title: Novel adaptation to hawkmoth pollinators in Clarkia reduces efficiency, not attraction of diurnal visitors.
abstract: OBJECTIVE
  Plant populations experiencing divergent pollination environments may be under selection to modify floral traits in ways that increase both attractiveness to and efficiency of novel pollinators. These changes may come at the cost of reducing overall effectiveness of other pollinators. The goal of this study was to examine differences in attractiveness and efficiency between Clarkia concinna and C. breweri, sister species of annual plants with parapatric distributions.
  
  
  METHODS
  An assessment was made as to whether observed differences in visitors between natural populations are driven by differences in floral traits or differences in the local pollination environment. Differences in floral attractiveness were quantified by setting out arrays of both species in the geographical range of each species and exposing both species to nocturnal hawkmoths (Hyles lineata) in flight cages. Differences in visitor efficiency were estimated by measuring stigma-visitor contact frequency and pollen loads for diurnal visitors, and pollen deposition on stigmas for hawkmoths.
  
  
  RESULTS
  The composition of visitors to arrayed plants was similar between plant species at any particular site, but highly divergent among sites, and reflected differences in visitors to natural populations. Diurnal insects visited both species, but were more common at C. concinna populations. Hummingbirds and hawkmoths were only observed visiting within the range of C. breweri. Despite attracting similar species when artificially presented together, C. concinna and C. breweri showed large differences in pollinator efficiency. All visitors except hawkmoths pollinated C. concinna more efficiently.
  
  
  CONCLUSIONS
  Differences in the available pollinator community may play a larger role than differences in floral traits in determining visitors to natural populations of C. concinna and C. breweri. However, floral traits mediate differences in pollinator efficiency. Increased effectiveness of the novel hawkmoth pollinator on C. breweri comes at relatively little cost in attractiveness to other visitors, but at large cost in their efficiency as pollinators.
authors.affiliation: Department of Ecology and Evolution, University of California, Santa Cruz, 1156 High Street, Santa Cruz, CA 95064, USA.
authors.forename: Timothy J
authors.initials: TJ
authors.lastname: Miller
authors.forename: Robert A
authors.initials: RA
authors.lastname: Raguso
authors.forename: Kathleen M
authors.initials: KM
authors.lastname: Kay
mesh_terms: D000222:Adaptation, Physiological; D000704:Analysis of Variance; D000818:Animals; D002140:California; D031663:Clarkia / Q000033:anatomy & histology / Q000502:physiology*; D003624:Darkness; D035264:Flowers / Q000033:anatomy & histology / Q000502:physiology; D005843:Geography; D009036:Moths / Q000502:physiology*; D011058:Pollen / Q000502:physiology; D054817:Pollination / Q000502:physiology*; D011157:Population Dynamics
keywords: Clarkia breweri; Clarkia concinna; Hyles lineata; floral traits; parapatric divergence; pollinator effectiveness; pollinator efficiency; pollinator shift; speciation