Plant Ecology

, Volume 192, Issue 2, pp 317–327 | Cite as

The adaptive value of young leaves being tightly folded or rolled on monocotyledons in tropical lowland rain forest: an hypothesis in two parts

  • Peter J. Grubb
  • Robyn V. Jackson


In tropical lowland rain forest, we find that species with the leaves tightly folded or rolled until they reach at least 50% of final length occur in 10 of the 15 monocot families with >100 species, and in 12 of the 24 monocot families with <100 species, but in only seven of the 212 dicot families (eudicots and magnoliids). Earlier researchers have described how examples of tightly folded and rolled leaves develop, but most have not considered the potentially adaptive value of this pattern of growth. We hypothesize that it is a protection against herbivorous invertebrates. For tropical and temperate dicots, the young leaves have been found to suffer much smaller losses to herbivores while folded and rolled than after they are unfolded or unrolled. Being folded or rolled until a late stage involves an ‘opportunity cost’ in the loss of photosynthesis. Among dicots, defences involving such a cost (notably late development of photosynthetic systems in pendent soft young leaves) are typical of shade-tolerant species, which have longer-lived leaves than light-demanders. In contrast, among monocots late folding and rolling are found in both shade-tolerators and light-demanders. We hypothesize that late folding and rolling bring a net advantage to monocots in general, whether shade-tolerant or light-demanding, despite the opportunity cost, because they mostly have fewer leaves per plant of a given size, and therefore an individual leaf is relatively more valuable to the plant. As a coda, we suggest that the ‘sleep movements’ of some tropical plants, and the circinate vernation of ferns and some cycads, provide protection against invertebrate herbivores through the apposition of two or more layers of leaf.


Herbivory Leaves Monocots Plant defences Tropical rain forest 



We thank Jennie Bee for the drawings in Fig. 1, Michael Lock for much help at Kew, Trevor Clifford, Peter Stevens, Jens-Christian Svenning, Ian Turner and Orlando Vargas for information about various plants and animals, William Foster, Julian Hibberd, Jennie Read, Lawren Sack and Mark Westoby for constructive criticism of earlier drafts, and three anonymous reviewers for their suggestions. The fieldwork by P.J.G. was made possible by a grant from the University of Cambridge, support from the Tropical Biology Association and a Mellon Fellowship.


  1. Benzing DH (2000) Bromeliaceae. Profile of an adaptive radiation. Cambridge University Press, CambridgeGoogle Scholar
  2. Bullock SH (1980) Demography of an undergrowth palm in littoral Cameroon. Biotropica 12:247–255CrossRefGoogle Scholar
  3. Bünning E, Moser I (1969) Interference of moonlight with photoperiodic measurement by plants, and their adaptive response. Proc Nat Acad Sci USA 62:1018–1022PubMedCrossRefGoogle Scholar
  4. Chapman RF (1974) Feeding in leaf-eating insects. Oxford University Press, OxfordGoogle Scholar
  5. Christophel DC, Hyland BPM (1993) Leaf atlas of Australian tropical rain forest trees. CSIRO, MelbourneGoogle Scholar
  6. Clayton WD (1978) Gramineae. In: Heywood VH (ed) Flowering plants of the world. Oxford University Press, OxfordGoogle Scholar
  7. Coley PD, Barrone JA (1996) Herbivory and plant defenses in tropical forests. Ann Rev Ecol Syst 27:305–335CrossRefGoogle Scholar
  8. Coley PD, Kursar TA (1996) Anti-herbivore defenses of young tropical leaves: physiological constraints and ecological trade-offs. In: Mulkey SS, Chazdon RL, Smith AP (eds) Tropical forest plant ecophysiology. Chapman and Hall, New YorkGoogle Scholar
  9. Corner EJH (1966) The natural history of palms. Weidenfeld and Nicolson, LondonGoogle Scholar
  10. Cullen J (1978) A preliminary survey of ptyxis (vernation) in the Angiosperms. Notes Roy Bot Gdn Edinb 37:161–214Google Scholar
  11. Darwin C (1880) The power of movement in plants. Murray, LondonGoogle Scholar
  12. de Steven D, Putz FE (1985) Mortality rates in rain forest palms. Principes 29:162–165Google Scholar
  13. Grubb PJ (1992) A positive distrust in simplicity—lessons from plant defences and from competition among plants and among animals. J Ecol 80:585–610CrossRefGoogle Scholar
  14. Hyland BPM, Whiffin T (1993) Australian tropical rain forest trees. CSIRO, MelbourneGoogle Scholar
  15. Jackson RV (1995) Insect herbivory on tropical Alphitonia (Rhamnaceae) species. Ph.D. thesis, James Cook University, TownsvilleGoogle Scholar
  16. Jackson RV, Kollmann J, Grubb PJ, Bee JN (1999) Insect herbivory on European tall-shrub species: the need to distinguish leaves before and after unfolding or unrolling, and the advantage of longitudinal sampling. Oikos 87:561–570CrossRefGoogle Scholar
  17. King M, Vincent JFV, Harris W (1996) Curling and folding of leaves of monocotyledons—a strategy for structural stiffness. N Z J Bot 34:411–416Google Scholar
  18. Lucas PW (2004) Dental functional morphology. Cambridge University Press, CambridgeGoogle Scholar
  19. Mabberley DJ (1997) The plant book, 2nd edn. Cambridge University Press, CambridgeGoogle Scholar
  20. McKenna DD, Farrell BD (2005) Molecular phylogenetics and evolution of host plant use in the neotropical rolled leaf ‘hispine’ beetle genus Cephaloleia (Chevrolat) (Chrysomelidae: Cassidinae). Mol Phylogen Evol 37:117–131CrossRefGoogle Scholar
  21. Reich PW, Ellsworth DS, Walters MB et al (1999) Generality of leaf traits relationships: a test across six biomes. Ecology 80:1955–1969CrossRefGoogle Scholar
  22. Richards PW (1996) Tropical rain forest, 2nd edn. Cambridge University Press, CambridgeGoogle Scholar
  23. Riley ND, Hahnewald E (1963) Insects in colour. Blandford Press, LondonGoogle Scholar
  24. Skutch AF (1927) Anatomy of leaf of banana, Musa sapientium L. var. Hort. Gros Michel. Bot Gaz 84:337–391CrossRefGoogle Scholar
  25. Sporne KR (1974) The morphology of gymnosperms, 2nd edn. Hutchinson, LondonGoogle Scholar
  26. Sporne KR (1975) The morphology of gymnosperms, 4th edn. Hutchinson, LondonGoogle Scholar
  27. Stevens PF (2007) Angiosperm phylogeny website, Version 7, May 2006, updated. http:// Cited 26 March 2007Google Scholar
  28. Stewart WN, Rothwell GW (1993) Paleobotany and the evolution of plants. Cambridge University Press, CambridgeGoogle Scholar
  29. Strong DR (1977) Rolled-leaf hispine beetles (Chrysomelidae) and Zingiberales host plants in Middle America. Biotropica 9:156–159CrossRefGoogle Scholar
  30. Ticktin T (2003) Relationships between El Niño Southern Oscillation and demographic patterns in a substitute food plant for collared peccaries in Panama. Biotropica 35:189–197Google Scholar
  31. Ueda MJ, Sugimoto T, Sawai Y, et al (2003) Chemical studies on plant leaf movement controlled by a biological clock. Pure Appl Chem 75:353–358CrossRefGoogle Scholar
  32. Venkatanaraya G (1957) On certain aspects of the development of the leaf of Cocos nucifera (L.). Phytomorphology 7:297–305Google Scholar
  33. Webb LJ (1959) A physiognomic classification of Australian rain forests. J Ecol 47:551–570CrossRefGoogle Scholar
  34. Wilder GJ (1976) Structure and development of leaves in Carludovica palmata (Cyclanthaceae) with reference to other Cyclanthaceae. Am J Bot 63:1237–1256CrossRefGoogle Scholar
  35. Wilder GJ (1981) Structure and development of Cyclanthus bipartitus Poit. (Cyclanthaceae) with reference to other Cyclanthaceae. II Adult leaf. Bot Gaz 142:222–236CrossRefGoogle Scholar
  36. Wilf P, Labandeira CC, Kress JW et al (2000) Timing the radiations of leaf beetles: hispines on gingers from the latest Cretaceous to recent. Science 289:291–294PubMedCrossRefGoogle Scholar
  37. Xiang H, Chen J (2004) Interspecific variation of plant traits associated with resistance to herbivory among four species of Ficus (Moraceae). Ann Bot 94:377–384PubMedCrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media B.V. 2007

Authors and Affiliations

  1. 1.Plant Sciences DepartmentUniversity of CambridgeCambridgeUK
  2. 2.19 Cananga CloseKamerungaAustralia

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