Skip to main content
SpringerLink
Log in

Shoals and schools: back to the heuristic definitions and quantitative references

  • Reviews
  • Published:
Reviews in Fish Biology and Fisheries Aims and scope Submit manuscript

Abstract

The terms ‘shoal’, ‘swarm’ and ‘school’ are very frequently used in research on collective behaviours in animals. Pitcher’s definitions are accepted as the authority in the field but are based on a conceptual criterion of sociability. Without call into question the basis of these definitions, they do not provide tools to determine these behaviours quantitatively. To compare studies between populations, species, taxa or different experimental treatments, and between different authors, quantitative references are necessary. Quantitative measurements of collective behaviours can also test and validate the predictive capacity of computer models by comparing real data from nature so that different models can be compared. The first part of this paper succinctly reviews the definitions and meanings of these behaviours, with particular attention paid to quantitative aspects. This review underlines a series of conceptual confusions concerning these behavioural terms observed in the scientific literature and oral scientific communications. The second part reviews the quantitative parameters developed by biologists studying collective fish behaviours, mainly fish shoals, and by theoretical biologists and physicists studying computer modelling of collective behaviours. The parameters reviewed herein make no attempt to explain the mechanisms and causes that create a shoal, a swarm or a school, but rather try to describe these collective behaviours, and to connect local and global properties with individual and collective behaviours. Recent development over the last decade in technology, data processing capacity, cameras, and video tracking tools have provided the opportunity to obtain quantitative measures of collective dynamic behaviours in animals both rapidly and precisely.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Log in via an institution

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Institutional subscriptions

Fig. 1
Fig. 2
Fig. 3
Fig. 4
Fig. 5
Fig. 6
Fig. 7
Fig. 8
Fig. 9
Fig. 10
Fig. 11
Fig. 12
Fig. 13
Fig. 14

Similar content being viewed by others

References

  • Adioui M, Treuil JP, Arino O (2003) Alignment in a fish school: a mixed Lagrangian-Eularian approach. Ecol Modell 167:19–32

    Google Scholar 

  • Aoki I (1982) A simulation study on the schooling mechanism in fish. Bull Jpn Soc Sci Fish 48:1081–1088

    Google Scholar 

  • Aoki I (1984) Internal dynamics of fish schools in relation to inter-fish distance. Bull Jpn Soc Sci Fish 50:751–758

    Google Scholar 

  • Ballerini M, Cabibbo N, Candelier R, Cavagna A, Cisbani E, Giardina I, Lecomte V, Orlandi A, Parisi G, Procaccini A, Viale M, Zdravkovic V (2008) Interaction ruling animal collective behaviour depends on topological rather than metric distance: evidence from a field study. Proc Nat Acad Sci USA (PNAS) 105:1232–1237

    CAS  Google Scholar 

  • Banks EM (1982) Behavioral research to answer questions about an animal welfare. J Anim Sci 54:434–446

    PubMed  CAS  Google Scholar 

  • Barlow GW (1991) Mating systems among cichlid fishes. In: Keenleyside MHA (ed) Cichlid fishes: behaviour, ecology and evolution. Fish and Fisheries series 2. Chapman & Hall, London, pp 173–190

  • Becco C, Vandewalle N, Delcourt J, Poncin P (2006) Experimental evidences of a structural and dynamical transition in fish school. Physica A Stat Mech Appl 367:487–493

    Google Scholar 

  • Boal JG, Gonzalez A (1998) The social behaviour of individual oval squids (Cephalopoda, Teurhoidea, Loliginidae, Sepioteuthis lessoniana) within a captive school. Ethology 104(161–1):78

    Google Scholar 

  • Bode NWF, Faria JJ, Franks DW, Krause J, Wood AJ (2010) How perceived threat increases synchronization in collectively moving animal groups. Proc R Soc B 227:3065–3070

    Google Scholar 

  • Breder CM (1959) Studies on the social groupings of fishes. Bull Museum Am Nat History 117:393–482

    Google Scholar 

  • Brown C, Laland K, Krause J (2011) Fish cognition and behaviour, 2nd edn. Fish and aquatic resources series 15, Blackwell, 472 p

  • Buhl J, Sumpter DJT, Couzin ID, Hale JJ, Despland E, Miller ER, Simpson SJ (2006) From disorder to order in marching locusts. Science 312:1402–1406

    PubMed  CAS  Google Scholar 

  • Bumann D, Krause J (1993) Front individuals lead in shoals of three-spined sticklebacks (Gasterosteus aculeatus) and juvenile roach (Rutilus rutilus). Behaviour 125:189–198

    Google Scholar 

  • Bumann D, Krause J, Rubenstein D (1997) Mortality risk of spatial positions in animal groups: the danger of being in the front. Behaviour 134:1063–1076

    Google Scholar 

  • Camazine S, Deneubourg JL, Franks NR, Sneyd J, Theraulaz G, Bonadeau E (2001) Self-organization in biological systems. Princeton studies in complexity. Princeton University Press, Princeton

  • Cardy J (1996) Scaling and renormalization in statistical physics. Cambridge lecture notes in physics no. 5, Cambridge University Press, Cambridge

  • Cavagna A, Cimarelli A, Giardina I, Orlandi A, Parisi G, Procaccini A, Santagati R, Stefanini F (2008a) New statistical tools for analyzing the structure of animal groups. Math Biosci 214:32–37

    PubMed  Google Scholar 

  • Cavagna A, Giardina I, Orlandi A, Parisi G, Procaccini A, Viale M, Zdravkovic V (2008b) The STARFLAG handbook on collective animal behaviour. 1: empirical methods. Anim Behav 76:217–236

    Google Scholar 

  • Cavagna A, Giardina I, Orlandi A, Parisi G, Procaccini A (2008c) The STARFLAG handbook on collective animal behaviour. 2: threedimensional analysis. Anim Behav 76:237–248

    Google Scholar 

  • Chivers DP, Brown GE, Smith RJF (1995) Familiarity and shoal cohesion in fathead minnows (Pimephales promelas): implications for antipredator behaviour. Can J Zool 73:955–960

    Google Scholar 

  • Clark PJ, Evans FC (1954) Distance to nearest neighbor as a measure of spatial relationships in populations. Ecology 35:445–453

    Google Scholar 

  • Clark PJ, Evans FC (1979) Generalization of a nearest neighbour measure of dispersion fro use in k dimensions. Ecology 60:316–317

    Google Scholar 

  • Codling EA, Hill NA, Pitchford JW, Simpson SD (2004) Random walk models for the movement and recruitment of reef fish larvae. Marine Ecol Progr Series 279:215–224

    Google Scholar 

  • Codling EA, Bearon RN, Thorn GJ (2010) Diffusion about the mean drift location in a biased random walk. Ecology 91:3106–3113

    PubMed  Google Scholar 

  • Couzin ID, Franks NR (2002) Self-organised lane formation and optimized traffic flow in army ants. Proc R Soc Lond B 270:139–146

    Google Scholar 

  • Couzin I, Krause J (2003) Self-organization and collective behavior in vertebrates. Adv Study Behav 32:1–75

    Google Scholar 

  • Couzin ID, Krause J, James R, Ruxton GD, Franks NR (2002) Collective memory and spatial sorting in animal groups. J Theor Biol 218:1–11

    PubMed  Google Scholar 

  • Couzin I, Krause J, Franks N, Levin S (2005) Effective leadership and decision-making in animal groups on the move. Nature 433:513–516

    PubMed  CAS  Google Scholar 

  • Cowen RK, Lwiza KMM, Sponauge S, Paris CB, Olson DB (2000) Connectivity of marine populations: open or closed? Science 287:857–859

    PubMed  CAS  Google Scholar 

  • Cullen JM (1974) Statics and dynamics of a fish school. Abstract XIVth International Ethological Congress, Parma, Italy

  • Czirok A, Vicsek M, Vicsek T (1999) Collective motion of organisms in three dimension. Phys A 264:299–304

    Google Scholar 

  • Delcourt J (2008) Structure et ontogenèse des comportements de banc chez deux poissons cichlidés: approche quantitative par l’utilisation du vidéotracking automatisé chez Oreochromis niloticus et Pelvicachromis pulcher. PhD thesis, University of Liège (Belgium), 333 p [in French]

  • Delcourt J, Denoël M, Ylieff M, Poncin P (2012) Video multitracking of fish behaviours: a review and future perspectives. Fish Fish. doi:10.1111/j.1467-2979.2012.00462.x

  • Dill LM, Dunbrack RL, Major PF (1981) A new stereophotographic technique for analysing the three-dimensional structure of fish schools. Environ Biol Fishes 6:7–13

    Google Scholar 

  • Dugatkin LA, Alfieri M (1991a) Guppies and the tit for tat strategy: preference based on past interaction. Behav Ecol Sociobiol 28:243–246

    Google Scholar 

  • Dugatkin LA, Alfieri M (1991b) Tit for tat in guppies (Poecilia reticulata): the relative nature of cooperation and defection during predator inspection. Evol Ecol 5:300–309

    Google Scholar 

  • Dyer JRG, Johansson A, Helbing D, Couzin ID, Krause J (2009) Leadership, consensus decision making and collective behaviours in humans. Philos Trans R Soc B 364:781–789

    Google Scholar 

  • Economakis AE, Lobel PS (1998) Aggregation behaviour of the grey reef shark, Carcharhinus amblyrhychos, at Johnston Atoll, Central Pacific Ocean. Environ Biol Fishes 51:129–139

    Google Scholar 

  • Eggers DM (1976) Theoretical effect of schooling by planktivorous fish predators on rate of prey consumption. J Fish Res Board Can 33:1964–1971

    Google Scholar 

  • Evans SR, Finnie M, Manica A (2007) Shoaling preferences in decapod crustacean. Anim Behav 74:1691–1696

    Google Scholar 

  • Fabre JH (1899) Chapitre XX: La processionnaire du pin: la procession. In souvenirs entomologiques 6ème série. Paris, pp 332–352

  • Faucher K, Parmentier E, Becco Ch, Vandewalle N, Vandewalle P (2010) Fish lateral system is required for accurate control of shoaling behaviour. Anim Behav 79:679–687

    Google Scholar 

  • Fukuda H, Torisawa S, Sawada Y, Takagi T (2010) Ontogenetic changes in schooling behaviour during larval and early juvenile stages of Pacific bluefin tuna Thunnus orientalis. J Fish Biol 76:1841–1847

    PubMed  CAS  Google Scholar 

  • Garnier S, Gautrais J, Theraulaz G (2007) The biological principles of swarm intelligence. Swarm Intell 1:3–31

    Google Scholar 

  • Gautrais J, Jost Ch, Theraulaz G (2008) Key factors in a self-organised fish school model. Ann Zool Fenn 45:415–428

    Google Scholar 

  • Gerlai R, Chatterjee D, Pereira T, Sawashima T, Krishnannair R (2009) Acute and chronic alcohol dose: population difference in behavior and neurochimestry of zebrafish. Genes Brain Behav 8:586–599

    PubMed  CAS  Google Scholar 

  • Géroudet P (1999) Les Palmipèdes d'Europe. 3rd edn, Delachaux et Niestlé, Lausanne, Paris, 510p

  • Gould SJ (1985) The Flamingo’s smile: reflections in natural history. Norton & Company, New York

    Google Scholar 

  • Gould SJ (2002) The structure of the evolutionary theory. Belknap Press, Cambridge

  • Gould SJ (2003) The Hedgehog, the Fox, and the Magister’s Pox. Harmony Books, New York

  • Grégoire G, Chaté H (2004) Onset of collective and cohesive motion. Phys Rev Lett 92:025702.1–025702.4

    Google Scholar 

  • Grégoire G, Chaté H, Tu Y (2003) Moving and staying together without a leader. Physica D 181:157

    Google Scholar 

  • Griffiths SW, Magurran AE (1997a) Schooling preferences for familiar fish vary with group size in a wild guppy population. Proc R Soc Lond B 264:547–551

    Google Scholar 

  • Griffiths SW, Magurran AE (1997b) Familiarity in schooling fish: how long does it take to acquire? Anim Behav 53:945–949

    Google Scholar 

  • Grünbaum D (1994) Translating stochastic density-dependent individual behavior with sensory constrainsts to an Eulerian model of animal swarming. J Math Biol 33:139–161

    PubMed  Google Scholar 

  • Grünbaum D (1998) Schooling as a strategy for taxis in a noisy environment. Evol Ecol 12:503–522

    Google Scholar 

  • Gueron S, Levin SA, Rubenstein DI (1996) The dynamics of the herds: from individuals to aggregations. J Theor Biol 182:85–98

    Google Scholar 

  • Haken H (1983) Synergetic, an introduction. Nonequilibrium phase transitions and self-organization in physics, chemistry, and biology. Springer, New York

  • Hamilton WD (1971) Geometry for the selfish herd. J Theor Biol 31:295–311

    PubMed  CAS  Google Scholar 

  • Hamner WM, Hamner PG (2000) Behavior of antartic krill (Euphasia superba): schooling, foraging, and antipredatory behaviour. Can J Fish Aquatic Sci 57:192–202

    Google Scholar 

  • Hansen J-P, Mc Donald IR (2006) Theory of simple liquids, 3rd edn. Academic Press, Elsevier Inc., Amsterdam, The Netherlands

  • Hemelrijk CK, Kunz H (2004) Density distribution and size sorting in fish schools: an individual-based model. Behav Ecol 16:178–187

    Google Scholar 

  • Hemelrijk CK, Hildenbrandt H, Reinders J, Stamhuis EJ (2010) Emergence of oblong school shape: models and empirical data of fish. Ethology 116:1099–1112

    Google Scholar 

  • Heursel (d’) A, Haddad CFB (2002) Schooling and swimming behaviors of Hyla semilineata tadpoles (Anura, Hylidae). Iheringia Sér. Zool Porto alegre 92:99–104

    Google Scholar 

  • Higgs DM, Fuiman LA (1996) Light intensity and schooling behaviour in larval gulf menhaden. J Fish Biol 48:979–991

    Google Scholar 

  • Hoare DJ, Krause J (2003) Social organisation, shoal structure and information transfer. Fish Fish 4:269–279

    Google Scholar 

  • Hunter JR (1966) Procedure for analysis of schooling behaviour. J Fish Res Board Can 23:547–562

    Google Scholar 

  • Hunter JR, Coyne KM (1982) The onset of schooling in northern anchovy larvae. Cal Coop Ocean Fish 23:246–251

    Google Scholar 

  • Hunter J, Nicholl R (1984) Visual threshold for schooling in northern anchovy Engraulis mordax. Fish Bull 83:235–242

    Google Scholar 

  • Hurley AC (1978) School structure of the squid Loligo opalescens. Fish Bull 76:433–442

    Google Scholar 

  • Huth A, Wissel C (1992) The simulation of the movement of fish schools. J Theor Biol 156:365–385

    Google Scholar 

  • Huth A, Wissel C (1994) The simulation of fish schools in comparison with experimental data. Ecol Modell 75(76):135–145

    Google Scholar 

  • Ikawa T, Okabe H, Mori T, Urabe K, Ikesshoji T (1994) A method for reconstructing the three-dimensional positions of swarming mosquitoes. J Insect Behav 7:237–248

    Google Scholar 

  • Ishizaki Y, Masuda R, Uematsu K, Shimizu K, Arimoto M, Takeuchi T (2001) The effect of dietary docosahexaenoic acid on schooling behaviour and brain development in larval yellowtail. J Fish Biol 58:1691–1703

    CAS  Google Scholar 

  • James MK, Armsworth PR, Mason LB, Bode L (2002) The structure of reef fish metapopulations: modelling larval dispersal and retention patterns. Proc R Soc Lond B 269:2079–2086

    Google Scholar 

  • Keenleyside MHA (1955) Some aspect of the schooling behaviour of fish. Behaviour 8:183–258

    Google Scholar 

  • Keenleyside MHA (1991) Parental care. In: Keenleyside MHA (1991) Cichlid fishes: behaviour, ecology and evolution. Fish and Fisheries series 2. Chapman & Hall, London, pp 191–208

  • Klimley AP (1985) Schooling in Sphyrna lewini, a species with low risk of predation: a non-egalitarian state. Zeitschrift für Tierpsychologie 70:297–319

    Google Scholar 

  • Krause J, Ruxton GD (2002) Living in group. Oxford University Press, Oxford

  • Krause J, Hoare D, Krause S, Hemelrijk CK, Rubenstein DI (2000) Leadership in fish shoals. Fish Fish 1:82–89

    Google Scholar 

  • Kunz H, Hemelrijk CK (2003) Artificial fish schools: collective effects of school size, body size, and body form. Artif Life 9:237–253

    PubMed  Google Scholar 

  • Lehner PN (1987) Design and execution of animal behaviour research: an overview. J Anim Sci 65:1213–1219

    PubMed  CAS  Google Scholar 

  • Lehner PN (1996) Handbook of ethological methods, 2nd edn. Cambridge University Press, Cambridge

  • Lukeman R, Li Y-X, Edelstein-Keshet L (2010) Inferring individual rules from collective behavior. Proc Nat Acad Sci USA 107:12576–12580

    PubMed  CAS  Google Scholar 

  • Magurran AE, Seghers BH (1990) Population differences in the schooling behaviour of newborn guppies, Poecilia reticulata. Ethology 84:334–342

    Google Scholar 

  • Major PF, Dill LM (1978) The three-dimensional structure of airborne bird flocks. Behav Ecol Sociobiol 4:111–122

    Google Scholar 

  • Martin P, Bateson P (2007) Measuring behaviour: an introductory guide, 3rd edn. Cambridge University Press, Cambridge

  • Masuda R, Tsukamoto K (1998) The ontogeny of schooling behaviour in the striped jack. J Fish Biol 52:483–493

    Google Scholar 

  • Masuda R, Tsukamoto K (1999) School formation and concurrent developmental changes in carangid fish with reference to dietary conditions. Environ Biol Fish 56:243–252

    Google Scholar 

  • Masuda R, Shoji J, Nakawama S, Tanaka M (2003) Development of schooling behaviour in Spanish mackerel Scomberomorus niphonius during early ontogeny. Fish Sci 69:772–776

    CAS  Google Scholar 

  • Mather JA, O’Dor RK (1984) Spatial organization of schools of the squid, Illex illecebrosus. Marine Behav Physiol 10:259–271

    Google Scholar 

  • Mauchline J (1971) Seasonal occurrence of mysids (Crustacea) and evidence of social behaviour. J Marine Biol Assoc UK 51:809–825

    Google Scholar 

  • Milinski M (1977a) Experiments on the selection by predators against spatial oddity of their prey. Zeitschrift für Tierpsychologie 43:311–325

    Google Scholar 

  • Milinski M (1977b) Do all members of a swarm suffer the same predation? Zeitschrift für Tierpsychologie 45:373–388

    Google Scholar 

  • Milinski M (1987) Tit for tat in sticklebacks and the evolution of cooperation. Nature 325:433–435

    Google Scholar 

  • Miller N, Gerlai R (2007) Quantification of shoaling behavior in zebrafish (Danio rerio). Behav Brain Res 184:157–166

    PubMed  Google Scholar 

  • Miller NY, Gerlai R (2008) Oscillations in shoal cohesion in zebrafish (Danio rerio). Behav Brain Res 193:148–151

    PubMed  Google Scholar 

  • Miyazaki T, Shiozawa S, Kogane T, Masuda R, Maruyama K, Tsukamoto K (2000) Developmental changes of the light intensity threshold for school formation in the striped jack Pseudocaranx dentex. Marine Ecol Progr Ser 192:267–275

    Google Scholar 

  • Mogilner A, Edelstein-Keshet L (1996) Spatio-angular order in populations of self-aligning objects: formation of oriented patches. Physica D 89:346–367

    Google Scholar 

  • Mogilner A, Bent L, Edelstein-Keshet L (1999) A non-local model for a swarm. J Math Biol 38:534–570

    Google Scholar 

  • Nakayama S, Masuda R, Shoji J, Takeuchi T, Tanaka M (2003) Effect of prey items on the development of schooling behavior in chub mackerel Scomber japonicus in the laboratory. Fish Sci 69:670–676

    CAS  Google Scholar 

  • Nakayama S, Masuda R, Tanaka M (2007) Onsets of schooling behaviour and social transmission in chub mackerel Scomber japonicus. Behav Ecol Sociobiol 61:1383–1390

    Google Scholar 

  • Newlands NK, Porcelli TA (2008) Measuring of the size, shape and structure of Atlantic bluefin tuna schools in the open ocean. Fish Res 91:42–55

    Google Scholar 

  • O’Brien DP (1988) Direct observations of clustering (schooling and swarming) behaviour in mysids (Crustacea: Mysidacea). Marine Ecol 42:235–246

    Google Scholar 

  • O’Connell CP (1972) The interrelation of biting and filtering in the feeding activity of the northern anchovy (Engraulis mordax). J Fish Res Board Can 29:285–293

    Google Scholar 

  • Olst (van) JC, Hunter JR (1970) Some aspects of the organization of fish shoals. J Fish Res Board Can 27:1225–1238

    Google Scholar 

  • Parrish JK, Viscido SV, Grünbaum D (2002) Self-organized fish schools: an examination of emergent properties. Biol Bull 202:296–305

    PubMed  Google Scholar 

  • Partridge BL (1980) Effect of school size on the structure and dynamic of minnow schools. Anim Behav 28:68–77

    Google Scholar 

  • Partridge BL (1981) Internal dynamics and the interrelations of fish in schools. J Comp Physiol 144:313–325

    Google Scholar 

  • Partridge BL (1982) The structure and function of fish schools. Sci Am 246:114–123

    PubMed  CAS  Google Scholar 

  • Partridge BL, Pitcher TJ (1980) The sensory basis of fish schools: relative roles of lateral line and vision. J Comp Physiol 135:315–325

    Google Scholar 

  • Partridge BL, Johansson J, Kalish J (1983) Structure of schools of giant bluefin tuna in Cape Cod Bay. Environ Biol Fish 9:253–262

    Google Scholar 

  • Pather S, Gerlai R (2009) Shuttle box learning in zebrafish (Danio rerio). Behav Brain Res 196:323–327

    PubMed  Google Scholar 

  • Paxton CGM (1996) Isolation and the development of shoaling in two populations of the guppy. J Fish Biol 49:514–520

    Google Scholar 

  • Pitcher TJ (1983) Heuristic definitions of fish shoaling behavior. Anim Behav 31:611–613

    Google Scholar 

  • Pitcher TJ, Parrish, JK (1993) functions of shoaling behaviour in teleosts. In: Pitcher TJ (ed) The behaviour of teleost fishes, 2nd edn. London, Croom Helm, pp 364–439

  • Pitcher TJ, Partridge BL, Wardle CS (1976) A blind fish can school. Science 194:963–965

    PubMed  CAS  Google Scholar 

  • Porch CE (1998) A numerical study of larval fish retention along the southeast Florida coast. Ecol Modell 109:35–59

    Google Scholar 

  • Reebs SG (2000) Can a minority of informed leaders determine the foraging movements in shoals? Anim Behav 59:403–409

    PubMed  Google Scholar 

  • Reebs SG (2001) Influence of body size on leadership in shoals of Golden Shiners, Notemigonus crysoleucas. Behaviour 138:797–809

    Google Scholar 

  • Reuter H, Breckling B (1994) Self-organisation of fish schools: an object-oriented model. Ecol Modell 75(76):147–159

    Google Scholar 

  • Rödel MO, Linsenmair KE (1997) Predator induced swarms in the tadpoles of an African Savannah frog, Phrynomantis microps. Ethology 103:902–914

    Google Scholar 

  • Romey WL (1996) Individual differences make a difference in the trajectories of simulated schools of fish. Ecol Modell 92:65–77

    Google Scholar 

  • Romey WL, Wallace AC (2007) Sex and the selfish herd: sexual segregation within nonmating whirligig groups. Behav Ecol 18:910–915

    Google Scholar 

  • Rosenthal GG, Ryan MJ (2005) Assortative preferences for stripes in danios. Anim Behav 70:1063–1066

    Google Scholar 

  • Ruxton GD, Jackson AL, Tosh CR (2007) Confusion of predators does not rely on specialist coordinated behaviour. Behav Ecol 18:590–596

    Google Scholar 

  • Ryer CH, Olla BL (1998) Effect of light on juvenile walleye pollock shoaling and their interaction with predators. Marine Ecol Progr Ser 167:215–226

    Google Scholar 

  • Salierno JD, Gipson GT, Kane AS (2008) Quantitative movement analysis of social behavior in mummichog, Fundulus heteroclitus. J Ethol 26:35–42

    Google Scholar 

  • Saverino C, Gerlai R (2008) The social zebrafish: behavioral responses to conspecific, heterospecific, and computer animated fish. Behav Brain Res 191:77–87

    PubMed  Google Scholar 

  • Sfakiotakis M, Lane DM, Davies JBC (1999) Review of fish swimming modes for aquatic locomotion. J Ocean Eng 24:237–252

    Google Scholar 

  • Sogard SM, Olla BL (1997) The influence of hunger and predation risk on group cohesion in a pelagic fish, walleye pollock Theragra chalcogramma. Environ Biol Fishes 50:405–413

    Google Scholar 

  • Soria M, Freon P, Chabanet P (2007) Schooling properties of an obligate and a facultative fish species. J Fish Biol 71:1257–1269

    Google Scholar 

  • Spieler M, Linsenmair KE (1999) Aggregation behaviour of Bufo maculatus tadpoles as an antipredator mechanism. Ethology 105:665–686

    Google Scholar 

  • Stöcker S (1999) Models for tuna school formation. Math Biosci 156:167–190

    PubMed  Google Scholar 

  • Sumpter DJT (2006) The principles of collective animal behaviour. Philos Trans R Soc B 361:5–22

    CAS  Google Scholar 

  • Thompson HR (1956) Distribution of distance to nth neighbour in a population of randomly distributed individuals. Ecology 37:391–394

    Google Scholar 

  • Tien JH, Levin SA, Rubenstein DI (2004) Dynamics of fish shoals: identifying key decision rules. Evol Ecol Res 6:555–565

    Google Scholar 

  • Torizawa S, Tagaki T, Fukuda H, Ishibashi Y, Sawada Y, Okada T, Miyashita S, Suzuki K, Yamane T (2007) Schooling behaviour and retinomotor response of juvenile Pacific bluefin tuna Thunnus orientalis under different light intensity. J Fish Biol 71:411–420

    Google Scholar 

  • Tosh CR, Jackson AL, Ruxton GD (2006) The confusion effect in predatory neural networks. Am Nat 167:E52–E65

    PubMed  Google Scholar 

  • Treherne JE, Foster WA (1981) Group transmission of predator avoidance behaviour in a marine insect: the Trafalgar effect. Anim Behav 29:911–917

    Google Scholar 

  • Vicsek T, Czirók A, Ben-Jacob E, Cohen I, Shochet O (1995) Novel type of phase transition in a system of self-driven particles. Phys Rev Lett 75:1226–1229

    PubMed  CAS  Google Scholar 

  • Viscido SV, Parrish JK, Grünbaum D (2004) Individual behaviour and emergent properties of fish schools: a comparison of observation and theory. Marine Ecol Progr Ser 273:239–249

    Google Scholar 

  • Viscido SV, Parrish JK, Grünbaum D (2005) The effect of population size and number of influential neighbors on the emergents properties of fish schools. Ecol Modell 183:347–363

    Google Scholar 

  • Waburton K, Lazarus J (1991) Tendency-distance models of social cohesion in animal groups. J Theor Biol 150:473–488

    Google Scholar 

  • Wark AR, Greenwood AK, Taylor EM, Yoshida K, Peichel CL (2011) Heritable differences in schooling behaviour among threespine stickleback populations revealed by a novel assay. PLoS ONE 6:e18316

    PubMed  CAS  Google Scholar 

  • Weihs D (1973) Hydromechanics of fish schooling. Nature 241:290–291

    Google Scholar 

  • Weihs D, Webb PW (1983) Optimalization of locomotion. In: Webb PW, Weihs D (eds) Fish biomechanics. Praeger, New York, pp 339–371

  • Williams MA (1997) The effect of experience on antipredator behavior of the coral reef fish, Caesio tile. Galaxea 13:167–179

    Google Scholar 

  • Williams MA, Coutant CC (2003) Modification of schooling behavior in larval atherinid fish Atherina mochon by heat exposure of eggs and larvae. Trans Am Fish Soc 132:638–645

    Google Scholar 

  • Wright D, Krause J (2006) Repeated measures of shoaling tendency in zebrafish (Danio rerio) and other small teleost fishes. Nat Protoc 1:1828–1831

    PubMed  CAS  Google Scholar 

Download references

Acknowledgments

Johann Delcourt is a PhD working for the FRS-FNRS (National Fund for Scientific Research of Belgium). This research was financially supported by the FNRS project 2.4617.08, 2.4569.06 and 2.4569.10F. We are also highly grateful to Jean-Louis Deneubourg (University of Brussels) for its comments and advices.

Author information

Authors and Affiliations

  1. Behavioral Biology Unit: Ethology and Animal Psychology, Laboratory of Fish and Amphibian Ethology, Department of Environmental Sciences and Management, Faculty of Sciences, Institute of Zoology, University of Liège, 22 Quai van Beneden, 4020, Liège, Belgium

    Johann Delcourt & Pascal Poncin

Authors
  1. Johann Delcourt
    View author publications

    You can also search for this author in PubMed Google Scholar

  2. Pascal Poncin
    View author publications

    You can also search for this author in PubMed Google Scholar

Corresponding author

Correspondence to Johann Delcourt.

Rights and permissions

Reprints and permissions

About this article

Cite this article

Delcourt, J., Poncin, P. Shoals and schools: back to the heuristic definitions and quantitative references. Rev Fish Biol Fisheries 22, 595–619 (2012). https://doi.org/10.1007/s11160-012-9260-z

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s11160-012-9260-z

Keywords

Search

Navigation