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Ethology is the
scientific Science is a systematic endeavor that builds and organizes knowledge in the form of testable explanations and predictions about the universe. Science may be as old as the human species, and some of the earliest archeological evidence for ...
study of
animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
behaviour, usually with a focus on behaviour under natural conditions, and viewing behaviour as an
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
arily adaptive trait. Behaviourism as a term also describes the scientific and objective study of animal behaviour, usually referring to measured responses to stimuli or to trained behavioural responses in a laboratory context, without a particular emphasis on evolutionary adaptivity. Throughout history, different naturalists have studied aspects of animal behaviour. Ethology has its scientific roots in the work of
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
and of American and German ornithologists of the late 19th and early 20th century, including Charles O. Whitman, Oskar Heinroth, and Wallace Craig. The modern discipline of ethology is generally considered to have begun during the 1930s with the work of Dutch biologist Nikolaas Tinbergen and Austrian biologists
Konrad Lorenz Konrad Zacharias Lorenz (; 7 November 1903 – 27 February 1989) was an Austrian zoologist, ethologist, and ornithologist. He shared the 1973 Nobel Prize in Physiology or Medicine with Nikolaas Tinbergen and Karl von Frisch. He is often regarde ...
and
Karl von Frisch Karl Ritter von Frisch, (20 November 1886 – 12 June 1982) was a German-Austrian ethologist who received the Nobel Prize in Physiology or Medicine in 1973, along with Nikolaas Tinbergen and Konrad Lorenz. His work centered on investigations o ...
, the three recipients of the 1973 Nobel Prize in Physiology or Medicine. Ethology combines laboratory and field science, with a strong relation to some other disciplines such as
neuroanatomy Neuroanatomy is the study of the structure and organization of the nervous system. In contrast to animals with radial symmetry, whose nervous system consists of a distributed network of cells, animals with bilateral symmetry have segregated, defi ...
, ecology, and
evolutionary biology Evolutionary biology is the subfield of biology that studies the evolutionary processes ( natural selection, common descent, speciation) that produced the diversity of life on Earth. It is also defined as the study of the history of life ...
. Ethologists typically show interest in a behavioural process rather than in a particular animal group, and often study one type of behaviour, such as aggression, in a number of unrelated species. Ethology is a rapidly growing field. Since the dawn of the 21st century researchers have re-examined and reached new conclusions in many aspects of animal communication,
emotions Emotions are mental states brought on by neurophysiological changes, variously associated with thoughts, feelings, behavioral responses, and a degree of pleasure or displeasure. There is currently no scientific consensus on a definition. ...
,
culture Culture () is an umbrella term which encompasses the social behavior, institutions, and norms found in human societies, as well as the knowledge, beliefs, arts, laws, customs, capabilities, and habits of the individuals in these groups ...
, learning and sexuality that the scientific community long thought it understood. New fields, such as
neuroethology Neuroethology is the evolutionary and comparative approach to the study of animal behavior and its underlying mechanistic control by the nervous system. It is an interdisciplinary science that combines both neuroscience (study of the nervous syste ...
, have developed. Understanding ethology or animal behaviour can be important in animal training. Considering the natural behaviours of different species or breeds enables trainers to select the individuals best suited to perform the required task. It also enables trainers to encourage the performance of naturally occurring behaviours and the discontinuance of undesirable behaviours.


Etymology

The term ''ethology'' derives from the
Greek language Greek ( el, label= Modern Greek, Ελληνικά, Elliniká, ; grc, Ἑλληνική, Hellēnikḗ) is an independent branch of the Indo-European family of languages, native to Greece, Cyprus, southern Italy ( Calabria and Salento), southe ...
:
ἦθος Ethos ( or ) is a Greek word meaning "character" that is used to describe the guiding beliefs or ideals that characterize a community, nation, or ideology; and the balance between caution, and passion. The Greeks also used this word to refer to ...
, ''
ethos Ethos ( or ) is a Greek word meaning "character" that is used to describe the guiding beliefs or ideals that characterize a community, nation, or ideology; and the balance between caution, and passion. The Greeks also used this word to refer to ...
'' meaning "character" and , ''
-logia ''-logy'' is a suffix in the English language, used with words originally adapted from Ancient Greek ending in ('). The earliest English examples were anglicizations of the French '' -logie'', which was in turn inherited from the Latin '' -logi ...
'' meaning "the study of". The term was first popularized by American myrmecologist (a person who studies ants) William Morton Wheeler in 1902.


History


The beginnings of ethology

Because ethology is considered a topic of biology, ethologists have been concerned particularly with the
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
of behaviour and its understanding in terms of
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
. In one sense, the first modern ethologist was
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
, whose 1872 book '' The Expression of the Emotions in Man and Animals'' influenced many ethologists. He pursued his interest in behaviour by encouraging his protégé
George Romanes George John Romanes FRS (20 May 1848 – 23 May 1894) was a Canadian-Scots evolutionary biologist and physiologist who laid the foundation of what he called comparative psychology, postulating a similarity of cognitive processes and mechanism ...
, who investigated animal learning and intelligence using an anthropomorphic method, anecdotal cognitivism, that did not gain scientific support. Other early ethologists, such as Eugène Marais, Charles O. Whitman, Oskar Heinroth, Wallace Craig and Julian Huxley, instead concentrated on behaviours that can be called instinctive, or natural, in that they occur in all members of a species under specified circumstances. Their beginning for studying the behaviour of a new species was to construct an
ethogram An ethogram is a catalogue or inventory of behaviours or actions exhibited by an animal used in ethology. The behaviours in an ethogram are usually defined to be mutually exclusive and objective, avoiding subjectivity and functional inference as ...
(a description of the main types of behaviour with their frequencies of occurrence). This provided an objective, cumulative database of behaviour, which subsequent researchers could check and supplement.


Growth of the field

Due to the work of
Konrad Lorenz Konrad Zacharias Lorenz (; 7 November 1903 – 27 February 1989) was an Austrian zoologist, ethologist, and ornithologist. He shared the 1973 Nobel Prize in Physiology or Medicine with Nikolaas Tinbergen and Karl von Frisch. He is often regarde ...
and Niko Tinbergen, ethology developed strongly in continental Europe during the years prior to
World War II World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the vast majority of the world's countries—including all of the great powers—forming two opposing ...
. After the war, Tinbergen moved to the
University of Oxford , mottoeng = The Lord is my light , established = , endowment = £6.1 billion (including colleges) (2019) , budget = £2.145 billion (2019–20) , chancellor ...
, and ethology became stronger in the UK, with the additional influence of William Thorpe, Robert Hinde, and Patrick Bateson at the Sub-department of Animal Behaviour of the
University of Cambridge The University of Cambridge is a public collegiate research university in Cambridge, England. Founded in 1209 and granted a royal charter by Henry III in 1231, Cambridge is the world's third oldest surviving university and one of its most pr ...
. In this period, too, ethology began to develop strongly in North America. Lorenz, Tinbergen, and von Frisch were jointly awarded the Nobel Prize in Physiology or Medicine in 1973 for their work of developing ethology. Ethology is now a well-recognized scientific discipline, and has a number of journals covering developments in the subject, such as '' Animal Behaviour'', '' Animal Welfare'', '' Applied Animal Behaviour Science'', ''
Animal Cognition Animal cognition encompasses the mental capacities of non-human animals including insect cognition. The study of animal conditioning and learning used in this field was developed from comparative psychology. It has also been strongly influen ...
'', '' Behaviour'', '' Behavioral Ecology'' and '' Ethology: International Journal of Behavioural Biology''. In 1972, th
International Society for Human Ethology
was founded to promote exchange of knowledge and opinions concerning human behaviour gained by applying ethological principles and methods and published their journal,
The Human Ethology Bulletin
'. In 2008, in a paper published in the journal ''Behaviour'', ethologist Peter Verbeek introduced the term "Peace Ethology" as a sub-discipline of Human Ethology that is concerned with issues of human conflict, conflict resolution, reconciliation, war, peacemaking, and peacekeeping behaviour.


Social ethology and recent developments

In 1972, the English ethologist John H. Crook distinguished comparative ethology from social ethology, and argued that much of the ethology that had existed so far was really comparative ethology—examining animals as individuals—whereas, in the future, ethologists would need to concentrate on the behaviour of social groups of animals and the social structure within them.
E. O. Wilson Edward Osborne Wilson (June 10, 1929 – December 26, 2021) was an American biologist, naturalist, entomologist and writer. According to David Attenborough, Wilson was the world's leading expert in his specialty of myrmecology, the study of an ...
's book '' Sociobiology: The New Synthesis'' appeared in 1975, and since that time, the study of behaviour has been much more concerned with social aspects. It has also been driven by the stronger, but more sophisticated, Darwinism associated with Wilson,
Robert Trivers Robert Ludlow "Bob" Trivers (; born February 19, 1943) is an American evolutionary biologist and sociobiologist. Trivers proposed the theories of reciprocal altruism (1971), parental investment (1972), facultative sex ratio determination (1973), ...
, and W. D. Hamilton. The related development of behavioural ecology has also helped transform ethology. Furthermore, a substantial rapprochement with comparative psychology has occurred, so the modern scientific study of behaviour offers a more or less seamless spectrum of approaches: from
animal cognition Animal cognition encompasses the mental capacities of non-human animals including insect cognition. The study of animal conditioning and learning used in this field was developed from comparative psychology. It has also been strongly influen ...
to more traditional comparative psychology, ethology,
sociobiology Sociobiology is a field of biology that aims to examine and explain social behavior in terms of evolution. It draws from disciplines including psychology, ethology, anthropology, evolution, zoology, archaeology, and population genetics. Within ...
, and behavioural ecology. In 2020, Dr. Tobias Starzak and Professor Albert Newen from the Institute of Philosophy II at the Ruhr University Bochum postulated that animals may have beliefs.


Relationship with comparative psychology

Comparative psychology also studies animal behaviour, but, as opposed to ethology, is construed as a sub-topic of
psychology Psychology is the scientific study of mind and behavior. Psychology includes the study of conscious and unconscious phenomena, including feelings and thoughts. It is an academic discipline of immense scope, crossing the boundaries between ...
rather than as one of
biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary i ...
. Historically, where comparative psychology has included research on animal behaviour in the context of what is known about human psychology, ethology involves research on animal behaviour in the context of what is known about animal
anatomy Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having it ...
, physiology,
neurobiology Neuroscience is the scientific study of the nervous system (the brain, spinal cord, and peripheral nervous system), its functions and disorders. It is a multidisciplinary science that combines physiology, anatomy, molecular biology, developme ...
, and
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
history. Furthermore, early comparative psychologists concentrated on the study of learning and tended to research behaviour in artificial situations, whereas early ethologists concentrated on behaviour in natural situations, tending to describe it as instinctive. The two approaches are complementary rather than competitive, but they do result in different perspectives, and occasionally conflicts of opinion about matters of substance. In addition, for most of the twentieth century, comparative psychology developed most strongly in North America, while ethology was stronger in
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a subcontinent of Eurasia and it is located entirel ...
. From a practical standpoint, early comparative psychologists concentrated on gaining extensive knowledge of the behaviour of very few
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
. Ethologists were more interested in understanding behaviour across a wide range of species to facilitate principled comparisons across taxonomic groups. Ethologists have made much more use of such cross-species comparisons than comparative psychologists have.


Instinct

The Merriam-Webster dictionary defines instinct as "A largely inheritable and unalterable tendency of an organism to make a complex and specific response to environmental stimuli without involving reason".


Fixed action patterns

An important development, associated with the name of Konrad Lorenz though probably due more to his teacher, Oskar Heinroth, was the identification of
fixed action pattern A fixed action pattern is an ethological term describing an instinctive behavioral sequence that is highly stereotyped and species-characteristic. Fixed action patterns are said to be produced by the innate releasing mechanism, a "hard-wired" neura ...
s. Lorenz popularized these as instinctive responses that would occur reliably in the presence of identifiable stimuli called sign stimuli or "releasing stimuli". Fixed action patterns are now considered to be instinctive behavioural sequences that are relatively invariant within the species and that almost inevitably run to completion. One example of a releaser is the beak movements of many bird species performed by newly hatched chicks, which stimulates the mother to regurgitate food for her offspring. Other examples are the classic studies by Tinbergen on the egg-retrieval behaviour and the effects of a " supernormal stimulus" on the behaviour of
graylag geese The greylag goose or graylag goose (''Anser anser'') is a species of large goose in the waterfowl family Anatidae and the type species of the genus '' Anser''. It has mottled and barred grey and white plumage and an orange beak and pink legs. A ...
. One investigation of this kind was the study of the
waggle dance Waggle dance is a term used in beekeeping and ethology for a particular figure-eight dance of the honey bee. By performing this dance, successful foragers can share information about the direction and distance to patches of flowers yielding nect ...
("dance language") in bee communication by
Karl von Frisch Karl Ritter von Frisch, (20 November 1886 – 12 June 1982) was a German-Austrian ethologist who received the Nobel Prize in Physiology or Medicine in 1973, along with Nikolaas Tinbergen and Konrad Lorenz. His work centered on investigations o ...
.


Learning


Habituation

Habituation is a simple form of learning and occurs in many animal taxa. It is the process whereby an animal ceases responding to a stimulus. Often, the response is an innate behaviour. Essentially, the animal learns not to respond to irrelevant stimuli. For example, prairie dogs (''Cynomys ludovicianus'') give alarm calls when predators approach, causing all individuals in the group to quickly scramble down burrows. When prairie dog towns are located near trails used by humans, giving alarm calls every time a person walks by is expensive in terms of time and energy. Habituation to humans is therefore an important adaptation in this context.


Associative learning

Associative learning in animal behaviour is any learning process in which a new response becomes associated with a particular stimulus. The first studies of associative learning were made by Russian physiologist Ivan Pavlov, who observed that dogs trained to associate food with the ringing of a bell would salivate on hearing the bell.


Imprinting

Imprinting enables the young to discriminate the members of their own species, vital for reproductive success. This important type of learning only takes place in a very limited period of time.
Konrad Lorenz Konrad Zacharias Lorenz (; 7 November 1903 – 27 February 1989) was an Austrian zoologist, ethologist, and ornithologist. He shared the 1973 Nobel Prize in Physiology or Medicine with Nikolaas Tinbergen and Karl von Frisch. He is often regarde ...
observed that the young of birds such as
geese A goose ( : geese) is a bird of any of several waterfowl species in the family Anatidae. This group comprises the genera '' Anser'' (the grey geese and white geese) and ''Branta'' (the black geese). Some other birds, mostly related to the she ...
and
chicken The chicken (''Gallus gallus domesticus'') is a domesticated junglefowl species, with attributes of wild species such as the grey and the Ceylon junglefowl that are originally from Southeastern Asia. Rooster or cock is a term for an adu ...
s followed their mothers spontaneously from almost the first day after they were hatched, and he discovered that this response could be imitated by an arbitrary stimulus if the eggs were incubated artificially and the stimulus were presented during a critical period that continued for a few days after hatching.


Cultural learning


Observational learning


Imitation

Imitation is an advanced behaviour whereby an animal observes and exactly replicates the behaviour of another. The National Institutes of Health reported that capuchin monkeys preferred the company of researchers who imitated them to that of researchers who did not. The monkeys not only spent more time with their imitators but also preferred to engage in a simple task with them even when provided with the option of performing the same task with a non-imitator. Imitation has been observed in recent research on chimpanzees; not only did these chimps copy the actions of another individual, when given a choice, the chimps preferred to imitate the actions of the higher-ranking elder chimpanzee as opposed to the lower-ranking young chimpanzee.


Stimulus and local enhancement

There are various ways animals can learn using observational learning but without the process of imitation. One of these is ''stimulus enhancement'' in which individuals become interested in an object as the result of observing others interacting with the object. Increased interest in an object can result in object manipulation which allows for new object-related behaviours by trial-and-error learning. Haggerty (1909) devised an experiment in which a monkey climbed up the side of a cage, placed its arm into a wooden chute, and pulled a rope in the chute to release food. Another monkey was provided an opportunity to obtain the food after watching a monkey go through this process on four occasions. The monkey performed a different method and finally succeeded after trial-and-error. Another example familiar to some cat and dog owners is the ability of their animals to open doors. The action of humans operating the handle to open the door results in the animals becoming interested in the handle and then by trial-and-error, they learn to operate the handle and open the door. In local enhancement, a demonstrator attracts an observer's attention to a particular location. Local enhancement has been observed to transmit foraging information among birds, rats and pigs. The stingless bee ('' Trigona corvina'') uses local enhancement to locate other members of their colony and food resources.


Social transmission

A well-documented example of social transmission of a behaviour occurred in a group of macaques on Hachijojima Island, Japan. The macaques lived in the inland forest until the 1960s, when a group of researchers started giving them potatoes on the beach: soon, they started venturing onto the beach, picking the potatoes from the sand, and cleaning and eating them. About one year later, an individual was observed bringing a potato to the sea, putting it into the water with one hand, and cleaning it with the other. This behaviour was soon expressed by the individuals living in contact with her; when they gave birth, this behaviour was also expressed by their young - a form of social transmission.


Teaching

Teaching is a highly specialized aspect of learning in which the "teacher" (demonstrator) adjusts their behaviour to increase the probability of the "pupil" (observer) achieving the desired end-result of the behaviour. For example, orcas are known to intentionally beach themselves to catch
pinniped Pinnipeds (pronounced ), commonly known as seals, are a widely range (biology), distributed and diverse clade of carnivorous, fin-footed, semiaquatic, mostly marine mammal, marine mammals. They comprise the extant taxon, extant family (biology ...
prey. Mother orcas teach their young to catch pinnipeds by pushing them onto the shore and encouraging them to attack the prey. Because the mother orca is altering her behaviour to help her offspring learn to catch prey, this is evidence of teaching. Teaching is not limited to mammals. Many insects, for example, have been observed demonstrating various forms of teaching to obtain food.
Ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...
s, for example, will guide each other to food sources through a process called "
tandem running Tandem running is a pair movement coordination observed in ants and termites. In ants, tandem running is used for social learning, by which one ant leads another native ant from the nest to the food source it has found. Tandem running is also us ...
," in which an ant will guide a companion ant to a source of food. It has been suggested that the pupil ant is able to learn this route to obtain food in the future or teach the route to other ants. This behaviour of teaching is also exemplified by crows, specifically
New Caledonian crow The New Caledonian crow (''Corvus moneduloides'') is a medium-sized member of the family Corvidae, native to New Caledonia. The bird is often referred to as the 'qua-qua' due to its distinctive call. It eats a wide range of food, including man ...
s. The adults (whether individual or in families) teach their young adolescent offspring how to construct and utilize tools. For example, '' Pandanus'' branches are used to extract insects and other larvae from holes within trees.


Mating and the fight for supremacy

Individual reproduction is the most important phase in the proliferation of individuals or genes within a species: for this reason, there exist complex
mating In biology, mating is the pairing of either opposite- sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. ''Copulation'' is the union of the sex organs of two sexually reprod ...
ritual A ritual is a sequence of activities involving gestures, words, actions, or objects, performed according to a set sequence. Rituals may be prescribed by the traditions of a community, including a religious community. Rituals are characterized ...
s, which can be very complex even if they are often regarded as fixed action patterns. The stickleback's complex mating ritual, studied by Tinbergen, is regarded as a notable example. Often in social life, animals fight for the right to reproduce, as well as social supremacy. A common example of fighting for social and sexual supremacy is the so-called
pecking order In biology, a dominance hierarchy (formerly and colloquially called a pecking order) is a type of social hierarchy that arises when members of animal social groups interact, creating a ranking system. A dominant higher-ranking individual is so ...
among poultry. Every time a group of poultry cohabitate for a certain time length, they establish a pecking order. In these groups, one chicken dominates the others and can peck without being pecked. A second chicken can peck all the others except the first, and so on. Chickens higher in the pecking order may at times be distinguished by their healthier appearance when compared to lower level chickens. While the pecking order is establishing, frequent and violent fights can happen, but once established, it is broken only when other individuals enter the group, in which case the pecking order re-establishes from scratch.


Living in groups

Several animal species, including humans, tend to live in groups. Group size is a major aspect of their social environment. Social life is probably a complex and effective survival strategy. It may be regarded as a sort of symbiosis among individuals of the same species: a
society A society is a group of individuals involved in persistent social interaction, or a large social group sharing the same spatial or social territory, typically subject to the same political authority and dominant cultural expectations. Soci ...
is composed of a group of individuals belonging to the same species living within well-defined rules on food management, role assignments and reciprocal dependence. When biologists interested in
evolution theory Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation t ...
first started examining social behaviour, some apparently unanswerable questions arose, such as how the birth of sterile castes, like in bees, could be explained through an evolving mechanism that emphasizes the reproductive success of as many individuals as possible, or why, amongst animals living in small groups like squirrels, an individual would risk its own life to save the rest of the group. These behaviours may be examples of altruism. Of course, not all behaviours are altruistic, as indicated by the table below. For example, revengeful behaviour was at one point claimed to have been observed exclusively in '' Homo sapiens''. However, other species have been reported to be vengeful including chimpanzees, as well as anecdotal reports of vengeful camels.
Altruistic Altruism is the principle and moral practice of concern for the welfare and/or happiness of other human beings or animals, resulting in a quality of life both material and spiritual. It is a traditional virtue in many cultures and a core asp ...
behaviour has been explained by the gene-centred view of evolution.


Benefits and costs of group living

One advantage of group living can be decreased predation. If the number of predator attacks stays the same despite increasing prey group size, each prey may have a reduced risk of predator attacks through the dilution effect. Further, according to the
selfish herd theory The selfish herd theory states that individuals within a population attempt to reduce their predation risk by putting other conspecifics between themselves and predators. A key element in the theory is the domain of danger, the area of ground in wh ...
, the fitness benefits associated with group living vary depending on the location of an individual within the group. The theory suggests that conspecifics positioned at the centre of a group will reduce the likelihood predations while those at the periphery will become more vulnerable to attack. Additionally, a predator that is confused by a mass of individuals can find it more difficult to single out one target. For this reason, the zebra's stripes offer not only camouflage in a habitat of tall grasses, but also the advantage of blending into a herd of other zebras. In groups, prey can also actively reduce their predation risk through more effective defence tactics, or through earlier detection of predators through increased vigilance. Another advantage of group living can be an increased ability to forage for food. Group members may exchange information about food sources between one another, facilitating the process of resource location. Honeybees are a notable example of this, using the
waggle dance Waggle dance is a term used in beekeeping and ethology for a particular figure-eight dance of the honey bee. By performing this dance, successful foragers can share information about the direction and distance to patches of flowers yielding nect ...
to communicate the location of flowers to the rest of their hive. Predators also receive benefits from hunting in groups, through using better strategies and being able to take down larger prey. Some disadvantages accompany living in groups. Living in close proximity to other animals can facilitate the transmission of parasites and disease, and groups that are too large may also experience greater competition for resources and mates.


Group size

Theoretically, social animals should have optimal group sizes that maximize the benefits and minimize the costs of group living. However, in nature, most groups are stable at slightly larger than optimal sizes. Because it generally benefits an individual to join an optimally-sized group, despite slightly decreasing the advantage for all members, groups may continue to increase in size until it is more advantageous to remain alone than to join an overly full group.


Tinbergen's four questions for ethologists

Niko Tinbergen argued that ethology always needed to include four kinds of explanation in any instance of behaviour: * Function – How does the behaviour affect the animal's chances of survival and reproduction? Why does the animal respond that way instead of some other way? * Causation – What are the stimuli that elicit the response, and how has it been modified by recent learning? * Development – How does the behaviour change with age, and what early experiences are necessary for the animal to display the behaviour? * Evolutionary history – How does the behaviour compare with similar behaviour in related species, and how might it have begun through the process of phylogeny? These explanations are complementary rather than mutually exclusive—all instances of behaviour require an explanation at each of these four levels. For example, the function of eating is to acquire nutrients (which ultimately aids survival and reproduction), but the immediate cause of eating is hunger (causation). Hunger and eating are evolutionarily ancient and are found in many species (evolutionary history), and develop early within an organism's lifespan (development). It is easy to confuse such questions—for example, to argue that people eat because they're hungry and not to acquire nutrients—without realizing that the reason people experience hunger is because it causes them to acquire nutrients.Barrett et al. (2002) Human Evolutionary Psychology. Princeton University Press.


See also

* Anthrozoology * Behavioral ecology *
Cognitive ethology Cognitive ethology is a branch of ethology concerned with the influence of conscious awareness and intention on the behaviour of an animal. Donald Griffin, a zoology professor in the United States, set up the foundations for researches in the cogn ...
* Deception in animals * Human ethology *
List of abnormal behaviours in animals Abnormal behaviour in animals can be defined in several ways. Statistically, abnormal is when the occurrence, frequency or intensity of a behaviour varies statistically significantly, either more or less, from the normal value. This means that th ...


References


Further reading


Burkhardt, Richard W. Jr. "On the Emergence of Ethology as a Scientific Discipline." Conspectus of History 1.7 (1981).


External links

* {{Authority control Ethology Articles containing video clips