title: The leaf-area shrinkage effect can bias paleoclimate and ecology research.
abstract: OBJECTIVE
  Leaf area is a key trait that links plant form, function, and environment. Measures of leaf area can be biased because leaf area is often estimated from dried or fossilized specimens that have shrunk by an unknown amount. We tested the common assumption that this shrinkage is negligible.
  
  
  METHODS
  We measured shrinkage by comparing dry and fresh leaf area in 3401 leaves of 380 temperate and tropical species and used phylogenetic and trait-based approaches to determine predictors of this shrinkage. We also tested the effects of rehydration and simulated fossilization on shrinkage in four species.
  
  
  RESULTS
  We found that dried leaves shrink in area by an average of 22% and a maximum of 82%. Shrinkage in dried leaves can be predicted by multiple morphological traits with a standard deviation of 7.8%. We also found that mud burial, a proxy for compression fossilization, caused negligible shrinkage, and that rehydration, a potential treatment of dried herbarium specimens, eliminated shrinkage.
  
  
  CONCLUSIONS
  Our findings indicate that the amount of shrinkage is driven by variation in leaf area, leaf thickness, evergreenness, and woodiness and can be reversed by rehydration. The amount of shrinkage may also be a useful trait related to ecologically and physiological differences in drought tolerance and plant life history.
authors.affiliation: Department of Ecology and Evolutionary Biology, University of Arizona, P.O. Box 210088, Tucson, Arizona 85721, USA. bblonder@e-mail.arizona.edu
authors.forename: Benjamin
authors.initials: B
authors.lastname: Blonder
authors.forename: Vanessa
authors.initials: V
authors.lastname: Buzzard
authors.forename: Irena
authors.initials: I
authors.lastname: Simova
authors.forename: Lindsey
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authors.lastname: Sloat
authors.forename: Brad
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authors.lastname: Boyle
authors.forename: Rebecca
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authors.lastname: Lipson
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authors.lastname: Aguilar-Beaucage
authors.forename: Angelina
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authors.lastname: Andrade
authors.forename: Benjamin
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authors.lastname: Barber
authors.forename: Chris
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authors.lastname: Barnes
authors.forename: Dharma
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authors.lastname: Bushey
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authors.lastname: Cartagena
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authors.lastname: Chaney
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authors.lastname: Contreras
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authors.lastname: Cox
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authors.lastname: Cueto
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authors.lastname: Curtis
authors.forename: Mariah
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authors.lastname: Fisher
authors.forename: Lindsey
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authors.lastname: Furst
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authors.lastname: Gallegos
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authors.lastname: Hall
authors.forename: Amelia
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authors.lastname: Hauschild
authors.forename: Alex
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authors.lastname: Jerez
authors.forename: Nadja
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authors.lastname: Jones
authors.forename: Aaron
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authors.lastname: Klucas
authors.forename: Anita
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authors.lastname: Kono
authors.forename: Mary
authors.initials: M
authors.lastname: Lamb
authors.forename: Jacob David Ruiz
authors.initials: JD
authors.lastname: Matthai
authors.forename: Colten
authors.initials: C
authors.lastname: McIntyre
authors.forename: Joshua
authors.initials: J
authors.lastname: McKenna
authors.forename: Nicholas
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authors.lastname: Mosier
authors.forename: Maya
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authors.lastname: Navabi
authors.forename: Alex
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authors.lastname: Ochoa
authors.forename: Liam
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authors.lastname: Pace
authors.forename: Ryland
authors.initials: R
authors.lastname: Plassmann
authors.forename: Rachel
authors.initials: R
authors.lastname: Richter
authors.forename: Ben
authors.initials: B
authors.lastname: Russakoff
authors.forename: Holden St
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authors.lastname: Aubyn
authors.forename: Ryan
authors.initials: R
authors.lastname: Stagg
authors.forename: Marley
authors.initials: M
authors.lastname: Sterner
authors.forename: Emily
authors.initials: E
authors.lastname: Stewart
authors.forename: Ting Ting
authors.initials: TT
authors.lastname: Thompson
authors.forename: Jake
authors.initials: J
authors.lastname: Thornton
authors.forename: Parker J
authors.initials: PJ
authors.lastname: Trujillo
authors.forename: Trevor J
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authors.lastname: Volpe
authors.forename: Brian J
authors.initials: BJ
authors.lastname: Enquist
mesh_terms: D015982:Bias; D002980:Climate; D004463:Ecology; D019684:Magnoliopsida / Q000033:anatomy & histology / Q000145:classification / Q000502:physiology; D008954:Models, Biological; D018515:Plant Leaves / Q000033:anatomy & histology* / Q000187:drug effects / Q000502:physiology*; D012106:Research / Q000592:standards*; D013045:Species Specificity; D014867:Water / Q000494:pharmacology
chemical_list: D014867:Water