title: The birds and the bees: pollinator behaviour and variation in the mating system of the rare shrub Grevillea macleayana.
abstract: OBJECTIVE
  In Australia, honey-bees have invaded systems that evolved without social insect pollinators, where many plants are adapted to vertebrate pollination. Behavioural differences between pollinators are likely to influence mating patterns, but few studies have examined this empirically in long-lived, woody, perennials. It was shown previously that outcrossing rates in Grevillea macleayana vary among populations. Here tests were conducted to determine whether the behaviour of birds and honey-bees differed between a population previously found to be highly outcrossed and two inbreeding populations.
  
  
  METHODS
  Visit frequencies and movement patterns of the visitors to inflorescences at three sites over two seasons were compared. A caging experiment was used to test the effects of excluding birds on pollen removal from newly opened flowers and on pollen deposition on stigmas that had been washed clean.
  
  
  RESULTS
  Honey-bees were the most frequent visitors overall, but honeyeaters were more frequent visitors in the population previously found to have a high outcrossing rate than they were in either of the other populations. More visits by honeyeaters were from distant plants. Pollen removal did not vary greatly among sites, and was not affected by bird exclusion; however, more pollen was deposited on the stigmas of cleaned pollen presenters in the population previously observed to be highly outcrossing than in the other two. This high level of pollen deposition was reduced by experimental bird exclusion.
  
  
  CONCLUSIONS
  Honey-bees were the most frequent visitors, by an order of magnitude, and excluding vertebrates revealed that bees were removing most of the pollen but deposited fewer pollen grains on stigmas. Birds were more frequent visitors at the site previously found to be outcrossing than the other two sites, and they moved further between plants and visited fewer inflorescences on a plant during a foraging bout than bees did. These characteristics of bird visits to G. macleayana would be sufficient to produce significant variation in outcrossing rates among sites.
authors.affiliation: University of Wollongong in Dubai, United Arab Emirates. robwhelan@uowdubai.ac.ae
authors.forename: Robert J
authors.initials: RJ
authors.lastname: Whelan
authors.forename: David J
authors.initials: DJ
authors.lastname: Ayre
authors.forename: Fiona M
authors.initials: FM
authors.lastname: Beynon
mesh_terms: D000704:Analysis of Variance; D000818:Animals; D001516:Bees / Q000502:physiology*; D001522:Behavior, Animal / Q000502:physiology*; D001717:Birds / Q000502:physiology*; D006722:Honey; D011058:Pollen / Q000502:physiology; D054817:Pollination / Q000502:physiology*; D039105:Proteaceae / Q000502:physiology*; D012098:Reproduction; D012621:Seasons