title: Monocot leaves are eaten less than dicot leaves in tropical lowland rain forests: correlations with toughness and leaf presentation.
abstract: OBJECTIVE
  In tropical lowland rain forest (TLRF) the leaves of most monocots differ from those of most dicots in two ways that may reduce attack by herbivores. Firstly, they are tougher. Secondly, the immature leaves are tightly folded or rolled until 50-100 % of their final length. It was hypothesized that (a) losses of leaf area to herbivorous invertebrates are generally greatest during leaf expansion and smaller for monocots than for dicots, and (b) where losses after expansion are appreciable any difference between monocots and dicots then is smaller than that found during expansion.
  
  
  METHODS
  At six sites on four continents, estimates were made of lamina area loss from the four most recently mature leaves of focal monocots and of the nearest dicot shoot. Measurements of leaf mass per unit area, and the concentrations of water and nitrogen were made for many of the species. In Panama, the losses from monocots (palms) and dicots were also measured after placing fully expanded palm leaflets and whole dicot leaves on trails of leaf-cutter ants.
  
  
  RESULTS
  At five of six sites monocots experienced significantly smaller leaf area loss than dicots. The results were not explicable in terms of leaf mass per unit area, or concentrations of water or nitrogen. At only one site was the increase in loss from first to fourth mature leaf significant (also large and the same in monocots and dicots), but the losses sustained during expansion were much smaller in the monocots. In the leaf-cutter ant experiment, losses were much smaller for palms than for dicots.
  
  
  CONCLUSIONS
  The relationship between toughness and herbivory is complex; despite the negative findings of some recent authors for dicots we hypothesize that either greater toughness or late folding can protect monocot leaves against herbivorous insects in tropical lowland rain forest, and that the relative importance varies widely with species. The difficulties of establishing unequivocally the roles of leaf toughness and leaf folding or rolling in a given case are discussed.
authors.affiliation: Plant Sciences Department, University of Cambridge, Downing Street, Cambridge CB2 3EA, UK. pjg12@cam.ac.uk
authors.forename: Peter J
authors.initials: PJ
authors.lastname: Grubb
authors.forename: Robyn V
authors.initials: RV
authors.lastname: Jackson
authors.forename: Ignacio M
authors.initials: IM
authors.lastname: Barberis
authors.forename: Jennie N
authors.initials: JN
authors.lastname: Bee
authors.forename: David A
authors.initials: DA
authors.lastname: Coomes
authors.forename: Nathaniel J
authors.initials: NJ
authors.lastname: Dominy
authors.forename: Marie Ann S
authors.initials: MA
authors.lastname: De La Fuente
authors.forename: Peter W
authors.initials: PW
authors.lastname: Lucas
authors.forename: Daniel J
authors.initials: DJ
authors.lastname: Metcalfe
authors.forename: Jens-Christian
authors.initials: JC
authors.lastname: Svenning
authors.forename: Ian M
authors.initials: IM
authors.lastname: Turner
authors.forename: Orlando
authors.initials: O
authors.lastname: Vargas
mesh_terms: D000821:Animal Feed; D000818:Animals; D007448:Invertebrates / Q000502:physiology; D019684:Magnoliopsida / Q000254:growth & development* / Q000469:parasitology / Q000502:physiology; D018515:Plant Leaves / Q000254:growth & development* / Q000469:parasitology / Q000502:physiology; D014197:Trees / Q000254:growth & development* / Q000469:parasitology / Q000502:physiology; D014329:Tropical Climate