] However, in 1851, Richard Lindon and William Gilbert, both shoemakers from the town of Rugby (near the school), exhibited both round and oval-shaped balls at the Great Exhibition in London. Richard Lindon's wife is said to have died of lung disease caused by blowing up pig's bladders.[57] Lindon also won medals for the invention of the "Rubber inflatable Bladder" and the "Brass Hand Pump".

In 1855, the U.S. inventor Charles Goodyear – who had patented vulcanized rubber – exhibited a spherical football, with an exterior of vulcanized rubber panels, at the Paris Exhibition Universelle. The ball was to prove popular in early forms of football in the U.S.A.[58]

The iconic ball with a regular pattern of hexagons and pentagons (see truncated icosahedron) did not become popular until the 1960s, and was first used in the World Cup in 1970.
Modern ball passing tactics
Main article: Passing (association football)

The earliest reference to a game of football involving players passing the ball and attempting to score past a goalkeeper was written in 1633 by David Wedderburn, a poet and teacher in Aberdeen, Scotland.[59] Nevertheless, the original text does not state whether the allusion to passing as 'kick the ball back' ('Repercute pilam') was in a forward or backward direction or between members of the same opposing teams (as was usual at this time)[60]

"Scientific" football is first recorded in 1839 from Lancashire[61] and in the modern game in Rugby football from 1862[62] and from Sheffield FC as early as 1865.[63][64] The first side to play a passing combination game was the Royal Engineers AFC in 1869/70[65][66] By 1869 they were "work[ing] well together", "backing up" and benefiting from "cooperation".[67] By 1870 the Engineers were passing the ball: "Lieut. Creswell, who having brought the ball up the side then kicked it into the middle to another of his side, who kicked it through the posts the minute before time was called"[68] Passing was a regular feature of their style[69] By early 1872 the Engineers were the first football team renowned for "play[ing] beautifully together"[70] A double pass is first reported from Derby school against Nottingham Forest in March 1872, the first of which is irrefutably a short pass: "Mr Absey dribbling the ball half the length of the field delivered it to Wallis, who kicking it cleverly in front of the goal, sent it to the captain who drove it at once between the Nottingham posts"[71] The first side to have perfected the modern formation was Cambridge University AFC[72][73][74] and introduced the 2–3–5 "pyramid" formation.[75][76]
Cambridge rules
Main article: Cambridge rules

In 1848, at Cambridge University, Mr. H. de Winton and Mr. J.C. Thring, who were both formerly at Shrewsbury School, called a meeting at Trinity College, Cambridge with 12 other representatives from Eton, Harrow, Rugby, Winchester and Shrewsbury. An eight-hour meeting produced what amounted to the first set of modern rules, known as the Cambridge rules. No copy of these rules now exists, but a revised version from circa 1856 is held in the library of Shrewsbury School.[77] The rules clearly favour the kicking game. Handling was only allowed when a player catches the ball directly from the foot entitling them to a free kick and there was a primitive offside rule, disallowing players from "loitering" around the opponents' goal. The Cambridge rules were not widely adopted outside English public schools and universities (but it was arguably the most significant influence on the Football Association committee members responsible for formulating the rules of Association football).
Sheffield rules
Main article: Sheffield rules

By the late 1850s, many football clubs had been formed throughout the English-speaking world, to play various codes of football. Sheffield Football Club, founded in 1857 in the English city of Sheffield by Nathaniel Creswick and William Prest, was later recognised as the world's oldest club playing association football.[78] However, the club initially played its own code of football: the Sheffield rules. The code was largely independent of the public school rules, the most significant difference being the lack of an offside rule.

The code was responsible for many innovations that later spread to association football. These included free kicks, corner kicks, handball, throw-ins and the crossbar.[79] By the 1870s they became the dominant code in the north and midlands of England. At this time a series of rule changes by both the London and Sheffield FAs gradually eroded the differences between the two games until the adoption of a common code in 1877.
Australian rules
Main article: Australian rules football
See also: Origins of Australian rules football
Tom Wills, widely regarded as the father of Australian football

There is archival evidence of "foot-ball" games being played in various parts of Australia throughout the first half of the 19th century. The origins of an organised game of football known today as Australian rules football can be traced back to 1858 in Melbourne, the capital city of Victoria.

In July 1858, Tom Wills, an Australian-born cricketer educated at Rugby School in England, wrote a letter to Bell's Life in Victoria & Sporting Chronicle, calling for a "foot-ball club" with a "code of laws" to keep cricketers fit during winter.[80] This is considered by historians to be a defining moment in the creation of Australian rules football. Through publicity and personal contacts Wills was able to co-ordinate football matches in Melbourne that experimented with various rules,[81] the first of which was played on July 31, 1858. One week later, Wills umpired a schoolboys match between Melbourne Grammar School and Scotch College. Following these matches, organised football in Melbourne rapidly increased in popularity.
Wood engraving of an Australian rules football match at the Richmond Paddock, Melbourne, 1866

Wills and others involved in these early matches formed the Melbourne Football Club (the oldest surviving Australian football club) on May 14, 1859. Club members Wills, William Hammersley, J. B. Thompson and Thomas H. Smith met with the intention of forming a set of rules that would be widely adopted by other clubs. The committee debated rules used in English public school games; Wills pushed for various rugby football rules he learnt during his schooling. The first rules share similarities with these games, and were shaped to suit to Australian conditions. H. C. A. Harrison, a seminal figure in Australian football, recalled that his cousin Wills wanted "a game of our own".[82] The code was distinctive in the prevalence of the mark, free kick, tackling, lack of an offside rule and that players were specifically penalised for throwing the ball.

The Melbourne football rules were widely distributed and gradually adopted by the other Victorian clubs. The rules were updated several times during the 1860s to accommodate the rules of other influential Victorian football clubs. A significant redraft in 1866 by H. C. A. Harrison's committee accommodated the Geelong Football Club's rules, making the game then known as "Victorian Rules" increasingly distinct from other codes. It soon adopted cricket fields and an oval ball, used specialised goal and behind posts, and featured bouncing the ball while running and spectacular high marking. The game spread quickly to other Australian colonies. Outside of its heartland in southern Australia the code experienced a significant period of decline following World War I but has since grown throughout Australia and in other parts of the world, and the Australian Football League emerged as the dominant professional competition.
Football Association
Main article: The Football Association
The first football international, Scotland versus England. Once kept by the Rugby Football Union as an early example of rugby football.

During the early 1860s, there were increasing attempts in England to unify and reconcile the various public school games. In 1862, J. C. Thring, who had been one of the driving forces behind the original Cambridge Rules, was a master at Uppingham School and he issued his own rules of what he called "The Simplest Game" (these are also known as the Uppingham Rules). In early October 1863 another new revised version of the Cambridge Rules was drawn up by a seven member committee representing former pupils from Harrow, Shrewsbury, Eton, Rugby, Marlborough and Westminster.

At the Freemasons' Tavern, Great Queen Street, London on the evening of October 26, 1863, representatives of several football clubs in the London Metropolitan area met for the inaugural meeting of The Football Association (FA). The aim of the Association was to establish a single unifying code and regulate the playing of the game among its members. Following the first meeting, the public schools were invited to join the association. All of them declined, except Charterhouse and Uppingham. In total, six meetings of the FA were held between October and December 1863. After the third meeting, a draft set of rules were published. However, at the beginning of the fourth meeting, attention was drawn to the recently published Cambridge Rules of 1863. The Cambridge rules differed from the draft FA rules in two significant areas; namely running with (carrying) the ball and hacking (kicking opposing players in the shins). The two contentious FA rules were as follows:

    IX. A player shall be entitled to run with the ball towards his adversaries' goal if he makes a fair catch, or catches the ball on the first bound; but in case of a fair catch, if he makes his mark he shall not run.
    X. If any player shall run with the ball towards his adversaries' goal, any player on the opposite side shall be at liberty to charge, hold, trip or hack him, or to wrest the ball from him, but no player shall be held and hacked at the same time.
    — [83]

At the fifth meeting it was proposed that these two rules be removed. Most of the delegates supported this, but F. M. Campbell, the representative from Blackheath and the first FA treasurer, objected. He said: "hacking is the true football". However, the motion to ban running with the ball in hand and hacking was carried and Blackheath withdrew from the FA. After the final meeting on 8 December, the FA published the "Laws of Football", the first comprehensive set of rules for the game later known as Association Football. The term "soccer", in use since the late 19th century, derives from an Oxford University abbreviation of "Association".[84]

The first FA rules still contained elements that are no longer part of association football, but which are still recognisable in other games (such as Australian football and rugby football): for instance, a player could make a fair catch and claim a mark, which entitled him to a free kick; and if a player touched the ball behind the opponents' goal line, his side was entitled to a free kick at goal, from 15 yards (13.5 metres) in front of the goal line.
Rugby football
Main article: History of rugby union
A rugby scrum in 1871

In Britain, by 1870, there were about 75 clubs playing variations of the Rugby school game. There were also "rugby" clubs in Ireland, Australia, Canada and New Zealand. However, there was no generally accepted set of rules for rugby until 1871, when 21 clubs from London came together to form the Rugby Football Union (RFU). The first official RFU rules were adopted in June 1871. These rules allowed passing the ball. They also included the try, where touching the ball over the line allowed an attempt at goal, though drop-goals from marks and general play, and penalty conversions were still the main form of contest.
North American football codes
Main articles: History of American football and Canadian football § History

As was the case in Britain, by the early 19th century, North American schools and universities played their own local games, between sides made up of students. For example, students at Dartmouth College in New Hampshire played a game called Old division football, a variant of the association football codes, as early as the 1820s.[85] They remained largely "mob football" style games, with huge numbers of players attempting to advance the ball into a goal area, often by any means necessary. Rules were simple, violence and injury were common.[86] The violence of these mob-style games led to widespread protests and a decision to abandon them. Yale University, under pressure from the city of New Haven, banned the play of all forms of football in 1860, while Harvard University followed suit in 1861.[86] In its place, two general types of football evolved: "kicking" games and "running" (or "carrying") games. A hybrid of the two, known as the "Boston game", was played by a group known as the Oneida Football Club. The club, considered by some historians as the first formal football club in the United States, was formed in 1862 by schoolboys who played the "Boston game" on Boston Common.[86][87] The game began to return to American college campuses by the late 1860s. The universities of Yale, Princeton (then known as the College of New Jersey), Rutgers, and Brown all began playing "kicking" games during this time. In 1867, Princeton used rules based on those of the English Football Association.[86]
The "Tigers" of Hamilton, Ontario, circa 1906. Founded 1869 as the Hamilton Foot Ball Club, they eventually merged with the Hamilton Flying Wildcats to form the Hamilton Tiger-Cats, a team still active in the Canadian Football League.[88]

In Canada, the first documented football match was a practice game played on November 9, 1861, at University College, University of Toronto (approximately 400 yards west of Queen's Park). One of the participants in the game involving University of Toronto students was (Sir) William Mulock, later Chancellor of the school.[89] In 1864, at Trinity College, Toronto, F. Barlow Cumberland, Frederick A. Bethune, and Christopher Gwynn, one of the founders of Milton, Massachusetts, devised rules based on rugby football.[89] A "running game", resembling rugby football, was then taken up by the Montreal Football Club in Canada in 1868.[90]

On November 6, 1869, Rutgers faced Princeton in a game that was played with a round ball and, like all early games, used improvised rules. It is usually regarded as the first game of American intercollegiate football.[86][91]

Modern North American football grew out of a match between McGill University of Montreal, and Harvard University in 1874. During the game, the two teams alternated between the rugby-based rules used by McGill and the Boston Game rules used by Harvard.[92][93][94] Within a few years, Harvard had both adopted McGill's rules and had persuaded other U.S. university teams to do the same. On November 23, 1876, representatives from Harvard, Yale, Princeton, and Columbia met at the Massasoit Convention in Springfield, Massachusetts, agreeing to adopt most of the Rugby Football Union rules, with some variations.[95]
Rutgers College Football Team, 1882

In 1880, Yale coach Walter Camp, who had become a fixture at the Massasoit House conventions where the rules were debated and changed, devised a number of major innovations. Camp's two most important rule changes that diverged the American game from rugby was replacing the scrummage with the line of scrimmage and the establishment of the down-and-distance rules.[95] American football still however remained a violent sport where collisions often led to serious injuries and sometimes even death.[96] This led U.S. President Theodore Roosevelt to hold a meeting with football representatives from Harvard, Yale, and Princeton on October 9, 1905, urging them to make drastic changes.[97] One rule change introduced in 1906, devised to open up the game and reduce injury, was the introduction of the legal forward pass. Though it was underutilized for years, this proved to be one of the most important rule changes in the establishment of the modern game.[98]

Over the years, Canada absorbed some of the developments in American football in an effort to distinguish it from a more rugby-oriented game. In 1903, the Ontario Rugby Football Union adopted the Burnside rules, which implemented the line of scrimmage and down-and-distance system from American football, among others.[99] Canadian football then implemented the legal forward pass in 1929.[100] American and Canadian football remain different codes, stemming from rule changes that the American side of the border adopted but the Canadian side has not.
Gaelic football
The All-Ireland Football Final in Croke Park, 2004.
Main article: History of Gaelic football

In the mid-19th century, various traditional football games, referred to collectively as caid, remained popular in Ireland, especially in County Kerry. One observer, Father W. Ferris, described two main forms of caid during this period: the "field game" in which the object was to put the ball through arch-like goals, formed from the boughs of two trees; and the epic "cross-country game" which took up most of the daylight hours of a Sunday on which it was played, and was won by one team taking the ball across a parish boundary. "Wrestling", "holding" opposing players, and carrying the ball were all allowed.

By the 1870s, Rugby and Association football had started to become popular in Ireland. Trinity College, Dublin was an early stronghold of Rugby (see the Developments in the 1850s section, above). The rules of the English FA were being distributed widely. Traditional forms of caid had begun to give way to a "rough-and-tumble game" which allowed tripping.

There was no serious attempt to unify and codify Irish varieties of football, until the establishment of the Gaelic Athletic Association (GAA) in 1884. The GAA sought to promote traditional Irish sports, such as hurling and to reject imported games like Rugby and Association football. The first Gaelic football rules were drawn up by Maurice Davin and published in the United Ireland magazine on February 7, 1887. Davin's rules showed the influence of games such as hurling and a desire to formalise a distinctly Irish code of football. The prime example of this differentiation was the lack of an offside rule (an attribute which, for many years, was shared only by other Irish games like hurling, and by Australian rules football).
Schism in Rugby football
An English cartoon from the 1890s lampooning the divide in rugby football which led to the formation of rugby league. The caricatures are of Rev. Frank Marshall, an arch-opponent of player payments, and James Miller, a long-time opponent of Marshall. The caption reads: Marshall: "Oh, fie, go away naughty boy, I don't play with boys who can’t afford to take a holiday for football any day they like!" Miller: "Yes, that's just you to a T; you’d make it so that no lad whose father wasn’t a millionaire could play at all in a really good team. For my part I see no reason why the men who make the money shouldn’t have a share in the spending of it."
Further information: History of rugby league

The International Rugby Football Board (IRFB) was founded in 1886, but rifts were beginning to emerge in the code. Professionalism was beginning to creep into the various codes of football.

In England, by the 1890s, a long-standing Rugby Football Union ban on professional players was causing regional tensions within rugby football, as many players in northern England were working class and could not afford to take time off to train, travel, play and recover from injuries. This was not very different from what had occurred ten years earlier in soccer in Northern England but the authorities reacted very differently in the RFU, attempting to alienate the working class support in Northern England. In 1895, following a dispute about a player being paid broken time payments, which replaced wages lost as a result of playing rugby, representatives of the northern clubs met in Huddersfield to form the Northern Rugby Football Union (NRFU). The new body initially permitted only various types of player wage replacements. However, within two years, NRFU players could be paid, but they were required to have a job outside sport.

The demands of a professional league dictated that rugby had to become a better "spectator" sport. Within a few years the NRFU rules had started to diverge from the RFU, most notably with the abolition of the line-out. This was followed by the replacement of the ruck with the "play-the-ball ruck", which allowed a two-player ruck contest between the tackler at marker and the player tackled. Mauls were stopped once the ball carrier was held, being replaced by a play-the ball-ruck. The separate Lancashire and Yorkshire competitions of the NRFU merged in 1901, forming the Northern Rugby League, the first time the name rugby league was used officially in England.

Over time, the RFU form of rugby, played by clubs which remained members of national federations affiliated to the IRFB, became known as rugby union.
Globalisation of association football
Main article: History of FIFA

The need for a single body to oversee association football had become apparent by the beginning of the 20th century, with the increasing popularity of international fixtures. The English Football Association had chaired many discussions on setting up an international body, but was perceived as making no progress. It fell to associations from seven other European countries: France, Belgium, Denmark, Netherlands, Spain, Sweden, and Switzerland, to form an international association. The Fédération Internationale de Football Association (FIFA) was founded in Paris on May 21, 1904. Its first president was Robert Guérin. The French name and acronym has remained, even outside French-speaking countries.
Further divergence of the two rugby codes

Rugby league rules diverged significantly from rugby union in 1906, with the reduction of the team from 15 to 13 players. In 1907, a New Zealand professional rugby team toured Australia and Britain, receiving an enthusiastic response, and professional rugby leagues were launched in Australia the following year. However, the rules of professional games varied from one country to another, and negotiations between various national bodies were required to fix the exact rules for each international match. This situation endured until 1948, when at the instigation of the French league, the Rugby League International Federation (RLIF) was formed at a meeting in Bordeaux.

During the second half of 20th century, the rules changed further. In 1966, rugby league officials borrowed the American football concept of downs: a team was allowed to retain possession of the ball for four tackles (rugby union retains the original rule that a player who is tackled and brought to the ground must release the ball immediately). The maximum number of tackles was later increased to six (in 1971), and in rugby league this became known as the six tackle rule.

With the advent of full-time professionals in the early 1990s, and the consequent speeding up of the game, the five metre off-side distance between the two teams became 10 metres, and the replacement rule was superseded by various interchange rules, among other changes.

The laws of rugby union also changed during the 20th century, although less significantly than those of rugby league. In particular, goals from marks were abolished, kicks directly into touch from outside the 22 metre line were penalised, new laws were put in place to determine who had possession following an inconclusive ruck or maul, and the lifting of players in line-outs was legalised.

In 1995, rugby union became an "open" game, that is one which allowed professional players. Although the original dispute between the two codes has now disappeared – and despite the fact that officials from both forms of rugby football have sometimes mentioned the possibility of re-unification – the rules of both codes and their culture have diverged to such an extent that such an event is unlikely in the foreseeable future.
A player takes a free kick, while the opposition form a "wall", in Association football
Use of the word "football"
For more details on this topic, see Football (word).

The word "football", when used in reference to a specific game can mean any one of those described above. Because of this, much friendly controversy has occurred over the term football, primarily because it is used in different ways in different parts of the English-speaking world. Most often, the word "football" is used to refer to the code of football that is considered dominant within a particular region. So, effectively, what the word "football" means usually depends on where one says it.
Players assemble at the line of scrimmage in an American football game.

In each of the United Kingdom, the United States, and Canada, one football code is known solely as "football", while the others generally require a qualifier. In New Zealand, "football" historically referred to rugby union, but more recently may be used unqualified to refer to association football. The sport meant by the word "football" in Australia is either Australian rules football or rugby league, depending on popularity (which largely conforms to what has been called the Barassi Line). In francophone Quebec, where Canadian football is more popular, the Canadian code is known as football while American football is known as Football américain and association football is known as le soccer.[101] Of the 45 national FIFA (Fédération Internationale de Football Association) affiliates in which English is an official or primary language, most currently use Football in their organizations' official names; the FIFA affiliates in Canada and the United States use Soccer in their names. A few FIFA affiliates have recently "normalized" to using "Football", including:

    Australia's association football governing body changed its name in 2005 from using "soccer" to "football"[102]
    New Zealand's governing body also changed in 2007, saying "the international game is called football."[103]
    Samoa changed from "Samoa Football (Soccer) Federation" to "Football Federation Samoa" in 2009.[104][105]

Popularity

Several of the football codes are the most popular team sports in the world.[7] Globally, association football is played by over 250 million players in over 200 nations,[106] and has the highest television audience in sport,[107] making it the most popular in the world,[108] American football is the most popular sport in the United States,[109] with the annual Super Bowl game accounting for seven of the top eight of the most watched broadcasts in U.S. television history.[110] Australian rules football has the highest spectator attendance of all sports in Australia.[111][112] Similarly, Gaelic football is the most popular sport in Ireland in terms of match attendance,[113] and the All-Ireland Football Final is the most watched event of that nation's sporting year.[114]