Leaves at low versus high rainfall: coordination of structure, lifespan and physiology.

•  Across species, leaf lifespan (LL) tends to be correlated with leaf mass per area (LMA). Previously we found that Australian perennial species from low-rainfall sites had c . 40% shorter LL at a given LMA than high-rainfall species. •  Here we relate indices of leaf strength (work to shear, W shear , and tissue toughness) to LL and LMA across the same suite of species. W shear is the work required to cut a leaf with a blade; W shear divided by leaf thickness gives tissue toughness. •  Low- and high-rainfall species did not differ in their LL at a given W shear , but dry-site species had lower W shear at a given LMA, leading to the observed LL - LMA shift with rainfall. These patterns were driven by 50% lower tissue toughness in dry-site species. •  The lower toughness was linked with high leaf N concentration, which is known to enhance water conservation during photosynthesis in low-rainfall species. Our results suggest that a significant cost of this strategy is reduced LL for a given investment in leaf tissue (LMA).

[1]  Mark Westoby,et al.  A leaf-height-seed (LHS) plant ecology strategy scheme , 1998, Plant and Soil.

[2]  A. Escudero,et al.  Effects of leaf longevity and retranslocation efficiency on the retention time of nutrients in the leaf biomass of different woody species , 1992, Oecologia.

[3]  P. Reich,et al.  Leaf lifespan as a determinant of leaf structure and function among 23 amazonian tree species , 1991, Oecologia.

[4]  V. Brown,et al.  Leaf palatability, life expectancy and herbivore damage , 1986, Oecologia.

[5]  C. Ohmart,et al.  Effects of food quality, particularly nitrogen concentrations, of Eucalyptus blakelyi foliage on the growth of Paropsis atomaria larvae (Coleoptera: Chrysomelidae) , 1985, Oecologia.

[6]  H. Mooney,et al.  Compromises between water-use efficiency and nitrogen-use efficiency in five species of California evergreens , 1983, Oecologia.

[7]  H. Mooney,et al.  Photosynthetic capacity and carbon allocation patterns in diverse growth forms of Eucalyptus , 2004, Oecologia.

[8]  P. Coley,et al.  Effects of plant growth rate and leaf lifetime on the amount and type of anti-herbivore defense , 2004, Oecologia.

[9]  P. Reich,et al.  Convergence towards higher leaf mass per area in dry and nutrient‐poor habitats has different consequences for leaf life span , 2002 .

[10]  N. Weber,et al.  Common Slope Tests for Bivariate Errors‐in‐Variables Models , 2002 .

[11]  P. Reich,et al.  Strategy shifts in leaf physiology, structure and nutrient content between species of high‐ and low‐rainfall and high‐ and low‐nutrient habitats , 2001 .

[12]  R. Villar,et al.  Comparison of leaf construction costs in woody species with differing leaf life-spans in contrasting ecosystems. , 2001, The New phytologist.

[13]  Ian J. Wright,et al.  Relationships between leaf lifespan and structural defences in a low-nutrient, sclerophyll flora , 2001 .

[14]  Ülo Niinemets,et al.  GLOBAL-SCALE CLIMATIC CONTROLS OF LEAF DRY MASS PER AREA, DENSITY, AND THICKNESS IN TREES AND SHRUBS , 2001 .

[15]  N. Dominy,et al.  Mechanical Defences to Herbivory , 2000 .

[16]  P. Ryser,et al.  Ecological significance of leaf life span among Central European grass species , 2000 .

[17]  I. Noble,et al.  A framework for understanding the relationship between environment and vegetation based on the surface area to volume ratio of leaves , 2000 .

[18]  J. Read,et al.  Characterising sclerophylly: some mechanical properties of leaves from heath and forest , 2000, Oecologia.

[19]  D. Soltis,et al.  Angiosperm phylogeny inferred from multiple genes as a tool for comparative biology , 1999, Nature.

[20]  Mark Westoby,et al.  EVOLUTIONARY DIVERGENCES IN LEAF STRUCTURE AND CHEMISTRY, COMPARING RAINFALL AND SOIL NUTRIENT GRADIENTS , 1999 .

[21]  N. Aranwela,et al.  Methods of assessing leaf‐fracture properties , 1999 .

[22]  I. Noble,et al.  A theoretical approach to linking the composition and morphology with the function of leaves , 1999 .

[23]  P. Reich,et al.  Generality of leaf trait relationships: a test across six biomes: Ecology , 1999 .

[24]  P. S. Karlsson,et al.  Leaf life span and nutrient resorption as determinants of plant nutrient conservation in temperate‐arctic regions , 1999 .

[25]  Sandra Díaz,et al.  Leaf structure and defence control litter decomposition rate across species and life forms in regional floras on two continents , 1999 .

[26]  Hendrik Poorter,et al.  A comparison of specific leaf area, chemical composition and leaf construction costs of field plants from 15 habitats differing in productivity , 1999 .

[27]  F. S. Chapin,et al.  The Mineral Nutrition of Wild Plants Revisited: A Re-evaluation of Processes and Patterns , 1999 .

[28]  Matthias Diemer,et al.  LEAF LIFESPANS OF HIGH-ELEVATION, ASEASONAL ANDEAN SHRUB SPECIES IN RELATION TO LEAF TRAITS AND LEAF HABIT , 1998 .

[29]  M. Diemer Life span and dynamics of leaves of herbaceous perennials in high‐elevation environments: ‘news from the elephant’s leg’ , 1998 .

[30]  Richard J. Williams,et al.  Carbon and nitrogen isotope discrimination and nitrogen nutrition of trees along a rainfall gradient in northern Australia , 1998 .

[31]  P. Reich,et al.  From tropics to tundra: global convergence in plant functioning. , 1997, Proceedings of the National Academy of Sciences of the United States of America.

[32]  J. Cornelissen,et al.  Functional leaf attributes predict litter decomposition rate in herbaceous plants. , 1997, The New phytologist.

[33]  M. F. Choong What makes a leaf tough and how this affects the pattern of Castanopsis fissa leaf consumption by caterpillars , 1996 .

[34]  W. Whitford,et al.  High Foliar Nitrogen in Desert Shrubs: An Important Ecosystem Trait or Defective Desert Doctrine? , 1996 .

[35]  Brian W. Darvell,et al.  A portable fracture toughness tester for biological materials , 1996 .

[36]  D. Hochuli The ecology of plant/insect interactions : implications of digestive strategy for feeding by phytophagous insects , 1996 .

[37]  D. Ackerly Canopy Structure and Dynamics: Integration of Growth Processes in Tropical Pioneer Trees , 1996 .

[38]  Robert Singerman Biomechanics—Materials—A practical approach , 1995 .

[39]  W. Wright,et al.  A comparative study of the fracture properties of five grasses , 1995 .

[40]  E. Garnier,et al.  Leaf anatomy, specific mass and water content in congeneric annual and perennial grass species , 1994 .

[41]  I. Turner Sclerophylly: primarily protective? , 1994 .

[42]  D. F. Parkhurst,et al.  Diffusion of CO2 and other gases inside leaves. , 1994, The New phytologist.

[43]  J. P. Grime,et al.  Methods in comparative plant ecology : a laboratory manual , 1993 .

[44]  J. P. Grime,et al.  Methods in Comparative Plant Ecology , 1993, Springer Netherlands.

[45]  P. Lucas,et al.  Leaf fracture toughness and sclerophylly: their correlations and ecological implications , 1992 .

[46]  P. Reich,et al.  Leaf Life‐Span in Relation to Leaf, Plant, and Stand Characteristics among Diverse Ecosystems , 1992 .

[47]  Julian F. V. Vincent,et al.  Biomechanics--materials : a practical approach , 1992 .

[48]  M. Pagel,et al.  The comparative method in evolutionary biology , 1991 .

[49]  A. Grafen The phylogenetic regression. , 1989, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[50]  F. Chapin,et al.  The Cost of Tundra Plant Structures: Evaluation of Concepts and Currencies , 1989, The American Naturalist.

[51]  A. Specht,et al.  Canopy structure in Eucalyptus-dominated communities in Australia along climatic gradients , 1989 .

[52]  Robert W. Pearcy,et al.  Plant Physiological Ecology , 1989, Springer Netherlands.

[53]  Steven Vogel,et al.  Life's Devices: The Physical World of Animals and Plants , 1988 .

[54]  S. Mori Insects and the Plant Surface , 1987 .

[55]  S. Caporn,et al.  THE INFLUENCE OF TEMPERATURE, WATER AND NUTRIENT CONDITIONS DURING GROWTH ON THE RESPONSE OF BRASSICA OLERACEA L. TO A SINGLE, SHORT TREATMENT WITH SIMULATED ACID RAIN , 1987 .

[56]  Christopher B. Field,et al.  photosynthesis--nitrogen relationship in wild plants , 1986 .

[57]  J. Felsenstein Phylogenies and the Comparative Method , 1985, The American Naturalist.

[58]  P. Coley,et al.  HERBIVORY AND DEFENSIVE CHARACTERISTICS OF TREE SPECIES IN A LOWLAND TROPICAL FOREST , 1983 .

[59]  David J. Hicks,et al.  The Ecology of Leaf Life Spans , 1982 .

[60]  Gordon H. Orians,et al.  A Cost-Income Model of Leaves and Roots with Special Reference to Arid and Semiarid Areas , 1977, The American Naturalist.

[61]  F. James Rohlf,et al.  Biometry: The Principles and Practice of Statistics in Biological Research , 1969 .

[62]  J. M. Cherrett A Simple Penetrometer for Measuring Leaf Toughness in Insect Feeding Studies , 1968 .

[63]  A. Loveless Further Evidence to Support a Nutritional Interpretation of Sclerophylly , 1962 .

[64]  E. Pitman A NOTE ON NORMAL CORRELATION , 1939 .

[65]  R. Knight The Plant in Relation to Water , 1932, Nature.

[66]  N. A. Maksimov,et al.  The plant in relation to water : a study of the physiological basis of drought resistance , 1929 .