title: Demographic consequences of chromatic leaf defence in tropical tree communities: do red young leaves increase growth and survival?
abstract: BACKGROUND
  Many tropical forest tree species delay greening their leaves until full expansion. This strategy is thought to provide newly flushing leaves with protection against damage by herbivores by keeping young leaves devoid of nutritive value. Because young leaves suffer the greatest predation from invertebrate herbivores, delayed greening could prevent costly tissue loss. Many species that delay greening also produce anthocyanin pigments in their new leaves, giving them a reddish tint. These anthocyanins may be fungicidal, protect leaves against UV damage or make leaves cryptic to herbivores blind to the red part of the spectrum.
  
  
  METHODS
  A comprehensive survey was undertaken of seedlings, saplings and mature trees in two diverse tropical forests: a rain forest in western Amazonia (Yasuní National Park, Ecuador) and a deciduous forest in Central America (Barro Colorado Island, Panamá). A test was made of whether individuals and species with delayed greening or red-coloured young leaves showed lower mortality or higher relative growth rates than species that did not.
  
  
  RESULTS
  At both Yasuní and Barro Colorado Island, species with delayed greening or red young leaves comprised significant proportions of the seedling and tree communities. At both sites, significantly lower mortality was found in seedlings and trees with delayed greening and red-coloured young leaves. While there was little effect of leaf colour on the production of new leaves of seedlings, diameter relative growth rates of small trees were lower in species with delayed greening and red-coloured young leaves than in species with regular green leaves, and this effect remained when the trade-off between mortality and growth was accounted for.
  
  
  CONCLUSIONS
  Herbivores exert strong selection pressure on seedlings for the expression of defence traits. A delayed greening or red-coloured young leaf strategy in seedlings appears to be associated with higher survival for a given growth rate, and may thus influence the species composition of later life stages.
authors.affiliation: Department of Evolution, Ecology & Organismal Biology, The Ohio State University, Columbus, OH 43210, USA. queenborough.1@osu.edu
authors.forename: Simon A
authors.initials: SA
authors.lastname: Queenborough
authors.forename: Margaret R
authors.initials: MR
authors.lastname: Metz
authors.forename: Renato
authors.initials: R
authors.lastname: Valencia
authors.forename: S Joseph
authors.initials: SJ
authors.lastname: Wright
mesh_terms: D000818:Animals; D000872:Anthocyanins / Q000502:physiology*; D017753:Ecosystem; D004484:Ecuador; D060434:Herbivory; D010176:Panama; D036226:Seedlings / Q000502:physiology*; D014197:Trees / Q000502:physiology*; D014329:Tropical Climate
chemical_list: D000872:Anthocyanins
keywords: Delayed greening; seedling herbivory; seedlings; tropical forest; young leaf colour