Skip to main content

Advertisement

Springer Nature Link
Log in

Social organisation of thornbill-dominated mixed-species flocks using social network analysis

  • Original Paper
  • Published:
Behavioral Ecology and Sociobiology Aims and scope Submit manuscript

Abstract

Mixed-species associations are a widespread phenomenon, comprising interacting heterospecific individuals which gain predator, foraging or social benefits. Avian flocks have traditionally been classified as monolithic species units, with species-wide functional roles, such as nuclear, active, passive, or follower. It has also been suggested that flocks are mutualistic interactions, where niches of participating species converge. However the species-level perspective has limited previous studies, because both interactions and benefits occur at the level of the individual. Social network analysis provides a set of tools for quantitative assessment of individual participation. We used mark-resighting methods to develop networks of nodes (colour-marked individuals) and edges (their interactions within flocks). We found that variation in flock participation across individuals within species, especially in the buff-rumped thornbill, encompassed virtually the entire range of variation across all individuals in the entire set of species. For example, female, but not male, buff-rumped thornbills had high network betweenness, indicating that they interact with multiple flocks, likely as part of a female-specific dispersal strategy. Finally, we provide new evidence that mixed-species flocking is mutualistic, by quantifying an active shift in individual foraging niches towards those of their individual associates, with implications for trade-off between costs and benefits to individuals derived from participating in mixed-species flocks. This study is, to our knowledge, the first instance of a heterospecific social network built on pairwise interactions.

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

Access this article

Log in via an institution

Subscribe and save

Springer+ Basic
$34.99 /Month
  • Get 10 units per month
  • Download Article/Chapter or eBook
  • 1 Unit = 1 Article or 1 Chapter
  • Cancel anytime
Subscribe now

Buy Now

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

Similar content being viewed by others

Explore related subjects

Discover the latest articles and news from researchers in related subjects, suggested using machine learning.

References

  • Bastian M, Heymann S (2010) Gephi: an open source software for exploring and manipulating networks, 0.8 alpha, https://gephi.org

  • Bejder L, Fletcher D, Brager S (1998) A method for testing association patterns of social animals. Anim Behav 56:719–725

    Article  PubMed  Google Scholar 

  • Bell HL (1980) Composition and seasonality of mixed-species feeding flocks of insectivorous birds in the Australian Capital Territory. Emu 80:227–232

    Article  Google Scholar 

  • Bell HL, Ford HA (1986) A comparison of the social-organization of 3 syntopic species of Australian Thornbill, Acanthiza. Behav Ecol Sociobiol 19:381–392

    Article  Google Scholar 

  • Boinski S, Scott PE (1988) Association of birds with monkeys in Costa-Rica. Biotropica 20:136–143

    Article  Google Scholar 

  • Buskirk WH (1976) Social-systems in a tropical forest Avifauna. Am Nat 110:293–310

    Article  Google Scholar 

  • Butts CT (2010) sna: Tools for Social Network Analysis, R package version 2.1, http://erzuli.ss.uci.edu/R.stuff/

  • Clutton-Brock T (2009) Cooperation between non-kin in animal societies. Nature 462:51–57

    Article  PubMed  CAS  Google Scholar 

  • Croft DP, James R, Thomas POR, Hathaway C, Mawdsley D, Laland KN, Krause J (2006) Social structure and co-operative interactions in a wild population of guppies (Poecilia reticulata). Behav Ecol Sociobiol 59:644–650

    Article  Google Scholar 

  • Croft DP, James R, Krause J (2008) Exploring animal social networks. Princeton University Press, Princeton

    Google Scholar 

  • Croft DP, Madden JR, Franks DW, James R (2011) Hypothesis testing in animal social networks. Trends Ecol Evol 26:502–507

    Article  PubMed  Google Scholar 

  • del Hoyo J, Elliott A, Christie DA (eds) (2007) Handbook of the birds of the world. Vol. 12. Picathartes to Tits and Chickadees. Lynx Edicions, Barcelona

    Google Scholar 

  • Dhondt AA (2012) Interspecific competition. Oxford Avian Biology Series. Oxford University Press, Oxford

    Google Scholar 

  • Diamond JM (1975) Assembly of species communities. In: Cody ML, Diamond JM (eds) Ecology and evolution of communities. Harvard University Press, Cambridge, pp 342–444

    Google Scholar 

  • Dolby AS, Grubb TC (1998) Benefits to satellite members in mixed-species foraging groups: an experimental analysis. Anim Behav 56:501–509

    Article  PubMed  Google Scholar 

  • Dolby AS, Grubb TC (2000) Social context affects risk taking by a satellite species in a mixed-species foraging group. Behav Ecol 11:110–114

    Article  Google Scholar 

  • Evans PGH (1982) Associations between seabirds and cetaceans — a review. Mammal Rev 12:187–206

    Article  Google Scholar 

  • Farley EA, Sieving KE, Contreras TA (2008) Characterizing complex mixed-species bird flocks using an objective method for determining species participation. J Ornithol 149:451–468

    Article  Google Scholar 

  • Franks DW, Ruxton GD, James R (2010) Sampling animal association networks with the gambit of the group. Behav Ecol Sociobiol 64:493–503

    Article  Google Scholar 

  • Goodale E, Kotagama SW (2005a) Alarm calling in Sri Lankan mixed-species bird flocks. Auk 122:108–120

    Article  Google Scholar 

  • Goodale E, Kotagama SW (2005b) Testing the roles of species in mixed-species bird flocks of a Sri Lankan rain forest. J Trop Ecol 21:669–676

    Article  Google Scholar 

  • Goodale E, Kotagama SW (2008) Response to conspecific and heterospecific alarm calls in mixed-species bird flocks of a Sri Lankan rainforest. Behav Ecol 19:887–894

    Article  Google Scholar 

  • Goodale E, Beauchamp G, Magrath RD, Nieh JC, Ruxton GD (2010) Interspecific information transfer influences animal community structure. Trends Ecol Evol 25:354–361

    Article  PubMed  Google Scholar 

  • Graves GR, Gotelli NJ (1993) Assembly of avian mixed-species flocks in Amazonia. Proc Natl Acad Sci USA 90:1388–1391

    Article  PubMed  CAS  Google Scholar 

  • Greenberg R (2000) Birds of many feathers: the formation and structure of mixed-species flocks of forest birds. In: Boinski S, Gerber PA (eds) On the move: how and why animals travel in groups. University of Chicago Press, Chicago, pp 521–558

    Google Scholar 

  • Harrison NM, Whitehouse MJ (2011) Mixed-species flocks: an example of niche construction? Anim Behav 81:675–682

    Article  Google Scholar 

  • Hartigan PM (1985) Computation of the dip statistic to test for unimodality. Appl Stat J Roy St C 34:320–325

    Google Scholar 

  • Hartigan JA, Hartigan PM (1985) The dip test of unimodality. Ann Stat 13:70–84

    Article  Google Scholar 

  • Hinde RA (1976) Interactions, relationships and social-structure. Man 11:1–17

    Article  Google Scholar 

  • Hino T (1998) Mutualistic and commensal organization of avian mixed-species foraging flocks in a forest of western Madagascar. J Avian Biol 29:17–24

    Article  Google Scholar 

  • Hino T (2000) Intraspecific differences in benefits from feeding in mixed-species flocks. J Avian Biol 31:441–446

    Article  Google Scholar 

  • Hino T (2002) Breeding bird community and mixed-species flocking in a deciduous broad-leaved forest in western Madagascar. Ornithol Sci 1:111–116

    Article  Google Scholar 

  • Horne JS, Garton EO, Krone SM, Lewis JS (2007) Analyzing animal movements using Brownian bridges. Ecology 88:2354–2363

    Article  PubMed  Google Scholar 

  • Hsieh FS, Chen CC (2011) Does niche-overlap facilitate mixed-species flocking in birds? J Ornithol 152:955–963

    Article  Google Scholar 

  • Hutchinson GE (1957) Population studies—animal ecology and demography—concluding remarks. Cold Spring Harb Sym 22:415–427

    Article  Google Scholar 

  • Hutto RL (1994) The composition and social-organization of mixed-species flocks in a tropical deciduous forest in Western Mexico. Condor 96:105–118

    Article  Google Scholar 

  • Krause J, Ruxton GD (2002) Living in groups. Oxford Series in Ecology and Environment. Oxford University Press, Oxford

    Google Scholar 

  • Krause J, Croft DP, James R (2007) Social network theory in the behavioural sciences: potential applications. Behav Ecol Sociobiol 62:15–27

    Article  Google Scholar 

  • Lack DL (1971) Ecological isolation in birds. Harvard University Press, Cambridge

    Google Scholar 

  • Latta SC, Wunderle JM (1996) The composition and foraging ecology of mixed-species flocks in pine forests of Hispaniola. Condor 98:595–607

    Article  Google Scholar 

  • Lepschi BJ (1993) Vegetation of Mulligans Flat, A.C.T. Cunninghamia 3:155–166

    Google Scholar 

  • Magrath RD, Pitcher BJ, Gardner JL (2007) A mutual understanding? Interspecific responses by birds to each other’s aerial alarm calls. Behav Ecol 18:944–951

    Article  Google Scholar 

  • Magrath RD, Pitcher BJ, Gardner JL (2009) An avian eavesdropping network: alarm signal reliability and heterospecific response. Behav Ecol 20:745–752

    Article  Google Scholar 

  • Maldonado-Coelho M, Marini MA (2004) Mixed-species bird flocks from Brazilian Atlantic forest: the effects of forest fragmentation and seasonality on their size, richness and stability. Biol Conserv 116:19–26

    Article  Google Scholar 

  • Manning AD, Wood JT, Cunningham RB, McIntyre S, Shorthouse DJ, Gordon IJ, Lindenmayer DB (2011) Integrating research and restoration: the establishment of a long-term woodland experiment in south-eastern Australia. Aust Zool 35:633–648

    Google Scholar 

  • Martin TE (1996) Fitness costs of resource overlap among coexisting bird species. Nature 380:338–340

    Article  CAS  Google Scholar 

  • McDonald DB (2007) Predicting fate from early connectivity in a social network. Proc Natl Acad Sci USA 104:10910–10914

    Article  PubMed  CAS  Google Scholar 

  • McIntyre S, Stol J, Harvey J, Nicholls AO, Campbell M, Reid A, Manning AD, Lindenmeyer D (2010) Biomass and floristic patterns in the ground layer vegetation of box-gum grassy eucalypt woodland in Goorooyarroo and Mulligans Flat Nature Reserves, Australian Capital Territory. Cunninghamia 11:319–357

    Google Scholar 

  • Morse DH (1970) Ecological aspects of some mixed-species foraging flocks of birds. Ecol Monogr 40:119–168

    Article  Google Scholar 

  • Moynihan M (1962) The organization and probable evolution of some mixed species flocks of neotropical birds. Smithson Misc Coll 143:1–140

    Google Scholar 

  • Newman MEJ (2003) Mixing patterns in networks. Phys Rev E 67

  • Newman MEJ (2010) Networks: an introduction. Oxford University Press, Oxford

    Google Scholar 

  • Nicholls JA, Double MC, Rowell DM, Magrath RD (2000) The evolution of cooperative and pair breeding in thornbills Acanthiza (Pardalotidae). J Avian Biol 31:165–176

    Article  Google Scholar 

  • Oh KP, Badyaev AV (2010) Structure of social networks in a Passerine bird: consequences for sexual selection and the evolution of mating strategies. Am Nat 176:E80–E89

    Article  PubMed  Google Scholar 

  • Oommen MA, Shanker K (2010) Shrewd alliances: mixed foraging associations between treeshrews, greater racket-tailed drongos and sparrowhawks on Great Nicobar Island, India. Biol Lett 6:304–307

    Article  PubMed  Google Scholar 

  • Powell GVN (1985) Sociobiology and adaptive significance of interspecific foraging flocks in the Neotropics. Ornithol Monogr 36:713–732

    Article  Google Scholar 

  • R Development Core Team (2010) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria, http://www.R-project.org

  • Schmidt KA, Dall SRX, van Gils JA (2010) The ecology of information: an overview on the ecological significance of making informed decisions. Oikos 119:304–316

    Article  Google Scholar 

  • Schoener TW (1974) Resource partitioning in ecological communities. Science 185:27–39

    Article  PubMed  CAS  Google Scholar 

  • Schoener TW (1983) Field experiments on interspecific competition. Am Nat 122:240–285

    Article  Google Scholar 

  • Sridhar H, Beauchamp G, Shanker K (2009) Why do birds participate in mixed-species foraging flocks? A large-scale synthesis. Anim Behav 78:337–347

    Article  Google Scholar 

  • Srinivasan U, Raza RH, Quader S (2010) The nuclear question: rethinking species importance in multi-species animal groups. J Anim Ecol 79:948–954

    Article  PubMed  Google Scholar 

  • Sundaresan SR, Fischhoff IR, Dushoff J (2009) Avoiding spurious findings of nonrandom social structure in association data. Anim Behav 77:1381–1385

    Article  Google Scholar 

  • Szekely T, Szep T, Juhasz T (1989) Mixed species flocking of tits (Parus spp.) — a field experiment. Oecologia 78:490–495

    Article  Google Scholar 

  • Thiollay JM (1999) Frequency of mixed species flocking in tropical forest birds and correlates of predation risk: an intertropical comparison. J Avian Biol 30:282–294

    Article  Google Scholar 

  • Thiollay JM (2003) Comparative foraging behavior between solitary and flocking insectivores in a Neotropical forest: does vulnerability matter? Ornithol Neotrop 14:47–65

    Google Scholar 

  • Thiollay JM, Jullien M (1998) Flocking behaviour of foraging birds in a neotropical rain forest and the antipredator defence hypothesis. Ibis 140:382–394

    Article  Google Scholar 

  • Valburg LK (1992) Flocking and frugivory — the effect of social groupings on resource use in the common bush-tanager. Condor 94:358–363

    Article  Google Scholar 

  • Waite TA, Grubb TC (1988) Copying of foraging locations in mixed-species flocks of temperate-deciduous woodland birds — an experimental-study. Condor 90:132–140

    Article  Google Scholar 

  • Wascher CAF, Szipl G, Boeckle M, Wilkinson A (2012) You sound familiar: carrion crows can differentiate between the calls of known and unknown heterospecifics. Anim Cogn 15:1015–1019

    Article  PubMed  Google Scholar 

  • Whitehead H (1995) Investigating structure and temporal scale in social organizations using identified individuals. Behav Ecol 6:199–208

    Article  Google Scholar 

  • Whitehead H (2008) Analyzing animal societies. University of Chicago Press, Chicago

    Google Scholar 

  • Whitehead H, Bejder L, Ottensmeyer CA (2005) Testing association patterns: issues arising and extensions. Anim Behav 69:e1–e6

    Article  Google Scholar 

  • Winterbottom JM (1943) On woodland bird parties in northern Rhodesia. Ibis 85:437–442

    Article  Google Scholar 

  • Zou FS, Chen GZ, Yang QF, Fellowes JR (2011) Composition of mixed-species flocks and shifts in foraging location of flocking species on Hainan Island, China. Ibis 153:269–278

    Article  Google Scholar 

Download references

Acknowledgements

We wish to thank the Australian Capital Territory Government and Mulligan’s Flat park rangers (P. Mills, G. Woodbridge, et al.) for providing access and continued support and the Fenner School for Environment and Society at the Australian National University for their collaboration. We thank Dr. Andrew Reeson, Dr. Louise Tierney and Lucy Aplin for their assistance, and Prof. Ben Sheldon, Dr. Colin Garroway, Dr. Eben Goodale and one anonymous reviewer for insightful comments. Funding assistance was provided by the European Research Council (AdG 250164) awarded to B. Sheldon, and Wolfson College Fieldwork Grant awarded to DRF.

Author information

Authors and Affiliations

  1. Edward Grey Institute of Field Ornithology, Department of Zoology, University of Oxford, South Parks Road, Oxford, OX1 3PS, UK

    Damien R. Farine

  2. College of Medicine, Biology & Environment, Australian National University, Canberra, Australia

    Peter J. Milburn

Authors
  1. Damien R. Farine
    View author publications

    Search author on:PubMed Google Scholar

  2. Peter J. Milburn
    View author publications

    Search author on:PubMed Google Scholar

Corresponding author

Correspondence to Damien R. Farine.

Additional information

Communicated by J. Krause.

Electronic supplementary material

Below is the link to the electronic supplementary material.

Online Resource 1

KML polygons showing and interactive overview of the study that can be viewed using Google Earth™. 1. Catching sites: two polygons showing the habitat patches in which all individuals were captured. 2. Overall home-range: home-range calculated from the observed locations of each group combined. 3. Individual home-range: home-range of each individual coloured by species and individually selectable with details of home-range size. (KML 205 kb).

Rights and permissions

Reprints and permissions

About this article

Cite this article

Farine, D.R., Milburn, P.J. Social organisation of thornbill-dominated mixed-species flocks using social network analysis. Behav Ecol Sociobiol 67, 321–330 (2013). https://doi.org/10.1007/s00265-012-1452-y

Download citation

  • Received:

  • Revised:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s00265-012-1452-y

Keywords