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Animal navigation

animal navigation, animal navigation national geographic
Animal navigation is the ability of many animals to find their way accurately without maps or instruments Birds such as the Arctic tern, insects such as the monarch butterfly and fish such as the salmon regularly migrate thousands of miles to and from their breeding grounds, and many other species navigate effectively over shorter distances

Dead reckoning, navigating from a known position using only information about one's own speed and direction, was suggested by Charles Darwin in 1873 as a possible mechanism In the 20th century, Karl von Frisch showed that honey bees can navigate by the sun, by the polarization pattern of the blue sky, and by the earth's magnetic field; of these, they rely on the sun when possible William Tinsley Keeton showed that homing pigeons could similarly make use of a range of navigational cues, including the sun, earth's magnetic field, olfaction and vision Ronald Lockley demonstrated that a species of small seabird, the Manx shearwater, could orient themselves and fly home at full speed, when released far from home, provided either the sun or the stars were visible

Several species of animal can integrate cues of different types to orient themselves and navigate effectively Insects and birds are able to combine learned landmarks with sensed direction from the earth's magnetic field or from the sky to identify where they are and so to navigate Internal 'maps' are often formed using vision, but other senses including olfaction and echolocation may also be used

The ability of wild animals to navigate may be adversely affected by products of human activity For example, there is evidence that pesticides may interfere with bee navigation, and that lights may harm turtle navigation


  • 1 Early research
  • 2 Mechanisms
    • 21 Remembered landmarks
    • 22 Orientation by the sun
    • 23 Orientation by the night sky
    • 24 Orientation by polarised light
    • 25 Magnetoreception
    • 26 Olfaction
    • 27 Other senses
  • 3 Path integration
  • 4 Effects of human activity
  • 5 See also
  • 6 Notes
  • 7 References
  • 8 Sources
  • 9 External links

Early research

Karl von Frisch 1953 discovered that honey bee workers can navigate, and indicate the range and direction to food to other workers with a waggle dance

In 1873, Charles Darwin wrote a letter to Nature magazine, arguing that animals including man have the ability to navigate by dead reckoning, even if a magnetic 'compass' sense and the ability to navigate by the stars is present:

With regard to the question of the means by which animals find their way home from a long distance, a striking account, in relation to man, will be found in the English translation of the Expedition to North Siberia, by Von Wrangell He there describes the wonderful manner in which the natives kept a true course towards a particular spot, whilst passing for a long distance through hummocky ice, with incessant changes of direction, and with no guide in the heavens or on the frozen sea He states but I quote only from memory of many years standing that he, an experienced surveyor, and using a compass, failed to do that which these savages easily effected Yet no one will suppose that they possessed any special sense which is quite absent in us We must bear in mind that neither a compass, nor the north star, nor any other such sign, suffices to guide a man to a particular spot through an intricate country, or through hummocky ice, when many deviations from a straight course are inevitable, unless the deviations are allowed for, or a sort of "dead reckoning" is kept All men are able to do this in a greater or less degree, and the natives of Siberia apparently to a wonderful extent, though probably in an unconscious manner This is effected chiefly, no doubt, by eyesight, but partly, perhaps, by the sense of muscular movement, in the same manner as a man with his eyes blinded can proceed and some men much better than others for a short distance in a nearly straight line, or turn at right angles, or back again The manner in which the sense of direction is sometimes suddenly disarranged in very old and feeble persons, and the feeling of strong distress which, as I know, has been experienced by persons when they have suddenly found out that they have been proceeding in a wholly unexpected and wrong direction, leads to the suspicion that some part of the brain is specialised for the function of direction

Later in 1873, Joseph John Murphy replied to Darwin, writing back to Nature with a description of how he, Murphy, believed animals carried out dead reckoning, by what is now called inertial navigation:

If a ball is freely suspended from the roof of a railway carriage it will receive a shock sufficient to move it, when the carriage is set in motion: and the magnitude and direction of the shock … will depend on the magnitude and direction of the force with which the carriage begins to move … … every change in … the motion of the carriage … will give a shock of corresponding magnitude and direction to the ball Now, it is conceivably quite possible, though such delicacy of mechanism is not to be hoped for, that a machine should be constructed … for registering the magnitude and direction of all these shocks, with the time at which each occurred … from these data the position of the carriage … might be calculated at any moment

Karl von Frisch 1886–1982 studied the European honey bee, demonstrating that bees can recognize a desired compass direction in three different ways: by the sun, by the polarization pattern of the blue sky, and by the earth’s magnetic field He showed that the sun is the preferred or main compass; the other mechanisms are used under cloudy skies or inside a dark beehive

William Tinsley Keeton 1933–1980 studied homing pigeons, showing that they were able to navigate using the earth's magnetic field, the sun, as well as both olfactory and visual cues

Donald Griffin 1915–2003 studied echolocation in bats, demonstrating that it was possible and that bats used this mechanism to detect and track prey, and to "see" and thus navigate through the world around them

Ronald Lockley 1903–2000, among many studies of birds in over fifty books, pioneered the science of bird migration He made a twelve-year study of shearwaters such as the Manx shearwater, living on the remote island of Skokholm These small seabirds make one of the longest migrations of any bird—10,000 kilometres—but return to the exact nesting burrow on Skokholm year after year This behaviour led to the question of how they navigated


Lockley began his book Animal Navigation with the words:

How do animals find their way over apparently trackless country, through pathless forests, across empty deserts, over and under featureless seas They do so, of course, without any visible compass, sextant, chronometer or chart

Many mechanisms have been proposed for animal navigation: there is evidence for a number of them Investigators have often been forced to discard the simplest hypotheses - for example, some animals can navigate on a dark and cloudy night, when neither landmarks nor celestial cues like sun, moon, or stars are visible The major mechanisms known or hypothesized are described in turn below

Remembered landmarks

Animals including mammals, birds and insects such as bees and wasps Ammophila and Sphex, are capable of learning landmarks in their environment, and of using these in navigation

Orientation by the sun

Main article: Sun compass in animals The sandhopper, Talitrus saltator, uses the sun and its internal clock to determine direction

Some animals can navigate using celestial cues such as the position of the sun Since the sun moves in the sky, navigation by this means also requires an internal clock Many animals depend on such a clock to maintain their circadian rhythm Animals that use sun compass orientation are fish, birds, sea-turtles, butterflies, bees, sandhoppers, reptiles, and ants

When sandhoppers such as Talitrus saltator are taken up a beach, they easily find their way back down to the sea It has been shown that this is not simply by moving downhill or towards the sight or sound of the sea A group of sandhoppers were acclimatised to a day/night cycle under artificial lighting, whose timing was gradually changed until it was 12 hours out of phase with the natural cycle Then, the sandhoppers were placed on the beach in natural sunlight They moved away from the sea, up the beach The experiment implied that the sandhoppers use the sun and their internal clock to determine their heading, and that they had learnt the actual direction down to the sea on their particular beach

Experiments with Manx shearwaters showed that when released "under a clear sky" far from their nests, the seabirds first oriented themselves and then flew off in the correct direction But if the sky was overcast at the time of release, the shearwaters flew around in circles

Monarch butterflies use the sun as a compass to guide their southwesterly autumn migration from Canada to Mexico

Orientation by the night sky

In a pioneering experiment, Lockley showed that warblers placed in a planetarium showing the night sky oriented themselves towards the south; when the planetarium sky was then very slowly rotated, the birds maintained their orientation with respect to the displayed stars Lockley observes that to navigate by the stars, birds would need both a "sextant and chronometer": a built-in ability to read patterns of stars and to navigate by them, which also requires an accurate time-of-day clock

In 2003, the African dung beetle Scarabaeus zambesianus was shown to navigate using polarization patterns in moonlight, making it the first animal known to use polarized moonlight for orientation In 2013, it was shown that dung beetles can navigate when only the Milky Way or clusters of bright stars are visible, making dung beetles the only insects known to orient themselves by the galaxy

Orientation by polarised light

Rayleigh sky model shows how polarization of light can indicate direction to bees Main article: Rayleigh sky model

Some animals, notably insects such as the honey bee, are sensitive to the polarisation of light Honey bees can use polarized light on overcast days to estimate the position of the sun in the sky, relative to the compass direction they intend to travel Karl von Frisch's work established that bees can accurately identify the direction and range from the hive to a food source typically a patch of nectar-bearing flowers A worker bee returns to the hive and signals to other workers the range and direction relative to the sun of the food source by means of a waggle dance The observing bees are then able to locate the food by flying the implied distance in the given direction, though other biologists have questioned whether they necessarily do so, or are simply stimulated to go and search for food However, bees are certainly able to remember the location of food, and to navigate back to it accurately, whether the weather is sunny in which case navigation may be by the sun or remembered visual landmarks or largely overcast when polarised light may be used


The homing pigeon can quickly return to its home, using cues such as the earth's magnetic field to orient itself Main article: Magnetoreception

Some animals, including mammals such as blind mole rats Spalax and birds such as pigeons, are sensitive to the earth's magnetic field

Homing pigeons use magnetic field information with other navigational cues Pioneering researcher William Keeton showed that time-shifted homing pigeons could not orient themselves correctly on a clear sunny day, but could do so on an overcast day, suggesting that the birds prefer to rely on the direction of the sun, but switch to using a magnetic field cue when the sun is not visible This was confirmed by experiments with magnets: the pigeons could not orient correctly on an overcast day when the magnetic field was disrupted


Returning salmon may use olfaction to identify the river in which they developed

Olfactory navigation has been suggested as a possible mechanism in pigeons Papi's 'mosaic' model argues that pigeons build and remember a mental map of the odours in their area, recognizing where they are by the local odour Wallraff's 'gradient' model argues that there is a steady, large-scale gradient of odour which remains stable for long periods If there were two or more such gradients in different directions, pigeons could locate themselves in two dimensions by the intensities of the odours However it is not clear that such stable gradients exist Papi did find evidence that anosmic pigeons unable to detect odours were much less able to orient and navigate than normal pigeons, so olfaction does seem to be important in pigeon navigation However, it is not clear how olfactory cues are used

Olfactory cues may be important in salmon, which are known to return to the exact river where they hatched Lockley reports experimental evidence that fish such as minnows can accurately tell the difference between the waters of different rivers Salmon may use their magnetic sense to navigate to within reach of their river, and then use olfaction to identify the river at close range

Other senses

Biologists have considered other senses that may contribute to animal navigation Many marine animals such as seals are capable of hydrodynamic reception, enabling them to track and catch prey such as fish by sensing the disturbances their passage leaves behind in the water Marine mammals such as dolphins, and many species of bat, are capable of echolocation, which they use both for detecting prey and for orientation by sensing their environment

Path integration

Path integration sums the vectors of distance and direction travelled from a start point to estimate current position, and so the path back to the start Main article: Path integration

Dead reckoning, in animals usually known as path integration, means the putting together of cues from different sensory sources within the body, without reference to visual or other external landmarks, to estimate position relative to a known starting point continuously while travelling on a path that is not necessarily straight Seen as a problem in geometry, the task is to compute the vector to a starting point by adding the vectors for each leg of the journey from that point

Since Darwin's On the Origins of Certain Instincts quoted above in 1873, path integration has been shown to be important to navigation in animals including ants, rodents and birds When vision and hence the use of remembered landmarks is not available, such as when animals are navigating on a cloudy night, in the open ocean, or in relatively featureless areas such as sandy deserts, path integration must rely on idiothetic cues from within the body

Studies by Wehner in the Sahara desert ant Cataglyphis bicolor demonstrate effective path integration to determine directional heading by polarized light or sun position and to compute distance by monitoring leg movement or optical flow

Path integration in mammals makes use of the vestibular organs, which detect accelerations in the three dimensions, together with motor efference, where the motor system tells the rest of the brain which movements were commanded, and optic flow, where the visual system signals how fast the visual world moves past the eyes Information from other senses such as echolocation and magnetoreception may also be integrated in certain animals The hippocampus is the part of the brain that integrates linear and angular motion to encode a mammal's relative position in space

David Redish states that "The carefully controlled experiments of Mittelstaedt and Mittelstaedt 1980 and Etienne 1987 have demonstrated conclusively that is a consequence of integrating internal cues from vestibular signals and motor efferent copy"

Effects of human activity

Neonicotinoid pesticides may impair the ability of bees to navigate Bees exposed to low levels of thiamethoxam were less likely to return to their colony, to an extent sufficient to compromise a colony's survival

Light pollution attracts and disorients photophilic animals, those that follow light For example, hatchling sea turtles follow bright light, particularly bluish light, altering their navigation Disrupted navigation in moths can easily be observed around bright lamps on summer nights Insects gather around these lamps at high densities instead of navigating naturally

See also

  • Animal migration
  • Salmon run


  1. ^ The book was A Journey on the Northern Coast of Siberia and the Icy Sea 2 vols, London, 1841 Wrangel is variously spelt Vrangel or Wrangell
  2. ^ JJ Murphy d 1894, of County Antrim, was treasurer and then president of the Belfast Literary Society He attempted to harmonise evolution and religion, publishing a book The Scientific Bases of Faith in 1872
  3. ^ A diagram of the experimental apparatus is available from JEB


  1. ^ Dingle, Hugh; Drake, V Alistair 2007 "What is migration" BioScience 57: 113–121 doi:101641/B570206 
  2. ^ a b Darwin, Charles 24 April 1873 "Origin of Certain Instincts" Nature 7 179: 417–418 doi:101038/007417a0 
  3. ^ Murphy, JJ Instinct: a Mechanical Analogy Nature 7:483 24 April 1873 doi:101038/007483b0
  4. ^ a b von Frisch, 1953, pages 93–96
  5. ^ Keeton, 1974
  6. ^ a b Yoon, Carol Kaesuk Donald R Griffin, 88, Dies; Argued Animals Can Think, The New York Times, November 14, 2003 Retrieved 27 February 2012
  7. ^ Ronald Lockley, 1942
  8. ^ a b Lockley, 1967 pages 114–117
  9. ^ Lockley, 1967 page 9
  10. ^ Tinbergen, 1984 pages 58–79
  11. ^ Collett, Thomas S; Graham, Paul Animal Navigation: Path Integration, Visual Landmarks and Cognitive Maps" Current Biology, Vol 14, R475–R477, June 22, 2004 doi:101016/jcub200406013
  12. ^ Dunlap, JC; Loros, J; DeCoursey, PJ 2003 Chronobiology: Biological Timekeeping Sinauer, Sunderland
  13. ^ a b Alcock, John 2009 Animal Behavior: An Evolutionary Approach Sinauer Associates pp 140–143 ISBN 978-0-87893-225-2 
  14. ^ Lockley, 1967 page 74
  15. ^ Lockley, 1967 page 136
  16. ^ Dacke, M; Nilsson, D E; Scholtz, C H; Byrne, M; Warrant, E J 2003 "Animal behaviour: Insect orientation to polarized moonlight" Nature 424 6944: 33 doi:101038/424033a 
  17. ^ Milius, Susan 2003 "Moonlighting: Beetles navigate by lunar polarity" Science News 164 1: 4 
  18. ^ Roach, John 2003 "Dung Beetles Navigate by the Moon, Study Says", National Geographic News Retrieved on 2007-08-02
  19. ^ Milius, S 2003 "Moonlighting" Science News 164: 4 doi:102307/3981988 JSTOR 3981988 
  20. ^ J Exp Biol May 2003 vol 206 no 9 1535-1543
  21. ^ Dacke, M; Baird, E; Byrne, M; Scholtz, C H; Warrant, E J 2013 "Dung Beetles Use the Milky Way for Orientation" Current Biology 23 4: 298–300 doi:101016/jcub201212034 PMID 23352694 
  22. ^ Dacke, Marie; Baird, Emily; Byrne, Marcus; Scholtz, Clarke H; Warrant, Eric J 2013 "Dung Beetles Use the Milky Way for Orientation" Current Biology 23 4: 298–300 doi:101016/jcub201212034 PMID 23352694 
  23. ^ Wits University 24 January 2013 "Dung Beetles Follow the Milky Way: Insects Found to Use Stars for Orientation" ScienceDaily Retrieved 25 January 2013 
  24. ^ von Frisch, 1953
  25. ^ Grüter, C; Balbuena, M; Farina, W 2008 "Informational conflicts created by the waggle dance" Proceedings of the Royal Society B 275 1640: 1321–1327 doi:101098/rspb20080186 PMC 2602683 PMID 18331980 
  26. ^ a b Kimchi, Tali; Etienne, Ariane S; Terkel, Joseph 2004 A subterranean mammal uses the magnetic compass for path integration PNAS, January 27, vol 101, no 4, 1105–1109
  27. ^ M Lindauer and H Martin, in SR Galler et al Animal Orientation and Navigation 559/1, 1972
  28. ^ Walcott, C 1996 Pigeon homing: observations, experiments and confusions Journal of Experimental Biology, 199, 21-7 Text
  29. ^ Keeton, WT 1971 "Magnets interfere with pigeon homing" Proceedings of the National Academy of Sciences of the United States of America 68 1: 102–6 doi:101073/pnas681102 PMC 391171 PMID 5276278 
  30. ^ Ioalè, P; Nozzolini, M; Papi, F 1990 "Homing pigeons do extract directional information from olfactory stimuli" Behav Ecol Sociobiol 26: 301–305 doi:101007/bf00171094 
  31. ^ Wallraff, HG 1974 Das Navigationssystem der Vögel Ein theoretischer Beitrag zur Analyse ungeklärter Orientierungsleistungen Schriftenreihe 'Kybernetik' München, Wien: R Oldenbourg Verlag
  32. ^ Wiltschko, W; Wiltschko, R 1996 "Magnetic Orientation in Birds" Journal of Experimental Biology 199: 29–38 
  33. ^ Lockley, 1967 page 180
  34. ^ Lohmann, KJ; Lohmann, CMF; Endres, CS 2008 The sensory ecology of ocean navigation J Exp Biol, 211: 1719–1728
  35. ^ Schulte-Pelkum, N; Wieskotten, S; Hanke, W; Dehnhardt, G; Mauck, B "Tracking of biogenic hydrodynamic trails in harbour seals Phoca vitulina" Journal of Experimental Biology 210, no 5 2007 781–7 doi:101242/jeb02708 PMID 17297138
  36. ^ Schevill, WE and McBride, AF 1956 Evidence for echolocation by cetaceans Deep-Sea Research 3:153–154
  37. ^ Breed, Michael D 2001 "Path Integration" Animal Behavior Online Retrieved 10 December 2012 
  38. ^ Gallistel The Organization of Learning 1990
  39. ^ Whishaw, IQ; Hines, DJ; Wallace, DG 2001 "Dead reckoning path integration requires the hippocampal formation: evidence from spontaneous exploration and spatial learning tasks in light allothetic and dark idiothetic tests" PDF Behavioural Brain Research 127: 49–69 doi:101016/s0166-43280100359-x 
  40. ^ Mittelstaedt, H; Mittelstaedt, M-L 1973 Mechanismen der orientierung ohne richtende aussenreize Forschr Zool, 21:46–58
  41. ^ Mittelstaedt, M-L; Mittelstaedt, H 1980 "Homing by path integration in a mammal" Naturwissenschaften 67: 566–567 doi:101007/bf00450672 
  42. ^ Wehner R 2003 "Desert ant navigation: how miniature brains solve complex tasks" PDF Journal of Comparative Physiology 189 8: 579–588 doi:101007/s00359-003-0431-1 PMID 12879352 
  43. ^ Gibson, JJ 1950 The Perception of the Visual World Houghton Mifflin 
  44. ^ McNaughton, BL; Battaglia FP; Jensen O; Moser EI; Moser MB August 2006 "Path integration and the neural basis of the 'cognitive map'" Nature Reviews Neuroscience 7 8: 663–678 doi:101038/nrn1932 PMID 16858394 
  45. ^ Redish, 1999 p67
  46. ^ Black, Richard 29 March 2012 "BBC News: Science & Environment" Pesticides hit queen bee numbers BBC Retrieved 30 March 2012 
  47. ^ Witherington, Blair E in Clemmons, Janine Rhea, and Buchholz, Richard editors 1997 Behavioral Approaches to Conservation in the Wild Cambridge University Press pp 301–328  CS1 maint: Uses authors parameter link


  • Gauthreaux, Sidney A 1980 Animal Migration, Orientation, and Navigation Academic Press 
  • Keeton, William 1972 Effects of magnets on pigeon homing pages 579–594 in Animal Orientation and Navigation NASA SP-262 Washington, DC
  • Keeton, William 1974 The orientational and navigational basis of homing in birds pages 47–132 in Advances in the Study of Behavior, Vol 5 Academic Press
  • Keeton, William 1977 Magnetic Reception biology In Encyclopedia of Science and Technology, 2nd Ed McGraw-Hill
  • Keeton, William 1979 Pigeon Navigation pages 5–20 in Neural Mechanisms of Behavior in the Pigeon A M Granda and J H Maxwell, Eds Plenum Publishing
  • Lockley, Ronald M 1967 Animal Navigation Pan Books 
  • Lockley, Ronald M 1942 Shearwaters J M Dent 
  • Redish, A David 1999 Beyond the Cognitive Map MIT Press  PDF
  • Tinbergen, Nico 1984 Curious Naturalists Revised ed University of Massachusetts Press 
  • von Frisch, Karl 1953 The Dancing Bees Harcourt, Brace & World 

External links

  • How Stuff Works: Animal Navigation
  • Oldenburg University: Animal Navigation
  • National Geographic: Animal Navigation resources for teachers

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