The leaf-area shrinkage effect can bias paleoclimate and ecology research

Benjamin Blonder, Vanessa Buzzard, Irena Simova, Lindsey Sloat, Brad Boyle, Rebecca Lipson, Brianna Aguilar-Beaucage, Angelina Andrade, Benjamin Barber, Chris Barnes, Dharma Bushey, Paulina Cartagena, Max Chaney, Karina Contreras, Mandarava Cox, Maya Cueto, Cannon Curtis, Mariah Fisher, Lindsey Furst, Jessica GallegosRuby Hall, Amelia Hauschild, Alex Jerez, Nadja Jones, Aaron Klucas, Anita Kono, Mary Lamb, Jacob David Ruiz Matthai, Colten Mcintyre, Joshua Mckenna, Nicholas Mosier, Maya Navabi, Alex Ochoa, Liam Pace, Ryland Plassmann, Rachel Richter, Ben Russakoff, Holden St Aubyn, Ryan Stagg, Marley Sterner, Emily Stewart, Ting Ting Thompson, Jake Thornton, Parker J. Trujillo, Trevor J. Volpe, Brian J. Enquist

Research output: Contribution to journalArticlepeer-review

51 Scopus citations


• Premise of the Study: 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. • Key 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.

Original languageEnglish (US)
Pages (from-to)1756-1763
Number of pages8
JournalAmerican journal of botany
Issue number11
StatePublished - 2012


  • Dry area
  • Fresh area
  • Leaf area
  • Leaf mass per area
  • Leaf size
  • Shrinkage specific leaf area
  • Stomatal density
  • Vein density

ASJC Scopus subject areas

  • Ecology, Evolution, Behavior and Systematics
  • Genetics
  • Plant Science


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