title: Roles of climate and functional traits in controlling toothed vs. untoothed leaf margins.
abstract: OBJECTIVE
  Leaf-margin state (toothed vs. untoothed) forms the basis of several popular methods for reconstructing temperature. Some potential confounding factors have not been investigated with large data sets, limiting our understanding of the adaptive significance of leaf teeth and their reliability to reconstruct paleoclimate. Here we test the strength of correlations between leaf-margin state and deciduousness, leaf thickness, wood type (ring-porous vs. diffuse-porous), height within community, and several leaf economic variables.
  
  
  METHODS
  We assembled a trait database for 3549 species from six continents based on published and original data. The strength of associations between traits was quantified using correlational and principal axes approaches.
  
  
  RESULTS
  Toothed species, independent of temperature, are more likely to be deciduous and to have thin leaves, a high leaf nitrogen concentration, a low leaf mass per area, and ring-porous wood. Canopy trees display the highest sensitivity between leaf-margin state and temperature; subcanopy plants, especially herbs, are less sensitive.
  
  
  CONCLUSIONS
  Our data support hypotheses linking the adaptive significance of teeth to leaf thickness and deciduousness (in addition to temperature). Toothed species associate with the "fast-return" end of the leaf economic spectrum, providing another functional link to thin leaves and the deciduous habit. Accounting for these confounding factors should improve climate estimates from tooth-based methods.
authors.affiliation: Department of Earth and Environmental Sciences, Wesleyan University, Middletown, Connecticut 06459, USA.
authors.forename: Dana L
authors.initials: DL
authors.lastname: Royer
authors.forename: Daniel J
authors.initials: DJ
authors.lastname: Peppe
authors.forename: Elisabeth A
authors.initials: EA
authors.lastname: Wheeler
authors.forename: Ulo
authors.initials: U
authors.lastname: Niinemets
mesh_terms: D018533:Biomass; D002980:Climate; D005843:Geography; D016015:Logistic Models; D018515:Plant Leaves / Q000033:anatomy & histology* / Q000502:physiology*; D025341:Principal Component Analysis; D019655:Quantitative Trait, Heritable; D018401:Sample Size; D013696:Temperature