title: The leaf size-twig size spectrum and its relationship to other important spectra of variation among species.
abstract: There is a spectrum from species with narrow, frequently branched twigs carrying small leaves and other appendages, to species with thick twigs carrying large leaves and appendages. Here we investigate the allometry of this spectrum and its relationship to two other important spectra of ecological variation between species, the seed mass-seed output spectrum and the specific leaf area-leaf lifespan spectrum. Our main dataset covered 33 woody dicotyledonous species in sclerophyll fire-prone vegetation on low nutrient soil at 1,200 mm annual rainfall near Sydney, Australia. These were phylogenetically selected to contribute 32 evolutionary divergences. Two smaller datasets, from 390 mm annual rainfall, were also examined to assess generality of cross-species patterns. There was two to three orders of magnitude variation in twig cross-sectional area, individual leaf size and total leaf area supported on a twig across the study species. As expected, species with thicker twigs had larger leaves and branched less often than species with thin twigs. Total leaf area supported on a twig was mainly driven by leaf size rather than by the number of leaves. Total leaf area was strongly correlated with twig cross-section area, both across present-day species and across evolutionary divergences. The common log-log slope of 1.45 was significantly steeper than 1. Thus on average, species with tenfold larger leaves supported about threefold more leaf area per twig cross-section, which must have considerable implications for other aspects of water relations. Species at the low rainfall site on loamy sand supported about half as much leaf area, at a given twig cross-section, as species at the low rainfall site on light clay, or at the high rainfall site. Within sites, leaf and twig size were positively correlated with seed mass, and negatively correlated with specific leaf area. Identifying and understanding leading spectra of ecological variation among species is an important challenge for plant ecology. The seed mass-seed output and specific leaf area-leaf lifespan spectra are each underpinned by a single, comprehensible trade-off and their consequences are fairly well understood. The leaf-size-twig-size spectrum has obvious consequences for the texture of canopies, but we are only just beginning to understand the costs and benefits of large versus small leaf and twig size.
authors.affiliation: Department of Biological Sciences, Macquarie University, NSW 2109, Australia. mwestoby@rna.bio.mq.edu.au
authors.forename: Mark
authors.initials: M
authors.lastname: Westoby
authors.forename: Ian J
authors.initials: IJ
authors.lastname: Wright
mesh_terms: D000704:Analysis of Variance; D018533:Biomass; D004777:Environment; D019684:Magnoliopsida / Q000033:anatomy & histology* / Q000235:genetics; D009517:New South Wales; D010802:Phylogeny; D018515:Plant Leaves / Q000033:anatomy & histology*; D018547:Plant Stems / Q000033:anatomy & histology*; D012639:Seeds / Q000502:physiology; D013045:Species Specificity