Springer Nature is making SARS-CoV-2 and COVID-19 research free. View research | View latest news | Sign up for updates

Traits and phylogenetic history contribute to network structure across Canadian plant–pollinator communities


Interaction webs, or networks, define how the members of two or more trophic levels interact. However, the traits that mediate network structure have not been widely investigated. Generally, the mechanism that determines plant-pollinator partnerships is thought to involve the matching of a suite of species traits (such as abundance, phenology, morphology) between trophic levels. These traits are often unknown or hard to measure, but may reflect phylogenetic history. We asked whether morphological traits or phylogenetic history were more important in mediating network structure in mutualistic plant-pollinator interaction networks from Western Canada. At the plant species level, sexual system, growth form, and flower symmetry were the most important traits. For example species with radially symmetrical flowers had more connections within their modules (a subset of species that interact more among one another than outside of the module) than species with bilaterally symmetrical flowers. At the pollinator species level, social species had more connections within and among modules. In addition, larger pollinators tended to be more specialized. As traits mediate interactions and have a phylogenetic signal, we found that phylogenetically close species tend to interact with a similar set of species. At the network level, patterns were weak, but we found increasing functional trait and phylogenetic diversity of plants associated with increased weighted nestedness. These results provide evidence that both specific traits and phylogenetic history can contribute to the nature of mutualistic interactions within networks, but they explain less variation between networks.

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

Fig. 1
Fig. 2
Fig. 3


  1. Almeida-Neto M, Ulrich W (2011) A straightforward computational approach for measuring nestedness using quantitative matrices. Environ Model Softw 26:173–178

  2. Bascompte J, Jordano P (2007) Plant-animal mutualistic networks: the architecture of biodiversity. Annu Rev Ecol Evol Syst 38:567–593

  3. Bascompte J, Jordano P, Melián CJ, Olesen JM (2003) The nested assembly of plant–animal mutualistic networks. Proc Natl Acad Sci 100:9383–9387

  4. Bascompte J, Jordano P, Olesen JM (2006) Asymmetric coevolutionary networks facilitate biodiversity maintenance. Science 312:431–433

  5. Bates D, Maechler M, Bolker B, et al. (2012) lme4: linear mixed-effects models using S4 classes. R package version 0.999999-0

  6. Blomberg SP, Garland T Jr, Ives AR (2003) Testing for phylogenetic signal in comparative data: behavioral traits are more labile. Evolution 57:717–745

  7. Blüthgen N, Menzel F, Blüthgen N (2006) Measuring specialization in species interaction networks. BMC Ecol 6:9. doi:10.1186/1472-6785-6-9

  8. Botta-Dukát Z (2005) Rao’s quadratic entropy as a measure of functional diversity based on multiple traits. J Veg Sci 16:533–540

  9. Bronstein JL, Alarcón R, Geber M (2006) The evolution of plant–insect mutualisms. New Phytol 172:412–428

  10. Cagnolo L, Salvo A, Valladares G (2011) Network topology: patterns and mechanisms in plant-herbivore and host-parasitoid food webs. J Anim Ecol 80:342–351

  11. Cane JH (1987) Estimation of bee size using intertegular span (Apoidea). J Kans Entomol Soc 60:145-147

  12. Cane JH, Sipes S (2006) Characterizing floral specialization by bees: analytical methods and a revised lexicon for oligolecty. Plant-Pollinator Interact Spec Gen Univ Chic Press Chic Lond, pp 99–122

  13. Cardinale BJ, Duffy JE, Gonzalez A, et al. (2012) Biodiversity loss and its impact on humanity. Nature 486:59–67

  14. Chamberlain SA, Holland JN (2009) Quantitative synthesis of context dependency in ant-plant protection mutualisms. Ecology 90:2384–2392. doi:10.1890/08-1490.1

  15. Clarke D, Whitney H, Sutton G, Robert D (2013) Detection and learning of floral electric fields by bumblebees. Science 340:66–69

  16. Cortis P, Vereecken N, Schiestl F, et al. (2009) Pollinator convergence and the nature of species’ boundaries in sympatric Sardinian Ophrys (Orchidaceae). Ann Bot 104:497–506

  17. Cribari-neto F, Zeileis A (2010) Beta regression in R. J Stat Softw 34:1–24

  18. Danieli-Silva A, De Souza JMT, Donatti AJ, et al. (2012) Do pollination syndromes cause modularity and predict interactions in a pollination network in tropical high-altitude grasslands? Oikos 121:35–43

  19. Donatti CI, Guimarães PR, Galetti M, et al. (2011) Analysis of a hyper-diverse seed dispersal network: modularity and underlying mechanisms. Ecol Lett 14:773–781

  20. Dormann CF (2011) How to be a specialist? Quantifying specialisation in pollination networks. Netw Biol 1:1–20

  21. Dunne JA, Williams RJ, Martinez ND (2002) Network structure and biodiversity loss in food webs: robustness increases with connectance. Ecol Lett 5:558–567

  22. Elle E, Elwell SL, Gielens GA (2012) The use of pollination networks in conservation. 1. Botany 90:525–534

  23. Encinas-Viso F, Revilla TA, Etienne RS (2012) Phenology drives mutualistic network structure and diversity. Ecol Lett 15:198–208

  24. Flora of North America Editorial (2002) Flora of North America: Magnoliophyta: Commelinidae (in Part): Cyperaceae, vol 23. Oxford University Press, Oxford

  25. Fontaine C, Collin CL, Dajoz I (2008) Generalist foraging of pollinators: diet expansion at high density. J Ecol 96:1002–1010

  26. Fritz SA, Purvis A (2010) Selectivity in mammalian extinction risk and threat types: a new measure of phylogenetic signal strength in binary traits. Conserv Biol 24:1042–1051

  27. Gibson RH, Knott B, Eberlein T, Memmott J (2011) Sampling method influences the structure of plant-pollinator networks. Oikos 120:822–831

  28. Greenleaf SS, Williams NM, Winfree R, Kremen C (2007) Bee foraging ranges and their relationship to body size. Oecologia 153:589–596

  29. Guimera R, Amaral LAN (2005a) Functional cartography of complex metabolic networks. Nature 433:895–900

  30. Guimera R, Amaral LAN (2005b) Cartography of complex networks: modules and universal roles. J Stat Mech Theory Exp 2005:P02001

  31. Harmon LJ, Glor RE (2010) Poor statistical performance of the Mantel test in phylogenetic comparative analyses. Evolution 64:2173–2178

  32. Hedges SB, Dudley J, Kumar S (2006) TimeTree: a public knowledge-base of divergence times among organisms. Bioinformatics 22:2971–2972

  33. Junker RR, Höcherl N, Blüthgen N (2010) Responses to olfactory signals reflect network structure of flower-visitor interactions. J Anim Ecol 79:818–823

  34. Junker RR, Blüthgen N, Brehm T, et al. (2013) Specialization on traits as basis for the niche-breadth of flower visitors and as structuring mechanism of ecological networks. Funct Ecol 27:329–341. doi:10.1111/1365-2435.12005

  35. Klinkenberg B (2012) E-Flora BC: atlas of the plants of British Columbia.

  36. Laliberté E, Legendre P (2010) A distance-based framework for measuring functional diversity from multiple traits. Ecology 91:299–305

  37. Maddison WP, Maddison DR (2011) Mesquite: a modular system for evolutionary analysis. Version 2.75.

  38. Michener CD (2007) The bees of the world, 2nd edn. John Hopkins University Press, Baltimore

  39. Oksanen J, Blanchet FG, Kindt R, et al. (2013) vegan: community ecology package. R package version 20-6.

  40. Olesen JM, Bascompte J, Dupont YL, Jordano P (2007) The modularity of pollination networks. Proc Natl Acad Sci 104:19891–19896

  41. Paine RT (1980) Food webs: linkage, interaction strength and community infrastructure. J Anim Ecol 49:667–685

  42. Paradis E, Claude J, Strimmer K (2004) APE: analyses of phylogenetics and evolution in R language. Bioinformatics 20:289–290

  43. Rezende EL, Lavabre JE, Guimarães PR, et al. (2007) Non-random coextinctions in phylogenetically structured mutualistic networks. Nature 448:925–928

  44. Sabo JL, Bastow JL, Power ME (2002) Length-mass relationships for adult aquatic and terrestrial invertebrates in a California watershed. J North Am Benthol Soc 21:336–343

  45. Santamaría L, Rodríguez-Gironés MA (2007) Linkage rules for plant–pollinator networks: trait complementarity or exploitation barriers? PLoS Biol 5:e31. doi:10.1371/journal.pbio.0050031

  46. Stang M, Klinkhamer PG, Van Der Meijden E (2006) Size constraints and flower abundance determine the number of interactions in a plant–flower visitor web. Oikos 112:111–121

  47. Thébault E, Fontaine C (2010) Stability of ecological communities and the architecture of mutualistic and trophic networks. Science 329:853–856

  48. Tylianakis JM, Tscharntke T, Lewis OT (2007) Habitat modification alters the structure of tropical host–parasitoid food webs. Nature 445:202–205

  49. Vázquez DP, Melián CJ, Williams NM, et al. (2007) Species abundance and asymmetric interaction strength in ecological networks. Oikos 116:1120–1127

  50. Vázquez DP, Chacoff NP, Cagnolo L (2009) Evaluating multiple determinants of the structure of plant-animal mutualistic networks. Ecology 90:2039–2046

  51. Webb CO, Donoghue MJ (2004) Phylomatic: tree assembly for applied phylogenetics. Mol Ecol Notes 5:181–183

  52. Webb CO, Ackerly DD, Kembel SW (2008) Phylocom: software for the analysis of phylogenetic community structure and trait evolution. Bioinformatics 24:2098–2100

  53. Wikstrom N, Savolainen V, Chase MW (2001) Evolution of the angiosperms: calibrating the family tree. Proc R Soc B Biol Sci 268:2211–2220. doi:10.1098/rspb.2001.1782

  54. Woodward G, Ebenman B, Emmerson M, et al. (2005) Body size in ecological networks. Trends Ecol Evol 20:402–409

Download references


We thank one anonymous reviewer and Handling Editor Steve Johnson for suggestions that greatly improved this manuscript. We acknowledge funding from NSERC-CANPOLIN, the Canadian Pollination Initiative. Additional funding was provided by an Alberta Conservation Association Grants in Biodiversity Program grant to Megan Evans, an Agriculture and AgriFood Canada grant to Mark Wonnick, NSERC-DG to Ralph Cartar, and NSERC-DG to Elizabeth Elle. This is publication 96 of NSERC-CANPOLIN. The experiments comply with the current laws of the country (Canada) in which the experiments were performed.

Author information

Correspondence to Scott A. Chamberlain.

Additional information

Communicated by Steven D. Johnson.

Electronic supplementary material

Rights and permissions

Reprints and Permissions

About this article

Verify currency and authenticity via CrossMark

Cite this article

Chamberlain, S.A., Cartar, R.V., Worley, A.C. et al. Traits and phylogenetic history contribute to network structure across Canadian plant–pollinator communities. Oecologia 176, 545–556 (2014).

Download citation


  • Mutualism
  • Interaction webs
  • Trophic levels
  • Morphological trait
  • Functional trait