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The horse'" ("Equus ferus caballus") is a hoofed (ungulate) mammal, a subspecies of one of seven extant species of the family Equidae. The horse has evolved over the past 45 to 55 million years from a small multi-toed creature into the large, single-toed animal of today. Humans began to domesticate horses around 4000 BC, and their domestication is believed to have been widespread by 3000 BC; by 2000 BC the use of domesticated horses had spread throughout the Eurasian continent. Although most horses today are domesticated, there are still endangered populations of the Przewalski's Horse, the only remaining true wild horse, as well as more common feral horses which live in the wild but are descended from domesticated ancestors. There is an extensive, specialized vocabulary used to describe equine-related concepts, covering everything from anatomy to life stages, size, colors, markings, breeds, locomotion, and behavior. Horses are anatomically designed to use speed to escape predators, and have a well-developed sense of balance and a strong fight-or-flight instinct. Related to this need to flee from predators in the wild is an unusual trait: horses are able to sleep both standing up and lying down. Female horses, called mares, carry their young for approximately 11 months, and a young horse, called a foal, can stand and run shortly following birth. Most domesticated horses begin training under saddle or in harness between the ages of two and four. They reach full adult development by age five, and have an average lifespan of between 25 and 30 years. Horse breeds are loosely divided into three categories based on general temperament: spirited "hot bloods" with speed and endurance; "cold bloods," such as draft horses and some ponies, suitable for slow, heavy work; and "warmbloods," developed from crosses between hot bloods and cold bloods, often focusing on creating breeds for specific riding purposes, particularly in Europe. There are over 300 breeds of horses in the world today, developed for many different uses. Horses and humans interact in many ways, not only in a wide variety of sport competitions and non-competitive recreational pursuits, but also in working activities including police work, agriculture, entertainment, assisted learning and therapy. Horses were historically used in warfare. A wide variety of riding and driving techniques have been developed, using many different styles of equipment and methods of control. Many products are derived from horses, including meat, milk, hide, hair, bone, and pharmaceuticals extracted from the urine of pregnant mares. Humans provide domesticated horses with food, water and shelter, as well as attention from specialists such as veterinarians and farriers. Biology. Horse anatomy is described by a large number of specific terms, as illustrated by the chart to the right. Specific terms also describe various ages, colors and breeds. Age. Depending on breed, management and environment, the domestic horse today has a life expectancy of 25 to 30 years. It is uncommon, but a few animals live into their 40s and, occasionally, beyond. The oldest verifiable record was "Old Billy," a 19th-century horse that lived to the age of 62. In modern times, Sugar Puff, who had been listed in the Guinness Book of World Records as the world's oldest living pony, died in 2007, aged 56. Regardless of a horse's actual birth date, for most competition purposes an animal is considered a year older on January 1 of each year in the northern hemisphere and August 1 in the southern hemisphere. The exception is in endurance riding, where the minimum age to compete is based on the animal's calendar age. A very rough estimate of a horse's age can be made from looking at its teeth. The following terminology is used to describe horses of various ages: In horse racing, the definitions of colt, filly, mare, and stallion may differ from those given above. In the UK, Thoroughbred horse racing defines a colt as a male less than five years old, and a filly as a female less than five years old. In the USA, both Thoroughbred racing and harness racing defines colts and fillies as four years old and younger. Size. The English-speaking world measures the height of horses in hands, abbreviated "h" or "hh," for "hands high," measured at the highest point of an animal's withers, where the neck meets the back, chosen as a stable point of the anatomy, unlike the head or neck, which move up and down; one hand is. Intermediate heights are defined by hands and inches, rounding to the lower measurement in hands, followed by a decimal point and the number of additional inches between 1 and 3. Thus a horse described as "15.2 h," is 15 hands, 2 inches in height. The size of horses varies by breed, but can also be influenced by nutrition. The general rule for cutoff in height between what is considered a horse and a pony at maturity is 14.2 hands. An animal 14.2 h or over is usually considered to be a horse and one less than 14.2 h a pony. However, there are exceptions to the general rule. Some breeds which typically produce individuals both under and over 14.2 h are considered horses regardless of their height. Conversely, some pony breeds may have features in common with horses, and individual animals may occasionally mature at over 14.2 h, but are still considered to be ponies. The distinction between a horse and pony is not simply a difference in height, but takes account of other aspects of "phenotype" or appearance, such as conformation and temperament. Ponies often exhibit thicker manes, tails and overall coat. They also have proportionally shorter legs, wider barrels, heavier bone, shorter and thicker necks, and short heads with broad foreheads. They often have calmer temperaments than horses and also a high level of equine intelligence that may or may not be used to cooperate with human handlers. In fact, small size, by itself, is sometimes not a factor at all. While the Shetland pony stands on average 10 hands high, the Falabella and other miniature horses, which can be no taller than, the size of a medium-sized dog, are classified by their respective registries as very small horses rather than as ponies. Light riding horses such as Arabians, Morgans, or Quarter Horses usually range in height from 14 to 16 hands and can weigh from. Larger riding horses such as Thoroughbreds, American Saddlebreds or Warmbloods usually start at about 15.2 hands and often are as tall as 17 hands, weighing from. Heavy or draft horses such as the Clydesdale, Belgian, Percheron, and Shire are usually at least 16 to 18 hands high and can weigh from about. The largest horse in recorded history was probably a Shire horse named Sampson, who lived during the late 1800s. He stood 21.2½ hands high, and his peak weight was estimated at. The current record holder for the world's smallest horse is Thumbelina, a fully mature miniature horse affected by dwarfism. She is tall and weighs. Colors and markings. Horses exhibit a diverse array of coat colors and distinctive markings, described with a specialized vocabulary. Often, a horse is classified first by its coat color, before breed or sex. Flashy or unusual colors are sometimes very popular, as are horses with particularly attractive markings. Horses of the same color may be distinguished from one another by their markings. The genetics that create many horse coat colors have been identified, although research continues on specific genes and mutations that result in specific color traits. Essentially, all horse colors begin with a genetic base of "red" (chestnut) or "black," with the addition of alleles for spotting, graying, suppression or dilution of color, or other effects acting upon the base colors to create the dozens of possible coat colors found in horses. Horses which are light in color are often misnamed as being "white" horses. A horse that looks pure white is, in most cases, actually a middle-aged or older gray. Grays have black skin underneath their white hair coat (with the exception of small amounts of pink skin under white markings). The only horses properly called white are those with pink skin under a white hair coat, a fairly rare occurrence. There are no truly albino horses, with pink skin and red eyes, as albinism is a lethal condition in horses. Reproduction and development. Pregnancy lasts for approximately 335–340 days and usually results in one foal. Twins are very rare. Colts are carried on average about 4 days longer than fillies. Horses are a precocial species, and foals are capable of standing and running within a short time following birth. Horses, particularly colts, may sometimes be physically capable of reproduction at about 18 months. In practice, individuals are rarely allowed to breed before the age of three, especially females. Horses four years old are considered mature, although the skeleton normally continues to develop until the age of six; the precise time of completion of development also depends on the horse's size, breed, gender, and the quality of care provided by its owner. Also, if the horse is larger, its bones are larger; therefore, not only do the bones take longer to actually form bone tissue, but the epiphyseal plates are also larger and take longer to convert from cartilage to bone. These plates convert after the other parts of the bones, but are crucial to development. Depending on maturity, breed, and the tasks expected, young horses are usually put under saddle and trained to be ridden between the ages of two and four. Although Thoroughbred race horses are put on the track at as young as two years old in some countries, horses specifically bred for sports such as dressage are generally not entered into top-level competition until they are a minimum of four years old, because their bones and muscles are not solidly developed, nor is their advanced training complete. For endurance riding competition, horses are not deemed mature enough to compete until they are a full 60 calendar months (5 years) old. Skeletal system. Horses have a skeleton that averages 205 bones. A significant difference between the horse skeleton, compared to that of a human, is the lack of a collarbone—the horse's front limb system is attached to the spinal column by a powerful set of muscles, tendons and ligaments that attach the shoulder blade to the torso. The horse's legs and hooves are also unique structures. Their leg bones are proportioned differently from those of a human. For example, the body part that is called a horse's "knee" is actually made up of the carpal bones that correspond to the human wrist. Similarly, the hock, contains the bones equivalent to those in the human ankle and heel. The lower leg bones of a horse correspond to the bones of the human hand or foot, and the fetlock (incorrectly called the "ankle") is actually the proximal sesamoid bones between the cannon bones (a single equivalent to the human metacarpal or metatarsal bones) and the proximal phalanges, located where one finds the "knuckles" of a human. A horse also has no muscles in its legs below the knees and hocks, only skin, hair, bone, tendons, ligaments, cartilage, and the assorted specialized tissues that make up the hoof. Hooves. The critical importance of the feet and legs is summed up by the traditional adage, "no foot, no horse". The horse hoof begins with the distal phalanges, the equivalent of the human fingertip or tip of the toe, surrounded by cartilage and other specialized, blood-rich soft tissues such as the laminae. The exterior hoof wall and horn of the sole is made of essentially the same material as a human fingernail. The end result is that a horse, weighing on average, travels on the same bones as a human on tiptoe. For the protection of the hoof under certain conditions, some horses have horseshoes placed on their feet by a professional farrier. The hoof continually grows, and needs to be trimmed (and horseshoes reset, if used) every five to eight weeks. Teeth. Horses are adapted to grazing. In an adult horse, there are 12 incisors, adapted to biting off the grass or other vegetation, at the front of the mouth. There are 24 teeth adapted for chewing, the premolars and molars, at the back of the mouth. Stallions and geldings have four additional teeth just behind the incisors, a type of canine teeth that are called "tushes." Some horses, both male and female, will also develop one to four very small vestigial teeth in front of the molars, known as "wolf" teeth, which are generally removed because they can interfere with the bit. There is an empty interdental space between the incisors and the molars where the bit rests directly on the bars (gums) of the horse's mouth when the horse is bridled. The incisors show a distinct wear and growth pattern as the horse ages, as well as change in the angle at which the chewing surfaces meet. The teeth continue to erupt throughout life as they are worn down by grazing, so a very rough estimate of a horse's age can be made by an examination of its teeth, although diet and veterinary care can affect the rate of tooth wear. Digestion. Horses are herbivores with a digestive system adapted to a forage diet of grasses and other plant material, consumed steadily throughout the day. Therefore, compared to humans, they have a relatively small stomach but very long intestines to facilitate a steady flow of nutrients. A horse will eat of food per day and, under normal use, drink to of water. Horses are not ruminants, so they have only one stomach, like humans, but unlike humans, they can also digest cellulose from grasses due to the presence of a "hind gut" called the cecum, or "water gut," which food goes through before reaching the large intestine. Unlike humans, horses cannot vomit, so digestion problems can quickly cause colic, a leading cause of death. Senses. The horse's senses are generally superior to those of a human. As prey animals, they must be aware of their surroundings at all times. They have the largest eyes of any land mammal, and because their eyes are positioned on the sides of their heads, horses have a range of vision of more than 350°, with approximately 65° of this being binocular (seen with both eyes) and the remaining 285° monocular (seen with only one eye). Horses have excellent day and night vision, but studies indicate that they have two-color, or dichromatic vision; their color vision is somewhat like red-green color blindness in humans. This means that certain colors, especially red and related colors, appear more green. Their hearing is good, and the pinna of each ear can rotate up to 180°, giving the potential for 360° hearing without having to move the head. Their sense of smell, while much better than that of humans, is not their strongest asset; they rely to a greater extent on vision. Horses have a great sense of balance, due partly to their ability to feel their footing and partly to highly developed proprioceptive abilities (the unconscious sense of where the body and limbs are at all times). A horse's sense of touch is well developed. The most sensitive areas are around the eyes, ears and nose. Via touch, horses perceive and respond immediately to changes in their environment, sensing contact as subtle as an insect landing anywhere on the body. Horses have an advanced sense of taste that allows them to sort through grains and grasses to choose what they would most like to eat, and their prehensile lips can easily sort even the smallest grains. Horses generally will not eat poisonous plants. However, there are exceptions and horses will occasionally eat toxic amounts of poisonous plants even when there is adequate healthy food. Movement. All horses move naturally with four basic gaits: the four-beat walk, which averages four miles per hour; the two-beat trot or jog, which averages per hour (faster for harness racing horses); and the leaping gaits known as the canter or lope (a three-beat gait that is per hour), and the gallop. The gallop averages per hour. The world record for a horse galloping over a short, sprint distance is per hour. Besides these basic gaits, some horses perform a two-beat pace, instead of the trot. In addition, there are several four-beat "ambling" gaits that are approximately the speed of a trot or pace, though smoother to ride. These include the lateral slow gait, rack, running walk, and tölt as well as the diagonal fox trot. Ambling gaits are often genetic traits in specific breeds, known collectively as gaited horses. In most cases, gaited horses replace the standard trot with one of the ambling gaits. Behavior. Horses are prey animals with a well-developed fight-or-flight instinct. Their first response to threat is to startle and usually flee, although they are known to stand their ground and defend themselves or their offspring in cases where flight is not possible, or when their young are threatened. They also tend to be curious; when startled, they will often hesitate an instant to ascertain the cause of their fright, and may not always flee from something that they perceive as non-threatening. Through selective breeding, some breeds of horses are quite docile, particularly certain large draft horses. Most light horse riding breeds were developed for speed, agility, alertness and endurance; natural qualities that extend from their wild ancestors. Horses are herd animals, with a clear hierarchy of rank, led by a dominant animal (usually a mare). They are also social creatures who are able to form companionship attachments to their own species and to other animals, including humans. They communicate in various ways, including vocalizations such as nickering or whinnying, mutual grooming, and body language. Many horses will become difficult to manage if they are isolated. Through proper training, it is possible to teach any horse to accept a human as a type of companion, and thus be comfortable away from other horses. When confined with insufficient companionship, exercise or stimulation, individuals may develop stable vices, an assortment of bad habits, mostly psychological in origin, that include wood chewing, wall kicking, "weaving" (rocking back and forth) and other problems. Intelligence and learning. In the past, horses were considered unintelligent, with no abstract thinking ability, unable to generalize, and driven primarily by a herd mentality. However, recent studies show that they perform a number of cognitive tasks on a daily basis, and frequently engage in mental challenges that include food procurement and social system identification. They have also been shown to have good spatial discrimination abilities. Studies have assessed equine intelligence in the realms of problem solving, learning speed, and knowledge retention. Results show that horses excel at simple learning, but also are able to solve advanced cognitive challenges that involve categorization and concept learning. They have been shown to learn from habituation, desensitization, Pavlovian conditioning, and operant conditioning. They respond to and learn from both positive and negative reinforcement. Domesticated horses tend to face greater mental challenges than wild horses, due to living in artificial environments that stifle instinctual behavior while learning tasks that are not natural. Horses are creatures of habit that respond and adapt well to regimentation, and respond best when the same routines and techniques are used consistently. Some trainers believe that "intelligent" horses are reflections of intelligent trainers who effectively use response conditioning techniques and positive reinforcement to train in the style that fits best with an individual animal's natural inclinations. Others who handle horses regularly note that personality also may play a role separate from intelligence in determining how a given animal responds to various experiences. Temperament. Thoroughbred race horses are a "hot blooded" breed. Horses are mammals, and as such are "warm-blooded" creatures, as opposed to reptiles, which are cold-blooded. However, these words have developed a separate meaning in the context of equine terminology, used to describe temperament, not body temperature. For example, the "hot-bloods," such as many race horses, exhibit more sensitivity and energy, while the "cold-bloods," such as most draft breeds, are quieter and calmer. "Hot blooded" breeds include "oriental horses" such as the Akhal-Teke, Barb, Arabian horse and now-extinct Turkoman horse, as well as the Thoroughbred, a breed developed in England from the older oriental breeds. Hot bloods tend to be spirited, bold, and learn quickly. They are bred for agility and speed. They tend to be physically refined—thin-skinned, slim, and long-legged. The original oriental breeds were brought to Europe from the Middle East and North Africa when European breeders wished to infuse these traits into racing and light cavalry horses. The "cold blooded" draft breeds are powerful and heavily-muscled Muscular, heavy draft horses are known as "cold bloods," as they are bred not only for strength, but also to have the calm, patient temperament needed to pull a plow or a heavy carriage full of people. They are sometimes nicknamed "gentle giants." Well-known draft breeds include the Belgian and the Clydesdale. Some, like the Percheron are lighter and livelier, developed to pull carriages or to plow large fields in drier climates. Others, such as the Shire, are slower and more powerful, bred to plow fields with heavy, clay-based soils. The cold-blooded group also includes some pony breeds. "Warmblood" breeds, such as the Trakehner or Hanoverian, developed when European carriage and war horses were crossed with Arabians or Thoroughbreds, producing a riding horse with more refinement than a draft horse, but greater size and more phlegmatic temperament than a lighter breed. Certain pony breeds with warmblood characteristics have been developed for smaller riders. A modern "Warmblood" horse is large, but agile and athletic. Today, the term "Warmblood" refers to a specific subset of sport horse breeds that have dominated the Olympic Games and international FEI competition in dressage and show jumping since the 1970s. Prior to that time, the term "warm blood" often referred to any cross between cold-blooded and hot-blooded breeds. Examples included breeds such as the Irish Draught or the Cleveland Bay. Less often, the term was even used to refer to breeds of light riding horse other than Thoroughbreds or Arabians, such as the Morgan horse. Sleep patterns. Horses are able to sleep both standing up and lying down. In an adaptation from life in the wild, horses are able to enter light sleep by using a "stay apparatus" in their legs, allowing them to doze without collapsing. Horses sleep better when in groups because some animals will sleep while others stand guard to watch for predators. A horse kept alone will not sleep well because its instincts are to keep a constant eye out for danger. Unlike humans, horses do not sleep in a solid, unbroken period of time, but take many short periods of rest. Horses may spend anywhere from four to fifteen hours a day in standing rest, and from a few minutes to several hours lying down. Total sleep time in a day may range from several minutes to a couple of hours, mostly in short intervals of about 15 minutes each. Horses must lie down to reach REM sleep. They only have to lie down for an hour or two every few days to meet their minimum REM sleep requirements. However, if a horse is never allowed to lie down, after several days it will become sleep-deprived, and in rare cases may suddenly collapse as it involuntarily slips into REM sleep while still standing. This condition differs from narcolepsy, although horses may also suffer from that disorder. Taxonomy and evolution. The horse as it is known today adapted by evolution to survive in areas of wide-open terrain with sparse vegetation, surviving in an ecosystem where other large grazing animals, especially ruminants, could not. Horses and other equids are odd-toed ungulates of the order Perissodactyla, a group of mammals that was dominant during the Tertiary period. In the past, this order contained 14 families and many species, but only three families—Equidae (the horse and related species), the tapir and the rhinoceros—containing 18 known species have survived to the present day. The earliest known member of the Equidae family was the "Hyracotherium", which lived between 45 and 55 million years ago, during the Eocene period and had 4 toes on each front foot, and 3 toes on each back foot. The extra toe on the front feet soon disappeared with the "Mesohippus", which lived 32 to 37 million years ago, and by about 5 million years ago, the modern Equus had developed. The extra side toes shrank in size until they have vanished in modern horses. All that remains is a set of small vestigial bones on the leg above the hoof, known informally as ergots, chestnuts, or splint bones. Their legs also lengthened as their toes disappeared and until they were a hoofed animal capable of running at great speed. Over millions of years, equid teeth also evolved from browsing on soft, tropical plants to adapt to browsing of drier plant material, and grazing of tougher plains grasses. Thus the proto-horses changed from leaf-eating forest-dwellers to grass-eating inhabitants of semi-arid regions worldwide, including the steppes of Eurasia and the Great Plains of North America. For reasons not fully understood, "Equus caballus" disappeared from North America around 10,000 years ago, at the end of the last ice age. The "Four Foundations" theory. Modern DNA evidence suggests that domesticated horses evolved from multiple wild populations. Specifically, the "Four Foundations" theory suggests that the modern horse evolved from multiple ancient wild prototypes, each adapted to a given habitat. However, an older theory holds that there was only one type of wild horse, the Tarpan subtype, and all other types diverged in form after domestication to meet human needs. Under the four foundations theory, all types and breeds of horses are thought to have developed from the following base prototypes: Domestication and surviving wild species. Competing theories exist as to the time and place of initial domestication. The earliest evidence for the domestication of the horse comes from Ukraine and dates to approximately 4,000 BC. It is thought that the horse was completely domesticated by 3000 BC, and by 2000 BC there was a sharp increase in the number of horse bones found in human settlements in northwestern Europe, indicating the spread of domesticated horses throughout the continent. Wild species surviving into modern times. A truly wild horse is a species or subspecies which has no ancestors that were ever domesticated. Therefore, most "wild" horses today are actually feral horses, animals that escaped or were turned loose from domestic herds and the descendants of those animals. Only two types of truly wild horses survived into recorded history. One, the Tarpan ("Equus ferus ferus") survived into the historical era, but became extinct in 1887. Its pure genetic line was lost, but three attempts have been made to re-create the Tarpan. In the early 1930s, Berlin Zoo Director Lutz Heck and Heinz Heck of the Munich Zoo began a program that by the 1960s produced the Heck horse. In 1936, Polish university professor Tadeusz Vetulani began a program using Konik horses, and in the mid-1960s Harry Hegard started a program in the United States using feral mustangs and local working ranch horses that has resulted in the Hegardt or Stroebel's Horse. None of the breeding programs were completely successful, although all three resulted in horses with many similarities to the Tarpan. There is only one true wild horse species alive today, the Przewalski's Horse ("Equus ferus przewalskii"). It is a rare Asian animal, also known as the Mongolian Wild Horse; Mongolian people know it as the "taki", and the Kyrgyz people call it a "kirtag". Small wild breeding populations of this animal, named after the Russian explorer Nikolai Przhevalsky, exist in Mongolia. There are also small populations maintained at zoos throughout the world. The species was considered extinct in the wild between 1969 and 1992, but a small breeding population was reestablished in the wild due to the conservation efforts of numerous zoos. Feral populations. Feral horses are born and live in the wild, but are descended from domesticated animals. Many populations of feral horses exist throughout the world. Studies of feral herds have provided useful insights into the behavior of prehistoric horses, as well as greater understanding of the instincts and behaviors that drive horses that live in domesticated conditions. Other modern equids. Besides the horse, there are seven other species of genus "equus" in the equidae family. These are the ass or donkey, "Equus asinus"; the Mountain Zebra, "Equus zebra"; Plains Zebra, "Equus burchelli"; Grévy's Zebra, "Equus grevyi"; the Kiang, "Equus kiang"; and the Kulan, "Equus hemionus", including its subspecies, the Onager, "Equus hemionus onager". Horses can crossbreed with other members of the "equus" genus. The most common hybrid is the mule, a cross between a "jack" (male donkey) and a mare. A related hybrid, a hinny, is a cross between a stallion and a jenny (female donkey). Other hybrids include the zorse, a cross between a zebra and a horse that is bred in Africa and used for trekking on Mount Kenya. With rare exceptions, most hybrids are sterile and cannot reproduce. Breeds. Horse breeds are groups of horses with distinctive characteristics that are transmitted consistently to their offspring, such as conformation, color, performance ability, or disposition. These inherited traits are usually the result of a combination of natural crosses and artificial selection methods aimed at producing horses for specific tasks. Certain breeds are known for certain talents. For example, Standardbreds are known for their speed in harness racing. Some breeds have been developed through centuries of crossings with other breeds, while others, such as Tennessee Walking Horses and Morgans, developed from a single sire from which all current breed members descend. There are more than 300 horse breeds in the world today. Origin of breeds. Modern horse breeds developed in response to a need for "form to function", the necessity to develop certain physical characteristics in order to perform a certain type of work. Thus, powerful but refined breeds such as the Andalusian or the Lusitano developed in the Iberian peninsula as riding horses that also had a great aptitude for dressage, while heavy draft horses such as the Clydesdale and the Shire developed out of a need to perform demanding farm work and pull heavy wagons. Ponies of all breeds originally developed mainly from the need for a working animal that could fulfill specific local draft and transportation needs while surviving in harsh environments. However, by the 20th century, many pony breeds had Arabian and other blood added to make a more refined pony suitable for riding. Other horse breeds developed specifically for light agricultural work, heavy and light carriage and road work, various equestrian disciplines, or simply as pets. Purebreds and registries. Horses have been selectively bred since their domestication. Today, there are over 300 breeds of horses in the world. However, the concept of purebred bloodstock and a controlled, written breed registry only became of significant importance in modern times. Today, the standards for defining and registration of different breeds vary. Sometimes purebred horses are called Thoroughbreds, which is incorrect; "Thoroughbred" is a specific breed of horse, while a "purebred" is a horse (or any other animal) with a defined pedigree recognized by a breed registry. An early example of people who practiced selective horse breeding were the Bedouin, who had a reputation for careful breeding practices, keeping extensive pedigrees of their Arabian horses and placing great value upon pure bloodlines. Though these pedigrees were originally transmitted via an oral tradition, written pedigrees of Arabian horses can be found that date to the 14th century. In the same period of the early Renaissance, the Carthusian monks of southern Spain bred horses and kept meticulous pedigrees of the best bloodstock; the lineage survives to this day in the Andalusian horse. One of the earliest formal registries was General Stud Book for Thoroughbreds, which began in 1791 and traced back to the Arabian stallions imported to England from the Middle East that became the foundation stallions for the breed. Some breed registries have a closed stud book, where registration is based on pedigree, and no outside animals can gain admittance. For example, a registered Thoroughbred or Arabian must have two registered parents of the same breed. Other breeds have a partially closed stud book but still allow certain infusions from other breeds. For example, the modern Appaloosa must have at least one Appaloosa parent, but may also have a Quarter Horse, Thoroughbred, or Arabian parent so long as the offspring exhibits appropriate color characteristics. The Quarter Horse normally requires both parents to be registered Quarter Horses, but allows "Appendix" registration of horses with one Thoroughbred parent, and the horse may earn its way to full registration by completing certain performance requirements. Others, such as most of the warmblood breeds used in sport horse disciplines, have open stud books to varying degrees. While pedigree is considered, outside bloodlines are admitted to the registry if the horses meet the set standard for the registry. These registries usually require a studbook selection process involving judging of an individual animal's quality, performance, and conformation before registration is finalized. A few "registries," particularly some color breed registries, are very open and will allow membership of all horses that meet limited criteria, such as coat color and species, regardless of pedigree or conformation. Breed registries also differ as to their acceptance or rejection of breeding technology. For example, all Jockey Club Thoroughbred registries require that a registered Thoroughbred be a product of a natural mating, so called "live cover". A foal born of two Thoroughbred parents, but by means of artificial insemination or embryo transfer, cannot be registered in the Thoroughbred studbook. On the other hand, since the advent of DNA testing to verify parentage, most breed registries now allow artificial insemination (AI), embryo transfer (ET), or both. The high value of stallions has helped with the acceptance of these techniques because they allow a stallion to breed more mares with each "collection," and greatly reduce the risk of injury during mating. Cloning of horses is highly controversial, and at the present time most mainstream breed registries will not accept cloned horses, though several cloned horses and mules have been produced. Interaction with humans. Around the world, horses play a role within human cultures. Horses are used for leisure activities, sports, and working purposes. The Food and Agriculture Organization (FAO) reports that in 2003, China had the largest number of horses in the world with over 8 million, followed by Mexico (6,260,000), Brazil (5,900,500), the United States (5,300,000), and Argentina (3,655,000). The American Horse Council estimates that horse-related activities have a direct impact on the economy of the United States of over $39 billion, and when indirect spending is considered, the impact is over $102 billion. In a 2004 "poll" conducted by Animal Planet, more than 50,000 viewers from 73 countries voted for the horse as the world's 4th favorite animal. Sport. Historically, equestrians honed their skills through competitions, games and races. Equestrian sports have the dual purpose of providing entertainment for crowds and creating and preserving the excellent horsemanship that was needed in battle. Many sports, such as dressage, eventing and show jumping, had origins in military training, which were focused on control and balance in both the horse and the rider. Other sports, such as rodeo, developed from practical skills such as those needed on working ranches. Sport hunting from horseback evolved from earlier practical hunting techniques. Horse racing, whether the horse was ridden or driven, evolved out of impromptu competitions between riders or drivers. The evolving competitions, requiring ever more demanding and specialized skills from both horse and rider, resulted in the systematic development of specialized breeds and equipment for each sport. The popularity of equestrian sports through the centuries has resulted in the preservation of skills that would otherwise have rapidly disappeared after horses stopped being used in combat. Horses are trained to be ridden or driven in many different sporting events and competitions. Examples include show jumping, dressage, three-day eventing, competitive driving, endurance riding, gymkhana, rodeos and fox hunting. Horse shows, which have their origins in medieval European fairs, are held around the world as venues in which horses are competed, exhibited and sold. They host a huge range of classes, covering all of the mounted and harness disciplines, as well as "In-hand" classes where the horses are led, rather than ridden, to exhibit their conformation. The method of judging classes varies depending on the discipline, but winning awards usually depends on style and ability of both horse and rider. Sports such as polo do not judge the horse itself, but rather use the horse as a partner for human competitors as a necessary part of the game. Although the horse assists this process and requires specialized training to do so, the details of its performance are not judged, only the result of the rider's actions—be it getting a ball through a goal or some other achievement. Examples of these sports of partnership between human and animal also include jousting (reenacting the skills used by medieval knights), in which the main goal is for one rider to unseat the other, and buzkashi, a team game played throughout Central Asia, the aim being to capture a goat carcass while on horseback. Horse racing is an equestrian sport and also a huge international industry, watched in almost every nation of the world. There are three types: "flat" racing; steeplechasing, i.e. racing over jumps; and harness racing, where horses trot or pace while pulling a driver in a small, light cart known as a sulky. A major part of horse racing's economic importance lies in the gambling associated with it. Communication between human and horse is paramount in any equestrian activity; to aid this process horses are usually ridden with a saddle on their backs to assist the rider with balance and positioning, and a bridle or related headgear to assist the rider in maintaining control. Sometimes horses are ridden without a saddle, and occasionally, horses are trained to perform without a bridle or other headgear. Many horses are also driven, which requires a harness, bridle and some type of vehicle. Work. There are certain jobs that horses do very well, and no technology has yet developed that can fully replace them. For example, mounted police horses are still effective for certain types of patrol duties and crowd control. Cattle ranches still require riders on horseback to round up cattle that are scattered across remote, rugged terrain. Search and rescue organizations in some countries depend upon mounted teams to locate people, particularly hikers and children, and to provide disaster relief assistance. Horses can also be used in other areas where it is necessary to avoid vehicular disruption to delicate soil. Examples include areas such as nature reserves. They may also be the only form of transport allowed in wilderness areas. They are also quieter than motorized vehicles. Law enforcement officers such as park rangers or game wardens may use horses for patrols, and horses or mules may also be used for clearing trails or other work in areas of rough terrain where vehicles are less effective. Some land management practices such as cultivating and logging can be efficiently performed with horses. In agriculture, less use of fossil fuels and increased environmental conservation can be seen over time with the use of draft animals such as horses. In forestry, logging can be done with horses and can result in reduced damage to soil structure and less damage to trees due to more selective logging. Although machinery has replaced horses in many parts of the world, an estimated 100 million | A phylogeny of the extant ant subfamilies. "'Ants'" are social insects of the family "'Formicidae'", and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,000 species are classified with upper estimates of about 14,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist. Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because ants appear to operate as a unified entity, collectively working together to support the colony. Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15–25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study. Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have establish themselves in new areas where they may be accidentally introduced. Taxonomy and evolution. The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago. In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant ("Sphecomyrma freyi") that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. "Sphecomyrma" was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface. During the Cretaceous period, only a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20–40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Termites, though sometimes called "white ants", are not ants and belong to the order Isoptera. The termites are actually more closely related to cockroaches and mantids. The fact that ants and termites are both eusocial came about by Convergent evolution. Velvet ants look like large ants, but are wingless female wasps. Etymology. The word "ant" is derived from "ante" of Middle English which is derived from "æmette" and "emmett" of Old English and is related to the Old High German "āmeiza" from which comes "Ameise", the German word for ant. The family name "Formicidae" is derived from the Latin "formīca" ("ant") from which derived Portuguese "formiga", Spanish "hormiga", Romanian "furnică", French "fourmi", etc. Distribution and diversity. Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15–20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates. Ants range in size from. Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies. Morphology. Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarisation. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae. All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species. The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests. Polymorphism. In the colonies of a few ant species, there are physical castes—workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution. Some other species show continuous variation in the size of workers. The smallest and largest workers in "Pheidologeton diversus" show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called "repletes". This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in "Acromyrmex" sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of "Solenopsis invicta" can decide whether the colony will have single or multiple queens. The Australian jack jumper ant ("Myrmecia pilosula"), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects. Development and reproduction. The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes. Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. During the short breeding period, the reproductives, excluding the colony queen, are carried outside where other colonies of similar species are doing the same. Then, all the winged breeding ants take flight. Mating occurs in flight and the males die shortly afterwards. Females of some species mate with multiple males. Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, a process akin to swarming in honeybees. Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size. Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity. Communication. Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path. Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group ("e.g.", foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species. Defence===. Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants ("Paraponera"), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, "Solenopsis" spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people. Trap-jaw ants of the genus "Odontomachus" are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of "Odontomachus bauri" recorded peak speeds of between 126 and 230 h (78 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: "Anochetus", "Orectognathus", and "Strumigenys", plus some members of the Dacetini tribe, which are viewed as examples of convergent evolution. In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or "necrophory", the disposal of dead nest-mates. Oleic acid has been identified as the compound released by dead ants that triggers undertaking behaviour in "Atta mexicana". Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of "Cataulacus muticus", an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Learning. Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of "Temnothorax albipennis" leads a naive nest-mate to newly discovered food by the excruciatingly slow process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close. Controlled experiments with colonies of "Cerapachys biroi" suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care. Nest construction. Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; "Temnothorax albipennis" will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats. The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Weaver ant ("Oecophylla" spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of "Polyrhachis". Food cultivation. Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants ("Atta" and "Acromyrmex") feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called "gongylidia". Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi. Navigation. Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants ("Cataglyphis fortis") use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant ("Cataglyphis bicolor") navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. Locomotion. Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant ("Harpegnathos saltator") is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including "Cephalotes atratus"; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling. Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. "Polyrhachis sokolova", a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests. Cooperation and competition. Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey. Some species (such as "Tetramorium caespitum") attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked. Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like "Strumigenys xenos" are entirely parasitic and do not have workers, but instead rely on the food gathered by their "Strumigenys perplexa" hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in "Wasmannia auropunctata", where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father. Relationships with other organisms. The spider "Myrmarachne plataleoides" (here a female) mimics weaver ants to avoid predators. Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines. Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies. Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae. Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the "Leucoagaricus" or "Leucocoprinus" genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant "Allomerus decemarticulatus" has evolved a three-way association with the host plant "Hirtella physophora" (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey. Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees ("Duroia hirsuta"). This modification of the forest provides the ants with more nesting sites inside the stems of the "Duroia" trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia ("Acacia cornigera") in Central America have hollow thorns that house colonies of stinging ants ("Pseudomyrmex ferruginea") that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the symbiotic ants. In return, the ants obtain food from protein-lipid Beltian bodies. Another example of this type of ectosymbiosis comes from the "Macaranga" tree, which has stems adapted to house colonies of "Crematogaster" ants. Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch. Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites ("Megaponera" and "Termitopone") while a few Cerapachyinae prey on other ants. Some termites form associations with certain ant species to keep away other predatory ant species.The tropical wasp "Mischocyttarus drewseni" coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees ("Trigona" and "Melipona") use chemical defences against ants. Flies in the Old World genus "Bengalia" (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly ("Vestigipoda myrmolarvoidea") live in the nests of ants of the genus "Aenictus" and are cared for by the ants. The fungus "Cordyceps" infects ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode ("Myrmeconema neotropicum") that infects canopy ants ("Cephalotes atratus") causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as "Hyeronima alchorneoides" and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode. South American poison dart frogs in the genus "Dendrobates" feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants. This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparastic, with the birds stealing prey. Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears ("Ursus arctos") have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants. Relationship with humans. Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. On the other hand, ants can become nuisances when they invade buildings, or cause economic losses. In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound. Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include "Paraponera clavata" ("Tocandira") and "Dinoponera" spp. (false "Tocandira"s) of South America and the "Myrmecia" ants of Australia. In South Africa, ants are used to help harvest rooibos ("Aspalathus linearis"), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them "en masse". Up to half a pound (200 g) of seeds can be collected from one ant-heap. As food. Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as "escamoles". They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90 kg) because they are seasonal and hard to find. In the Colombian department of Santander, "hormigas culonas" (roughly interpreted as "large-bottomed ants") "Atta laevigata" are toasted alive and eaten. In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant ("Oecophylla smaragdina") is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, "yum" (ยำ), in a dish called "yum khai mod daeng" (ยำไข่มดแดง) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the "Naturalist in Australia" wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates". In his "First Summer in the Sierra", John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant ("Myrmecocystus"). As pests. Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions. Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound. In science and technology. Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours. The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms. In culture. Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of "Pseudomyrmex" is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance. Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his "A Tramp Abroad". Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel "The Once and Future King". The plot in French entomologist and writer Bernard Werber's "Les Fourmis" science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include "Antz", "A Bug's Life", "The Ant Bully", "The Ant and the Aardvark", "Atom Ant", and there is a comic book superhero called Ant-Man. From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program". Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of "Ender's Game", the Bugs of "Starship Troopers", the giant ants in the film "Them!", and ants mutated into super intelligence in "Phase IV". In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the "Master of Orion" series of games or the ChCht in "Deadlock II". These characters are often credited with a hive mind, a common misconception about ant colonies. |