dog |
ant |
top 10 words in brain distribution (in article): species animal breed male cat wolf hunt female human dog |
top 10 words in brain distribution (in article): species animal breed male hunt female human size population bird |
top 10 words in brain distribution (not in article): bird lion elephant egg feed deer forest hybrid habitat water |
top 10 words in brain distribution (not in article): cat wolf dog wild lion elephant blade iron tiger steel |
times more probable under dog 30 20 10 6 4 2.5 1.25 1 1.25 2.5 4 6 10 20 30 times more probable under ant (words not in the model) | |
The dog'" ("Canis lupus familiaris",) is a domesticated subspecies of the gray wolf, a member of the Canidae family of the order Carnivora. The term is used for both feral and pet varieties. The domestic dog has been one of the most widely kept working and companion animals in human history. The domestication of the gray wolf took place in a handful of events roughly 15,000 years ago in central Asia. The dog quickly became ubiquitous across culture in all parts of the world, and was extremely valuable to early human settlements. For instance, it is believed that the successful emigration across the Bering Strait might not have been possible without sled dogs. As a result of the domestication process, the dog developed a sophisticated intelligence that includes unparalleled social cognition and a simple theory of mind that is important to their interaction with humans. These social skills have helped the dog to perform in myriad roles, such as hunting, herding, protection, and, more recently, assisting handicapped individuals. Currently, there are estimated to be 400 million dogs in the world. Over the 15,000 year span that the dog had been domesticated, it diverged into only a handful of landraces, groups of similar animals whose morphology and behavior have been shaped by environmental factors and functional roles. Humans did not take an active, intentional role in this process until the last few hundred years. As the modern understanding of genetics developed, humans began to intentionally breed dogs for a wide range of specific traits. Through this process, the dog has developed into hundreds of varied breeds, and shows more behavioral and morphological variation than any other land mammal. For example, height measured to the withers ranges from a few inches in the Chihuahua to a few feet in the Irish Wolfhound; color varies from white through grays (usually called "blue'") to black, and browns from light (tan) to dark ("red" or "chocolate") in a wide variation of patterns; coats can be short or long, coarse-haired to wool-like, straight, curly, or smooth. It is common for most breeds to shed this coat, but non-shedding breeds are also popular. Etymology and related terminology. "Dog" is the common use term that refers to members of the subspecies "Canis lupus familiaris". The term is sometimes used to refer to a wider range of species: it can be used to refer to any mammal belonging to the family Canidae, which includes wolves, foxes, jackals, and coyotes; it can be used to refer to the subfamily of Caninae, or the genus Canis, also often called the "true dogs". Some members of the family have "dog" in their common names, such as the raccoon dog and the African wild dog. A few animals have "dog" in their common names but are not canids, such as the prairie dog and the dog fish. The English word "dog" can be traced back to the Old English "docga", a "powerful breed of canine". The term may derive from Proto-Germanic "*dukkōn", represented in Old English "finger-docce" ("finger-muscle"). Due to the linguistically archaic structure of the word, the term "dog" may ultimately derive from the earliest layer of Proto-Indo-European vocabulary, reflecting the role of the dog as the earliest domesticated animal. The English word "hound", which refers to a specific breed group in English, means "dog" in general in other Germanic languages; it is cognate to German "hund", Dutch "hond", common Scandinavian "hund", and Icelandic "hundur". "Hound" itself is derived from the Proto-Indo-European "*kwon-", which is also the direct root of the Greek κυων (kuōn) and the indirect root of the Latin "canis" through the variant form "*kani-". In breeding circles, a male canine is referred to as a "dog", while a female is called a "bitch". A group of offspring is a "litter". The father of a litter is called the "sire", and the mother is called the "dam". Offspring are generally called "pups" or "puppies" until they are about a year old. The process of birth is "whelping". Taxonomy and evolution. The domestic dog was originally classified as "Canis familiaris" and "Canis familiarus domesticus" by Linnaeus in 1758, and is currently classified as "Canis lupus familiaris", a subspecies of the gray wolf "Canis lupus", by the Smithsonian Institution and the American Society of Mammalogists. Overwhelming evidence from behavior, vocalizations, morphology, and molecular biology led to the contemporary scientific understanding that a single species, the gray wolf, is the common ancestor for all breeds of domestic dogs, however the timeframe and mechanisms by which dogs diverged are controversial. The current consensus among biologists and archaeologists is that no one can be sure when dogs were domesticated. There is conclusive evidence that dogs genetically diverged from their wolf ancestors at least 15,000 years ago but most believe domestication to have occurred much earlier. The evidence cited for an earlier divergence comes from archaeological findings and mitochondrial DNA studies, both of which are inconclusive. The archaeological evidence demonstrates that the domestication of dogs occurred prior to 15,000 years ago. Some genetic evidence indicates that the domestication of dogs from their wolf ancestors began in the late Upper Paleolithic close to the Pleistocene Holocene boundary, between 17,000 and 14,000 years ago. The earliest dog fossils, two large skulls from Russia and a mandible from Germany, date from roughly 14,000 years ago. Their likely ancestor is the large Eurasian wolf ("Canis lupus lupus"). Remains of smaller dogs from Natufian cave deposits in the Middle East have been dated to around 10,000 to 12,000 years ago. There is a great deal of archealogical evidence for dogs throughout Europe and Asia around this period and through the next two thousand years (roughly 8,000 to 10,000 years ago), with fossils uncovered in Germany, the French Alps, and Iraq, and cave paintings in Turkey. DNA studies have provided a wider range of possible divergence dates, from 15,000 to 40,000 years ago, to as much as 100,000 to 140,000 years ago. This evidence depends on a number of assumptions that others claim are violated. Genetic studies are based in comparisons of genetic diversity between species, and depend on a calibration date, such as the wolf-coyote divergence date, which is estimated to be roughly 1 million years ago. If this divergence date is closer to 750,000 or 2 million years ago, then genetic analyses would be interpreted very differently. Furthermore, it is believed that the genetic diversity of wolves has been in decline for the last 200 years, and that the genetic diversity of dogs has been reduced by selective breeding, which could bias DNA analyses to support an earlier divergence. The genetic evidence for the domestication event occurring in East Asia is also subject to violations of assumptions. These conclusions are based on the location of maximal genetic divergence, assumes that hybridization does not occur, and that breeds remain geographically localized. Although these assumptions hold for many species, there is good reason to believe that they do not hold for canines. Genetic analyses indicate all dogs are likely descended from a handful of domestication events with a small number of founding females, although there is evidence that domesticated dogs interbred with local populations of wild wolves on several occasions. Data suggests that dogs first diverged from wolves in East Asia, and that these domesticated dogs then quickly migrated throughout the world, reaching the North American continent around 8000 B.C. The oldest groups of dogs, which show the greatest genetic variability and are the most similar to their wolf ancestors, are primarily Asian and African breeds, including the Basenji, Saluki, Afghan Hound, Tibetan Terrier, Lhasa Apso, Chow Chow, Pekingese, Shar-Pei, Shi Tzu, Akita, Shiba Inu, Alaskan Malamute, Siberian Husky, and Samoyed. Some breeds that were thought to be very old, such as the Pharaoh Hound, Ibizan Hound, and Norwegian Elkhound, are now known to have been recreated more recently. There is a great deal of controversy surrounding the evolutionary framework for the domestication of dogs. At least three early species of the "Homo" genus began spreading out of Africa roughly 400,000 years ago, and thus lived for a considerable period in contact with canine species. Despite this, there is no evidence of any adaptation of these canine species to the presence of the close relatives of modern man. If dogs were domesticated, as believed, roughly 15,000 years ago, the event (or events) would have coincided with a large expansion in human territory and the development of agriculture. This has led some biologists to suggest that one of the forces that led to the domestication of dogs was a shift in human lifestyle in the form of established human settlements. Permanent settlements would have coincided with a greater amount of disposable food and would have created a barrier between wild and anthropogenic canine populations. Biology. Domestic dogs have been selectively bred for millennia for various behaviors, sensory capabilities, and physical attributes. Modern dog breeds show more variation in size, appearance, and behavior than any other domestic animal. Nevertheless, their morphology is based on that of their wild ancestors, gray wolves. Dogs are predators and scavengers, and like many other predatory mammals, the dog has powerful muscles, fused wrist bones, a cardiovascular system that supports both sprinting and endurance, and teeth for catching and tearing. Dogs are highly variable in height and weight. The smallest known dog was a Yorkshire Terrier, who stood only 6.3 cm (2.5 in) at the shoulder, 9.5 cm (3.75 in) in length along the head-and-body, and weighed only 113 grams (4 ounces). The largest known dog was an English Mastiff which weighed 155.6 kg (343 lbs) and was 250 cm (8.2 feet) from the snout to the tail. The tallest dog is a Great Dane that stands 106.7 cm (42.2 in) at the shoulder. Sight. The dog's visual system is engineered to serve the purposes of a hunter. While a dog's visual acuity is poor (that of a poodle's has been estimated to translate to a Snellen rating of 20 75), their visual discrimination for moving objects is very high; dogs have been shown to be able to discriminate between humans (i.e., identifying their owner) from distances up to a mile. As crepuscular hunters, dogs often rely on their vision in low light situations: they have very large pupils, a high density of rods in the fovea, an increased flicker rate, and a tapetum lucidum. The tapetum is a reflective surface behind the retina that reflects light back to give the photoreceptors a second chance to catch the photons. Like most mammals, dogs are dichromats and have color vision equivalent to red-green color blindness in humans. The eyes of different breeds of dogs have different shapes, dimensions, and retina configurations. Many long-nosed breeds have a "visual streak" — a wide foveal region that runs across the width of the retina and gives them a very wide field of excellent vision. Some long-muzzled breeds, particularly the sighthounds, have a field of vision up to 270° (compared to 180° for humans). Short-nosed breeds, on the other hand, have an "area centralis": a central patch with up to three times the density of nerve endings as the visual streak, giving them detailed sight much more like a human's. Some broad-headed breeds with short noses have a field of vision similar to that of humans. Most breeds have good vision, but some show a genetic predisposition for myopia — such as Rottweilers, where one out of every two has been found to be myopic. Hearing. The frequency range of dog hearing is approximately 40 Hz to 60,000 Hz, which means that dogs can detect sounds outside both ends of the human auditory spectrum. Additionally, dogs have ear mobility which allows them to rapidly pinpoint the exact location of a sound. Eighteen or more muscles can tilt, rotate, raise, or lower a dog's ear. A dog can identify a sound's location much faster than a human can, as well as hear sounds at four times the distance. Smell. While the human brain is dominated by a large visual cortex, the dog brain is largely dominated by an olfactory cortex. The olfactory bulb in dogs is roughly forty times bigger than the olfactory bulb in humans, relative to total brain size, with 125 to 220 million smell-sensitive receptors. The bloodhound exceeds this standard with nearly 300 million receptors. Dogs can discriminate odors at concentrations nearly 100 million times lower than humans can. Coat. The coats of domestic dogs are either "double", made up of a coarse topcoat and a soft undercoat, like a wolf, or "single", with the topcoat only. Dogs with double coats tend to originate in colder climates. Domestic dogs often display the remnants of countershading, a common natural camouflage pattern. The general theory of countershading is that an animal that is lit from above will appear lighter on its upper half and darker on its lower half, where it will usually be in its own shade. This is a pattern that predators can learn to watch for. A countershaded animal will have dark coloring on its upper surfaces and light coloring below, which reduces its general visibility. Thus many breeds will have an occasional "blaze", stripe, or "star" of white fur on their chest or underside. Tail. There are many different shapes for dog tails: straight, straight up, sickle, curled, or cork-screw. In some breeds, the tail is traditionally docked to avoid injuries (especially for hunting dogs). In some breeds, puppies can be born with a short tail or no tail at all. This occurs more frequently in those breeds that are frequently docked and thus have no breed standard regarding the tail. Types and breeds. While all dogs are genetically very similar, natural selection and selective breeding have reinforced certain characteristics in certain populations of dogs, giving rise to dog types and dog breeds. Dog types are broad categories based on function, genetics, or characteristics. Dog breeds are groups of animals that possess a set of inherited characteristics that distinguishes them from other animals within the same species. Modern dog breeds are non-scientific classifications of dogs kept by modern kennel clubs. Purebred dogs of one breed are genetically distinguishable from purebred dogs of other breeds, but the means by which kennel clubs classify dogs is unsystematic. Systematic analyses of the dog genome has revealed only four major types of dogs that can be said to be statistically distinct. These include the "old world dogs" (e.g., Malamute and Shar-Pei), "Mastiff"-type (e.g., Labrador Retriever), "herding"-type (e.g., St. Bernard), and "all others" (also called "modern"- or "hunting"-type). Health. Dogs are susceptible to various diseases, ailments, and poisons, some of which can affect humans. To defend against many common diseases, dogs are often vaccinated. Some breeds of dogs are prone to certain genetic ailments such as elbow or hip dysplasia, blindness, deafness, pulmonic stenosis, cleft palate, and trick knees. Two serious medical conditions particularly affecting dogs are pyometra, affecting unspayed females of all types and ages, and bloat, which affects the larger breeds or deep-chested dogs. Both of these are acute conditions, and can kill rapidly. Dogs are also susceptible to parasites such as fleas, ticks, and mites, as well as hookworms, tapeworms, roundworms, and heartworms. Dogs are also vulnerable to some of the same health conditions as humans, including diabetes, dental and heart disease, epilepsy, cancer, hypothyroidism, and arthritis. Mortality. The typical lifespan of dogs varies widely among breeds, but for most the median longevity, the age at which half the dogs in a population have died and half are still alive, ranges from 10 to 13 years. Individual dogs may live well beyond the median of their breed. The breed with the shortest lifespan (among breeds for which there is a questionnaire survey with a reasonable sample size) is the Dogue de Bordeaux, with a median longevity of about 5.2 years, but several breeds, including Miniature Bull Terriers, Bulldogs, Nova Scotia Duck-Tolling Retrievers, Bloodhounds, Irish Wolfhounds, Greater Swiss Mountain Dogs, Great Danes, and Mastiffs, are nearly as short-lived, with median longevities of 6 to 7 years. The longest-lived breeds, including Toy Poodles, Border Terriers, Miniature Dachshunds, Miniature Poodles, and Tibetan Spaniels, have median longevities of 14 to 15 years. The median longevity of mixed breed dogs, taken as an average of all sizes, is one or more years longer than that of purebred dogs when all breeds are averaged. The dog widely reported to be the longest-lived is "Bluey," who died in 1939 and was claimed to be 29.5 years old at the time of his death; however, the Bluey record is anecdotal and unverified. The longest verified records are of dogs living for 24 years. Predation. Although wild dogs, like wolves, are apex predators, they can be killed in territory disputes with wild animals. Furthermore, in areas where both dogs and other large predators live, dogs can be a major food source for big cats or canines. Reports from Croatia indicate that dogs are killed more frequently than sheep. Wolves in Russia apparently limit feral dog populations. In Wisconsin, more compensation has been paid for dog losses than livestock. Some wolf pairs have been reported to prey on dogs by having one wolf lure the dog out into heavy brush where the second animal waits in ambush. In some instances, wolves have displayed an uncharacteristic fearlessness of humans and buildings when attacking dogs, to the extent that they have to be beaten off or killed. Coyotes and big cats have also been known to attack dogs. Leopards in particular are known to have a predilection for dogs, and have been recorded to kill and consume them regardless of the dog's size or ferocity. Tigers in Manchuria, Indochina, Indonesia, and Malaysia, are reputed to kill dogs with the same vigor as leopards. Striped Hyenas are major predators of village dogs in Turkmenistan, India, and the Caucasus. Diet. Despite its descent from wolves, the domestic dog is an omnivore, though it is classified in the order Carnivora. Unlike an obligate carnivore, such as a member of the cat family with its shorter small intestine, a dog is neither dependent on meat-specific protein nor a very high level of protein in order to fulfill its basic dietary requirements. Dogs are able to healthily digest a variety of foods, including vegetables and grains, and can consume a large proportion of these in their diet. In the wild, canines often eat available plants and fruits. Reproduction. In domestic dogs, sexual maturity begins to happen around age six to twelve months for both males and females, although this can be delayed until up to two years old for some large breeds. This is the time at which female dogs will have their first estrous cycle. They will experience subsequent estrous cycles biannually, during which the body prepares for pregnancy. At the peak of the cycle, females will come into estrus, being mentally and physically receptive to copulation. Because the ova survive and are capable of being fertilized for a week after ovulation, it is possible for a female to mate with more than one male. Adolescence for most domestic dogs is around 12 to 15 months, beyond which they are for the most part more adult than puppy. Domestication has selectively bred for higher libido and earlier and more frequent breeding cycles in dogs than in their wild ancestors, and dogs remain reproductively active until old age. Dogs bear their litters roughly 56 to 72 days after fertilization, with an average of 63 days, although the length of gestation can vary. An average litter consists of about six puppies, though this number may vary widely based on the breed of dog. Toy dogs generally produce from one to four puppies in each litter, while much larger breeds may average as many as twelve. Some dog breeds have acquired traits through selective breeding that interfere with reproduction. Male French Bulldogs, for instance, are incapable of mounting the female. For many dogs of this breed, the female must be artificially inseminated in order to reproduce. Spaying and neutering. Neutering refers to the sterilization of animals, usually by removal of the male's testicles or the female's ovaries and uterus, in order to eliminate the ability to procreate and reduce sex drive. Because of the overpopulation of dogs in some countries, animal control agencies, such as the American Society for the Prevention of Cruelty to Animals (ASPCA), advise that dogs not intended for further breeding should be neutered, so that they do not have undesired puppies that may have to be destroyed later. According to the Humane Society of the United States, 3–4 million dogs and cats are put down each year in the United States and many more are confined to cages in shelters because there are many more animals than there are homes. Spaying or castrating dogs helps keep overpopulation down. Local humane societies, SPCAs, and other animal protection organizations urge people to neuter their pets and to adopt animals from shelters instead of purchasing them. Several notable public figures have spoken out against animal overpopulation, including Bob Barker. On his game show, "The Price Is Right", Barker stressed the problem at the end of every episode, saying: "Help control the pet population. Have your pets spayed or neutered." The current host, Drew Carey, makes a similar plea at the conclusion of each episode. Neutering reduces problems caused by hypersexuality, especially in male dogs. Spayed female dogs are less likely to develop some forms of cancer, affecting mammary glands, ovaries, and other reproductive organs. However, neutering increases the risk of urinary incontinence in female dogs, and prostate cancer in males, as well as osteosarcoma, hemangiosarcoma, cruciate ligament rupture, obesity, and diabetes mellitus in either gender. The hormonal changes involved with sterilization can change the animal's personality and metabolism. Recent studies proved that spayed and neutered dogs in general are more aggressive towards people and other dogs, and more fearful and sensitive to touch than dogs than had not been sterilized. Spaying or neutering very young animals ("early-age spay"), can result in increased health concerns later on in life for both sexes. Incontinence in female dogs is made worse by spaying too early. In both males and females, alteration causes changes in hormones during development. This inhibits the natural signals needed for proper body development, leading to larger animals with greater risk for hip dysplasia, osteoporosis, and other joint disorders. Other studies have shown, however, that early-age neutering of male dogs is associated with no major risks compared to neutering at the more traditional age of six months. Behavior and intelligence. Although dogs have been the subject of a great deal of Behaviorist psychology (e.g.,Pavlov's Dog), they do not enter the world with a psychological "blank slate". Rather, dog behavior is affected by genetic factors as well as environmental factors. Domestic dogs exhibit a number of behaviors and predispositions that were inherited from wolves. The grey wolf is a social animal that has evolved a sophisticated means of communication and social structure. The domestic dog has inherited some of these predispositions, but many of the salient characteristics in dog behavior have been largely shaped by selective breeding by humans. Thus, some of these characteristics, such as the dog's highly developed social cognition, are found only in primitive forms in grey wolves. Intelligence. "Intelligence" is an umbrella term that encompasses the faculties involved in a wide range of mental tasks, such as learning, problem-solving, and communication. The domestic dog has a predisposition to exhibit a social intelligence that is uncommon in the animal world. Dogs are capable of learning in a number of ways, such as through simple reinforcement learning (e.g.,classical or operant conditioning) and by observation. Dogs go through a series of stages of cognitive development. They are not born with an understanding of object permanence; the understanding that objects which are not being actively perceived still remain in existence. The understanding of object permanence occurs during as the infant is learning the coordination of secondary circular reactions, as described by Jean Piaget. That is, as the infants learn to interact intentionally with objects around it. For dogs, this occurs at roughly 8 weeks of age. Handlers of working dogs such as herding dogs and sled dogs know that pups learn behaviors quickly by following examples set by experienced dogs. Many of these behaviors are allelomimetic in that they depend on a genetic predisposition to learn and imitate behaviors of other dogs. This form of intelligence is not peculiar to those tasks dogs have been bred to perform, but can be generalized to myriad abstract problems. Adler & Alder demonstrated this by giving Dachshund puppies the task of learning to pull a cart by tugging on an attached piece of ribbon in order to get a reward in the cart. Puppies that watched an experienced dog successfully retrieve the reward in this way learned the task fifteen times faster than those who were left to solve the problem on their own. In addition to learning by example from other dogs, dogs have also been shown to learn by mimicking human behaviors. In another study, puppies were presented with a box, and shown that when a handler pressed a lever, a ball would roll out of the box. The handler then allowed the puppy to play with the ball, making it an intrinsic reward. The pups were then allowed to interact with the box. Roughly three-quarters of the puppies subsequently touched the lever, and over half successfully released the ball, compared to only 6 percent in a control group that did not watch the human manipulate the lever. Furthermore, the ability for dogs to learn by example has been shown to be as effective as operant conditioning. McKinley and Young have demonstrated that dogs show equivalent accuracy and learning times when taught to identify an object by operant conditioning as they do when taught by human example. Their study found that handing an object between experimenters who then use its name in a sentence successfully taught an observing dog each object's name, allowing the dog to subsequently retrieve the item. Dogs also demonstrate sophisticated social cognition by associating behavioral cues with abstract meanings. One such class of social cognition involves the understanding that others are conscious agents. This is called a theory of mind, and is an area in which dogs excel. Research has shown that dogs are capable of interpreting subtle social cues, and appear to recognize when a human or dog's attention is focused on them. To test this, researchers devised a task in which a reward was hidden underneath one of two buckets. The experimenter then attempted to communicate with the dog to indicate the location of the reward by using a wide range of signals: tapping the bucket, pointing to the bucket, nodding to the bucket, or simply looking at the bucket. The results showed that domestic dogs were better than chimpanzees, wolves, and human infants at this task, and even young puppies with limited exposure to humans performed well. Dr. Stanley Coren, an expert on dog psychology, states that these results demonstrated the social cognition of dogs can exceed that of even our closest genetic relatives, and that this capacity is a recent genetic acquisition which distinguishes the dog from its ancestor, the wolf. Studies have also investigated whether dogs engaged in partnered play change their behavior depending on the attention-state of their partner. Those studies showed that play signals were only sent when the dog was holding the attention of its partner. If the partner was distracted, the dog instead engaged in attention-getting behavior before sending a play signal. Dr. Coren has also argued that dogs demonstrate a sophisticated theory of mind by engaging in deception, which he supports with a number of anecdotes, including one example where a dog hid a stolen treat by sitting on it until the rightful owner of the treat left the room. Although this could have been accidental, Coren suggests that the thief understood that the treat's owner would be unable to find the treat if it were out of view. Together, the empirical data and anecdotal evidence points to dogs possessing at least a limited form of theory of mind. Interactions with humans. Domestic dogs inherited a complex social hierarchy and behaviors from their ancestor, the wolf. Dogs are pack animals with a complex set of behaviors related to determining each dog's position in the social hierarchy, and they exhibit various postures and other means of nonverbal communication that reveal their states of mind. These sophisticated forms of social cognition and communication may account for their trainability, playfulness, and ability to fit into human households and social situations, and these attributes have earned dogs a unique relationship with humans despite being potentially dangerous apex predators. Although experts largely disagree over the details of dog domestication, it is agreed that human interaction played a significant role in shaping the subspecies. Shortly after domestication, dogs became ubiquitous in human populations, and spread throughout the world. Emigrants from Siberia likely crossed the Bering Strait with dogs in their company, and some experts suggest that use of sled dogs may have been critical to the success of the waves that entered North America roughly 12,000 years ago. Dogs were an important part of life for the Athabascan population in North America, and were their only domesticated animal. Dogs also carried much of the load in the migration of the Apache and Navajo tribes 1,400 years ago. Use of dogs as pack animals in these cultures often persisted after the introduction of the horse to North America. Work. Dogs have lived and worked with humans in so many roles that they have earned the unique nickname, "man's best friend", a phrase which is used in other languages as well. They have been bred for herding livestock, hunting (e.g.,pointers, hounds), keeping living spaces clear of rats, guarding, helping fishermen with nets, and pulling loads, in addition to their roles as companions. More recently, dogs have taken on a number of roles under the general classification of service dogs, which provide assistance to individuals with physical or mental disabilities. Service dogs include guide dogs, utility dogs, assistance dogs, hearing dogs, and psychological therapy dogs. Some dogs owned by epileptics have even been shown to alert their handler when the handler shows signs of an impending seizure, sometimes well in advance of onset, allowing the owner to seek safety, medication or medical care. Sports and shows. Owners of dogs often enter them in competitions, whether show (breed conformation shows) or sports, including dog racing and dog sledding. Conformation shows, also referred to as breed shows, are a type of dog show in which a judge familiar with a specific dog breed evaluates individual purebred dogs for how well the dogs conform to their established breed type as described in their breed standard. As the breed standard only deals with the externally observable qualities of the dog (such as appearance, movement, and temperament), separately tested-for qualities are not part of the judging in conformation shows (such as tests for ability, health, or any other specific tests for characteristics that cannot be directly observed). Physical characteristics. Compared to equally sized wolves, dogs tend to have 20% smaller skulls, 10% smaller brains, as well as proportionately smaller teeth than other canid species. Dogs require fewer calories to function than wolves. Their diet of human refuse in antiquity made the large brains and jaw muscles needed for hunting unnecessary. It is thought by certain experts that the dog's limp ears are a result of atrophy of the jaw muscles. The skin of domestic dogs tends to be thicker than that of wolves, with some Inuit tribes favoring the former for use as clothing due to its greater resistance to wear and tear in harsh weather. The paws of a dog are half the size of those of a wolf, and their tails tend to curl upwards, another trait not found in wolves. Behavior. Dogs tend to be poorer than wolves at observational learning, being more responsive to instrumental conditioning. Feral dogs show little of the complex social structure or dominance hierarchy present in wolf packs. For dogs, other members of their kind are of no help in locating food items, and are more like competitors. Feral dogs are primarily scavengers, with studies showing that unlike their wild cousins, they are poor ungulate hunters, having little impact on wildlife populations where they are sympatric. However, feral dogs have been reported to be effective hunters of reptiles in the Galapagos islands, and free ranging pet dogs are more prone to predatory behavior toward wild animals. Despite common belief, domestic dogs can be monogamous. Breeding in feral packs can be, but does not have to be restricted to a dominant alpha pair (despite common belief, such things also occur in wolf packs). Male dogs are unusual among canids by the fact that they mostly seem to play no role in raising their puppies, and do not kill the young of other females to increase their own reproductive success. Some sources say that dogs differ from wolves and most other large canid species by the fact that they do not regurgitate food for their young, nor the young of other dogs in the same territory. However, this difference was not observed in all domestic dogs. Regurgitating of food by the females for the young as well as care for the young by the males has been observed in domestic dogs, dingos as well as in other feral or semi-feral dogs. Regurgitating of food by the females and direct choosing of only one mate has been observed even in those semi-feral dogs of direct domestic dog ancestry. Also regurgitating of food by males has been observed in free-ranging domestic dogs. Trainability. Dogs display much greater tractability than tame wolves, and are generally much more responsive to coercive techniques involving fear, aversive stimuli, and force than wolves, which are most responsive toward positive conditioning and rewards. Unlike tame wolves, dogs tend to respond more to voice than hand signals. Although they are less difficult to control than wolves, they can be comparatively more difficult to teach than a motivated wolf. | 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. |