Professional Documents
Culture Documents
Contents
Examples
History
Proposed functions
Social interaction
Protection from predators
Enhanced foraging
Increased locomotion efficiency
Costs of group living
Ectoparasitism and disease
Intraspecific competition
Reproduction and development
Stress
Inbreeding
Group structure
Experimental approach
Modeling approach
Collective decision making
See also
References
Further reading
External links
Examples
Examples of collective animal behavior include:
Flocking birds
Herding ungulates
Shoaling and schooling fish
Schooling Antarctic krill
Pods of dolphins
Marching locusts
Nest building ants
History
The basis of collective animal behaviour originated from the study of collective phenomena[1]; that is,
repeated interactions among individuals that produce large scale patterns. The foundation of collective
phenomena originates from the idea that collective systems can be understood from a set of techniques.
For example, Nicolis and Prigogine (1977)[2] employed the use of non-linear thermodynamics to help
explain similarities between collective systems at different scales. Other studies aim to use physics,
mathematics and chemistry to provide frameworks to study collective phenomena.[3][4][5]
Proposed functions
Many functions of animal aggregations have been proposed. These proposed functions may be grouped
into the four following categories: social and genetic, anti-predator, enhanced foraging, and increased
locomotion efficiency.
Social interaction
Support for the social and genetic function of aggregations, especially those formed by fish, can be seen
in several aspects of their behavior. For instance, experiments have shown that individual fish removed
from a school will have a higher respiratory rate than those found in the school. This effect has been
partly attributed to stress, although hydrodynamic factors were considered more important in this
particular study.[6] The calming effect of being with conspecifics may thus provide a social motivation
for remaining in an aggregation. Herring, for instance, will become very agitated if they are isolated from
conspecifics.[7] Fish schools have also been proposed to serve a reproductive function since they provide
increased access to potential mates.
Enhanced foraging
A third proposed benefit of animal groups is that of enhanced foraging. This ability was demonstrated by
Pitcher and others in their study of foraging behavior in shoaling cyprinids.[21] In this study, the time it
took for groups of minnows and goldfish to find a patch of food was quantified. The number of fishes in
the groups was varied, and a statistically significant decrease in the amount of time necessary for larger
groups to find food was established. Further support for an enhanced foraging capability of schools is
seen in the structure of schools of predatory fish. Partridge and others analyzed the school structure of
Atlantic bluefin tuna from aerial photographs and found that the school assumed a parabolic shape, a fact
that was suggestive of cooperative hunting in this species (Partridge et al., 1983).[22]
Another example can be seen in homing pigeons. When a homing pigeon is released with other
individuals from its roost, these pigeon groups showed increased efficiency and decision making to
shorten the distance of the route taken to return home, thus saving energy when flying between
locations.[24]
For example, cliff swallows that are commonly parasitized by swallow bugs incur a cost when forming
colonies, as these parasitic bugs increase the mortality rates of cliff swallow nestlings[26]. A study shows
that the number of swallow bugs found in cliff swallow nests increased with the increase of cliff swallow
colony size, thus reducing overall success of these colonies.[26]
Larger groups of animals tend to harbour an increased number of pathogens and are at a higher risk of
epidemics[27]. This is particularly due to the large amount of waste material produced by larger groups,
allowing for a favourable environment for pathogens to thrive.
Intraspecific competition
Another cost to group living is the competition over food resources. As individuals group together, there
is an increased nutritional requirement of the larger group compared to smaller groups. This causes an
increased energetic cost as individuals now travel farther to visit resource patches[28].
An example of intraspecific competition can be seen within groups of whales and dolphins. Female
bottle-nose dolphins with similar home ranges tend to have varied foraging habits in an effort to reduce
and negate the intraspecific competition of resources[29]. Benefits of group living on defence from
predators is very evident in nature, however in locations of high resource competition poses an effect on
the mortality of certain individuals. This can be seen in species of shoaling fish, where the initial
aggregation of individuals to a group initially allowed for the protection from predators, however the
limiting resources available changes over time, and mortality rates of these fish begin to increase[30],
showing that resource competition is an important regulator of reef fish groups after the initial benefits of
refuge grouping and predatory protection.
Interesting contrasts to the benefit of increased group size on foraging efficiency can be seen in nature
particularly due to intraspecific interactions. A study conducted on the Alaskan moose shows that with
increasing group size, there is a decrease in foraging efficiency[31]. This is result of increased social
aggression in the groups, as the individuals of the group spent most of its time in alert-alarm postures,
thus spending less time foraging and feeding, reducing its foraging efficiency.
Stress
Another cost to group living is stress levels within individuals of a group. Stress levels within group
living varies dependent on the size of the colony or group. A large group of animals may suffer larger
levels of stress arising from intraspecific food competition. In contrast, smaller groups may have
increased stress levels arising from the lack of adequate defense from predators as well as a reduced
foraging efficiency[34].
An example can be seen in a study conducted on a species of ring-tail lemurs (Lemur catta). This study
found that an optimum group size of around 10-20 individuals produces the lowest level of cortisol (an
indicator of stress), while groups with smaller or larger than 10-20 individuals showed an increased level
of cortisol production, thus an increased level of stress within the individuals of the larger and smaller
groups[35].
Inbreeding
Another proposed cost to group living is the cost incurred to avoid inbreeding. Individuals may it be male
or females in groups may disperse in an effort to avoid inbreeding[36]. This poses a more detrimental
effect on smaller, isolated groups of individuals, as they are at a greater risk of inbreeding and thus
suppressing the group’s overall fitness[27].
Group structure
The structure of large animal groups has been difficult to study because of the large number of animals
involved. The experimental approach is therefore often complemented by mathematical modeling of
animal aggregations.
Experimental approach
Experiments investigating the structure of animal aggregations seek to determine the 3D position of each
animal within a volume at each point in time. It is important to know the internal structure of the group
because that structure can be related to the proposed motivations for animal grouping. This capability
requires the use of multiple cameras trained on the same volume in space, a technique known as
stereophotogrammetry. When hundreds or thousands of animals occupy the study volume, it becomes
difficult to identify each one. In addition, animals may block one another in the camera views, a problem
known as occlusion. Once the location of each animal at each point in time is known, various parameters
describing the animal group can be extracted.
Density: The density of an animal aggregation is the number of animals divided by the volume (or area)
occupied by the aggregation. Density may not be a constant throughout the group. For instance, starling
flocks have been shown to maintain higher densities on the edges than in the middle of the flock, a
feature that is presumably related to defense from predators.[37]
Polarity: The group polarity describes if the group animals are all pointing in the same direction or not.
In order to determine this parameter, the average orientation of all animals in the group is determined.
For each animal, the angular difference between its orientation and the group orientation is then found.
The group polarity is then the average of these differences (Viscido 2004).[38]
Nearest Neighbor Distance: The nearest neighbor distance (NND) describes the distance between the
centroid of one animal (the focal animal) and the centroid of the animal nearest to the focal animal. This
parameter can be found for each animal in an aggregation and then averaged. Care must be taken to
account for the animals located at the edge of an animal aggregation. These animals have no neighbor in
one direction.
Nearest Neighbor Position: In a polar coordinate system, the nearest neighbor position describes the
angle and distance of the nearest neighbor to a focal animal.
Packing Fraction: Packing fraction is a parameter borrowed from physics to define the organization (or
state i.e. solid, liquid, or gas) of 3D animal groups. It is an alternative measure to density. In this
parameter, the aggregation is idealized as an ensemble of solid spheres, with each animal at the center of
a sphere. The packing fraction is defined as the ratio of the total volume occupied by all individual
spheres divided by the global volume of the aggregation (Cavagna 2008). Values range from zero to one,
where a small packing fraction represents a dilute system like a gas. Cavagna found that the packing
fraction for groups of starlings was 0.012.[39]
Integrated Conditional Density: This parameter measures the density at various length scales and
therefore describes the homogeneity of density throughout an animal group.[39]
Pair Distribution Function: This parameter is usually used in physics to characterize the degree of
spatial order in a system of particles. It also describes the density, but this measures describes the density
at a distance away from a given point. Cavagna et al. found that flocks of starlings exhibited more
structure than a gas but less than a liquid.[39]
Modeling approach
The simplest mathematical models of animal aggregations generally instruct the individual animals to
follow three rules:
A recent investigation showed that small groups of fish used consensus decision-making when deciding
which fish model to follow. The fish did this by a simple quorum rule such that individuals watched the
decisions of others before making their own decisions. This technique generally resulted in the 'correct'
decision but occasionally cascaded into the 'incorrect' decision. In addition, as the group size increased,
the fish made more accurate decisions in following the more attractive fish model.[46] Consensus
decision-making, a form of collective intelligence, thus effectively uses information from multiple
sources to generally reach the correct conclusion.
Some simulations of collective decision-making use the Condorcet method to model the way groups of
animals come to consensus.
See also
Biomimetics
Collective cell migration
Collective intelligence
Emergence
Swarm intelligence
References
1. Sumpter, D.J., 2010. Collective animal behavior. Princeton University Press.
2. Nicolis, G. & Prigogine, I. 1977. Self-organization in nonequilibrium systems . New York:
John Wiley & Sons.
3. Wiener, N. 1948. Cybernetics . New York: John Wiley & Sons.
4. Ashby, W. R. 1947. Principles of the self-organizing dynamic system. Journal of General
Psychology 125– 128.
5. von Bertalanffy, L. 1968. General system theory . New York: George Braziller.
6. Abrahams, M.; Colgan, P. (1985). "Risk of predation, hydrodynamic efficiency, and their
influence on school structure". Environmental Biology of Fishes. 13 (3): 195–202.
doi:10.1007/bf00000931 (https://doi.org/10.1007%2Fbf00000931).
7. Partridge, B.; Pitcher, T.; Cullen, M.; Wilson, J. (1980). "The three-dimensional structure of
fish schools". Behav Ecol and Sociobiology. 6 (4): 277–288. doi:10.1007/bf00292770 (http
s://doi.org/10.1007%2Fbf00292770).
8. Milinski, H.; Heller, R. (1978). "Influence of a predator on the optimal foraging behavior of
sticklebacks". Nature. 275 (5681): 642–644. Bibcode:1978Natur.275..642M (https://ui.adsab
s.harvard.edu/abs/1978Natur.275..642M). doi:10.1038/275642a0 (https://doi.org/10.1038%
2F275642a0).
9. Jeschke JM, Tollrian R (2007). "Prey swarming: which predators become confused and
why?". Animal Behaviour. 74 (3): 387–393. doi:10.1016/j.anbehav.2006.08.020 (https://doi.o
rg/10.1016%2Fj.anbehav.2006.08.020).
10. Ioannou CC, Tosh CR, Neville L, Krause J (2008). "The confusion effect—from neural
networks to reduced predation risk" (http://beheco.oxfordjournals.org/content/19/1/126).
Behavioral Ecology. 19 (1): 126–130. doi:10.1093/beheco/arm109 (https://doi.org/10.1093%
2Fbeheco%2Farm109).
11. Krakauer DC (1995). "Groups confuse predators by exploiting perceptual bottlenecks: a
connectionist model of the confusion effect". Behavioral Ecology and Sociobiology. 36 (6):
421–429. doi:10.1007/BF00177338 (https://doi.org/10.1007%2FBF00177338).
12. Olson RS, Hintze A, Dyer FC, Knoester DB, Adami C (2013). "Predator confusion is
sufficient to evolve swarming behaviour" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC40
43163). J. R. Soc. Interface. 10 (85): 20130305. arXiv:1209.3330 (https://arxiv.org/abs/120
9.3330). doi:10.1098/rsif.2013.0305 (https://doi.org/10.1098%2Frsif.2013.0305).
PMC 4043163 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4043163). PMID 23740485
(https://www.ncbi.nlm.nih.gov/pubmed/23740485).
13. Demsar J, Hemelrijk CK, Hildenbrandt H, Bajec IL (2015). "Simulating predator attacks on
schools: Evolving composite tactics" (https://pure.rug.nl/ws/files/85452752/Simulating_pred
ator_attacks_on_schools_Evolving_composite_tactics.pdf) (PDF). Ecological Modelling.
304: 22–33. doi:10.1016/j.ecolmodel.2015.02.018 (https://doi.org/10.1016%2Fj.ecolmodel.2
015.02.018).
14. Roberts, G. (1996). "Why individual vigilance declines as group size increases". Anim
Behav. 51 (5): 1077–1086. CiteSeerX 10.1.1.472.7279 (https://citeseerx.ist.psu.edu/viewdo
c/summary?doi=10.1.1.472.7279). doi:10.1006/anbe.1996.0109 (https://doi.org/10.1006%2
Fanbe.1996.0109).
15. Lima, S. (1995). "Back to the basics of anti-predatory vigilance: the group-size effect".
Animal Behaviour. 49 (1): 11–20. doi:10.1016/0003-3472(95)80149-9 (https://doi.org/10.101
6%2F0003-3472%2895%2980149-9).
16. Olson RS, Haley PB, Dyer FC, Adami C (2015). "Exploring the evolution of a trade-off
between vigilance and foraging in group-living organisms" (https://www.ncbi.nlm.nih.gov/pm
c/articles/PMC4593673). Royal Society Open Science. 2 (9): 150135. arXiv:1408.1906 (http
s://arxiv.org/abs/1408.1906). Bibcode:2015RSOS....250135O (https://ui.adsabs.harvard.ed
u/abs/2015RSOS....250135O). doi:10.1098/rsos.150135 (https://doi.org/10.1098%2Frsos.1
50135). PMC 4593673 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4593673).
PMID 26473039 (https://www.ncbi.nlm.nih.gov/pubmed/26473039).
17. Hamilton, W. D. (1971). "Geometry for the selfish herd". J. Theor Biology. 31 (2): 295–311.
doi:10.1016/0022-5193(71)90189-5 (https://doi.org/10.1016%2F0022-5193%2871%299018
9-5). PMID 5104951 (https://www.ncbi.nlm.nih.gov/pubmed/5104951).
18. Olson RS, Knoester DB, Adami C (2013). Critical Interplay Between Density-dependent
Predation and Evolution of the Selfish Herd (http://dl.acm.org/citation.cfm?doid=2463372.24
63394). Proceedings of GECCO 2013. Gecco '13. pp. 247–254.
doi:10.1145/2463372.2463394 (https://doi.org/10.1145%2F2463372.2463394).
ISBN 9781450319638.
19. Turner, G.; Pitcher, T. (1986). "Attack abatement: a model for group protection by combined
avoidance and dilution". American Naturalist. 128 (2): 228–240. doi:10.1086/284556 (http
s://doi.org/10.1086%2F284556).
20. Krause, J.; Ruxton, G.; Rubenstein, D. (1998). "Is there always an influence of shoal size on
predator hunting success?". Journal of Fish Biology. 52 (3): 494–501.
doi:10.1006/jfbi.1997.0595 (https://doi.org/10.1006%2Fjfbi.1997.0595).
21. Pitcher, T.; Magurran, A.; Winfield, I. (1982). "Fish in larger shoals find food faster". Behav.
Ecol. And Sociobiology. 10: 2.
22. Partridge, B. Johansson; Kalish, J. (1983). "The structure of schools of giant bluefin tuna in
Cape Cod Bay". Environmental Biology of Fishes. 9 (3–4): 253–262.
doi:10.1007/bf00692374 (https://doi.org/10.1007%2Fbf00692374).
23. Fish, F.E. Kinematics of ducklings swimming in formation: consequences of position. (1995)
Journal of Experimental Zoology 273:1-11.
24. Biro, D., Sumpter, D.J., Meade, J. and Guilford, T., 2006. From compromise to leadership in
pigeon homing. Current Biology, 16(21), pp.2123-2128.
25. Alexander, R.D., 1974. The evolution of social behavior. Annual review of ecology and
systematics, 5(1), pp.325-383.
26. Brown, C.R. and Brown, M.B., 1986. Ectoparasitism as a cost of coloniality in cliff swallows
(Hirundo pyrrhonota). Ecology, 67(5), pp.1206-1218.
27. Loehle, C., 1995. Social barriers to pathogen transmission in wild animal populations.
Ecology, 76(2), pp.326-335.
28. Chapman, C.A. and Valenta, K., 2015. Costs and benefits of group living are neither simple
nor linear. Proceedings of the National Academy of Sciences, 112(48), pp.14751-14752.
29. Rossman, S., Ostrom, P.H., Stolen, M., Barros, N.B., Gandhi, H., Stricker, C.A. and Wells,
R.S., 2015. Individual specialization in the foraging habits of female bottlenose dolphins
living in a trophically diverse and habitat rich estuary. Oecologia, 178(2), pp.415-425.
30. Ford, J.R. and Swearer, S.E., 2013. Two's company, three's a crowd: Food and shelter
limitation outweigh the benefits of group living in a shoaling fish. Ecology, 94(5), pp.1069-
1077.
31. Molvar, E.M. and Bowyer, R.T., 1994. Costs and benefits of group living in a recently social
ungulate: the Alaskan moose. Journal of Mammalogy, 75(3), pp.621-630.
32. Borries, C., Larney, E., Lu, A., Ossi, K. and Koenig, A., 2008. Costs of group size: lower
developmental and reproductive rates in larger groups of leaf monkeys. Behavioral Ecology,
19(6), pp.1186-1191.
33. Silva, J.D., Macdonald, D.W. and Evans, P.G., 1994. Net costs of group living in a solitary
forager, the Eurasian badger (Meles meles). Behavioral Ecology, 5(2), pp.151-158.
34. Sterck, E.H., Watts, D.P. and van Schaik, C.P., 1997. The evolution of female social
relationships in nonhuman primates. Behavioral ecology and sociobiology, 41(5), pp.291-
309.
35. Pride, R.E., 2005. Optimal group size and seasonal stress in ring-tailed lemurs (Lemur
catta). Behavioral Ecology, 16(3), pp.550-560.
36. Clutton-Brock, T.H., 1989. Female transfer and inbreeding avoidance in social mammals.
Nature, 337(6202), p.70.
37. M. Ballerini, N. Cabibbo, R. Candelier, A. Cavagna, E. Cisbani, I. Giardina, A. Orlandi, G.
Parisi, A. Procaccini, M. Viale, and V. Zdravkovic (2008) ‘Empirical investigation of starling
flocks: a benchmark study in collective animal behaviour’ Animal Behavior 76(1): 201-215.
38. Viscido, S.; Parrish, J.; Grunbaum, D. (2004). "Individual behavior and emergent properties
of fish schools: a comparison of observation and theory". Marine Ecology Progress Series.
273: 239–249. Bibcode:2004MEPS..273..239V (https://ui.adsabs.harvard.edu/abs/2004ME
PS..273..239V). doi:10.3354/meps273239 (https://doi.org/10.3354%2Fmeps273239).
39. Cavagna, A.; Cimarelli, Giardina; Orlandi, Parisi; Procaccini, Santagati; Stefanini (2008).
"New statistical tools for analyzing the structure of animal groups". Mathematical
Biosciences. 214 (1–2): 32–37. doi:10.1016/j.mbs.2008.05.006 (https://doi.org/10.1016%2F
j.mbs.2008.05.006). PMID 18586280 (https://www.ncbi.nlm.nih.gov/pubmed/18586280).
40. M. Ballerini, N. Cabibbo, R. Candelier, A. Cavagna, E. Cisbani, I. Giardina, V. Lecomte, A.
Orlandi, G. Parisi, A. Procaccini, M. Viale, and V. Zdravkovic (2008) ‘Interaction ruling
animal collective behavior depends on topological rather than metric distance: Evidence
from a field study’ PNAS 105:1232-1237.
41. Hildenbrandt, H; Carere, C; Hemelrijk, CK (2010). "Self-organized aerial displays of
thousands of starlings: a model" (http://beheco.oxfordjournals.org/content/21/6/1349.full).
Behavioral Ecology. 21 (6): 1349–1359. doi:10.1093/beheco/arq149 (https://doi.org/10.109
3%2Fbeheco%2Farq149).
42. Hemelrijk, CK; Hildenbrandt, H (2011). "Some causes of the variable shape of flocks of
birds" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3150374). PLOS ONE. 6 (8): e22479.
Bibcode:2011PLoSO...622479H
(https://ui.adsabs.harvard.edu/abs/2011PLoSO...622479H).
doi:10.1371/journal.pone.0022479 (https://doi.org/10.1371%2Fjournal.pone.0022479).
PMC 3150374 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3150374). PMID 21829627
(https://www.ncbi.nlm.nih.gov/pubmed/21829627).
43. Project Starflag (http://www.rug.nl/sciencelinx/exhibits/swarming/index)
44. Swarm behaviour model by University of Groningen (http://www.rug.nl/fmns-research/beso/
_people/hemelrijk)
45. Sumpter, D. http://www.collective-behavior.com/index.html
46. Sumpter, D.; Krause, J; James, R.; Couzin, I.; Ward, A. (2008). "Consensus decision
making by fish". Current Biology. 18 (22): 1773–1777. doi:10.1016/j.cub.2008.09.064 (http
s://doi.org/10.1016%2Fj.cub.2008.09.064). PMID 19013067 (https://www.ncbi.nlm.nih.gov/p
ubmed/19013067).
Further reading
Camazine, S., Deneubourg, J.L., Franks, N.R., Sneyd, J., Theraulaz, G. and Bonabeau, E.
(2001) Self-Organization in Biological Systems Princeton University Press, Princeton, N.J.
ISBN 0-691-01211-3 (especially Chapter 11)
Sumpter, D. J. T. (2010) "Collective Animal Behavior" Princeton University Press, Princeton,
ISBN 978-0-691-14843-4
External links
Collective Animal Behavior website organized around David Sumpter's book (2008) by the
same name (http://www.collective-behavior.com/index.html)
STARFLAG project: Description of starling flocking project (http://hal.elte.hu/starling/)
Center for Biologically Inspired Design at Georgia Tech (http://www.cbid.gatech.edu/)
David Sumpter's research website (http://www.math.uu.se/~david/web/index.html)
Iain Couzin's research website (http://icouzin.princeton.edu/)
Website of Julia Parrish, an animal aggregation researcher (http://www.fish.washington.edu/
people/parrish/)
Research for this Wikipedia entry was conducted as a part of a Locomotion
Neuromechanics course (APPH 6232) offered in the School of Applied Physiology at
Georgia Tech (https://web.archive.org/web/20110719200641/http://www.ap.gatech.edu/Cha
ng/Lab/CNL/APPH6232.html)
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