Association tendency and preference for heterospecifics in an invasive species

in Behaviour
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Animals gain benefits by forming groups with phenotypically and behaviourally similar individuals. The most common groups are homogenous, composed by conspecifics, although in some cases associations of similar organisms of different species have been reported when individuals benefit from it. In this study, we tested the prediction that the Trinidadian guppy, Poecilia reticulata, a fish that has successfully invaded at least 70 countries, will shoal with heterospecifics. We measured shoaling tendency and shoal companion preference in wild-caught female guppies when they encounter two heterospecific species: the native Poecilia picta and the non-native Poecilia sphenops, a poeciliid recently introduced in Trinidad. Our results show that guppies have a higher tendency to shoal with conspecifics; if the alternative is be alone, they readily shoal with both species even when they have had no previous experience with other poeciliids. Individuals in these associations could benefit from safety in numbers along with other advantages of group living. This predisposition to associate with other species that share similar ecological conditions could explain the guppy’s success as invasive species as it enables them to increase their shoal size during the first stages of invasion and thus avoid Allee effects.



AlcarazC.Vila-GispertA.Garcia-BerthouE. (2005). Profiling invasive fish species, the importance of phylogeny and human use. — Divers. Distrib. 11: 289-298.

BarberI. (2003). Parasites and size-assortative schooling in three-spined sticklebacks. — Oikos 101: 331-337.

CampobelloD.SaraM.HareJ.F. (2012). Under my wing, lesser kestrels and jackdaws derive reciprocal benefits in mixed-species colonies. — Behav. Ecol. 23: 425-433.

ChapmanB.B.WardA.J.W.KrauseJ. (2008). Schooling and learning: early social environment predicts social learning ability in the guppy, Poecilia reticulata. — Anim. Behav. 76: 923-929.

CouzinI.D. (2009). Collective cognition in animal groups. — Trends Cogn. Sci. 13: 36-43.

CouzinI.D.KrauseJ. (2003). Self-organization and collective behavior in vertebrates. — Adv. Stud. Behav. 32: 1-75.

CroftD.P.ArrowsmithB.J.BielbyJ.SkinnerK.WhiteE.CouzinI.D.MagurranA.E.RamnarineI.KrauseJ. (2003). Mechanisms underlying shoal composition in the Trinidadian guppy, Poecilia reticulata. — Oikos 100: 429-438.

CroftD.P.KrauseJ.JamesR. (2004). Social networks in the guppy (Poecilia reticulata). — Proc. Roy. Soc. Lond. B: Biol. Sci. 271: S516-S519.

CroftD.P.JamesR.ThomasP.O.R.HathawayC.MawdsleyD.LalandK.N.KrauseJ. (2006). Social structure and co-operative interactions in a wild population of guppies (Poecilia reticulata). — Behav. Ecol. Sociobiol. 59: 644-650.

CroftD.P.DardenS.K.RuxtonG.D. (2009a). Predation risk as a driving force for phenotypic assortment: a cross-population comparison. — Proc. Roy. Soc. Lond. B: Biol. Sci. 276: 1899-1904.

CroftD.P.KrauseJ.DardenS.K.RamnarineI.W.FariaJ.J.JamesR. (2009b). Behavioural trait assortment in a social network: patterns and implications. — Behav. Ecol. Sociobiol. 63: 1495-1503.

DeaconA.E.RamnarineI.W.MagurranA.E. (2011). How reproductive ecology contributes to the spread of a globally invasive fish. — PLOS One 6: e24416.

DrakeJ.M.KramerA.M. (2011). Allee effects. — Nat. Educ. Knowl. 3(10): 2.

DyerJ.R.G.CroftD.P.MorrellL.J.KrauseJ. (2009). Shoal composition determines foraging success in the guppy. — Behav. Ecol. 20: 165-171.

EdenbrowM.DardenS.K.RamnarineI.W.EvansJ.P.JamesR.CroftD.P. (2011). Environmental effects on social interaction networks and male reproductive behaviour in guppies, Poecilia reticulata. — Anim. Behav. 81: 551-558.

GriffithsS.W.MagurranA.E. (1997). Familiarity in schooling fish, how long does it take to acquire?Anim. Behav. 53: 945-949.

GriffithsS.W.MagurranA.E. (1998). Sex and schooling behaviour in the Trinidadian guppy. — Anim. Behav. 56: 689-693.

GriffithsS.W.MagurranA.E. (1999). Schooling decisions in guppies (Poecilia reticulata) are based on familiarity rather than kin recognition by phenotype matching. — Behav. Ecol. Sociobiol. 45: 437-443.

HamiltonW.D. (1971). Geometry for the selfish herd. — J. Theor. Biol. 31: 295-311.

HolwayD.A.SuarezA.V. (1999). Animal behavior, an essential component of invasion biology. — Trends Ecol. Evol. 14: 328-330.

HurlbertS.H. (1984). Pseudoreplication and the design of ecological field experiments. — Ecol. Monogr. 54: 187-211.

KennyJ.S. (1995). Views from the bridge, a memoir on the freshwater fishes of Trinidad. — Julian S. Kenny, Maracas, St. Joseph, Trinidad and Tobago.

KrauseJ.RuxtonG.D. (2002). Living in groups. — Oxford University Press, New York, NY.

KrauseJ.WardA.J.W.JacksonA.L.RuxtonG.D.JamesR.CurrieS. (2005). The influence of differential swimming speeds on composition of multi-species fish shoals. — J. Fish Biol. 67: 866-872.

LandeauL.TerborghJ. (1986). Oddity and the confusion effect in predation. — Anim. Behav. 34: 1372-1380.

MagurranA.E. (2005). Evolutionary ecology, the Trinidadian guppy. — Oxford University Press, Oxford.

MagurranA.E.SeghersB.H. (1991). Variation in schooling and aggression amongst guppy (Poecilia reticulata) populations in Trinidad. — Behaviour 118: 214-234.

MagurranA.E.SeghersB.H.ShawP.W.CarvalhoG.R. (1995). The behavioral diversity and evolution of guppy, Poecilia reticulata, populations in Trinidad. — Adv. Stud. Behav. 24: 155-202.

MorseD.H. (1977). Feeding behaviour and predator avoidance in heterospecific groups. — BioScience 27: 332-339.

PavlovD.S.KasumyanA.O. (2000). Patterns and mechanisms of schooling behaviour in fish: a review. — J. Ichthyol. 40: 163-231.

PitcherT.J. (1983). Heuristic definitions of fish shoaling behaviour. — Anim. Behav. 31: 611-613.

PiyapongC.ButlinR.K.FariaJ.J.ScrutonK.J.WangJ.KrauseJ. (2011). Kin assortment in juvenile shoals in wild guppy populations. — Heredity 106: 749-756.

PowellG.V.N. (1989). On the possible contribution of mixed species flocks to species richness in neotropical avifaunas. — Behav. Ecol. Sociobiol. 24: 387-393.

SazimaC.KrajewskiJ.P.BonaldoR.M.SazimaI. (2007). Nuclear-follower foraging associations of reef fishes and other animals at an oceanic archipelago. — Environ. Biol. Fish. 80: 351-361.

SchluppI.RyanM.J. (1996). Mixed-species shoals and the maintenance of a sexual-asexual mating system in mollies. — Anim. Behav. 52: 885-890.

SieversC.WillingE.M.HoffmannM.DreyerC.RamnarineI.MagurranA.E. (2012). Reasons for the invasive success of a guppy (Poecilia reticulata) population in Trinidad. — PLOS One 7: e38404.

SokalR.R.RohlfF.J. (1981). Biometry, the principles and practice of statistics in biological reserch. — W.H. Freeman, San Francisco, CA.

StenslandE.AngerbornA.BerggrenP. (2003). Mixed-species groups in mammals. — Mamm. Rev. 33: 205-223.

StephensP.A.SutherlandW.J. (1999). Consequences of the Allee effect for behaviour, ecology and conservation. — Trends Ecol. Evol. 14: 401-405.

ValeroA.Macías GarciaC.MagurranA.E. (2008). Heterospecific harassment of native endangered fishes by invasive guppies in Mexico. — Biol. Lett. 4: 149-152.

WarburtonK.LeesN. (1996). Species discrimination in guppies, learned responses to visual cues. — Anim. Behav. 52: 371-378.

WardA.J.W.KrauseJ. (2001). Body length assortative shoaling in the European minnow, Phoxinus phoxinus. — Anim. Behav. 62: 617-621.

WardA.J.W.AxfordS.KrauseJ. (2002). Mixed-species shoaling in fish: the sensory mechanisms and costs of shoal choice. — Behav. Ecol. Sociobiol. 52: 182-187.


  • Diagram of the tank set up. For the shoaling tendency trials, one of the bottles remained empty and for the shoaling preference part each bottle contained a shoal. Time spent shoaling was recorded whenever the fish was within one body length of the bottle containing a shoal.

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  • Shoaling tendency. White bars represent the time (out of a possible maximum of 600 s) focals from Acono (where guppies were the only poeciliid present), Charlieville (where guppies coexist with P. picta) or Maraval (where guppies coexist with P. sphenops) spent with P. reticulata shoals, light grey is for P. picta shoals and dark grey for P. sphenops shoals. Dotted line shows the expected time of association if randomly swimming (30 s). Horizontal lines in the bars represent the median and boxes the 50% of the data values.

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  • Preference between conspecifics and heterospecifics. Vertical axis shows the difference in time spent with one over an other shoal. Positive numbers show preference for conspecifics and negative numbers show preference for heterospecifics. White bars represent the difference in time between P. reticulata and P. picta, grey bars represent the difference between P. reticulata and P. sphenops. Horizontal lines in the bars represent the median and boxes the 50% of data values. Guppies from Acono had no preference between conspecifics and heterospecifics (one-way t-test, t13>1.51, p>0.13); guppies from Charlieville (where they coexist with P. picta) had no preference between P. picta and conspecifics (one-way t-test, t13=1.586, p=0.137) but preferred conspecifics over P. sphenops (one-way t-test, t13=2.62, p=0.021), and focals from Maraval (where they coexist with P. sphenops) showed no preference between P. sphenops and conspecifics (one-way t-test, t13=2.11, p=0.054) but preferred conspecifics over P. picta (one-way t-test, t13=2.88, p=0.013).

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