Ethology

Not to be confused with ethnology or ecology.

Animal Behavior redirects here. For the journal, see
Animal Behaviour (journal). For the band Praxis single,
see Transmutation (Mutatis Mutandis).
Ethology is the scientic and objective study of ani-

joint awardees of the 1973 Nobel Prize in Physiology or

Medicine.[2] Ethology is a combination of laboratory and
eld science, with a strong relation to some other disciplines such as neuroanatomy, ecology, and evolution.
Ethologists are typically interested 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 animals.
Ethology is a rapidly growing eld. Since the dawn of
the 21st century, many aspects of animal communication,
emotions, culture, learning and sexuality that the scientic community long thought it understood have been reexamined, and new conclusions reached. New elds, such
as neuroethology, have developed.
Understanding ethology or animal behaviour can be important in animal training. Considering the natural behaviours of dierent species or breeds enables the trainer
to select the individuals best suited to perform the required task. It also enables the trainer to encourage the
performance of naturally occurring behaviours and also
the discontinuance of undesirable behaviours.[3]

1 Etymology
The term ethology derives from the Greek language:
, ethos meaning character and -, -logia
meaning the study of. The term was rst popularized
by American myrmecologist (a person who studies ants)
William Morton Wheeler in 1902.[4] An earlier, slightly
dierent sense of the term was proposed by John Stuart
Mill in his 1843 System of Logic.[5] He recommended the
development of a new science, ethology, the purpose
of which would be explanation of individual and national
dierences in character, on the basis of associationistic
psychology. This use of the word was never adopted.

A range of animal behaviours

mal behaviour, usually with a focus on behaviour under

natural conditions, and viewing behaviour as an evolutionarily adaptive trait.[1] Behaviourism is a term that also
describes the scientic and objective study of animal behaviour, but it usually refers to the study of trained be- Other words that derive from ethos include ethics[6] and
havioural responses in a laboratory context, and without ethical.
a particular emphasis on evolutionary adaptivity.
Many naturalists have studied aspects of animal behaviour throughout history. Ethology has its scientic
roots in the work of Charles Darwin 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 by
Austrian biologists Konrad Lorenz and Karl von Frisch,

2 Relationship with comparative

psychology
Comparative psychology also studies animal behaviour,
but, as opposed to ethology, is construed as a sub-topic
of psychology rather than as one of biology. Historically,
where comparative psychology has included research on
1

3 HISTORY

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, physiology, neurobiology, and phylogenetic history. Furthermore, early comparative psychologists concentrated on the study of learning and tended to research
behaviour in articial situations, whereas early ethologists
concentrated on behaviour in natural situations, tending
to describe it as instinctive.

animal species were eternal and immutable, created with

a specic purpose, as this seemed the only possible explanation for the incredible variety of living beings and
their surprising adaptation to their habitats.[4]
Jean-Baptiste Lamarck (1744 - 1829) was the rst biologist to describe a complex theory of evolution. His theory
substantially comprised two statements: rst, that animal
organs and behaviour can change according to the way
they are used; and second, that those characteristics can
transmit from one generation to the next (the example of
the girae whose neck becomes longer while trying to
reach the upper leaves of a tree is well-known). The second statement is that every living organism, humans included, tends to reach a greater level of perfection. When
Charles Darwin went to the Galapagos Islands, he was
well aware of Lamarcks theories and was inuenced by
them.

The two approaches are complementary rather than competitive, but they do result in dierent perspectives, and
occasionally conicts 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.
From a practical standpoint, early comparative psychologists concentrated on gaining extensive knowledge of the
behaviour of very few species. Ethologists were more interested in understanding behaviour across a wide range
of species to facilitate principled comparisons across 3.2
taxonomic groups. Ethologists have made much more
use of a truly comparative method than comparative
psychologists have.

3
3.1

Theory of evolution by natural selection and the beginnings of ethology

History
Scala naturae and Lamarcks theories

Jean-Baptiste Lamarck (17441829)

See also: Great Chain of Being

Until the 19th century, the most common theory among
scientists was still the concept of scala naturae, proposed
by Aristotle. According to this theory, living beings were
classied on an ideal pyramid that represented the simplest animals on the lower levels, with complexity increasing progressively toward the top, occupied by human beings. In the Western world of the time, people believed

Charles Darwin (18091882)

Because ethology is considered a topic of biology, ethologists have been concerned particularly with the evolution
of behaviour and the understanding of behaviour in terms
of the theory of natural selection. In one sense, the rst
modern ethologist was Charles Darwin, whose book The
Expression of the Emotions in Man and Animals inuenced many ethologists. He pursued his interest in behaviour by encouraging his protg George Romanes,

3
who investigated animal learning and intelligence using Madingley.[8] In this period, too, ethology began to dean anthropomorphic method, anecdotal cognitivism, that velop strongly in North America.
did not gain scientic support.
Lorenz, Tinbergen, and von Frisch were jointly awarded
Other early ethologists, such as Charles O. Whitman, the Nobel Prize in Physiology or Medicine in 1973 for
Oskar Heinroth, Wallace Craig and Julian Huxley, in- their work of developing ethology.[9]
stead concentrated on behaviours that can be called in- Ethology is now a well-recognised scientic discipline,
stinctive, or natural, in that they occur in all members
and has a number of journals covering developments
of a species under specied circumstances. Their begin- in the subject, such as Animal Behaviour, Animal Welning for studying the behaviour of a new species was to fare, Applied Animal Behaviour Science, Behaviour,
construct an ethogram (a description of the main types Behavioral Ecology and Journal of Ethology. In 1972, the
of behaviour with their frequencies of occurrence).[4] International Society for Human Ethology was founded to
This provided an objective, cumulative data-base of be- promote exchange of knowledge and opinions concerning
haviour, which subsequent researchers could check and human behaviour gained by applying ethological princisupplement.
ples and methods and published their journal, The Human
Ethology Bulletin. In 2008, in a paper published in the
journal Behaviour, ethologist Peter Verbeek introduced
3.3 Social ethology and recent develop- the term Peace Ethology as a sub-discipline of Human
ments
Ethology that is concerned with issues of human conict,
conict resolution, reconciliation, war, peacemaking, and
In 1970, the English ethologist John H. Crook published peacekeeping behaviour.[10]
an important paper in which he distinguished comparative ethology from social ethology, and argued that much Today, along with ethologists, many biologists, zooloof the ethology that had existed so far was really com- gists, primatologists, anthropologists, veterinarians, and
parative ethologyexamining animals as individuals physicians study ethology and other related elds such as
whereas, in the future, ethologists would need to concen- animal psychology, the study of animal social groups, antrate on the behaviour of social groups of animals and the imal cognition and animal welfare science.
social structure within them.
Also in 1970, Robert Ardrey's book The Social Contract:
A Personal Inquiry into the Evolutionary Sources of Order and Disorder was published.[7] The book and study
investigated animal behaviour and then compared human
behaviour to it as a similar phenomenon.

4 Instinct

E. O. Wilson'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, and William Hamilton. The related development of behavioural ecology has also helped transform
ethology. Furthermore, a substantial reapprochement
with comparative psychology has occurred, so the modern scientic study of behaviour oers a more or less
seamless spectrum of approaches: from animal cogniKelp gull chicks peck at red spot on mothers beak to stimulate
tion to more traditional comparative psychology, etholregurgitating reex.
ogy, sociobiology, and behavioural ecology.
The Merriam-Webster dictionary denes instinct as A
largely inheritable and unalterable tendency of an organ3.4 Growth of the eld
ism to make a complex and specic response to environ[11]
Due to the work of Lorenz and Tinbergen, ethology mental stimuli without involving reason.
developed strongly in continental Europe during the
years prior to World War II.[4] After the war, Tinber4.1 Fixed action patterns
gen moved to the University of Oxford, and ethology
became stronger in the UK, with the additional inuence of William Thorpe, Robert Hinde, and Patrick Main article: Fixed action pattern
Bateson at the Sub-department of Animal Behaviour of
the University of Cambridge, located in the village of An important development, associated with the name of

LEARNING

Konrad Lorenz though probably due more to his teacher, 5.3 Imprinting
Oskar Heinroth, was the identication of xed action patterns (FAPs). Lorenz popularized FAPs as instinctive re- Main article: Imprinting (psychology)
sponses that would occur reliably in the presence of iden- Being able to discriminate the members of ones own
tiable stimuli called sign stimuli or releasing stimuli.
FAPs are now considered to be instinctive behavioural
sequences that are relatively invariant within the species
and almost inevitably run to completion.[12]
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
ospring.[13] Other examples are the classic studies by
Tinbergen on the egg-retrieval behaviour and the eects
of a "supernormal stimulus" on the behaviour of graylag
geese.[14][15]
One investigation of this kind was the study of the waggle
dance (dance language) in bee communication by Karl
von Frisch.[16] Lorenz subsequently developed a theory of Example of imprinting in a moose
the evolution of animal communication based on his observations of xed action patterns and the circumstances species is also of fundamental importance for reproductive success. Such discrimination can be based on a numin which they are expressed.
ber of factors. However, this important type of learning
only takes place in a very limited period of time. This
kind of learning is called imprinting,[22] and was a second
5 Learning
important nding of Lorenz. Lorenz observed that the
young of birds such as geese and chickens followed their
mothers spontaneously from almost the rst day after they
5.1 Habituation
were hatched, and he discovered that this response could
be imitated by an arbitrary stimulus if the eggs were incuMain article: Habituation
bated articially and the stimulus were presented during
a critical period that continued for a few days after hatchHabituation is a simple form of learning and occurs in ing.
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 5.4 Cultural learning
respond to irrelevant stimuli. For example, prairie dogs
(Cynomys ludovicianus) give alarm calls when predators Main article: Cultural transmission in animals
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 5.4.1 Observational learning
energy. Habituation to humans is therefore an important
Main article: Observational learning
adaptation in this context.[17][18][19]

5.2

Associative learning

5.4.2 Imitation

Main article: Association (psychology)

Main article: Imitation

Associative learning, in animal behaviour, is any learning process in which a new response becomes associated
with a particular stimulus.[20] The rst studies of associative learning were made by Russian physiologist Ivan
Pavlov.[21] Examples of associative learning include when
a goldsh swims to the water surface when a human is going to feed it, or the excitement of a dog whenever it sees
a collar as a prelude for a walk.

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

5.5

Teaching

same task with a non-imitator.[23] 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.[24]

5.4.3

Stimulus and local enhancement

5.5 Teaching
See also: Animal culture Teaching
Teaching is a highly specialised 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, killer whales are known to intentionally
beach themselves to catch pinniped prey.[31] Mother
killer whales teach their young to catch pinnipeds by
pushing them onto the shore and encouraging them to
attack the prey. Because the mother killer whale is altering her behaviour to help her ospring learn to catch
prey, this is evidence of teaching.[31] Teaching is not limited to mammals. Many insects, for example, have been
observed demonstrating various forms of teaching to obtain food. Ants, for example, will guide each other to
food sources through a process called tandem running,
in which an ant will guide a companion ant to a source of
food.[32] 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.

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.[25] Increased
interest in an object can result in object manipulation
which allows for new object-related behaviours by trialand-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 separate occasions. The monkey performed a dierent method and 6 Mating
and the ght for
nally succeeded after trial-and-error.[26] Another examsupremacy
ple 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 Individual reproduction is the most important phase in
becoming interested in the handle and then by trial-and- the proliferation of individuals or genes within a species:
error, they learn to operate the handle and open the door. for this reason, there exist complex mating rituals, which
In local enhancement, a demonstrator attracts an ob- can be very complex even if they are often regarded as
servers attention to a particular location.[27] Local en- FAPs. The stickleback's complex mating ritual, studied
hancement has been observed to transmit foraging infor- by Tinbergen, is regarded as a notable example of a FAP.
mation among birds, rats and pigs.[28]

5.4.4

Social transmission

See also: Cultural transmission in animals

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.[29] 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.[30]

Often in social life, animals ght for the right to reproduce, as well as social supremacy. A common example
of ghting for social and sexual supremacy is the so-called
pecking order 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 rst, and
so on. Higher level chickens are easily distinguished by
their well-cured aspect, as opposed to lower level chickens. While the pecking order is establishing, frequent and
violent ghts 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.

7 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 is composed of a group of individuals belonging to the same species living within well-dened rules
on food management, role assignments and reciprocal dependence.

TINBERGENS FOUR QUESTIONS FOR ETHOLOGISTS

7.2 Group size

Theoretically, social animals should have optimal group
sizes that maximize the benets and minimize the costs of
group living. However, in nature, most groups are stable
at slightly larger than optimal sizes.[36] Because it generally benets 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.[40]

When biologists interested in evolution theory rst 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.[33] Of course,
not all behaviours are altruistic, as indicated by the ta- 8 Tinbergens four questions for
ble below. For example, revengeful behaviour was at one
ethologists
point claimed to have been observed exclusively in Homo
sapiens. However, other species have been reported to
be vengeful, including reports of vengeful camels[34] and Main article: Tinbergens four questions
chimpanzees.[35]
Altruistic behaviour has been explained by the gene- Niko Tinbergen argued that ethology always needed to
include four kinds of explanation in any instance of becentered view of evolution.
haviour:

7.1

Benets 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
eect.[36] Additionally, a predator that is confused by a
mass of individuals can nd it more dicult to single out
one target. For this reason, the zebras stripes oer not
only camouage in a habitat of tall grasses, but also the
advantage of blending into a herd of other zebras.[37] In
groups, prey can also actively reduce their predation risk
through more eective defense tactics, or through earlier
detection of predators through increased vigilance.[36]
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.[36] Honeybees
are a notable example of this, using the waggle dance to
communicate the location of owers to the rest of their
hive.[38] Predators also receive benets from hunting in
groups, through using better strategies and being able to
take down larger prey.[36]
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.[39]

Function How does the behaviour aect the animals 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 modied 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 exclusiveall 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 organisms lifespan (development). It is easy to confuse such
questionsfor example, to argue that people eat because
they're hungry and not to acquire nutrientswithout realizing that the reason people experience hunger is because
it causes them to acquire nutrients.[41]

List of ethologists

[14] Tinbergen, N. (1951). The Study of Instinct. Oxford University Press, New York.

People who have made notable contributions to ethology [15] Tinbergen, N. (1953). The Herring Gulls World. Collins,
(many listed here are actually comparative psychologists):
London.

10

Gallery of videos

Play media
Food manipulation behaviour by fox squirrels.

11

See also

12

References

[1] Denition of ETHOLOGY. Merriam-Webster. Retrieved 30 October 2012. 2 : the scientic and objective
study of animal behaviour especially under natural conditions
[2] Nobel Prize page for 1973 Medicine Award to Tinbergen,
Lorenz, and von Frisch for contributions in ethology
[3] McGreevy P and Boakes R,"'Carrots and sticks: Principles of animal training'".(Sydney: Sydney University
Press., 2011), pp. xi-23.

[16] Buchmann, Stephen (2006). Letters from the Hive: An Intimate History of Bees, Honey, and Humankind. Random
House of Canada. p. 105. ISBN 978-0-553-38266-2.
[17] M.D.Breed (2001). Habituation. Retrieved 9 September 2014.
[18] Keil, Frank C.; Robert Andrew Wilson (2001). The MIT
encyclopedia of the cognitive sciences. MIT Press. p. 184.
ISBN 978-0-262-73144-7.
[19] Bouton, M.E. (2007). Learning and behavior: A contemporary synthesis. MA, Sinauer: Sunderland.
[20] Associative learning. Encyclopdia Britannica. Retrieved 9 September 2014.
[21] Hudmon, Andrew (2005). Learning and memory. Infobase Publishing. p. 35. ISBN 978-0-7910-8638-4.
[22] Mercer, Jean (2006). Understanding attachment: parenting, child care, and emotional development. Greenwood
Publishing Group. p. 19. ISBN 978-0-275-98217-1.
[23] Imitation Promotes Social Bonding in Primates, August
13, 2009 News Release - National Institutes of Health
(NIH)". Nih.gov. 2009-08-13. Retrieved 2011-11-08.

[4] Matthews, Janice R.; Robert W. Matthews (2009). Insect

Behaviour. Springer. p. 13. ISBN 978-90-481-2388-9.

[24] http://www.plosone.org/article/info%3Adoi%2F10.
1371%2Fjournal.pone.0010625#pone-0010625-g003

[5] Bourg, Julian (2007). From revolution to ethics: May 1968

and contemporary French thought. McGill-Queens Press
- MQUP. p. 155. ISBN 978-0-7735-3199-4.

[25] Spence, K.W. (1937). Experimental studies of learning and higher mental processes in infra-human primates.
Psychological Bulletin, 34: 806-850

[6] Janes, Sharyn; Karen Saucier Lundy (2009). Community

Health Nursing: Caring For The Publics Health. Jones &
Bartlett Learning. p. 251. ISBN 978-0-7637-1786-5.

[26] Haggerty, M.E., (1909). Imitation in monkeys. Journal

of Comparative Neurology and Psychology, 19: 337-455

[7] Ardrey, Robert (1970). The Social Contract: A Personal

Inquiry into the Evolutionary Sources of Order and Disorder. Atheneum.
[8] Bateson, P. (1991). The Development and Integration of
Behaviour: Essays in Honour of Robert Hinde. Cambridge
University Press. p. 479. ISBN 978-0-521-40709-0.
[9] Encyclopaedia Britannica (1975). Yearbook of science
and the future. p. 248.
[10] Verbeek, Peter. (2008) Peace Ethology. Behaviour 145,
1497-1524
[11] Instinct. Merriam-Webster Dictionary.
[12] Campbell, N. A. (1996). Chapter 50. Biology (4 ed.).
Benjamin Cummings, New York. ISBN 0-8053-1957-3.
[13] Bernstein, W M (2011). A Basic Theory of Neuropsychoanalysis. Karnac Books. p. 81. ISBN 978-1-85575-8094.

[27] Hoppitt, W. and Laland, K.N. (2013). Social Learning: An Introduction to Mechanisms, Methods, and Models.
Princeton University Press. ISBN 978-1-4008-4650-4.
[28] Galef, B. G.; Giraldeau, L.-A. (2001). Social inuences on foraging in vertebrates: Causal mechanisms and
adaptive functions. Animal Behaviour 61 (1): 315.
doi:10.1006/anbe.2000.1557. PMID 11170692.
[29] Wilson, Edward O. (2000). Sociobiology: the new synthesis. Harvard University Press. p. 170. ISBN 978-0-67400089-6.
[30] Japanese Macaque - Macaca fuscata.
biomes.org. Retrieved 2011-11-08.

Blueplanet-

[31] Rendell, L. and Whitehead, H. (2001). Culture in whales

and dolphins. Behavioral and Brain Sciences. 24: 309-24
[32] Hoppitt, W.J., G.R. Brown, R. Kendal, L. Rendell, A.
Thornton, M,M, Webster, and K,N, Laland, (2008).
Lessons from animal teaching. Trends in Ecology & Evolution, 23: 486-93

14

EXTERNAL LINKS

[33] Cummings, Mark; Carolyn Zahn-Waxler; Ronald Iannotti

(1991). Altruism and aggression: biological and social origins. Cambridge University Press. p. 7. ISBN 978-0521-42367-0.

Ethology in the world of the horse whisperers and

new masters

[34] The Ape and the Sushi Master. Books.google.com. 200112-27. ISBN 978-0-465-04176-3. Retrieved 2011-1108.

Center for Avian Cognition University of Nebraska

(Alan Kamil, Alan Bond)

[35] Beyond Revenge. Books.google.com. 2008-04-11. ISBN

978-0-7879-7756-6. Retrieved 2011-11-08.
[36] Davies, Nicholas B., Krebs, John R., and West, Stuart A.
An Introduction to Behavioral Ecology, 4th ed. Oxford:
Wiley-Blackwell, 2012.
[37] How do a zebras stripes act as camouage?" 15
April 2008. HowStuWorks.com. Accessed 22 April
2014.
<http://animals.howstuffworks.com/mammals/
question454.htm>
[38] Riley, J.; Greggers, U.; Smith, A.; Reynolds, D.; Menzel,
R. (2005) The ight paths of honeybees recruited by the
waggle dance. Nature, 435 (7039): 205207. Bibcode:
2005Natur.435..205R. doi:10.1038/nature03526
[39] A Look at Animal Social Groups. Rathads, Triana. Science 360. R. R. Donnelly, 29 Aug. 2007. Web. Retrieved
22 Apr. 2014. <http://www.sciences360.com/index.php/
a-look-at-animal-social-groups-6-23511/>.
[40] Sibley, R.M. (1983) Optimal group size is unstable. Animal Behavior, 31: 947-948.
[41] Barrett et al. (2002) Human Evolutionary Psychology.
Princeton University Press.

13

Further reading

Burkhardt, Richard W. Jr. On the Emergence of

Ethology as a Scientic Discipline. Conspectus of
History 1.7 (1981).
Klein, Z. (2000). The ethological approach to the
study of human behavior (PDF). Neuroendocrinology Letters (21): 477481.
Karen Shanor and Jagmeet Kanwal: Bats Sing, Mice
Giggle: Revealing the Secret Lives of Animals, Icon
(2009). 'Accessible to the lay reader and acceptable
to the scientic community' (The Daily Telegraph),
10 October 2009.

14

External links

Konrad Lorenz Institute for Evolution and Cognitive

Research
Center for the Integrative Study of Animal behaviour
Applied Ethology

Abstracts of the XXIX Ethological Conference

Multimedia site about Applied Animal Behaviour

topics by Michigan State University professor
Diagrams on Tinbergens four questions
The Four Areas of Biology
The Four Areas of Biology AND levels of inquiry

15

Text and image sources, contributors, and licenses

15.1

Text

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15.2

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15

TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

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