title: Geographical differentiation in floral traits across the distribution range of the Patagonian oil-secreting Calceolaria polyrhiza: do pollinators matter?
abstract: OBJECTIVE
  The underlying evolutionary processes of pollinator-driven floral diversification are still poorly understood. According to the Grant-Stebbins model speciation begins with adaptive local differentiation in the response to spatial heterogeneity in pollinators. Although this crucial process links the micro- and macroevolution of floral adaptation, it has received little attention. In this study geographical phenotypic variation was investigated in Patagonian Calceolaria polyrhiza and its pollinators, two oil-collecting bee species that differ in body size and geographical distribution.
  
  
  METHODS
  Patterns of phenotypic variation were examined together with their relationships with pollinators and abiotic factors. Six floral and seven vegetative traits were measured in 45 populations distributed across the entire species range. Climatic and edaphic parameters were determined for 25 selected sites, 2-16 bees per site of the most frequent pollinator species were captured, and a critical flower-bee mechanical fitting trait involved in effective pollination was measured. Geographical patterns of phenotypic and environmental variation were examined using uni- and multivariate analyses. Decoupled geographical variation between corolla area and floral traits related to the mechanical fit of pollinators was explored using a Mantel test.
  
  
  RESULTS
  The body length of pollinators and the floral traits related to mechanical fit were strongly correlated with each other. Geographical variation of the mechanical-fit-related traits was decoupled from variation in corolla size; the latter had a geographical pattern consistent with that of the vegetative traits and was mainly affected by climatic gradients.
  
  
  CONCLUSIONS
  The results are consistent with pollinators playing a key role in shaping floral phenotype at a geographical scale and promoting the differentiation of two floral ecotypes. The relationship between the critical floral-fit-related trait and bee length remained significant even in models that included various environmental variables and an allometric predictor (corolla area). The abiotic environment also has an important role, mainly affecting floral size. Decoupled geographical variation between floral mechanical-fit-related traits and floral size would represent a strategy to maintain plant-pollinator phenotypic matching in this environmentally heterogeneous area.
authors.affiliation: Laboratorio de Ecología Evolutiva y Biología Floral, Instituto Multidisciplinario de Biología Vegetal (IMBIV), CONICET-Universidad Nacional de Córdoba, Córdoba, Argentina.
authors.forename: Andrea
authors.initials: A
authors.lastname: Cosacov
authors.forename: Andrea A
authors.initials: AA
authors.lastname: Cocucci
authors.forename: Alicia N
authors.initials: AN
authors.lastname: Sérsic
mesh_terms: D000818:Animals; D001118:Argentina; D001516:Bees / Q000502:physiology*; D002980:Climate; D035264:Flowers / Q000033:anatomy & histology* / Q000502:physiology; D005843:Geography; D015999:Multivariate Analysis; D010641:Phenotype; D010938:Plant Oils / Q000378:metabolism*; D054817:Pollination; D019655:Quantitative Trait, Heritable; D012044:Regression Analysis; D028083:Scrophulariaceae / Q000033:anatomy & histology* / Q000502:physiology*
chemical_list: D010938:Plant Oils
keywords: Abiotic environmental gradients; Calceolaria; Patagonia; bee morphology; floral ecotypes; geographical range; local adaptation; oil-collecting bees; oil-offering flowers; phenotypic covariance; specialized pollination; speciation; vegetative morphology